The Subfamily Mutisioideae (Asteraceae) - Springer Link

Bot. Rev. (2008) 74:469–716 DOI 10.1007/s12229-008-9016-6

The Subfamily Mutisioideae (Asteraceae) Liliana Katinas1,54 & John Pruski2 & Gisela Sancho1 & María Cristina Tellería3 1

División Plantas Vasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, USA 3 Laboratorio de Sistemática y Biología Evolutiva (LASBE), Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina 4 Author for Correspondence; e-mail: [email protected] 2

Published online: 9 December 2008 # The New York Botanical Garden 2008

Abstract The subfamily Mutisioideae (74 genera, ca. 865 species), which comprises three tribes, Mutisieae (43 genera, ca. 500 species), Nassauvieae (25 genera, ca. 315 species), and Stifftieae (six genera, 48 species), is analyzed at the generic level. A total of 87 genera traditionally related to Mutisioideae were studied. The genera of Mutisioideae are described and illustrated, and keys to tribes and genera are given. Corollas, anthers, styles, and pollen, are analyzed and discussed. The styles (smooth, rugulose to papillose, papillae short and rounded) constitute the main characters for delimiting the subfamily. The presence and distribution of the style papillae support the recognition of three tribes, although other characters as corolla shape contribute to their definition. The pollen of Nassauvieae can be clearly differentiated from the pollen of Mutisieae and Stifftieae. The morphology supports the exclusion of Brachylaena, Cloiselia, Dicoma, Duseniella, Erythrocephalum, Gladiopappus, Hesperomannia, Macledium, Moquinia, Pasaccardoa, Pleiotaxis, Tarchonanthus, and Warionia but not the exclusion of other taxa, for example, Hecastocleis, the Gochnatia complex, or the Ainsliaea group members. Resumen La subfamilia Mutisioideae (74 géneros, ca. 865 especies), que comprende tres tribus, Mutisieae (43 géneros, ca. 500 especies), Nassauvieae (25 géneros, ca. 315 especies), y Stifftieae (6 géneros, 48 especies), es analizada a nivel genérico. Se estudió un total de 87 géneros que tradicionalmente fueron relacionados a Mutisioideae. Se describen e ilustran los géneros de Mutisioideae, y se proveen claves para las tribus y los géneros. Se analizan y discuten las corolas, anteras y estilos. Los caracteres del estilo (glabro, ruguloso a papiloso, las papilas cortas y redondeadas) son los más importantes para delimitar la subfamilia. La presencia y distribución de las papilas del estilo ayudan al reconocimiento de las tres tribus, aunque otros caracteres como la forma de la corola contribuyen a su definición. El polen de Nassauvieae se puede diferenciar claramente del polen de Mutisieae y Stifftieae. La morfología apoya la exclusión de Brachylaena, Cloiselia, Dicoma, Duseniella, Erythrocephalum, Gladiopappus, Hesperomannia, Macledium, Moquinia, Pasaccardoa, Pleiotaxis, Tarchonanthus, y Warionia, pero no la exclusión de otros taxa, por ejemplo, Hecastocleis, los miembros del complejo Gochnatia o del grupo Ainsliaea.

470

L. Katinas, et al.

Keywords Asteraceae . Mutisioideae . Corolla . Style . Pollen

Introduction In the taxonomic history of the family Asteraceae, two significant classification systems, one by Cassini (1819) and the other by Bentham (Bentham & Hooker, 1873), were proposed. Cassini (1816, formally 1819) established 19 tribes, among them the Mutisieae and Nassauvieae. Bentham (Bentham & Hooker, 1873) proposed a system of 13 tribes, including the tribe Mutisieae (as Mutisiaceae), which was divided into five subtribes. Bentham’s system mostly prevailed years, with changes in the circumscription of the subtribes (e.g., Hoffmann, 1894; Cabrera, 1961, 1977; Hansen, 1990; Bremer, 1994; Table 1), until recent years with the appearance of molecular phylogenies. Cassini (1816, 1819) proposed the tribe Mutisieae to include the genera with bilabiate corollas considered by Lagasca (1811) in his Chaenanthophorae and by de Candolle (1812) in his Labiatiflorae. Cassini emphasized the style characteristics over the corolla morphology and established two related tribes: Mutisieae and Nassauvieae. The tribe Mutisieae included genera with short style branches internally uniformly covered by stigmatic papillae and externally covered by short hairs: “…ses deux stigmatophores courts, non divergentes, un peu arqués en dedans, demi-cylindriques, arrondies au sommet qui est un peu épaissi; leur face interne plane porte deux très-petits bourrelets stigmatiques marginaux confluens au sommet,

Table 1 Main Taxonomical Changes in the Classification and Nomenclature of the Subfamily Mutisioideae Lagasca (1811)

de Candolle (1812)

“Subfamily” Chaenanthophorae

“Subfamily” Labiatiflorae

Sect. III Sect. II

Sect. I

Cassini (1819)

Cassini (1824, 1825)

Lindley (1829)

Tribe Mutisieae

Tribe Mutisieae

–

– Sect. I: Archetypae

“Subfamily” Mutisioideae (as “Suborder Mutisieae”) – Tribe Archetypae

Sects. II, III, IV Sect. I

Tribe Nassauvieae

Sect. II: Gerberieae Tribe Nassauvieae

Sect. I: Archetypae Sect. II: Trixideae

Higher taxonomic categories are in bold

Tribe Gerberieae “Subfamily” Nassauvioideae (as “Suborder Nassauvieae”) Tribe Archetypae Tribe Trixideae

Mutisioideae (Asteraceae)

471

et souvent imperceptibles; leur face externe convexe porte sur sa partie supérieure quelques petits collecteurs papilliformes, épars” (Cassini, 1819: 199). The Nassauvieae, on the other hand, included genera with long, truncate style branches, with a crown of collector hairs at the apex: “…ses deux stigmatophores d’une longueur moyenne, divergens, arqués en dehors, demi-cylindriques, tronqués au sommet qui est un peu épaissi; leur face interne plane porte deux très-petits bourrelets stigmatiques marginaux, souvent imperceptibles; leur troncature terminale est garnie d’une touffé de collecteurs piliformes” (Cassini, 1819: 198). The Mutisioideae were first proposed by Lindley (1829) as his suborder Mutisieae with two tribes, Archetypae (containing Chaetanthera Ruiz & Pav.) and Gerbereae (with Chaptalia Vent., Gerbera L., and Perdicium L.) (Table 1). Lindley (1829) also established the Nassauvioideae as his suborder Nassauvieae, with the tribes Archetypae (including Triptilion Ruiz & Pav.) and Trixideae (including Jungia L. f. and Trixis P. Browne). Lindley’s subfamily ranks were not used by further authors until Bremer in 1996. In 1830, Don established the tribe Stifftieae characterized by bisexual florets with tubular five-lobed corollas, stamens inserted in the corolla throat, with caudate anthers and separate and recurved style branches. Don’s tribe Stifftieae included the genera Stifftia J. C. Mikan, Anastraphia D. Don [= Gochnatia Kunth], and Pentaphorus D. Don [= Gochnatia]. After Cassini, Lessing (1832) made another important contribution to the tribal classification of Asteraceae. Lessing also emphasized style features and reduced the number of Cassini’s tribes from 19 to eight. Two of Lessing’s tribes were Mutisiaceae, divided into three subtribes (Mutisieae, Lerieae, and Facelidae), and Nassauviaceae, divided into two subtribes (Nassauvieae and Trixideae) (Table 1). Table 1 (continued) Don (1830)

Lessing (1832)

“Subfamily” Labiatiflorae Tribe Mutisiaceae Tribe Barnadeseae Tribe Mutiseae

de Candolle (1838) “Subfamily” Labiatiflorae Tribe Mutisiaceae

Subtribe Mutisieae

Tribe Chaetanthereae Tribe Stifftieae

Tribe Perdiceae Tribe Polyachyreae Tribe Trixideae Tribe Jungeae

Hoffmann (1894)

Tribe Mutisiaceae Subtribe Barnadesieae Subtribe Gerbereae

Tribe Mutisieae

Subtribe Onoserideae Subtribe Gochnatieae Subtribe Nassauvieae

Tribe Diazeuxeae Subtribe Facelideae Subtribe Lerieae Tribe Nassauviaceae Subtribe Nassauvieae Subtribe Trixideae

Bentham & Hooker (1873)

Tribe Nassauviaceae

Subtribe Mutisinae (in text, Gerberinae in key)

Subtribe Gochnatinae SubtribeNassauvinae

472

L. Katinas, et al.

As previously pointed out, Bentham, another great contributor to the family’s classification, produced in 1873 a system of 13 tribes. His tribe Mutisiaceae included the five subtribes Barnadesieae, Onoserideae, Gochnatieae, Gerberieae, and Nassauvieae, which were reduced to three by Hoffmann in 1894 (Gochnatinae, Mutisinae [Gerberinae in the key] and Nassauvinae) (Table 1). Bentham used a wider spectrum of different morphological characters than did previous authors. He emphasized types of capitula in his classification and included in the Mutisieae genera with other types of corolla than the bilabiate, such as ligulate and tubular corollas. In the twentieth century, Jeffrey (1967) revised African Mutisieae and established 12 informal series mainly based on style morphology. Some of Jeffrey’s series, such as the Dicoma Cass., Pertya Sch. Bip., and Pleiotaxis Steetz series, were the basis, with much re-arrangement, of some generic groups that would be recognized by future authors (e.g., Hansen 1991a; Bremer, 1994). On the other hand, the Stifftia series, defined by rounded, glabrous, and short style branches and glabrous corollas, has been substantially modified from Jeffrey’s concept. The genera included by Jeffrey in the Stifftia series were Achnopogon Maguire, Steyerm. & Wurdack, Cnicothamnus Griseb., Duidaea S. F. Blake, Gochnatia, Glossarion Maguire & Wurdack, Gongylolepis R. H. Schomb., Nouelia Franch., Oldenburgia Less., Quelchia N. E. Br., Stifftia, Stomatochaeta (S. F. Blake) Maguire & Wurdack, and Wunderlichia Riedel ex Benth. Some of these genera were later studied in more detail (e.g., Pruski, 1997; Ortiz, 2000; Freire et al., 2002) and shown to have papillose styles and/or pubescent corollas instead of glabrous as treated by Jeffrey.

Table 1 (continued) Cabrera (1961, 1977)

Subfamily Cichorioideae Tribe Mutisieae Subtribe Barnadesiinae Subtribe Mutisiinae

Subtribe Gochnatiinae Subtribe Nassauviinae

Bremer & Jansen (1992) Subfamily Barnadesioideae Subfamily Cichorioideae Tribe Mutisieae – (subtribal categories not treated)

Bremer (1994)

Panero & Funk (2002)

Subfamily Barnadesioideae Subfamily Cichorioideae Tribe Mutisieae

Subfamily Barnadesioideae

Subfamily Mutisioideae

Subfamily Mutisioideae Tribe Mutisieae

–

–

–

Subtribe Mutisiinae s.l.

Subfamily Gochnatioideae Subfamily Hecastocleidoideae “Stifftia Group”

Tribe Stifftieae

Subfamily Mutisioideae

Tribe Nassauvieae

Subtribe Nassauviinae

Pruski (2004) and present paper Subfamily Barnadesioideae


473

Crisci (1974a, 1980) treated the subtribe Nassauviinae, performing detailed morphological and palynological studies and establishing its monophyly and the relationships among its genera. Cabrera’s paper on Mutisieae (1977), first presented at the International Symposium on the Biology and Chemistry of Asteraceae in 1975, can be viewed as the first, modern systematic review of the tribe Mutisieae. Cabrera divided Mutisieae into four subtribes: Barnadesiinae, Gochnatiinae, Mutisiinae, and Nassauviinae, following his previous classification (Cabrera, 1961). In the subtribe Barnadesiinae, he included several genera from Hoffmann’s subtribes Gochnatiinae and Mutisiinae. Of the four subtribes, he considered the Nassauviinae as the most natural in the tribe, morphologically homogeneous and geographically well-defined. Cabrera defined the Mutisieae as having bilabiate corollas, caudate anthers, and characteristic style shape. He pointed out, however, that several genera of Mutisieae differ in having actinomorphic or ligulate corollas and ecaudate anthers (the last feature in some genera of Barnadesiinae). Cabrera concluded that “…there is no doubt that certain genera must be excluded from the tribe or transferred to other subtribes; perhaps genera of other tribes should be included in the Mutisieae” (Cabrera, 1977: 1041). As circumscribed by Cabrera (1977), the Mutisieae included 83 genera (Table 2) and about 950 mainly American species, with some representatives in southern Asia and Africa and one species in Australia. The four subtribes were mainly distinguished as follows: central florets actinomorphic in Barnadesiinae and Gochnatiinae versus bilabiate in Mutisiinae and Nassauviinae; corollas villous and axillary spines in Barnadesiinae versus non villous and absence of axillary spines in Gochnatiinae; style branches rounded at the apex without collector hairs in Mutisiinae versus truncate with a crown of collector hairs in Nassauviinae. Cabrera distinguished papillae (very short, conical to rounded hairs, spread on the outer side of the style branches) from collector hairs (longer, cylindrical, obtuse at the apex hairs, that form a truncate brush terminally at the style branches), restricting the use of the term collector hairs to Nassauviinae. Jansen and Palmer (1987) found that a large chloroplast DNA inversion present in Asteraceae is absent in some genera of the subtribe Barnadesiinae, which were then put in their own subfamily (Bremer & Jansen, 1992). With this change, the Asteraceae were divided into three subfamilies: Barnadesioideae with one tribe, Cichorioideae with six tribes (one the Mutisieae), and Asteroideae with ten tribes. At this point, Mutisieae began to attract the attention of many workers because of its still unresolved taxonomy and its ancestral position, highlighting its importance for understanding the classification and evolution of the whole family. Also in 1987, Bremer presented the first cladistic analysis of Asteraceae involving 81 characters of morphology, anatomy, chromosomes, chemistry, and the chloroplast DNA inversion. This study showed the Mutisieae as a paraphyletic grade at the base of the family. One year later, Jansen and Palmer (1988) obtained phylogenies of Mutisieae based on chloroplast DNA restriction site mapping. In this second study, the Barnadesiinae were basal, the Nassauviinae monophyletic, and the Mutisiinae and Gochnatiinae were either monophyletic (using ten restriction enzymes) or paraphyletic (using nine additional restriction enzymes). Hansen (1991a), considered the tribe Mutisieae as monophyletic on the basis of petal “mutisioid” epidermal pattern (tabular cells with crested outer cell walls) if the

474

L. Katinas, et al.

Table 2 Comparative List of the Genera Within Mutisioideae Considered by Cabrera (1977) and Bremer (1994) and in the Present Work Cabrera (1977)

Changes considered by Bremer (1994)

Changes considered in this work

Subtribe Barnadesiinae Arnaldoa Barnadesia Chuquiraga Dasyphyllum Doniophyton Fulcaldea Huarpea Schlechtendahlia Subtribe Gochnatiinae Achyrothalamus Actinoseris Ainsliaea Aphyllocladus Not considered Chimantaea Chucoa Cnicothamnus Cyclolepis Diaspananthus Dicoma Duseniella Erythrocephalum Gochnatia Gypothamnium Hecastocleis Hesperomannia Hochstetteria Hyalis Not considered Lycoseris Macroclinidium Moquinia Nouelia Oldenburgia Onoseris Pasaccardoa Pertya Plazia Pleiotaxis Quelchia Stenopadus Stifftia Stomatochaeta Not considered Urmenetea Warionia Wunderlichia Subtribe Mutisiinae Achnopogon Not considered Brachyclados Cardonaea Catamixis

To Barnadesioideae To Barnadesioideae To Barnadesioideae To Barnadesioideae To Barnadesioideae To Barnadesioideae To Barnadesioideae To Barnadesioideae Subtribe Mutisiinae s. l. Achyrothalamus Actinoseris Ainsliaea Aphyllocladus Brachylaena Chimantaea Chucoa Cnicothamnus Cyclolepis = Ainsliaea Dicoma To Barnadesioideae Erythrocephalum Gochnatia Gypothamnium Hecastocleis Hesperomannia = Dicoma Hyalis Not considered Lycoseris Macroclinidium To Vernonieae Nouelia Oldenburgia Onoseris Pasaccardoa Pertya Plazia Pleiotaxis Quelchia Stenopadus Stifftia Stomatochaeta Tarchonanthus Urmenetea Unassigned to tribe Wunderlichia Subtribe Mutisiinae s. s. Achnopogon To Nassauviinae Brachyclados = Gongylolepis Catamixis

–a –a –a –a –a –a –a –a Tribe Mutisieae (= Erythrocephalum)a Actinoseris Ainsliaea Aphyllocladus –a To Stifftieae Chucoa Cnicothamnus Cyclolepis = Ainsliaea –a –a –a Gochnatia Gypothamnium Hecastocleis –a –a Hyalis Ianthopappus Lycoseris Macroclinidium –a Nouelia Oldenburgia Onoseris –a Pertya Plazia –a To Stifftieae To Stifftieae To Stifftieae To Stifftieae –a Urmenetea –a To Stifftieae Tribe Mutisieae Achnopogon Adenocaulon Brachyclados Gongylolepis Catamixis


475

Table 2 (continued) Cabrera (1977)



Chaetanthera Chaptalia Dinoseris Duidaea Not considered Eurydochus Gerbera Glossarion Gongylolepis Guaicaia Hyaloseris Leibnitzia Not considered Mutisia Myripnois Neblinaea Pachylaena Perdicium Piloselloides Not considered Trichocline Uechtritzia

Chaetanthera Chaptalia = Hyaloseris Duidaea To Nassauviinae =Gongylolepis Gerbera Glossarion Gongylolepis = Glossarion Hyaloseris Leibnitzia Lulia Mutisia Myripnois Neblinaea Pachylaena Perdicium = sect. of Gerbera Not considered Trichocline Uechtritzia

To Gochnatiinae To Gochnatiinae To Gochnatiinae To Gochnatiinae To Gochnatiinae To Gochnatiinae Subtribe Nassauviinae Acourtia Not considered Ameghinoa Not considered Not considered Calopappus Cephalopappus Not considered Dolichlasium Not considered Holocheilus Jungia Leucheria Leunisia Lophopappus Macrachaenium Marticorenia Moscharia Nassauvia Oxyphyllum Panphalea Perezia Pleocarphus Polyachyrus Proustia

To Mutisiinae s. l. To Mutisiinae s. l. To Mutisiinae s. l. To Mutisiinae s. l. To Mutisiinae s. l. To Mutisiinae s. l. Subtribe Nassauviinae Acourtia Adenocaulon Ameghinoa Not considered Burkartia = Nassauvia Cephalopappus Not considered Dolichlasium Eriachaenium Holocheilus Jungia Leucheria Leunisia Lophopappus Macrachaenium Marticorenia Moscharia Nassauvia Oxyphyllum Panphalea Perezia Pleocarphus Polyachyrus Proustia

Chaetanthera Chaptalia Dinoseris Duidaea Eriachaenium Eurydochus Gerbera Glossarion Gongylolepis Glossarion Hyaloseris Leibnitzia Lulia Mutisia Myripnois Neblinaea Pachylaena Perdicium = sect. of Gerbera Salcedoa Trichocline Uechtritzia Tribe Stifftieae Chimantaea Quelchia Stenopadus Stifftia Stomatochaeta Wunderlichia Tribe Nassauvieae Acourtia To Mutisieae Ameghinoa Berylsimpsonia Burkartia Calopappus Cephalopappus Criscia Dolichlasium To Mutisieae Holocheilus Jungia Leucheria Leunisia Lophopappus Macrachaenium Marticorenia Moscharia Nassauvia Oxyphyllum Panphalea Perezia Pleocarphus Polyachyrus Proustia

476

L. Katinas, et al.

Table 2 (continued) Cabrera (1977)



Triptilion Trixis Total genera: 89

Triptilion Trixis 76

Triptilion Trixis 74

a

Excluded herein from Mutisioideae

Barnadesiinae and several Old World genera of Gochnatiinae (sensu Cabrera, 1977) were excluded from the tribe. Hansen proposed to exclude from Mutisieae the genera Dicoma, Erythrocephalum Benth., Gladiopappus Humbert, Pasaccardoa Kuntze, and Pleiotaxis (and their current synonyms), later called the Dicoma group by Bremer (1994). Hansen also suggested the genera Adenocaulon Hook., Brachylaena R. Br., Tarchonanthus L., and Warionia Benth. & Coss. to be excluded from Mutisieae. Mutisioid ray corolla epidermal cells appear to be not exclusive of Mutisieae, they also occur in the Barnadesioideae and in some Arctoteae (Bremer, 1994). Robinson (1991) and formally Bremer (1994) abolished the limits between the Mutisiinae and the Gochnatiinae. Thus, the Mutisieae included only two subtribes: the Mutisiinae s.l. (Mutisiinae s. str. plus Gochnatiinae) and the Nassauviinae. The artificial taxonomic separation of the Mutisiinae and the Gochnatiinae was also confirmed by palynological evidence (Tellería et al., 2003). Karis et al. (1992) performed a morphology-based cladistic analysis of the family Asteraceae with emphasis on Cichorioideae on the basis of 53 taxa of Asteraceae and 72 characters, using the Barnadesioideae as outgroup. Their tree supported the placement of a paraphyletic tribe Mutisieae as a basal assemblage to the rest of Asteraceae. Genera of the Mutisieae (Stenopadus S. F. Blake, Hesperomannia A. Gray, Wunderlichia, Ainsliaea DC., and Gochnatia) appeared as isolated clades at the base of one of the two equally parsimonious final trees, other genera (Actinoseris (Endl.) Cabrera, Cnicothamnus, and Brachylaena) were sister to most Mutisieae, and others (Stifftia, Gongylolepis, Hyaloseris Griseb., and Pertya) were clades basal to the rest of Cichorioideae and Asteroideae. In Bremer’s (1994) treatment of the family Asteraceae, Mutisieae included 76 genera and ca. 970 species (Table 2). This treatment represented, after Cabrera (1977), a second, modern systematic review of the tribe. Bremer included in Mutisieae the subtribes Mutisiinae s.l. and Nassauviinae. Within Mutisiinae, he presented a tentative arrangement of generic groups, some already suggested by previous authors: the Stenopadus group (Achnopogon, Chimantaea Maguire, Steyerm. & Wurdack, Duidaea, Glossarion, Gongylolepis, Neblinaea Maguire & Wurdack, Stenopadus, Stomatochaeta); Gochnatia and related genera (Actinoseris, Chucoa Cabrera, Cyclolepis D. Don, Gochnatia, Hyalis D. Don ex Hook. & Arn., Nouelia); the Ainsliaea group (Ainsliaea, Macroclinidium Maxim., Myripnois Bunge, Pertya); Brachylaena and Tarchonanthus; the Plazia group (Aphyllocladus Wedd., Gypothamnium Phil., Plazia Ruiz & Pav.); the Dicoma group (Achyrothalamus O. Hoffm., Dicoma, Erythrocephalum, Gladiopappus, Pasaccardoa, Pleiotaxis); Onoseris Willd. and Urmenetea Phil.; Mutisia L. f. and Hyaloseris; the Chaetanthera group (Brachyclados D. Don, Chaetanthera, Pachylaena D. Don ex


477

Hook. & Arn.); the Gerbera group (Chaptalia, Gerbera, Leibnitzia Cass., Perdicium, Trichocline Cass., Uechtritzia Freyn). Further, Bremer’s Nassauviinae included 24 genera with ca. 320 species. The genera Catamixis Thomson, Cnicothamnus, Hecastocleis A. Gray, Hesperomannia, Lulia Zardini, Lycoseris Cass., Oldenburgia, Stifftia, and Wunderlichia were regarded as isolated in the tribe Mutisieae. Again, the monophyly of Nassauviinae and the paraphyly of Mutisiinae s. l. were recognized. Bremer (1996) later modified the traditional scheme of the subfamily Cichorioideae, which, until then, was constituted by the tribes Arctoteae, Cardueae, Lactuceae [= Cichorieae], Liabeae, Mutisieae, and Vernonieae. He treated the subfamily Cichorioideae as only containing Arctoteae, Lactuceae, Liabeae, and Vernonieae, with the tribe Cardueae the sole member of the subfamily Carduoideae. Although implicit in the cladogram, the Mutisieae were not raised to subfamilial level: it remained as an unresolved grade awaiting their reclassification, at the base of the family tree above the Barnadesioideae. Molecular phylogenies based on ndhF sequences (Jansen & Kim, 1996) mostly supported Bremer’s scheme but with a paraphyletic Cardueae. Pruski (2004) and Jeffrey (2004; for Stifftieae) resurrected the tribes Stifftieae and Nassauvieae. Pruski and Sancho (2004), following Bremer’s (1996) scheme, recognized the subfamily Mutisioideae as one of the five subfamilies of Asteraceae: Barnadesioideae, Mutisioideae, Carduoideae, Cichorioideae, and Asteroideae. In this contribution, morphologically based, we follow this scheme and consider the subfamily Mutisioideae to be comprised of three tribes: Mutisieae, Nassauvieae, and Stifftieae. Two molecular phylogenetic studies (Kim et al., 2002; Panero & Funk, 2002) investigated the circumscription of Mutisieae. Kim et al. (2002) used DNA sequences of the chloroplast gene ndhF from 31 genera and obtained an unsolved strict consensus tree showing that the Mutisieae are polyphyletic and that the genera Tarchonanthus, Brachylaena, Dicoma, Pasaccardoa, Ainsliaea, Pertya, and Myripnois are related to the Cardueae. Results of Panero and Funk (2002), using several chloroplast genes and molecular markers (ndhF, trnL-trnF, matK, ndhD, rbcL, rpoB, rpoC1, exon1, 23S-trnI, and ndhI), supported the establishment of the new subfamilies Gochnatioideae (Cnicothamnus, Cyclolepis, Gochnatia, Richterago Kuntze, possibly Chucoa), Hecastocleidoideae (Hecastocleis), and Pertyoideae (Ainsliaea, Diaspananthus Miq. [= Ainsliaea; Freire, 2007], Macroclinidium, Myripnois, Pertya). The tribes Dicomeae (Dicoma, Erythrocephalum, Gladiopappus, Macledium Cass., Cloiselia S. Moore, Pasaccardoa, Pleiotaxis), and Tarchonantheae (Brachylaena, Tarchonanthus), containing members of the Mutisieae s. l., were included in Carduoideae. Remaining Mutisieae were recognized as the Stifftia group and a basal, unresolved clade referred to as the Mutisioideae. No tribes, subtribes, or constituting genera were indicated by Panero and Funk within the Stifftia group and the Mutisioideae. Further, more defined molecular phylogenies of the family Asteraceae (Funk et al., 2005; Panero & Funk, 2008) showed approximately the same branching sequence of Panero and Funk’s (2002), with a Mutisieae clade containing three clades: the Onoseris clade, the Mutisia clade, and the Nassauvia clade. Based on these results, Panero & Funk (2007) established the following new taxa: the subfamily Stifftioideae (with Dinoseris, Duidaea, Eurydochus, Glossarion, Gongylolepis, Hyaloseris, Neblinaea, Quelchia, and

478

L. Katinas, et al.

Stifftia), the subfamily Wunderlichioideae with the tribes Wunderlichieae (Chimantaea, Stenopadus, Stomatochaeta, and Wunderlichia) and Hyalideae (Ianthopappus, Hyalis, Leucomeris, and Nouelia), and the tribe Onoserideae of Mutisioideae (with the genera Aphyllocladus, Gypothamnium, Lycoseris, Plazia, Onoseris, and Urmenetea). The systematic treatment by Hind (2007) regarded a tribe Mutisieae s. l., which was divided into five subtribes (Nassauviinae, Mutisiinae, Gerberinae, Gochnatiinae, and Tarchonanthinae) and seven groups (Catamixis, Dicoma, Hecastocleis, Nouelia, Pertya, Stenopadus, and Stifftia groups) and recognized two genera, Adenocaulon and Eriachaenium, as having a problematic placement. Finally, Katinas et al. (2008a) performed a molecular phylogenetic analysis of Nassauviinae based on the nuclear ITS and plastid trnL-trnF regions, with a sampling of 46 species of Mutisieae s. l. The study shows the monophyly of the Nassauviinae and a partial agreement with the tree of Funk et al. (2005) concerning generic relationships within the subtribe. Despite previous phylogenetic morphological and molecular studies, consensus has not been reached for the limits of Mutisioideae. Since its establishment in 1816 by Cassini and until recent years, the broadest concept of the tribe (Mutisieae sensu lato) was the prevailing one, with little fluctuation in the number of genera and species according to the different authors. After the two central revisionary works of Cabrera (1977) and Bremer (1994), much morphological and palynological work has been done to elucidate the taxonomic limits of Mutisioideae or at least to define generic groups (e.g., Pruski, 1991; Ortiz, 2000; Sancho, 2000, 2004; Freire et al., 2002; Tellería & Katinas, 2004). Moreover, new genera have been described (e.g., Zardini, 1980; Crisci & Freire, 1986; Katinas, 1994; Ortiz, 2001; Roque & Hind, 2001; Jiménez Rodríguez et al., 2004), and generic systematic revisions have been published (e.g., Egeröd & Stähl, 1991; Harling, 1995; Katinas, 1996b; Freire, 2007). There are currently two main taxonomic concepts of the Mutisieae that are in use. One is the concept of the tribe Mutisieae sensu lato, based mainly on morphological grounds. This is the concept employed, for example, by Cabrera (1977), Bremer (1994), Jeffrey (2007, as subfamily Mutisioideae), Hind (2007), and in this contribution (as subfamily Mutisioideae) with some genera variation in each circumscription. These authors recognized groups that were proposed at various times as distinctive ‘generic groups’ or associations, tribes, or subfamilies based on morphological and/or molecular data. The other concept, based on phylogenies obtained with chloroplast molecular data (e.g., Panero & Funk, 2002, 2007, 2008 as subfamily Mutisioideae; Funk et al., 2005), is that of the tribe Mutisieae sensu stricto (or subfamily Mutisioideae sensu stricto). This contribution is the result of the extensive work by the authors on the Mutisioideae, toward a comprehensive treatment of the subfamily. We refrain from prematurely creating taxonomic units within tribes. No new formal subtribes are established here, and generic groups are not given ranked status, since several lineages received taxonomic status as a result of the last molecular studies mentioned above. The objectives are (1) to describe and evaluate morphological and palynological characters of taxonomic value within the subfamily Mutisioideae and (2) to describe and illustrate the genera of Mutisioideae.


479

Material and Methods Information on the morphology of each genus was obtained from the literature and from our observations of herbarium and fresh specimens. These data result from original as well as from our previous work. In all, 73 of the 74 genera of Mutisioideae (specimens of Catamixis were not seen), ca. 700 of the ca. 860 species of Mutisioideae, 12 of the 13 excluded genera (specimens of Gladiopappus were not seen), and ca. 35 of the 127 excluded species were studied. The source literature is cited below each generic description, and types and representative herbarium specimens studied are cited under the Species list and in the corresponding figure legends. In general, the terminology follows Harris and Wolf Harris (1994) and Stearn (1996). Nomenclatural, taxonomic, and other observations were made only when pertinent. In general, the plant illustrations were drawn by Gisela Sancho mostly as modifications from earlier publications, although some drawings are originals; the sources of the figures that were redrawn are cited in the corresponding figure legend. A Wild M5 stereomicroscope and a Leitz SM Lux microscope both with the camera lucida technique were used for drawing the details. The style of Tables 5, 6, 7, 8, 9 and 10 is that of the tables of Harold Robinson in his work on Vernonieae (Robinson, 1999). Vegetative and floral parts were dissected and observed after boiling in water and staining with 2% safranin. For the scanning electron microscopy (SEM), portions of styles of FAA-fixed samples were taken, critical-point dried in CO2, and examined in a JEOL/EO JSM6360 microscope at Museo de La Plata, Argentina. Pollen grains were removed from herbarium specimens and were acetolyzed under standard methods and then chlorinated (Erdtman, 1960). For light microscopy (LM), slides were prepared by mounting the pollen in glycerol jelly and sealing with paraffin. Measurements of the polar (P) and equatorial (E) diameters of pollen grains were based on 25 grains, and measurements of the thickness of the exine were based on at least 15 grains. The P/E ratio was calculated for each specimen. Acetolyzed pollen grains were suspended in 90% ethanol, mounted on stubs for SEM, sputtercoated with gold palladium, and examined with a JEOL JSM T-100 scanning electron microscope. Palynological terminology, in general, follows Punt et al. (1994); the size classification was taken from Erdtman (1969). Pollen sources are cited in the figure legend of each genus and marked with an asterisk (*) in the Species list. Because the pollen of a broad sample of Nassauvieae was studied by Crisci (1974a), we emphasized the study of the pollen of tribes Mutisieae and Stifftieae herein. Two types of pollen descriptions are under each genus: (1) those containing measurements of the polar and equatorial diameters and exine thickness made for this study on the basis of the material marked with an asterisk (*) in the Species list, (2) those lacking measurements already published in previous papers where more detailed descriptions can be found, in which case we cite the relevant literature and important pollen features of the genus.

Morphological Characters The subfamily Mutisioideae has been subject to previous morphological and palynological studies, among them those of Wodehouse (1929a, b), Koch (1930),

480

L. Katinas, et al.

Carlquist (1957b, c, 1958), Parra and Marticorena (1972), Crisci (1974a), Baagøe (1977), Cabrera (1977), Skvarla et al. (1977), Grau (1980), Hansen (1991a, b), Freire and Katinas (1995), and Freire et al. (2002). Key characters for the classification of Mutisioideae are discussed here. Corolla Variation in Mutisioideae The bilabiate corolla was generally considered the typical form in the Mutisioideae. However, as noted by Cabrera (1977), corolla shape is highly variable in the group (Table 3, Fig. 1a–k). Four basic types of corollas are found in the group under study: (a) Bilabiate (3+2 corolla lips): zygomorphic, with an external three-dentate lip, radiating or not, and an internal two-cleft lip (Fig. 1d,f). (b) Tubular (0+5 corolla lobes): actinomorphic, limb five-lobed at the apex, the lobes usually deeply or very deeply incised (Fig. 1j) and less commonly shortly incised (Fig. 1k). These corollas are usually funnelform with a tube gradually expanding into a limb or less frequently sub-campanulate with the tube expanding into the limb. (c) Ligulate (5+0 ligule lobes): zygomorphic, with a short and narrow tube expanding into a ligule five-lobed at the apex, the lobes deeply (Fig. 1c) or shallowly incised (Fig. 1b). (d) True ray (3+0 limb lobes): zygomorphic, with a short and narrow tube expanding into a shallowly three-lobed limb (Fig. 1a). Modifications of these four basic types occur: (e) Tubular-bilabiate (intermediate between 3+2 corolla lips and 0+5 corolla lobes): sub-zygomorphic, transitional type where the lip-lobes are erect and short and the incisions in the corolla are unequal, resulting in a slightly bilabiate corolla (Fig. 1g,i). (f) Sub-bilabiate (or pseudobilabiate; 4+1, 3+1 corolla lips): zygomorphic, probably derived from a bilabiate corolla with different degrees of fusion among the teeth. The most common types are corollas with an external threedentate or four-dentate lip and one entire, untoothed, internal lip (Fig. 1e). (g) Filiform: irregular or zygomorphic, a very narrow, reduced tubular corolla with a short apical portion irregularly cleft, approaching other types of corollas (e.g., filiform-bilabiate, filiform-tubular, filiform-true ray) (Fig. 1h). The genera of Nassauvieae, with few exceptions (Acourtia, Lophopappus Rusby), have capitula with all bilabiate corollas. Most genera of Stifftieae have capitula with all tubular corollas; Quelchia is the only genus of the tribe that has tubular and bilabiate corollas. In Mutisieae, corolla morphology is more variable (Table 3), including genera that have capitula containing solely bilabiate corollas (e.g., Achnopogon, Pachylaena) or solely tubular corollas (e.g., Chucoa, Gochnatia). Some genera of Mutisieae are highly variable in corolla morphology (e.g., Lycoseris, Onoseris), and sometimes the variation appears within the same capitulum with


481

Table 3 Corolla Types in Genera of the Subfamily Mutisioideae Type of corolla

Tribe

Bilabiate

Mutisieae

Tubular

Ligulate 5-deeply lobed Ligulate 5-shortly lobed Sub-bilabiate Tubular and bilabiate Ligulate 5-shortly lobed and bilabiate Sub-bilabiate to tubular 4–5lobed/tubular True ray to sub-bilabiate/ bilabiate, tubular-bilabiate Bilabiate/bilabiate, tubular-bilabiate Tubular/tubular-bilabiate Bilabiate/tubular to tubular-bilabiate Broadly bilabiate, sub-bilabiate/tubular Broadly bilabiate/bilabiate/ bilabiate True ray/filiform (or absent)/ bilabiate, tubular-bilabiate True ray/tubular, tubular-bilabiate

Genera

Achnopogon, Brachyclados, Chaetanthera, Duidaea, Eurydochus, Gerbera, Glossarion, Gongylolepis, Lulia, Mutisia, Neblinaea, Trichocline, Uechtritzia Nassauvieae Acourtia, Ameghinoa, Berylsimpsonia, Burkartia, Calopappus, Cephalopappus, Criscia, Dolichlasium, Holocheilus, Jungia, Leucheria, Leunisia, Lophopappus, Macrachaenium, Marticorenia, Moscharia, Nassauvia, Oxyphyllum, Panphalea, Perezia, Pleocarphus, Polyachyrus, Proustia, Triptilion, Trixis Stifftieae Quelchia Mutisieae Chucoa, Cyclolepis, Gochnatia, Hecastocleis Nassauvieae Acourtia, Lophopappus Stifftieae Chimantaea, Quelchia, Stenopadus, Stifftia, Stomatochaeta, Wunderlichia Mutisieae Ainsliaea, Macroclinidium, Myripnois, Pertya Mutisieae Dinoseris, Glossarion, Mutisia, Hyaloseris Mutisieae Onoseris Nassauvieae Lophopappus Mutisieae Glossarion Mutisieae

Adenocaulon, Eriachaenium Chaetanthera, Lycoseris

Mutisieae

Perdicium, Leibnitzia

Mutisieae Mutisieae

Gochnatia Aphyllocladus, Lycoseris, Onoseris, Urmenetea

Mutisieae Mutisieae

Actinoseris, Aphyllocladus, Cnicothamnus, Gypothamnium, Hyalis, Ianthopappus, Nouelia, Oldenburgia, Onoseris, Plazia Chaptalia, Gerbera

Mutisieae

Chaptalia, Leibnitzia

Mutisieae

Lycoseris

Corollas separated by bars represent the different types of corolla in the same capitulum, from the marginal to the central corollas. In boldface are indicated genera that exhibit more than one type of corolla (or corolla arrangement)

transitional forms from the marginal to the central florets (e.g., Leibnitzia). Therefore, corolla morphology is not the main taxonomic character circumscribing the Mutisioideae, despite its use in previous treatments. The combination of the different types of corollas results in some basic types of capitulum (Bremer, 1994): (a) Discoid: with morphologically similar, non-radiating florets throughout the capitulum. The capitula may be composed by florets with all tubular (Fig. 1j) or all bilabiate (Fig. 1f) corollas.

482

L. Katinas, et al.

Fig. 1 Floral characters of Mutisioideae: A–K, corolla. A True ray (Chaptalia nutans). B Ligulate shallowly lobed (Dinoseris salicifolia). C Ligulate deeply lobed with one deeper split and unilateral lobes (Ainsliaea latifolia). D Bilabiate radiating (Onoseris alata). E Sub-bilabiate radiating (Cnicothamnus lorentzii). F Bilabiate (Criscia stricta). G Tubular-bilabiate (Chaptalia nutans). H Filiform (Chaptalia nutans). I Sub-bilabiate (Urmenetea atacamensis). J Tubular deeply-lobed (Gochnatia polymorpha). K Tubular shallowly-lobed (Onoseris hastata). L–O Anther apical appendages. L Truncate (Plazia daphnoides). M Acute (Brachyclados caespitosus). N Apiculate (Oldenburgia intermedia). O Constricted (Adenocaulon chilense). P, Q Anther tails. P Anthers caudate, with long tails (Cnicothamnus lorentzii). Q Anthers auriculate, with short tails (Adenocaulon chilense). R, S Styles. R Bifid, with long branches (Dinoseris salicifolia). S Bilobed, with short branches (Ianthopappus corymbosus)


483

(b) Radiate: with true ray corollas or some kind of distinctly radiating marginal florets (e.g., bilabiate, Fig. 1d). Pruski (1997) uses the term “bilabiate” for genera centered in the Guayana Highlands that have generally few-flowered capitula, with all bilabiate corollas, all of them with a large outer lip. (c) Ligulate: with all corollas possessing a strap-shaped limb with five apical teeth (Fig. 1b) or with deeply five-lobed corollas with one deeper split (Fig. 1c), these slightly ligulate and with unilateral lobes. (d) Disciform: with at least two morphological types of non radiating florets. Examples include: (1) marginal bilabiate and central tubular (some species of Aphyllocladus), (2) marginal bilabiate, sub-bilabiate, or true ray and central tubular-bilabiate (some species of Chaetanthera), (3) marginal subbilabiate, tubular four- or five-lobed and central tubular five-lobed (Adenocaulon), (4) marginal tubular four-lobed and central five-lobed (Eriachaenium Sch. Bip.). There are genera with three types of corollas in the same capitula (capitula trimorphic). For example, Chaptalia has true ray or bilabiate marginal corollas, filiform intermediate corollas, and tubularbilabiate central corollas. Comparison with Closer Subfamilies Corolla morphology helps to differentiate Mutisioideae from related subfamilies only to some degree. Bilabiate-like corollas exist in other Cichorioideae (e.g., Heterolepis Cass. in Arctoteae, Dipterocypsela S. F. Blake in Vernonieae), but they are not truly mutisioid bilabiate corollas (Bremer, 1994: 220). Tubular corollas in general gradually dilate into a limb (funnelform) in Mutisioideae. In Nouelia and Wunderlichia, the corollas abruptly dilate into a limb (campanulate) as occurs in Arctoteae, most Cardueae, and some Vernonieae and Liabeae. The Barnadesioideae are the most similar subfamily to the Mutisioideae regarding corolla shape, although in the former the corollas are covered by barnadesioid hairs. For instance, the genus Dasyphyllum Kunth (Cabrera, 1959b) has tubular, sub-bilabiate, and ligulate corollas; Fulcaldea Poir., Doniophyton Wedd., and Duseniella K. Schum. have tubular, funnelform corollas; Barnadesia Mutis has sub-bilabiate (4+1 corolla lips) marginal corollas and disc corollas that show all of the transitional morphological states including tubular-funnelform five-lobed, tubular three–four-lobed, subbilabiate (3+1, 4+1 corolla lips), sub-ligulate (with one deeper split), and ligulate five-dentate (Urtubey, 1999). Arnaldoa Cabrera, Chuquiraga Juss., and Schlechtendalia Less. have sub-bilabiate (4+1 corolla lips) corollas. Anthers Variation in Mutisioideae The anthers are typically caudate, with the pollen-bearing anther thecae extending below the attachment point of the filament (i.e., calcarate or spurred anthers; Robinson, 1983, 1999) (Fig. 1p). The anther collar was only found in Adenocaulon and Eriachaenium, and the antheropodium is common in some genera of

484

L. Katinas, et al.

Nassauvieae. The tails are short only in Adenocaulon and Eriachaenium (Fig. 1q). Tails are smooth (glabrous) or papillose, sometimes showing variation within the same genus (e.g., Chaptalia, Gochnatia, Onoseris, Trixis). Tails are usually free, but in some cases (e.g., Glossarion, Gongylolepis, Stenopadus) the tails of adjacent anthers are partially or completely connate. According to Karis et al. (1992), the endothecial thickening is polarized (Dormer, 1962) in Mutisioideae. The apical appendage is also well developed. In most genera, the appendage is acute (Fig. 1m), but apiculate (e.g., Gochnatia, Cnicothamnus, Chimantaea, some species of Ainsliaea) (Fig. 1n) and truncate apices (e.g., Aphyllocladus, Macrachaenium Hook. f., Plazia) (Fig. 1l) are also found. The apical appendages are neither constricted nor demarcated from the thecae, as commonly occurs in the subfamily Asteroideae, except in Adenocaulon and Eriachaenium (Fig. 1o). Exerted anthers are typical of some Guayana Highland-centered genera and are also present, for example, in Cnicothamnus and species of Gochnatia and Mutisia. In many genera of Mutisieae the anthers are reduced to staminodes in the functionally female florets. The staminodes range from diminute and filiform to very well developed with the appearance of sterile anthers, in some cases containing few pollen grains. We did not find important morphological differences among the anthers of Mutisieae, Stifftieae, and Nassauvieae. Comparison with Closer Subfamilies As occurs with the corolla, the anthers help to differentiate the Mutisioideae only to some extent. Long-caudate anthers are characteristic of Mutisioideae, most Carduoideae, some genera of Barnadesioideae, and part of tribes Lactuceae and Liabeae of Cichorioideae (vs. auriculate). In Barnadesioideae, anthers that are auriculate or decurrent with the filament, such as those of Barnadesia (Urtubey, 1999), are very common. Robinson (1999) considers stamens of most Cichorioideae s.l. (i.e., Mutisioideae, Carduoideae, and Cichorioideae s. str.) as calcarate. The polarized endothecial thickening of Mutisioideae is shared with Carduoideae and Cichorioideae, although in some tribes other types of thickening may exist (Bremer, 1994). Style Distinction of Tribes Style features constitute one of the most important characters to define Mutisioideae and to circumscribe tribes within the subfamily. The subfamily Mutisioideae is characterized by its glabrous to papillose styles, with the papillae or the relatively short hairs apically rounded. The stigmatic area is continuous, covering the whole inner surface of the style branches. In the tribe Mutisieae, the style branches are usually very short (styles bilobed, Fig. 1s, e.g., Ainsliaea, Trichocline), but there are also genera with long style branches (styles bifid, Fig. 1r, e.g., Dinoseris Griseb., Hyaloseris, Lycoseris). Variation of the style branch length, short or long, occurs also within the same genus


485

(e.g., Chaetanthera, Chaptalia, Gochnatia). The style branch apex is rounded in most genera, rarely truncate (e.g., Eurydochus Maguire & Wurdack, Plazia) or acute (e.g., Cnicothamnus, Chucoa, Macroclinidium). Sometimes the stigmatic surface apically projects upwards, forming an edge that gives an apiculate aspect to the branches (e.g., some species of Gochnatia, Salcedoa F. Jiménez Rodr. & Katinas) (Fig. 1s). Dorsally, the style branches are papillose in most genera (Fig. 77a), although smooth (e.g., Gochnatia, Hecastocleis, Gongylolepis) (Fig. 77b) or rugulose branches with swollen dorsal cells (Chimantaea) are present (Fig. 77d). The papillae or short hairs may be variable in length, but they are always rounded at the apex (Fig. 78a–c); acute hairs are completely lacking in Mutisioideae. The papillae are distributed above the style branch bifurcation point in most taxa, covering the whole branch or restricted to the terminal, apical portion of the branch (Fig. 77a). In a few cases, the papillae extend below the style branch bifurcation point (e.g., Pachylaena, Pertya). In the tribe Stifftieae (e.g., Stenopadus, Stomatochaeta, Wunderlichia), the papillae extend below the style branch bifurcation point toward the style base (Fig. 77c–d); the only exception is the genus Quelchia, which has smooth styles. In the tribe Nassauvieae, most of the styles have relatively long branches and papillae or collector hairs (Crisci, 1974a; Cabrera, 1977) that form a tuft restricted to the branch apex (Fig. 77e–f). Exceptions are Lophopappus and Macrachaenium, which have styles with dorsally papillose branches. Crisci (1974a) performed a careful study in all genera of Nassauviinae and determined four style types according to differences in branch length, apex shape, and papilla length and considered the papillae in the styles of Nassauvieae as “collector hairs.” The first type has truncate stylar branches (Ameghinoa Speg., Burkartia Crisci, Calopappus Meyen, Dolichlasium Lag., Holocheilus Cass., Jungia, Leucheria, Marticorenia Crisci, Moscharia Ruiz & Pav., Nassauvia, Oxyphyllum Phil., Perezia, Pleocarphus D. Don, Polyachyrus Lag., Triptilion, and Trixis). The second type has the branches less than 1 mm long, rounded at the apex, bearing long collector hairs on the distal half (Cephalopappus Nees & Mart., Lophopappus, Macrachaenium, and Proustia). The third type has the branches more than 1 mm long, rounded and slightly expanded at the apex, with long collector hairs on the distal fifth (Acourtia, Berylsimpsonia B. L. Turner, Leunisia Phil., and Perezia). The fourth type has style branches spathulate, with very long collector hairs in the distal third (Criscia Katinas). Considering the morphology and pubescence of styles, Erbar and Leins (1995) distinguished different mechanisms of secondary pollen presentation in the Campanulales–Asterales-complex. Two different mechanisms were established for Asteraceae: the brushing and the pump mechanisms. In the brushing mechanism, the back of the style branches and the upper part of the style bear short hairs. The anthers open and pollen is partly loaded onto the hairs of the style. As the style elongates, the pollen grains are completely swept out of the anthers by the brushing hairs. In the pump mechanism, the style branches are truncate with hairs only at their tips. The growing style acts as a piston and pushes the pollen gradually out of the anther tube. Given the style morphology of the tribes, the brushing mechanism would characterize members of Mutisieae and Stifftieae, and the pump mechanism would characterize Nassauvieae.

486

L. Katinas, et al.

Comparison with Other Subfamilies Unlike the styles found in Mutisioideae, the arctotoid and the carduoid styles (Bremer, 1987, 1994) have a ring of collector or sweeping hairs on the shaft below the style branch bifurcation point. In addition, the Arctoteae and many Cardueae have apically thickened styles, whereas the style of Mutisioideae is usually uniform in width. A few genera of Cardueae, such as Amphoricarpus Spruce ex Miers, Chardinia Desf., and Tugarinovia Iljin (subtribe Carlininae), were described as having shortly bilobed and apically minutely pilose styles (Bremer, 1994), thus resembling many genera of Mutisioideae. However, close examination with the light microscope shows that the style branches of these genera of Cardueae have their subapical papillae longer than their apical ones, thus approaching the subapical tuft of hairs typical of the tribe. The tribes Lactuceae, Liabeae, and Vernonieae of Cichorioideae have vernonioid styles (Bremer, 1987), with style branches apically tapered or rounded and with acute collector hairs (in some Vernonieae, there are rounded, multicelled collector hairs; Cabrera, 1944) distributed from the apex to below the branch bifurcation point. Although a continuous stigmatic surface on the style branches is shared among Mutisioideae, Carduoideae, and Cichorioideae, differences in style shape, papillae arrangement, and especially papilla shape (i.e., rounded in Mutisioideae vs. acute in Carduoideae and Cichorioideae, Fig. 78d–f) in the external side of the branches set the subfamily Mutisioideae apart from the other two subfamilies. Styles of the members of the subfamily Asteroideae have collector hairs, with different hair arrangements. In Asteroideae, the stigmatic surface is usually restricted to two bands (Carlquist, 1976), thus strongly differing from the condition in the Mutisioideae. However, styles of genera of Mutisioideae are more similar to those of the Barnadesioideae than to any other group of Asteraceae. For instance, smooth styles are found in Dasyphyllum, styles dorsally papillose below the branch bifurcation point such as those of tribe Stifftieae may be found in Chuquiraga, Dasyphyllum, Doniophyton, and Schlechtendalia (Cabrera, 1977), and styles papillose above the bifurcation point are found in Barnadesia (Cabrera, 1977; Urtubey, 1999). One might argue that these style characters are symplesiomorphic, however primitive and advanced states recognition for these features depends on the choice of the outgroup (Calyceraceae, Goodeniaceae, Menyanthaceae). Pollen Distinction of Tribes The pollen of Mutisioideae is a useful character mainly to distinguish the subfamily from other subfamilies and to distinguish Nassauvieae from Mutisieae and Stifftieae. The Mutisioideae would be considered an eurypalynous subfamily because of its wide range of variation in grain size, shape (P/E), aperture type, and exine sculpture and structure. This agrees with Skvarla et al. (1977) who concluded that the pollen of Mutisioideae is diverse at the exo- and endomorphic levels. It should be noted, however, that the major variation occurs in the pollen of the tribes Mutisieae and Stifftieae, whereas the pollen in the tribe Nassauvieae is more homogeneous.


487

The pollen of the genera of Nassauvieae is tricolporate, microechinate with few developed microspines, the exine distinctly bilayered, and both the ectosexine and the endosexine clearly columellate (Crisci, 1974a) (see 109B of Figs. 107–111 and 112B of Figs. 112–116). The pollen of Mutisieae and Stifftieae is microechinate, echinate, or almost psilate in some members of Mutisieae; the exine is distinctly bilayered but commonly with a compact ectosexine constituted by columellae very thickly disposed and an endosexine with columellae clearly distinguishable, stout and ramified (Hansen, 1991b; Tellería et al., 2003; Tellería & Katinas, 2004; Zao et al., 2006) (see 80C of Figs. 80–85, 98A of Figs. 95–100, 118B, C of Figs. 117–118 and Fig. 119c). Besides the characteristic exine structure, additional pollen characters reinforce the distinction of the Nassauvieae from the Mutisieae and Stifftieae: (1) the pollen of Nassauvieae and Stifftieae is always tricolporate, whereas in Mutisieae it can be tricolporate (most genera) or tricolpate (Hecastocleis and some species of Ainsliaea); (2) polar caps are more frequent in pollen of Nassauvieae (e.g., Ameghinoa, Leunisia, Trixis) than in pollen of Mutisieae and Stifftieae; (3) the spheroidal and suboblate pollen shape mainly occurs in Nassauvieae (e.g., Leucheria, Triptilion), whereas the pollen of Mutisieae and Stifftieae have generally subprolate to prolate shape; (4) in Mutisieae (e.g., Chaetanthera, Lulia, Mutisia) the mesoaperture is either clearly distinguishable or diffuse, whereas in Stifftieae it rarely occurs, and it was never reported in Nassauvieae; and (5) the pollen of Nassauvieae is generally smaller (see Material and Methods) than that of Mutisieae and Stifftieae (Parra & Marticorena, 1972; Crisci 1976b; Hansen, 1991b; Zao et al., 2006). Exine Types The organization of the exine consists of tectum, ectosexine, endosexine, and nexine. The ectosexine and the endosexine are columellate and they are separated by an internal tectum (clearly distinguishable or not); the nexine is usually thickened at level of aperture. The pollen of the genera of Nassauvieae may be distinguished on the basis of the thickness of the external and internal tectum and the orientation of the internal tectum. Crisci (1974a) distinguished five exine types; all of them have a tectum surface with very few developed microspines: (1) Oxyphyllum type: ectosexine and endosexine equally thick, separated from each other by zigzag (non-parallel to nexine) internal tectum (Leucheria, Moscharia, Nassauvia, Oxyphyllum, Polyachyrus, and Triptilion). (2) Trixis type: ectosexine thinner than endosexine, both layers columellate (Acourtia, Ameghinoa, Berylsimpsonia, Burkartia, Criscia, Dolichlasium, Holocheilus, Jungia, Leunisia, Lophopappus, Macrachaenium, Marticorenia, Pleocarphus, Proustia, and Trixis). (3) Proustia type: ectosexine and endosexine equally thick, both layers columellate (Lophopappus and Proustia). (4) Calopappus type: ectosexine thinner than endosexine, both layers columellate and separated by a zigzag internal tectum (Calopappus).

488

L. Katinas, et al.

(5) Cephalopappus type: ectosexine thicker than endosexine, both layers columellate and separated by an internal tectum more or less parallel to nexine (Cephalopappus). In the genera of Mutisieae and Stifftieae, differences in exine sculpture led us to classify six types of exine (Table 4). Three of them, the Gongylolepis, Macroclinidium, and Stenopadus types, are proposed here, increasing the typification made by Stix (1960), Crisci (1974a) and Tellería et al. (2003). The exine types are as follows: (1) Ainsliaea type (82B of Figs. 80–85) (Tellería et al., 2003; Tellería & Katinas, 2005): exine microechinate, tectum imperforate. Ectosexine and endosexine equally thick, both layers scarcely differentiated, with compact aspect or slightly columellate. Ratio ectosexine/endosexine: ca. 1:1. Exclusively present in some species of Ainsliaea of the tribe Mutisieae. (2) Gongylolepis type (94B of Figs. 92–94) (Tellería, 2008): exine echinate, with short or long spines (until ca. 6 μm length), spines with apical channel, tectum very perforate. Ectosexine thinner than endosexine, ectosexine slightly columellate, endosexine with stout and ramified columellae, both layers separated by a conspicuous internal tectum. Ratio ectosexine/endosexine among spines: 1:4; 1:5. Present in Glossarion rhodanthum Maguire & Wurdack, Gongylolepis benthamiana R. H. Schomb., G. huachamacari

Table 4 Distribution of Main Pollen Features Within Mutisieae and Stifftieae: Exine Type and Aperture Type

Shaded area indicates pollen of genera closest to the Carduoideae a Genera with tricolporate and tricolpate pollen. In boldface: Pollen of genera with two exine types.


(3)

(4)

(5)

(6)

489

Maguire, and Duidaea marahuacensis Steyerm. of the tribe Mutisieae, and in Quelchia of the tribe Stifftieae. Mutisia type (98A of Figs. 95–100) (Tellería et al., 2003; Tellería & Katinas, 2004): exine scabrate, microechinate or echinate with short spines (usually less than 2 μm), tectum imperforate or scarcely perforate. Broad range of exine thickness (4–18 μm). Ectosexine equal, thinner or thicker than endosexine, ectosexine slightly columellate or with compact aspect, endosexine with stout and ramified columellae, both layers separated by an inconspicuous or conspicuous internal tectum. Ratio ectosexine/endosexine: 1:1; 1:2; 1:2.5; 1.5:1; 2:1; 2.5:1. The most common type in the tribe Mutisieae and also present in the tribe Stifftieae. Macroclinidium type (100B of Figs. 95–100): exine echinate, with long spines (4–5 μm length), spines usually with compact tip, tectum very perforate. Ectosexine thinner than endosexine; ectosexine slightly columellate–granulate, endosexine with ramified columellae. Internal tectum conspicuous. Ratio ectosexine/endosexine among spines: ca. 1:3; 1:2. Exclusively present in Macroclinidium of the tribe Mutisieae. Stenopadus type (118B, C of Figs. 117–118) (Tellería, 2008): exine psilate (with light microscope), very perforate, with scarce and inconspicuous microspines (with SEM). Ectosexine and endosexine equally thick, ectosexine and endosexine columellate (apparently single columellae). Ratio ectosexine/ endosexine: 1:1. Exclusively present in species of Stenopadus and Stomatochaeta of the tribe Stifftieae. Wunderlichia type (121 of Figs. 120–121) (Tellería et al., 2003; Tellería, 2008): exine echinate, with short spines (less than 3 μm length), spines with apical channel, tectum very perforate. Ectosexine columellate, equal to or slightly thinner than endosexine, with compact inner layer and thinner scarcely columellate outer layer. Ratio ectosexine/endosexine: 1:1; 1:1.5. The Wunderlichia exine type is very similar to the Gongylolepis type but with less developed spines. The Wunderlichia type can be viewed as transitional between the Mutisia and the Gongylolepis types. Present in the tribe Mutisieae in Eurydochus, species of Gochnatia, and in Salcedoa and in the tribe Stifftieae in Wunderlichia.

Comparison with Closer Subfamilies Despite the diversity of pollen types that occurs in Asteraceae (e.g., Wodehouse, 2008; Erdtman, 1966), and despite the fact that the Anthemoid pattern pollen is present in Mutisioideae and also in the tribes Anthemideae (subfamily Asteroideae) and Cardueae (subfamily Carduoideae) (Skvarla & Turner, 1966; Skvarla et al., 1977; Bolick, 1978), the pollen of Mutisioideae can be distinguished from the pollen of other subfamilies. In the tribe Anthemideae, Skvarla et al. (1977) recognized two basic pollen types: Anthemis L. and Artemisia L. types, distinguished by the presence or absence (or great reduction) of spines respectively (see also Vezey et al., 1994). The spines in Anthemideae, when present, are generally large in proportion to the size of the grain (Wodehouse, 1935). In contrast, in the genera of Mutisioideae with echinate pollen, the spines are short when compared with the overall size of the pollen grain. The Artemisia exine type resembles the Mutisia exine type, but the

490

L. Katinas, et al.

grains of Anthemideae are small and spheroidal with a thin exine, whereas those of Mutisioideae (excluding the tribe Nassauvieae) are usually large and elliptic with a thick exine. The closest genera of Mutisioideae to the Artemisia exine type are Adenocaulon and Eriachaenium, which were once considered members of the tribe Anthemideae (Stix, 1960; Leins, 1968; Skvarla et al., 1977 for Adenocaulon). In comparison with the pollen in the tribes of Cichorioideae, pollen in the subfamily Mutisioideae is clearly different. Arctoteae, Lactuceae, Liabeae, and Vernonieae have echinate and echinolophate pollen, with caveate exine in Arctoteae and most Lactuceae, and an ecaveate exine in Liabeae and Vernonieae (Skvarla et al., 1977; Robinson & Marticorena, 1986; Robinson, 1999). Moreover, pollen of Liabeae is prominently echinate and the exine has much broader columellae than in the Anthemoid pattern (Skvarla et al., 1977), which characterizes the pollen of Mutisioideae. On the other hand, the Carduoideae have pollen resembling some genera of the Mutisieae and Stifftieae (Table 4). For instance, the exine of Macroclinidium (100 of Figs. 95–100) resembles that of Carduoideae, differing only in having solid spines rather than spines with an apical channel (Skvarla et al., 1977; Tormo-Molina & Ubera-Jiménez, 1995). The Guayana Highland-centered genera of Mutisieae and Stifftieae include the major number of pollen types close to the Carduoideae types (Tellería, 2008). Four exine types in Carduoideae (Tormo-Molina & Ubera-Jiménez, 1995) resemble those in Mutisioideae. (1) The Arctium L., Cyanopsis Cass., and Onopordum L. exine type is characterized by long spines with apical channels and an intricate layer of columellae under the tectum. This type closely resembles the Gongylolepis type and the Wunderlichia exine type (see Exine types). (2) The Carlina L. and Cyanus Mill. exine type is characterized by a microechinate surface, ectosexine with compact aspect, an endosexine thicker than the ectosexine, and stout and ramified columellae. This type is similar to the Mutisia exine type. (3) The ecaveate pollen of some species of Centaurea L. (Wagenitz, 1955; Nordenstam & El-Ghazaly, 1977; Pehlivan, 1995) is also very similar to the Mutisia exine type. (4) The particular Echinops L. pollen type, with its very thick exine (Dimon, 1971; Garnatje & Martin, 2007), resembles the pollen of some species of Chaetanthera (type II; Tellería & Katinas, 2004). These observations reinforce Wodehouse’s (1929b) suggestion of a possible relationship between genera of Mutisioideae and Carduoideae. The pollen of Barnadesioideae strongly differs from that of Mutisioideae. In Barnadesioideae, three main pollen types occur (Urtubey & Tellería, 1998): two of them are lophate or with a variable number of intercolpal depressions, both generally caveate; the third lacks the intercolpal depressions, the exine columellate–granulate or with compact aspect. All of these features are absent in Mutisioideae. Summary of the Morphological Observations We conclude the following: (1) Style features (smooth, rugulose to papillose styles, the papillae being relatively short and rounded) constitute the main characters delimiting the subfamily Mutisioideae. (2) Pollen morphology (i.e., pollen surface psilate, microechinate or echinate with a densely columellate exine) is secondarily helpful for delimitating the subfamily. (3) The presence and distribution of style papillae differentiate the three tribes: (a) those genera with smooth or with dorsally papillose styles with the papillae distributed above the branch bifurcation point


491

belong to the tribe Mutisieae (43 genera, ca. 500 species); (b) those with styles rugulose or papillose, both above and below the bifurcation point (almost reaching the base), belong to the tribe Stifftieae (six genera, 48 species); and (c) those with the papillae clustered in an apical tuft belong to the tribe Nassauvieae (25 genera, ca. 320 species). (4) The pollen of genera of Nassauvieae can be clearly differentiated from that of Mutisieae and Stifftieae, being microechinate, with few developed microspines, having a distinctly bilayered exine, and having both the ectosexine and the endosexine clearly columellate. The pollen of Mutisieae and Stifftieae is psilate, microechinate, or echinate, with a compact ectosexine constituted by very thickly disposed columellae and a clearly columellate endosexine. (5) The overall morphology of tribes Nassauvieae and Stifftieae is homogeneous, whereas tribe Mutisieae is heterogeneous for most of its morphological features (e.g., corolla shape, pollen morphology). (6) Morphological data support the molecular studies that exclude certain genera (see Excluded taxa) and disagree with the exclusion of other genera, for example Hecastocleis, and the members of the Gochnatia complex and the Ainsliaea group (see Circumscription of the tribes of Mutisioideae). (7) The morphologically closest subfamilies to Mutisioideae are Barnadesioideae in corolla shape (both have the same range of corolla types except that true ray and filiform corollas are present only in Mutisioideae) and style features (smooth, rugulose to papillose styles, the papillae being relatively short and rounded), and Carduoideae in pollen features (both have the Anthemoid exine pattern as well as some genera with echinate and highly perforate surfaces).

Taxonomic Circumscription Circumscription of the Subfamily Mutisioideae The subfamily Mutisioideae can be morphologically defined (Fig. 2) to include genera characterized by (1) frequent presence of bilabiate florets and tubular florets that are funnelform to sub-campanulate (vs. truly campanulate as in most Arctoteae, Cardueae, Liabeae, and Vernonieae); (2) styles with the same width from base to tip (vs. apically widened in Arctoteae and some genera of Cardueae); (3) styles smooth, rugulose, or with short and rounded papillae, the papillae distributed uniformly either only above or both above and below the branch bifurcation point, or constituting a tuft restricted to the branch apex (vs. vernonioid, carduoid, or arctotoid styles); (4) the anthers caudate with long tails, except for Adenocaulon and Eriachaenium (vs. shortly caudate or ecaudate in Arctoteae, Vernonieae, and certain genera of Liabeae and Barnadesioideae); (5) lack of barnadesioid hairs (Bremer & Jansen, 1992) which are restricted to Barnadesioideae; (6) pollen ecaveate (vs. caveate in Lactuceae, Arctoteae, and some genera of Cardueae and Barnadesioideae); (7) pollen surface almost psilate, microechinate or echinate (vs. lophate in some Arctoteae, Barnadesioideae, Lactuceae, and Vernonieae); and (8) exine structure always densely columellate with stout columellae (vs. with sparse and broad columellae in Liabeae and Vernonieae, never with the sponge-like complex of freely branching and anastomosing columellae structure as in the lophate type of Lactuceae and some Barnadesioideae).

492

L. Katinas, et al.

Fig. 2 Diagram of the classification of the subfamily Mutisioideae and excluded genera. The main morphological characters defining the Subfamily Mutisioideae, the tribes Mutisieae, Nassauvieae, and Stifftieae, and the generic groups of Mutisieae are illustrated

The style of Mutisioideae is obviously different from the carduoid, arctotoid, and vernonioid styles (Bremer, 1987) that characterize the subfamilies Carduoideae and Cichorioideae. However, styles of Mutisioideae and those of the subfamily Barnadesioideae are similar. Despite this resemblance, the subfamily Barnadesioideae is distinguished from Mutisioideae by a combination of two other morphological characters: exclusive presence of barnadesioid hairs and a pollen morphology very different from that of the Mutisioideae. The resemblance in style features between Mutisioideae and Barnadesioideae supports Cabrera’s (1961, 1977) inclusion of the Barnadesioideae as a subtribe of the tribe Mutisieae (= Mutisioideae). Circumscription of the Tribes of Mutisioideae Three tribes are distinguished within Mutisioideae: tribe Mutisieae, tribe Nassauvieae, and tribe Stifftieae (Fig. 2). Tribe Mutisieae The tribe Mutisieae is much more heterogeneous when compared with Nassauvieae and Stifftieae. A central core (Fig. 2, Tables 5 and 6) of genera have mostly bilabiate corollas, generally combined with other types of corolla, and dorsally papillose style branches with the papillae uniformly distributed above the branch bifurcation point


493

Table 5 Tribe Mutisieae, Central Core of Genera: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

Crossed out corolla indicates: Absent. 1 shrubs, 2 subshrubs, 3 perennial herbs, 4 annual herbs, 5 scapiform or sub-scapiform perennial herbs, 6 trees, 7 caulescent, perennial herbs, 8 trailing shrubs

494

L. Katinas, et al.

Table 5 (continued)

(Fig. 77a). Around these genera, a number of “satellite” genera exist, most of them defined as generic groups by Bremer (1994). Although with some distinguishing features, all of these groups have morphological characters that link them to Mutisioideae. Some of these groups of genera have been recently excluded from Mutisioideae on the basis of molecular phylogenetic analyses (Panero & Funk, 2002, 2008; Funk et al., 2005). A discussion of those generic groups follows: Gochnatia complex. The Gochnatia complex (Freire et al., 2008) comprises the genera Actinoseris, Cnicothamnus, Cyclolepis, Gochnatia, Hyalis, Ianthopappus Roque & D. J. N. Hind, and Nouelia and is defined by the combination of apiculate anther appendages and smooth style branches (Fig. 77b; Table 7). Hansen (1991a) and then Bremer (1994) treated some of these genera as the Gochnatia group. Cabrera (1971a) related Pleiotaxis and Chucoa to Actinoseris, Cyclolepis, and Gochnatia. Pleiotaxis and Chucoa, however, lack the characters typical of the Gochnatia complex, such as apiculate anther appendages (acute in Chucoa) and


495

Table 6 Tribe Nassauvieae: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

Crossed out corolla indicates: Absent. 1 perennial herbs, 2 scapiform, perennial herbs, 3 subshrubs, 4 shrubs, 5 clambering shrubs, 6 annual herbs, 7 scapiform annual herbs, 8 trees

496

L. Katinas, et al.

Table 6 (continued)

smooth styles (papillose in Chucoa and Pleiotaxis) (Freire et al., 2002; Sancho et al., 2005). Phylogenetic molecular data supported relationships between Gochnatia and Cnicothamnus (Kim et al., 2002). Panero and Funk (2002) created the subfamily Gochnatioideae, based also on molecular data, that includes Cnicothamnus, Cyclolepis, Gochnatia, Richterago (= Gochnatia sect. Discoseris (Endl.) Cabrera and Actinoseris pro parte), and possibly Chucoa. Nouelia and Gochnatia sect. Leucomeris (D. Don) Cabrera (sensu Cabrera, 1971a) were not included in the subfamily Gochnatioideae. Panero and Funk (2007) established the tribes Wunderlichieae and Hyalideae of the subfamily Wunderlichioideae, the latter including genera of the Gochnatia complex, i.e., Ianthopappus, Hyalis, Leucomeris (= Gochnatia sect. Leucomeris), and Nouelia. From a morphological point of view, the relationships between Hyalideae and Wunderlichieae are not strongly supported. There are other genera of the tribe Mutisieae s. l. with glabrous styles (e.g., Hecastocleis) or apiculate anthers (e.g., Ainsliaea, Wunderlichia), but they do not have both features together. On the other hand, some genera of the Guayanas (e.g., Gongylolepis, Quelchia) have smooth style branches and acuminate anthers, providing a link between the two generic groups. The genera of the Gochnatia complex in general share the Mutisia exine type, supporting their placement in Mutisioideae, tribe Mutisieae. However, some species of Gochnatia (e.g., G. amplexifolia (Gardner) Cabrera, G. buchii (Urb.) J. Jiménez Alm., G. glutinosa (D. Don) D. Don ex Hook. & Arn., G. magna M. C. Johnst. ex Cabrera, G. shaferi (Britton) Jervis & Alain) show a close palynological relationship with some Guayana Highland-centered genera because both have the Wunderlichia exine type (Table 4). Guayana Highland-Centered Genera. Members of this group have been generally viewed as natural by some authors (Maguire, 1956; Carlquist, 1958; Pruski, 1991),


497

Table 7 Tribe Mutisieae, Gochnatia Complex: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

1 perennial, caulescent herbs, 2 scapiform, perennial herbs, 3 shrubs, 4 trees, 5 subshrubs

with Pruski (1989a, b, 1991) circumscribing the group as comprising Achnopogon, Chimantaea, Duidaea, Eurydochus, Glossarion, Gongylolepis, Neblinaea, Quelchia, Stenopadus, Stifftia, Stomatochaeta, and Wunderlichia. The group, also known as the Stenopadus group (Bremer, 1994), is mainly centered in the Guayana Highlands of northern South America, with representatives in eastern Brazil and the Andes. Because of their high degree of endemism and biogeographic isolation, the Guayana Highland-centered genera have generated considerable interest. Recently, the genus Salcedoa from the Dominican Republic has been linked to some of the genera centered in the Guayana-Highland by Jiménez Rodríguez et al. (2004). Despite the fact that some authors regarded the Guayana-Highland centered genera as a group, some others (e.g., Maguire et al., 1957a, b; Cabrera, 1977) placed the genera of this complex in different subtribes. From a morphological point of view, two main groups are distinguished within the Guayana-Highland centered genera and relatives: (1) the zygomorphic-flowered genera with epaleate, homogamous capitula of all bilabiate corollas (rarely ligulate by abortion of the inner two corolla lips) and smooth style branches (Table 8), and (2) the actinomorphic-flowered genera with epaleate to paleate, homogamous capitula of all tubular corollas and rugulose to papillose styles beyond the branches bifurcation point. The genera with bilabiate corollas were included in the subtribe Mutisiinae and those with tubular corollas in the subtribe Gochnatiinae (Maguire et al., 1957a, b; Cabrera, 1977). The latter group is currently treated as the tribe Stifftieae (see below). Molecular studies (Kim et al., 2002; Funk et al., 2005) have shown the two sampled Guayana Highland-centered genera Duidaea and Stifftia in separate branches. Panero and Funk (2002), in their molecular analysis based on several regions from the chloroplast genome, proposed the circumscription of the Stifftia group, which includes genera of South America

498

L. Katinas, et al.

Table 8 Tribe Mutisieae, Guayana Highland-Centered Bilabiate Genera: Comparative Table Showing the Distinguishing Morpho logical Characters and the Distribution of These Genera

Crossed out corolla indicates: Absent. 1 shrubs, 2 trees

and Asia. A morphology-based phylogeny of the group (Jiménez Rodríguez et al., 2004) produced clades with partially contradictory branching patterns when Gochnatia was used to root the cladogram. In the alternative minimal length trees, the genera with bilabiate florets are either monophyletic or polyphyletic, whereas the genera with tubular florets are either monophyletic or paraphyletic. The relationships of the Guayana Highland-centered genera, according to the morphology, seem to point in two different directions: (1) they resemble genera or species of the Gochnatia complex in their discoid capitula with all tubular corollas (tribe Stifftieae), acuminate to apiculate anther appendages (e.g., Chimantaea, Wunderlichia), styles with smooth branches (e.g., Achnopogon, Duidaea), and pollen morphology (see Gochnatia complex); and (2) they resemble the tribe Cardueae in the pollen morphology (e.g., Gongylolepis, Quelchia). The group is taxonomically complex and is currently under study by John Pruski. After Pruski’s (2004) work, the Guayana Highland-centered genera were placed into two groups: those with bilabiate florets belonging to the tribe Mutisieae and those with tubular florets belonging to the tribe Stifftieae. In taxonomic and molecular works, Panero and Funk (2007, 2008) established the subfamilies Stifftioideae and Wunderlichioideae, both containing Guayana- and non Guayana-centered genera. There are not morphological characters that support these taxa. Ainsliaea Group. The Asian genera Ainsliaea, Macroclinidium, Myripnois, and Pertya were considered as a monophyletic group within the Mutisioideae by Hansen (1991a) and Bremer (1994; Table 9). The group was principally supported by its trinerved leaves, few-flowered capitula, and style branches short and dorsally pilose. Kim et al. (2002), using a molecular approach, confirmed the monophyly of the group


499

(adding two morphological synapomorphies: homogamous capitula arranged laterally on the branches) and suggested that the Asian clade could be recognized as a distinct tribe. Simultaneously, Panero and Funk’s (2002) molecular studies produced results congruent with Kim et al.’s (2002). Panero and Funk proposed the new subfamily Pertyoideae, sister to most tribes of Asteraceae, excepting Mutisieae and its segregates and the tribe Cardueae. Morphologically, the corolla of these Asian genera deviates from the typical bilabiate or tubular corolla of Mutisioideae (Fig. 1c). Although bilabiate corollas have been mentioned as occurring in the group (Mattfeld, 1934 for Myripnois; Panero & Funk, 2002), a closer examination reveals deeply five-lobed corollas with one deeper split (6 of Figs. 3–6, 31 and 33 of Figs. 31–34, 40 of Figs. 39– 42), these slightly ligulate and with unilateral lobes (Koyama, 1975; Freire, 2007). The pollen of Macroclinidium has a particular exine type, and some species of Ainsliaea have tricolpate pollen, which is unusual in Asteraceae with the Mutisia or the Ainsliaea exine type (Tellería et al., 2003; Tellería & Katinas, 2004, 2005) (82 of Figs. 80–85). The Mutisia exine type in species of Ainsliaea, Pertya, and Myripnois links the group to other members of the tribe Mutisieae. Most importantly, the style branches covered by short papillae above the branch bifurcation point (Fig. 79a) or above and somewhat below (Pertya) is the main character that supports keeping the Asian group within Mutisioideae. This generic group is unique within Mutisioideae by its particular corollas, but the complete morphology of the group is still unknown mainly because of the lack of revisionary studies in some of its genera. Hecastocleis. The genus Hecastocleis, with its only species H. shockleyi A. Gray, is endemic to western United States in the states of California and Nevada (Hickman, 1993; Table 10). It has a distinctive and peculiar morphology when compared with other genera of Mutisieae, with its single-flowered capitula aggregated into compound heads and with each aggregation of one–five heads surrounded by spiny, ovate bracts. In the most recent molecular phylogenetic studies (Panero & Funk, 2002, 2008; Funk et al., 2005), Hecastocleis appears as an independent clade separated (subfamily Hecastocleidoideae) from the rest of Mutisioideae. The unique occurrence of tricolpate pollen in Hecastocleis (and some species of Ainsliaea) would provide additional evidence to support that this genus stands apart from other genera of Mutisioideae (95 of Figs. 95–100). But the Mutisia exine type of the pollen of Hecastocleis, together with its smooth style branches (Fig. 79b), does not suggest a complete independence of this genus from the subfamily (Tellería & Katinas, 2005). Adenocaulon and Eriachaenium (Table 10). These two genera are problematic because their tribal and subfamilial placement in Asteraceae is uncertain. Both genera are in many respects so different from other genera of Asteraceae that they have been described and tentatively placed in several tribes both in the subfamily Cichorioideae and subfamily Asteroideae (Katinas, 2000). Some characters link them to Cichorioideae (e.g., deeply lobed central corollas, ecaveate pollen), whereas other characters are typical of Asteroideae (e.g., anther appendage basally constricted, short anther tails). The peculiarity of both genera led Rydberg (1917) to create the tribe Adenocauleae (for Adenocaulon), which was tentatively accepted by some authors (Robinson, 1994; Pruski, 1997; Katinas, 2000). Previous cladistic analyses based on molecular data undertaken to analyze relationships within

500

L. Katinas, et al.

Asteraceae place Adenocaulon in the tribe Mutisieae (Jansen & Kim, 1996; Kim et al., 2002). A cladistic analysis based on morphological data (Katinas, 2000) shows that Adenocaulon and Eriachaenium are sister taxa and constitute an isolated clade nested within Cichorioideae s.l. Adenocaulon and Eriachaenium are the genera of Mutisioideae that have the most number of uncommon characters for the tribe, i.e., tubular four-lobed corollas, very short anther tails, anther collar, anther appendages constricted and demarcated from the thecae, and spheroidal, small pollen grains. On the other hand, their papillose, mutisioid styles (Fig. 79c) and the Mutisia exine type of their pollen grains would suggest a link with the tribe Mutisieae of Mutisioideae. Therefore, from the informal generic groups and isolated taxa of Mutisioideae pointed out by Bremer (1994), we recognize the seven above-mentioned as sharply distinctive taxa. The other groups of Bremer, the Chaetanthera group (Brachyclados, Chaetanthera, and Pachylaena), the Gerbera group (Chaptalia, Gerbera, Leibnitzia, Perdicium, Trichocline, and Uechtritzia), the pair MutisiaHyaloseris, the pair Onoseris–Urmenetea, and the Plazia group (Aphyllocladus, Gypothamnium, and Plazia), all share bilabiate florets and papillose style branches, and therefore we place them in the central core of Mutisioideae (Fig. 2, Table 5). Tribe Nassauvieae Genera of Nassauvieae (Table 6) are an important component of the Andean flora of South America. It is a well defined tribe that constitutes a natural group defined by capitula with all the florets bilabiate and by style branches truncate at the apex, glabrous on the outer surface, and apically crowned by elongate papillae or collector hairs (Figs. 77e, f and 78c). The pollen grains have the exine distinctly bilayered, with both ectosexine and endosexine clearly columellate, a pattern that is consistent throughout the tribe (Crisci, 1974a; Cabrera, 1977; Tellería et al., 2003) (112B of Figs. 112–116). A few genera display some features that deviate from this traditional concept of Nassauvieae: Lophopappus, Macrachaenium, and Proustia have rounded style branch apices that are dorsally papillose, and some species of Acourtia and Lophopappus have capitula with tubular florets. However, the exine stratification, typical of pollen grains of Nassauvieae (Crisci, 1974a; Tellería et al., 2003), confirms the position of these genera in the tribe. Tribe Stifftieae Stifftieae (Table 11), a small tribe of six genera (Chimantaea, Quelchia, Stenopadus, Stifftia, Stomatochaeta, and Wunderlichia) and 48 species, was established by David Don in 1830 and then resurrected by Pruski (2004) and Jeffrey (2004) to include genera with strictly discoid capitula and dorsally rugulose to papillose styles. The genera of Stifftieae have a unique type of style, dorsally covered by papillae that extend almost to the base of the style (Figs. 77c and 78b). In Stifftia, the papillae are restricted to the apex, and the surface below consists of swollen cells that give a rugulose aspect to the style (Fig. 77d). Quelchia is the only genus of the tribe with a smooth style, but because it has mostly tubular florets it was included in Stifftieae by Pruski (2004). The


501

Table 9 Tribe Mutisieae, Ainsliaea Group: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

1 perennial herbs with rosulate leaves, 2 perennial herbs with leaves clustered at the median part of the stems, 3 shrubs, 4 subshrubs

genera of this tribe show relationships with the bilabiate Guayana Highland-centered genera of tribe Mutisieae and with genera of Cardueae in their pollen.

The Mutisioideae as a Primitive Subfamily One of the recent, most important discoveries in the systematics of Asteraceae is the lack of the 22 kb inversion in the chloroplast genome of three species of some South American genera (Jansen & Palmer, 1987). These genera now belong to the subfamily Barnadesioideae, previously known as subtribe Barnadesiinae of the tribe Mutisieae (Cabrera, 1977). The findings of Jansen and Palmer (1987) reinforced an already growing body of evidence for primitiveness of the tribe Mutisieae, as suggested for example by Carlquist (1961), Poljakov (1967), Skvarla et al. (1977), and Jeffrey (1977).

502

L. Katinas, et al.

Because Barnadesioideae are believed to be sister to the rest of Asteraceae, the subfamily has been a focus of attention and a model to understand the origins and evolution of the whole family. Although Barnadesioideae are considered the sister to the rest of Asteraceae, the subfamily seems most closely related to Mutisioideae, as traditionally viewed by Cabrera (1977) and as is corroborated in this contribution. Both taxa share some morphological and chemical characters in their styles, corollas, and flavonoids (Stuessy et al., 1996). According to Robinson (1987), it seems unlikely that the Barnadesioideae are so remote from Mutisieae that the remainder of the Asteraceae could evolve from within this gap. Robinson suggests that the more immediate descendents of the earliest Asteraceae with the inverted chloroplast DNA carried the two forms (with and without the chloroplast DNA inversion) mixed in their populations through many generations, and that mixture characterized much of the initial diversity of the family. It was from these mixed ancestral populations that the modern taxa of Asteraceae were derived, but the older form of DNA was retained in only one group, the Barnadesioideae. It has been suggested (Pruski, 1991; Bremer, 1994) that the basal group of the tribe Mutisieae could be among the Guayana Highland-centered genera (e.g., Stenopadus). The assemblages of largely Guayana Highland-centered genera could be modern representatives of an ancient lineage of long extinct progenitors, with some extant taxa themselves bordering on the edge of extinction. Most genera and species that occur on mid- to high-elevation, nutrient-poor, sandstone tepuis that are actively eroding away are known only by few individuals in small populations. According to Bremer (1993), among the archaic, relict genera of the basal complex of Asteraceae are Stenopadus and its immediate relatives from the Guayana Highland in Venezuela, Hesperomannia (currently placed in Vernonieae; Kim et al., 1998) from the Hawaiian Islands, Wunderlichia from Brazil, Ainsliaea from eastern Asia, and Gochnatia from throughout America and part of southeast Asia. Bremer (1994) provided a hypothesized “Stenopadus like” ancestral morphology for the Asteraceae. This hypothesis suggests that the ancestor was a woody plant with alternate leaves, large and discoid capitula, actinomorphic deeply five-lobed corollas, caudate and calcarate anthers with sterile apical appendages, shortly bilobed styles glabrous or papillose with continuous stigmatic areas, and a pappus of scabrid bristles. These are the characteristics of the members of the tribe Stifftieae. Stuessy et al. (1996) and Urtubey and Stuessy (2001) in general suggested similar characteristics for the ancestor, except that it would have been a perennial herb similar to Schlechtendalia (Barnadesioideae) and members of Calyceraceae and would have a scaly pappus. Until recently, the oldest known fossil record in the family corresponded to pollen of Mutisioideae type from the Eocene to middle Oligocene (to possibly Miocene) strata from the province of Malleco in Chile and Patagonia in Argentina, and from the late Oligocene of Louisiana, Texas, California, and Alaska (Barreda, 1993; Graham, 1996). This led some modern authors to propose South America as the likely place of origin for the family. Bremer (1992) applied the ancestral areas method, an approach of historical biogeography, to a cladogram of Asteraceae to determine a probable place of origin for the family. The results indicated that Brazil south of the Amazon Basin was most probably part of the ancestral area of Asteraceae. Bremer concluded that the ancestral area was part of a Pacific distribution pattern including South America and excluding Africa and most of Eurasia. Stuessy et al. (1996) presented a detailed


503

scenario based on a morphological cladogram of Barnadesioideae. These authors postulated that the Asteraceae and Calyceraceae, the sister family of Asteraceae, may have diverged as separate families in a southern South American (Argentina and Chile) transitional area of tropical and temperate forests in the early Oligocene. During the Miocene, the initial differentiation of Barnadesioideae began, which then gave rise to other groups of Asteraceae. According to Stuessy et al. (1996), the best candidate for this early lineage would be found in Mutisioideae, tribe Mutisieae, due to its similarity with Barnadesioideae. The results of Gustafsson et al. (2001), on the basis of a molecular cladogram of Barnadesioideae, support Stuessy et al.’s (1996) idea of a southern South American origin for the subfamily. Bremer and Gustafsson (1997) proposed a Cretaceous origin for the sunflower alliance—the monophyletic group made up by Asteraceae, Campanulaceae, and ten smaller families—with the ancestor occurring in East Gondwana, i.e., western Antarctica, Africa, and South Table 10 Tribe Mutisieae, the Generic Pair Adenocaulon and Eriachaenium and the Independent Genus Hecastocleis: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

1 perennial herbs with rosulate leaves, 2 perennial herbs, 3 shrubs

504

L. Katinas, et al.

America before the breakup of this supercontinent. The Australasian-South American connection displayed by the Goodeniaceae and the Calyceraceae and Asteraceae is, according to Bremer and Gustafsson (1997), a reflection of the early distribution of the alliance in both East and West Gondwana. In their cladogram, inferred by parsimony analysis of 35 rbcL sequences from all of the families of the alliance, the age of the Asteraceae is estimated to be at least 38 Myr. The occurrence of two asteraceous pollen types with wall structure types like those in the Mutisieae has been reported from the Upper Paleocene–Eocene of South Africa (Zavada & de Villiers, 2000; Scott et al., 2006). Interestingly, this pollen was recovered from offshore and onshore localities of southern Africa. According to Zavada and de Villiers (2000), an African origin of Asteraceae is more likely, with subsequent dispersal to southern South America, where the earliest occurrence of Asteraceae pollen is in the Oligocene (Barreda, 1993). However, these Eocene pollen records need taxonomic and/or stratigraphic confirmation (Scott et al., 2006). Those reported as Mutisiapollis spp. from the Late Eocene of the Gulf coast of Mexico (Elsik & Yancey, 2000) might be the oldest ones, but neither description nor illustration was provided to date (T. E. Yancey, personal communication) to confirm their identification. A second, smaller chloroplast DNA inversion occurring at one end of the larger inversion was found in Asteraceae, excepting Barnadesioideae (Kim et al., 2005). Two clock calibrations from Poaceae and Oleaceae were employed to calibrate the Asteraceae molecular clock, resulting in a mid-Eocene (42–48 Myr) origin of the family. The two chloroplast genome inversions occurred, according to this study, in the Late Eocene/Early Oligocene (38–42 Myr) when the Barnadesioideae diverged from the rest of the Asteraceae. Funk et al. (2005) proposed that the origin of the extant members of Asteraceae was in southern South America and a subsequent radiation in Africa gave rise to most of the tribes we know today. New fossil pollen discoveries of Mutisieae s. l. and Barnadesioideae were reported from marine deposits of Patagonia, in southern Argentina (Katinas et al., 2007; Barreda et al., 2008). A minimum age of 28–23 Myr (Late Oligocene, Chattian) for fossil pollen related to the extant genera of the Gochnatia complex, and 23–20 Myr (Early Miocene, Aquitanian) for fossil pollen of the Nassauvieae and Barnadesioideae were reported (Katinas et al., 2007). Fossil pollen records of the Late Miocene (11–9 Myr) in Patagonia show high abundance and diversity of Nassauvieae associated with pollen of Astereae and Barnadesioideae. It is interesting to note that the chronological order of appearance of these groups in Patagonia, first Mutisieae s. l. and second Barnadesioideae, differs from the branching order of the current accepted phylogenies, i.e., first Barnadesioideae and second Mutisieae s. l. Many molecular biologists have used and are using fossils in their analyses as a way of providing a calibration point for evolutionary models used to approximate ages for the nodes of the phylogenetic trees. Opinion remains divided as to how best to use temporal (stratigraphic) data in phylogenetic reconstruction (see the online debate from 1998 at http://www.nature.com/nature/debates/fossil/). Some authors think that the order of stratigraphic appearance of taxa in the fossil record should play a role in phylogenetic tree-building. In such cases stratigraphic order is used from the outset to influence and direct branching relationships by maximizing the congruence between the age of a taxa’s first occurrence in the fossil record and how early it branches from a phylogenetic tree. Others see stratigraphy as irrelevant to


505

Table 11 Tribe Stifftieae: Comparative Table Showing the Distinguishing Morphological Characters and the Distribution of These Genera

1 shrubs, 2 subshrubs, 3 trees, 4 vines

phylogeny, and dismiss mismatches between the fossil record and phylogenetic hierarchy as a result of an inadequate fossil record. The role of stratigraphy is then restricted to post hoc dating of branching events. If we see the last pollen findings (e. g., Katinas et al., 2007) through the former position (despite the fact that the age of fossils is considered minimum age), they contradict the branching pattern of the current hypotheses on the evolution of Asteraceae. If we see these findings through the latter position, hopefully, they will help to calibrate more precisely the phylogenetic trees involving Asteraceae.

Distributional Patterns Genera of Mutisioideae are found on all continents except Europe and Antarctica, with its major concentration in South America. The tribe Stifftieae, with its low number of genera, is very restricted in distribution compared to the tribes Nassauvieae and Mutisieae. The Nassauvieae and Mutisieae, themselves, have very different distributional patterns. The tribe Nassauvieae is exclusively Neotropical, with mostly monotypic, endemic, Andean, Chilean, and Patagonian genera, only a few being widespread. Largely Andean–Patagonian-centered genera are Dolichlasium (one species), Leucheria (46 species), Lophopappus (6 species), Nassauvia (38 species), and Perezia (31 species). Exclusively Patagonian genera are Ameghinoa (one species), Burkartia (one species), and Macrachaenium (one species), the last from the Nothofagus Blume forest. Central and northern Chile, a hot spot in South America,

506

L. Katinas, et al.

hosts many endemic genera such as Calopappus (one species), Leunisia (one species), Marticorenia (one species), Moscharia (two species), Oxyphyllum (one species), Pleocarphus (one species), Polyachyrus (seven species), and six of the seven species of Triptilion. Only Jungia (27 species) and Trixis (39 species) are widespread genera ranging from North and Central America to southern Argentina. On the other hand, Acourtia and Berylsimpsonia are not South American, occurring in North and Central America and in the West Indies, respectively. The genera of Mutisieae are mainly South American but they are also represented in Asia, Africa, and Australia by only one species (Trichocline spathulata (A. Cunn. ex DC.) J. H. Willis). The genera are in general not centered in one particular area, but there are some generic groups concentrated in several different restricted areas. For instance, the Guayana Highland-centered genera belonging to tribes Mutisieae and Stifftieae inhabit the Guayana Highlands of northern Brazil, Colombia, Guyana, and Venezuela, but also extend to the Andes of Colombia, Ecuador, and Peru, Dominican Republic, and eastern Brazil. Unlike the Nassauvieae, there are few monotypic genera or genera with few species endemic to restricted areas: Gypothamnium (one species) from northern Chile, Hecastocleis (one species) from western United States, Perdicium (two species) from southern Africa, and Lulia (one species) from southern Brazil. There are many widespread genera, for instance Chaptalia in America, from southern United States to central Argentina; Gerbera in Africa, southern Asia, and America; Gochnatia, with two Asian species, from Mexico to Argentina; and Mutisia from the Andes of Colombia to southern Argentina and Chile, and in southern Brazil and adjacent regions. Some genera well represented in the Andean mountain ranges are Aphyllocladus, Chaetanthera, Chucoa, Onoseris, Pachylaena, and Plazia. Finally, the Asian Ainsliaea group ranges from Afghanistan to Japan and southeastern Asia. The genera of Stifftieae have their major diversity in the Guayana Highlands. Chimantaea is endemic to the eastern Venezuelan Guayana Highland. All species of Quelchia are endemic to the Guayana Highland in southern Venezuela. Stenopadus is a genus of 15 species, 14 of which are found in the sandstone areas of the Guayana Highland in Colombia, Venezuela, Guyana, Brazil, with one species in the Andes of Ecuador and Peru. All species of Stomatochaeta are endemic to the Guayana Highland of southern Venezuela, Guyana, and northern Brazil. Stifftia occurs in Brazil and French Guiana and is the only genus of the tribe occurring both north and south of the Amazon River. Wunderlichia does not inhabit the Guayana Highlands; it occurs in eastern Brazil. In general, the genera of Mutisioideae have their highest concentration in the Andes and Patagonia (25 genera), with other important centers in the Guayana shield (11 genera), northern and central Chile (seven genera), and southeastern Asia (four genera).

Taxonomy Subfamily Mutisioideae (Cass.) Lindl. Lindley in Loudon, Encycl. pl.: 1074. 1829, based on the tribe Mutisieae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 199. 1819.


507

Herbs, shrubs, subshrubs, vines, or trees; monoecious, dioecious, or gynodioecious. Leaves alternate, rarely opposite, spreading along the stem or clustered in a basal rosette, in the median part of the stem, or at the stem apex; sessile, clasping, decurrent to petiolate, occasionally with stipules; blades simple, rarely compound, entire to divided, unarmed to spiny, predominantly pinnately veined, less commonly three-veined, palmately veined, or parallel-veined, glabrous to pubescent. Capitulescences monocephalous or few at the end of the branches, or laxly to densely corymbose, racemose to paniculate, rarely glomerulose or pseudocephalia. Capitula homogamous or heterogamous; discoid, radiate, ligulate, or disciform; receptacle epaleate, less frequently paleate, glabrous to pubescent, alveolate, ciliate, setose to fimbriate; involucre uniseriate to multiseriate. Florets one to more than 100; corollas isomorphic, dimorphic, sub-dimorphic, or rarely trimorphic, white, yellow, cream, pink, purple, blue, orange, or red; marginal florets functionally female, neuter, or bisexual; corollas bilabiate, sub-bilabiate, true ray, ligulate, or tubular and subzigomorphic, rarely tubular four-lobed; in trimorphic capitula, internal marginal florets female, with or without staminodes, corollas filiform to bilabiate; central florets bisexual, functionally female, or male, corollas tubular-funnelform to rarely sub-campanulate, deeply to very deeply-lobed, bilabiate, ligulate, rarely subbilabiate; anthers calcarate, caudate, with long, rarely short tails and thus anthers auriculate, smooth to papillose; connectival or anther appendage ovate to lanceolate, acute, truncate to apiculate, not constricted or demarcated from the thecae, rarely demarcated; style shortly bilobed to bifid and cleft into two long branches, rounded to truncate at the apex, smooth throughout, or dorsally papillose only above the style branch bifurcation to papillose both above and below the bifurcation, papillae rounded, never acute, or with an apical crown of short collector hairs, inner surface of the branches completely covered by stigmatic papillae. Cypselae oblong, ellipsoid, ovoid, obovoid to turbinate, truncate, attenuate to rostrate at the apex, smooth to ribbed; glabrous to pubescent, with typical or modified twin hairs, glandular hairs and leaf-like hairs (e.g., two-armed, flagellate); pappus of one to more series of bristles, equal to unequal in length, scabrid to plumose, capillary to somewhat rigid and paleaceous, reduced to a crown of fringed scales, or absent. Pollen tricolporate or rarely tricolpate, suboblate to prolate, usually of medium to large size (25–100 μm); exine almost psilate, microechinate to echinate, exine structure with Anthemoid pattern. x=9. Characters that define the subfamily are in italics. TYPE: Mutisieae Cass. The Mutisioideae occur mainly in the New World, with the major concentration of genera in the mountains and arid regions of South America, and some genera distributed in Central America, Mexico, and southern United States, southern and eastern Asia, Africa (including Madagascar), Arabia, and Australia. It comprises 74 genera and ca. 865 species. Key to Tribes of Mutisioideae 1a. Style branches smooth or with papillae sparsely distributed, never clustered at the branch apices. Pollen microechinate or echinate, the ectosexine with columellae very thickly disposed (compact aspect), and the endosexine clearly columellate.

508

L. Katinas, et al.

2a. Capitula isomorphic, dimorphic, or trimorphic. Style branches dorsally smooth or with papillae completely covering the branches or only the branch apex, very rarely extending below the branch bifurcation point ................................................................................................ Tribe Mutisieae 2b. Capitula isomorphic with all tubular corollas (bilabiate in Quelchia bracteata). Style branches rugulose with swollen cells or papillose, exceptionally smooth (Quelchia); the swollen cells or the papillae covering the branches and also extending much below the branch bifurcation point (ca. 3/4 of the style) .............................................................. Tribe Stifftieae 1b. Style branches apically crowned by a tuft of papillae (collector hairs). Pollen microechinate, both ectosexine and endosexine clearly columellate ............ ................................................................................................ Tribe Nassauvieae

Tribe Mutisieae Cass. H. Cassini, J. Phys. Chim. Hist. Nat. Arts 88: 199. 1819. Mutisiinae Dumort., Anal. fam. pl.: 31. 1829; Less., Linnaea 5: 241. 1830. Mutisiaceae Burnett, Outlines Bot.: 934. 1835. TYPE: Mutisia L. f. Onoserideae Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 4. 1818, as “Onoseridae.” Onoseridinae Benth., in Benth. & Hook. f., Gen. pl. 2: 215. 1873, as “Onoserideae.” Onoserideae (Benth.) Panero & V. A. Funk, Phytologia 89: 359. 2007. TYPE: Onoseris Willd. Chaetantherae Dumort., Anal. fam. pl.: 31. 1829. TYPE: Chaetanthera Ruiz & Pav. Gerbereae Lindl. in Loudon, Encycl. pl.: 1074. 1829, as “Gerberieae.” Gerberinae Benth., in Benth. & Hook. f., Gen. pl. 2: 217. 1873, as “Gerbereae.” TYPE: Gerbera L. Perdiciaceae Link, Handbuch 1: 728. 1829. Perdicieae D. Don, Trans. Linn. Soc. London 16: 239. 1830, as “Perdiceae.” TYPE: Perdicium L. Adenocauleae Rydb., Fl. Rocky Mts.: 833. 1917. Adenocaulinae A. Gray, Syn. fl. N. Amer. 1: 59. 1884. TYPE: Adenocaulon Hook. Gochnatiinae Benth., in Benth. & Hook. f., Gen. pl. 2: 216. 1873, as “Gochnatieae.” Gochnatioideae (Benth. & Hook. f.) Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 913. 2002. Gochnatieae (Benth. & Hook. f.) Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 913. 2002. TYPE: Gochnatia Kunth. Hecastocleidoideae Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 914. 2002, as “Hecastocleioideae.” Hecastocleideae Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 914. 2002. TYPE: Hecastocleis A. Gray. Pertyoideae Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 915. 2002. Pertyeae Panero & V. A. Funk, Proc. Biol. Soc. Wash. 115: 915. 2002. TYPE: Pertya Sch. Bip. Hyalideae Panero, Phytologia 89: 358. 2007. TYPE: Hyalis D. Don ex Hook. & Arn. Subshrubs, shrubs, perennial herbs, vines or small trees, monoecious, dioecious, or gynodioecious. Leaves alternate to rarely opposite, rosulate, clustered at the


509

branch apices to clustered in the axils of the old leaves; petiolate, sessile to sheathing at the base; blades entire, lyrate, pinnatisect to pinnately compound, unarmed to spiny, pinnately, palmately to three-veined, rarely parallel-veined, glabrous to pubescent. Capitulescences monocephalous, two- to three-headed at the end of the branches, laxly to densely corymbose, racemose, paniculate to glomerulose or pseudocephalia. Capitula homogamous or heterogamous, discoid, disciform, ligulate, or radiate, isomorphic or heteromorphic with dimorphic, sub-dimorphic, or rarely with trimorphic florets, with one to numerous florets; receptacle epaleate to paleate, glabrous, pilose, setose to fimbriate; involucre uniseriate to multiseriate. Isomorphic capitula with all the florets bisexual or rarely functionally female, corolla ligulate, shallowly to deeply five-dentate, sub-bilabiate to tubular-funnelform, and deeply five-lobed. Heteromorphic capitula with marginal florets bisexual, female, or neuter, corolla bilabiate, sub-bilabiate (3+1 corolla lips), true ray to tubular four- to five-lobed; intermediate florets in the trimorphic capitula female, corolla filiform to bilabiate, with or without staminodes; central florets bisexual, functionally female or male, corolla bilabiate, sub-bilabiate, tubular-bilabiate, tubular-funnelform to subcampanulate, and five-lobed; anthers with apical appendage lanceolate, rounded, acute, truncate to apiculate, rarely basally constricted and demarcated from the thecae, tails long, rarely short, smooth to papillose, adjacent tails free to connate; style cleft into two short or bilobed to long or bifid branches, branches dorsally glabrous to papillose, with the papillae covering the branches above the branch bifurcation point, occasionally extending somewhat below, papillae short and rounded at the apex. Cypselae oblong to sub-obovate, truncate, attenuate to rostrate at the apex, glabrous to pubescent, sometimes dimorphic with the marginal cypselae conspicuously larger than the central ones; pappus of scabrid to plumose bristles, capillary to narrowly paleaceous, often plumose only at the apex, equal to different in length, less commonly reduced to a crown of fringed scales, or pappus absent. Pollen tricolporate, rarely tricolpate, microechinate with small puncta to echinate, ectosexine very compact, endosexine with stout and ramified columellae. Characters that define the tribe are in italics. Mutisieae include 43 genera and ca. 500 species principally South American with a concentration in the Andes and some endemics of the Guayana Highland. Five genera occur in North America, eleven genera in Asia, and three in Africa; one species of Trichocline occurs in Australia. Genera: Achnopogon Maguire, Steyerm. & Wurdack, Actinoseris (Endl.) Cabrera, Adenocaulon Hook., Ainsliaea DC., Aphyllocladus Wedd., Brachyclados D. Don, Catamixis Thomson, Chaetanthera Ruiz & Pav., Chaptalia Vent., Chucoa Cabrera, Cnicothamnus Griseb., Cyclolepis D. Don, Dinoseris Griseb., Duidaea S. F. Blake, Eriachaenium Sch. Bip., Eurydochus Maguire & Wurdack, Gerbera L., Glossarion Maguire & Wurdack, Gochnatia Kunth, Gongylolepis R. H. Schomb., Gypothamnium Phil., Hecastocleis A. Gray, Hyalis D. Don ex Hook. & Arn., Hyaloseris Griseb., Ianthopappus Roque & D. J. N. Hind, Leibnitzia Cass., Lulia Zardini, Lycoseris Cass., Macroclinidium Maxim., Mutisia L. f., Myripnois Bunge, Neblinaea Maguire & Wurdack, Nouelia Franch., Oldenburgia Less., Onoseris Willd., Pachylaena D. Don ex Hook. & Arn., Perdicium L., Pertya Sch. Bip., Plazia Ruiz & Pav., Salcedoa F. Jiménez Rodr. & Katinas, Trichocline Cass., Uechtritzia Freyn, Urmenetea Phil.

510

L. Katinas, et al.

Key to Genera 1a Style branches dorsally smooth. 2a Capitula with all bilabiate or rarely ligulate corollas. 3a Leaves one-veined; capitula pendulous; phyllaries reddish ............ Duidaea 3b Leaf venation pinnate or less commonly sub-palmate; capitula not obviously pendulous, commonly erect; phyllaries not brightly colored. 4a Leaves tomentose abaxially; corollas bilabiate or ligulate. 5a Capitulescence monocephalous; corollas ligulate and/or bilabiate; cypselae glabrous to scarcely glandular ......................... Glossarion 5b Capitulescence corymbose; corollas bilabiate; cypselae villose ................................................................................................ Salcedoa 4b Leaves frequently glabrous, less commonly slightly pubescent beneath; corollas bilabiate. 6a Capitulescence terminal to subterminal; capitula commonly large, > (6)10 mm wide, six- to many-flowered; involucre hemispherical. 7a Phyllaries subequal, decurrent onto peduncle; cypselae puberulent; pappus bristles fragile, sometimes individually deciduous .................................................................................... Eurydochus 7b Phyllaries graduated, peduncles sometimes bracteate, but the bracts leaf-like and not resembling the phyllaries; cypselae glabrous; pappus bristles persistent ……...…………………… Gongylolepis 6b Capitulescence lateral, three- to nine-headed; capitula commonly small, < 6 mm wide, two- to six-flowered; involucre cylindrical. 8a Leaves densely spirally arranged, except sometimes on capitulescence; capitula subsessile; phyllaries woolly; corolla tube puberulent ………………..………………..………………..……. Achnopogon 8b Leaves alternate, mostly at stem apex; capitula pedunculate; phyllaries glabrous; corollas glabrous ………………. Neblinaea 2b Capitula with all or some of the corollas tubular. 9a Capitula with only tubular corollas. 10a Capitulescences with spiny bracts enclosing several, cylindrical, oneflowered capitula ……………………………………...… Hecastocleis 10b Capitulescences without spiny bracts enclosing the many-flowered capitula. 11a Branches unarmed; plants monoecious, gynodioecious, or polygamous; when gynodioecious corolla tubes in female florets short (1/3 of corolla length) ……………………...… Gochnatia 11b Branches modified in thorns; plants gynodioecious, corolla tubes of female florets long and narrow (up to 2/3 of corolla length) ………………………………………..……………… Cyclolepis 9b Capitula with marginal florets bilabiate to sub-bilabiate and central florets tubular. 12a. Leaves three-veined. 13a. Central florets numerous; pappus purplish …………………Ianthopappus 13b. Central floret 1; pappus pale ……………………………………….. Hyalis 12b. Leaves pinnately veined.


511

14a. Perennial herbs …………...…………………………….. Actinoseris 14b. Shrubs or small trees. 15a. Marginal florets with corollas sub-bilabiate (3+1 corolla lips) ... ………………………………………………… Cnicothamnus 15b. Marginal florets with corollas bilabiate (3+2 corolla lips) ……………………......……………………………… Nouelia 1b. Style branches papillose. 16a. Anthers auriculate at the base, with tails very short and inconspicuous. 17a. Leaves alternate, distributed more or less evenly along stem; capitulescence monocephalous ….…………………… Eriachaenium 17b. Leaves rosulate to sub-rosulate; capitulescences of few capitula …………………………………………………...…….. Adenocaulon 16b. Anthers caudate at the base, with tails long and conspicuous. 18a. Capitula usually with all florets homomorphic. 19a. All florets usually with tubular corollas. 20a. Leaves reduced, linear-oblong to linear-lanceolate, caducous ……………………………...………………… Aphyllocladus 20b. Leaves well developed, lanceolate to obovate, persistent …………………………………………………….. Chucoa 19b. All florets with ligulate, bilabiate, or sub-bilabiate corollas. 21a. Corollas ligulate. 22a. Corollas ligulate, deeply five-lobed limb, with one deeper split; style bilobed. 23a. Pappus bristles plumose ……………………. Ainsliaea 23b. Pappus bristles scabrid. 24a. Dioecious shrubs; phyllaries ca. 5 …… Myripnois 24b. Monoecious perennial herbs, vines, subshrubs, or shrubs; phyllaries six to more. 25a. Herbs; leaves clustered at the median part of the stem ...………………… Macroclinidium 25b. Shrubs; leaves alternate, more or less evenly distributed along the stem or clustered on brachyblasts (short shoots) ..……….. Pertya 22b. Corollas ligulate with a five-dentate limb (occasionally some corollas are sub-bilabiate); style bifid. 26a. Stems with brachyblasts (short shoots) and macroblasts (long shoots) …...…………………………. Hyaloseris 26b. Stems without brachyblasts. 27a. Leaves alternate ……………………… Catamixis 27b. Leaves opposite ..……….……………. Dinoseris 21b. All corollas bilabiate to sub-bilabiate (except Mutisia linifolia with ligulate corollas), rarely marginal true ray corollas. 28a. Herbs (except Chaetanthera glandulosa); pappus bristles simple, scabrid to barbellate ………………. Chaetanthera 28b. Stout to trailing shrubs or subshrubs; pappus bristles plumose ...………………………………………… Mutisia 18b. Capitula with heteromorphic florets.

512

L. Katinas, et al.

29a. Central corollas tubular or tubular-bilabiate. 30a. Plants dioecious ………………………………………. Lycoseris 30b. Plants monoecious. 31a. Anther apical appendages acute to long-apiculate. 32a. Anther apical appendages long-apiculate; involucre globose-urceolate to cyathiform; central corollas deeply five-lobed ..………………………….. Oldenburgia 32b. Anther apical appendages acute to subapiculate; involucre cylindrical to campanulate; central corollas shallowly five-lobed. 33a. Leaves rugose ..……………………… Urmenetea 33b. Leaves smooth ………………………… Onoseris 31b. Anther apical appendages truncate. 34a. Leaves reduced, caducous …………….. Aphyllocladus 34b. Leaves well developed, persistent. 35a. Leaves obovate-oblong; cypselae glabrous ………………………………………...… Plazia 35b. Leaves linear; cypselae villose …. Gypothamnium 29b. Central corollas bilabiate to tubular-bilabiate. 36a. Herbs usually scapose. 37a. Pappus bristles connate at the base ……………… Perdicium 37b. Pappus bristles free. 38a. Plants dimorphic: a vernal stage with reduced or no leaves and chasmogamous capitula, and an autumnal stage with fully developed leaves and cleistogamous capitula ……………………………………. Leibnitzia 38b. Plants not dimorphic. 39a. Cypselae truncate at the apex. 40a. Cypselae villose with long, shaggy hairs ………………………………… Uechtritzia 40b. Cypselae shortly pubescent ....… Trichocline 39b. Cypselae distinctly rostrate to attenuate at the apex. 41a. Capitula with dimorphic or trimorphic florets; when trimorphic, the intermediate florets with corollas bilabiate to filiformbilabiate, as long as the styles or longer; staminodes generally present …… Gerbera 41b. Capitula with trimorphic florets; the intermediate florets with corollas very reduced, filiform (irregularly tubular, true ray to bilabiate), shorter than the styles; staminodes absent ………………………. Chaptalia 36b. Shrubs, vines, subshrubs or not scapose herbs. 42a. Pappus bristles scabrid to barbellate. 43a. Shrubs .………………………………… Brachyclados 43b. Caulescent herbs (except Chaetanthera glandulosa).


513

44a. Leaves linear, graminiform, parallel-veined ........... ....................................................................... Lulia 44b. Leaves not graminiform, reticulate-veined ……… ………...………………………….. Chaetanthera 42b. Pappus bristles plumose. 45a. Herbs; leaves orbicular to spathulate …… Pachylaena 45b. Stout or trailing shrubs to subshrubs; leaves neither spathulate nor orbicular ……………………… Mutisia Achnopogon Maguire, Steyerm. & Wurdack Mem. New York Bot. Gard. 9: 437. 1957. TYPE: Achnopogon virgatus Maguire, Steyerm. & Wurdack. 3 of Figs. 3–6. Etymology. From the Greek achno, hair, and pogon, beard, in reference to the lanate pubescence on young leaves. Shrubs with stems either erect and simple or divaricately branched. Leaves apically clustered and densely spirally inserted, alternate; sessile to subsessile, petioles, when present, clasping; blades lanceolate to obovate, coriaceous, pinnately veined, margin entire, lanate when young, glabrous with age. Capitulescences monocephalous or lateral and cymose of three to seven heads; capitula short-pedunculate, homogamous, bilabiate, two- to six-flowered; receptacle epaleate; involucre multiseriate, cylindrical, phyllaries woolly. Florets isomorphic, bisexual, corolla bilabiate, outer lip expanded; anthers reddish to cream-colored, exserted, apical appendages acute, tails papillose; style bifid, red, branches dorsally smooth. Cypselae glabrous to sparsely pilose at the apex; pappus of scabrid, capillary bristles. Pollen Description. Pollen not seen. According to Carlquist (1957b), the pollen of Achnopogon is similar to that of Gongylolepis in its sculpture and structure. Habitat and Distribution. The genus contains two species, and is endemic to the eastern Venezuelan Guayana Highland. Both species occur only at about 2,000 m elevation, one on Auyan-Tepui and the second on the more southerly Chimanta Massif. Species list 1. Achnopogon steyermarkii Aristeg., Acta Bot. Venez. 2: 350. 1967. TYPE: Venezuela. State Bolívar: Auyan-tepui, J. A. Steyermark 93512 (holotype, VEN not seen). Additional specimen examined: Venezuela. State Bolívar: Auyan-tepui, J. A. Steyermark 93497 (US). 2. Achnopogon virgatus Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 438. 1957. TYPE: Venezuela. State Bolívar: Torono-tepui, Chimantá Massif, J. A. Steyermark & J. J. Wurdack 681 (isotype, NY!). Additional specimen examined: Venezuela. State Bolívar, Dist. Piar, macizo del Chimantá, O. Hubber 11508 (US).

514

L. Katinas, et al.

Literature. Maguire et al. (1957a), Aristeguieta (1964), Pruski (1997).

Actinoseris (Endl.) Cabrera Bol. Soc. Argent. Bot. 13: 46. 1970, based on Actinoseris Endl., Gen. pl.: 483. 1838, as group “a” of Seris Less. TYPE: Seris polymorpha Less. [= Actinoseris polymorpha (Less.) Cabrera]. 4 of Figs. 3–6. Ingenhusia Vell., Fl. flumin.: 351. 1829 (1825). TYPE: Ingenhusia radiata Vell. [= Actinoseris radiata (Vell.) Cabrera]. Etymology. From the Greek aktinos, star like, radiating from a centre, and seris, seridos, chicory, lettuce. Herbs perennial, scapose or caulescent, with simple to few branched stems. Leaves rosulate or alternate; sessile to petiolate; blades elliptic, obovate to linear, pinnately veined, margin entire to denticulate, glabrous, scarcely pubescent to lanate. Capitulescences monocephalous or corymbose; capitula pedunculate, homogamous or heterogamous, radiate; receptacle epaleate; involucre three- to six-seriate. Florets dimorphic; marginal florets bisexual or functionally female with staminodes, corolla bilabiate to sub-bilabiate (3+1, 4+1 corolla lips), outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform to slightly campanulate, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bifid to bilobed, branches dorsally smooth. Cypselae villose; pappus of scabrid, capillary bristles. Pollen Description. Actinoseris angustifolia (Gardner) Cabrera (80 of Figs. 80–85). Pollen subprolate, elliptic or spheroidal in equatorial view, circular in polar view, medium to large size, P × E = (44–52×34–43) μm. Tricolporate, colpi long with margin and psilate or scarcely microgranulate membrane, mesoaperture diffuse? Exine Mutisia type with compact ectosexine, echinate, 8–10 μm thick, spines 2.5 μm length. Ratio ectosexine/endosexine: 1:1; 1.5:1. Nexine ca. 2 μm thick. Note: pollen close to that of type II of Gochnatia (see also Roque & Silvestre-Capelato, 2001). Habitat and Distribution. Genus of seven species of rocky and grassy places of Brazil. Species list 1. Actinoseris angustifolia (Gardner) Cabrera, Bol. Soc. Argent. Bot. 13: 50. 1970. Basionym: Seris angustifolia Gardner. TYPE: Brazil. State Minas Geraes: Diamont District, G. Gardner 4955 (lectotype, designated by Roque & Pirani, 2001: 1158, K not seen). Additional specimens examined: Brazil. State Minas Geraes: Jaboticatuba, G. Hatschbach 28756 (LP); Jaboticatuba, Serra do Cipó, G. Hatschbach 29986 (LP*). 2. Actinoseris arenaria (Baker) Roque, Kew Bull. 52: 199. 1997. Basionym: Trichocline arenaria Baker. TYPE: Brazil. State Minas Geraes: Itambe, G. Gardner 4956 (lectotype, designated by Roque, 1997: 199, K not seen; isotype, NY not seen, NY photo at LP!).


515

3. Actinoseris hatschbachii Zardini, Bol. Mus. Bot. Munic. 23: 1. 1975. TYPE: Brazil. State Minas Geraes: Santa Ana do Riacho, Serra do Cipó, G. Hatschbach 35366 (holotype, LP!). 4. Actinoseris polymorpha (Less.) Cabrera, Bol. Soc. Argent. Bot. 13: 48. 1970. Basionym: Seris polymorpha Less. TYPE: Brazil: s.d., Sello s.n. (lectotype, designated by Cabrera, 1970: 48, P not seen). Additional specimen examined: Brazil. State Minas Geraes: Serra do Cipó, A. C. Brade 2007 (LP). 5. Actinoseris polyphylla (Baker) Cabrera, Bol. Soc. Argent. Bot. 13: 50. 1970. Basionym: Seris polyphylla Baker. TYPE: Brazil. State Minas Geraes: Serra da Lapa, Riedel 1077 (lectotype, designated by Roque & Pirani, 2001: 1159, P not seen). Additional specimen examined: Brazil. State Minas Geraes: Santa Ana do Riacho, G. Hatschbach 35304 (LP). 6. Actinoseris radiata (Vell.) Cabrera, Bol. Soc. Argent. Bot. 13: 50. 1970. Basionym: Ingenhusia radiata Vell. TYPE: Lectotype, designated by Cabrera, 1970: 50, the illustration in Fl. flumin. Icon. 8: tab. 93. 1831 (1827). Additional specimen examined: Brazil. State Paraná: Campo Largo, G. Hatschbach 690 (LP). 7. Actinoseris stenophylla Cabrera, Bol. Mus. Bot. Munic. 15: 3. 1974. TYPE: Brazil. State Minas Geraes: Jaboticatubas, Serra do Cipó, G. Hatschbach 30061 (holotype, LP!).

Observations. Cabrera (1970) considered that the capitula of Actinoseris are homogamous with all the florets bisexual. However, in some species (e.g., A. arenaria, A. hatschbachii, A. radiata) the capitula are heterogamous with the outer florets functionally female, with staminodes, and with the central florets bisexual (Freire et al., 2002). The genus Richterago, which was regarded a synonym of Gochnatia by Cabrera (1970), was recently reinstated and recircumscribed (Roque & Pirani, 2001) to include the species of Actinoseris and the species of Gochnatia sect. Discoseris. Further, new species of Richterago were described: R. campestris Roque & J. N. Nakaj., R. petiolata Roque & J. N. Nakaj. (Roque & Nakajima, 2001), R. caulescens Roque, R. conduplicata Roque, R. elegans Roque, R. lanata Roque, R. riparia Roque (Roque, 2001a). Recent molecular studies recovered Richterago amplexifolia (= Gochnatia amplexifolia) and Richterago angustifolia (= Actinoseris angustifolia) as sister taxa (Panero & Funk, 2008). A more detailed phylogenetic analysis including more species of Actinoseris as well as more species of Gochnatia may provide the final answers about the identity of Richterago. Literature. Cabrera (1970, 1974), Roque (1997, 2001a), Roque and Nakajima (2001), Roque and Pirani (2001), Freire et al. (2002).

Adenocaulon Hook. Bot. Misc. 1: 19. 1829. TYPE: Adenocaulon bicolor Hook. 5 of Figs. 3–6. Etymology. From the Greek aden, gland, and kaulos, stalk, stem, by the stalked glandular hairs.

516

L. Katinas, et al.

Figs. 3–6 3 Achnopogon steyermarkii Aristeg. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Aristeguieta, 1967: 350). Scale bars: A=5 cm; B, C, E= 5 mm; D=0.8 mm. 4 Actinoseris angustifolia (Gardner) Cabrera. A Habit. B Marginal corolla. C Central corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Roque & Pirani, 1997: 167). Scale bars: A=3 cm; B, C, F=2 mm; D, E=1 mm. 5 Adenocaulon chilense Less. A Habit. B, C Marginal corollas. B Corolla sub-bilabiate. C Corolla tubular four-lobed. D Marginal cypsela. E Central cypsela. A. bicolor Hook. F Central corolla. G Stamen. H Upper part of the style. (Redrawn from Katinas, 2000: 230). Scale bars: A=5 cm; B, C, E, F=1 mm; D=2 mm; G, H=0.3 mm. 6 Ainsliaea latifolia (D. Don) Sch. Bip. A Habit. B Floret. C Stamen. D Upper part of the style. A. acerifolia Sch. Bip. E Habit. F Stamen. (A–D, from Drummond 25596, UC; E, F from Maximowicz s.n., MO). Scale bars: A, E=4 cm; B=5 mm; C, F=2.5 mm; D=1.2 mm


517

Herbs perennial, scapiform with stout rhizomes, stems simple, erect, with stipitate-glandular hairs. Leaves glabrous to subglabrous above, tomentose beneath; basal leaves rosulate to sub-rosulate; petiolate to pseudopetiolate; blades elliptic, ovate, obovate, to deltoid, margin entire to lyrate, pinnately to palmately veined; upper leaves similar to the basal ones but few and reduced. Capitulescences terminal, laxly racemose to corymbose, on long peduncles; capitula pedunculate, heterogamous, disciform; receptacle epaleate; involucre uniseriate. Florets dimorphic; marginal florets female, without staminodes, corolla sub-bilabiate (3+1 corolla lips), tubular-funnelform four- to five-lobed, rarely bilabiate; central florets male with a rudimentary ovary, corolla tubular-funnelform, deeply five-lobed; anther apical appendages rounded to acute at the apex, basally constricted and demarcated from the thecae, basally auriculate with tails very short, smooth; style bilobed, branches dorsally papillose. Cypselae truncate at the apex, pubescent (glandular multiseriate capitate hairs), dimorphic, marginal cypselae conspicuously bigger than the central ones; pappus absent. Pollen Description. Adenocaulon (81 of Figs. 80–85). Pollen spheroidal, circular in polar view, medium size, P × E = (26–32×26–30) μm. Tricolporate, colpi long with thin margin and microgranulate membrane, mesoaperture diffuse. Exine Mutisia type, microechinate, 4–5 μm thick, slightly slender at the poles. Ratio ectosexine/ endosexine: 1:1.5; 1:2. Nexine 1.5 μm thick. SEM: tectum punctate. Note: pollen of A. bicolor is identical to that of A. chilense Less. and similar to that of Eriachaenium magellanicum Sch. Bip. (Mutisieae) and Artemisia verlotorum Lamotte (Anthemideae). Some grains of A. chilense have endosexine with compact aspect as occurs in E. magellanicum (see also Skvarla et al., 1977; Bittmann, 1990a). Habitat and Distribution. Genus of five species with disjunct distribution in the temperate forests and Patagonia of Argentina and Chile, Guatemala and Mexico, northern United States and southern Canada, and Asia. Species list 1. Adenocaulon bicolor Hook., Bot. Misc. 1: 19. 1830. TYPE: Canada. British Columbia: Fort Vancouver, s.d., D. Scouler s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: United States. California: Sierra Nevada, Placer Co., between Iowa Hill and Forest Hill, J. H. Hunziker 4943 (LP*). 2. Adenocaulon chilense Less., Linnaea 6: 107. 1831. TYPE: Chile. Antuco, s.d., E. F. Poeppig s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Ayzen: Los Mallines near Balmaceda, R. M. Bruzzone 36 (LP*). 3. Adenocaulon himalaicum Edgew., Trans. Linn. Soc. London 20: 64. 1851. TYPE: Nepal. Himalaya, Nagkanda, 1844, M. P. Edgeworth s.n. (holotype, K not seen). Additional specimen examined: Japan. Hokkaido: Moiwayama region, near Sapporo, P. Dorsett & W. Morse 1179 (US). 4. Adenocaulon lyratum S. F. Blake, J. Wash. Acad. Sci. 24: 435 1934. TYPE: Guatemala. Dept. Chimaltenango: Chichauac, 20 Sept 1933, A. F. Sckutch 622 (holotype, US not seen, US photo at LP!). Additional specimen examined:

518

L. Katinas, et al.

Figs. 7–10 7 Aphyllocladus spartioides Wedd. A Habit (branch). B Branch with leaves. C Marginal floret (without cypsela). D Central floret (without cypsela). E Stamen. F Upper part of the style. G Cypsela with pappus. (Redrawn from Cabrera, 1978: 583). Scale bars: A=6 cm; B=3 cm; C, D, G=4 mm; E=2 mm; F=0.8 mm. 8 Brachyclados caespitosus (Phil.) Speg. A Habit. B Leaf, abaxial view. C Marginal floret (without cypsela). D Staminode. E Central floret (without cypsela). F Stamen. G Upper part of the style. H Cypsela with pappus. (Redrawn from Cabrera, 1971b: 317). Scale bars: A=1 cm; B, F=3 mm; C, E, H= 10 mm; D=4 mm; G=1 mm. 9 Catamixis baccharoides Thomson. A Habit (branch). B Corolla. C Upper part of the style. D Cypsela with pappus. (Redrawn from Thomson, 1865: 342). Scale bars: A=4 cm; B, D=3 mm; C=0.8 mm. 10 Chaetanthera splendens (J. Rémy) B. L. Rob. A Habit. B Phyllary of the involucre. C Marginal floret (without pappus). D Staminode. E Central floret. F Stamen. G Upper part of the style. (Redrawn from Cabrera, 1937: 152). Scale bars: A=1 cm; B=6 mm; C, E=4 mm; D=1 mm; F= 3 mm; G=0.6 mm

Mexico. State Chiapas: Mun. of San Cristóbal las Casas, above Rancho Nuevo, 9 mi SE of San Cristóbal las Casas, D. Breedlove 14027 (US). 5. Adenocaulon nepalense M. Bittmann, Candollea 45: 403. 1990. TYPE: Nepal. Jaljale Himal: SE of Jaljale, Farille & Lachard 847509 (holotype, GOET not seen).


519

Literature. Bittmann (1990a, b), Katinas (2000).

Ainsliaea DC. Prodr. 7: 13. 1838. TYPE: Ainsliaea pteropoda DC. [= Ainsliaea latifolia (D. Don) Sch. Bip.]. 6 of Figs. 3–6. Diaspananthus Miq., Ann. Mus. Bot. Lugduno-Batavi 2: 186. 1866, as Diaspanthus. TYPE: Diaspananthus uniflorus (Miq.) Kitam. [= Ainsliaea uniflora Sch. Bip.]. Etymology. Named after the British physician Sir Whitelaw Ainslie (1767–1836), author of Materia medica Indica. Herbs perennial, scapose or rarely small shrubs. Leaves usually rosulate or clustered at the median part of the stem; sessile to distinctly petiolate with petioles winged to wingless; blades linear, elliptic, obovate, deltoid to subrounded, palmately, pinnately, three- to five-veined, margin entire to partite, glabrous to pubescent on both surfaces or only beneath. Capitulescences unilaterally spicate, racemose to paniculate; capitula sessile to pedunculate, homogamous, ligulate, one- to five-flowered; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, chasmogamous or cleistogamous; chasmogamous florets with corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; cleistogamous florets with corolla pseudoligulate, tubular, four- to five-lobed, with one deeper slit but without an expanded lip; anther apical appendages rounded to apiculate, tails slightly to long-papillose; style bilobed, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of plumose, capillary bristles, occasionally absent. Pollen Description. Ainsliaea (82 of Figs. 80–85) (Tellería & Katinas, 2005). Pollen spheroidal to prolate, medium to large size, tricolpate (e.g., A. lancangensis Y. Y. Qian, A. latifolia, and A. macroclinidioides Hayata) or tricolporate (e.g., A. acerifolia Sch. Bip., A. pertyoides Franch., A. spanocephala Y. C. Tseng, and A. uniflora), exine Ainsliaea type or Mutisia type, microechinate (see also Lin et al., 2005). Habitat and Distribution. Genus of 49 Asian species distributed in Afghanistan, Bhutan, China, India, Indonesia, Japan, Kashmir, Korea, Union of Myanmar, Nepal, Pakistan, Philippines, Taiwan, Thailand, and Vietnam. Species list 1. Ainsliaea acerifolia Sch. Bip., Jahresber. Pollichia 18–19: 188. 1861. TYPE: Japan: Sine loc., H. Bürger & H. Zollinger 272 (holotype, P!). Additional specimens examined: Japan. Kyushu: Kund-sho-jan, C. J. Maximowicz s.n. (MO); Honshu: Miyagi Prefecture, Boufford 19871 (CM*). 2. Ainsliaea angustata C. C. Chang, Sin. Contr. 5: 158. 1934. TYPE: China. Szechuan: Ma-Ping hsien, W. P. Fang 1641 (isotype, PE not seen, PE photo at

520

3.

4.

5.

6.

7. 8. 9. 10. 11.

12.

13.

14.

15.

16.

L. Katinas, et al.

LP!). Additional specimens examined: China. Sichuan: Szechuan, Tien-chuanhsien, K. L. Chu 2273 (PE, W). Ainsliaea apiculata Sch. Bip., Jahresber. Pollichia 18–19: 188. 1861. TYPE: Japan: sine loc., H. Bürger & H. Zollinger 266 (lectotype, designated by Freire, 2007: 162, P!). Additional specimens examined: Japan. Honshu: Aichi Prefecture, Mikawa, Kita-shitara-gun, Nagura-mura, Oo-nagura, along Kamsa R., R. M. Furuse s.n. (S); Kyushu: Kagoshina Prefecture, Mt. Kaimon, Saito 1846 (US*). Ainsliaea aptera DC., Prodr. 7: 14. 1838. TYPE: Nepal. India orientale, 1833, Royle s.n. (lectotype, designated by Freire, 2007: 105, K!). Additional specimen examined: India. Punjab: Simla, J. R. Drummond 22635 (UC). Ainsliaea apteroides (C. C. Chang) Y. C. Tseng, Acta Phytotax. Sin. 31: 363. 1993. Basionym: Ainsliaea pteropoda DC. var. apteroides C. C. Chang. TYPE: China. Yunnan: Mi lêh district, A. Henry 9907A (holotype, K not seen; isotype, US!). Additional specimens examined: China. Yunnan: Mengtze, A. Henry 9907 (MO, NY). Ainsliaea bonatii Beauverd, Bull. Soc. Bot. Genève 1: 377. 1909. TYPE: China. Yunnan: Tsong-chan, F. Ducloux 3646 (type, NY!). Additional specimens examined: China. Yunnan: Mi lê, A. Henry 9955 (MO, NY). Ainsliaea brandisiana Kurz, J. Asiat. Soc. Bengal 41: 318. 1872. TYPE: Myanmar. Martaban, S. Kurz 865 (holotype, G!). Ainsliaea caesia Hand.-Mazz., Beih. Bot. Centralbl. 56: 469. 1937. TYPE: China. Guangdong: Kwangtung, R. Mell 874 (holotype, WU!). Ainsliaea cavaleriei H. Lév., Fl. Kouy-Tchéou: 82. 1914. TYPE: China. Sud de Tin-fan, J. Cavalerie 1907 (holotype, E!). Ainsliaea chapaensis Merr., J. Arnold Arbor. 21: 387. 1940. TYPE: Vietnam. Tonkin: Chapa, A. Pételot 2068 (isotype, MO!). Ainsliaea cordifolia Franch. & Sav., Enum. pl. Jap. 2: 416. 1879. TYPE: Japan. Honshu: Mie Prefecture, 7 Oct. 1956, M. Furuse s.n. (epitype, designated by Freire, 2007: 167, S!). Additional specimen examined: Japan. Chiba Prefecture, Awa, Awa-gun, Amatsu-machi, Mt. Kiyosumi on S side, M. Furuse s.n. (S). Ainsliaea crassifolia C. C. Chang, Sin. Contr. 6: 549. 1935. TYPE: China. Yunnan: Yung-pe Mtns., G. Forrest 11070 (lectotype, designated by Freire, 2007: 113, E not seen, E photo at LP!, PE!). Ainsliaea dissecta Franch. & Sav., Enum. pl. Jap. 2: 416. 1879. TYPE: Japan. Honshu: Tottori Prefecture, Mt. Hakko, J. Sugimoto 10912 (epitype, designated by Freire, 2007: 169, S!). Additional specimen examined: Japan. Honshu: Aichi Prefecture, Prov. Mikawa, Mt. Hooraiji, Hooraiji-machi, Minami-shitaragun, M. Furuse s.n. (S). Ainsliaea elegans Hemsl., Icon. pl. 28: t. 2747. 1902. TYPE: China. Yunnan: Mengtze, A. Henry 9108 (lectotype, designated by Freire, 2007: 116, K!). Additional specimen examined: China. Yunnan: Mar-li-po, Kwang-jin-in, K. M. Feng 12964 (KUN). Ainsliaea foliosa Hand.-Mazz., Acta Horti Gothob. 12: 348. 1938. TYPE: China. [Sichuan?]: Sikang, “inter Taining (Ngata) et Taofu (Dawo), Sunglingku,” H. Smith 12120 (holotype, W!). Ainsliaea fragrans Champ. ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 4: 236. 1852. TYPE: China. Guangdong: Hong Kong, J. G. Champion 161 (holotype, K!).


521

17. Ainsliaea fulvipes Jeffrey & W. W. Sm., Notes Roy. Bot. Gard. Edinburgh 8: 175. 1914. TYPE: China. Yunnan: Teng-yueh, G. Forrest 7862 (lectotype, designated by Freire, 2007: 123, S!). 18. Ainsliaea glabra Hemsl., J. Linn. Soc., Bot. 23: 471. 1888. TYPE: China. Sichuan: Szechuan, Mt. Omei, E. Faber 391 (lectotype, designated by Freire, 2007: 123, K!). 19. Ainsliaea glumacea (Fr.) Sch. Bip., Jahresber. Pollichia 20–21: 407. 1863. Basionym: Hieracium glumaceum Fr. TYPE: India. Silhet, 1830, Wallich s.n. (lectotype, designated by Freire, 2007, K-WALL not seen). Additional specimen examined: India. Assam: Tserapúndzi till near Máirong, ex herb. Schlagintweit, Cat. No. 391 (US). 20. Ainsliaea gracilis Franch., J. Bot. (Morot) 8: 297. 1894. TYPE: China. Sichuan: Su-tchuen oriental, Tchen-keou-tin, R. P. Farges 1034A (type, P!). 21. Ainsliaea grossedentata Franch., J. Bot. (Morot) 8: 297. 1894. TYPE: China. Sichuan: Su-tchuen oriental, Tchen- keou-tin, R. P. Farges 333 (isotype, P!). 22. Ainsliaea henryi Diels, Bot. Jahrb. Syst. 29: 628. 1901. TYPE: China. Hubei: Hupeh, A. Henry 6639 (type, K!). Additional specimen examined: China. Hubei: Hupeh, A. Henry 7461 (NY). 23. Ainsliaea hypoleuca Diels ex H. Limpr., Repert. Spec. Nov. Regni Veg. 12: 514. 1922. TYPE: China. Sichuan: Szetchuan, Kwan Hsien, H. Limpricht 1251 (lectotype, designated by Freire, 2007: 131, W!). 24. Ainsliaea kawakamii Hayata, Icon. pl. formos. 8: 72. 1919. TYPE: China. Taiwan: Formosa, s.d., T. Kawakami s.n. (holotype, TI not seen, TI photo at LP!). 25. Ainsliaea lancangensis Y. Y. Qian, J. Trop. Subtrop. Bot. 8: 161. 2000. TYPE: China. Yunnan: Lancang, Y. Y. Qian 3185 (holotype, HITBC not seen). Additional specimen examined: China. Yunnan: Lancang, Y. Y. Qian 3343 (SMAO*). 26. Ainsliaea latifolia (D. Don) Sch. Bip., Jahresber. Pollichia 18: 190. 1861. Basionym: Liatris latifolia D. Don. TYPE: Nepal. Narainhetty Nepalensium, 24 Feb. 1803, Hamilton s.n. (holotype, BM not seen). Additional specimens examined: India, Punjab, Kasauli, J. R. Drummond 20055 (UC). China, Prov. Yunnan, Handel & Mazzetti 8501 (W*). 27. Ainsliaea linearis Makino, Bot. Mag. 23: 250. 1909. TYPE: Japan. Kyushu: Kagoshima Prefecture, Osuma, M. Togasi 1448 (neotype, designated by Freire, 2007: 175, MO!). Additional specimen examined: Japan. Kyushu: Kagoshima Prefecture, Isl. Yakushima, Yaku-cho, from Yodoko Hut along the Yodoko River, D. E. Boufford & S. Mitsuta 20062 (CM). 28. Ainsliaea macrocephala (Mattf.) Y. C. Tseng, Acta Phytotax. Sin. 31: 364. 1993. Basionym: Ainsliaea pteropoda DC. var. macrocephala Mattf. TYPE: China. Yunnan: Likiang, C. Schneider 2328 (lectotype, designated by Freire, 2007: 136, K!). 29. Ainsliaea macroclinidioides Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25, 19: 141. 1908. TYPE: Taiwan, Suizan, Nagasawa 665 (lectotype, designated by Freire, 2007: 177, TI not seen, duplicate, TI photo at LP!). Additional specimen examined: Japan. Ryukyu: Okinawa Prefecture, Isl. Ishigaki, Mt. Omoto-dake, G. Murata 17066 (AAU*).

522

L. Katinas, et al.

30. Ainsliaea mairei H. Lév., Le Monde des Plantes 18: 31. 1916. TYPE: China. Yunnan: Mt. Io-Chan, Nov 1912, E. E. Maire s.n. (holotype, E!). 31. Ainsliaea nana Y. C. Tseng, Acta Phytotax. Sin. 31: 365. 1993. TYPE: China. Sichuan: Tianquan, Chung-luen Wu 12307 (holotype, NAS not seen).. 32. Ainsliaea nervosa Franch., Bull. Mus. Hist. Nat. (Paris) 1: 64. 1895. TYPE: China. Yunnan: Long-Ki, J. M. Delavay 4939 (holotype, P not seen, P photo at LP!). 33. Ainsliaea oblonga Koidz., Bot. Mag. (Tokyo) 28: 150. 1914. TYPE: Japan. Yaeyama archipelago: insl. Nisiomotesima, Aug 1889, Tashiro s.n. (holotype, TI not seen, TI photo at LP!). 34. Ainsliaea parvifolia Merr., Philipp. J. Sci. 12: 110. 1917. TYPE: China. Guangdong: Loh Fau Mtn. (Lofaushan), E. D. Merrill 10237 (holotype, US!). 35. Ainsliaea paucicapitata Hayata, Icon. pl. formos. 8: 71. 1919. TYPE: Taiwan. Nökösan, Jan 1908, U. Mori s.n. (type, TI not seen, TI photo at LP!). 36. Ainsliaea pentaflora S. E. Freire, Novon 12: 451. 2002. TYPE: Vietnam. Tonkin: Massif du Tom Dav, A. Pételot 417 (holotype, C!). 37. Ainsliaea pertyoides Franch., J. Bot. (Morot) 2: 70. 1888. TYPE: China. Yunnan: ad fauces Kien-min-keou, prope Mo-so-yn, J. M. Delavay 1218 (lectotype, designated by Freire, 2007: 143, P!). Additional specimen examined: China. Yunnan, 28 Jan 1889, J. M. Delavay s.n. (UC*). 38. Ainsliaea pingbianensis Y. C. Tseng, Acta Phytotax. Sin. 31: 365. 1993. TYPE: China. Yunnan: Pingbian, P. Mao 4186 (holotype, IBSC not seen, IBSC photo at LP!). 39. Ainsliaea qianiana S. E. Freire, Novon 12: 453. 2002. TYPE: China. Yunnan: Colline de Kin-tchong-chan, E. E. Maire 2529 (holotype, W!; isotypes, NY!, UC!). 40. Ainsliaea ramosa Hemsl., J. Linn. Soc., Bot. 23: 471. 1888. TYPE: China. Hubei: Hupeh, Nanto and mtns. to the N, A. Henry 4422 (holotype, K not seen; isotype, US!). 41. Ainsliaea reflexa Merr., Philipp. J. Sci. 1: 242. 1906. TYPE: Philippines. Luzon: Lepanto, Mt. Data, E. D. Merrill 4571 (holotype, NY!). 42. Ainsliaea rubrinervis C. C. Chang, Sin. Contr. 4: 226. 1934. TYPE: China. Sichuan: Kwan-hsien, W. P. Fang 2076 (holotype, N not seen; isotype, NY!). 43. Ainsliaea smithii Mattf., Acta Horti Gothob. 8: 79. 1933. TYPE: China. Sichuan: Sze-chuan, Drogochi, H. Smith 4698 (isotype, W!). 44. Ainsliaea spanocephala Y. C. Tseng, Acta Phytotax. Sin. 26: 74. 1988. TYPE: Thailand. Loei: Phu Luang, Wildlife Sanctuary, H. Koyama, H. Terao & W. Thawatchai 33699 (isotypes, BKF!, GH!, KYO!). Additional specimens examined: Thailand. Loei: Phu Luang, Wildlife Sanctuary, 1982, T. Smitinand s.n. (BKF 077323*); id., Nuijomdha & Vidal 438 (C*). 45. Ainsliaea spicata Vaniot, Bull. Acad. Int. Géogr. Bot. 12: 117. 1903. TYPE: China. Yunnan: environs de Yun-nan-sen, F. Ducloux 9D (holotype, E!; isotype, W!). 46. Ainsliaea trinervis Y. C. Tseng, Acta Phytotax. Sin. 31: 367. 1993. TYPE: China. Guangxi: Rongshui, S. Q. Chen 16409 (holotype, IBSC not seen; isotype, TI not seen, TI photo at LP!).


523

47. Ainsliaea uniflora Sch. Bip., Jahresber. Pollichia 18: 187. 1861. TYPE: Japan. H. Burger & H. Zollinger 283 (type, K!). Additional specimen examined: Honshu, Mie, Ise, Yamada, 10 Oct 1911, s. leg. (US 1132181*). 48. Ainsliaea walkeri Hook. f., Bot. Mag. 102: tab. 6225. 1876. TYPE: China. Guangdong: Hong Kong, Dec 1875, Capt. A. L. Wilkes s.n. (holotype, K!). 49. Ainsliaea yunnanensis Franch., J. Bot. (Morot) 2: 70. 1888. TYPE: China. Yunnan: Che-tcho-tze, supra Tapin-tze, J. M. Delavay 608 (type, NY!).

Literature. Freire (2002, 2007), Mitsui et al. (2007).

Aphyllocladus Wedd. Chlor. andina 1: 11. 1855. TYPE: Aphyllocladus spartioides Wedd. 7 of Figs. 7–10. Jobaphes Phil., Fl. atacam.: 27. 1860. TYPE: Jobaphes virgatus Phil. [= Aphyllocladus denticulatus (J. Rémy) Cabrera]. Etymology. From the Greek a, without, phyllon, leaf, and clados, branch, with reference to the aphyllous branches. Shrubs. Leaves alternate, reduced, early caducous; sessile; blades linear-oblong to linear-ovate, pinnately or three- to five-veined, margin entire, subglabrous to lanuginous on both surfaces. Capitulescences monocephalous, terminal; capitula pedunculate, homogamous or heterogamous, radiate or disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic or isomorphic; marginal florets bisexual or female, corolla bilabiate, outer lip expanded or not; central florets or all florets bisexual, with corolla tubular-funnelform, deeply five-lobed; anther apical appendages truncate, reddish, tails papillose; style bifid, branches dorsally papillose. Cypselae densely long-villose; pappus of scabrid to barbellate bristles, the innermost flat, longer, and wider than the others and barbellate at the apex. Pollen Description. Aphyllocladus (83 of Figs. 80–85) (Tellería & Katinas, 2004). Pollen prolate to subprolate, large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Genus of five species from the Andes of Argentina, Bolivia, and Chile. Species list 1. Aphyllocladus decussatus Hieron., Icon. Descr. Pl.: 28. 1855. TYPE: Argentina. Prov. La Rioja: Los Corrales, Sierra de Famatina, G. H. E. W. Hieronymus & G. Niederlein 831 (holotype, B, not seen, B photo F n. 15873 at LP!; isotype, GH not seen). 2. Aphyllocladus denticulatus (J. Rémy) Cabrera, Darwiniana 9: 367. 1951. Basionym: Cyclolepis denticulata J. Rémy. TYPE: Chile. Copiapo, C. Gay 239 (holotype, P not seen, P photo F n. 38078 at LP!). Additional specimen examined: Chile. Prov. Atacama: Bandurrias, s.d., N. Geisse s.n. (LP).

524

L. Katinas, et al.

3. Aphyllocladus ephedroides Cabrera, Darwiniana 9: 370. 1951. TYPE: Argentina. Prov. La Rioja: Dept. General Sarmiento, Vinchina, G. Covas 1229 (holotype, LP!). 4. Aphyllocladus sanmartinianus Molfino, Bol. Soc. Argent. Bot. 5: 30. 1953. TYPE: Argentina. Prov. San Juan: Iglesias, Cuesta de la Casitas, R. A. Spegazzini 230 (holotype, BAB, not seen). Additional specimen examined: Argentina. Prov. San Juan: camino a Las Hornillas, A. L. Cabrera et al. 24290 (LP*). 5. Aphyllocladus spartioides Wedd., Chlor. andina 1: 11. 1855. TYPE: Bolivia. Chuquisaca and Potosí, 1843–1847, H. A. Weddell s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, Humahuaca, A. Ruiz Leal 14386 (LP). Literature. Cabrera (1951, 1978), Molfino (1953), Freire (1996). Brachyclados D. Don Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Brachyclados lycioides D. Don. 8 of Figs. 7–10. Lavidia Phil., Anales Univ. Chile 85: 837. 1894. TYPE: Lavidia caespitosa Phil. [= Brachyclados caespitosus (Phil.) Speg.]. Etymology. From the Greek brachys, short, and klados, branch, with reference to the short shoots or brachyblasts. Shrubs stout to cushion-like, with brachyblasts and macroblasts. Leaves alternate, rigid; sessile, clasping; blades linear, ovate, subulate to oblong, parallel veins in the sheathing portion and one prominent central vein in the blade, margin entire, revolute, glabrous above, pubescent beneath. Capitulescences monocephalous, terminal; capitula subsessile, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bilobed, branches dorsally papillose. Cypselae truncate to shortly attenuate at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Brachyclados (84 of Figs. 80–85) (Tellería & Katinas, 2004). Pollen subprolate to prolate, large size, tricolporate, exine Mutisia type, microechinate or almost psilate (see also Parra & Marticorena, 1972). Habitat and Distribution. Genus of three species of central Chile, Patagonia, and western Argentina. Species list 1. Brachyclados caespitosus (Phil.) Speg., Anales Soc. Ci. Argent. 53: 28. 1902. Basionym: Lavidia caespitosa Phil. TYPE: Chile. Patagonia, Lago Pinto, s.d., H. Ibar s.n. (type, probably SGO). Additional specimen examined: Argentina. Prov. Chubut: cerca de Facundo, A. L. Cabrera et al. 23227 (LP).


525

2. Brachyclados lycioides D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido de Patagones, cerca de Cardenal Cagliero, A. L. Cabrera & H. A. Fabris 14865 (LP*). 3. Brachyclados megalanthus Speg., Anales Soc. Ci. Argent. 53: 29. 1902. TYPE: Argentina. Prov. Buenos Aires: Tres Cerros to Río Negro, Carmen de Patagones, Feb 1898, C. L. Spegazzini s.n. (syntypes, LP!).

Observation. Brachyclados lycioides constitutes a very important element in the diet composition of the guanaco (Lama guanicoe) (Candia & Dalmasso, 1995). Literature. Cabrera (1971b), Katinas (1996a).

Catamixis Thomson J. Linn. Soc., Bot. 9: 342. 1867. TYPE: Catamixis baccharoides Thomson. 9 of Figs. 7–10. Etymology. From the Greek katamixis, katameixis, admixture, or kata and mixis, a mixing, mingling because the affinities in Catamixis point in many directions. Subshrubs branched, erect. Leaves alternate; short-petiolate; blades obovate to oblong, pinnately veined, margin obtusely dentate-serrate, pubescent to glabrate. Capitulescences corymbose; capitula pedunculate, homogamous, ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb shallowly five-lobed; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae truncate at the apex, densely pubescent; pappus of scabrid, capillary bristles, unequal in length. Pollen Description. Pollen not seen. Habitat and Distribution. Monotypic genus endemic to the narrow range in the Siwalik belt and Theri district of Garwhal Himalayas in India and Nepal. Species list 1. Catamixis baccharoides Thomson, J. Linn. Soc., Bot. 9: 343. 1867. TYPE: India. Uttaranchal: District Garhwal, Siwalicis Himalaya, s.d., s. leg. (syntypes, K not seen, photos at LP!). Observation. The ligulate corolla of Catamixis results unusual for Mutisioideae, where there is a predominance of bilabiate corollas (although some species of Mutisia have also ligulate florets). According to Small (1919), all five petals in the corolla of Catamixis are fused and the split is posterior (the posterior lip corresponds to the two-lobed lip in the typical bilabiate corolla) so that something like a ligulate

526

L. Katinas, et al.

corolla results. But this is just one more variation in the manner of splitting of the variable corollas of Mutisioideae and cannot be taken as an intermediate type leading to the Cichorieae (Small, 1919). The genus and its only species could be on the verge of extinction (Nayar & Ahmedullah, 1985). Literature. Nayar and Ahmedullah (1985), Rao (1995).

Chaetanthera Ruiz & Pav. Fl. peruv. prodr.: 106. 1794. TYPE: Chaetanthera ciliata Ruiz & Pav. 10 of Figs. 7–10. Aldunantea J. Rémy, Fl. chil. 3: 320. 1849. TYPE: not designated, based on the three species Aldunantea chilensis J. Rémy, A. gnaphalioides J. Rémy, and A. lycopodioides J. Rémy. Carmelita Gay ex DC., Prodr. 7: 14. 1838. [= Chaetanthera subgen. Carmelita (Gay) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 97. 1937]. TYPE: Carmelita formosa Gay ex DC. [= Chaetanthera villosa D. Don]. Cherina Cass., Dict. sci. nat. 8: 437. 1817. TYPE: Cherina micropylla Cass. [= Chaetanthera microphylla (Cass.) Hook. & Arn.]. Chondrochilus Phil., Linnaea 28: 711. 1858. TYPE: Chondrochilus crenatus Phil. [= Chaetanthera renifolia (J. Rémy) Cabrera]. Egania J. Rémy, Fl. chil. 3: 324. 1849. [= Chaetanthera subgen. Egania (J. Rémy) Reiche, Fl. chil. 4: 355. 1905]. TYPE: not designated, based on the three species Egania acerosa J. Rémy, E. apiculata J. Rémy, and E. dioica J. Rémy. Elachia DC., Prodr. 7: 256. 1838. TYPE: Elachia euphrasioides DC. [= Chaetanthera euphrasioides (DC.) F. Meigen]. Euthrixia D. Don, Trans. Linn. Soc. London 16: 257. 1830. TYPE: Euthrixia salsoloides D. Don [= Chaetanthera microphylla (Cass.) Hook. & Arn.]. Minythodes Phil., Anales Univ. Chile 115: 317. 1904, nom. nudum. Oriastrum Poepp. & Endl., Nov. gen. sp. pl. 3: 50. 1842. [= Chaetanthera subgen. Oriastrum (Poepp. & Endl.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 96. 1937]. TYPE: Oriastrum pusillum Poepp. & Endl. [= Chaetanthera pulvinata (Phil.) Hauman]. Proselia D. Don, Trans. Linn. Soc. London 16: 234. 1830. [= Chaetanthera subgen. Proselia D. Don, l. c.]. TYPE: Proselia serrata D. Don [= Chaetanthera serrata Ruiz & Pav.]. Tiltilia Phil., Anales Univ. Chile 115: 323. 1904, nom. nudum. Tylloma D. Don, Trans. Linn. Soc. London 16: 238. 1830. [= Chaetanthera subgen. Tylloma (D. Don) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 97. 1937]. TYPE: Tylloma limbatum D. Don [= Chaetanthera limbata (D. Don) Less.]. Etymology. From the Greek chaite, bristle, long hair and anthera, anther, alluding to the papillose tails of the anthers. Herbs caulescent, stout to cushion-like, annual or perennial, rarely shrubs, monoecious, gynodioecious, or dioecious. Leaves alternate, sparse to densely


527

imbricate; sessile to clasping; blades linear, subulate, oblanceolate to spathulate, pinnately, three-veined, or with a conspicuous central vein, margin entire, ciliate to dentate, glabrous to densely pubescent. Capitulescences monocephalous, terminal; capitula sessile to pedunculate, homogamous or heterogamous, discoid, disciform, or radiate; receptacle epaleate; involucre multiseriate, phyllaries acute at the apex or with an apical, showy appendage. Florets isomorphic in dioecious and gynodioecious plants, or dimorphic in monoecious plants; marginal florets female or neuter, with or without staminodes, corolla bilabiate, outer lip expanded or not, inner lip developed to absent, sub-bilabiate (3+1 corolla lips), or true ray; central florets bisexual or male, corolla bilabiate to tubular-bilabiate; female capitula with all florets female, bilabiate; anther apical appendages acute, tails papillose; style bilobed to bifid, branches dorsally papillose. Cypselae truncate at the apex, glabrous to pubescent; pappus of scabrid, simple to barbellate, capillary bristles. Pollen Description. Chaetanthera (85 of Figs. 80–85) (Tellería & Katinas, 2004). Pollen spheroidal–subprolate to prolate, size medium to large, tricolporate, with endocingulus, exine Mutisia type, microechinate. Two pollen types were recognized, Type I (found in 14 species) with pollen subrectangular in equatorial view, subcircular or subtriangular in polar view, medium size, nexine outline elliptic to subrectangular in shape, exine 5–7 μm thick; and Type II (found in 25 species) with pollen elliptic in equatorial view, large size, nexine outline dumbbell-shaped, exine 11–18 μm thick. In the type II, two subtypes were recognized (see also Parra & Marticorena, 1972; Markgraf & D’Antoni, 1978; Wingenroth & Heusser, 1983). Habitat and Distribution. Genus of 45 South American species distributed mainly in the Andes, from Peru to southern Argentina and Chile. Species list 1. Chaetanthera acerosa (J. Rémy) Benth. & Hook. f. ex Griseb., Symb. fl. argent.: 214. 1879. Basionym: Egania acerosa J. Rémy. TYPE: Sine loc., s.d., C. Gay s.n. (types probably GH, P). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Rafael, valle de Las Leñas, A. Ruiz Leal 24661 (LP). 2. Chaetanthera acheno-hirsuta (Tombesi) Arroyo, A. M. R. Davies & Till-Bottr., Gayana, Bot. 61: 28. 2004. Basionym: Chaetanthera pulvinata (Phil.) Hauman var. acheno-hirsuta Tombesi. TYPE: Argentina. Prov. San Juan: Dept. Iglesia, Mina Las Carachas, E. Nicora et al. 8217 (holotype, SI!). 3. Chaetanthera apiculata (J. Rémy) F. Meigen, Bot. Jahrb. Syst. 18: 456. 1894. Basionym: Egania apiculata J. Rémy. TYPE: Chile. Coquimbo, Pasto Blanco, Los Patos, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Santiago: cordillera Río San Francisco, Fierro Carrera, E. Werdermann 627 (LP). 4. Chaetanthera australis Cabrera, Notas Mus. La Plata, Bot. 1: 64. 1935. TYPE: Argentina. Prov. Río Negro: San Carlos de Bariloche, arroyo Ñirico, A. Burkart 6599 (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquén: near San Martín de los Andes, Böcher et al. 1658 (LP).

528

L. Katinas, et al.

5. Chaetanthera aymarae Martic. & Quezada, Bol. Soc. Biol. Concepción 48: 107. 1974. TYPE: Chile. Prov. Tarapacá: Dept. Arica, camino de Zaphahuira a Putre, Quebrada de Socoroma, M. Ricardi et al. 166 (holotype, CONC!). 6. Chaetanthera boliviensis J. Kost., Blumea 5: 673. 1945. TYPE: Bolivia. Choquecatachico, T. Herzog 2339 (type, probably M). 7. Chaetanthera brachylepis Phil., Anales Univ. Chile 87: 12. 1894. TYPE: Chile. Prov. Arauco: Cordillera de Nahuelbuta, vega de Rucapillán, Jan 1877, R. Philippi s.n. (holotype, S not seen, S photo F n. 15960 at LP!). Additional specimen examined: Chile. Prov. Curicó: Potrero Grande, Lomas Blancas, E. Barros 7443 (LP*). 8. Chaetanthera chilensis (Willd.) DC., Ann. Mus. Natl. Hist. Nat. 19: 70. 1812. Basionym: Perdicium chilense Willd. TYPE: Chile. Prov. Ñuble: Cucha Cucha, s.d., s. leg., herb. Willdenow n. 16092 (holotype, BD not seen, BD photo at LP!). Additional specimen examined: Chile. Prov. Ñuble: Chillán, Bureo, E. Barros 7441 (LP). 9. Chaetanthera chiquianensis Ferreyra, Publ. Mus. Hist. Nat. “Javier Prado,” Ser. B, Bot. 6: 5. 1953. TYPE: Peru. Dept. Ancash: Prov. Bolognesi, cerro al S de Chiquián, E. Cerrate 1323 (isotype, LP!). 10. Chaetanthera ciliata Ruiz & Pav., Syst. veg. fl. peruv. chil.: 190. 1798. TYPE: Chile. Prov. Maule: Huiquilemu, s.d., s. leg. (type, probably MA). Additional specimen examined: Chile. Prov. Ñuble, Brochers 1071c (LP). 11. Chaetanthera cochlearifolia (A. Gray) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Oriastrum cochlearifolium A. Gray. TYPE: Peru. Alpamarca, s.d., leg. U. S. Explor. Exped. under Cap. Wilkes s.n. (holotype, GH not seen). Additional specimen examined: Peru. Prov. Junín: Casapalca, Macbride & Featherstone 845 (LP). 12. Chaetanthera dioica (J. Rémy) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Egania dioica J. Rémy. TYPE: Chile. Cordilleras de Coquimbo, cerca de Pasto Blanco, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Argentina. Prov. La Rioja: Sierra de Famatina, La Mesada, Río Amarillo superior, F. Kurtz 13717 (LP). 13. Chaetanthera elegans Phil., Linnaea 28: 712. 1856. TYPE: Chile. Prov. Concepción: Los Angeles, C. Gay 806 (isotype, S not seen). Additional specimen examined: Chile. Prov. Linares: Cordillera de Linares, Los Hualis, E. Barros 2410 (LP). 14. Chaetanthera euphrasioides (DC.) F. Meigen, Bot. Jahrb. Syst. 17: 284. 1893. Basionym: Elachia euphrasioides DC. TYPE: Chile. Altos Andes, s.d., C. Gay s.n. (type, probably GH, P). Additional specimen examined: Chile. Prov. Santiago: Quebrada de Jorquera, cerca del Refugio de Farellones, cordillera de Las Condes, G. Looser 5751 (LP). 15. Chaetanthera flabellata D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Chile. Cordilleras, Cummings 291 (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: Piuquencillos, valle del Río Colorado, Pisano et al. 1641 (LP). 16. Chaetanthera flabellifolia Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 153. 1937, nom. novum. Basionym: Pachylaena elegans Phil. TYPE: Chile. Prov. Coquimbo, Baños del Toro, 1860–1861, H. Volckmann s.n. (holotype, S not


17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

529

seen). Additional specimen examined: Chile. Prov. Coquimbo: Baños del Toro, E. Werdermann 189 (LP). Chaetanthera glabrata (DC.) F. Meigen, Bot. Jahrb. Syst. 23: 456. 1894. Basionym: Tylloma glabratum DC. TYPE: Chile. Collina, s.d., C. Macrae s.n. (type, probably K). Additional specimen examined: Chile. Prov. Elqui: Dept. Elqui, Vicuña, Cuesta de Elqui, G. Looser 4389 (LP). Chaetanthera glandulosa J. Rémy, Fl. chil. 3: 311. 1847. TYPE: Chile: s.d., C. Gay s.n. (isotype, B not seen, B photo F n. 15968 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Morro Blanco, Cabrería, C. Jiles 1237 (LP). Chaetanthera gnaphalioides (J. Rémy) I. M. Johnst., Physis (Buenos Aires) 9: 325. 1929. Basionym: Aldunatea gnaphalioides J. Rémy. TYPE: Chile. Prov. Coquimbo: cerros de las cordilleras de Doña Ana, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Dept. Elqui: 3 km al interior de Rivadavia, en el camino a Huanta, Wagenknecht 4388 (LP). Chaetanthera incana Poepp. ex Less., Linnaea 5: 284. 1830. TYPE: Chile. Prov. Aconcagua: Concón, E. F. Poeppig 218 (holotype, B not seen, B photo F n. 15969 at LP!; isotype, BD not seen). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Illapel, al N de Los Vilos, C. Jiles 2326 (LP). Chaetanthera kalinae A. M. R. Davies, Novon 16: 51. 2006. TYPE: Chile. Prov. Elqui: Vicuña, Embalse La Laguna, C. Ehrhart 61 (isotype, MO not seen). Chaetanthera lanata (Phil.) I. M. Johnst., Physis (Buenos Aires) 9: 325. 1929. Basionym: Tylloma lanatum Phil. TYPE: Chile. Prov. Atacama: Vaquillas, s.d., R. Philippi s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Coquimbo: Baños del Toro, A. L. Cabrera 3550 (LP!). Chaetanthera leptocephala Cabrera, Notas Mus. La Plata, Bot. 17: 78. 1954. TYPE: Chile. Prov. Coquimbo: Quebrada del Toro, 20 km E de Hurtado, hacienda El Bosque, E. Wagenknecht 18490 (holotype, CONC!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Illapel, hacienda Cuncumen, Cuncumen, a 77 km al E de Illapel, Muñoz & Johnson 2192 (LP). Chaetanthera limbata (D. Don) Less., Syn. gen. Compos.: 116. 1832. Basionym: Tylloma limbatum D. Don. TYPE: Chile. Prov. Coquimbo: Coquimbo, s.d., Caldcleugh s.n. (holotype, G not seen, G photo F n. 28856 at LP!). Additional specimen examined: Chile. Prov. El Loa: Atacama, Estación Cuba, E. Barros 2402 (LP). Chaetanthera linearis Poepp. ex Less., Syn. gen. Compos.: 112. 1832. TYPE: Chile: s.d., C. Bertero 912 (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: cordillera de Santiago, Refugio de Farellones, G. Looser 5214 (LP). Chaetanthera lycopodioides (J. Rémy) Cabrera, Revista Centro Estud. Doct. Ci. Nat. 1: 58. 1935. Basionym: Aldunantea lycopodioides J. Rémy. TYPE: Chile. Santiago, Cordillera de Los Patos, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Los Andes: Potrero Escondido, O. Boelcke 2468 (LP). Chaetanthera microphylla (Cass.) Hook. & Arn., Comp. Bot. Mag. 1: 104. 1835. Basionym: Cherina microphylla Cass. TYPE: Chile, s.d., s. leg. herb. M.

530

28.

29.

30.

31.

32.

33.

34.

35.

36.

37.

L. Katinas, et al.

de Jussieu s.n. (type, probably P). Additional specimen examined: Argentina. Prov. Neuquén: Andacollo, Mina Guaraco, A. L. Cabrera 11120 (LP). Chaetanthera minuta (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 134. 1937. Basionym: Tylloma minutum Phil. TYPE: Chile. Prov. Atacama: Pastos Largos, 6 Jan 1885, F. Philippi s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Atacama: Dept. Copiapó, Mondaca, A. Krapovickas & J. H. Hunziker 5700 (LP). Chaetanthera moenchioides Less., Syn. gen. Compos.: 113. 1832. TYPE: Chile. Prov. Concepción: Antuco, s.d., E. F. Poeppig 208 (holotype, BD not seen, BD photo F n. 15974 at LP!). Additional specimen examined: Chile. Chacabuco: Colina, s.d., s. leg. (LP). Chaetanthera pentacaenoides (Phil.) Hauman, Anales Soc. Ci. Argent. 86: 317. 1918. Basionym: Oriastrum pentacaenoides Phil. TYPE: Argentina. Prov. Mendoza: entre Mendoza y Santa Rosa, 1869, s. leg. (holotype, S not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Alto de los Manantiales, A. Ruiz Leal & F. Roig 23617 (LP). Chaetanthera perpusilla (Wedd.) A. Anderb. & S. E. Freire, Taxon 39: 431. 1990. Basionym: Luciliopsis perpusilla Wedd. TYPE: Bolivia. Dept. La Paz: Ravin de Chuquiaguillo, La Lancha, 1851, H. A. Weddell s.n. (holotype, P!). Chaetanthera peruviana A. Gray, Proc. Amer. Acad. Arts 5: 144. 1861. TYPE: Peru. Andes, Baños, s.d., leg. U. S. Explor. Exped. under Capt. Wilkes s.n. (holotype, GH not seen). Additional specimen examined: Peru. Dept. Ancash: Prov. Bolognesi, Rumipufuio, E. Cerrate et al. 6490 (LP). Chaetanthera planiseta Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 128, nom. novum. Basionym: Oriastrum pusillum Poepp. & Endl. TYPE: Chile. Andes, Cordillera de San Jago, s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Cachapoal: Rancagua, cordillera de Codegua, E. Barros 9849 (LP). Chaetanthera pulvinata (Phil.) Hauman, Physis (Buenos Aires) 3: 420. 1917. Basionym: Oriastrum pulvinatum Phil. TYPE: Chile. Prov. Coquimbo: Cordillera de Doña Ana, 1861, H. Volckmann s.n. (holotype, S not seen). Additional specimen examined: Argentina. Prov. San Juan: Valle del Cura, pirca de los Ingenieros, R. Pérez Moreau 158 (LP). Chaetanthera pusilla (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 106. 1835. Basionym: Tylloma pusillum D. Don. TYPE: Argentina. Prov. Mendoza: Andes de Mendoza, s.d., J. Gillies s.n. (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: cordillera de Santiago, Jan 1872, Germain s.n. (LP). Chaetanthera renifolia (J. Rémy) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 149. 1937. Basionym: Elachia renifolia J. Rémy. TYPE: Chile. Prov. Santiago: La Polvadera, s.d., s. leg. (holotype, P? not seen, photo F n. 38089 at LP!). Additional specimen examined: Chile. Area Metropolitana: Cajón de la Yerba Loca, Feb 1891, R. Philippi s.n. (LP). Chaetanthera revoluta (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 106. 1937. Basionym: Egania revoluta Phil. TYPE: Chile. Prov. Antofagasta: entre Aguas Calientes y Socaire, 5 Feb 1885, F. Philippi s.n. (isotype, S not seen, S photo F n. 15979 at LP!). Additional specimen examined: Argentina.


38.

39.

40.

41.

42.

43.

44.

45.

531

Prov. Salta: Dept. San Antonio de los Cobres, abra Alto Chorrillos, A. L. Cabrera 8874 (LP*). Chaetanthera serrata Ruiz & Pav., Syst. veg. fl. peruv. chil.: 191. 1798. TYPE: Chile. Prov. Concepción: Concepción, s.d., H. Pavón s.n. (syntype, BM not seen, photo at LP!). Additional specimen examined: Chile. Prov. Concepción: Tomé, Lomas de Cocholzué, Junge 1282 (LP). Chaetanthera spathulifolia Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 141. 1937. TYPE: Chile. Prov. Coquimbo: Illapel, El Rodel, Jan 1888, Z. Vergara s.n. (isotype, S not seen). Additional specimen examined: Argentina. Prov. Mendoza: Cuchilla del Diablo, 15 Feb 1934, R. Pérez Moreau s.n. (LP). Chaetanthera sphaeroidalis (Reiche) Hicken, Darwiniana 1: 41. 1922. Basionym: Oriastrum sphaeroidale Reiche. TYPE: Chile. Prov. Atacama: desierto de Atacama, 1897, E. Delaigne s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Huasco: Atacama, Vallenar, cordillera Laguna Chica, E. Werdermann 253 (LP). Chaetanthera splendens (J. Rémy) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Elachia splendens J. Rémy. TYPE: Chile. Prov. Santiago: colina Chimborango, etc., s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Cabrería, C. Jiles 1567 (LP). Chaetanthera stuebelii Hieron., Bot. Jahrb. Syst. 21: 368. 1896. TYPE: Bolivia. Sicasica, entre Tomarapé y La Paz, A. Stuebel 15a (holotype, BD not seen). Additional specimen examined: Argentina. Prov. La Rioja: Sierra de Famatina, ladera este, A. Krapovickas & J. H. Hunziker 5332 (LP). Chaetanthera tenella Less., Syn. gen. Compos.: 114. 1832. TYPE: Chile. Cachapual: Rancagua and Quillota, C. G. Bertero 162, 910 (probable syntypes, NY, NY photos at LP!). Chaetanthera valdiviana Phil., Linnaea 28: 712. 1856. TYPE: Chile. Prov. Valdivia: San Juan, Jan 1855, R. Philippi s.n. (isotype S, not seen). Additional specimen examined: Chile. Prov. Valdivia: San Juan, R. Philippi 1068 (LP). Chaetanthera villosa D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Argentina, Prov. Mendoza: El Planchón, s.d., J. Gillies s.n. (syntype, GH not seen). Additional specimen examined: Argentina. Prov. Río Negro: Cerro Utne, C. C. Hosseus 435 (LP).

Literature. Cabrera (1937), Koster (1945), Ferreyra (1953), Cabrera (1954), Marticorena and Quezada (1974), Anderberg and Freire (1990), Arroyo et al. (2004), Davies (2006), Hershkovitz et al. (2006).

Chaptalia Vent., nom. cons. Descr. pl. nouv. 7: pl. 61. 1802. TYPE: Chaptalia tomentosa Vent. 11 of Figs. 11–14. Leria DC., Ann. Mus. Natl. Hist. Nat. 19: 68. 1812, non Adans., 1763, nom. illeg. [= Chaptalia sect. Leria (DC.) Burkart, Darwiniana 6: 560. 1944]. TYPE: Leria nutans (L.) DC., based on Tussilago nutans L. [= Chaptalia nutans (L.) Pol.].

532

L. Katinas, et al.

Figs. 11–14 11 Chaptalia nutans (L.) Pol. A Habit. B Outermost marginal floret (without cypsela). C Innermost marginal floret (without cypsela). D Central corolla. E Stamen. F Upper part of the style. G Cypsela. H Pappus. (Redrawn from Burkart, 1944: 566). Scale bars: A=4 cm; B–D=4 mm; E, F=1 mm; G= 5 mm; H=6 mm. 12 Chucoa ilicifolia Cabrera. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (From López 1090, LP). Scale bars: A=5 cm; B, E=4 mm; C=2 mm; D= 1.5 mm. 13 Cnicothamnus lorentzii Griseb. A Habit (branch). B Phyllary. C Marginal corolla. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (Redrawn from Cabrera, 1978: 577). Scale bars: A=4 cm; B–D, G=10 mm; E, F=5 mm. 14 Cyclolepis genistoides D. Don. A Habit (branch). B Corolla of female capitula. C Corolla of bisexual capitula. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1978: 591). Scale bars: A=4 cm; B, C, F=5 mm; D, E=2 mm

Lieberkuhna Cass., Dict. sci. nat. 26: 286. 1823. [= Chaptalia sect. Lieberkuhna (Cass.) Burkart, Darwiniana 6: 539. 1944]. TYPE: Lieberkuhna bracteata Cass., nom.illeg., based on Perdicium piloselloides Vahl [= Chaptalia piloselloides (Vahl) Baker].


533

Loxodon Cass., Dict. sci. nat. 27: 253. 1823. [= Chaptalia sect. Loxodon (Cass.) Burkart, Darwiniana 6: 534. 1944]. TYPE: Loxodon brevipes Cass., nom. illeg., based on Tussilago exscapa Pers. [= Chaptalia exscapa (Pers.) Baker]. Oxydon Less., Linnaea 5: 357. 1830. TYPE: Oxydon bicolor Willd. ex Less., nom. illeg., based on Chaptalia runcinata Kunth. Thyrsanthema Neck. ex Kuntze, Revis. gen. pl. 1: 369. 1891, nom. rejic. (see obs. of Nesom, 1995: 154). Etymology. In honor of the French chemist Jean Antoine Chaptal (1756–1832). Herbs perennial, acaulescent, scapose. Leaves rosulate; sessile to pseudopetiolate; blades entire to lyrate, lanceolate, elliptical, obovate to spathulate, pinnately veined, margin entire to runcinate, glabrous to glabrescent above, densely pubescent beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform or radiate, chasmogamous or cleistogamous, erect or nodding; receptacle epaleate; involucre multiseriate. Florets trimorphic; marginal florets female, without staminodes, corolla true ray to bilabiate with minute inner lips, outer lip expanded or not; intermediate florets female, staminodes absent, corolla filiform irregularly tubular, true ray to bilabiate, shorter than the style; central florets bisexual, corolla bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth to papillose; style bifid, branches dorsally papillose. Cypselae rostrate to long-attenuate at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Chaptalia (86 of Figs. 86–91) (Parra & Marticorena, 1972; Salgado-Labouriau, 1982). Pollen subprolate to prolate, medium to large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Chaptalia has currently ca. 70 species distributed in America, from southern United States to Central Argentina, although the number of species could be reduced to 20–35 (see Observation). Species list 1. Chaptalia albicans (Sw.) Vent. ex Steud, Nomencl. bot. 1: 344. 1840. Basionym: Tussilago albicans Sw. TYPE: Jamaica? s.d., P. Browne s.n. (lectotype, designated by Nesom, 1984: 398, LINN 953.16, microfiche 537 not seen, photo at LP!). Additional specimen examined: Bahamas. New Providence: near Nassau, Curtis 181 (MO). 2. Chaptalia angustata Urb., Symb. antill. 7: 432. 1912. TYPE: Dominican Republic. Santo Domingo: Constanza, H. von Türckheim 2908 (syntype, US!). 3. Chaptalia anisobasis S. F. Blake, J. Wash. Acad. Sci. 25: 324. 1935. TYPE: Colombia. Dept. Magdalena: Páramos of the Sierra Nevada de Santa Marta, about 30 mi. inland from Dibulla, W. Seifriz 484 (holotype, US!). 4. Chaptalia arechavaletae Hieron. ex Arechav., Anales Mus. Nac. Montevideo 2: 14. 1904. TYPE: Uruguay: s.d., J. Arechavaleta s.n. (holotype, B not seen, B photo F n. 15998 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Belgrano bajo, A. Burkart 1696 (SI).

534

L. Katinas, et al.

5. Chaptalia azuensis Urb. & Ekman, Ark. Bot. 23: 95. 1930. TYPE: Dominican Republic. Prov. de Azua: Cordillera Central, Loma Nalga de Maco, E. L. Ekman H 6305 (isotype, US!). 6. Chaptalia callacallensis Cuatrec., Fontqueria 9: 5. 1985. TYPE: Peru. Dept. Amazonas: Prov. Chachapoyas, cerros Calla Calla, 18 km above Leimebamba, on road to Balsas, P. C. Hutchinson & J. K. Wright 5589 (type, US!). 7. Chaptalia chapadensis D. J. N. Hind, Kew Bull. 54: 993. 2000. TYPE: Brazil. State Bahia: Mun. Palmeiras, Pai Inácio, BR242W of Lençois at km 232, Mori & Boom 14388 (holotype, K not seen, K photo at LP!). 8. Chaptalia cipoensis Roque, Kew Bull. 60: 133. 2005. TYPE: Brazil. State Minas Gerais: Mun. Santana do Riacho, Serra do Cipó, Rodovia Belo Horizonte-Conceiçao do Mato Dentro, km 132, Semir & Sazima CESC 4758 (holotype, UEC not seen). 9. Chaptalia comptonioides Britton & P. Wilson, Mem. Torrey Bot. Club 16: 117. 1920. TYPE: Cuba. Prov. Oriente: Ensenada Mora, Britton et al. 12937 (isotype, MO!). 10. Chaptalia cordata Hieron., Bot. Jahrb. Syst. 21: 370. 1896. TYPE: Ecuador. Prov. Carchi, Páramo del Angel, near El Voladero, G. Harling & Anderson 12104 (neotype, designated by Harling, 1991: GB not seen, photo in Fl. Ecuador n. 42: 49, Fig. 7. 1991). Additional specimen examined: Peru. Dept. Amazonas: Prov. Bongará, along Yambrasbamba-Pomacocha trail, between Yanayacu and Pomacocha, J. J. Wurdack 1062 (LP). 11. Chaptalia cordifolia (Baker) Cabrera, Fl. Ilustr. Catarinense 1: 67. 1973. Basionym: Trichocline cordifolia Baker. TYPE: Brazil. Río de Janeiro, A. Glaziou 18320 (lectotype, designated by Cabrera 1973: 67, K not seen). Additional specimen examined: Brazil. State Santa Catarina: Campo dos Padres, Bom Retiro, P. Reitz 2531 (LP). 12. Chaptalia crassiuscula Urb., Repert. Spec. Nov. Regni Veg. 21: 228. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, E. L. Ekman 9127 (holotype, S not seen, S photo at LP!). 13. Chaptalia crispata Urb. & Ekman, Ark. Bot. 23: 97. 1930. TYPE: Haiti. Massif de la Selle: Marigot, Rivière Chotard, E. L. Ekman H 8004a (type, probably in HAC). 14. Chaptalia dentata (L.) Cass., Dict. sci. nat. 26: 104. 1823. Basionym: Tussilago dentata L. TYPE: Lectotype, designated by Nesom, 1984: 397, the illustration in Pl. Amer. 2: tab. 40, Fig. 77. 1755. Additional specimen examined: Bahamas. Andros Island: Conch Sound, J. Northrop & A. Northrop 400 (GH). 15. Chaptalia denticellata Urb. & Ekman, Ark. Bot. 23: 98. 1930. TYPE: Haiti. Massif de la Selle: Port au Prince, near Bassin-Laval, E. L. Ekman H 5493 (isotype, US!). 16. Chaptalia denticulata (Baker) Zardini, Darwiniana 19: 728. 1975. Basionym: Trichocline denticulata Baker. TYPE: Brazil. State Rio de Janeiro: ad fontem fluminis Soberbo, Glaziou 6031 (holotype, B not seen, B photo F n. 16015 at LP!). Additional specimen examined: Brazil. State Bahia: Pico das Almas, Río de Contas, SE de Campo do Queiroz, R. M. Harley et al. 26584 (US). 17. Chaptalia dolichopoda Urb. & Ekman, Ark. Bot. 23: 99. 1930. TYPE: Haiti. Massif de la Selle: Croix-des-Bousquets, Badeau, inter Morne Mérillon and Morne Badeau, E. L. Ekman H 7796a (isotype, GH!).


535

18. Chaptalia eggersii Urb., Symb. antill. 3: 418. 1903. TYPE: Dominican Republic. Santo Domingo: Valle Nuevo, Eggers 2220 (isotype, SI!). 19. Chaptalia ekmanii Urb., Repert. Spec. Nov. Regni Veg. 21: 227. 1925. TYPE: Cuba. Prov. Pinar del Río: Pinar de Cajálbana, E. L. Ekman 17340 (holotype, S not seen, photo LP!). 20. Chaptalia estribensis G. L. Nesom, Phytologia 78: 160. 1995. TYPE: Mexico. State Hidalgo: Municipio Tenango de Doria, 8 to 11 km SW of Tenango de Doria, D. E. Breedlove & Almeda 59571 (holotype, CAS not seen). Additional specimen examined: Mexico. State Hidalgo: El Cirio, al E de Tenango de Doria, Municipio Tenango de Doria, Torres C. & García 8237 (MEXU). 21. Chaptalia exscapa (Pers.) Baker, Fl. bras. 6: 379. 1884. Basionym: Tussilago exscapa Pers. TYPE: Uruguay. Montevideo, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Avellaneda, Wilde, A. Burkart 2191 (SI). 22. Chaptalia flavicans Urb. & Ekman, Ark. Bot. 23: 101. 1930. TYPE: Haiti. Massif de la Selle: Morne des Commissaires ad Grand Gosier, E. L. Ekman H 6903 (holotype, US!). 23. Chaptalia graminifolia (Dusén) Cabrera, Fl. Ilustr. Catarinense 1: 60. 1973. Basionym: Chaptalia piloselloides (Vahl) Baker var. graminifolia Dusén. TYPE: Brazil. State Paraná: entre Capão Grande y Villa Velha, P. Dusén 4062 (syntype, B not seen, B photo F n. 16003 at LP!). Additional specimen examined: Brazil. State Minas Gerais: Serra do Caparao, Pico Luez Inacio, Brade 16890 (LP). 24. Chaptalia hermogenis M. D. Moraes, Novon 8: 173. 1998. TYPE: Brazil. State São Paulo: Eldorado, Núcleo Caverna do Diablo, Parque Estadual de Jacupiranga, M. D. Moraes & F. Arzolla 367 (isotype, LP!). 25. Chaptalia hidalgoensis L. Cabrera & G. L. Nesom, Sida 20: 1363. 2003. TYPE: Mexico. State Hidalgo: Mun. Zimapan, Verdosas, V. M. Huerta 1352 (holotype, IEB not seen). 26. Chaptalia hololeuca Greene, Leafl. Bot. Observ. Crit. 1: 192. 1906. TYPE: Mexico. State Coahuila: and vicinity, Saltillo, E. Palmer 192 (holotype, US!). Additional specimen examined: Mexico. State Coahuila: Saltillo, Palmer 528 (GH). 27. Chaptalia ignota Burkart, Physis (Buenos Aires) 11: 102. 1932. TYPE: Argentina. Prov. Buenos Aires: San Isidro, A. Burkart 2153 (isotype, LP!). 28. Chaptalia incana Cuatrec., Proc. Biol. Soc. Wash. 74: 24. 1961. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, Hoya del Río Donachuí, Meollaca (or Meuyaca), J. Cuatrecasas & R. Romero Castañeda 24438 (holotype, US!). 29. Chaptalia integerrima (Vell.) Burkart, Darwiniana 6: 576. 1944. Basionym: Tussilago integerrima Vell. TYPE: Lectotype, selected by Burkart, 1944: 576, the illustration at Fl. flumin icon. 8: tab. 140. 1831 (1827). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Bolívar, Pirovano, A. Burkart 7064 (SI). 30. Chaptalia isernina Cuatrec., Anales Ci. Univ. Madrid 4: 212. 1935. TYPE: Peru. Prov. Chanchamayo: Chanchamayo, J. Isern 42 (holotype, F!). 31. Chaptalia latipes Urb. & Ekman, Ark. Bot. 23: 99. 1930. TYPE: Haiti. Massif du Nord: Gros-Morne, Belanse, E. L. Ekman H 4921 (holotype, S not seen, S photo at LP!).

536

L. Katinas, et al.

32. Chaptalia leiocarpa (DC.) Urb., Symb. antill. 8: 747. 1921. Basionym: Leria leiocarpa DC. TYPE: Cuba. s.d., R. de la Sagra s.n. (isotype, F!). 33. Chaptalia leptophylla Urb., Repert. Spec. Nov. Regni Veg. 21: 229. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, Río Jimbambay, E. L. Ekman 9559 (isotype, SI!). 34. Chaptalia lyratifolia Burkart, Darwiniana 6: 527. 1944. TYPE: Mexico. State Nuevo León: Sierra Madre, above Monterrey, C. G. Pringle 10217 (isotype, SI!). 35. Chaptalia madrensis G. L. Nesom, Phytologia 78: 158. 1995. TYPE: Mexico. Nuevo León: ca. 35 km ENE of Dr. Arroyo, NNW of Cerro Pena Nevada, G. Nesom 4758A (holotype, US!). 36. Chaptalia malcabalensis Cuatrec., Ciencia (México) 24: 123. 1965. TYPE: Peru. Dept. Amazonas: Prov. Chachapoyas, cerro Malcabal (Cerro Tumbe), 3– 6 km SW of Molino-pampa, J. J. Wurdack 141 (holotype, US!). 37. Chaptalia mandonii Sch. Bip. ex Burkart, Darwiniana 6: 551. 1944. TYPE: Bolivia. Larecaja, en la vecindad de Sorata, monte Chilica, Mandon 11 (isotype, GOET not seen). Additional specimen examined: Argentina. Prov. Catamarca: Dept. Andalgalá, Cuesta de Mina Capillitas, A. L. Cabrera et al. 24738 (LP). 38. Chaptalia media (Griseb.) Urb., Symb. antill. 3: 419. 1903. Basionym: Leria media Griseb. TYPE: Cuba. Cuba oriental C. Wright 332 (isotype, MO!). 39. Chaptalia membranacea Urb., Symb. antill. 3: 418. 1903. TYPE: Dominican Republic. Santo Domingo: Puerto Plata, Loma Isabel, Eggers 1582 (isotype, SI!). 40. Chaptalia meridensis S. F. Blake, Contr. U. S. Natl. Herb. 20: 540. 1924. TYPE: Venezuela. Mérida: Páramo del Molino, A. Jahn 954 (holotype, US!). 41. Chaptalia modesta Burkart, Physis (Buenos Aires) 11: 100. 1932. TYPE: Argentina. Prov. Salta: Pampa Grande, Jan 1897, C. L. Spegazzini s.n. (holotype, LP!). 42. Chaptalia montana Britton, Bull. Torrey Bot. Club 50: 51. 1923. TYPE: Cuba. Prov. Oriente: Sierra Maestra, Fr. Léon 10802 (type, probably in HAC). 43. Chaptalia mornicola Urb. & Ekman, Ark. Bot. 23: 97. 1930. TYPE: Haiti. Massif de la Selle: Nouvelle Touraine, Chapelle Faure, E. L. Ekman H 1413 (holotype, S not seen, S photo at LP!). 44. Chaptalia nipensis Urb., Repert. Spec. Nov. Regni Veg. 21: 229. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, E. L. Ekman 3094 (holotype, S not seen, S photo at LP!). 45. Chaptalia nutans (L.) Pol., Linnaea 41: 582. 1877. Basionym: Tussilago nutans L. TYPE: America: s.d., s. leg. (lectotype, designated by Nesom, 1995: 162, LINN n. 995.5 not seen, photo at LP!). Additional specimen examined: Brazil. State São Paulo: Campos de Butantã, Joly 145AB3 (LP*). 46. Chaptalia oblonga D. Don, Trans. Linn. Soc. London 16: 240. 1830. TYPE: Peru. s.d., H. Ruiz & J. A. Pavón s.n. (holotype, MA not seen, MA photo at LP!). 47. Chaptalia obovata C. Wright, Anales Acad. Ci. Méd. Habana 6: 212. 1869. TYPE: Cuba. s.d., C. Wright 3617 (isotype, US!). 48. Chaptalia ovalis D. Don, Trans. Linn. Soc. London 16: 241. 1830. TYPE: Peru. s.d., H. Pavón s.n. (holotype, MA not seen, MA photo at LP!).


537

49. Chaptalia paramensis Cuatrec., Proc. Biol. Soc. Wash. 74: 25. 1961. TYPE: Colombia. Dept. Magdalena: Sierra de Perijá, between Cerro Venado and Cerro Avión, J. Cuatrecasas & R. Romero Castañeda 25117 (holotype, US!). 50. Chaptalia piloselloides (Vahl) Baker, Fl. bras. 6: 378. 1884. Basionym: Perdicium piloselloides Vahl. TYPE: Uruguay. Montevideo, s.d., Commerson s.n. (type, not localized). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Avellaneda, Sarandí, A. Burkart 2208 (SI). 51. Chaptalia primulacea Greene, Leafl. Bot. Observ. Crit. 1: 195. 1906. TYPE: Dominican Republic. Santo Domingo, C. Wright et al. 261 (isotype, US!). 52. Chaptalia pringlei Greene, Leafl. Bot. Observ. Crit. 1: 192. 1906. TYPE: Mexico. State Oaxaca: above Dominguillo, C. G. Pringle 5796 (lectotype, designated by Nesom, 1995: 161, US!). Additional specimen examined: Mexico. State Oaxaca: ca. 4 km W of Tamazulapan on road to Chilapa de Diaz, G. Nesom 4405 (MO). 53. Chaptalia pumila (Sw.) Urb., Symb. antill. 3: 420. 1903. Basionym: Tussilago pumila Sw. TYPE: Jamaica. s.d., A. Swartz s.n. (holotype, M!). 54. Chaptalia rocana Britton & P. Wilson, Mem. Torrey Bot. Club 16: 118. 1920. TYPE: Cuba. Mtns. of southern Santa Clara, Río Caracusey, Banao mtns., Léon & Roca 7904 (type, probably in HAC). 55. Chaptalia rotundifolia D. Don, Trans. Linn. Soc. London 16: 242. 1830. TYPE: Peru. Pillao, s.d., H. Ruiz & J. A. Pavón s.n. (holotype, MA not seen). 56. Chaptalia runcinata Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 5. 1818. TYPE: Colombia: F. Humboldt & A. Bonpland 2031 (lectotype, designated by Nesom, 1995: 168, P not seen, P photo F n. 38167 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido La Plata, alrededores de La Plata, Elizalde, A. L. Cabrera 907 (SI). 57. Chaptalia shaferi Britton & P. Wilson, Mem. Torrey Bot. Club 16: 118. 1920. TYPE: Cuba. Prov. Oriente: valley of Río Yamaniguey, J. A. Shafer 4203 (holotype, GH!). 58. Chaptalia similis R. E. Fr., Nova Acta Regiae Soc. Sci. Upsal. 1: 95. 1905. TYPE: Argentina. Prov. Jujuy: Santa Catalina, Kurtz 11460 (syntype, BAF not seen). Additional specimen examined: Argentina. Prov. Catamarca: Dept. Andalgalá, cuesta de Mina Capillitas, A. L. Cabrera et al. 24738 (LP). 59. Chaptalia sinuata (Less.) Baker, Fl. bras. 6: 378. 1884. Basionym: Leria nutans (L.) DC. var. sinuata Less. TYPE: Uruguay. Montevideo, Sellow 1413, 2212 (syntypes, B not seen, B photo F n. 16006 at LP!). Additional specimen examined: Argentina. Prov. Misiones: Dept. Candelaria, entre Candelaria y Santa Ana, 20 Nov 1896, N. Alboff s.n. (LP). 60. Chaptalia spathulata (D. Don) Hemsl., Biol. Cent.-Amer., Bot. 2: 255. 1881– 1882. Basionym: Leria spathulata D. Don. TYPE: Mexico. State Guerrero: Mazatlán, M. Sessé & J. M. Moçiño 2028 (holotype, MA not seen). 61. Chaptalia stenocephala (Griseb.) Urb., Symb. antill. 3: 420. 1903. Basionym: Leria stenocephala Griseb. TYPE: Cuba. s.d., C. Wright 2874 (isotype, MO!). 62. Chaptalia stuebelii Hieron., Bot. Jahrb. Syst. 21: 371. 1896. TYPE: Ecuador. Prov. Pichincha: Cerro Pichincha, Holmgren 297 (neotype, designated by Harling, 1991: 53, S not seen). Additional specimen examined: Ecuador. Prov.

538

63.

64.

65.

66.

67.

68.

L. Katinas, et al.

Azuay: valley of the Río Paute, between Paute and Cuenca, Parroquia José Víctor Izquierdo at “Pupazche,” Camp 2573 (US). Chaptalia texana Greene, Leafl. Bot. Observ. Crit. 1: 191. 1906. TYPE: United States. State Texas: W Texas, G. C. Neally 297 (holotype, US!). Additional specimen examined: Mexico. State Baja California: La Champagna, Sierra de Las Palmas, S of Sta. Rosalía, Gentry & Fox 11768 (MEXU). Chaptalia tomentosa Vent., Descr. pl. nouv. 7: tab. 61. 1802. TYPE: Walter, Carol. Bosc (holotype G-DC, microfiche 1238, frame 15!). Additional specimen examined: United States. State Alabama: Spring Hill, Graves 1950 (MO). Chaptalia transiliens G. L. Nesom, Rhodora 86: 127. 1984. TYPE: Mexico. Nuevo León: ca. 35 km ENE of Dr. Arroyo, NNW of Cerro Pena Nevada, G. Nesom 4759 (holotype, US!). Chaptalia turquinensis Borhidi & O. Muñiz, Acta Bot. Acad. Sci. Hung. 17: 35. 1971. TYPE: Cuba. Prov. Oriente: Sierra Maestra, Pico Turquino, Paso de los Angustios, V. Samek 815 (type, probably in HAC). Chaptalia undulata Urb. & Ekman, Ark. Bot. 23: 100. 1930. TYPE: Haiti. Massif du Nord: St. Louis du Nord, Morne Baron, s. leg. H 4696 (type, probably in HAC). Chaptalia vegaensis Urb. & Ekman, Ark. Bot. 23: 96. 1930. TYPE: Dominican Republic. Santo Domingo: Prov. La Vega, Bonao, Loma Peguera, E. L. Ekman H 11480 (holotype, S not seen, S photo at LP!).

Observation. A revision in progress by L. Katinas shows that many of the species under the name Chaptalia (e.g., C. anisobasis, C. estribensis, C. meridensis) deviate from the generic concept mainly by the presence of staminodes. Thus, they should be excluded from the genus and moved to Gerbera. Other species, such as many of those of the Antilles (e.g., C. crispata, C. dolichopoda, C. membranacea), are variants of one morphologically heterogeneous species and their names should be treated as synonyms. As a result, the probable number of species of the genus will be 20–35 (Katinas, 2004a). Literature. Burkart (1944), Nesom (1995).

Chucoa Cabrera Bol. Soc. Argent. Bot. 6: 40. 1955. TYPE: Chucoa ilicifolia Cabrera. 12 of Figs. 11–14. Weberbaueriella Ferreyra, Bol. Soc. Peruana Bot. 5: 2. 1955. TYPE: Weberbaueriella johnstoniana Ferreyra [= Chucoa ilicifolia Cabrera]. Etymology. From the type locality in the province of Santiago Chuco, Peru. Shrubs. Leaves alternate; sessile to subsessile; blades lanceolate to obovate, pinnately veined, margin denticulate to spinose, lanuginous to glabrous above, lanose beneath. Capitulescences monocephalous or two- to three-headed, terminal; capitula pedunculate, homogamous or occasionally heterogamous, discoid; recepta-


539

cle epaleate; involucre multiseriate. Florets isomorphic, bisexual, rarely the marginal ones female with staminodes, corolla actinomorphic, tubular-funnelform to subcampanulate, shallowly to deeply five-lobed; anther apical appendages acute, tails smooth to papillose, filaments smooth to papillose at the base; style bifid to bilobed, branches dorsally papillose. Cypselae glabrous to subglabrous; pappus of scabrid bristles, all capillary and equal in length or the innermost longer and somewhat flattened at the base. Pollen Description. Chucoa ilicifolia (87 of Figs. 86–91). Pollen subprolate to prolate, elliptic in equatorial view, subtriangular in polar view, large size, P × E = (70–85×72– 69) μm, tricolporate, colpi long, with margins and psilate membrane, mesoaperturate. Exine Mutisia type, microechinate, 10–13.5 μm thick at the equator, slender at the poles, ectosexine compact. Ratio ectosexine∕endosexine: 1: 1. Nexine 2–2.5 μm thick. Habitat and Distribution. Genus of two species endemic to rocky dry slopes of northwestern Peru. Species list 1. Chucoa ilicifolia Cabrera, Bol. Soc. Argent. Bot. 6: 42. 1955. TYPE: Peru. Dept. La Libertad: Prov. Santiago Chuco, Angasmarca-Tulpo, A. López Miranda 1090 (holotype, LP!*). 2. Chucoa lanceolata (H. Beltrán & Ferreyra) G. Sancho, S. E. Freire & Katinas, Taxon 54: 86. 2005. Basionym: Gochnatia lanceolata H. Beltrán & Ferreyra. TYPE: Peru. Dept. Arequipa: Prov. Castilla, Andagua, Carolina 1 (holotype, USM not seen; isotype, US!). Observation. There are some problems in trying to sort out priority between the names Chucoa and Weberbaueriella since both names were published in the same year, 1955, and date of priority has not been conclusively established. After first published, Weberbaueriella has not appeared in any other modern literature and Chucoa is the name most commonly used for the taxon. If Weberbauerella Ulbrich, a genus of Fabaceae established in 1906, and Weberbaueriella Ferreyra were ruled out to be treated as homonyms, it would establish Chucoa as unquestionably the correct name for the Asteraceae genus (Brummitt, 2005). The current recommendation of the General Committee, following the opinion of the Committee for Spermatophyta, is that Weberbauerella (Fabaceae) and Weberbaueriella (Asteraceae) are treated as confusable names (Barrie, 2006). Literature. Ferreyra (1995), Sancho et al. (2005). Cnicothamnus Griseb. Abh. Königl. Ges. Wiss. Göttingen 19: 196. 1874. TYPE: Cnicothamnus lorentzii Griseb. 13 of Figs. 11–14. Lefrovia Franch., J. Bot. (Morot) 2: 377. 1888. TYPE: Lefrovia rhaponticoides Franch. [= Cnicothamnus lorentzii Griseb.].

540

L. Katinas, et al.

Etymology. From the Greek knecos, yellow, yellowish, a name applied to Carthamus tinctorius with deep orange flowers, and thamnos, shrub, by the resemblance of the flower color of both species. Shrubs to small trees. Leaves alternate; short-petiolate; blades wide-elliptic to ovate, pinnately veined, margin denticulate to dentate, glabrous above and floccosepubescent beneath. Capitulescences monocephalous, terminal; capitula sessile to short-pedunculate, homogamous, radiate; receptacle epaleate; involucre multiseriate, six- to ten-seriate, phyllaries acute or with showy, orbicular to spatulate, laciniate apical appendages. Florets dimorphic, bisexual; marginal florets with corolla subbilabiate (3+1 corolla lips), outer lip expanded; central florets with corolla tubularfunnelform, deeply five-lobed; anther apical appendages apiculate, tails smooth; style bilobed, branches dorsally smooth. Cypselae densely villose; pappus of scabrid bristles, half capillary and short and half somewhat paleaceous, clavate at the apex and longer. Pollen Description. Cnicothamnus lorentzii (88 of Figs. 86–91). Pollen subprolate to prolate, elliptic in equatorial view, circular in polar view, large size, P × E = (66– 76×48–54) μm. Tricolporate, colpi long, with margin and scabrate membrane, mesoaperture diffuse? Exine Mutisia type, echinate, 8–11 μm thick, spines of 2– 2.5 μm length. Ratio ectosexine/endosexine: 1.5:1; 1:1. Nexine ca. 2 μm thick. Note: pollen similar to that of type II of Gochnatia. Habitat and Distribution. Genus of two species from the subtropical forests of Bolivia and northwestern Argentina. Species list 1. Cnicothamnus azafran (Cabrera) Cabrera, Notas Mus. La Plata, Bot. 9: 256. 1944. Basionym: Cnicothamnus lorentzii Griseb. var. azafran Cabrera. TYPE: Argentina. Prov. Salta: Río Maíz Gordo, A. Ragonese 334 (holotype, LP!). 2. Cnicothamnus lorentzii Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 197. 1874. TYPE: Argentina. Prov. Tucumán, P. G. Lorentz 182 (isotype, CORD not seen). Additional specimens examined: Argentina. Prov. Jujuy: Ledesma, Sierra de Calilegua, S. Venturi 5310 (LP); Prov. Salta, 50 km SE from Salta city, S. Maldonado 408 (LP*).

Literature. Cabrera (1978), Freire (1996), Freire et al. (2002).

Cyclolepis D. Don Philos. Mag. Ann. Chem. 11: 392. 1832. TYPE: Cyclolepis genistoides D. Don. 14 of Figs. 11–14. Cyclopis Gillies, Arch. Bot. (Paris) 2: 468. 1833, an ortographic variant of the name Cyclolepis.


541

Etymology. From the Greek kyclos, circle, ring, and lepis, scales, by the imbricate disposition of the phyllaries. Shrubs branched, gynodioecious, short branches modified in thorns. Leaves alternate, caducous; short-petiolate; blades elliptic, pinnately veined, margin entire, pubescent on both surfaces. Capitulescences racemose to spicate; capitula subsessile, homogamous, discoid; receptacle epaleate; involucre three- to sixseriate. Florets isomorphic in bisexual capitula, corolla actinomorphic, tubularfunnelform, deeply five-lobed, lobes coiled; anther apical appendages apiculate, tails slightly papillose; style bifid, branches dorsally smooth; female capitula with florets without staminodes, corolla tubular-filiform, shallowly five-lobed, tube long, up to 2/3 of corolla length, lobes straight. Cypselae villose; pappus of scabrid bristles, half capillary and short and half somewhat paleaceous, plumose at the apex and longer. Pollen Description (Tellería & Forcone, 2002) (89 of Figs. 86–91). Pollen spheroidal to subprolate, elliptic to spheroidal in equatorial view, circular in polar view, medium to large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Monotypic genus that grows in salty soils from Paraguay to northern Patagonia in Argentina. Species list 1. Cyclolepis genistoides D. Don, Philos. Mag. Ann. Chem. 11: 392. 1832. TYPE: Sine loc., s. d., J. Gillies s.n. (holotype, probably K not seen). Additional specimens examined: Argentina. Prov. Salta: Quebrada de las Conchas, A. L. Cabrera 3782 (LP); Prov. Chubut: Trelew, A. Forcone 459 (CORD*). Literature. Cabrera (1951, 1971b), Freire et al. (2002).

Dinoseris Griseb. Symb. fl. argent.: 213. 1879. TYPE: Dinoseris salicifolia Griseb. 15 of Figs. 15–18. Etymology. From the Greek deinos, terrible, powerful, mighty, and seris, chicory, and lettuce. Shrubs to little trees. Leaves opposite; petiolate; blades elliptic to lanceolate, pinnately veined, margin entire to serrulate, glabrous. Capitulescences monocephalous or two- to three-headed, terminal; capitula pedunculate, homogamous, ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb shallowly five-dentate; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae truncate to slightly attenuate at the apex, glabrous; pappus of scabrid, capillary bristles.

542

L. Katinas, et al.

Figs. 15–18 15 Dinoseris salicifolia Griseb. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Cabrera, 1978: 610). Scale bars: A=7 cm; B, E=15 mm; C= 10 mm; D=3.5 mm. 16 Duidaea rubriceps S. F. Blake. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (A redrawn from Pruski, 1997: 262; B–E redrawn from Gleason, 1931: 498). Scale bars: A=3.5 cm; B, E=6 mm; C=7.6 mm; D=2.5 mm. 17 Eriachaenium magellanicum Sch. Bip. A Habit. B Marginal corolla. C Staminode. D Marginal cypsela. E Disc corolla. F Stamen. G Upper part of the style. H Central cypsela. (Redrawn from Katinas, 2000: 232). Scale bars: A=5 cm; B–F =2 mm; G=0.5 mm. 18 Eurydochus bracteatus Maguire & Wurdack. A Habit (branch). B Corolla. C Stamen. D Stamen apex curved. E Upper part of the style. F Cypsela with pappus. (Redrawn from Maguire & Wurdack, 1958: 57). Scale bars: A=5.4 cm; B=7 mm; C=2.5 mm; D=2.5 mm; E=2 mm; F= 10 mm

Pollen Description (Tellería & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Monotypic genus that grows on the slopes in the subtropical forests of Bolivia and northern Argentina.


543

Species list 1. Dinoseris salicifolia Griseb., Symb. fl. argent.: 214. 1879. TYPE: Argentina. Prov. Tucumán: Sierra de Tucumán, Suciliaco, P. G. Lorentz & G. H. E. W. Hieronymus 862 (isosyntype, CORD not seen). Additional specimen examined: Argentina. Prov. Salta: Campo Quijano, July 1954, Corte s.n. (LP). Observation. Some authors (e.g., Ariza Espinar, 1973; Bremer, 1994) consider the only species of Dinoseris (D. salicifolia) a species of Hyaloseris. However, other authors (e.g., Cabrera, 1978; Katinas, 1996a) prefer to maintain both genera as separate. Some characters that distinguish both taxa are: brachyblasts and three- to six-flowered capitula in Hyaloseris versus lack of brachyblasts and capitula with more than 20 florets in Dinoseris. Literature. Ariza Espinar (1973), Cabrera (1978), Katinas (1996a).

Duidaea S. F. Blake Bull. Torrey Bot. Club 58: 496. 1931. TYPE: Duidaea tatei S. F. Blake. 16 of Figs. 15–18. Etymology. Named for Mount Duida (state Amazonas, Venezuela), which towers above Rio Orinoco at La Esmeralda. Shrubs. Leaves alternate, clustered at the stem apex; sessile to subsessile, the petioles, when present, slightly clasping; blades linear to oblanceolate, subcoriaceous to coriaceous, one-veined, margin entire, revolute, glabrous to densely pubescent beneath. Capitulescences monocephalous, rarely several heads per branch, either axillar or seemingly terminal; capitula short- to longpedunculate, pendulous, homogamous, bilabiate; receptacle epaleate; involucre multiseriate, phyllaries reddish. Florets isomorphic, bisexual, corolla bilabiate; anthers commonly light purple, generally exserted, apical appendages acuminate, tails papillose; style deep purple, bifid, branches dorsally smooth. Cypselae glabrous to densely pubescent; pappus of scabrid bristles, the outermost shorter. Pollen Description. Duidaea marahuacensis (90 of Figs. 86–91). Pollen spheroidal– subprolate, spheroidal or elliptic in equatorial view, circular in polar view, large size, P × E = (62–72×52–64) μm. Tricolporate, colpi long, with margin and scabrate membrane. Exine Gongylolepis type, 11–14 μm thick in the whole grain, spines 2.5 μm length. Ratio ectosexine/endosexine among spines: 1:3; 1:4. SEM: tectum very perforate. Note: nexine scarcely distinguishable. Pollen similar to that of Glossarion rhodanthum and Quelchia bracteata (see also Carlquist, 1957b; Tellería, 2008). Habitat and Distribution. Duidaea is endemic to two isolated tepuis in Amazonas, Venezuela and contains four species.

544

L. Katinas, et al.

Species list 1. Duidaea marahuacensis Steyerm., Brittonia 30: 50. 1978. TYPE: Additional specimens examined: Venezuela. Terr. Fed. Amazonas: Cerro Marahuaca, S. S. Tillet et al. 752.348 (holotype, VEN!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dtpo. Atabapo, Cerro Marahuaca, R. Liesner 24644 (MO, US*). 2. Duidaea pinifolia S. F. Blake, Bull. Torrey Bot. Club 58: 499. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Summit of Mt. Duida, G. H. H. Tate 530 (isotype, US!). 3. Duidaea rubriceps S. F. Blake, Bull. Torrey Bot. Club 58: 500. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida: Brocchinia Hills, G. H. H. Tate 589 (isotype, US!). 4. Duidaea tatei S. F. Blake. Bull, Torrey Bot. Club. 58: 498. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida: Gorge of Caño Negro, Savanna Hills, G. H. H. Tate 814 (isotype, US!). Literature. Maguire et al. (1957a), Aristeguieta (1964), Pruski (1997).

Eriachaenium Sch. Bip. Flora 38: 120. 1855. TYPE: Eriachaenium magellanicum Sch. Bip. 17 of Figs. 15–18. Etymology. From the Greek erion, wool, and from the Latin achaenium, a type of fruit, by the villose fruits. Herbs perennial, dwarf, with stout, oblique to vertical rhizomes, stems prostrate to ascending. Leaves alternate; sessile, clasping; blades oblanceolate, pinnately veined, margin entire to undulate-dentate, glabrous to subglabrous above, tomentose beneath. Capitulescences monocephalous, axillar; capitula pedunculate, heterogamous, disciform; receptacle epaleate; involucre uniseriate. Florets dimorphic; marginal florets female with staminodes, corolla tubular-funnelform, deeply fourlobed; central florets bisexual or male with a rudimentary ovary, corolla tubularfunnelform, deeply five-lobed; anther apical appendages rounded to acute, basally constricted and demarcated from the thecae, basally auriculate with tails very short, smooth to slightly papillose; style bilobed, dorsally papillose. Cypselae truncate at the apex, shaggy, dimorphic, the marginal cypselae conspicuously bigger than the central ones; pappus absent. Pollen Description (91 of Figs. 86–91). Pollen spheroidal to prolate, spheroidal or elliptic in equatorial view, circular in polar view, medium size, P × E = (30–36×24– 30) μm. Tricolporate, colpi long with thin margin and microgranulate membrane, mesoaperture diffuse. Exine Mutisia type, microechinate, 2–6 μm thick, slightly slender at the poles. Ratio ectosexine/endosexine: 1:1.5; 1:2. Nexine 1.5 μm thick. SEM: tectum punctate. Note: grains with pyramidal or rectangular shape and grains biaperturate were observed. Some grains have the endosexine with compact aspect,


545

as occurs in Adenocaulon chilense (Mutisieae). Pollen similar to that of Adenocaulon and Artemisia verlotorum (Anthemideae). Habitat and Distribution. Monotypic genus endemic to Patagonia in Argentina and Chile. Species list 1. Eriachaenium magellanicum Sch. Bip., Flora 38: 121. 1855. TYPE: Chile. Prov. Magallanes: Oazy Harbour, Lechler 1256 (type, probably SGO). Additional specimens examined: Argentina. Prov. Santa Cruz: Jan 1903, C. L. Spegazzini s.n. (LP*); Río Gallegos, O’Donell 4091 (LP*); id., Estancia Guer Aike, H. Sleumer 908 (LP*). Literature. Cabrera (1971b), Robinson and Brettell (1973), Moore (1983), Katinas (2000).

Eurydochus Maguire & Wurdack Bol. Soc. Venez. Ci. Nat. 20: 57. 1958. TYPE: Eurydochus bracteatus Maguire & Wurdack. 18 of Figs. 15–18. Etymology. Derived from the Greek eurys, broad, large, and dochos, a receptacle, dechomai, to receive, in reference to the broad receptacles. Trees or shrubs. Leaves alternate, often clustered apically; petiolate, the petioles clasping; blades elliptic to oblanceolate, chartaceous to sub-coriaceous, pinnately veined, margin entire, glabrous. Capitulescences monocephalous, heads subterminal; capitula pedunculate, homogamous, bilabiate, six- to many-flowered; receptacle epaleate; involucre multiseriate, hemispherical, phyllaries subequal, the outer phyllaries commonly decurrent onto the peduncle. Florets isomorphic, bisexual, corolla bilabiate, outer lip expanded; anthers brown to purple, exserted, apical appendages acute, slightly coiled at the apex, tails papillose, connate; style rosecolored, bifid, branches dorsally smooth. Cypselae puberulent; pappus of fragile, scabrid, capillary bristles, sometimes individually deciduous, the outermost shorter. Pollen Description. Pollen prolate, elliptic in equatorial view, circular in polar view, large size, P × E = (72–80×54–60) μm. Tricolporate, long colpi with microgranulate membrane. Exine Wunderlichia type, 8–10 μm thick in the whole grain; spines 3 μm length. Ratio ectosexine/endosexine among spines: 1:1; 1.5:1; 1:1.5. Nexine 1.5 μm thick. Note: pollen grains similar to those of Salcedoa mirabaliarum F. Jiménez Rodr. & Katinas and Wunderlichia azulensis and W. mirabilis Riedel ex Baker (see also Jiménez Rodríguez et al., 2004; Tellería, 2008). Habitat and Distribution. Eurydochus is a poorly known monotypic genus known only from two collections in the western Guayana Highland. It occurs on Cerro de la

546

L. Katinas, et al.

Neblina on the border of Venezuela and Brazil and on nearby Serra Pirapucu in Brazil. Species list 1. Eurydochus bracteatus Maguire & Wurdack, Bol. Soc. Venez. Ci. Nat. 20: 57. 1958. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 42186 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Río Yatua, Cañón Grande, B. Maguire et al. 42238 (US*). Literature. Maguire and Wurdack (1958), Aristeguieta (1964), Pruski (1991, 1997).

Gerbera L., nom. cons. Sp. pl.: 885. 1753. TYPE: Arnica gerbera L. [= Gerbera linnaei Cass. (type cons.)]. 19 of Figs. 19–22. Aphyllocaulon Lag., Amen. nat. Españ.: 38. 1811, species name not designated. Atasites Neck., Elem. bot. 1: 7. 1790, nom. illeg. Berniera DC., Prodr. 7: 18. 1838, nom. illeg. TYPE: Berniera nepalensis DC. [= Gerbera maxima (D. Don) Beauverd]. Epiclinastrum Bojer ex DC., Prodr. 7: 18. 1838. TYPE: Epiclinastrum candollei Bojer ex DC. [= Gerbera piloselloides (L.) Cass.]. Gerberia Cass., Bull. Sci. Soc. Philom. Paris: 34. 1817, an orthographic variant of the name Gerbera. Lasiopus Cass., Bull. Sci. Soc. Philom. Paris: 152. 1817, non D. Don, 1836. [= Gerbera sect. Lasiopus (Cass.) Sch. Bip., Flora 27: 780. 1844]. TYPE: Lasiopus ambiguus Cass. [= Gerbera ambigua (Cass.) Sch. Bip.]. Leptica E. Mey. ex DC., Prodr. 7: 16. 1838. TYPE: Leptica ciliata E. Mey ex DC. [= Gerbera piloselloides (L.) Cass.]. Oreoseris Wall. ex DC., Prodr. 7: 17. 1838. TYPE: Oreoseris lanuginosa Wall. ex DC. [= Gerbera gossypina (Royle) Beauverd]. Etymology. In honor of the eighteenth century German naturalist Traugott Gerber (d. 1743), physician, traveler, and plant collector. Herbs perennial, acaulescent, scapose. Leaves rosulate; sessile, petiolate to pseudopetiolate; blades linear-oblanceolate to orbicular, pinnately veined, margin entire to deeply pinnatisect, densely pubescent on both surfaces or only beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate or rarely disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic to trimorphic; marginal florets female generally with filiform to well developed staminodes, corolla bilabiate with an expanded outer lip, rarely minute; intermediate florets (when present) female with or without staminodes, corolla bilabiate, rarely filiformbilabiate, as long as the style or longer; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bilobed to bifid, dorsally


547

Figs. 19–22 19 Gerbera crocea (L.) Kuntze. A Habit. Gerbera jamesonii Bolus ex Adlam. B Outermost marginal corolla. C Staminode of marginal floret. D Innermost marginal corolla. E Central corolla. F Stamen. G Upper part of the style. H Cypsela with pappus. (A from Williams 3704, WIS; B–H from Hu 6385, US). Scale bars: A=5 cm; B=5 mm; C–F, H=2 mm; G=0.3 mm. 20 Glossarion rhodanthum Maguire & Wurdack. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Aristeguieta, 1964: 916). Scale bars: A=4 cm; B=6 mm; C=6.7 mm; D=3 mm. 21 Gochnatia discolor Baker. A Habit (branch). B Corolla. C Staminode. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Sancho, 2000: 89). Scale bars: A=5 cm; B, F=4 mm; C, D= 2 mm; E=0.8 mm. 22 Gongylolepis paniculata Maguire & K. D. Phelps. A Habit (branch). G. benthamiana R. H. Schomb. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (A redrawn from Pruski, 1997: 291; B–E from Anderson 13840, MO). Scale bars: A=3.5 cm; B=6 mm; C= 5 mm; D=2 mm; E=12 mm

papillose. Cypselae rostrate to long-attenuate at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Gerbera hieracioides (Kunth) Zardini (92 of Figs. 92–94) (Tellería & Katinas, 2004). Pollen subprolate to spheroidal, large size, tricolporate,

548

L. Katinas, et al.

exine Mutisia type, slightly microechinate (see also Southworth, 1983; Lin et al., 2005). Habitat and Distribution. Genus of 29 species distributed in Africa, southern Asia, and America. Species list 1. Gerbera ambigua (Cass.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus ambiguus Cass. TYPE: Africa. South Africa: Cape Sonnerat, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Africa. South Africa: Natal, NW Pietermaritzburg, N side of Old Howick road, E of Ling road, T. Mc Farland 128 (WIS). 2. Gerbera aurantiaca Sch. Bip., Flora 27: 781. 1844. TYPE: Africa. South Africa: Natal, Pietermaritzburg, Krauss 388 (holotype, P? not seen; isotype, BM not seen). Additional specimen examined: Africa. South Africa: Natal, Old Howick road, E of Link road, about 4 km NW of Pietermaritzburg business district, T. Mc Farland 265 (WIS). 3. Gerbera bojeri (DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus bojeri DC. TYPE: Africa. Madagascar: s.d., Bojer s.n. (holotype, G-DC not seen). Additional specimen examined: Africa. Madagascar: vicinity of Station Forestiere Antsampandrano, T. Croat 29094 (MO). 4. Gerbera cordata (Thunb.) Less., Linnaea 5: 297. 1830. Basionym: Arnica cordata Thunb. TYPE: Africa. South Africa: Cape province, Capb. Spei, s.d., s. leg. herb. Thunberg 19988 (paratype, UPS not seen). Additional specimen examined: Africa. South Africa: W Cape province, Keurbooms River Forest Res., Willosoman, Dahlstrand 9528 (MO). 5. Gerbera crocea (L.) Kuntze, Revis. gen. pl. 3: 149. 1898. Basionym: Arnica crocea L. TYPE: Sine loc., s.d., s. leg. (holotype, LINN 1001, n. 7 not seen). Additional specimens examined: Africa. South Africa Cape, Hermanus, F. Rogers 26453 (US); Vogelgat, W of base camp, I. Williams 3704 (WIS). 6. Gerbera delavayi Franch., J. Bot. (Morot) 2: 68. 1888. TYPE: China. Yunnan: Choui-tsin-yn, J. M. Delavay 1918 (isotype, GH!). 7. Gerbera diversifolia Humbert, Mem. Soc. Linn. Normandie 25: 151, 307. 1923. TYPE: Africa. Madagascar: Tampoketsa d’Analamaitso, haut bassin du Bemarivo, Perrier de la Bâthie 3230 (holotype, P not seen, P photo at LP!). 8. Gerbera elliptica Humbert, Mem. Soc. Linn. Normandie 25: 154, 307. 1923. TYPE: Africa. Madagascar: Perrier de la Bâthie 3190 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!). Additional specimen examined: Africa. Madagascar: Massif de l’Andringitra, Iratsy, vallées de la Riambava et de la Antsifotra, H. Humbert 3654 (GH). 9. Gerbera emirnensis Baker, J. Linn. Soc., Bot. 22: 498. 1887. TYPE: Africa. Madagascar: Central Madagascar, Baron 3457 (lectotype, designated by Hansen, 1985b: 452, K not seen, K photo at LP!). 10. Gerbera galpinii Klatt, Bull. Herb. Boissier 4: 844. 1896. TYPE: Africa. South Africa: Transvaal, Barberton, Umotali valley, E. E. Galpin 1132 (isotype, K not seen, K photo at LP!).


549

11. Gerbera gossypina (Royle) Beauverd, Bull. Soc. Bot. Genève 2: 40. 1910. Basionym: Chaptalia gossypina Royle. TYPE: It is found at Mussooree, and similar elevations throughout Gurhual and Simore (type not localized). Additional specimen examined: India. Landour, Mussoorie, April 1929, F. Cleland s.n. (US). 12. Gerbera henryi Dunn, J. Linn. Soc., Bot. 35: 511. 1903. TYPE: China. Yunnan: Mentzge, A. Herny 9111 (isotype, US not seen, US photo at LP!). 13. Gerbera hieracioides (Kunth) Zardini, Bol. Soc. Argent. Bot. 16: 105. 1974. Basionym: Onoseris hieracioides Kunth. TYPE: Ecuador. Prov. Chimborazo: Alausi, Sitzan, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Peru. Dept. Cajamarca: Prov. Contumasá, Ditstrito Guzmango, Herilla, A. Sagástegui 106 (LP*). 14. Gerbera hintonii (Bullock) Katinas, Novon 8: 383. 1998. Basionym: Chaptalia hintoni Bullock. TYPE: Mexico. Mexico: District Temascaltepec, Nanchititla, G. B. Hinton 3098 (isotype, MO!). 15. Gerbera hypochaeridioides Baker, J. Linn. Soc., Bot. 20: 193. 1883. TYPE: Africa. Madagascar: Central Madagascar, Baron 2088 (lectotype, designated by Hansen, 1985b: 452, K not seen, K photo at LP!). 16. Gerbera jamesonii Bolus ex Adlam, Gard. Chron. 3: 775. 1888. TYPE: Africa. South Africa: Natal, s.d., Adlam s.n. (lectotype, designated by Hansen, 1985a: 21, SAM not seen). Additional specimens examined: Africa. South Africa: Transvaal, 2 Oct 1925, E. M. Young s.n. (WIS); Hong Kong, Chung Chi College, S. Y. Hu 6385 (US). 17. Gerbera leandrii Humbert, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 197. 1955. TYPE: Sine loc., H. Humbert 23599 (holotype, P not seen, P photo at LP!). 18. Gerbera linnaei Cass., Dict. sci. nat. 18: 460. 1821, nom. novum. Basionym: Arnica gerbera L. TYPE: Sine loc., s.d., s. leg. (lectotype, designated by Hansen, 1985a: 9, LINN 1001 not seen). Additional specimen examined: Africa, South Africa: C. Peninsula, Kogelfontein, T. Salter 8532B (MO). 19. Gerbera maxima (D. Don) Beauverd, Bull. Soc. Bot. Genève 2: 44. 1910. Basionym: Chaptalia maxima D. Don. TYPE: Nepal, Narainhetty, Hamilton & Wallich s.n. (isosyntypes, K not seen). 20. Gerbera nivea (Wall. ex DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Oreoseris nivea Wall. ex DC. TYPE: Nepal. Gossin-Than, Wallich 3143/253 (lectotype, designated by Hansen, 1988: 66, K not seen). Additional specimen examined: China. Yunnan: between Li-Kang and Joung-Ning, J. Rock 17333 (US). 21. Gerbera parva N. E. Br., Bull. Misc. Inform. Kew: 27. 1895. TYPE: Africa. South Africa: Natal, Drakensberg, near Bushman’s river, Evans 57 (lectotype, designated by Hansen, 1985a: 16, K not seen, K photo at LP!). 22. Gerbera perrieri Humbert, Mem. Soc. Linn. Normandie 25: 151, 307. 1923. TYPE: Sine loc., Perrier de la Bâthie 3219 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!). 23. Gerbera petasitifolia Humbert, Mem. Soc. Linn. Normandie 25: 152, 307. 1923. TYPE: Sine loc., Perrier de la Bâthie 2877 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!).

550

L. Katinas, et al.

24. Gerbera piloselloides (L.) Cass., Dict. sci. nat. 18: 461. 1821. Basionym: Arnica piloselloides L. TYPE: Sine loc., s.d., Burmann s.n., herb. Oldenland (holotype?, G-DC not seen). Additional specimen examined: Africa. Cameroon: Northwest province, near Verkovi village, Mt. Oku, D. Thomas & H. McLeod 6046 (MO). 25. Gerbera raphanifolia Franch., J. Bot. (Morot) 2: 67. 1888. TYPE: China. Yunnan: Mo-che-tsin, J. M. Delavay 1922 (isotype, GH!). 26. Gerbera serrata (Thunb.) Druce, Bot. Exch. Club Brit. Isles Rep.: 624. 1917. Basionym: Arnica serrata Thunb. TYPE: Sine loc., s.d., s. leg. herb. Thunberg 20009 (holotype, UPS not seen). Additional specimen examined: Africa. South Africa: Cape, Knysne Distr., Afgunst Flats, Fourcade 5309 (MO). 27. Gerbera tomentosa DC., Prodr. 7: 16. 1838. TYPE: Cap. Bonae-Spei, s.d., s. leg. (type, not localized). Additional specimens examined: Africa. South Africa: Cape Province, s.d., Zagler s.n. (MO); Kirstenbosch, Newlands, Wemmers Hoek, Dist. Paarl, R. Compton 745 (US). 28. Gerbera viridifolia (DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus viridifolius DC. var. hirsutus DC. TYPE: Africa. South Africa: Keiskama and Buffelriver, Drège 5110 (lectotype, designated by Hansen, 1985a: 24, G-DC not seen). Additional specimen examined: Africa. South Africa: Natal, Greytown, Aug 1934, J. Wylie s.n. (WIS). 29. Gerbera wrightii Harv., Fl. cap. 3: 521. 1865. TYPE: Africa. South Africa: Cape peninsula, near Simonstown, C. Wright 343 (holotype, TCD not seen).

Observation. There are many cultivars of Gerbera, an ornamental genus grown widely for the cut-flower market. An example is Gerbera hybrida, a functional diploid carrying chromosomes or chromosomal fragments from other Gerbera species such as G. jamesonii and G. viridifolia (Teeri et al., 2006). Literature. Zardini (1974), Hansen (1985a,b, 1988), Katinas (1998).

Glossarion Maguire & Wurdack Mem. New York Bot. Gard. 9: 390. 1957, emend. Pruski, Brittonia 41: 39. 1989. TYPE: Glossarion rhodanthum Maguire & Wurdack. 20 of Figs. 19–22. Guaicaia Maguire, Mem. New York Bot. Gard. 17: 437. 1967. TYPE: Guaicaia bilabiata Maguire [= Glossarion bilabiatum (Maguire) Pruski]. Etymology. From the Greek diminutive of glossa, tongue, in reference to the ligulate corollas that are occasionally found in the genus, these corollas being “diminutive” bilabiate corollas lacking the inner lips. Shrubs. Leaves alternate; petiolate, the petioles clasping; blades narrowly lanceolate to elliptic, coriaceous, pinnately veined, margin entire, abaxially bufftomentose. Capitulescences monocephalous, heads terminal or sometimes axillar;


551

capitula pedunculate, homogamous, isomorphic or dimorphic; receptacle epaleate; involucre multiseriate. Florets bisexual, isomorphic capitula with all ligulate or all bilabiate corollas; dimorphic capitula with ligulate and bilabiate corollas; ligulate corolla five-dentate; bilabiate corollas, with the inner lip either deeply two- to threecleft; anthers generally exserted, apical appendages acute, tails slightly papillose, connate with the tails of adjacent stamens; style red, bifid, branches dorsally smooth. Cypselae scarcely glandular or glabrous; pappus of scabrid bristles, somewhat flattened at the apex, the outermost shorter. Pollen Description. Glossarion rhodanthum. Pollen prolate, elliptic in equatorial view, circular in polar view, large size P × E = ca. (74–80×56–60) μm. Tricolporate, long colpi with psilate membrane. Exine Gongylolepis type, 12– 16 μm thick in the whole grain, spines 2.5 μm length. Ratio ectosexine/ endosexine: ca. 1:1. Nexine ca. 2.5 μm thick. Note: pollen is similar to that of Duidaea marahuacensis and Quelchia bracteata (see also Carlquist, 1957b; Barroso & Maguire, 1973; Tellería, 2008). Habitat and Distribution. Glossarion is a genus of two species endemic to Cerro de la Neblina in the Guayana Shield of southern Venezuela. Species list 1. Glossarion bilabiatum (Maguire) Pruski, Brittonia 41: 39. 1989. Basionym: Guaicaia bilabiata Maguire. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, below Pico Phelps, B. Maguire et al. 60475 (holotype, NY!). 2. Glossarion rhodanthum Maguire & Wurdack, Mem. New York Bot. Gard. 9: 390. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37027 (holotype, US!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Río Yatua, ca. 2 km N Cumbé camp, B. Maguire et al. 37149 (US*). Observation. Guaicaia was described as being most closely related to Glossarion, differing from it mainly by capitula having only bilabiate florets. However, capitula of Glossarion frequently contain not only ligulate florets, but also bilabiate ones. Therefore, Glossarion has been redefined by Pruski (1989a) to include Guaicaia. Literature. Maguire et al. (1957a), Aristeguieta (1964), Maguire (1967), Pruski (1989a, 1991, 1997). Gochnatia Kunth in Humb., Bonpl. & Kunth Nov. gen. sp. 4 (folio ed.): 15. 1818. TYPE: Gochnatia vernonioides Kunth. 21 of Figs. 19–22. Anastraphia D. Don, Trans. Linn. Soc. London 16: 295. 1830. TYPE: Anastraphia ilicifolia D. Don [= Gochnatia sagraeana Jervis & Alain].

552

L. Katinas, et al.

Leucomeris D. Don, Prodr. fl. nepal.: 169. 1825. [= Gochnatia sect. Leucomeris (D. Don) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 128. 1971]. TYPE: Leucomeris spectabilis D. Don [= Gochnatia spectabilis (D. Don) Less.]. Pentaphorus D. Don, Trans. Linn. Soc. London 16: 296. 1830. [= Gochnatia sect. Pentaphorus (D. Don) DC., Prodr. 7: 24. 1838]. TYPE: Pentaphorus foliolosus D. Don [= Gochnatia foliolosa (D. Don) D. Don ex Hook. & Arn.]. Richterago Kuntze, Revis. gen. pl. 1: 360. 1891. TYPE: Seris discoidea Less. [= Gochnatia discoidea (Less.) Cabrera]. Seris Less., Linnaea 5: 253. 1830, non Willd., 1807, nom. illeg. Etymology. In honor of Dr. Caroli Gochnat, a physician from Strasburg who studied the Cichorioideae ca. 1808. Shrubs or subshrubs, rarely trees, monoecious, gynodioecious, or polygamous. Leaves alternate; petiolate; blades ovate, elliptic to linear-elliptic, pinnately to threeveined, margin entire, serrulate to strongly dentate-spinose, subglabrous to glabrous above, densely tomentose beneath. Capitulescences monocephalous, corymbose, racemose, paniculate to glomerulose cymes; capitula pedunculate to subsessile, homogamous or less commonly heterogamous, discoid or disciform; receptacle epaleate; involucre multiseriate, occasionally the phyllaries decurrent onto the peduncle. Florets isomorphic, less commonly sub-dimorphic, bisexual or female with staminodes, corolla actinomorphic, tubular-funnelform, deeply five-lobed, or marginal corollas tubular-bilabiate with tube short (1/3 of corolla length), two incisions deeper than the others; anther apical appendages apiculate, tails smooth to papillose; style bilobed, branches dorsally smooth. Cypselae villose; pappus of scabrid, capillary bristles occasionally somewhat flattened, equal to unequal in length. Pollen Description. Gochnatia (93 of Figs. 92–94). Pollen spheroidal to prolate, circular in polar view, medium or large size. Tricolporate, colpi long with scabrate membrane, with or without mesoaperture (see also Parra & Marticorena, 1972; Roque & Silvestre-Capelato, 2001; Lin et al., 2005). On the basis of the pollen size and the exine thickness, two pollen types were recognized: type I and type II. Type I: pollen medium size, exine Mutisia type, microechinate, 2–6 μm thick. Found in G. argentina (Cabrera) Cabrera, G. barrosoae Cabrera (93A of Figs. 92–94), G. blanchetiana (DC.) Cabrera, G. hatschbachii Cabrera, G. haumaniana Cabrera, G. polymorpha (Less.) Cabrera (93B, C of Figs. 92–94), G. pulchra Cabrera, G. rusbyana Cabrera, G. sordida (Less.) Cabrera, and G. velutina (Bong.) Cabrera. Gochnatia barrosoae (93A of Figs. 92–94). Pollen spheroidal–subprolate, size P × E = (39–44×30–34) μm. Exine 5–6 μm thick, microechinate. Nexine 1.5 μm thick. Gochnatia blanchetiana. Pollen subprolate, medium size, P × E = (30–38×26–32) μm. Exine 2–4 μm thick, scabrate. Nexine ca. 1 μm thick. Gochnatia haumaniana. Pollen spheroidal– subprolate, size P × E = (32–36×25–29) μm. Mesoaperturate. Exine 2–4 μm thick, microechinate. Nexine ca. 1 μm thick. Type II: pollen large or medium size, exine Wunderlichia or Mutisia type, 6–10 μm thick. Found in G. amplexifolia, G. angustifolia G. Sancho, S. E. Freire & Katinas, G. buchii, G. glutinosa, G. magna, G. palosanto Cabrera, G. rotundifolia Less., G. shaferi, and G. vernonioides Kunth. Gochnatia amplexifolia. Pollen prolate, medium to large size, P × E = (47–51×32–


553

36) μm. Mesoaperturate. Exine Wunderlichia type, 6–7 μm thick, echinate, spines 2.5 μm length. Nexine ca. 1.5 μm thick. Gochnatia angustifolia (Sancho et al., 2005). Pollen subprolate, large size, exine Mutisia type, microechinate. Gochnatia buchii (93E of Figs. 92–94). Pollen subprolate-prolate, large size, P × E = (54–58×38– 47) μm. Mesoaperturate. Exine Wunderlichia type, 8–10 μm thick, echinate, spines 2.5 μm length. Nexine 2 μm thick. Gochnatia glutinosa. Pollen subprolate, medium size, P × E = ca. (44–49×36–37) μm. Exine Wunderlichia type, ca. 6 μm thick, microechinate. Nexine ca. 1.5 μm thick. Gochnatia magna. Pollen subprolate-prolate, medium to large size, P × E = (45–52×33–38) μm. Exine Wunderlichia type, 8– 9.5 μm thick, echinate, spines 2.5 μm length. Nexine ca. 2 μm thick. Gochnatia palosanto. Pollen subprolate, medium to large size, P × E = (49–56×37–44) μm. Exine Wunderlichia type, 8 μm thick, slightly thickened at the poles, microechinate. Nexine ca. 1.5 μm thick. Gochnatia rotundifolia. Pollen prolate, large size, P × E = ca. (56×38) μm. Exine Wunderlichia type, ca. 8 μm thick, echinate, spines ca. 1.5 μm length. Nexine ca. 2 μm thick. Gochnatia shaferi (93D of Figs. 92–94). Pollen prolate, large size, P × E = (55–61×41–47) μm. Exine Wunderlichia type, 8–9 μm thick, microechinate. Nexine 1.5–2 μm thick. Gochnatia vernonioides. Pollen prolate, medium to large size, P × E = (48–62×35–43) μm. Exine Wunderlichia type, 7–9 μm thick, microechinate. Nexine 1.5 μm thick. Habitat and Distribution. Genus of 70 species most from Mexico to Argentina, two species from Asia (China, India, Nepal, Pakistan, and southeastern Asia). Species list 1. Gochnatia amplexifolia (Gardner) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 155. 1971. Basionym: Seris amplexifolia Gardner. TYPE: Brazil. State Minas Geraes: Diamond District, elevated grassy tracts, G. Gardner 4787 (lectotype, designated by Roque & Pirani, 2001: 1157, BM not seen). Additional specimen examined: Brazil. State Minas Geraes: Santa Ana do Riacho, Serra do Cipó, G. Hatschbach 35312 (LP*). 2. Gochnatia angustifolia G. Sancho, S. E. Freire & Katinas, Taxon 54: 89. 2005. TYPE: Bolivia. Potosí: José M. Linares Lizarazu, Jatum Palmar, Torrico & Peca 336 (holotype, LPB!). 3. Gochnatia arborescens Brandegee, Zoe 5: 163. 1903. TYPE: Mexico. State Baja California: Cañon de Santa María, El Juste, 1 March 1898, C. A. Purpus s.n. (holotype, UC not seen). Additional specimen examined: Mexico. State Baja California: between San Antonio and Port of Bahía de los Muertos, Wiggins 5632 (US). 4. Gochnatia arequipensis Sandwith, Bull. Misc. Inform. Kew: 100. 1934. TYPE: Peru. Arequipa: Volcán de Arequipa, Lechler 2751 (holotype, K not seen). Additional specimen examined: Peru. Arequipa: Monte Chiwata, Eyerdam & Beetle 22120 (LP). 5. Gochnatia argentina (Cabrera) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Moquinia argentina Cabrera. TYPE: Argentina. Prov. Chaco: Colonia Benítez, Schulz 173 (holotype, LP!). Additional specimen examined: Argentina. Prov. Tucumán: Campo Alegre, Cruz Alta, S. Venturi 703 (LP).

554

L. Katinas, et al.

6. Gochnatia argyrea (Dusén ex Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Moquinia argyrea Dusén ex Malme. TYPE: Brazil. State Paraná: Capão Grande, Vila Velha, P. K. H. Dusén 4035 (lectotype, designated by Cabrera, 1971: 93, S!). 7. Gochnatia attenuata (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 8. 1960. Basionym: Anastraphia attenuata Britton. TYPE: Cuba. Prov. Oriente: Serra Nipe, J. A. Shafer 3113 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Sierra Nipe, E. L. Ekman 19174 (LP). 8. Gochnatia barrosoae Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. TYPE: Brazil. State Minas Geraes: Ituitaba, facenda do fundão, Macedo 1138 (holotype, LP!). Additional specimens examined: Brazil. State São Paulo, Mathes 3 (LP*); id., Mogí-Guaçu, Mantovani 903 (LP*). 9. Gochnatia blanchetiana (DC.) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Baccharis blanchetiana DC. TYPE: Brazil. State Bahia: Serra Jacobina, Blanchet 2569 (holotype, G-DC!). Additional specimen examined: Brazil. State Minas Gerais, Serra do Espinaço, Irwin et al. 28329 (SP*). 10. Gochnatia boliviana S. F. Blake, Contr. U. S. Natl. Herb. 22: 651. 1924. TYPE: Bolivia. Santa Cruz: Sierra de Santa Cruz, May 1892, C. E. O. Kuntze s.n. (holotype, US!). Additional specimen examined: Bolivia. Dept. Santa Cruz: Florida, A. L. Cabrera & Gutiérrez 33804 (LP). 11. Gochnatia buchii (Urb.) J. Jiménez Alm., Phytologia 8: 327. 1962. Basionym: Anastraphia buchii Urb. TYPE: Haiti. Prope Poste Coudau, W. Buch 1017 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Dominican Republic. Falda del Morro: Montecristo, Jiménez 1598 (LP); Montecristo, Jiménez 3613 (LP*). 12. Gochnatia calcicola (Britton) Jervis & Alain, Candollea 17: 120. 1960. Basionym: Anastraphia calcicola Britton. TYPE: Cuba. Prov. Oriente: Guantánamo, L. Britton 2042 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Guantánamo, E. L. Ekman 15764 (S). 13. Gochnatia cardenasii S. F. Blake, J. Wash. Acad. Sci. 25: 324. 1935. TYPE: Bolivia. Potosí: Quechisla, M. Cárdenas 326 (holotype, US!). Additional specimen examined: Argentina. Prov. Jujuy: Tafna, Cordo & Ferrer 88-B-17 (SI). 14. Gochnatia cordata Less., Linnaea 5: 263. 1830. TYPE: Brazil: sine loc, s. d., Sellow s.n. (holotype, B not seen, B photo F n. 15855 at LP!). Additional specimen examined: Argentina. Prov. Entre Ríos: Federación, A. Burkart & Crespo 23169 (LP). 15. Gochnatia cowellii (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 8. 1960. Basionym: Anastraphia cowellii Britton. TYPE: Cuba. Prov. Santa Clara: Palm Barren, Britton & Cowell 10183 (holotype, NY!). 16. Gochnatia crassifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia crassifolia Britton. TYPE: Cuba. Prov. Oriente: Río Yamanigüey, J. A. Shafer 4261 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Baracoa, E. L. Ekman 4023 (S).


555

17. Gochnatia cubensis (Carabia) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia cubensis Carabia. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, J. P. Carabia 3788 (holotype, NY not seen, NY photo at LP!). 18. Gochnatia curviflora (Griseb.) O. Hoffm., Nat. Pflanzenfam. 5: 337. 1893. Basionym: Moquinia curviflora Griseb. TYPE: Bolivia. Tarija: Cuesta del Tambo, P. G. Lorentz & G. Hieronymus 879 (isotype, CORD not seen). Additional specimen examined: Bolivia. Tarija, Fiebrig 2838 (LP). 19. Gochnatia decora (Kurz) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 131. 1971. Basionym: Leucomeris decora Kurz. TYPE: Burma. Pegu, Yomah, S. Kurz 2241 (isotype, G not seen). Additional specimen examined: Myanmar. Bhamo: Lweji, Maung Mya 5309 (LP). 20. Gochnatia densicephala (Cabrera) G. Sancho, Novon 9: 559. 1999. Basionym: Gochnatia paniculata (Less.) Cabrera var. densicephala Cabrera. TYPE: Brazil. State Rio de Janeiro, A. F. M. Glaziou 11072 (holotype, K!). 21. Gochnatia discoidea (Less.) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 153. 1971. Basionym: Seris discoidea Less. TYPE: Brazil. Serra da Gacheira, Sellow 794, 994 [lectotype (sic), designated by Roque & Pirani, 2001: 1158, not seen]. Additional specimen examined: Brazil. State Bahia: Igregia Velha, Blanchet 3345 (LP). 22. Gochnatia discolor Baker, Fl. bras. 6: 350. 1884. TYPE: Brazil. State Minas Geraes, 1840, Claussen s.n. (isotype, G!). 23. Gochnatia ekmanii (Urb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia ekmanii Urb. TYPE: Cuba. Prov. Pinar del Río: Sierra de los Organos, E. L. Ekman 13865 (isotype, NY not seen, NY photo at LP!). 24. Gochnatia elliptica (Léon) Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia elliptica Léon. TYPE: Cuba. Prov. Oriente: Ovando, Maisí, Fr. Léon 17580 (isotype, NY not seen, NY photo at LP!). 25. Gochnatia enneantha (S. F. Blake) Alain, Bull. Torrey Bot. Club 92: 304. 1965. Basionym: Anastraphia enneantha S. F. Blake. TYPE: Dominican Republic. Lajama, Samaná Península, W. L. Abbott 2287 (holotype, US!). Additional specimen examined: Dominican Republic. Cordillera Central: Samaná, Los Haitises, E. L. Ekman H-15498 (S). 26. Gochnatia floribunda Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 125. 1971. TYPE: Brazil. State Minas Geraes: Cerro do Frio, G. Gardner 4806 (isotype, G!). 27. Gochnatia foliolosa (D. Don) D. Don ex Hook. & Arn., Comp. Bot. Mag. 1: 108. 1835. Basionym: Pentaphorus foliolosus D. Don. TYPE: Chile: sine loc., s. d., Caldcleugh s.n. (types, probably BM, K), Additional specimen examined: Chile. Prov. Santiago: Santiago, Cerro Renca, A. L. Cabrera 3451 (LP). 28. Gochnatia gardneri (Baker) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia gardneri Baker. TYPE: Brazil. State Goyaz: Capella da Passe, G. Gardner 4183 (lectotype, designated by Cabrera, 1971: 109, K!). 29. Gochnatia glutinosa (D. Don) D. Don ex Hook. & Arn., Comp. Bot. Mag. 1: 108. 1835. Basionym: Pentaphorus glutinosus D. Don. TYPE: Argentina.

556

30.

31.

32.

33.

34.

35.

36.

37.

38.

39.

40.

L. Katinas, et al.

Prov. Mendoza: near San Isidro, s.d., J. Gillies s.n. (type, probably K). Additional specimens examined: Argentina. Prov. Tucumán: Tafí, H. A. Fabris 1343 (LP); Prov. Salta: Trancas, Cuesta del Anca, Jan 1897, C. L. Spegazzini s.n. (LP*). Gochnatia gomezii (Léon) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia gomezii Léon. TYPE: Cuba. Prov. Oriente: Cerro de Miraflores, Fr. León & Gómez 28876 (holotype, HAC not seen). Additional specimen examined: Cuba. Prov. Oriente: Moa, Cerro Miraflores, Marie-Victorin et al. 21591 (GH). Gochnatia hatschbachii Cabrera, Bol. Mus. Bot. Munic. 15: 3. 1974. TYPE: Brazil. State Minas Geraes: Jaboticatubas, Serra do Cipó, G. Hatschbach 29951 (holotype, LP!). Gochnatia haumaniana Cabrera, Notas Mus. La Plata, Bot. 15: 44. 1950. TYPE: Paraguay. Pedro Juan Caballero: Sierra de Amambay, T. Rojas 6575 (holotype LP!). Additional specimen examined: Paraguay. Pedro Juan Caballero: Sierra de Amambay, T. Rojas 10391 (K*). Gochnatia hiriartiana Medrano, Villaseñor & Medina, Novon 14: 434. 2004. TYPE: Mexico. Hidalgo: Meztitlán, González-Medrano, G. G. Hernández & G. Rodríguez 17920 (holotype, MEXU not seen). Gochnatia hypoleuca (DC.) A. Gray, Proc. Amer. Acad. Arts 19: 57. 1883. Basionym: Moquinia hypoleuca DC. TYPE: Mexico. State Nueva León: Monterrey, Berlandier 1391 (isotype, US!). Additional specimen examined: Mexico. State Hidalgo: Cañada del Vaquero, Gonzalez Quintero 3215 (LP). Gochnatia ilicifolia Less., Linnaea 5: 261. 1830. TYPE: America (Guiane?), sine loc., s.d., Richard s.n. (holotype, B not seen). Additional specimen examined: Bahamas Islands. Andros Island: Coppice, Small & Carter 8526 (K). Gochnatia intertexta (C. Wright ex Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia inertexta C. Wright ex Griseb. TYPE: Cuba. Bahia Onda, prope San Marcos, C. Wright 2877 (isotype, BM not seen). Additional specimen examined: Cuba. Prov. Pinar del Río: Cajálbana, Alain A-1680 (NY). Gochnatia magna M. C. Johnst. ex Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 147. 1971. TYPE: Mexico. State Tamaulipas: half mile E from Nuevo Morelos, J. Graham & M. C. Johnston 4485 (holotype, MEXU not seen). Additional specimen examined: Mexico. San Luís Potosí, A. Cronquist 11277 (NY). Gochnatia maisiana (Léon) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia maisiana Léon. TYPE: Cuba. Prov. Oriente: Mesa del Chivo, Fr. Léon 18243 (isotype, NY not seen, NY photo at LP!). Gochnatia mantuensis (C. Wright ex Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia mantuensis C. Wright ex Griseb. TYPE: Cuba. Nueva Filipina, prope Mantua, C. Wright 2876 (isotype, BM not seen). Additional specimen examined: Cuba. Prov. Pinar del Río: Guane, Shafer 11208 (LP). Gochnatia microcephala (Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia microcephala Griseb. TYPE: Cuba. Ex Catae, 1860–64, C. Wright s.n. (isotype, NY not


41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

51.

52.

557

seen). Additional specimen examined: Cuba. Boca Guantánamo a Mantua, Bro. Hioram 4874 (NY). Gochnatia mollisima (Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia mollissima Malme. TYPE: Brazil. Río Grande do Sul, G. A. Malme 648 (holotype, S!). Gochnatia montana (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia montana Britton. TYPE: Cuba. Prov. Pinar del Río: Sierra Caliente, J. A. Shafer 13781 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Guane, E. L. Ekman 18725 (S). Gochnatia obtusifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia obtusifolia Britton. TYPE: Cuba. Prov. Oriente: camp La Gloria, J. A. Shafer 8189 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Southern Baracoa region, Fr. Léon 11963 (NY). Gochnatia oligantha (Urb.) R. A. Howard, J. Arnold Arbor. 42: 137. 1961. Basionym: Anastraphia oligantha Urb. TYPE: Dominican Republic. Santo Domingo, 1871, C. Wright et al. 248 (isotype, NY!). Additional specimen examined: Dominican Republic. Montecristo, N of Villa Isabel, Jiménez 3614 (US). Gochnatia oligocephala (Gardner) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia oligocephala Gardner. TYPE: Brazil. State Ceará: Serra de Araripe, G. Gardner 2422 (holotype, BM not seen). Additional specimen examined: Brazil. State Bahia: Rio Branco, C. H. Curran 284 (NY). Gochnatia orbiculata (Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia orbiculata Malme. TYPE: Brazil. State Paraná: Vilha Velha, P. Dusén 9437 (lectotype, designated by Cabrera 1971: 105, S!). Additional specimen examined: Brazil. State Rio Grande do Sul, Tweedie 998 (K). Gochnatia palosanto Cabrera, Notas Mus. La Plata, Bot. 1: 62. 1935. TYPE: Argentina. Prov. Tucumán: Trancas, Vipos, C. Schreiter 85 (holotype, LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Candelaria, Sierra de la Candelaria, S. Venturi 9793 (LP*). Gochnatia paniculata (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia paniculata Less. TYPE: Brazil. São Paulo: Casa Branca, Loefgren 16994 (neotype, designated by Sancho, 1999: 557, US!). Gochnatia parvifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia parvifolia Britton. TYPE: Cuba. Prov. Oriente: Barren Savannas, Shafer 2938 (holotype, NY!). Gochnatia patazina Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 28. 1971. TYPE: Peru. La Libertad: Pataz, between Hualillas and Tayabamba, A. López & A. Sagástegui 3409 (holotype, LP!). Gochnatia pauciflosculosa (C. Wright ex Hitchc.) Jervis ex Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 48. 1971. Basionym: Anastraphia pauciflosculosa C. Wright ex Hitchc. TYPE: Bahamas Islands. Inagua: 12 April 1890, A. S. Hitchcock s.n. (lectotype, designated by Cabrera, 1971: 50, MO not seen). Additional specimen examined: Bahamas Islands. Fortune Island, Eggers 3866 (K). Gochnatia picardae (Urb.) J. Jiménez Alm., Phytologia 8: 327. 1962. Basionym: Anastraphia picardae Urb. TYPE: Haiti. Prope Cadets, Picarda

558

53.

54. 55.

56. 57.

58.

59. 60.

61.

62.

63.

64.

L. Katinas, et al.

1073 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Haiti. Massif de la Selle: Croix-des Bauquets, gorge of Grande-Rivière de cul de sac, E. L. Ekman H-5385 (K). Gochnatia polymorpha (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia polymorpha Less. TYPE: Brazil: s.d., Sellow s. n., (holotype, B not seen). Additional specimens examined: Brazil. State Bahia: Jacobina, Igreja Velha, Blanchet 3251 (F); State Rio de Janeiro: Tijuca, A. F. M. Glaziou 3039 (LP*). Gochnatia pulchra Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 106. 1971. TYPE: Brazil. São Paulo: Ityrapina, 28 April 1923, Gehrt s.n. (holotype, LP!). Gochnatia purpusii Brandegee, Zoe 5: 240. 1906. TYPE: Mexico. State Puebla: Tehuacán, C. A. Purpus 1164 (holotype, UC not seen). Additional specimen examined: Mexico. Tehuacán, Purpus 4248 (NY). Gochnatia ramboi Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 98. 1971. TYPE: Brazil. State Rio Grande do Sul: Palmeira, B. Rambo 51961 (holotype, LP!). Gochnatia recurva (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. Basionym: Anastraphia recurva Britton. TYPE: Cuba. Prov. Oriente: Serpentine hills near mouth of Rio Yamaniguey, J. A. Shafer 4257 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: between Moa and Punta Andén, Léon 20946 (LP). Gochnatia rotundifolia Less., Syn. gen. Compos.: 102. 1832. TYPE: Brazil: sine loc., s.d., Sellow s.n. (holotype, B not seen). Additional specimens examined: Brazil. State São Paulo: Capital, Jabaquara, O. Handro 157 (LP); id., Ypiranga, Brade 6265 (LP*). Gochnatia rusbyana Cabrera, Notas Mus. La Plata, Bot. 15: 41. 1950. TYPE: Bolivia. Yungas, M. Bang 2252 (isotypes, G!, LP!, MO!). Gochnatia sagraeana Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 9. 1960. TYPE: Cuba. Habana, Jibacoa, Fr. León et al. 19708 (type, probably HAC). Additional specimen examined: Cuba. Habana, Fr. Léon 7094 (NY). Gochnatia shaferi (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 10. 1960. Basionym: Anastraphia shaferi Britton. TYPE: Cuba. Prov. Oriente: Río Naranja, J. A. Shafer 3865 (isotype, US!). Additional specimens examined: Cuba. Prov. Oriente: La Caridad, López Figueiras 1738 (NY); id., Sierra de Nipe, Victorin & Clemente 22031 (LP, NY*). Gochnatia smithii B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 32: 50. 1896. TYPE: Mexico. State Oaxaca: Cuicatlan, L. C. Smith 363 (lectotype, designated by Cabrera, 1971: 140, K not seen). Additional specimen examined: Mexico. State Oaxaca: Cuesta de Coyula, Conzatti 4135 (US). Gochnatia sordida (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia polymorpha Less. var. sordida Less. TYPE: Brazil. s.d., Sello s.n. (isotype, K!). Gochnatia spectabilis (D. Don) Less., Linnaea 5: 261. 1830. Basionym: Leucomeris spectabilis D. Don. TYPE: Nepal. Sirinagur, 1821, Wallich s.n. (isotype, G not seen). Additional specimen examined: India. Uttar Pradesh: Rajpur, Dehra Dun District, Galrola 32 (LP).


559

65. Gochnatia suffrutescens Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 150. 1971. TYPE: Brazil. State Minas Geraes: São João del Rey, Mello Barreto 4744 (holotype, F not seen). 66. Gochnatia tortuensis (Urb.) J. Jiménez Alm., Phytologia 8: 328. 1962. Basionym: Anastraphia tortuensis Urb. TYPE: Haiti: La Tortue Island, E. L. Ekman H-4313 (isotype, US!). Additional specimen examined: Haiti. Presquile du Nord-Ouest: Port-de Paix, E. L. Ekman H-3553 (S). 67. Gochnatia vargasii Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 19. 1971. TYPE: Peru. Apurimac: Abancay, C. Vargas 16317 (holotype, LP!). 68. Gochnatia velutina (Bong.) Cabrera, Notas Mus. La Plata, Bot. 15: 44. 1950. Basionym: Moquinia velutina Bong. TYPE: Brazil. State São Paulo: s.d., Langsdorff s.n. (holotype, LE not seen, LE photo at LP!). Additional specimen examined: Brazil. State Paraná: Ponta Grossa, Vila Velha, R. M. Klein & S. B. Smith 14885 (LP). 69. Gochnatia vernonioides Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 16. 1818. TYPE: Peru. Dept. Amazonas: s.d., F. Humboldt & A. Bonpland s.n. (isotype, K not seen). Additional specimens examined: Peru. Dept. Amazonas: Prov. Chachapoyas, Tingo, R. Ferreyra 7097 (LP); Prov. Chachapoyas, Leimebamba, A. López et al. 4364 (LP*). 70. Gochnatia wilsonii (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 18: 10. 1960. Basionym: Anastraphia wilsonii Britton. TYPE: Cuba. Santa Clara: Río Tobaya, Trinidad, N. L. Britton et al. 5573 (holotype, NY not seen, NY photo at LP!).

Observation. See Observations in Actinoseris. The Asian species of Gochnatia belonging to the sect. Leucomeris (G. decora and G. spectabilis) as well as those belonging to the sect. Pentaphorus (G. foliolosa and G. glutinosa) have been lately regarded as independent genera from Gochnatia (i.e., Leucomeris and Pentaphorus respectively; Hind, 2007; Panero & Funk, 2008). Literature. Jervis (1954), Cabrera (1971a, 1974), Sancho (1996, 1999, 2000), Freire et al. (2002), González-Medrano et al. (2004), Sancho et al. (2005).

Gongylolepis R. H. Schomb. Linnaea 20: 759. 1847. TYPE: Gongylolepis benthamiana R. H. Schomb. 22 of Figs. 19–22. Cardonaea Aristeg., Maguire & Steyerm., Mem. New York Bot. Gard. 23: 892. 1972. TYPE: Cardonaea jauaensis Aristeg., Maguire & Steyerm. [= Gongylolepis jauaensis (Aristeg., Maguire & Steyerm.) V. M. Badillo]. Neocaldasia Cuatrec. Revista Acad. Colomb. Ci. Exact. 6: 16. 1944. TYPE: Neocaldasia colombiana Cuatrec. [= Gongylolepis colombiana (Cuatrec.) Cuatrec.]. Etymology. From the Greek gongylos, round, and lepis, scale, in reference to the rounded phyllary apices.

560

L. Katinas, et al.

Shrubs or trees, less commonly subshrubs. Leaves alternate, often clustered at the apex; sessile to petiolate, petioles, when present, commonly clasping; blades oblanceolate, lanceolate to obovate, coriaceous, venation pinnate, sub-palmate or sub-parallel, margin entire, sometimes revolute, glabrous to abaxially pubescent. Capitulescences monocephalous or cymose; capitula sessile to pedunculate, peduncles sometimes with leaf-like bracts, homogamous, bilabiate, six- to manyflowered; receptacle epaleate; involucre multiseriate, campanulate to hemispherical, phyllaries graduate. Florets isomorphic, bisexual, corolla bilabiate; anthers purple-brown, mostly exserted, apical appendages acuminate, tails slightly papillose, often connate; style white to tinged with purple, bifid, branches dorsally smooth. Cypselae commonly glabrous; pappus of scabrid, persistent capillary bristles. Pollen Description. Gongylolepis (94 of Figs. 92–94). Pollen spheroidal to subprolate, spheroidal or elliptic in equatorial view, circular in polar view, medium or large size. Tricolporate, colpi long or short with margin and psilate membrane. Exine Gongylolepis type, uniformly thickened. Note: pollen very similar to that of Achnopogon (see also Carlquist, 1957b; Barroso & Maguire, 1973). Gongylolepis benthamiana. Pollen subprolate, elliptic in equatorial view, large size, P × E = (72– 94×61–78) μm. Colpi 47–60 μm length. Exine 10–11 μm thick in the whole grain, spines 2.5–4 μm length. Nexine 2–4 μm thick. Gongylolepis huachamacari (94 of Figs. 92–94). Pollen spheroidal–subprolate, spheroidal in equatorial view, large size P × E = (74–88×70–80) μm. Long colpi. Exine 7–10 μm thick, spines 5–6 μm length. Nexine ca. 3 μm thick. SEM: tectum very perforate with striate aspect, apertural membrane scabrate (see also Tellería, 2008). Habitat and Distribution. The genus contains 14 species, 13 of which are endemic to the Guayana region of Colombia, Venezuela, Guyana, extending southward toward Manaus, Brazil. However, one species, Gongylolepis colombiana (Cuatrec.) Cuatrec., occurs in the northern Andes of Colombia and Venezuela. Species list 1. Gongylolepis benthamiana R. H. Schomb., Linnaea 20: 760. 1847. TYPE: Guyana: Schomburgk 11 (holotype, K not seen). Additional specimen examined: Venezuela. State Bolívar: Distr. Roscio, 13 km al N de Santa Elena de Vaivén, J. A. Steyermark & R. Liesner 127536 (MO*). 2. Gongylolepis bracteata Maguire, Mem. New York Bot. Gard. 8: 156. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Sipapo, B. Maguire & L. Politi 27604 (isotype, US). 3. Gongylolepis colombiana (Cuatrec.) Cuatrec., Fieldiana, Bot. 27: 51. 1950. Basionym: Neocaldasia colombiana Cuatrec. TYPE: Colombia. Norte de Santander, J. Cuatrecasas 13435 (isotype, US!). Additional specimen examined: Venezuela. State Tachira, H. van der Werff & F. Ortega 8077 (MO). 4. Gongylolepis cortesii (S. Díaz) Pruski & S. Díaz, Fl. Venez. Guayana 3: 285. 1997. Basionym: Eurydochus cortesii S. Díaz. TYPE: Colombia. Guinia: Serranía del Naquén, camino de Caño Rico al Campamento 15, Cortés et al. 238 (isotype, US!).


561

5. Gongylolepis erioclada S. F. Blake, Bull. Torrey Bot. Club 58: 495. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Duida, G. H. H. Tate 758 (holotype, US!). 6. Gongylolepis fruticosa Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 437. 1957. TYPE: Venezuela. State Bolívar: cerro Chimantá, J. A. Steyermark & J. J. Wurdack 1136 (holotype, NY!). 7. Gongylolepis glaberrima S. F. Blake, Bull. Torrey Bot. Club 58: 494. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida, G. H. H. Tate 617 (holotype, NY!). 8. Gongylolepis huachamacari Maguire, Mem. New York Bot. Gard. 8: 159. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, B. Maguire et al. 30135 (holotype, NY!). Additional specimens examined: Venezuela. Dept. Río Negro: Cerro Aracámuni, 1° 26′ N, 65° 47′ W, R. Liesner & F. Delascio 21960 (MO*); Terr. Fed. Amazonas, Cerro Huachamacari, B. Maguire et al. 30080 (MO*). 9. Gongylolepis jauaensis (Aristeg., Steyerm. & Wurdack) V. M. Badillo, Ernstia 46: 8. 1988. Basionym: Cardonaea jauaensis Aristeg., Steyerm. & Wurdack. TYPE: Venezuela. State Bolívar: Meseta de Jaua, Cerro Jaua, J. A. Steyermark 97945 (holotype, VEN!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atabapo, cerro Marahuaca, below Salto Los Hornos, R. Liesner 25112 (MO). 10. Gongylolepis martiana (Baker) Steyerm. & Cuatrec., Fieldiana, Bot. 28: 640. 1953. Basionym: Stifftia martiana Baker. TYPE: Colombia. Terr. Fed. Amazonas: Araracuara, s.d., C. Martius s.n. (holotype, M not seen). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atures, 10 km al S del Río Guayapo, O. Huber 3293 (MO). 11. Gongylolepis oblanceolata Pruski, Ann. Missouri Bot. Gard. 76: 994. 1989. TYPE: Brazil. State Amazonas: Barcelos, Platô da Serra Aracá, M. G. Silva et al. 7169 (holotype, INPA!). Additional specimen examined: Brazil. State Amazonas: Municipio de Barcelos, plato da Serra Aracá, I. L. Amaral 1557 (US). 12. Gongylolepis paniculata Maguire & K. D. Phelps, Mem. New York Bot. Gard. 8: 157. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Guanay, B. Maguire et al. 31655 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro Yaví, K. Phelps & C. Hitchcock 8 (MO). 13. Gongylolepis paruana Maguire, Mem. New York Bot. Gard. 8: 159. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Serranía Parú, Río Parú, Caño Asísa, Río Ventuari, R. S. Cowan & J. J. Wurdack 31239 (holotype, NY!). 14. Gongylolepis pedunculata Maguire, Mem. New York Bot. Gard. 8: 156. 1953. TYPE: Venezuela. Amazonas: Serranía Parú, Río Parú, Caño Asísa, Río Ventuari, R. S. Cowan & J. J. Wurdack 31131 (holotype, NY!).

Literature. Maguire et al. (1957a), Aristeguieta (1964), Steyermark et al. (1972), Badillo (1988), Pruski (1989b, 1991, 1997), Díaz-Piedrahita and Vélez-Nauer (1993).

562

L. Katinas, et al.

Gypothamnium Phil. Fl. atacam.: 27. 1860. TYPE: Gypothamnium pinifolium Phil. 23 of Figs. 23–26. Etymology. From the Greek gyps, gypos, vulture, and thamnos, shrub. Shrubs. Leaves alternate, densely arranged in the upper branches; sessile; blades linear, margin entire, one-veined, glabrous. Capitulescences monocephalous, terminal; capitula sessile, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages reddish, truncate, tails papillose; style bilobed, branches dorsally papillose. Cypselae villose; pappus of scabrid bristles, the innermost flattened, ciliate at the apex, the outermost shorter, not ciliate, rigid. Pollen Description (Tellería & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Parra & Marticorena, 1972). Habitat and Distribution. Monotypic genus endemic to northern Chile. Species list 1. Gypothamnium pinifolium Phil., Fl. atacam.: 27. 1860. TYPE: Chile. Prov. Antofagasta: cerca de las Animas, Cachinal de la Costa, Taltal, en el valle de Guanillo más arriba de Paposo, etc., R. Philippi 2349, s.n. (syntypes, SGO not seen, SGO photos at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Taltal, Francke 1044 (LP). Literature. Cabrera (1951).

Hecastocleis A. Gray Proc. Amer. Acad. Arts 17: 220. 1882. TYPE: Hecastocleis shockleyi A. Gray. 24 of Figs. 23–26. Etymology. From the Greek hekastos, each, and kleis, lock, key, to shut up, which refers to each floret enclosed by the phyllaries of the involucre. Shrubs. Leaves alternate; sessile; blades linear-elliptic to oblanceolate, pinnately veined, margin spiny, glabrous above, subglabrous beneath. Capitulescence a pseudocephalium, several-headed, surrounded by spiny wide-ovate bracts; capitula sessile, one-flowered; receptacle epaleate; involucre multiseriate, cylindrical, phyllaries coriaceous with spiny apex. Florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages subacute, tails


563

Figs. 23–26 23 Gypothamnium pinifolium Phil. A Habit (branch). B Marginal corolla. C Staminode. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (From Francke 1044, LP). Scale bars: A=5 cm; B, D=5.5 mm; C, E=2.5 mm; F=1.25 mm; G=4 mm. 24. Hecastocleis shockleyi A. Gray. A Habit (branch). B One-flowered capitulum. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (From Train 3973, LP). Scale bars: A=3 cm; B=5 mm; C=3.5 mm; D=3 mm; E= 0.6 mm; F=2.5 mm. 25 Hyalis lancifolia Baker. A Habit. B Marginal corolla. C Central corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1978: 583). Scale bars: A= 6 cm; B, C, F=6 mm; D, E=2.3 mm. 26 Hyaloseris cinerea (Griseb.) Griseb. A Habit (branch). B Stamen. C Upper part of the style. D Cypsela with pappus. (Redrawn from Ariza Espinar, 1973: 202). H. camataquiensis Hieron. ex J. Kost. E Corolla. (Redrawn from Cabrera, 1978: 612). Scale bars: A=2.5 cm; B=5 mm, C=2.5 mm; D=10 mm; E=9 mm

smooth; style bilobed, branches dorsally smooth. Cypselae glabrous; pappus a ring of short, laciniate scales. Pollen Description (Tellería & Katinas, 2005) (95 of Figs. 95–100). Pollen spheroidal, medium size, tricolpate, exine Mutisia type, scabrate.

564

L. Katinas, et al.

Habitat and Distribution. Monotypic genus endemic to dry, rocky slopes in western United States. Species list 1. Hecastocleis shockleyi A. Gray, Proc. Amer. Acad. Arts 17: 221. 1882. TYPE: United States. State Nevada: Esmeralda County, Candelaria, W. H. Schockley 55 (isotype, US not seen, US photo at LP!). Additional specimens examined: United States. State Nevada: 5 mi. of Basalt, on Mt. Montgomery-Tonopah highway, Mineral-Esmeralda Co. line, Train 3973 (LP*); State California, Inyo Co., Eureka valley to Fish Lake valley, Alexander & Kellogg 5301 (US*); Death valley region, Leadfield, Gilman 1144 (US*). Literature. Simpson (1978), Panero and Funk (2002).

Hyalis D. Don ex Hook. & Arn. Comp. Bot. Mag. 1: 108. 1835. TYPE: Hyalis argentea D. Don ex Hook. & Arn. 25 of Figs. 23–26. Etymology. From the Greek hyaleos, of glass, transparent, probably because of the light color of the leaves or the shiny, scarious involucral bracts. Subshrubs rhizomatous, single to few-branched stems. Leaves alternate; sessile; blades oblong to linear-elliptic, margin entire to serrulate, three-veined, glabrous to pubescent. Capitulescences densely corymbose; capitula short-pedunculate, homogamous, radiate, five- to six-flowered; receptacle epaleate; involucre three- to six-seriate. Florets dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded; central floret 1, tubular-funnelform corolla, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bifid, branches dorsally smooth. Cypselae villose; pappus of scabrid to smooth, capillary bristles, unequal in length. Pollen Description. Hyalis argentea (Tellería & Forcone, 2002). Pollen prolate– subprolate, medium to large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Genus of two species that grow in sandy areas and in forests of Schinopsis Engl. (Anacardiaceae) and Copernicia Mart. (Arecaceae) of northern Argentina, Bolivia, and Paraguay. Species list 1. Hyalis argentea D. Don ex Hook. & Arn., Comp. Bot. Mag. 1: 108. 1835. TYPE: Argentina. Prov. Mendoza: Mendoza, s.d., J. Gillies s.n. Prov. San Luis: Río Cuarto, s.d., J. Gillies s.n.. Prov. Córdoba: Villa del Río Cuarto, pampas of Córdoba, s.d., J. Gillies s.n. Prov. Buenos Aires: salt plains of Bahía Blanca,


565

north Patagonia, s.d., Tweedie s.n. (syntypes, probably K). Additional specimen examined: Argentina. Prov. Mendoza: Tupungato, A. Ruiz Leal 3701 (LP). 2. Hyalis lancifolia Baker, Fl. bras. 6: 368. 1882. TYPE: Argentina. Prov. Salta: s.d., Pearce s.n. (type, not localized). Additional specimen examined: Argentina. Prov. Chaco: San Fernando, A. L. Cabrera 4083 (LP). Literature. Cabrera (1951, 1978), Freire et al. (2002).

Hyaloseris Griseb. Symb. fl. argent.: 212. 1879. TYPE: Gochnatia cinerea Griseb. [= Hyaloseris cinerea (Griseb.) Griseb.]. 26 of Figs. 23–26. Etymology. From the Greek hyaleos, of glass, and seris, chicory, lettuce, probably because of the shiny, scarious involucral bracts. Shrubs or little trees, with brachyblasts and macroblasts. Leaves alternate to opposite, clustered in the brachyblasts; short-petiolate; blades linear-oblong, elliptic to ovate, pinnately veined, margin entire to dentate, glabrous to pubescent. Capitulescences monocephalous or two- to four-headed, racemose to spicate; capitula sessile to pedunculate, homogamous, discoid or ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic or with transitional corollas in the same capitulum, bisexual, ligulate, shallowly five-dentate limb, occasionally some segments partially connate, sub-bilabiate (4+1 corolla lips), rarely bilabiate; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Hyaloseris cinerea (96 of Figs. 95–100) (Tellería & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Genus of seven species of dry and rocky soils in Bolivia and Argentina. Species list 1. Hyaloseris andrade-limae Cristóbal & Cabrera, Hickenia 1: 255. 1982. TYPE: Argentina. Prov. Santiago del Estero: Dept. Guasayán, Sierra de Guasayán, ruta 64, 70 km de Santiago del Estero, A. Krapovickas et al. 37415 (isotype, LP!). 2. Hyaloseris boliviensis J. Kost., Blumea 5: 668. 1945. TYPE: Bolivia. Pulquina, T. Herzog 1798 (isotype, LP!). Additional specimen examined: Bolivia. Dept. Cochabamba: Prov. Totora, Higuerita, Steinbach 3877 (LIL, not seen). 3. Hyaloseris camataquiensis Hieron. ex J. Kost., Blumea 5: 667. 1945. TYPE: Bolivia. Catamaqui, Fiebrig 3067 (isotype, S not seen). Additional specimen examined. Bolivia. Quechisla, M. Cárdenas 11 (LP). 4. Hyaloseris cinerea (Griseb.) Griseb., Symb. fl. argent.: 213. 1879. Basionym: Gochnatia cinerea Griseb. TYPE: Argentina. Prov. Catamarca: Fuerte de

566

L. Katinas, et al.

Andalgalá, P. G. Lorentz 690 (isotype, CORD not seen). Additional specimen examined: Argentina, Prov. Córdoba, entre Taninga y Chancaní, Sierra de Pocho, A. Ragonese 9335 (LP*). 5. Hyaloseris longicephala B. L. Turner, Phytologia 59: 317. 1986. TYPE: Bolivia. Chuquisaca: Prov. Oropeza, an der strasse von Sucre Richtung Cochabamba, vor Surima, T. Feuerer & N. Hohne 45762 (holotype, TEX not seen). 6. Hyaloseris quadriflora J. Kost., Blumea 5: 667. 1945. TYPE: Bolivia. Cochabamba: Cerro de San Pedro, T. Herzog 2461 (isotypes, CONC!, LP!). 7. Hyaloseris rubicunda Griseb., Symb. fl. argent.: 213. 1879. TYPE: Argentina. Prov. Catamarca: Dept. Belén, Portezuelo, entre Cortadera y Ciénaga, Schickendantz 284 (lectotype, designated by Ariza Espinar, 1973: 205 as “isotype", CORD not seen). Additional specimen examined: Argentina. Prov. Salta: alrededores de Cachi, Jan 1897, C. L. Spegazzini s.n. (LP). Literature. Ariza Espinar (1973), Cristóbal and Cabrera (1982), Turner (1986).

Ianthopappus Roque & D. J. N. Hind Novon 11: 97. 2001. TYPE: Gochnatia corymbosa Less. [= Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind]. 27 of Figs. 27–30. Etymology. From the Latin ianthum, purple, and pappus, apical tuft of bristles on the fruit, referring to the color of the pappus. Subshrubs unbranched. Leaves alternate; short-petiolate; blades wide-obovate to orbicular, three-veined, margin entire, glabrous above, densely white-tomentose beneath. Capitulescences corymbose, two- to 18-headed; capitula long-pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate, three- to sixseriate. Florets dimorphic; marginal florets female with staminodes, corollas bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally smooth. Cypselae villose; pappus purplish, scabrid bristles, the innermost longer and plumose at the apex. Pollen Description. Ianthopappus corymbosus (97 of Figs. 95–100) (Roque & Silvestre Capelato, 2001). Pollen prolate, large size, tricolporate, exine Mutisia type, microechinate. Pollen close to that of type II of Gochnatia. Habitat and Distribution. Monotypic genus from eastern Argentina, southeastern Brazil, and northwestern Uruguay in “campestre” vegetation with predominance of steppe. Species list 1. Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind, Novon 11: 97. 2001. Basionym: Gochnatia corymbosa Less. TYPE: Brazil. Sello 3479 (neotype,


567

Figs. 27–30 27 Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind. A Habit (branch). B Marginal corolla. C Staminode. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (A, B, D– G redrawn from Roque & Hind, 2001: 99; C from Palacios-Cuezzo 2304, LP). Scale bars: A=4 cm; B=3 mm; C, E=1 mm; D=2 mm; F=0.5 mm; G=4 mm. 28 Leibnitzia seemanii (Sch. Bip.) G. L. Nesom. A–H Cleistogamous form. A Habit. B, C Outermost marginal florets (without cypselae). D Innermost marginal floret (without cypsela). E Central corolla. F Stamen. G Upper part of the style. H Cypsela with pappus. I–L Chasmogamous form. I Habit. J, K Marginal florets (without cypselae). L Central floret (without cypsela). (A–E, J–L redrawn from Nesom, 1983: 134–135; F–I from Pringle 13544, WIS). Scale bars: A, I=6 cm; B–E, J–L= 5 mm; F=2 mm; G=0.5 mm; H=8 mm. 29 Lulia nervosa (Less.) Zardini. A Habit. B Marginal corolla. C Staminode. D Central floret (without pappus). E Stamen. F Upper part of the style. G Cypsela with pappus. (Redrawn from Zardini, 1975: 656). Scale bars: A=6 cm; B=7 mm; C=5 mm; D, E, G=4 mm; F=1 mm. 30 Lycoseris trinervis (Ruiz & Pav. ex D. Don) S. F. Blake. A Habit (branch). L. crocata (Bertol.) S. F. Blake. B–F Male capitulum. B Marginal corolla. C Staminode. D Central corolla. E Stamen. F Upper part of the style. G–L Female capitulum. G Marginal corolla. H Staminode of the marginal floret. I Central corolla. J Staminode of central floret. K Upper part of the style. L Cypsela of central floret with pappus. (A from Woytkowski 7402, US; B–F from Araque Molina & Barkley 19AT063, LP; G–L Bro. Paul 931, LP). A=4 cm; B, D, G, I=5 mm; C, E= 2 mm; F, H, J, K=1 mm; L=7.5 mm

568

L. Katinas, et al.

designated by Cabrera, 1970: 52, K not seen). Additional specimens examined: Argentina. Prov. Corrientes: Paso Troncón, Palacios-Cuezzo 2304 (LP); Uruguay. Dept. Artigas: Río Cuareim, J. Chebataroff 3015 (LP*). Literature. Cabrera (1970), Roque and Hind (2001), Freire et al. (2002). Leibnitzia Cass. Dict. sci. nat. 25: 420. Nov. 1822. TYPE: Tussilago anandria L. [= Leibnitzia anandria (L.) Turcz.]. 28 of Figs. 27–30. Anandria Less., Linnaea 5: 241. 1830, nom. illeg. Cleistanthium Kunze, Bot. Zeitung (Berlin) 9: 40. 1851. TYPE: Cleistanthium nepalense Kunze [= Leibnitzia nepalensis (Kunze) Kitam.]. Etymology. In honor of the German scientist Gottfried Wilhelm Leibnitz (1646–1716), an eminent eighteenth century philosopher and mathematician interested in botany. Herbs perennial, acaulescent, scapose, with plant dimorphism: a vernal (spring) stage with reduced or no leaves and chasmogamous capitula, and an autumnal (summer and fall) stage with fully developed leaves and cleistogamous capitula. Leaves rosulate; pseudopetiolate, blades lanceolate, elliptical to obovate, pinnately veined, margin denticulate to lyrate, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform or radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic with a gradual transition from marginal to central florets, marginal florets female, corolla true ray, filiform to bilabiate with a minute inner lip, outer lip expanded or not, with or without staminodes; central florets bisexual, bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Leibnitzia occimadrensis G. L. Nesom. Pollen spheroidal, circular in polar view, medium size, P × E = (35–42×36–40) μm. Tricolporate, colpi long with microgranulate membrane. Exine Mutisia type, microechinate, 7 μm thick at the equator and 4–5 μm thick at the poles, ectosexine slightly columellate. Ratio ectosexine/endosexine: 1:2. Nexine ca. 2 μm thick (see also Lin et al., 2005). Habitat and Distribution. Genus of six species distributed in North and Central America and Asia. Species list 1. Leibnitzia anandria (L.) Turcz., Ukaz. Otkryt. 8: 404. 1831. Basionym: Tussilago anandria L. TYPE: Sine loc., s.d., s. leg. (holotype, LINN 995 not seen). Additional specimen examined: Russia. Siberia, Tuva Autonomous Republic, near Lake Chagiti and Tanv Ola mtn. range, T. Elias et al. 8183 (US).


569

2. Leibnitzia knorringiana (B. Fedtsch.) Pobed., Fl. U.S.S.R. 28: 594. 1963. Basionym: Gerbera knorringiana B. Fedtsch. TYPE: Soviet Central Asia. Tien Shan: Lake Sary-Chilek, s.d., s. leg. (type, probably LE). 3. Leibnitzia nepalensis (Kunze) Kitam., J. Jap. Bot. 14: 297. 1938. Basionym: Cleistanthium nepalense Kunze. TYPE: Sine loc., Flor. Aug. 1850, s. leg. (holotype, based on a cultivated specimen in LZ not seen). 4. Leibnitzia occimadrensis G. L. Nesom, Brittonia 35: 130. 1983. TYPE: Mexico. State Sinaloa: Sierra Surutato, Los Pucheros, H. S. Gentry 7189 (isotype, GH!). 5. Leibnitzia ruficoma (Franch.) Kitam., J. Jap. Bot. 14: 297. 1938. Basionym: Gerbera ruficoma Franch. TYPE: China. Che-tcho-tze, 10 Oct 1882, Delavay s.n. (holotype, P? not seen). Additional specimen examined: China. Sikang Province: Sacred Mt., Karwar-kar-boo, Tsa-wa-rung, C. W. Wang 66451 (GH). 6. Leibnitzia seemannii (Sch. Bip.) G. L. Nesom, Brittonia 35: 127. 1983. Basionym: Chaptalia seemannii Sch. Bip. TYPE: Mexico. Durango: Sierra Madre, Seemann 1988 (isotype, K not seen, K photo at LP!); Federal District, Cima Station, C. G. Pringle 13544 (WIS!).

Literature. Nesom (1983), Hansen (1988).

Lulia Zardini Bol. Soc. Argent. Bot. 19: 255. 1980. TYPE: Trichocline nervosa Less. [= Lulia nervosa (Less.) Zardini]. 29 of Figs. 27–30. Etymology. In honor of the Spanish botanist, resident in Argentina, Angel Lulio Cabrera (1908–1999) who made enormous contributions to the study of Asteraceae. Herbs caulescent, perennial. Leaves alternate; sessile; blades linear, graminiform, long, parallel-veined, margin entire, glabrous. Capitulescences monocephalous, terminal; capitula long-pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with well developed staminodes, corolla bilabiate, with an expanded outer lip; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bilobed, branches dorsally papillose. Cypselae rostrate at the apex, pubescent; pappus with barbellate, capillary bristles. Pollen Description (Tellería & Katinas, 2004) (98 of Figs. 95–100). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate. Habitat and Distribution. Monotypic genus endemic to Sphagnum L. bogs in southern Brazil.

570

L. Katinas, et al.

Species list 1. Lulia nervosa (Less.) Zardini, Bol. Soc. Argent. Bot. 19: 257. 1980. Basionym: Trichocline nervosa Less. TYPE: Brazil. State São Paulo: Campos da Bocaina, A. F. M. Glaziou 8110 (neotype, designated by Zardini, 1980: 258, P not seen). Additional specimen examined: Brazil. State Santa Catarina: Mun. Campo Alegre, fazenda de Ernesto Scheide, S. B. Smith & R. M. Klein 10576 (LP*).

Literature. Zardini (1975, 1980).

Lycoseris Cass. Dict. sci. nat. 33: 463. 1824. TYPE: Atractylis mexicana L. f. [= Lycoseris mexicana (L. f.) Cass.]. 30 of Figs. 27–30. Diazeuxis D. Don, Trans. Linn. Soc. London 16: 251. 1830. TYPE: Diazeuxis mutisiana D. Don [= Lycoseris mexicana (L. f.) Cass.]. Etymology. From the Greek lykos, wolf, and seris, chicory, lettuce. Subshrubs dioecious. Leaves alternate; subsessile; blades elliptic to ovate, threeveined, margin entire to minutely dentate, glabrescent to glabrous above, floccose to dense-arachnoid beneath. Capitulescences monocephalous or paniculate to corymbose, terminal; capitula pedunculate, homogamous or heterogamous, radiate; receptacle epaleate, setose; involucre multiseriate. Florets of female capitula dimorphic, with staminodes, style sometimes widened above, bifid, branches dorsally papillose; marginal florets neuter, corolla true ray three-dentate, bilabiate to sub-bilabiate (3+1, 4+1 corolla lips), style rudimentary; central florets female, corolla actinomorphic, narrowly tubular, shallowly five-lobed, or sub-zigomorphic with unequal insicions; male capitula with florets dimorphic; marginal florets neuter, corolla true ray threedentate to sub-bilabiate (3+1 corolla lips), with staminodes, style rudimentary, branches dorsally papillose; central florets functionally male, corolla tubular-bilabiate to actinomorphic, tubular-funnelform, shallowly five-lobed, anther apical appendage acute, tails smooth, style widened above, dorsally papillose, papillae longer than in the female capitula. Cypselae glabrous; pappus of numerous, scabrid bristles in female florets, and flatter, seta-like, and relatively few scabrid bristles in male florets. Pollen Description. Lycoseris trinervis (Ruiz & Pav. ex D. Don) S. F. Blake (99 of Figs. 95–100). Pollen subprolate–prolate, elliptic in equatorial view, circular in polar view, medium to large size, P × E = (47–55×33–39.5) μm. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, microechinate, 5–6 μm thick, slightly slender at the poles. Ratio ectosexine/endosexine: 1:1; 1.5:1. Nexine ca. 1 μm thick (see also Egeröd & Stähl, 1991). Habitat and Distribution. Genus of 11 species of tropical and subtropical America, from Guatemala to Brazil and Bolivia.


571

Species list 1. Lycoseris boliviana Britton, Bull. Torrey Bot. Club 19: 266. 1892. TYPE: Bolivia. Dept. La Paz: Guanay, H. H. Rusby 1557 (lectotype, designated by Egeröd & Ståhl, 1991: 569, NY not seen, NY photo at LP!) 2. Lycoseris colombiana K. Egeröd, Nordic J. Bot. 11: 564. 1991. TYPE: Colombia. Dept. Cauca: El Bordo, Archer 3388 (holotype, US not seen). 3. Lycoseris crocata (Bertol.) S. F. Blake, Bull. Torrey Bot. Club 53: 218. 1926. Basionym: Aster crocatus Bertol. TYPE: Guatemala. Vulcano d’ acqua, 1837, Velásquez s.n. (holotype, BOLO not seen). Additional specimens examined: Colombia. Dept. Atlántico: cerca a Usiacurí, J. Araque Molina & F. A. Barkley 19 At063 (LP); Puerto Colombia, Bro. Paul 931 (LP). 4. Lycoseris eggersii Hieron., Bot. Jahrb. Syst. 29: 75. 1901. TYPE: Ecuador. Prov. Manabí: El Recreo, H. F. A. Eggers 14952 (isotype, LP!). Additional specimens examined: Ecuador. Prov. Manabí: south of Santa Ana, O. Haught 3462 (LP). Bolivia. Dept. Santa Cruz: Prov. Velazco, San Miguelito, 23 km N de San Ignacio, A. Krapovickas & A. Schinini 32419 (LP). 5. Lycoseris grandis Benth., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn: 111. 1852. TYPE: Costa Rica. San Matheo, A. S. Öersted 10517 (isotype, US not seen, US photo at LP!). 6. Lycoseris mexicana (L. f.) Cass., Dict. sci. nat. 33: 474. 1824. Basionym: Atractylis mexicana L. f. TYPE: Sine loc., s.d., J. C. Mutis s.n. (holotype, LINN 971 not seen). 7. Lycoseris minor K. Egeröd, Nordic J. Bot. 11: 559. 1991. TYPE: Colombia. Dept. Huila: Cordillera Oriental, valley of Río Fortalecillas, 32–33 km of Neiva, F. R. Fosberg 19778 (holotype, US not seen, US photo at LP!). 8. Lycoseris peruviana K. Egeröd, Nordic J. Bot. 11: 564. 1991. TYPE: Peru. Dept. San Martín: Pasaraya, near Saposoa, F. Woytkowski 5080 (isotype, NY not seen, NY photo at LP!). 9. Lycoseris retroflexa J. Kost., Blumea 5: 664. 1945. TYPE: Bolivia. Dept. Santa Cruz: near Paliza, T. Herzog 1263 (holotype, L not seen). 10. Lycoseris trinervis (Ruiz & Pav. ex D. Don) S. F. Blake, Contr. U. S. Natl. Herb. 22: 653. 1924. Basionym: Diazeuxis trinervis Ruiz & Pav. ex D. Don. TYPE: Ecuador. Huayaquil, s.d., Tafalla s.n. (lectotype, designated by Egeröd & Ståhl, 1991: 559, BM not seen). Additional specimens examined: Peru. Junín: San Ramón, Chanchamayo, L. Diers 1249 (LP*); id., Woytkowski 7402 (US). 11. Lycoseris triplinervia Less., Linnaea 5: 257. 1830. TYPE: Colombia. Santa Marta, von Rohr 27 (holotype, C not seen). Additional specimen examined: Venezuela. State Barinas: 80 km from Barinas, along road to San Cristóbal, bank of Río Bumbum, F. J. Breteler 4414 (LP).

Observation. Lycoseris displays some style characters that are not common in the tribe Mutisieae such as apically somewhat widened styles and style branches of male capitula covered by long papillae. These modifications of the style are presumably

572

L. Katinas, et al.

due to the dioecious condition of this genus. The pollen features, on the other hand, reveal the typical structure of most members of the tribe Mutisieae. Literature. Egeröd and Stähl (1991), Díaz-Piedrahita and Vélez-Nauer (1993), Ferreyra (1995).

Macroclinidium Maxim. Bull. Acad. Imp. Sci. Saint-Pétersbourg, ser. 3, 15: 375. 1871. TYPE: Macroclinidium robustum Maxim. 31 of Figs. 31–34. Pertya sect. Macroclinidium (Maxim.) Makino, Bot. Mag. (Tokyo) 14: 144. 1900. TYPE: Macroclinidium robustum Maxim. Etymology. From the Greek makros, long, large, and cline, a bed, klinidion, little bed, probably referring to the large receptacle. Herbs perennial, unbranched. Leaves clustered near the median part of the stem; sessile to long-petiolate; blades ovate, obovate, three-lobed, palmately, pinnately, or three-veined to subpalmately veined, margin dentate to serrate, glabrous to slightly pilose especially beneath. Capitulescences spicate to paniculate; capitula sessile, homogamous, ligulate, one- to ten-flowered; receptacle epaleate, glabrous to setose; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Macroclinidium (100 of Figs. 95–100). Pollen spheroidal or suboblate, circular in polar view, medium or large size. Tricolporate, colpi very short with microgranulate membrane. Exine Macroclinidium type, equally thickened in the whole grain. Note: the Macroclinidium exine type is similar to the Cardueae exine type; they differ in the morphology of the spines, being solid in the Macroclinidium type and with an apical channel in the Cardueae type. Macroclinidium rigidulum (Miq.) Makino. Pollen spheroidal, large size, P × E = (51–60×52–56) μm. Colpi ca. 23 μm length. Exine ca. 14 μm thick in the whole grain, spines ca. 5 μm length. Nexine 2–2.5 μm thick. Macroclinidium robustum. Pollen suboblate, medium size, P × E = (44–50×50–64) μm. Colpi ca. 16 μm length, with psilate membrane. Exine 9–12 μm thick, spines 4–5 μm length. Nexine ca. 1.5 μm thick (see also Lin et al., 2005). Habitat and Distribution. Genus of six species endemic to Japan. Species list 1. Macroclinidium hybridum (Makino) Matsum., Index pl. jap. 3: 658. 1912. Basionym: Pertya hybrida Makino. TYPE: Japan. Musashi: Omiya-hachiman in Wada-mura, 1897, Makino s.n. (type, MAK not seen, MAK photo at LP!).


573

Figs. 31–34 31 Macroclinidium robustum Maxim. A Habit. B Floret (without cypsela). C Stamen. D Cypsela with pappus. E Upper part of the style. (From Wawra 1527, NY). Scale bars: A=2 cm; B, D= 5 mm; C=2.5 mm; E=1.2 mm. 32 Mutisia speciosa Aiton ex Hook. A Habit (branch). B Marginal floret (without cypsela). C Staminode. D Central floret (without cypsela). E Stamen. F Upper part of the style. G Cypsela with pappus. (Redrawn from Cabrera, 1965: 51). Scale bars: A=4 cm; B, D, G=14 mm; C, F= 2 mm; E=9 mm. 33 Myripnois dioica Bunge. A Habit (branch). B Female floret. C Male floret. D Stamen. E Upper part of the style. (A from Wang 20106, NY; B–E from Sheehan s.n., NY). Scale bars: A=2 cm; B, C=5 mm; D, E=1.2 mm. 34 Neblinaea promontoriorum Maguire & Wurdack. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Aristeguieta, 1964: 918). Scale bars: A=3.5 cm; B, E=6 mm; C=2 mm; D=1 mm

574

L. Katinas, et al.

2. Macroclinidium koribanum, Nakai, Bot. Mag. (Tokyo) 26: 249. 1912. TYPE: Japan. Sendai: Taihakusan, 1901, K. Koriba s.n. (type, probably TI). 3. Macroclinidium rigidulum (Miq.) Makino, Bot. Mag. (Tokyo) 12: 194. 1898. Basionym: Eupatorium rigidulum Miq. TYPE: not localized. Additional specimens examined: Japan. Honshu: Ishikawa, Nunohashi-machi, Komatsushi, Tsugaru & Takeuchi 7324 (NY); id., Deguchi et al. 6382 (MO*). 4. Macroclinidium robustum Maxim., Bull. Acad. Imp. Sci. Saint-Pétersbourg, ser. 3, 15: 376. 1871. TYPE: Japan. Jokohama, Wawra 1527 (type, NY!). 5. Macroclinidium suzuki Kitam., J. Jap. Bot. 14: 385. 1938. TYPE: Japan. Hondo: Shirakawa, Kinshoji, 20 Oct 1930, S. Suzuki s.n. (type, probably TI). 6. Macroclinidium trilobum (Makino) Makino, Bot. Mag. (Tokyo) 7: 302. 1894; Bot. Mag. (Tokyo) 12: 194. 1989. Basionym: Ainsliaea triloba Makino. TYPE: not localized. Additional specimen examined: Japan. Rikugen, 1926, Masamune s.n. (NY).

Literature. Kitamura (1937), Ohwi (1965).

Mutisia L. f. Suppl. pl.: 57. 1781 (1782). TYPE: Mutisia clematis L. f. 32 of Figs. 31–34. Aplophyllum Cass., Dict. sci. nat. 33: 472. 1824. TYPE: not designated, based on the three species Mutisia decurrens Cav., M. ilicifolia Cav., and M. linearifolia Cav. Guariruma Cass., Dict. sci. nat. 33: 472. 1824. [= Mutisia sect. Guariruma (Cass.) Cabrera, Opera Lilloana 13: 138. 1965]. TYPE: Mutisia hastata Cav. (lectotype designated by Cabrera, 1965). Etymology. In honor of the Spanish botanist José Celestino Bruno Mutis y Bosio (1732–1808), physician, clergyman, and first director of the Real Expedición del Nuevo Reino de Granada. Shrubs or subshrubs stout to trailing, stem winged to unwinged. Leaves alternate; sessile to short-petiolate; blades simple to pinnately compound, with or without terminal tendrils, linear, subulate, lanceolate to ovate, pinnately to one-veined, margin entire to runcinate, sometimes revolute, glabrous to pubescent. Capitulescences monocephalous, terminal, erect to pendulous; capitula pedunculate, heterogamous, rarely homogamous, discoid or radiate; receptacle epaleate; involucre multiseriate, phyllaries erect to recurved downward. Florets dimorphic or rarely isomorphic; marginal florets female, with staminodes, corolla bilabiate, outer lip expanded or not, inner lip developed to very reduced, or true ray; central florets bisexual, corolla bilabiate; or all the florets bisexual, corolla bilabiate, sub-bilabiate (4+1 corolla lips), or ligulate; anther apical appendages acute, tails smooth; style bilobed, dorsally papillose. Cypselae truncate to attenuate at the apex, glabrous; pappus of plumose, capillary bristles.


575

Pollen Description. Mutisia (101 of Figs. 101–106) (Tellería & Katinas, 2004). The two species examined (M. kurtzii R. E. Fr. and M. retrorsa Cav.) have pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Cabrera, 1965; Skvarla & Turner, 1966; Heusser, 1971; Parra & Marticorena, 1972; Skvarla et al., 1977). Note: pollen of Mutisia is highly variable in size and exine thickness (Tellería & Katinas, in press). Habitat and Distribution. Genus of 63 South American species, from the Andes of Colombia to southern Argentina and Chile, also southern Brazil and adjacent regions. Species list 1. Mutisia acerosa Poepp. ex Less., Syn. gen. Compos.: 107. 1832. TYPE: Chile. Near Las Cruces et La Guardia de los Hornillos, E. F. Poeppig 26 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, El Maitén, C. Jiles 2451 (LP). 2. Mutisia acuminata Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 192. 1798. TYPE: Peru. Ceuchin, May 1779, s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Volcán, A. L. Cabrera et al. 13894 (LP). 3. Mutisia alata Hieron., Bot. Jahrb. Syst. 19: 73. 1894. TYPE: Ecuador. Inter Huasi-huaico et Contrayerba, Lehmann 4571 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cajamarca: Cajamarca, Huambos, Ochoa 1594 (LP). 4. Mutisia andersonii Sodiro ex Hieron., Bot. Jahrb. Syst. 29: 80. 1900. TYPE: Ecuador. Prov. Pichincha: Cerro Pichincha, E. Sodiro s.n. (neotype, designated by Harling, 1991: 74, W not seen). Additional specimen examined: Ecuador. Prov. Chimborazo: camino Río Bamba a Huamboya- De Púngala a Cusñipaccha, R. Scolnick 1532 (LP). 5. Mutisia araucana Phil., Anales Univ. Chile 41: 737. 1872. TYPE: Chile. Prov. Arauco: pinares de Nahuelbuta, s.d., H. Volckmann s.n. (holotype, SGO not seen, SGO photo at LP!). 6. Mutisia arequipensis Cabrera, Opera Lilloana 13: 171. 1965. TYPE: Peru. Dept. Arequipa: A. Weberbauer 6857 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Arequipa: Prov. Cailloma, distr. Coporaque, valle de Colca, ca. 100 km N de Arequipa, J. Treacy 840 (WIS). 7. Mutisia brachyantha Phil., Anales Univ. Chile 85: 825. 1894. TYPE: Chile. Andes, Talca, s.d., L. H. Silva s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Morro Blanco, C. Jiles 1590 (LP). 8. Mutisia burkartii Cabrera, Darwiniana 20: 210. 1976. TYPE: Argentina. Prov. Neuquén: Nuco Pehuén, 25 km al W de Loncopue, M. Gentili 215 (holotype, LP!). 9. Mutisia campanulata Less., Linnaea 5: 269. 1830. TYPE: Brazil: s.d., Sello s.n. (holotype, B not seen, B photo F n. 15925 at LP!). Additional specimen examined: Argentina. Prov. Misiones: El Paraíso, M. A. Torres 67 (LP).

576

L. Katinas, et al.

10. Mutisia cana Poepp. & Endl., Nov. gen. sp. pl. 1: 17. 1835. TYPE: Chile. Río Colorado, Santa Rosa, s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Santiago: Peñalolén, E. Navas 959 (LP). 11. Mutisia clematis L. f., Suppl. pl.: 373. 1781 (1782). TYPE: Habitat in Nova Granada, J. C. Mutis 21 (holotype, LINN 1004.1 not seen). Additional specimen examined: Colombia. Dept. Boyacá: Río Gomeza (Río Arzobispo), 7–10 km al E de Socha, F. R. Fosberg 22294 (LP). 12. Mutisia coccinea A. St.-Hil., Voy. distr. diam. 1: 386. 1833. TYPE: Brazil. State São Paulo: d’Hytu, St. Hilaire 1474bis (holotype, P not seen). Additional specimen examined: Argentina. Prov. Misiones: Loreto, G. Grüner 516 (LP). 13. Mutisia cochabambensis Hieron., Bot. Jahrb. Syst. 19: 74. 1894. TYPE: Bolivia. Cochabamba, M. Bang 881 (isotype, WIS). 14. Mutisia comptoniaefolia Rusby, Mem. Torrey Bot. Club 3: 65. 1893. TYPE: Bolivia. Talca Chugiaguillo, M. Bang 795 (isotype, WIS!). 15. Mutisia decurrens Cav., Icon. 5: 65. 1799. TYPE: Chile et Cordillera del Planchón, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquén: Cañadón de la Mosca, Pampa del Toro, G. Dawson 3287 (LP). 16. Mutisia discoidea Harling, Fl. Ecuador 42: 70. 1991. TYPE: Ecuador. Prov. Cañar: road Cañar-El Tambo, ca. 4 km from Cañar, J. Madsen 50281 (holotype, AAU not seen). 17. Mutisia friesiana Cabrera, Notas Mus. La Plata, Bot. 13: 19. 1948. TYPE: Argentina. Prov. Salta: Dept. Poma, Quebrada de Cobres, A. L. Cabrera 8716 (holotype, LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Susques, Quebrada de Potrero, L. Catalano 25 (LP). 18. Mutisia grandiflora Humb. & Bonpl., Pl. aequinoct. 1: 177. 1808 TYPE: Colombia. Andibus Quinduensibus, inter urbem Ibague et Carthago, s.d., F. Humboldt & A. Bonpland s.n. (holotype, P not seen). Additional specimen examined: Colombia. Dept. del Valle: cordillera central, Hoya del Río Bugalagrande, Loma de Barragán, entre la población y Albania, J. Cuatrecasas 20917 (F). 19. Mutisia hamata Reiche, Anales Univ. Chile 115: 103. 1904. TYPE: Chile. Prov. Tarapacá: Amincha, s.d., s. leg. (type, probably SGO). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, camino de El Angosto al Chañi, A. L. Cabrera et al. 22495 (LP). 20. Mutisia hastata Cav., Icon. 5: 64. 1799. TYPE: Peru. Cordillera del Planchón, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Peru. Dept. Lima: Prov. Canta, Lachaqui, Toma, G. Vilcapoma S. 1307 (LP). 21. Mutisia hieronymii Sodiro ex Cabrera, Notas Prelim. Mus. La Plata 3: 167. 1934. TYPE: Ecuador. Prov. Napo-Pastaza: Cuyuja-Baeza, Feb 1905, A. Sodiro s.n. (isotype, BAF not seen). 22. Mutisia homoeantha Wedd., Chlor. andina 1: 21. 1855. TYPE: Bolivia. Chuquisaca, M. Weddell 4075 (holotype, P not seen). Additional specimen examined: Argentina. Prov. Salta: Dept. Santa Victoria, Lizoite, T. Meyer 33143 (LP). 23. Mutisia ilicifolia Cav., Icon. 5: 63. 1799. TYPE: Chile. Prov. Coquimbo: Coquimbo, s.d., M. Nee s.n. (lectotype, designated by Cabrera, 1965: 198, MA


24.

25.

26.

27.

28.

29.

30.

31.

32.

33.

34.

577

not seen, MA photo at LP!). Additional specimen examined: Chile. Prov. Curicó: Cordillera del Planchón, E. Barros 1751 (LP). Mutisia intermedia Hieron., Bot. Jahrb. Syst. 19: 72. 1894. TYPE: Colombia. Nariño: cerca de Túquerres, s.d., J. Idrobo & M. Ospina 2421 (neotype, designated by Díaz-Piedrahita & Vélez-Nauer, 1993: 96, COL not seen). Mutisia involucrata Phil., Linnaea 28: 710. 1856. TYPE: Chile in collibus subandinis, Talcaregue, Feb 1831, C. Gay 786 (type, probably P). Additional specimen examined: Chile. Prov. Curicó: Teno, La Montaña, E. Barros 3804 (LP). Mutisia kurtzii R. E. Fr., Nova Acta Regiae Soc. Sci. Upsal. 1: 94. 1905. TYPE: Argentina. Prov. Jujuy: Tilcara, Quebrada de Juella, F. Claren 11732 (isotype, BAF not seen, BAF photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, cerro de Zenta, Feb 1931, E. Budin s.n. (LP*). Mutisia lanata Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 192. 1798. TYPE: Peru: s.d., s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Bolivia. Dept. Yungas: Tusilmaya a Chlumani, R. Scolnick & R. Luti 519 (LP). Mutisia lanigera Wedd., Chlor. andina 1: 16. 1855. TYPE: Bolivia. Provincie de Carangas, C. D’Orbigny 1341 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Toconce, M. Ricardi & C. Marticorena 4677/1062 (LP). Mutisia latifolia D. Don, Trans. Linn. Soc. London 16: 270. 1830. TYPE: Chile. Prov. Valparaíso: Valparaíso, s.d., D. Bridges s.n. (holotype, G not seen, G photo F n. 28874 at LP!). Additional specimen examined: Chile. Prov. Santiago: Cuesta La Dormida, G. Looser 1323 (LP). Mutisia ledifolia Decne. ex Wedd., Chlor. andina 1: 20. 1855. TYPE: Bolivia. Dept. Potosí: cumbre de la Cuesta de Lagunillas, C. D’Orbigny 1272 (lectotype, designated by Cabrera, 1965: 134, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Rinconada, Quebrada de Fundiciones, A. L. Cabrera 9438 (LP). Mutisia lehmannii Hieron., Bot. Jahrb. Syst. 19: 73. 1894. TYPE: Ecuador. Prov. Azuay: Cerro Soldados, W of Sayausí, G. Harling et al. 8743 (neotype, designated by Harling, 1991: 67, GB not seen). Additional specimen examined: Ecuador. Prov. Azuay: area nacional de Recreacion “Cajas,” Sayausi-Molleturo road W of Cuenca, L. J. Dorr & I. Valdepino 6382 (F). Mutisia linearifolia Cav., Icon. 5: 66. 1799. TYPE: Chile. Cordillera del Planchón, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquén: Cordillera del Viento, J. Frenguelli 521 (LP). Mutisia linifolia Hook., Bot. misc. 1: 12. 1830. TYPE: Argentina. Prov. Mendoza: camino de las Minas de Uspallata, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Quebrada del Toro, 1934, A. Pérez Moreau (LP). Mutisia lutzii G. M. Barroso, Mem. Inst. Oswaldo Cruz 53: 357. 1955. TYPE: Brazil. State Minas Geraes: Serra do Caparao, A. Lutz 1200 (holotype, R not seen).

578

L. Katinas, et al.

35. Mutisia macrophylla Phil., Anales Univ. Chile 18: 62. 1861. TYPE: Chile. Entre Lota y Chillán, s.d., Pearce s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Chillán: San Carlos, San Fabián, E. Barros 1772 (LP). 36. Mutisia magnifica C. Ulloa & P. Jørg., Novon 6: 131. 1996. TYPE: Ecuador. Loja: Cerro Uritusinga, Loja-La Palma, km 18.4, P. M. Jörgensen et al. 1010 (holotype, MO!). 37. Mutisia mandoniana Wedd. ex Cabrera, Opera Lilloana 13: 163. 1965. TYPE: Bolivia. Prov. Larecaja: inter Sorata et Lacatia, G. Mandon 8 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Bolivia. Near Tarcacama, Larecaja, M. Cardenas 4869 (F). 38. Mutisia mathewsii Hook. & Arn., Comp. Bot. Mag. 1: 107. 1835. TYPE: Peru: s.d., Mathews 1119 (isotype, P not seen, P photo F n. 38030 at LP!). Additional specimen examined: Peru. Dept. Junín: Yauli, Macbride & Featherstone 907 (LP). 39. Mutisia microcephala Sodiro ex Cabrera, Notas Prelim. Mus. La Plata 3: 168. 1934. TYPE: Ecuador. Cuyuja-Baeza, Feb 1905, A. Sodiro s.n. (isotype, BAF not seen, BAF photo at LP!). Additional specimen examined: Ecuador. Prov. Imbabura: Lake Cuicocha, C. W. Penland & R. H. Summers 731 (F). 40. Mutisia microneura Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 121. 1953. TYPE: Ecuador. Prov. Chañar: Tipacocha, C. W. Penland & R. H. Summers 1007 (holotype, F not seen, F photo at LP!). 41. Mutisia microphylla Willd. ex DC., Prodr. 7: 6. 1838. TYPE: Ecuador. Inter Quito et Pichincha, s.d., F. Humboldt & A. Bonpland s.n. (holotype, G not seen). Additional specimen examined: Ecuador. Prov. Pichincha: faldas de Pichincha, C. Firmin 524 (F). 42. Mutisia ochroleuca Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 122. 1953. TYPE: Colombia. Dept. Nariño: Volcan de Chiles, Tambo, NW shoulder of Volcan de Chiles, J. A. Ewan 16089 (holotype, F!). 43. Mutisia oligodon Poepp. & Endl., Nov. gen. sp. pl. 1: 18. 1835. TYPE: Chile. Chile meridionalis, Antuco, Castillo de Ballenar, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Río Negro: Bariloche, L. R. Parodi 11564 (LP). 44. Mutisia orbignyana Wedd., Chlor. andina 1: 22. 1855. TYPE: Bolivia. St. Bartolo, C. D’Orbigny 1387 (holotype, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Valle Grande, Caspalá, A. Burkart & N. S. Troncoso 11700 (LP). 45. Mutisia pulcherrima Muschl., Bot. Jahrb. Syst. 50: 102. 1913. TYPE: Peru. Prov. Chachapoyas: inter Tambos Bajazan et Almirante, A. Weberbauer 4444 (holotype, B not seen, B photo F n. 15495 at LP!). Additional specimen examined: Peru. Prov. Chachapoyas: Dept. Amazonas, Jalca de Calla-Calla, A. Sagástegui A. 7469 (LP). 46. Mutisia rauhii Ferreyra, Bol. Soc. Peruana Bot. 8: 75. 1980. TYPE: Peru. Dept. Cuzco: al E de la Cordillera de Salcantay, W. Rauh 1451 (holotype, USM!). 47. Mutisia retrorsa Cav., Icon. 5: 65. 1799. TYPE: Argentina. Prov. Santa Cruz: Puerto Deseado, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Argentina. Prov. Santa Cruz: Lago Argentino, Jan 1902, R. Hauthal s.n. (LP).


579

48. Mutisia rimbachii Sodiro ex S. K. Harris, Rhodora 35: 257. 1933. TYPE: Ecuador. Andes, 1850–1857, R. Spruce 5456 (lectotype, designated by Harling, 1991: 59, BM not seen; isolectotype, NY not seen, NY photo at LP!). Additional specimen examined: Ecuador. Prov. Tungurahua: Mocha, alrededores del pueblo, P. Villacres 234 (F). 49. Mutisia rosea Poepp. ex Less., Syn. gen. Compos.: 106. 1832. TYPE: Chile. Inter Las Cruces and Guardia de los Hornillos, E. F. Poeppig 113 (27) (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Santiago: San Gabriel, G. Montero O. 515 (F). 50. Mutisia saltensis Cabrera, Darwiniana 9: 373. 1951. TYPE: Argentina. Prov. Salta: Dept. Orán, Cerro Santa Bárbara, Carmelich et al. 190 (holotype, LP!). 51. Mutisia sinuata Cav., Icon. 5: 66. 1799. TYPE: Chile. Cordilleras del Planchón y Portillo, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Tunuyán, Valle del Alto Tunuyán, A. Ruiz Leal 2146 (LP). 52. Mutisia sodiroi Hieron., Bot. Jahrb. Syst. 29: 79. 1900. TYPE: Ecuador. Prov. Pichincha, Cerro Pichincha, s.d., Sodiro s.n. (neotype, designated by Harling, 1991: 62, G not seen). Additional specimen examined: Colombia. Dept. Nariño: cordillera occidental, Oscurana, on S slope of Volcan Gualcalá, 14 km E of Piedrancha, F. R. Fosbert 21188 (F). 53. Mutisia speciosa Aiton ex Hook., Bot. Mag. 54: 2705. 1827. TYPE: Brazil: s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Misiones: Arroyo Chapa, San Ignacio, G. Grüner 1077 (LP). 54. Mutisia spectabilis Phil., Linnaea 33: 107. 1864. TYPE: Chile. Prov. Aconcagua: Calabozo, Catemu, Sept 1860, R. A. Philippi s.n. (holotype, SGO not seen, SGO photo at LP!). 55. Mutisia spinosa Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 193. 1798. TYPE: Chile. Arauco, Colo-Colo, s.d., s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Argentina. Prov. Río Negro: Bariloche, A. L. Cabrera & M. M. Job 51 (LP). 56. Mutisia splendens Renjifo, Anales Univ. Chile 65: 299. 1884. TYPE: Chile: Dept. Quillota, Vichiculen, Dec 1881, C. Renjifo (type, probably SGO). Additional specimen examined: Chile. Prov. Santiago, Claude-Joseph 974 (US). 57. Mutisia stuebelii Hieron., Bot. Jahrb. Syst. 21: 367. 1895. TYPE: Colombia. Cerro Munchique, cerca de Popayán, A. Stübel 318a (holotype, B not seen, B photo F n. 15951 at LP!). Additional specimen examined: Colombia. Dept. del Cauca: cordillera central, cabeceras del río Palo, quebrada de Santo Domingo, campamento de los quineros, J. Cuatrecasas 19156 (F). 58. Mutisia subspinosa Cav., Icon. 5: 64. 1799. TYPE: Peru. Guamauranga, Planchón, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Estancia San Isidro, cerca del Mogote Aspero, A. Ruiz Leal 22532 (LP). 59. Mutisia subulata Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 193. 1798. TYPE: Chile. Concepción, s.d., s. leg. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquén: Cordillera del Viento, J. Frenguelli 516 (LP).

580

L. Katinas, et al.

60. Mutisia tridens Poepp. ex Less., Syn. gen. Compos.: 105. 1832. TYPE: Chile. Prov. Bío Bío: Antuco, E. F. Poeppig 220 (isotype, NY not seen, NY photo at LP!). 61. Mutisia venusta S. F. Blake, Proc. Biol. Soc. Wash. 46: 191. 1933. TYPE: Peru. Cuzco, Piñasniocj, Panticalla Pass, O. F. Cook & G. B. Gilbert 1235 (holotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: Prov. Urubamba, entre Piri y Peñas, C. Vargas C. 4420 (LP). 62. Mutisia vicia J. Kost., Blumea 5: 670. 1945. TYPE: Bolivia. Comarapa, T. Herzog 1974 (isotype, LP!). Additional specimen examined: Bolivia. Dept. Cochabamba: Pocona, M. Cárdenas 2256 (LP). 63. Mutisia wurdackii Cabrera, Opera Lilloana 13: 43. 1965. TYPE: Peru. Dept. Amazonas: Prov. Bongará: 2–4 km W–SW of Pomacocha, J. J. Wurdack 840 (holotype, LP!). Additional specimen examined: Peru. Dept. Amazonas: Prov. Chachapoyas, entre Ingenio y Pomacocha, A. López et al. 4303 (LP). Literature. Cabrera (1965, 1976), Ferreyra (1980), Harling (1991), Ulloa Ulloa and Jørgensen (1996); Tellaría and Katinas (in press). Myripnois Bunge Enum. pl. China bor.: 38. 1833 (1831). TYPE: Myripnois dioica Bunge. 33 of Figs. 31–34. Etymology. From the Greek myri, ointment or resin, and pnois, to smell, to have an odor, referring to the aromatic compounds in the plant. Shrubs dioecious. Leaves clustered on brachyblasts; subsessile; blades ovate-elliptic, palmately, pinnately to three-veined, margin entire, slightly pubescent on both surfaces. Capitulescences monocephalous, terminal or axillar; capitula sessile on the brachyblasts, homogamous, ligulate, five- to seven-flowered; receptacle epaleate; involucre uniseriate, phyllaries ca. 5. Florets isomorphic, male or female, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae pubescent; pappus of scabrid, capillary bristles, absent in male florets. Pollen Description (102 of Figs. 101–106). Pollen spheroidal–subprolate, spheroidal to elliptic in equatorial view, circular in polar view, medium size, P × E = (38–46×30– 36) μm. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, 5–6 μm thick. Ratio ectosexine/endosexine: ca. 1:1. Nexine 2.5 μm thick (see also Lin et al., 2005). Habitat and Distribution. Monotypic genus endemic to China. Species list 1. Myripnois dioica Bunge, Enum. pl. China bor.: 38. 1833 (1831). TYPE: not localized. Additional specimens examined: China. Hopei: Peking, Wang 20106 (NY); id., s.d., Sheehan s.n. (NY*).


581

Literature. Von Bunge (1835).

Neblinaea Maguire & Wurdack Mem. New York Bot. Gard. 9: 391. 1957. TYPE: Neblinaea promontoriorum Maguire & Wurdack. 34 of Figs. 31–34. Etymology. Named for Cerro de la Neblina. This mountain, which forms part of the border between Brazil and Venezuela, contains the tallest peak in Brazil. Shrubs. Leaves alternate, mostly clustered at stem apex; petiolate, petioles clasping; blades oblanceolate to spathulate, pinnately veined, margin entire, partially revolute, glabrous, rarely puberulent beneath. Capitulescences cymose, two- to nineheaded, subterminal to lateral, rarely monocephalous; capitula pedunculate, homogamous, bilabiate, two- to five-flowered; receptacle epaleate; involucre multiseriate, cylindrical, phyllaries glabrous. Florets isomorphic, bisexual, corolla bilabiate, glabrous; anthers tan to brown, often white at the apex, exserted, apical appendages acuminate, tails slightly papillose, connate with the tails of adjacent stamens; style bifid, branches dorsally smooth. Cypselae glabrous; pappus of slightly scabrid bristles, half are capillary, half somewhat paleaceous and longer. Pollen Description (103 of Figs. 101–106). Pollen subprolate, elliptic in equatorial view, circular in polar view, large size, P × E = (62–70×46–55) μm. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, microechinate, ca. 7 μm thick, ectosexine slightly columellate, slightly thickened at the poles. Ratio ectosexine/ endosexine: 1:2. Nexine ca. 2 μm thick. SEM: tectum perforate. Note: pollen similar to that of Chimantaea cinerea, C. huberi, Stifftia uniflora, Stenopadus campestris, S. colveii, and S. huachamacari. (see also Carlquist, 1957b; Tellería, 2008). Habitat and Distribution. Neblinaea is monotypic and endemic to Cerro de la Neblina, where it occurs on both sides of the Venezuelan–Brazilian frontier. Species list 1.

Neblinaea promontoriorum Maguire & Wurdack, Mem. New York Bot. Gard. 9: 392. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37009 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, N. T. Silva & Brazzao 60895 (NY*).

Literature. Pruski (1991, 1997).

Nouelia Franch. J. Bot. (Morot) 2: 67. 1888. TYPE: Nouelia insignis Franch. 35 of Figs. 35–38.

582

L. Katinas, et al.

Etymology. In honor of M. A. -A. Nouel (d. 1887), naturalist and director of the Musée d’Orléans, France. Shrubs or small trees. Leaves alternate; petiolate; blades ovate-elliptic, pinnately veined, margin entire, tomentose beneath. Capitulescences monocephalous, rarely two- to three-headed, terminal; capitula short-pedunculate, homogamous, radiate; receptacle epaleate; involucre multiseriate, six- to ten-seriate. Florets dimorphic, bisexual; marginal florets with corollas bilabiate, outer lip, expanded; central florets with corolla actinomorphic, tubular-campanulate, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bifid, branches dorsally smooth. Cypselae villose; pappus of scabrid, capillary bristles, the outermost shorter. Pollen Description. Pollen subprolate–prolate, elliptic in equatorial view, circular in polar view, large size, P × E = (62–69×40–50) μm. Tricolporate, colpi long with margin and psilate membrane, mesoaperturate. Exine Mutisia type, scabrate, 6– 8 μm, ectosexine slightly columellate, thick internal tectum. Ratio ectosexine/ endosexine: 2:1. Nexine ca. 2.5 μm thick. Habitat and Distribution. Monotypic genus endemic to Sichuan–Yun-nan region in China. Species list 1.

Nouelia insignis Franch., J. Bot. (Morot) 2: 68. 1888. TYPE: China. Yun-Nan: Tapin-tze, J. M. Delavay 2498, 619, 245 (syntypes, probably P). Additional specimen examined: China, Maire 2516 (NY*).

Observation. Most populations of Nouelia insignis are seriously threatened and are even at the brink of extinction in China (Heng, 1995). In recent decades, N. insignis has become endangered due to rapid habitat fragmentation. Most of the extant populations consist of fewer than 50 individuals, and no more than 5,000 individuals survive in the wild (Luan et al., 2006). Literature. Franchet (1888), Heng (1995), Freire et al. (2002).

Oldenburgia Less. Linnaea 5: 252. 1830. TYPE: Oldenburgia paradoxa Less. 36 of Figs. 35–38. Etymology. In honor of the Swedish soldier and traveler Franz Pehr Oldenburg (1740–1774) who accompanied Thunberg and Masson on some of their travels in the Cape in the early 1770s. Shrubs or small trees, often cushion-like shrubs. Leaves alternate, clustered at the end of the branches; subsessile; blades obovate to elliptic, coriaceous, pinnately veined, margin entire, revolute, glabrous above, tomentose beneath. Capitulescences


583

Figs. 35–38 35 Nouelia insignis Franch. A Habit (branch). B Marginal corolla. C Central corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (A, F redrawn from Franchet, 1888: plate II; B, C redrawn from Freire et al., 2002: 535; D, E from Maire 2516, NY). Scale bars: A=4 cm; B, C, F=8 mm; D=1 mm; E=0.6 mm. 36 Oldenburgia intermedia Bond. A Habit (branch). B Marginal floret (without cypsela). C Staminode. D Central floret (without cypsela). E Cypsela with pappus. O. grandis (Thunb.) Baill. F Stamen. G Upper part of the style. (A–E redrawn from Bond, 1987: 497; F, G from Humbert 10298, MO). Scale bars: A=2 cm; B, D, E=7 mm; C, F=4.5 mm; G=1 mm. 37 Onoseris speciosa Kunth. A Habit. O. alata Rusby. B Marginal floret (without cypsela). C Staminode. D Central floret (without cypsela). E Stamen. F Upper part of the style. G Cypsela with pappus. (A redrawn from Sancho, 2004: 438; B–G redrawn from Cabrera, 1978: 593). Scale bars: A=4 cm; B, D=9 mm; C, E=1.5 mm; F= 0.5 mm; G=6 mm. 38 Pachylaena atriplicifolia D. Don ex Hook. & Arn. A Habit (without rhizome). B Marginal floret (without cypsela). C Staminode. D Central floret (without cypsela). E Stamen. F Upper part of the style. G Cypsela with pappus. (A, B, D, G redrawn from Cabrera, 1971b: 328; C, E, F from Chicchi 67, LP). Scale bars: A=2 cm; B, D, G=10 mm; C, E=5 mm; F=0.5 mm

584

L. Katinas, et al.

monocephalous or two- to five-headed, corymbose; capitula sessile to pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate, ovoid, globoseurceolate to cyathiform. Florets dimorphic; marginal florets female, well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages long-apiculate, tails smooth; style bilobed, branches dorsally rugulose to papillose. Cypselae glabrous to villose; pappus of plumose to barbellate capillary bristles, widened at the apex. Pollen Description. Pollen not seen. Habitat and Distribution. Genus of four species from southern Africa (Cape Province). Species list 1. Oldenburgia grandis (Thunb.) Baill., Hist. pl. 8: 97. 1882. Basionym: Arnica grandis Thunb. TYPE: Cap, s.d., Masson s.n., herb. Thunb. (holotype, UPS microfiche not seen). Additional specimen examined: Africa. South Africa, H. Humbert 10298 (MO). 2. Oldenburgia intermedia Bond, S. African J. Bot. 53: 496. 1987. TYPE: Africa. South Africa: Bergriviernek, Jonkershoek, P. Bond 1723 (isotype, K not seen). Additional specimen examined: South Africa, Western Cape Province, Tulbagh District, Voëlvlei Mts., Saddle Mt. on the way to Ontongskpo from Voëlvlei, S. Ortiz 832a (SANT). 3. Oldenburgia papionum DC., Prodr. 7: 12. 1838. TYPE: Berge bei Tulbagh R III, Drège 1835 (lectotype, designated by Bond, 1987: 499, G not seen). Additional specimen examined: Africa. South Africa, Western Cape Province, Tulbagh District, Voëlvlei Mts., Saddle Mt. on the way to Ontongskpo from Voëlvlei, S. Ortiz 831a (SANT). 4. Oldenburgia paradoxa Less., Linnaea 5: 252. 1830. TYPE: Lectotype, designated by Bond, 1987: 496, the illustration in Linnaea 5: Table 3, Figs. 67–70 (1830). Observation. In the molecular analysis of Funk et al. (2005), Oldenburgia appears nested in the subfamily Carduoideae, sister to Tarchonantheae and Cardueae. Ortiz (2006b) suggested that it could be established as a tribe containing only the genus Oldenbugia. Our observations show that the species of this genus have the typical mutisioid style.

Literature. Bond (1987).

Onoseris Willd. Sp. pl. 3: 1702. 1803. TYPE: Atractylis purpurea L. f. [= Onoseris purpurea (L. f.) S. F. Blake]. 37 of Figs. 35–38.


585

Caloseris Benth., Pl. hartw.: 88. 1841. TYPE: Caloseris rupestris Benth. [= Onoseris onoseroides (Kunth) B. L. Rob.]. Cataleuca Koch & Fintelm., Wochenschr. Gärtnerei Pflanzenk. 2: 163. 1859, nom. nudum. Centroclinium D. Don, Trans. Linn. Soc. London 16: 254. 1830. TYPE: Centroclinium albicans D. Don [= Onoseris albicans (D. Don) Ferreyra]. Chaetachlaena D. Don, Trans. Linn. Soc. London 16: 256. 1830. TYPE: Chaetachlaena odorata D. Don [= Onoseris odorata (D. Don) Hook. & Arn.]. Cladoseris (Less.) Spach, Hist. nat. vég. 10: 35. 1841. TYPE: Cladoseris annua (Less.) Spach, based on Onoseris annua Less. Cursonia Nutt., Trans. Amer. Philos. Soc. 7: 422. 1841. TYPE: Cursonia peruviana Nutt. [= Onoseris odorata (D. Don) Hook. & Arn.]. Hilairia DC., Prodr. 7: 33. 1838, nom. nudum. Hipposeris Cass., Dict. sci. nat. 33: 464. 1824. [= Onoseris subgen. Hipposeris (Cass.) Less., Syn. gen. Compos.: 119. 1832]. TYPE: Onoseris salicifolia Kunth (lectotype designated by Sancho, 2004). Isotypus Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 8. 1818. TYPE: Isotypus onoseroides Kunth [= Onoseris onoseroides (Kunth) B. L. Rob.]. Pereziopsis J. M. Coult., Bot. Gaz. (London) 20: 52. 1895. TYPE: Pereziopsis donnell-smithii J. M. Coult. [= Onoseris donnell-smithii (J. M. Coult.) Ferreyra]. Rhodoseris Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 95. 1851. TYPE: Rhodoseris conspicua Turcz. [= Onoseris onoseroides (Kunth) B. L. Rob.]. Schaetzellia Klotzch, Allg. Gartenzeitung. 17: 82. 1849. TYPE: Schaetzellia deckeri Klotzsch [= Onoseris onoseroides (Kunth) B. L. Rob.]. Seris Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 1: 139. 1807. TYPE: Seris onoseroides Willd. [= Onoseris onoseroides (Kunth) B. L. Rob.]. Etymology. From the Greek onos, donkey, and seris, seridos, chicory, lettuce, probably because the rosulate leaves, lanceolate and tomentose beneath, which resemble the ears of a donkey. Herbs annual or perennial or subshrubs, branched to unbranched. Leaves alternate, often clustered near the base; sessile to long-petiolate; blades ovate to elliptic, linear, hastate to rarely orbicular, pinnately to palmately veined, margin entire to dentate, glabrescent above, densely pubescent beneath. Capitulescences monocephalous, terminal, sometimes on scapes, corymbose to paniculate; capitula heterogamous, radiate or discoid; involucre multiseriate; receptacle epaleate, pilose to setose. Florets usually dimorphic, when isomorphic, all slightly zygomorphic with corollas subbilabiate (4+1 corolla lips), lip not expanded; when dimorphic marginal florets female, with staminodes, corolla bilabiate to sub-bilabiate (3+1 corolla lips), outer lip expanded or not; central florets actinomorphic, tubular-funnelform, corolla shallowly five-lobed, lobes recurved, or slightly zygomorphic, five-lobed, lobes straight, with one or two deeper insicions; anther apical appendages acute, tails smooth to papillose, filaments usually papillose at the base; style bifid, branches dorsally papillose, sometimes glabrescent. Cypselae pubescent to glabrescent; pappus of all capillary, scabrid bristles, unequal in length, sometimes the inner ones longer, wider, and flattened.

586

L. Katinas, et al.

Pollen Description. Onoseris alata Rusby (Tellería & Katinas, 2004). Pollen subprolate–prolate, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen of O. odorata has polar caps. Habitat and Distribution. Genus of 31 species from southern Mexico to northwestern Argentina, with the highest concentration in the western slopes of the Andes of Peru and few species in Brazil. Species list 1. Onoseris acerifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 6. 1818. TYPE: Peru. Paramo de Yamoca, F. Humboldt & A. Bonpland 2568 (holotype, P not seen). Additional specimen examined: Bolivia. Santa Cruz, Steinbach 8200 (US). 2. Onoseris alata Rusby, Descr. S. Amer. pl.: 163. 1920. TYPE: Bolivia. Dept. La Paz: Coroico, Yungas, M. Bang 2365 (isotypes, US!, GH, K not seen, GH and K photos at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Humahuaca, M. Venturi 8359 (US). 3. Onoseris albicans (D. Don) Ferreyra, J. Arnold Arbor. 25: 389. 1944. Basionym: Centroclinium albicans D. Don. TYPE: Peru: s.d., H. Ruiz & J. A. Pavón s.n. (holotype, P not seen). Additional specimen examined: Peru. Apurimac, Vargas 5816 (LP). 4. Onoseris amplexicaulis Ferreyra, J. Arnold Arbor. 25: 364. 1944. TYPE: Peru. Dept. Lima: Matucana, J. F. Macbride & W. Featherstone 310 (isotypes US!, GH not seen, GH photo at LP!). Additional specimen examined: Peru. Dept. Lima: Matucana, J. F. Macbride & W. Featherstone 131 (F). 5. Onoseris annua Less., Linnaea 5: 341. 1830. TYPE: Peru. Lima, s.d., J. Dombey s.n. (isotypes, B, F not seen, B photo F n. 15887 in LP!). Additional specimen examined: Peru. Ancash, R. Ferreyra 7376 (US). 6. Onoseris brasiliensis Cabrera, Bol. Mus. Bot. Munic. 19: 1. 1975. TYPE: Brazil. Mato Grosso: Rondonopolis, G. Hatschbach 34719 (holotype, LP!). 7. Onoseris cabrerae Ferreyra, Bol. Soc. Argent. Bot. 19: 19. 1980. TYPE: Peru. Dept. Amazonas: Chachapoyas, R. Ferreyra & C. Acleto 15339 (holotype, USM!, USM photo LP!). 8. Onoseris castelnaeana Wedd., Chlor. andina 1: 10. 1855. TYPE: Peru. Cusco, 1847, M. F. Castelnau s.n. (holotype, P not seen, P photo F n. 38105 at LP!). Additional specimen examined: Peru. Apurimac, Vargas 404 (LP). 9. Onoseris chrysactinioides Sagást. & M. O. Dillon, Brittonia 37: 8. 1985. TYPE: Peru. Dept. Cajamarca: Contumazá, A. Sagástegui et al. 10839 (holotype, HUT!, HUT photo at LP!; isotype MO!). Additional specimen examined: Peru. Dept. Cajamarca: Contumazá, Becker & Terrones 1011 (US). 10. Onoseris costaricensis Ferreyra, J. Arnold Arbor. 25: 367. 1944. TYPE: Costa Rica. San Ramón, Brenes 6520 (holotype, F not seen). Additional specimen examined: Costa Rica. Pittier 10596 (US). 11. Onoseris cumingii Hook. & Arn., Comp. Bot. Mag. 1: 103. 1835. TYPE: Peru. Lima, Cuming 995 (holotype, K not seen, K photo at LP!). Additional specimen examined: Peru. Piura, O. Haught 30 (US).


587

12. Onoseris donnell-smithii (J. M. Coult.) Ferreyra, J. Arnold Arbor. 25: 368. 1944. Basionym: Pereziopsis donnell-smithii J. M. Coult. TYPE: Guatemala. Santa Rosa: Río de los Esclavos, E. T. Heyde & E. J. D. S. Lux 4527 (isotype, US!). Additional specimen examined: El Salvador. Santa Ana, Standley 19701 (US). 13. Onoseris drakeana André, Rev. Hort.: 181. 1882. TYPE: Colombia. Cauca, E. F. André 2917 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Colombia. Nariño, Uribe 5355 (US). 14. Onoseris fraterna S. F. Blake, J. Wash. Acad. Sci. 33: 368. 1943. TYPE: Bolivia. Dept. La Paz: Sur Yungas, B. A. Krukoff 10266 (holotype, US!). Additional specimen examined: Bolivia. Dept. La Paz: Sur Yungas, Fournet 472 (US). 15. Onoseris gnaphalioides Muschl., Bot. Jahrb. Syst. 50: 94. 1913. TYPE: Peru. Ancash, Caraz, A. Weberbauer 3012 (isotype, F not seen). Additional specimen examined: Peru. Ayacucho, Hjerting 1415 (LP). 16. Onoseris hastata Wedd., Chlor. andina 1: 9. 1855. TYPE: Bolivia. Dept. Chuquisaca: Monte Curi, H. A. Weddell 3763 (isotype, G not seen). Additional specimen examined: Argentina. Prov. Jujuy: Tumbaya, Volcán, A. L. Cabrera et al. 18286 (LP). 17. Onoseris humboldtiana Ferreyra, Publ. Mus. Hist. Nat. “Javier Prado,” Ser. B, Bot. 11: 1. 1959. TYPE: Peru. Ica: Nazca, Lomas de San Nicolás, R. Ferreyra 13400 (holotype, USM!, USM photo at LP!). 18. Onoseris hyssopifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 7. 1818. TYPE: Ecuador. Imbabura, F. Humboldt & A. Bonpland 2200 (holotype, P not seen; isotype, B not seen, B photo F n. 15893 at LP!). Additional specimen examined: Ecuador. Azuay, King 6648 (US). 19. Onoseris linearifolia Sagást., Phytologia 57: 415. 1985. TYPE: Peru. Dept. Cajamarca: Cajabamba, A. Sagástegui et al. 11264 (holotype, HUT!, HUT photo at LP!). Additional specimen examined: Peru. Dept. Cajamarca: Cajabamba, I. Sanchez Vega et al. 956 (US). 20. Onoseris lopezii Ferreyra, Publ. Mus. Hist. Nat. “Javier Prado,” Ser. B, Bot. 11: 4. 1959. TYPE: Peru. La Libertad: Santiago de Chuco, A. López Miranda 439 (holotype, USM!, USM photo at LP!). Peru. Cajamarca, Hudson 1073 (MO). 21. Onoseris macbridei Ferreyra, Publ. Mus. Hist. Nat. “Javier Prado,” Ser. B, Bot. 1: 2. 1949. TYPE: Peru. Cajamarca: Yamaluc, R. Ferreyra 846 (holotype, USM!, USM photo at LP!). Additional specimen examined: Peru. Ancash, López et al. 7473 (LP). 22. Onoseris minima Domke, Notizbl. Bot. Gart. Berlin-Dahlem 13: 247. 1936. TYPE: Peru. Dept. Arequipa: Socosani, D. B. Stafford 295a (holotype, K not seen). Additional specimen examined: Peru. Moquegua, A. Weberbauer 7440 (LP). 23. Onoseris odorata (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 103. 1835. Basionym: Chaetachlaena odorata D. Don. TYPE: Peru. Guayaquilia Peruvianorum, s.d., J. Tafalla s.n. (holotype, P not seen). Additional specimen examined: Peru. A. Weberbauer 1492 (US). 24. Onoseris onoseroides (Kunth) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Isotypus onoseroides Kunth. TYPE: Venezuela. Near Tui River, F. Humboldt & A. Bonpland s.n. (holotype, P not seen, P photo at LP!). Additional specimen examined: Colombia. Guajira, O. Haught 3937 (LP).

588

L. Katinas, et al.

25. Onoseris peruviana Ferreyra, J. Arnold Arbor. 25: 373. 1944. TYPE: Peru. Junín: La Merced, J. F. Macbride 5425 (holotype, US!; isotype, F not seen, F photo at LP!). 26. Onoseris purpurea (L. f.) S. F. Blake, Proc. Biol. Soc. Wash. 38: 85. 1925. Basionym: Atractylis purpurea L. f. TYPE: Colombia. “Nova Granada,” J. C. Mutis s.n. (syntype, US!). Colombia. Cundinamarca, Killip et al. 38365 (US). 27. Onoseris sagittata (Rusby) Rusby, Descr. S. Amer. pl.: 164. 1920. Basionym: Seris sagittatus Rusby. TYPE: Bolivia. Dept. Cochabamba: Toredon, M. Bang 1139 (isotypes, MO!, G not seen). Additional specimen examined: Bolivia. Samaipata, Fosberg 28652 (US). 28. Onoseris salicifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 7. 1818. TYPE: Ecuador. Chimborazo: Alausi, F. Humboldt & A. Bonpland 3226 (holotype, P not seen; isotype, B not seen, B photo F n. 15896 at LP!). Additional specimen examined: Ecuador. Chimborazo, O. Haught 3318 (LP). 29. Onoseris silvatica Greenm., Proc. Amer. Acad. Arts 40: 51. 1904. TYPE: Costa Rica. Alto del Rodeo, Pittier 1622 (holotype, GH not seen). Additional specimen examined: Colombia. Cundinamarca, King 6027 (US). 30. Onoseris speciosa Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 6. 1818. TYPE: Ecuador. Chimborazo, Alausi, A. Bonpland 3235 (isotype, F not seen). Additional specimen examined: Peru. Amazonas, King & Bishop 9197 (US). 31. Onoseris weberbaueri Ferreyra, J. Arnold Arbor. 25: 386. 1944. TYPE: Peru. Cajamarca: hills between Tabaconas and Marañon rivers, A. Weberbauer 6163 (holotype, F not seen, F photo n. 49246 at LP!). Additional specimen examined: Peru. Amazonas, Gentry et al. 61362 (MO). Literature. Ferreyra (1944, 1949, 1959, 1995), Cabrera (1975), Sagástegui Alva and Dillon (1985), Sancho (2004).

Pachylaena D. Don ex Hook. & Arn. Comp. Bot. Mag. 1: 106. 1835. TYPE: Pachylaena atriplicifolia D. Don ex Hook. & Arn. 38 of Figs. 35–38. Chionoptera DC., Prodr. 7: 14. 1838. TYPE: Chionoptera gayophyta DC. [= Pachylaena atriplicifolia D. Don ex Hook. & Arn.]. Etymology. From the Greek pachys, thick, stout, and chlaena, overcoat, blanket, because of the rigid phyllaries surrounding the florets, or with reference to the crass and thick leaves. Herbs postrate, caulescent to semi-acaulescent, perennial. Leaves nearly rosulate; pseudopetiolate; blades orbicular to spathulate, crass, rugose, pinnately veined, margin denticulate, glabrous. Capitulescences monocephalous, terminal; capitula


589

pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bilobed, dorsally papillose, papillae occasionally extending below the bifurcation point. Cypselae truncate at the apex, glabrous; pappus of plumose, capillary bristles. Pollen Description. Pachylaena atriplicifolia (104 of Figs. 101–106) (Tellería & Katinas, 2004). Pollen spheroidal–subprolate, medium to large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Urmenetea atacamensis Phil. (see also Parra & Marticorena, 1972). Habitat and Distribution. Genus of one species distibuted in the Andes of Argentina and Chile. Species list 1. Pachylaena atriplicifolia D. Don ex Hook. & Arn., Comp. Bot. Mag. 1: 106. 1835. TYPE: Argentina. Prov. Mendoza: near Agua del Ceno [Cerro] Pelado, on the ascent to El alto de los Manantides [Manantiales], s.d., J. Gillies s.n. (holotype, K not seen, K photo at LP!). Additional specimens examined: Argentina. Prov. Neuquén: Dept. Pehuenches, Cerro Chacaycó, A. Chicchi 67 (LP); id., Cerro Piedra Santa, 7 Feb 1944, J. Frenguelli s.n. (LP*).

Literature. Johnston (1929), Cabrera (1971b), Katinas (1996a, 2008). Perdicium L. Pl. rar. afr.: 22. 1760. TYPE: Perdicium capense L. 39 of Figs. 39–42. Idicium Neck., Elem. bot. 1: 28. 1790, nom. illeg. Pardisium Burm. f., Fl. indica: 26. 1768, an orthographic variant of the name Perdicium. Etymology. From the Greek perdikion, the name of a plant applied by Theophrastus to a species of Polygonum, perdikion is the diminutive of perdix, partridge. Herbs perennial, acaulescent, scapose. Leaves rosulate; pseudopetiolate; blades oblong to obovate, pinnately veined, margin sinuate to runcinate, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with staminodes, corolla bilabiate; central florets bisexual, corolla bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae constricted to shortly rostrate at the apex, glabrous to pubescent; pappus of scabrid, basally connate, capillary bristles.

590

L. Katinas, et al.

Figs. 39–42 39 Perdicium leiocarpum DC. A Habit. B Marginal corolla. C Staminode. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (From Drège s.n., MO 1981647). Scale bars: A=1 cm; B, D, G=2 mm; C, E, F=1 mm. 40 Pertya ovata Maxim. A Habit (branch). B Floret. C Stamen. D Upper part of the style. (From Takahashi et al. 1387, MO). P. sinensis Oliv. E Habit (branch). (From Sino-Amer. Exped. 1537, MO). Scale bars: A, E=2 cm; B=5 mm; C=2.5 mm; D=1.2 mm. 41 Plazia daphnoides Wedd. A Habit (branch). B Marginal floret (without cypsela). C Central floret (without cypsela). D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1978: 580). Scale bars: A=4 cm; B, F=10 mm; C=5 mm; D=4 mm; E=2 mm. 42 Salcedoa mirabaliarum F. Jiménez Rodr. & Katinas. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Jiménez Rodríguez et al., 2004: 993). Scale bars: A=2 cm; B, C=5 mm; D= 3 mm; E=10 mm


591

Pollen Description. Perdicium capense. Pollen spheroidal to prolate, spheroidal or elliptic in equatorial view, subcircular in polar view, large size, P × E = (50–65× 41–47) μm. Tricolporate, long colpi with microgranulate membrane. Exine Mutisia type, microechinate, 6–8 μm thick at the equator and 4–5 μm thick at the poles, ectosexine slightly columellate. Ratio ectosexine/endosexine: 1:1; 1:2; 1:3. Nexine ca. 2–3 μm thick. Habitat and Distribution. Genus of two species endemic to the western Cape in southern Africa. Species list 1. Perdicium capense L., Pl. rar. afr.: 22. 1760. TYPE: Sine loc., s.d., Burmann s.n. (holotype?, G not seen). Additional specimen examined: Africa. South Africa: terra Capensis, regio occidentalis, Tarkinsberg, Schlechter 8616 (MO). 2. Perdicium leiocarpum DC., Prodr. 7: 39. 1838. TYPE: Africa. Africa capensi, Camisberge, s.d., Drège s.n. (holotype, G-DC not seen). Additional specimen examined: Africa. s.d., Drège s.n. (MO). Literature. Hansen (1985c).

Pertya Sch. Bip. Bonplandia 10: 109. 1862. TYPE: Pertya glabrescens Sch. Bip. [= Pertya scandens Sch. Bip.]. 40 of Figs. 39–42. Etymology. In honor of Joseph Anton Maximilian Perty (1804–1884), German botanist and zoologist. Shrubs or subshrubs, stems branched. Leaves alternate or clustered on brachyblasts; sessile to petiolate; blades commonly ovate, oblong, elliptic, lanceolate to linear, palmately, pinnately to three-veined, margin entire to lobed, subglabrous to pubescent beneath. Capitulescences monocephalous, capitula terminal on the brachyblasts to paniculate; capitula pedunculate, homogamous, ligulate, one-, three- to 16-flowered; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae, glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Pertya (105 of Figs. 101–106). Pollen spheroidal to prolate, elliptic or subrectangular in equatorial view, circular in polar view, medium or large size. Tricolporate, colpi long with scabrate–microgranulate membrane. Exine Mutisia type, microechinate, equally thickened in the whole grain or slightly thickened at the poles. Nexine ca. 1.5 μm thick. Pertya cordifolia Mattf. Pollen spheroidal–subprolate, elliptic in equatorial view, medium size, P × E = (41–42×38– 40) μm. Exine 5.5–7 μm thick. Ectosexine with compact aspect. Ratio ectosexine/

592

L. Katinas, et al.

endosexine: 1:1.5. Pertya discolor Rehder. Pollen subprolate–prolate, elliptic in equatorial view, medium size, P × E = (37–43×29–33). Exine 5–6 μm thick. Ectosexine with granulate aspect or slightly columellate, thick internal tectum. Ratio ectosexine/endosexine: ca. 1.5:1. Pertya ovata Maxim. Pollen spheroidal–subprolate, subrectangular, medium to large size, P × E = (48–52×33–42) μm. Exine 6–7 μm thick, slightly thickened at the poles. Ratio ectosexine/endosexine: ca. 1:1.5 (see also Lin et al., 2005). Habitat and Distribution. Genus of ca. 22 species distributed in Afghanistan, China, Japan, Taiwan, and Thailand. Species list 1. Pertya aitchisoni C. B. Clarke, J. Linn. Soc., Bot. 18: 72. 1881. TYPE: Afghanistan. Kurrum Valley, Aitchison 392, 721 (syntypes, P!). 2. Pertya angustifolia Y. C. Tseng, Guihaia 5: 328. 1985. TYPE: China. Sichuan: Daofu, Songlinkou, Guan 50236 (type, SCBI not seen). 3. Pertya berberidoides (Hand.-Mazz.) Y. C. Tseng, Guihaia 5: 328. 1985. Basionym: Pertya bodinieri Vaniot var. berberidoides Hand.-Mazz. TYPE: China. Yunnan: Tonschieferfelsen de str. St. unter Meti über dem Yangtse sw von Dschungdien, s. leg. n. 7781 (type, WU not seen). 4. Pertya bodinieri Vaniot, Bull. Acad. Int. Géogr. Bot. 12: 116. 1903. TYPE: not localized. Additional specimen examined: China. Yunnan: Yun nan sen, Ducloux 257 (UC). 5. Pertya cordifolia Mattf., Notizbl. Bot. Gart. Berlin-Dahlem 11: 103. 1931. TYPE: China. Hunan: Yünshan, S. S. Sin n. S. H. 956 (type, probably B). Additional specimen examined: Kwansi, Steward & Cheo 972 (NY*). 6. Pertya corymbosa Y. C. Tseng, Guihaia 5: 332. 1985. TYPE: China. Guangxi: Rongshui, S. H. Chun 16893 (type, SCBI not seen). 7. Pertya desmocephala Diels, Notizbl. Bot. Gart. Berlin-Dahlem 9: 1032. 1926. TYPE: China. Chekiang: Lung chiung hsien, Hu 409 (type, probably M). 8. Pertya discolor Rehder, J. Arnold Arbor. 10: 135. 1929. TYPE: China. Gansu: en route to Linhua Shan, from Choni via Daozhou, J. F. Rock 12667 (type, S!). Additional specimen examined: Shansi, Chieh-hsiu distr., Mien-shan-ye, Smith 5786 (MO*). 9. Pertya henanensis Y. C. Tseng, Guihaia 5: 330. 1985. TYPE: China. Henan: Songxian County, Forest. Henan Exped. 973 (types, PE not seen, IBSC not seen). 10. Pertya hossei Craib ex Hosseus, Kew Bull.: 402. 1911. TYPE: Thailand. Doi Djieng Dao-Hochland, C. C. Hosseus 405 (type, MO!). 11. Pertya macrophylla Nakai, Icon. pl. koisikav. 4: 77. 1920. TYPE: Japan. Hondo: Sagami, prope Zushi, 19 Oct 1919, Nakai s.n. (type, TI not seen, TI photo at P!). 12. Pertya mattfeldii Bornm., Repert. Spec. Nov. Regni Veg. Beih. 108: 65. 1938. TYPE: not localized. Additional specimen examined: Afghanistan. Nuristan: between Pushuki and Kushimaicot, 1955, Kitamura s.n. (KYO). 13. Pertya monocephala W. W. Sm., Notes Roy. Bot. Gard. Edinburgh 8: 212. 1914. TYPE: China. Yunnan: Jalong, Oui-chu valley, F. Kingdon Ward 1141 (type, E not seen, E photo at LP!).


593

14. Pertya ovata Maxim., Bull. Acad. Imp. Sci. Saint-Pétersbourg. 16: 217. 1871. TYPE: Japan. Yokohama, 1862, Maximowicz s.n. (type, P!). Additional specimen examined: Honshu, Pref. Wakayama, near Mizugamine, Takahashi et al. 1387 (MO*). 15. Pertya phylicoides Jeffrey, Notes Roy. Bot. Gard. Edinburgh 5: 200. 1912. TYPE: China. Yunnan: from Zhongdian plateau to Yangzi, near Jiao-tou, G. Forrest 112 (type, E not seen). 16. Pertya pungens Y. C. Tseng, Guihaia 5: 334. 1985. TYPE: China. Guangdong: Yangchun, Jinshan, Wang 38778 (type, MO!). 17. Pertya scandens (Thunb. ex Murray) Sch. Bip., Bonplandia 10: 109. 1862. Basionym: Erigeron scandens Thunb. ex Murray. TYPE: Sine loc., Faurie 6756 (type of Pertya scandens (Thunb. ex Murray) Sch. Bip. f. schultziana Franch., P!). 18. Pertya simozawai Masam., Trans. Nat. Hist. Soc. Taiwan 30: 37. 1940. TYPE: not localized. Additional specimen examined: Taipei. Sanshia: Wuliaochien, Peng 8683 (TAI). 19. Pertya sinensis Oliv., Icon. pl. 23: pl. 2214. 1892. TYPE: not localized. Additional specimens examined: China. Hubei: Shennongjia Forerst, S of Jiuhuping Forest Farm along the Jizigou canyon bottom, Sino-Amer. Exped. 1537 (MO, NY). 20. Pertya tsoongiana Ling, Contr. Bot. Surv. N. W. China 1: 40. 1939. TYPE: not localized. Additional specimen examined: China. Szechuan: Fushan, T. P. Wang 10293 (PE). 21. Pertya uniflora (Maxim.) Mattf., Notizbl. Bot. Gart. Berlin-Dahlem 11: 105. 1931. Basionym: Myripnois uniflora Maxim. TYPE: China. Kansu, Przewalski 1880 (type, K not seen). 22. Pertya yakushimensis H. Koyama & Nagam., Acta Phytotax. Geobot. 39: 67. 1988. TYPE: Japan. Kyushu: Kagoshima-ken, Yakushima, Tainoko River, 1984, Miura s.n. (type, KYO!).

Literature. Kitamura (1937), Koyama (1973, 1975), Tseng (1985), Hind (2001a).

Plazia Ruiz & Pav. Fl. peruv. prodr.: 104. 1794. TYPE: Plazia conferta Ruiz & Pav. 41 of Figs. 39–42. Aglaodendron J. Rémy, Ann. Sci. Nat., Bot. 12: 175. 1849. TYPE: Aglaodendron cheiranthifolium J. Rémy [= Plazia cheiranthifolia (J. Rémy) Wedd.]. Harthamnus H. Rob., Phytologia 45: 452. 1980. TYPE: Harthamnus boliviensis H. Rob. [= Plazia daphnoides Wedd.]. Etymology. In honor of Juan Plaza (1526–1609), a Spanish botanist and medical doctor. Shrubs. Leaves alternate, clustered at the end of the branches; sessile; blades obovateoblong, one-veined, margin entire, glabrous to densely pubescent on both surfaces. Capitulescences monocephalous, terminal; capitula sessile, surrounded by leaves, homogamous, radiate; receptacle epaleate; involucre three- to four-seriate. Florets

594

L. Katinas, et al.

dimorphic, bisexual; marginal florets with corolla bilabiate, often geniculate, outer lip expanded; central florets with corolla tubular sub-campanulate, deeply five-lobed; anther apical appendages reddish, truncate, tails papillose; style bifid, branches dorsally papillose, rounded to truncate at the apex. Cypselae glabrous; pappus of scabrid bristles, some capillary, the innermost flattened and longer. Pollen Description. Plazia daphnoides (106 of Figs. 101-106) (Tellería & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Parra & Marticorena, 1972). Habitat and Distribution. Genus of three species from the Andes of Argentina, Bolivia, Chile, and Peru. Species list 1. Plazia cheiranthifolia (J. Rémy) Wedd., Chlor. andina 1: 12. 1855. Basionym: Aglaodendron cheiranthifolium J. Rémy. TYPE: Chile. s.d., C. Gay s.n. (holotype, P not seen, P photo F n. 38107 at LP!). 2. Plazia conferta Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 187. 1798. TYPE: Peru. Tarma, s.d., H. Ruiz & J. A. Pavón s.n. (holotype, MA not seen). 3. Plazia daphnoides Wedd., Chlor. andina 1: 13. 1855. TYPE: Peru. Cordillera de Tacora, entre Tacna et La Paz, s.d., H. A. Weddell s.n. (lectotype, selected by Ferreyra, 1995: 83, P not seen). Additional specimens examined: Argentina. Mendoza: Paramillo de Uspallata, A. Ruiz Leal 4839 (LP); Prov. Jujuy: Casabindo, A. L. Cabrera 9315 (LP*). Literature. Cabrera (1951, 1978), Robinson (1980), Ferreyra (1995), Freire (1996).

Salcedoa F. Jiménez Rodr. & Katinas Syst. Bot. 29: 991. 2004. TYPE: Salcedoa mirabaliarum F. Jiménez Rodr. & Katinas. 42 of Figs. 39–42. Etymology. For the Salcedo province, the type locality, in the Dominican Republic. Trees small. Leaves alternate; petiolate; blades oblanceolate to elliptical, pinnately veined, margin entire, glabrous above, tomentose beneath. Capitulescences corymbose; capitula pedunculate, homogamous, bilabiate; receptacle epaleate; involucre multiseriate, outermost phyllaries decurrent onto the stalk. Florets isomorphic, bisexual; corolla bilabiate, deeply incised; anther apical appendages apiculate, tails papillose; style bilobed, branches reddish, dorsally smooth. Cypselae villose; pappus of scabrid bristles, the outermost capillary, the innermost somewhat paleaceous, with tips slightly broadened, and longer. Pollen Description (Jiménez Rodríguez et al., 2004). Pollen prolate, large size, tricolporate, exine Wunderlichia type, echinate.


595

Habitat and Distribution. Monotypic genus endemic to Cordillera Septentrional, northern Dominican Republic. Species list 1. Salcedoa mirabaliarum F. Jiménez Rodr. & Katinas, Syst. Bot. 29: 991. 2004. TYPE: Dominican Republic. Prov. Salcedo: Cordillera Septentrional, el Peñón del Nuevo Mundo, F. Jiménez et al. 3345 (isotype, LP!). Additional specimen examined: Dominican Republic. Prov. Salcedo: Mun. Tenares, Distr. Blanco Arrriba, Veloz et al. 2383 (LP*).

Literature. Jiménez Rodríguez et al. (2004).

Trichocline Cass. Bull. Sci. Soc. Philom. Paris: 13. 1817. TYPE: Doronicum incanum Lam., Encycl. 2: 315. 1786, non L., 1753, nec Hill, 1759, nom. illeg. [= Trichocline incana Cass.]. 43 of Figs. 43–46. Amblysperma Benth., Enum. pl.: 67. 1837. [= Trichocline sect. Amblysperma (Benth.) Zardini, Darwiniana 19: 657. 1975]. TYPE: Amblysperma scapigera Benth. [= Trichocline spathulata (A. Cunn. ex DC.) J. H. Willis]. Bichenia D. Don, Trans. Linn. Soc. London 16: 236. 1830. TYPE: Bichenia aurea D. Don [= Trichocline aurea (D. Don) Reiche]. Etymology. From the Greek thrix, trichos, hair, and kline, bed in reference to the fimbriate receptacle. Herbs perennial, acaulescent, usually scapose. Leaves rosulate; sessile to petiolate; blades linear to ovate, pinnately veined, margin entire to pinnatisect, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle epaleate, smooth to fimbriate; involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth to papillose, filaments smooth to papillose; style bilobed, dorsally papillose to sometimes glabrescent. Cypselae truncate at the apex, shortly pubescent; pappus of scabrid, capillary bristles. Pollen Description. Trichocline reptans (Wedd.) B. L. Rob. (107 of Figs. 107–111) (Tellería & Katinas, 2004). Pollen prolate, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Lulia nervosa. Habitat and Distribution. Genus of 22 species most of them in South America from southern Peru to central Argentina and Chile, and one species in Australia (see Observation).

596

L. Katinas, et al.

Figs. 43–46 43 Trichocline sinuata (D. Don) Cabrera. A Habit. B Marginal floret (without cypsela). C Staminode. D Central floret (without cypsela). E Stamen. F Upper part of the style. G Cypsela with pappus. (A, B, D, F, G redrawn from Zardini, 1975: 672; C, E from Cabrera & Fabris 14847, LP). Scale bars: A=1.5 cm; B, D, G=5 mm; C=1 mm; E=3 mm; F=0.8 mm. 44 Uechtrizia kokanica (Regel & Schmalh.) Pobed. A Habit. B Marginal corolla. C Staminode. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (From Zaprjagaev s.n., WIS). Scale bars: A=5 cm; B=4 mm; C, E= 2 mm; D=5 mm; F=1 mm; G=10 mm. 45 Urmenetea atacamensis Phil. A Habit. B Marginal floret (without cypsela). C Staminode. D Central corolla. E Stamen. F Upper part of the style. G Cypsela with pappus. (A, B, G redrawn from Cabrera, 1978: 596; C–F from Cabrera & Schwabe 80, LP). Scale bars: A=3 cm; B=7 mm; C=1.5 mm; D=6 mm; E=3 mm; F=2 mm; G=4 mm. 46 Acourtia bravohollisiana Rzed. A Habit. B Floret (without cypsela). C Cypsela with pappus. A. huajuapana B. L. Turner var. actinomorpha Rzed. D Habit. E Floret (without cypsela). (Redrawn from Rzedowski, 1983: 99). A. nana (A. Gray) Reveal & R. M. King. F Stamen. G Upper part of the style. (From Supernaugh s.n., LP 303725). Scale bars: A=2 cm; B, C, E, F=2 mm; D=1 cm; G=1 mm

Species list 1. Trichocline aurea (D. Don) Reiche, Anales Univ. Chile 115: 342. 1904. Basionym: Bichenia aurea D. Don. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably GH). Additional specimen examined: Chile. Prov. Concepción: La Florida, E. Barros 244 (LP).


597

2. Trichocline auriculata (Wedd.) Hieron., Bot. Jahrb. Syst. 21: 369. 1896. Basionym: Bichenia auriculata Wedd. TYPE: Bolivia. Environs de Potosí, D’Orbigny 1429 (holotype, P not seen, P photo F n. 38073 at SI not seen). Additional specimen examined: Argentina. Prov. Salta: Dept. Iruya, quebrada de Iruya, Luti & Montaña 5358 (LP). 3. Trichocline beckeri H. Rob., Phytologia 65: 47. 1988. TYPE: Peru. Dept. Cajamarca: surroundings of Cajamarca, B. Becker & F. M. Terrones H. 1305 (holotype, US not seen, US photo at LP!). 4. Trichocline boecheri Cabrera, Bol. Soc. Argent. Bot. 15: 122. 1973. TYPE: Argentina. Prov. Mendoza: ruta 40, km 512, T. W. Böcher et al. 1551 (holotype, LP!). Additional specimen examined: Argentina. Prov. San Juan: Dept. Iglesia, Malimán, A. L. Cabrera et al. 24526 (LP). 5. Trichocline catharinensis Cabrera, Fl. Ilustr. Catarinense 1: 44. 1973. TYPE: Brazil. State Santa Catarina: Caçador, Fazenda Carneiros, L. B. Smith & R. Reitz 9041 (holotype, LP!). Additional specimen examined: Brazil. State Santa Catarina: Mun. Curitibanos, entre Curitibanos e Lages, Pereira 8398 & Pabst 7673 (LP). 6. Trichocline caulescens Phil., Anales Mus. Nac. Santiago de Chile 8: 1. 1891. TYPE: Chile. Médano, pr. Pica, F. Philippi 2069 (holotype, SGO not seen; isotype, B not seen, B photo F n. 16013 at LP!). Additional specimen examined: Chile. Prov. Tarapacá: camino al Portezuelo de Chapiquiña, M. Ricardi et al. 35 (LP). 7. Trichocline cineraria (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 103. 1835. Basionym: Bichenia cineraria D. Don. TYPE: Argentina. Prov. Mendoza: near el Portezuelo de Uspallata, Andes of Mendoza, s.d., J. Gillies s.n. (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, camino a Laguna del Diamante, Barkley & Cuezzo 20Mz398 (LP). 8. Trichocline dealbata (Hook. & Arn.) Benth. & Hook. f. ex Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 215. 1879. Basionym: Chaetanthera dealbata Hook. & Arn. TYPE: Argentina. Prov. Mendoza: ascens to Peuquenes, Andes of Mendoza, s.d., J. Gillies s.n. (lectotype, designated by Zardini 1975: 704, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Tupungato, Portezuelo Morado, Semper 11842 (LP). 9. Trichocline deserticola Zardini, Darwiniana 20: 304. 1976, nom. novum. Basionym: Onoseris spathulata Phil. TYPE: Chile. Prov. Antofagasta: Ascotán, 23 Feb 1885, F. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Antofagasta: camino a Ascotán, NW volcán San Pedro, Martin 449 (LP). 10. Trichocline exscapa Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 197. 1874. TYPE: Argentina. Prov. Tucumán: sierra de Tucumán, P. G. Lorentz 132 (isotype, LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Metán, cima del cerro de Las Mesadas Largas (continuación del Crestón), S. Maldonado 2962 (LP). 11. Trichocline heterophylla (Spreng.) Less., Linnaea 5: 289. 1830. Basionym: Onoseris heterophylla Spreng. TYPE: Brazil. Brasilia meridionalis, Sellow 907 (isotype, LP!). Additional specimen examined: Uruguay. Dept. Florida: Cerro Colorado, estancia San Pedro, Gallinal et al. 2810 (LP).

598

L. Katinas, et al.

12. Trichocline humilis Less., Linnaea 5: 288. 1830. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Entre Ríos: Dept. Federación, Chajarí, A. L. Cabrera 12361 (LP). 13. Trichocline incana Cass., Dict. sci. nat. 55: 216. 1826, nom. novum. Basionym: Doronicum incanum Lam., nom. illeg. TYPE: Uruguay. Montevideo, s.d., D. Commerson s.n. (holotype, P not seen, P photo F n. 22642 at SI not seen). Additional specimen examined: Uruguay. Dept. Maldonado: Punta del Este, Scott de Birabén 24 (LP). 14. Trichocline linearifolia Malme, Kongl. Svenska Vetensk. Acad. Handl. 12: 114. 1933. TYPE: Brazil. State Paraná: Tamanduá, P. Dusén 7714 (isotype, LP!). Additional specimen examined: Brazil. State Paraná: Mun. Curitiba, Río Atuba, Kummrow 198 (LP). 15. Trichocline macrocephala Less., Linnaea 5: 288. 1830. TYPE: Brazil. Brasilia, s. d., Sellow s.n. (neotype, designated by Zardini, 1975: 666, K not seen). Additional specimen examined: Brazil. State Santa Catarina: Mun. Joaçaba, campos of Palmas, 62 km W of Caçador, S. B. Smith & R. M. Klein 11409 (LP). 16. Trichocline macrorhiza Cabrera, Darwiniana 9: 51. 1949. TYPE: Argentina. Prov. Jujuy: Tumbaya, Volcán, Cerro Bola Loma, 21 July 1948, s. leg. (holotype, BAB not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, subida de Purmamarca, Abra de Pives, A. L. Cabrera et al. 26403 (LP). 17. Trichocline maxima Less., Linnaea 5: 290. 1830. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (isotype, K not seen, K photo at LP!). Additional specimen examined: Uruguay. Dept. Rivera: Paso Ataques, J. Chebataroff 9112 (LP). 18. Trichocline plicata Hook. & Arn., Comp. Bot. Mag. 1: 103. 1835. TYPE: Argentina. Prov. San Luis: Portezuela and Cerro de Achiras, s.d., s. leg. (holotype, OXF not seen, OXF photo at LP!). Additional specimen examined: Argentina. Prov. Córdoba: Dept. Pocho, Sierra de Pocho, A. L. Cabrera et al. 29649 (LP). 19. Trichocline reptans (Wedd.) Hieron., Bot. Jahrb. Syst. 21: 369. 1896. Basionym: Bichenia reptans Wedd. TYPE: Bolivia. Environs de Chuquisaca, s.d., s. leg. (type, probably P). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Santa Bárbara, Abra de los Morteros, A. L. Cabrera et al. 24099 (LP*). 20. Trichocline sinuata (D. Don) Cabrera, Man. Fl. Alrededores Buenos Aires: 531. 1953. Basionym: Bichenia sinuata D. Don. TYPE: Argentina. Las Pampas del río Desaguadero, s.d., J. Gillies s.n. (lectotype, designated by Zardini, 1975: 673, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido de Patagones, Igarzabal, A. L. Cabrera & H. A. Fabris 14847 (LP*). 21. Trichocline spathulata (A. Cunn. ex DC.) J. H. Willis, W. Austral. Naturalist 10: 157. 1967. Basionym: Celmisia spathulata A. Cunn. ex DC. TYPE: Australia. SW coast Australia, Oyster Harbour, King George’s Sound, Jan 1818, A. Cunningham s.n. (holotype, G-DC not seen, G-DC photo at LP!). Additional specimen examined: Australia. Murray District, Pritzel 140 (LP). 22. Trichocline speciosa Less., Syn. gen. Compos.: 117. 1832. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (neotype, designated by Zardini, 1975: 696, P not


599

seen). Additional specimen examined: Brazil. State Paraná: Mun. P. Grossa, Parque V. Velha, G. Hatschbach 16048 (LP).

Observation. A new species of Amblysperma, A. minor Keighery, was described for western Australia (Keighery, 2005). The genus Amblysperma was reduced to the synonymy of Trichocline (Katinas, 2004b), and therefore this new species could belong to Trichocline. However, because the protologue of this new species lacks a description and a detailed illustration of corollas and style, we prefer to maintain it as a dubious species until we analyze this taxon in more detail. Literature. Zardini (1975), Robinson (1988), Hind (2001b), Katinas (2004b).

Uechtritzia Freyn Oesterr. Bot. Z. 42: 240. 1892. TYPE: Uechtritzia armena Freyn & Sint. 44 of Figs. 43–46. Etymology. In honor of the German (Silesian) botanist Rudolf Karl von Uechtritz (1838–1886). Herbs perennial, acaulescent, scapose. Leaves rosulate; pseudopetiolate; blades elliptic to ovate, pinnately veined, margin entire to lyrate or pinnatifid, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle fimbriate-laciniate; involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae truncate at the apex, villose with shaggy hairs; pappus of scabrid, capillary bristles. Pollen Description. Uechtritzia kokanica (Regel & Schmalh.) Pobed. (Tellería & Katinas, 2004). Pollen prolate, large size, tricolporate, exine Mutisia type, scabrate. Habitat and Distribution. Genus of three species distributed in southern Russia, north to Afghanistan, Kashmir, northwestern China, Armenia, and northwestern India. Species list 1. Uechtritzia armena Freyn & Sint., Oesterr. Bot. Z. 42: 241. 1892. TYPE: Armenia. Sipikordagh, Sinentis 3185 (holotype, BRNU not seen). 2. Uechtritzia kokanica (Regel & Schmalh.) Pobed., Fl. U.S.S.R. 28: 597. 1963. Basionym: Gerbera kokanica Regel & Schmalh. TYPE: Bangladesh. In Kokania, prope Schahimardan, Karassu, s.d., O. Fedtschenko s.n. (holotype?, LE not seen). Additional specimen examined: Russia. Tadzhikistania: Hissaricum, Unou, 13 Aug 1934, F. Zaprjagaev s.n. (WIS).

600

L. Katinas, et al.

3. Uechtritzia lacei (Watt) C. Jeffrey, Kew Bull. 21: 213. 1967. Basionym: Gerbera lacei Watt. TYPE: India. Nachar forest, Lace 935 (lectotype, designated by Hansen, 1988: 74, K not seen).

Literature. Hansen (1988).

Urmenetea Phil. Fl. atacam.: 26. 1860. TYPE: Urmenetea atacamensis Phil. 45 of Figs. 43–46. Etymology. In honor of Hieronymus Urmeneta (1816–1881) who promoted agricultural science in Chile. Herbs perennial, subscapose. Leaves nearly rosulate; petiolate; blades spathulate to orbicular, palmately veined, margin serrate, rugose above, woolly beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with reduced, filiform staminodes, corollas bilabiate, outer lip expanded; central florets with corollas tubular-bilabiate, shallowly five-lobed, lobes straight, one incision deeper than the others; anther apical appendages acute to apiculate, tails papillose, filaments papillose at the base; style bifid, branches dorsally papillose. Cypselae glabrous; pappus of scabrid, capillary bristles, a few inner ones flattened and wider. Pollen Description (Tellería & Katinas, 2004) (108 of Figs. 107–111). Pollen spheroidal, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Pachylaena atriplicifolia (see also Parra & Marticorena, 1972). Habitat and Distribution. Monotypic genus from the Andes of Chile and northwestern Argentina. Species list 1. Urmenetea atacamensis Phil., Fl. atacam.: 26. 1860. TYPE: Chile. Prov. Atacama: cerca de la población de San Pedro de Atacama, s.d., R. Philippi s.n. (type, probably SGO). Additional specimen examined: Argentina. Prov. Salta: Los Andes, Abra de Navarro, A. L. Cabrera & H. Schwabe 80 (LP*). Literature. Cabrera (1978), Freire (1996), Sancho (2004).

Tribe Nassauvieae Cass. H. Cassini, J. Phys. Chim. Hist. Nat. Arts. 88: 198. 1819. Nassauvioideae (Cass.) Lindl. in Loudon, Encycl. pl.: 1074. 1829. Nassauviinae Dumort., Anal. fam. pl.: 31.


601

1829, as “Nassauvieae”; Less., Linnaea 5: 2. 1830. Nassauviaceae Burmeist., Handb. Naturgesch. 1: 290. 1837. TYPE: Nassauvia Comm. ex Juss. Trixideae (Cass.) Lindl. in Loudon, Encycl. pl.: 1074. 1829. Trixidinae Less., Linnaea 5: 6. 1830, as “Trixidea.” TYPE: Trixis P. Browne. Jungeae D. Don, Trans. Linn. Soc. London 16: 224. 1830, as “Jungieae.” TYPE: Jungia L. f. Polyachyreae D. Don, Trans. Linn. Soc. London 16: 229. 1830. Polyachyrinae Endl., Gen. pl. 3: 489. 1838. TYPE: Polyachyrus Lag. Herbs, subshrubs, shrubs or vines. Leaves alternate to rosulate, entire to pinnatisect, rarely spiny, glabrous to pubescent. Capitulescences monocephalous, laxly to densely corymbose, racemose, paniculate to glomerulose or pseudocephalia. Capitula homogamous, discoid or radiate by extension of the outer lip in marginal corollas, rarely disciform; involucre uniseriate to multiseriate; receptacle epaleate to paleate, glabrous to pubescent. Florets isomorphic, rarely dimorphic, bisexual, corolla bilabiate with the outer lip three-toothed and the inner bifid, rarely tubular; anthers with apical appendage lanceolate, acute, long, smooth to papillose tails; style cleft into two truncate branches, each branch with an apical crown of collector or sweeping hairs, rarely branches dorsally papillose and papillae not in a crown. Cypselae truncate to rostrate at the apex, glabrous to pubescent; pappus of capillary bristles to paleaceous, scabrose to plumose, equal in length, rarely pappus absent. Pollen microechinate, exine distinctly bilayered, with the ectosexine and the endosexine clearly columellate. n=4, 11, 12, 20, 22, 27, 28; 2n=16, 24, 36, 47±3, 54. Characters that define the tribe are in italics. Tribe with 25 genera and ca. 320 species of the Neotropical biogeographic province (Cabrera & Willink, 1973). Plants belonging to this tribe are an important component of the Andean and Patagonian flora. Genera: Acourtia D. Don, Ameghinoa Speg., Berylsimpsonia B. L. Turner, Burkartia Crisci, Calopappus Meyen, Cephalopappus Nees & Mart., Criscia Katinas, Dolichlasium Lag., Holocheilus Cass., Jungia L. f., Leucheria Lag., Leunisia Phil., Lophopappus Rusby, Macrachaenium Hook. f., Marticorenia Crisci, Moscharia Ruiz & Pav., Nassauvia Comm. ex Juss., Oxyphyllum Phil., Panphalea Lag., Perezia Lag., Pleocarphus D. Don, Polyachyrus Lag., Proustia Lag., Triptilion Ruiz & Pav., Trixis P. Browne. Literature. Crisci (1974a, 1980), Cabrera (1977), Katinas (1995).

Key to Genera 1a. Paleae embracing all or only the central florets of the capitula. 2a. Leaves linear to linear-lanceolate …………………………….. Pleocarphus 2b. Leaves ovate-elliptic, suborbicular to triangular-ovate. 3a. Leaves ovate-elliptic, pinnately veined, sessile, decurrent to clasping ……………..……………………………...……………………...... Marticorenia 3b. Leaves elliptic to orbicular-cordate, palmately veined, petiolate (except Jungia stuebelii with leaves ovate, pinnately veined, sessile) …. Jungia 1b. Paleae only in the marginal florets of the capitula, or completely absent.

602

L. Katinas, et al.

4a. Pappus absent. 5a. Leaves spathulate, entire; cypselae constricted to rostrate at the apex ..……………………………………………………….... Cephalopappus 5b. Leaves linear-lanceolate, orbicular, entire to lyrate to pinnatisect; cypselae truncate at the apex ……………………………… Panphalea 4b. Pappus present. 6a. Pappus bristles narrowly paleaceous, as long as the florets or longer (except in Nassauvia coronipappa with crown-like pappus). 7a. Pappus elements longitudinally plicate, apically plumose .... Triptilion 7b. Pappus elements flat, ciliate to laciniate throughout their length. 8a. Involucre multiseriate; capitulescence monocephalous...Calopappus 8b. Involucre two-seriate; capitulescences monocephalous, racemose, spicate to pseudocephalium …………….……………. Nassauvia 6b. Pappus capillary, if paleaceous very short. 9a. Capitula arranged in pseudocephalia. 10a. Pseudocephalia spiciform …………..……………. Polyachyrus 10b. Pseudocephalia capituliform ..……..………………. Moscharia 9b. Capitulescences monocephalous or cymose. 11a. Most leaves reduced to three-parted spines; paleae subtending all the marginal florets ...……..…………………….. Oxyphyllum 11b. Leaves not reduced to three-parted spines; capitula with or without paleae. 12a. Style branches dorsally papillose. 13a. Shrubs or little trees. 14a. Plants with thorns to unarmed; capitula in racemes to panicles ....……………….. Proustia 14b. Plants unarmed; capitula solitary or few at the apex of the branches ..………..… Lophopappus 13b. Perennial, acaulescent herbs ...…….. Macrachaenium 12b. Style branches smooth, crowned by a tuft of collector hairs. 15a. Shrubs or subshrubs. 16a. Leaves pinnatisect ..……………... Dolichlasium 16b. Leaves entire to lobulate. 17a. Clambering plants, with recurved, bifurcate, pseudostipulate spines ...… Berylsimpsonia 17b. Erect plants, without spines or with thorns formed by foreshortened stems. 18a. Pappus paleaceous–setaceous, unequally fused at the base ………… Leunisia 18b. Pappus bristles scabrid to plumose 19a. Dwarf shrubs with subulate leaves; capitulescences monocephalous or two-headed .…………. Burkartia 19b. Erect plants with broad leaves; capitulescences monocephalous to cymose with numerous capitula.


603

20a. Stems with brachyblasts and macroblasts; cypselae usually truncate at the apex ..…………..…. Ameghinoa 20b. Stems without brachyblasts; cypselae usually rostrate .... …………...……..…. Trixis 15b. Perennial leafy caulescent or scapose herbs. 21a. Plants covered with a woolly to silky white pubescence ..……………………….. Leucheria 21b. Plants glabrous, subglabrous or with a reddish pubescence. 22a. Corollas and pappus orange-red. 23a. Scapose herbs; capitulescences monocephalous ..………..…………………… Criscia 23b. Caulescent herbs; capitulescences corymbose, paniculate to spicate …. Trixis 22b. Corollas pink, purple, white, rarely yellow; pappus white, brown, reddish, bronzepurplish. 24a. Involucre one- to two-seriate ……..... …………………………. Holocheilus 24b. Involucre with three to more series of phyllaries. 25a. Stem base with a tuft of rustcolored woolly hairs; capitula discoid ..…………….. Acourtia 25b. Stem base glabrous or at least not densely rust-colored woolly hairs; capitula radiate, occasionally capitula discoid ….. Perezia Acourtia D. Don Trans. Linn. Soc. London 16: 203. 1830. TYPE: Acourtia formosa D. Don [= Acourtia fruticosa (La Llave & Lex.) B. L. Turner]. 46 of Figs. 43–46. Neoshinneria B. L. Turner, Phytologia 38: 456. 1978, nom. nudum. Etymology. In honor of Mrs. Mary Elizabeth Catherine Gibbes A’Court (1792– 1878), a British amateur botanist. Herbs perennial or subshrubs, caulescent or acaulescent, base of the stem with a tuft of rust-colored woolly hairs. Leaves rosulate to spreading; sessile, petiolate, pseudopetiolate to clasping; blades lanceolate, broadly orbicular to cordate, pinnately veined, margin usually spinulose, entire to lyrate, glabrous to pubescent. Capitulescences monocephalous, or corymbose to paniculate; capitula pedunculate to subsessile,

604

L. Katinas, et al.

homogamous, discoid; receptacle epaleate, occasionally fimbrillate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate, occasionally tubular-funnelform five-lobed; anther apical appendages acute to slightly apiculate, tails smooth; style bifid, branches linear, apically rounded, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus of simple to scabrid bristles, soft to rigid. Pollen Description. Acourtia (Crisci, 1974a; Cabrera-Rodríguez, 1992 sub Rzedowskiela; Cabrera-Rodríguez & Dieringer, 2003). Pollen subprolate, oblate-spheroidal, prolate-spheroidal or spheroidal, medium size, tricolporate, exine Trixis and Proustia type, slightly microechinate. Habitat and Distribution. Genus of ca. 85 species distributed from southern United States to Central America reaching El Salvador, with a high concentration in the forested middle and higher slopes of the Sierra Madre in Mexico and the high plateaus east of the mountains in central and northern Mexico. Species list 1. Acourtia acevedoi M. González, Phytologia 61: 117. 1986. TYPE: Mexico. Durango: Municipio Vicente Guerrero, Rincón de las Mulas, 3 km al SW de San Isidro, S. Acevedo 163 (isotype, NY, NY photo at LP!). 2. Acourtia aspera (Bacig.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia aspera Bacig. TYPE: Mexico. Coahuila: Saltillo, Bro. Arsène 3395 (isotype, MO, MO photo at LP!). 3. Acourtia belizeana B. L. Turner, Phytologia 74: 138. 1993. TYPE: Belize. Toledo District: lower part of Richardson Creek, affluent of Bladen Branch, lower part of Maya Mtns., G. Davidse & A. E. Brant 31904 (isotype, US not seen). 4. Acourtia bravohollisiana Rzed., Bol. Soc. Bot. Mexico 45: 98. 1983. TYPE: Mexico. Guerrero: camino a la torre de microondas Tuxpan, +- 10 km al NE de Iguala, J. Rzedowski 37162 (isotype, NY, NY photo at LP!). 5. Acourtia butandae R. L. Cabrera, Brittonia 53: 423. 2001. TYPE: Mexico. Chihuahua: 18 km of Villa Matamoros, hwy 45, L. Cabrera & G. Dieringer 650 (isotype, US not seen, US photo at LP!). 6. Acourtia caltepecana B. L. Turner, Phytologia 74: 386. 1993. TYPE: Mexico. Puebla: Mpio. Caltepec, Cerro El Tambor, al NE de Caltepec, P. Tenorio 7634 (holotype, TEX not seen). 7. Acourtia carpholepis (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia carpholepis A. Gray. TYPE: Mexico. Puebla,: Chapulco, Liebmann 351 (lectotype, designated by Turner, 1993a: 387, GH, not seen). 8. Acourtia carranzae R. L. Cabrera, Brittonia 53: 416. 2001. TYPE: Mexico. Querétaro: Mun. Arroyo Seco, 1 km al S de Arroyo Seco, E. Carranza 1488 (holotype, IEB not seen). 9. Acourtia ciprianoi Panero & Villaseñor, Brittonia 51: 87. 1999. TYPE: Mexico. Oaxaca: Dto. Tehuantepec, Mun. Santiago Astata, carretera Pochutla-Salina Cruz, C. Martínez 1978 (isotype, TEX not seen).


605

10. Acourtia collina (S. Watson) Crisci, J. Arnold Arbor. 55: 605. 1974. Basionym: Perezia collina S. Watson. TYPE: Mexico. Jalisco: near Guadalajara, C. G. Pringle 2123 (holotype, GH not seen). 11. Acourtia cordata (Cerv.) B. L. Turner, Phytologia 74: 388. 1993. Basionym: Perdicium cordatum Cerv. TYPE: Mexico. DF?, montibus del desierto Mexico vicinis, M. Sessé & J. M. Moçiño 3736 (lectotype, designated by Turner, 1993a: 388, M, not seen). 12. Acourtia coulteri (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia coulteri A. Gray. TYPE: Mexico. State Hidalgo: Zimapán, Coulter 234 (holotype, GH not seen). 13. Acourtia cuernavacana (B. L. Rob. & Greenm.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia cuernavacana B. L. Rob. & Greenm. TYPE: Mexico. Morelos: above Cuernavaca, C. G. Pringle 6196 (isotype, MO, MO photo at LP!). 14. Acourtia dieringeri R. L. Cabrera, Sida 13: 419. 1989. TYPE: Mexico. Chihuahua: 20 km N of Basaseachic, along the Basaseachic-Yepachic dirt road, L. Cabrera & G. Dieringer 628 (isotype, GH not seen). 15. Acourtia discolor Rzed., Bol. Soc. Bot. Mexico 45: 100. 1983. TYPE: Mexico. Oaxaca: 30 km al NNW de Telixtlahauaca, sobre la carretera de Cuicatlán, J. Rzedowski 37181 (holotype, ENCB not seen). 16. Acourtia dissiticeps (Bacig.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia dissiticeps Bacig. TYPE: Mexico. San Luis Potosí: Sierra Tablón, Minas de San Rafael, C. A. Purpus 5018 (isotype, MO, MO photo at LP!). 17. Acourtia dugesii (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia dugesii A. Gray. TYPE: Mexico. State Guanajuato: Guanajuato, s.d., A. Dugès s.n. (holotype, GH not seen). 18. Acourtia durangensis B. L. Turner, Phytologia 74: 389. 1993. TYPE: Mexico. State Durango: Cienega bottomland near small lake, 40 mi N of Cd. Durango, H. Gentry 8594 (holotype, MICH not seen). 19. Acourtia elizabethiae Rzed. & Calderon, Acta Bot. Mex. 12: 16. 1990. TYPE: Mexico. Querétaro: alrededores de Maconí, municipio de Cadereyta, J. Rzedowski 47625 (holotype, ENCB not seen). 20. Acourtia erioloma (S. F. Blake) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia erioloma S. F. Blake. TYPE: Mexico. Oaxaca: near Tlaxiaco, W. H. Camp 2225 (holotype, NY not seen, NY photo at LP!). 21. Acourtia fragrans Rzed., Bol. Soc. Bot. Mexico 45: 102. 1983. TYPE: Mexico. Puebla: Salinas Grandes, cerca de Zapotitlán de las Salinas, J. Rzedowski 37174 (isotype, NY not seen, NY photo at LP!). 22. Acourtia fruticosa (La Llave & Lex.) B. L. Turner, Phytologia 74: 390. 1993. Basionym: Perezia fruticosa La Llave & Lex. TYPE: Mexico. Michoacán: Jesus del Monte near Morelia, C. G. Pringle 10410 (neotype, designated by Turner, 1993a: 390, LL not seen; isoneotype, US not seen, US photo at LP!). 23. Acourtia gentryi R. L. Cabrera, Acta Bot. Mex. 20: 35. 1992. TYPE: Mexico. Sinaloa: above la Jolla, Sierra Surotato, H. S. Gentry 7258 (isotype, NY not seen, NY photo at LP!). 24. Acourtia glandulifera (D. L. Nash) B. L. Turner, Phytologia 38: 461. 1978. Basionym: Perezia glandulifera D. L. Nash. TYPE: Guatemala. Dept.

606

25.

26.

27.

28.

29.

30.

31.

32.

33.

34.

35.

36.

37.

38.

L. Katinas, et al.

Huehuetenango: canyon of Río Selequa, 40 km NW of Huehuetenango, L. O. Williams et al. 41167 (isotype, NY not seen, NY photo at LP!). Acourtia glomeriflora (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Gochnatia glomeriflora A. Gray. TYPE: Mexico. Cuernavaca, D. Bilimek 512 (holotype, GH not seen). Acourtia gracilis R. L. Cabrera, Acta Bot. Mex. 20: 36. 1992. TYPE: Mexico. Puebla: Barranca de Patla, Villa Juárez, E. Matuda 38750 (holotype, MICH not seen). Acourtia grandifolia (S. Watson) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia grandifolia S. Watson. TYPE: Mexico. Jalisco: hillsides near Guadalajara, C. G. Pringle 1858 (isotype, MO, MO photo at LP!). Acourtia guatemalensis B. L. Turner, Phytologia 74: 393. 1993. TYPE: Guatemala. Dept. Solola: above Lake Atitlán, ca. 3–5 km W of Panojachel, L. O. Williams et al. 25318 (isotype, US not seen, US photo at LP!). Acourtia hidalgoana B. L. Turner, Phytologia 74: 394. 1993. TYPE: Mexico. State Hidalgo: 7 km al NE de Mesquititlán, sobre la carretera a Zacualtipan, J. Rzedowski 32510 (holotype, MEXU not seen). Acourtia hondurana B. L. Turner, Phytologia 38: 466. 1978. TYPE: Honduras. El Paraíso: cerros El Zapotillo, road from Zamorano to Guinope, G. Webster et al. 11982 (holotype, MICH not seen). Acourtia hooveri (McVaugh) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia hooveri McVaugh. TYPE: Mexico. Jalisco: Nevada de Colima, 11 mi. from Atenquique on the Tonila, R. McVaugh 11750 (isotype, NY not seen, NY photo at LP!). Acourtia huajuapana B. L. Turner, Phytologia 38: 455. 1978. TYPE: Mexico. State Oaxaca: 37 mi NW of Huajuapan de León, along highway 125, B. L. Turner P-59 (holotype, LL not seen). Acourtia humboldtii (Less.) B. L. Turner, Phytologia 74: 395. 1993. Basionym: Dumerilia humboldtii Less. TYPE: Mexico. sine loc., s.d., A. Humboldt s.n. (holotype B, microfiche Herb. Willdenow 16095 not seen). Acourtia intermedia R. L. Cabrera, Brittonia 53: 418. 2001. TYPE: Mexico. Durango: 166 mi NW of Durango, along hwy 145, L. Cabrera & G. Dieringer 653 (isotype, US not seen, US photo at LP!). Acourtia joaquinensis R. L. Cabrera, Brittonia 53: 423. 2001. TYPE: Mexico. Querétaro: Mun. San Joaquín, ca. 1 km al S de Santo Tomás, Huerta 1324 (holotype, IEB not seen). Acourtia lepidopoda (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lepidopoda B. L. Rob. TYPE: Mexico. Morelos: near Cuernavaca, C. G. Pringle 9253 (holotype, GH not seen). Acourtia lobulata (Bacig.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lobulata Bacig. TYPE: Mexico. Oaxaca: San Juan del Estado, L. C. Smith 888 (holotype, GH not seen, photo in Bacigalupi, 1931: plate 2). Acourtia longifolia (S. F. Blake) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia longifolia S. F. Blake. TYPE: Mexico. Jalisco: Calabaza, B. P. Reko 4872 (holotype, US not seen).


607

39. Acourtia lozanii (Greenm.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lozanii Greenm. TYPE: Mexico. Hidalgo: between Metepec and Zontecomate stations, C. G. Pringle 8871 (isotype, MO, MO photo at LP!). 40. Acourtia macvaughii B. L. Turner, Phytologia 74: 398. 1993. TYPE: Mexico. Michoacán: 8–12 km SE of Aserradero Dos Aguas and nearly W of Aguililla, R. McVaugh 22789 (isotype, NY not seen, NY photo at LP!). 41. Acourtia macrocephala Sch. Bip. ex Seeman, Bot. Voy. Herald. 55: 315. 1856. TYPE: Mexico. Durango?: Sierra Madre, Dec–Feb 1838–1839, Seeman s.n. (holotype, K, not seen). 42. Acourtia mexiae R. L. Cabrera, Acta Bot. Mex. 20: 32. 1992. TYPE: Mexico. Jalisco: Real Alto, trail to El Tajo de Santiago, Y. Mexia 1749 (isotype, NY not seen, NY photo at LP!). 43. Acourtia mexicana (Lag. ex D. Don) H. Rob., Phytologia 69: 106. 1990. TYPE: Mexico. s.d., M. Sessé & J. M. Moçiño s.n. (holotype, G not seen). 44. Acourtia michoacana (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia michoacana B. L. Rob. TYPE: Mexico. Michoacán: near Patzcuaro, C. G. Pringle 3988 (isotype, MO, MO photo at LP!). 45. Acourtia microcephala DC., Prodr. 7: 66. 1838. TYPE: United States. California: Douglas 62 (holotype, GH not seen). 46. Acourtia moctezumae Rzed. & Calderon, Acta Bot. Mex. 12: 14. 1990. TYPE: Mexico. Querétaro: 4 km al SE de Matzacintla, municipio de Landa, L. M. Chávez 217 (holotype, ENCB not seen). 47. Acourtia molinana B. L. Turner, Phytologia 38: 461. 1978. TYPE: Honduras. Dept. Morazán, Río Guarabuquí, A. Molina R. 3048 (holotype, US not seen). 48. Acourtia moschata (La Llave & Lex.) DC., Prodr. 7: 66. 1838. Basionym: Perezia moschata La Llave & Lex. TYPE: Mexico. Michoacán de Ocampo: 24 km W of Morelia, H. D. D. Ripley & R. C. Barneby 14849 (neotype, designated by Turner, 1993a: 398, NY not seen, NY photo at LP!). 49. Acourtia nana (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia nana A. Gray. TYPE: Mexico. Chihuahua: high, dry valley near Chihuahua, J. Gregg 45 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Sine loc., s.d., Supernaugh s.n. (LP 303725). 50. Acourtia nelsonii (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia nelsonii B. L. Rob. TYPE: Mexico. State Jalisco: mountains near Talpa, E. Nelson 4037 (holotype, GH not seen). 51. Acourtia nudicaulis (A. Gray) B. L. Turner, Phytologia 38: 467. 1978. Basionym: Perezia nudicaulis A. Gray. TYPE: Guatemala. Sine loc., s.d., Skinner s.n. (holotype, GH not seen). 52. Acourtia nudiuscula (B. L. Rob.) B. L. Turner, Phytologia 38: 467. 1978. Basionym: Perezia nudiuscula B. L. Rob. TYPE: Mexico. Nayarit: Tepic, E. Palmer 2018 (isotype, NY not seen, NY photo at LP!). 53. Acourtia oaxacana R. L. Cabrera, Acta Bot. Mex. 20: 31. 1992. TYPE: Mexico. Oaxaca: 62 km al SE de Oaxaca, sobre la carretera a Tehuantepec, J. Panero & A. Salinas 2495 (isotype, US not seen). 54. Acourtia ovatifolia R. L. Cabrera, Sida 14: 141. 1990. TYPE: Mexico. Guerrero: 9 mi by road N of Iguala, W. R. Anderson & C. Anderson 5656 (holotype, MICH not seen).

608

L. Katinas, et al.

55. Acourtia oxylepis (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia oxylepis A. Gray. TYPE: Mexico. San Luis Potosí, C. C. Parry & E. Palmer 547 (isotype, MO, MO photo at LP!). 56. Acourtia palmeri (S. Watson) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia palmeri S. Watson. TYPE: Mexico. Baja California: near Los Angeles Bay, Gulf of California, E. Palmer 527 (isotype, NY not seen, NY photo at LP!). 57. Acourtia parryi (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia parryi A. Gray. TYPE: Mexico. San Luis Potosí, C. C. Parry & E. Palmer 545 (isotype, MO, MO photo at LP!). 58. Acourtia patens (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia patens A. Gray. TYPE: Mexico. Sine loc., 1839–40, Seeman s.n. (lectotype, designated by Turner 1993a: 401, GH not seen). 59. Acourtia pilulosa (Bacig.) B. L. Turner, Phytologia 74: 402. 1993. Basionym: Perezia dugesii A. Gray var. pilulosa Bacig. TYPE: Mexico. Oaxaca: below Jayacatlin, L. Smith 373 (holotype, GH not seen). 60. Acourtia pinetorum (Brandegee) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia pinetorum Brandegee. TYPE: Mexico. Baja California: Sierra de la Laguna, 22 Jan 1899, T. S. Brandegee s.n. (isotype, GH not seen). 61. Acourtia platyphylla (A. Gray) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia platyphylla A. Gray. TYPE: Mexico. Chihuahua, A. Wislizenus 165 (isotype, MO, MO photo at LP!). 62. Acourtia platyptera (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia platyptera B. L. Rob. TYPE: Mexico. Michoacán or Guerrero: Sierra Madre, E. Langlassé 773 (isotype, US not seen). 63. Acourtia potosina R. L. Cabrera, Brittonia 53: 420. 2001. TYPE: Mexico. San Luis Potosí: ca. 32.2 km E of San Luis Potosí, 1.1 km S of jct. with Rt. 70 along road to microwave station, A. Prather & J. Soule 914 (isotype, TEX not seen). 64. Acourtia pringlei (B. L. Rob. & Greenm.) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia pringlei B. L. Rob. & Greenm. TYPE: Mexico. Michoacán: near Morelia, C. G. Pringle 5464 (holotype, GH not seen). 65. Acourtia pulchella R. L. Cabrera, Acta Bot. Mex. 20: 33. 1992. TYPE: Mexico. Aguascalientes: 3 km al E de La Congoja, sobre el camino a San José de Gracia, J. Rzedowski & R. McVaugh 897 (holotype, MICH not seen). 66. Acourtia purpusii (Brandegee) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia purpusii Brandegee. TYPE: Mexico. San Luis Potosí: Minas de San Rafael, C. A. Purpus 4787 (isotype, MO, MO photo at LP!). 67. Acourtia queretarana B. L. Turner, Phytologia 74: 402. 1993. TYPE: Mexico. Querétaro: Mpio. Pinal de Amoles, 13 km al NE de Pinal de Amoles, sobre la carretera a Jalpan, J. Rzedowski 48108 (holotype, TEX not seen). 68. Acourtia reticulata (Lag. ex D. Don) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Proustia reticulata Lag. ex. D. Don. TYPE: Mexico. Sine loc., 1787–1804, M. Sessé & M. Moçiño 3082 (lectotype, designated by Turner 1993a: 403, M not seen). 69. Acourtia runcinata (Lag. ex D. Don) B. L. Turner, Phytologia 38: 460. 1978. Basionym: Clarionia runcinata D. Don. TYPE: Mexico. Sine loc., s.d., M. Sessé & M. Moçiño s.n. (holotype, BM not seen). Additional specimen


70.

71.

72.

73.

74.

75.

76.

77. 78.

79.

80.

81.

82. 83.

84.

609

examined: United States. Texas: Neueces county, Corpus Christy bay, A. Heller 1537 (LP). Acourtia rzedowskii B. L. Turner, Phytologia 74: 405. 1993. TYPE: Mexico. Puebla: 8 km al NNE de Azumbilla, sobre la carretera a Esperanza, J. Rzedowski 37173 (isotype, US not seen). Acourtia scapiformis (Bacig.) B. L. Turner, Phytologia 38: 463. 1978. Basionym: Perezia scapiformis Bacig. TYPE: Mexico. Oaxaca: Las Sedas, C. G. Pringle 6015 (isotype, MO, MO photo at LP!). Acourtia scaposa (S. F. Blake) B. L. Turner, Phytologia 38: 463. 1978. Basionym: Perezia scaposa S. F. Blake. TYPE: Mexico. Michoacán: Distrito Coalcoman, Aquila, G. B. Hinton et al. 15838 (holotype, US not seen). Acourtia simulata (S. F. Blake) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia simulata S. F. Blake. TYPE: Mexico. Michoacán: Distrito [District] Coalcoman, G. B. Hinton 13654 (isotype, NY not seen, NY photo at LP!). Acourtia sinaloana B. L. Turner, Phytologia 74: 406. 1993. TYPE: Mexico. Sinaloa: summit of Sierra Tacuichamona, H. S. Gentry 5679 (isotype, NY not seen, NY photo at LP!). Acourtia souleana B. L. Turner, Phytologia 74: 406. 1993. TYPE: Mexico. Oaxaca: 7.5 mi NW of Huahuapán de León, 4.5 mi SE of Puebla, J. A. Soule & L. A. Prather 3187 (holotype, TEX not seen). Acourtia tenoriensis B. L. Turner, Phytologia 58: 225. 1985. TYPE: Mexico. Pueblo: Mpio. Tamazulapan, Cerro Pericon, al NW de San Pedro Nopala, Tenorio & Romero de T. 7871 (holotype, TEX not seen). Acourtia thurberi (A. Gray) Reveal & R. M. King, Phytologia 27: 231. 1973. TYPE: Mexico. State Sonora: Santa Cruz, Thurber 939 (holotype, GH not seen). Acourtia tomentosa (Brandegee) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia tomentosa Brandegee. TYPE: Mexico. Puebla: Esperanza, C. A. Purpus 2632 (isotype, MO, MO photo at LP!). Acourtia turbinata (La Llave & Lex.) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia turbinata La Llave & Lex. TYPE: Mexico. State Michoacán: Municipio Senguio, 5 km al S de Chincua, J. Rzedowski 48261 (neotype, designated by Turner 1993a: 407, TEX not seen). Acourtia umbratalis (B. L. Rob. & Greenm.) B. L. Turner, Phytologia 38: 465. 1978. Basionym: Perezia umbratalis B. L. Rob. & Greenm. TYPE: Mexico. Oaxaca: Tomellin Cañon, C. G. Pringle 5966 (holotype, GH not seen). Acourtia venturae R. L. Cabrera, Brittonia 53: 420. 2001. TYPE: Mexico. Guanajuato: Mun. Victoria, Puerto del Aire, por la Sierra de Victoria, E. Ventura & E. López 9797 (holotype, IEB not seen). Acourtia veracruzana B. L. Turner, Phytologia 74: 408. 1993. TYPE: Mexico. State Veracruz: Maltrata, E. Matuda 1236 (holotype, MEXU not seen). Acourtia wislizenii (A. Gray) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia wislizenii A. Gray. TYPE: Mexico. Chihuahua: base of Sierra Madre, Wislizenus 198 (isotype, MO, MO photo at LP!). Acourtia wrightii (A. Gray) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia wrightii A. Gray. TYPE: United States. Texas: on the Río Seco and westward, also on the Río Grande, C. Wright 411 (holotype, GH not seen).

610

L. Katinas, et al.

85. Acourtia zacatecana B. L. Turner, Phytologia 74: 411. 1993. TYPE: Mexico. Zacatecas: 6 km antes de Monte Escobedo, a 36 km de Huejucar, J. D. García P. & S. Delgado 856 (isotype, NY not seen, NY photo at LP!). Literature. Bacigalupi (1931), Reveal and King (1973), Cabrera-Rodríguez (1992, 2001), Cabrera-Rodríguez and Dieringer (2003).

Ameghinoa Speg. Revista Fac. Agron. Univ. Nac. La Plata 3: 539. 1897. TYPE: Ameghinoa patagonica Speg. 47 of Figs. 47–50. Etymology. In honor of the Argentine geologist Carlos Ameghino (1865–1936) who collected plants in Patagonia. Shrubs stout, with brachyblasts and macroblasts. Leaves alternate; short-petiolate; blades obovate, coriaceous, palmately veined, margin dentate to lobate, glabrescent. Capitulescences corymbose; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre one- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description (Tellería & Forcone, 2002) (109 of Figs. 107–111). Pollen prolate to subprolate, medium size, tricolporate, exine Trixis type, slightly microechinate, with polar caps (see also Crisci, 1974a). Habitat and Distribution. Monotypic genus endemic to the Patagonian steppe in Argentina. Species list 1. Ameghinoa patagonica Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 539. 1897. TYPE: Argentina. Prov. Santa Cruz: Dept. Deseado, Puerto Deseado, 1894, C. Ameghino s.n. (holotype, LP!). Additional specimens examined: Argentina. Prov. Neuquén: Cerro Negro, R. Maldonado 681 (LP); Prov. Chubut: Dept. Río Senguerr, Río Senguerr, A. Soriano 3208 (LP*). Literature. Cabrera (1971b), Katinas (1995).

Berylsimpsonia B. L. Turner Phytologia 74: 351. 1993. TYPE: Proustia vanillosma C. Wright [= Berylsimpsonia vanillosma (C. Wright) B. L. Turner]. 48 of Figs. 47–50.


611

Figs. 47–50 47 Ameghinoa patagonica Speg. A Habit (branch). B Floret (without cypsela). C Stamen. D Upper part of the style. E Cypsela with pappus. (A–C redrawn from Cabrera, 1971b: 376; D, E, from Maldonado 681, LP). Scale bars: A=2 cm; B, E=4 mm; C=2.5 mm; D=1.2 mm. 48 Berylsimpsonia vanillosma (C. Wright) B. L. Turner. A Habit (branch). B Floret (without cypsela). C Cypsela with pappus. B. crassinervis (Urb.) B. L. Turner. D Stamen. E Upper part of the style. (A–C redrawn from Fabris, 1968: 32; D, E from Leonard 7754, US). Scale bars: A=3 cm; B, C=3 mm; D=1.5 mm; E=1 mm. 49 Burkartia lanigera (Hook. & Arn.) Crisci. A Habit. B Floret (without cypsela). C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Crisci, 1976a: 244). Scale bars: A=2 cm; B, E=2.5 mm; C, D= 1 mm. 50 Calopappus acerosus Meyen. A Habit (branch). B Leaves. C Floret (without cypsela). D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Crisci & Freire, 1986: 60). Scale bars: A=1 cm; B, C, F=5 mm; D=2.5 mm; E=1 mm

Etymology. In honor of the botanist Beryl Simpson (b. 1942), from United States, who made important contributions to the study of Asteraceae. Shrubs clambering, with bifurcate, short, pseudostipulate, recurved spines. Leaves alternate; short-petiolate; blades oblong, cuneate to suborbicular, pinnately veined, margin serrulate to spinulose, pubescent. Capitulescences densely

612

L. Katinas, et al.

racemose to paniculate; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate to rounded, crowned by collector hairs. Cypselae truncate or slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Pollen not seen. Habitat and Distribution. Genus of two species of Cuba, Haiti, Puerto Rico, and Dominican Republic. Species list 1. Berylsimpsonia crassinervis (Urb.) B. L. Turner, Phytologia 74: 352. 1993. Basionym: Proustia crassinervis Urb. TYPE: Unknown data (type, probably B). Additional specimen examined: Haiti. Dept. du Nord: vicinity of St. Michel de l’Atalaye, E. Leonard 7754 (US). 2. Berylsimpsonia vanillosma (C. Wright) B. L. Turner, Phytologia 74: 352. 1993. Basionym: Proustia vanillosma C. Wright. TYPE: Cuba. C. Wright 3616 (isotype, US not seen, US photo at LP!). Additional specimen examined: Puerto Rico. Municipio de Salinas: Barrio Lapa, Tetas de Cayey, R. King & G. Proctor 10601 (US). Literature. Fabris (1968), Turner (1993b).

Burkartia Crisci Bol. Soc. Argent. Bot. 17: 242. 1976. TYPE: Perezia lanigera Hook. & Arn. [= Burkartia lanigera (Hook. & Arn.) Crisci]. 49 of Figs. 47–50. Etymology. In honor of the Argentine botanist Arturo Burkart (1906–1975) who made important contributions to the knowledge of the flora of Argentina. Shrubs dwarf, cushion-like. Leaves densely imbricate; sessile, clasping; blades subulate, coriaceous, three to five parallel veins in the sheathing portion, reduced to one vein in the blade, margin entire, revolute, pubescent. Capitulescences monocephalous or two-headed; capitula sessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Pollen spheroidal, elliptic in equatorial view, circular in polar view, medium size, P × E = (43–47×34–37) μm. Tricolporate, colpi long with microgranulate membrane. Exine Trixis type, 5 μm at the equator, slightly slender at


613

the poles, occasionally with polar elevations. Nexine 1–1.5 μm thick (see also Crisci, 1974a sub Perezia lanigera). Habitat and Distribution. Monotypic genus endemic to the Patagonian steppe in Argentina. Species list

1. Burkartia lanigera (Hook. & Arn.) Crisci, Bol. Soc. Argent. Bot. 17: 243. 1976. Basionym: Perezia lanigera Hook. & Arn. TYPE: Argentina. Prov. Santa Cruz: Puerto Deseado (“Port Desire”), C. Darwin 314 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Chubut: Comodoro Rivadavia, Cañadón del Lagarto, A. Ruiz Leal 14704 (LP).

Literature. Crisci (1976a).

Calopappus Meyen Reise Chile 1: 315. 1834. TYPE: Calopappus acerosus Meyen. 50 of Figs. 47–50. Etymology. From the Greek kalos, beauty, and pappos, apical tuft of bristles on the fruit, by the appearance of the pappus. Shrubs dwarf, cushion-like. Leaves densely imbricate; sessile, clasping; blades subulate, coriaceous, three to five parallel veins in the sheathing portion, reduced to one vein in the blade, margin entire to dentate-spiny, glabrous to pubescent. Capitulescences monocephalous, terminal; capitula sessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae slightly constricted at the apex, glabrous; pappus of scabrid, narrowly paleaceous bristles. Pollen Description (Crisci, 1974a; Crisci & Freire, 1986). Pollen spheroidal-prolate to spheroidal-oblate, large pollen, tricolporate, exine Calopappus type. Habitat and Distribution. Monotypic genus endemic to the arid biogeographic region of Central Chile. Species list 1. Calopappus acerosus Meyen, Reise Chile 1: 315. 1834. TYPE: Chile. Prov. Colchagua: Talcaregue, 19..?, J. Gay s.n. (holotype, P not seen, P photo F n.

614

L. Katinas, et al.

38131 at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Dept. Los Andes, Saladillo, Río Blanco, O. Zöllner 1406 (CONC).

Literature. Crisci and Freire (1986). Cephalopappus Nees & Mart. Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 5. 1824. TYPE: Cephalopappus sonchifolius Nees & Mart. 51 of Figs. 51–54. Etymology. From the Greek kephale, head, and pappos, apical tuft of bristles on the fruit, probably in reference to the cypsela lacking a pappus. Herbs perennial, acaulescent. Leaves rosulate; pseudopetiolate to petiolate; blades spathulate, pinnately veined, margin sinuate-denticulate, glabrous to pubescent. Capitulescences loosely racemose; capitula long-pedunculate, sometimes scapiform, homogamous, discoid; receptacle epaleate; involucre one- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted to rostrate at the apex, glabrous, tuberculate; pappus absent. Pollen Description (110 of Figs. 107–111). Pollen spheroidal, circular in polar view, medium size, P = E = (33–38) μm. Tricolporate, colpi long with microgranulate membrane. Exine Cephalopappus type, 6 μm thick, slightly slender at the poles. Nexine ca.1.5 μm thick (see also Crisci, 1974a). Habitat and Distribution. Monotypic genus endemic to eastern Brazil, generally growing close to the rivers. Species list 1. Cephalopappus sonchifolius Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 5, tab. 1. 1824. TYPE: Brazil. Minas Gerais: ad flumen Ilhéos, Dec 1816, C. Martius s.n. (holotype, M not seen, M photo F n. 20737 at LP!). Additional specimen examined: Brazil. State Río de Janeiro: Río de Janeiro, Jardín Botánico, A. L. Cabrera 12243 (LP*). Literature. Baker (1884).

Criscia Katinas Bol. Soc. Argent. Bot. 30: 60. 1994. TYPE: Onoseris stricta Spreng. [= Criscia stricta (Spreng.) Katinas]. 52 of Figs. 51–54.


615

Figs. 51–54 51 Cephalopappus sonchifolius Nees & Mart. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela. (From Cabrera 12243, LP). Scale bars: A=6 cm; B, E=2 mm; C, D=1 mm. 52 Criscia stricta (Spreng.) Katinas. A Habit. B Marginal floret (without cypsela). C Central corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Katinas, 1994: 61). Scale bars: A=8 cm; B, C, F=7 mm; D, E=4 mm. 53 Dolichlasium lagascae D. Don. A Habit (branch). B Leaf. C Floret (without cypsela). D Stamen. E Upper part of the style. F Cypsela with pappus. (A–C, F redrawn from Cabrera, 1971b: 376; D, E from Spegazzini 1484, LP). Scale bars: A=2 cm; B=1 cm; C, F=10 mm; D, E=2.5 mm. 54 Holocheilus hieracioides (D. Don) Cabrera. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (From Nuñez 6, MCNS). Scale bars: A=5 cm; B, E=2.5 mm; C= 1.5 mm; D=1 mm

Etymology. In honor of the Argentine botanist Jorge V. Crisci (b. 1945) for his important contributions to our knowledge of the tribe Nassauvieae. Herbs perennial, acaulescent, scapose to scapiform. Leaves rosulate; pseudopetiolate; blades obovate to orbicular, pinnately veined, margin entire to denticulate, pubescent. Capitulescences monocephalous or few capitula, on scapes; capitula homogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid,

616

L. Katinas, et al.

branches apically rounded, crowned by collector hairs. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description (111 of Figs. 107–111). Pollen prolate, medium to large size, tricolporate, exine Trixis type (see also Crisci, 1974a). Habitat and Distribution. Monotypic genus from rocky and sandy places in northeastern Argentina, southern Brazil, and Uruguay. Species list 1. Criscia stricta (Spreng.) Katinas, Bol. Soc. Argent. Bot. 30: 62. 1994. Basionym: Onoseris stricta Spreng. TYPE: Uruguay. Montevideo, s.d., Sello s.n. (lectotype, designated by Katinas, 1994: 62, BM!). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Tandil, Villa del Lago, J. V. Crisci 545 (LP*). Literature. Katinas (1994).

Dolichlasium Lag. Amen. nat. Españ.: 33. 1811. TYPE: Dolichlasium lagascae D. Don. 53 of Figs. 51–54. Etymology. From Greek dolichos, long, and lasios, woolly, shaggy, because of the long, papillose anther tails. Shrubs dwarf. Leaves alternate; petiolate; blades oblong to elliptic, pinnatisect, pinnately veined, lobes palmatifid, glabrescent. Capitulescences monocephalous, terminal; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Exine Trixis type (see Crisci 1974a). Habitat and Distribution. Monotypic genus from the Andes of western Argentina. Species list 1. Dolichlasium lagascae D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Probably from Argentina. Prov. Mendoza: s.d., J. Gillies s.n. (types probably at BM, K, not seen). Additional specimen examined: Argentina. Prov. Mendoza: serranías de Uspallata, C. L. Spegazzini 1484 (LP).


617

Literature. Cabrera (1936, 1971b).

Holocheilus Cass. Bull. Sci. Soc. Philom. Paris: 73. 1818. TYPE: Perdicium brasiliense L. [= Holocheilus brasiliensis (L.) Cabrera]. 54 of Figs. 51–54. Cleanthes D. Don, Trans. Linn. Soc. London 16: 194. 1830. TYPE: Cleanthes hieracioides D. Don [= Holocheilus hieracioides (D. Don) Cabrera]. Platycheilus Cass., Dict. sci. nat. 34: 212. 1825. TYPE: Platycheilus ochroleucus Cass. [= Holocheilus brasiliensis (L.) Cabrera]. Etymology. From Greek holos, whole, all, entire, and cheilos, lip, because of the capitula with all the corollas bilabiate. Herbs perennial, caulescent to acaulescent, scapose. Leaves pinnately veined, margin entire to runcinate, glabrous to pubescent; basal leaves rosulate; pseudopetiolate; blades elliptic, obovate, oblanceolate to spathulate, entire to pinnatisect; upper leaves alternate, reduced to bracts; sessile to clasping. Capitulescences monocephalous or loosely to densely corymbose; capitula pedunculate to subsessile, homogamous, discoid; receptacle epaleate; involucre one- to two-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Exine Trixis type (see Crisci 1974a). Habitat and Distribution. Genus of seven species distributed in southern Brazil, Paraguay, Uruguay, and northern Argentina. Species list 1. Holocheilus brasiliensis (L.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 14. 1968. Basionym: Perdicium brasiliense L. TYPE: Brazil. s.d., Arduini s.n. (lectotype, designated by Katinas in Jarvis & Turland, 1998: 365, LINN 1003.4 not seen, LINN photo at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Pdo. Tornquist, Abra de la Ventana, L. Pertusi 214 (LP). 2. Holocheilus fabrisii Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 9. 1968. TYPE: Argentina. Prov. Jujuy: Dept. Santa Bárbara, cerro Centinela, H. A. Fabris et al. 5145 (holotype, LP!). 3. Holocheilus hieracioides (D. Don) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 7. 1968. Basionym: Cleanthes hieracioides D. Don. TYPE: Brazil: s.d., Sello s.n. (type, probably P). Additional specimen examined: Argentina. Prov. Salta: Dept. Capital, ruta 9 entre Castañares y Río Vaqueros, V. Nuñez 6 (MCNS).

618

L. Katinas, et al.

4. Holocheilus illustris (Vell.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 6. 1968. Basionym: Castra illustris Vell. TYPE: Tab. 81. T.8 (type, not localized). Additional specimen examined: Brazil. State Paraná: Mun. Punta Grossa, est. Velha, rod. Do Café-Itaiacoca, G. Hatschbach 11593 (LP). 5. Holocheilus monocephalus Mondin, Napaea 11: 31. 1995. TYPE: Brazil. State Rio Grande do Sul: mun. São José dos Ausentes, Rocinha, C. A. Mondin et al. 982 (holotype, ICN not seen). 6. Holocheilus pinnatifidus (Less.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 4. 1968. Basionym: Trixis pinnatifida Less. TYPE: Brazil: s.d., Sello s.n. (syntypes, probably P). Additional specimen examined: Brazil. State São Paulo: Bananal, serra da Bocaina-Sertão do río Vermelho, A. C. Brade & A. P. Duarte 20126 (LP). 7. Holocheilus schulzii (Cabrera) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 12. 1968. Basionym: Trixis schulzii Cabrera. TYPE: Argentina. Prov. Chaco: Colonia Benítez, A. Schulz 132 (holotype, LP!). Additional specimen examined: Argentina. Prov. Corrientes: estancia Garruchos, A. L. Cabrera 11840 (LP).

Literature. Cabrera (1936, 1968), Mondin (1995), Mondin and Vasques (2004), Katinas and Crisci (2008).

Jungia L. f., nom. cons. Suppl. pl.: 58. 1781 (1782). TYPE: Jungia ferruginea L. f. 55 of Figs. 55–58. Dumerilia Lag. ex DC., Ann. Mus. Natl. Hist. Nat. 19: 71. 1812, non Less., 1830. [= Jungia sect. Dumerilia (Lag. ex DC.) Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 32. 1995]. TYPE: Dumerilia axillaris Lag. ex DC. [= Jungia axillaris (Lag. ex DC.) Spreng.]. Martrasia Lag., Amen. nat. Españ.: 36. 1811. TYPE: not designated, the genus was based on the four species M. auriculata Lag., M. crenata Lag., M. pubescens Lag., and M. sericea Lag., all of them nomina nuda (Harling, 1995). Rhinactina Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 1: 139. 1807, nom. nudum. Tostimontia S. Díaz, Revista Acad. Colomb. Ci. Exact. 25: 9. 2001. TYPE: Tostimontia gunnerifolia S. Díaz (see Observation). Trinacte Gaertn., Fruct. sem. pl.: 415. 1791. TYPE: based on Jungia ferruginea L. f. Etymology. In honor of the German botanist Joachim Jungius (1587–1657), professor in Hamburg. Herbs perennial, subshrubs, or shrubs, erect to trailing. Leaves alternate to rosulate, stipulate to exstipulate; petiolate, rarely sessile; blades elliptic to suborbicular-cordate, palmately veined, rarely pinnately veined, margin entire to lobate, glabrous to pubescent. Capitulescences corymbose to paniculate; capitula


619

Figs. 55–58 55 Jungia pauciflora Rusby. A Habit (branch). B Palea. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (From Novara 339, MCNS). Scale bars: A=5 cm; B, C, F=2.5 mm; D=2 mm; E=0.8 mm. 56 Leucheria pteropogon (Griseb.) Cabrera. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Cabrera, 1978: 640). Scale bars: A=7 cm; B=5 mm; C=2 mm; D=1.5 mm; E=6 mm. 57 Leunisia laeta Phil. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (From Zöllner 429, LP). Scale bars: A=4 cm; B= 10 mm; C, D=3 mm; E=12 mm. 58. Lophopappus berberidifolius Cabrera. A Habit (branch). B Bilabiate corolla. C Tubular corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1953a: 48). Scale bars: A=4 cm; B, C, F=4 mm; D=2.5 mm; E=1 mm

pedunculate to subsessile, homogamous, discoid or radiate; receptacle paleaceous, paleae thin, scarious, embracing the central florets; involucre one- to two-seriate. Florets dimorphic or isomorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate, slightly constricted to attenuate at the apex, glabrous to pubescent; pappus of plumose, capillary bristles, occasionally barbellate to ciliate. Pollen Description. Jungia stuebelii (Hieron.) Crisci (Crisci, 1971). Pollen spheroidal, medium size, tricolporate, exine Trixis type, microechinate.

620

L. Katinas, et al.

Habitat and Distribution. Neotropical genus of 28 species, ranging from southwestern Mexico to northern Argentina, including the Galápagos Islands. Species list 1. Jungia axillaris (Lag. ex DC.) Spreng., Syst. veg. 4: 301. 1827. Basionym: Dumerilia axillaris Lag. ex DC. TYPE: In Peruvia, Chile et Panamaide, s. leg., ex herb. M. Lagasca s.n. (holotype, G-DC not seen). Additional specimen examined: Peru. s.d., C. Gay s.n. (LP). 2. Jungia beckii Harling, Novon 7: 246. 1997. TYPE: Bolivia. La Paz: Prov. Nor Yungas, ca. 4 km from Chuspipata towards Puente Villa, S. Beck 18672 (holotype, LPB not seen). 3. Jungia brachyphylla Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 124. 1953. TYPE: Colombia. Dept. Valle, cordillera al N de las Brisas, Carrizales, J. Cuatrecasas 22546 (holotype F, not seen). 4. Jungia calyculata Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 125. 1953. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, H. H. Smith 677 (isotype not seen, US not seen, US photo at LP!). 5. Jungia coarctata Hieron., Bot. Jahrb. Syst. 29: 81. 1900. TYPE: Ecuador. Prov. Pichincha: Lloa, A. Sodiro 184 (neotype, designated by Harling, 1991: 84, W not seen). Additional specimen examined: Peru. Dept. Amazonas: Prov. Chachapoyas, Puna-urcu, E-SE of Chachapoyas, J. J. Wurdack 690 (LP). 6. Jungia crenatifolia Harling, Fl. Ecuador 42: 93. 1991. TYPE: Ecuador. Prov. Azuay, Páramo de Tinajillas, ca. 10 km S of Cumbe, G. Harling 5681 (holotype S, not seen). 7. Jungia ferruginea L. f., Suppl. Pl.: 390. 1781 (1782). TYPE: Sine loc., J. C. Mutis 70 (holotype, LINN n. 1046.1 not seen). Additional specimen examined: Colombia. Bogotá: south side of ravine back of Parque Nacional, H. Schiefer 452 (LP). 8. Jungia fistulosa Hieron., Bot. Jahrb. Syst. 29: 83. 1900. TYPE: Ecuador. Prov. Cotopaxi: above Pilaló, G. Harling et al. 9003 (neotype, designated by Harling, 1991: 92, GB not seen). 9. Jungia floribunda Less., Linnaea 5: 38. 1830. TYPE: Brazil: 1926, Sello s.n. (lectotype, designated by Harling, 1995: 74, K not seen). Additional specimen examined: Paraguay. Dept. La Cordillera: camino Caacupé a Tobaty, A. Schinini 4683 (LP). 10. Jungia glandulifera Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 86. 1995. TYPE: Ecuador. Prov. Loja: Cariamanga, along road to Gonzanamá, G. Harling 6014 (holotype, S not seen). 11. Jungia gracilis Harling, Novon 2: 18. 1992. TYPE: Peru. Dept. Amazonas: Río Utcubamba valley, 31 km along road S of Tingo, L. R. M. King & L. E. Bishop 9278 (holotype, US not seen, US photo at LP!). Additional specimen examined: Brazil. Brasilia, DF: Reserva Ecológica do IBGE, B. A. S. Pereira 594 (LP). 12. Jungia hirsuta Cuatrec., Brittonia 8: 48. 1954. TYPE: Ecuador. Prov. Chimborazo: canyon of Río Chanchán, near Huigra, W. H. Camp 3011 (isotype, NY not seen, NY photo at LP!).


621

13. Jungia karstenii Cuatrec., Brittonia 8: 186. 1956. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, San Miguel, s.d., Karsten s.n. (holotype, P not seen). 14. Jungia mitis Benoist, Bull. Soc. Bot. France 84: 634. 1938. TYPE: Ecuador. Prov. Pichincha: Valle de Lloa, slopes of Unguí, Benoist 2677 (holotype, P not seen). 15. Jungia ovata Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 116. 1995. TYPE: Ecuador. Prov. Napo: road Quito-Baeza, 13 km E of Papallacta, Bohlin & Bohlin 676 (holotype, GB not seen). 16. Jungia paniculata (DC.) A. Gray, Proc. Amer. Acad. Arts 5: 145. 1861. Basionym: Dumerilia paniculata DC. TYPE: Peru: s.d., J. de Jussieu s.n., herb. Jussieu 9524 (holotype, P not seen). Additional specimen examined: Ecuador. Chimborazo: 9.7 km S of Chunchi, T. Stuessy & G. Nesom 5935a (LP). 17. Jungia pauciflora Rusby, Bull. New York Bot. Gard. 4: 401. 1907. TYPE: Bolivia. Dept. Cochabamba: vicinity of Cochabamba, 1891, M. Bang 2048 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Capital, San Lorenzo, L. Novara 339 (MCNS). 18. Jungia polita Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 215. 1879. TYPE: Bolivia. Dept. Tarija: between Tarija and Santa Ana, P. G. Lorentz & G. H. E. W. Hieronymus 952 (holotype, GOET, not seen). Additional specimen examined: Argentina. Prov. Córdoba: Dept. Minas, C. Sayago 3145 (LP). 19. Jungia pringlei Greenm., Publ. Field Columbian Mus., Bot. Ser. 2: 286. 1907. TYPE: Mexico. State Michoacán: near Uruapán, C. G. Pringle 10357 (isotype, US not seen, US photo at LP!). 20. Jungia rugosa Less., Linnaea 5: 36. 1830. TYPE:, Herb. Willdenow 15992 (lectotype, designated by Harling, 1995: 52, B not seen). Additional specimen examined: Peru. Cuzco: Prov. Paucastambo, quebrada de Llulluchayoc, C. Vargas C. 4309 (LP). 21. Jungia schuerae Harling, Novon 2: 16. 1992. TYPE: Peru. Dept. Lima: Prov. Huarochirí, Matucana, E. Asplund 11101 (holotype, S not seen). Additional specimen examined: Peru. Prov. Yungay: Dept. Ancash, arriba de Yungay, A. López M. et al. 7407 (LP). 22. Jungia sellowii Less., Syn. gen. Compos.: 416. 1832. TYPE: Sine loc., s.d., Sello s.n. [47] (neotype, designated by Harling, 1995: 79, P not seen). Additional specimen examined: Brazil. State Santa Catarina: Morro de Campo Alegre, São Francisco do Sul, P. Reitz & R. M. Klein 10958 (LP). 23. Jungia sordida J. Kost., Blumea 5: 683. 1945. TYPE: Bolivia. Abra de la Senda, T. Herzog 1839 (lectotype, designated by Harling, 1995: 64, L not seen). Additional specimen examined: Argentina. Prov. Jujuy: Laguna Yala, A. L. Cabrera & M. Marchionni 12968 (LP). 24. Jungia spectabilis D. Don, Trans. Linn. Soc. London 16: 227. 1830. TYPE: Ecuador. Prov. Guayas: Guayaquila, s.d., Tafalla s.n. (lectotype, designated by Harling, 1995: 83, BM not seen). Additional specimen examined: Ecuador. Prov. Manabi: near Jipijapa, O. Haught 3449 (LP). 25. Jungia stuebelii (Hieron.) Crisci, Bol. Soc. Argent. Bot. 13: 341. 1971. Basionym: Leucheria stuebelii Hieron. TYPE: Peru. Dept. Cajamarca: Prov.

622

L. Katinas, et al.

Cajamarca, Quinuamayo, entre Encañada y Celendín, A. Sagástegui 7385 (neotype, designated by Harling, 1995: 29, LP!). 26. Jungia vitocensis Cuatrec., Anales Ci. Univ. Madrid 4: 230. 1935. TYPE: Peru. Dept. Junín: Prov. Tarma, camino de Vitoc a los altos de Palca, J. Isern 462 (holotype, MA not seen). Additional specimen examined: Peru. Prov. Contumazá: Dept. Cajamarca, vicinity of Guzmando, A. Sagástegui 2901 (LP). 27. Jungia weberbaueri Cerrate, Publ. Mus. Hist. Nat. “Javier Prado,” Ser. B, Bot. 4: 22. 1951. TYPE: Peru. Dept. Huánuco: Prov. Pachitea, near Chaglla, A. Weberbauer 6701 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: Prov. Calea, Manto, Marin 2406 (LP). 28. Jungia woodii D. J. N. Hind, Kew Bull. 59: 311. 2004. TYPE: Bolivia. La Paz: Murillo, ca. 15–20 km E of summit on descent into Zongo valley, Wood 15029 (holotype, K not seen). Observation. The monospecific genus Tostimontia (Díaz-Piedrahita, 2001) from Sierra de Santa Marta in Colombia was distinguished from Jungia by the cespitose habit, peltate leaves with foliaceous bracts, and solitary capitula. All these characters, however, are found in Jungia and thus Tostimontia becomes a synonym of Jungia. Literature. Harling (1995, 1997), Hind (2004). Leucheria Lag. Amen. nat. Españ.: 32. 1811. TYPE: Leucheria hieracioides Cass. 56 of Figs. 55–58. Bertolonia DC. ex Cass., Opusc. phytol. 2: 153. 1826, nom. nudum, a name attributed to de Candolle by Cassini, but apparently never published by de Candolle (Crisci, 1976b). Cassiopea D. Don, Trans. Linn. Soc. London 16: 215. 1830, a name frequently listed as a synonym of Leucheria, but it was actually used only as a sectional name by Don and by de Candolle (Crisci, 1974a). Clybatis Phil., Anales Univ. Chile 41: 742. 1872. TYPE: Clybatis volkmanni Phil. [= Leucheria nutans (J. Rémy) Reiche]. Chabraea DC., Ann. Mus. Natl. Hist. Nat. 19: 65, 71. 1812. TYPE: Chabraea purpurea (Vahl) DC., based on Perdicium purpureum Vahl [= Leucheria purpurea (Vahl) Hook. & Arn.]. Eizaguirrea J. Rémy, Fl. chil. 3: 401. 1849. TYPE: Eizaguirrea candollei J. Rémy [= Leucheria floribunda DC.]. Frageria Delile ex Steud., Nomencl. bot. 1: 645. 1840, nom. nudum. Lasiorrhiza Lag., Amen. nat. Españ.: 32. 1811, pro parte, on the basis of two species: Perdicium purpureum (Vahl) Hook. & Arn. and Perdicium brasiliense L. [= Holocheilus brasiliensis (L.) Cabrera] (Crisci, 1976b). Leucaeria DC., Ann. Mus. Natl. Hist. Nat. 19: 66. 1812, an orthographic variant of the name Leucheria. Leuchaeria Less., Linnaea 5: 10. 1830, an orthographic variant of the name Leucheria.


623

Leuceria D. Don, Trans. Linn. Soc. London 16: 212. 1830, an orthographic variant of the name Leucheria. Leukeria Endl., Ench. bot.: 249. 1841, an orthographic variant of the name Leucheria. Maclovia DC. ex Steud., Nomencl. bot. 1: 338. 1840, nom. nudum. Mimela Phil., Anales Univ. Chile 27: 336. 1865. TYPE: Mimela pedicularifolia Phil. [= Leucheria scrobiculata D. Don]. Ptilurus D. Don, Trans. Linn. Soc. London 16: 218. 1830. TYPE: Ptilurus daucifolius D. Don [= Leucheria daucifolia (D. Don) Crisci]. Etymology. From the Greek leukon, white, and erion, wool, because of the pubescence of these plants. Herbs annual or perennial, caulescent to acaulescent and scapose, woolly to silky. Leaves pubescent; basal leaves rosulate to sub-rosulate; petiolate; blades linear to ovate, pinnately veined, margin entire to partite; upper leaves numerous, similar to the basal ones but reduced, or lacking; sessile. Capitulescences monocephalous, or corymbose to paniculate; capitula pedunculate, homogamous, discoid or radiate; involucre one- to five-seriate; receptacle epaleate to paleate, paleae embracing the marginal florets, with the open part facing either the center of the capitulum or the outer part of the capitulum. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of scabrid to plumose, capillary bristles. Pollen Description. Leucheria (112 of Figs. 112–116) (Crisci 1974a, 1976b). Pollen spheroidal, small or medium size, tricolporate, exine Oxyphyllum type (see also Parra & Marticorena, 1972; Markgraf & D’Antoni, 1978; Wingenroth & Heusser, 1983). Habitat and Distribution. Genus of 47 South American species distributed in the Andean region from Peru to the Islas Malvinas (Falkland Islands) and in Patagonia. Species list 1. Leucheria achillaeifolia Hook. & Arn., Comp. Bot. Mag. 2: 43. 1836. TYPE: Argentina. Prov. Santa Cruz: Port Desire, s.d., C. Darwin 391 (type, probably K). Additional specimen examined: Argentina. Prov. Neuquén: Sierra Auca Mahuida, H. A. Fabris 914 (LP). 2. Leucheria amoena Phil., Linnaea 28: 714. 1856. TYPE: Chile. Prov. Linares: cordillera de Linares, Jan 1857, R. Germain s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquén: Andacollo, Arroyo Guaraco, A. L. Cabrera 11152 (LP). 3. Leucheria apiifolia Phil., Anales Univ. Chile 87: 111. 1894. TYPE: Chile. Prov. Curicó: s.d., Vidal s.n. (holotype, SGO not seen, SGO photo at LP!). Additional specimen examined: Chile. Prov. Curicó: Dept. Curicó, a orillas de la Laguna de Teno, C. Marticorena & O. Matthei 892 (LP).

624

L. Katinas, et al.

4. Leucheria bridgesii Hook. & Arn., Comp. Bot. Mag. 1: 36. 1835. TYPE: Cordillera, Bridges 486 (holotype, P not seen, P photo in LP!). Additional specimen examined: Chile. Prov. Santiago: Pérez Caldera, C. Marticorena & O. Matthei 664 (LP). 5. Leucheria candidissima D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Chile. Sine loc., s.d., J. Gillies s.n. (isotype, GH not seen, GH photo at LP!). Additional specimen examined: Argentina. Prov. Neuquén: Andacollo, Cordillera del Viento, Cerro Corona, M. Jereb 17 (LP). 6. Leucheria cerberoana J. Rémy, Fl. chil. 3: 386. 1847. TYPE: Chile. Coquimbo, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Coquimbo: carretera Panamericana, a 3 km al N de Puerto Oscuro, C. Marticorena et al. 1423 (LP). 7. Leucheria coerulescens J. Rémy, Fl. chil. 3: 382. 1849. TYPE: Chile: s.d., C. Gay s.n. (isotype, P not seen). Additional specimen examined: Argentina. Prov. Río Negro: Isla Victoria, A. L. Cabrera 11503 (LP). 8. Leucheria congesta D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. San Juan: Dept. Calingasta, Cerro Castaño, H. A. Fabris & J. M. Marchionni 2339 (LP). 9. Leucheria cumingii Hook. & Arn., Comp. Bot. Mag. 1: 36. 1835. TYPE: Chile. Prov. Coquimbo: s.d., Cuming 906 (holotype, W not seen, W photo F n. 3286 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Punta Teatino, La Serena, Behn 8571 (LP). 10. Leucheria daucifolia (D. Don) Crisci, Darwiniana 20: 52. 1976. Basionym: Ptilurus daucifolius D. Don. TYPE: not localized. Additional specimen examined: Peru. Dept. Puno: Prov. Carabaya, Fauchinta, Allinccapacc, C. Vargas C. 7179 (LP). 11. Leucheria diemii Cabrera, Bol. Soc. Argent. Bot. 11: 288. 1969. TYPE: Argentina. Prov. Neuquén: Parque Nacional Nahuel Huapí, S del Cerro Fía, W. Diem 3311 (holotype, LP!). Additional specimen examined: Argentina. Prov. Neuquén: Parque Nacional Nahuel Huapi, Cerro Mesa, W. Diem 3602 (LP). 12. Leucheria eriocephala Speg., Anales Soc. Ci. Argent. 53: 31. 1902. TYPE: Argentina. Prov. Santa Cruz: Karr-aik, Lago Argentino, 1898, C. Ameghino s.n. (holotype, LP!). 13. Leucheria floribunda DC., Icon. sel. pl. 4: 88. 1839. TYPE: In excelsiis Andibus Chilensium legit cl. Gay (type, not localized). Additional specimens examined: Chile. Prov. Aconcagua: Valle Alista, al N de Río Blanco, O. Zöllner 3784 (LP); Cordillera de Santiago, Grandjot 3353 (GH*). 14. Leucheria garciana J. Rémy, Fl. chil. 3: 381. 1849. TYPE: Chile. Prov. Colchagua: Colchagua, C. Gay 314 (holotype, P not seen, P photo F n. 38113 at LP!). Additional specimen examined: Chile. Prov. Curicó: a orillas de la Laguna de Teno, C. Marticorena & O. Matthei 887 (LP). 15. Leucheria gayana (J. Rémy) Reiche, Fl. chil. 4: 429. 1905. Basionym: Chabraea gayana J. Rémy. TYPE: Chile: s.d., C. Gay s.n. (isotype, GH not seen, GH photo F n. 16045 at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Tunuyán, Valle del Alto Tunuyán, A. Ruiz Leal 2063 (LP).


625

16. Leucheria gilliesii Hook. & Arn., Comp. Bot. Mag. 1: 35. 1835. TYPE: Sine loc., s.d., s. leg. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Valle Hermoso, cerca del Refugio, Lagiglia 668 (LP). 17. Leucheria glabriuscula (Phil.) Reiche, Fl. chil. 4: 414. 1905. Basionym: Chabraea glabriuscula Phil. TYPE: Chile. Prov. Santiago: Cordillera de las Arañas, Dec 1854, Germain s.n. (isotype, LP!). 18. Leucheria glacialis (Poepp. ex Less.) Reiche, Anales Univ. Chile 116: 203. 1905. Basionym: Lasiorrhiza glacialis Poepp. ex Less. TYPE: Chile. Antuco, E. F. Poeppig 217 (holotype, P not seen, P photo F n. 20733 at LP!). Additional specimen examined: Chile. Prov. Talca: entre Laguna del Maule y Paso Pehuenches, M. Ricardi et al. 961 (LP). 19. Leucheria glandulosa D. Don, Trans. Linn. Soc. London 16: 217. 1830. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably G). Additional specimen examined: Chile. Prov. Valparaíso: Cuesta de Zapata, A. L. Cabrera 11445 (LP). 20. Leucheria graui Katinas, M. C. Tellería & Crisci, Novon 18: 366. 2008. TYPE: Chile. Prov. Maule, Laguna del Maule, J. Grau 2893 (holotype, LP!). 21. Leucheria hahnii Franch., Miss. Sci. Cape Horn, Bot. 5: 349. 1889. TYPE: Argentina. Prov. Tierra del Fuego: Canal del Beagle, Hahn 89 (holotype, P not seen, P photo F n. 38114 at LP!). Additional specimen examined: Argentina. Prov. Santa Cruz: Cerro Cazador, M. Gentili 417 (LP). 22. Leucheria hieracioides Cass., Dict. sci. nat. 55: 392. 1828. TYPE: Chile. s.d., d’Urville s.n. (holotype, P not seen). Additional specimen examined: Chile. Prov. Santiago, Dept. Santiago, Cuesta de Chacabuco, C. Marticorena & E. Weldt 615 (LP). 23. Leucheria integrifolia (Phil.) Crisci, Darwiniana 16: 628. 1971. Basionym: Chabraea integrifolia Phil. TYPE: Chile. Prov. Linares: Andes de Linares, s.d., R. Germain s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Linares: precordillera, O. Zöllner 2461 (LP). 24. Leucheria landbeckii (Phil.) Reiche, Fl. chil. 4: 423. 1905. Basionym: Chabraea landbeckii Phil. TYPE: Chile. Prov. Santiago: Cordillera de Las Arañas, 1861, Landbeck s.n. (holotype, LP!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, Real de los Trece, Estancia Lloncha, A. Ruiz Leal 7154 (LP). 25. Leucheria leontopodioides (Kuntze) K. Schum., Just’s Bot. Jahresber. 26: 378. 1900. Basionym: Lasiorrhiza leontopodioides Kuntze. TYPE: Argentina. Patagonia, s.d., Moreno & Tonini 149 (type, not localized). Additional specimen examined: Argentina. Prov. Santa Cruz: Cerro Calafate, H. Sleumer 1163 (LP). 26. Leucheria lithospermifolia (Less.) Reiche, Anales Univ. Chile 116: 208. 1905. Basionym: Lasiorrhiza lithospermifolia Less. TYPE: Chile. Prov. Bío Bío: Sierra Velluda, Andes de Antuco, E. F. Poeppig 215 (holotype, P not seen). Additional specimen examined: Chile. Prov. Bío Bío: Laguna Laja, O. Zöllner 5526 (LP). 27. Leucheria magna Phil., Anales Univ. Chile 87: 108. 1894. TYPE: Chile. Prov. Aysén: Valle del Río Palena, s.d., F. Delfín s.n. (holotype, SGO, not

626

28.

29.

30.

31.

32.

33.

34.

35.

36.

37.

38.

L. Katinas, et al.

seen, SGO photo at LP!). Additional specimen examined: Chile. Prov. Aysén: Aysén, M. Bruzzone 149 (LP). Leucheria menana J. Rémy, Fl. chil. 3: 387. 1849. TYPE: Chile. Prov. Coquimbo: peñascos marítimos de Coquimbo, C. Gay 916 (holotype, P not seen, P photo F n. 16053 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: La Serena, E. Barros 2653 (LP). Leucheria millefolium Dusén & Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 333. 1916. TYPE: Argentina. Prov. Chubut: región del Lago General Pico, Valle Frías, Hogberg 12 (isolectotype designated by Crisci, 1976b: 48, BAF not seen). Additional specimen examined: Argentina. Prov. Río Negro: Cerro Anecón, Ferruglio 54 (LP). Leucheria multiflora Phil., Anales Univ. Chile 41: 743. 1872. TYPE: Chile. Prov. Aconcagua: Los Molles, Philippi 1126 (isotype, LP!). Additional specimen examined: Chile. Prov. Aconcagua: Quintero, El Durazno, Junge 6846 (LP). Leucheria nutans (J. Rémy) Reiche, Anales Univ. Chile 116: 208. 1905. Basionym: Chabraea nutans J. Rémy. TYPE: Chile. Prov. Concepción: cordillera, C. Gay 386 (holotype, P not seen). Additional specimen examined: Argentina. Prov. Río Negro: Cerro Catedral, U. Eskuche 02–0 (LP). Leucheria oligocephala J. Rémy, Fl. chil. 3: 383. 1849. TYPE: Chile: s.d., C. Gay s.n. (isotype, P not seen). Additional specimen examined: Chile. Prov. Santiago: Cerros de Peñalolén, Mahu 997 (LP). Leucheria paniculata Poepp. ex Less., Syn. gen. Compos.: 403. 1832. TYPE: Chile. Prov. Bío Bío: Cordillera de Antuco, E. F. Poeppig 218 (isotype, NY, NY photo at LP!). Leucheria papillosa Cabrera, Revista Sudamer. Bot. 3: 58. 1936. TYPE: Argentina. Prov. Río Negro: Cerro López, A. L. Cabrera & M. M. Job 339 (holotype, LP!). Additional specimen examined: Argentina. Prov. Río Negro: región del Lago Nahuel Huapi, Paso de las Nubes, A. L. Cabrera 5905 (LP). Leucheria polyclados (J. Rémy) Reiche, Fl. chil. 4: 415. 1905. Basionym: Chabraea polyclados J. Rémy. TYPE: Without data (holotype, P 17, 1/70 not seen). Additional specimen examined: Chile. Prov. Atacama: cercanías de Plaza, A. Krapovickas & J. Hunziker 5792 (LP). Leucheria pteropogon (Griseb.) Cabrera, Darwiniana 9: 62. 1949. Basionym: Senecio pteropogon Griseb. TYPE: Argentina. Prov. Salta: alrededores del Nevado del Castillo, P. G. Lorentz & G. H. E. W. Hieronymus 101 (isotype, LP!). Additional specimen examined: Argentina. Prov. Catamarca: faldeo S de las Cumbres de la Sierra de Granadillas, Quebrada de los Potrerillos, H. Sleumer & Vervoorst 2633 (LP). Leucheria purpurea (Vahl) Hook. & Arn., Comp. Bot. Mag. 2: 43. 1836. Basionym: Perdicium purpureum Vahl. TYPE: Argentina. Prov. Tierra del Fuego: Magallanes, s.d., D. Commerson s.n. (lectotype, designated by Crisci, 1976b: 43, P not seen). Additional specimen examined: Argentina. Prov. Santa Cruz: Esperanza, H. Sleumer 1085 (LP). Leucheria rosea Poepp. ex Less., Syn. gen. Compos.: 402. 1832. TYPE: Chile. Puente Vizcachas-La Guardia, s.d., E. F. Poeppig 574 (holotype, P not


39.

40.

41.

42.

43.

44.

45.

46.

47.

627

seen, P photo F n. 20732 at LP!). Additional specimen examined: Chile. Prov. Santiago: Com. Las Condes, Cerro Naranjo, A. L. Cabrera 483 (LP). Leucheria runcinata D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: entre Las Cuevas y Cristo Redentor, H. A. Fabris 1254 (LP). Leucheria salina (J. Rémy) Hieron., Bol. Acad. Nac. Ci. 4: 50. 1881. Basionym: Chabraea salina J. Rémy. TYPE: Chile: s.d., C. Gay s.n. (isotype, GH not seen, GH photo F n. 16058 at LP!). Additional specimen examined: Argentina. Prov. La Rioja: Dept. General Sarmiento, Quebrada La Hedionda, J. H. Hunziker 2207 (LP). Leucheria scrobiculata D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: Arroyo de los Leones, A. Ruiz Leal 11743 (LP). Leucheria senecioides Hook. & Arn., Bot. Beechey Voy. 1: 28. 1830. TYPE: Chile. Concepción, s.d., s. leg. (type, probably K). Additional specimen examined: Chile. Prov. Concepción: Fundo Valencia, G. Looser 3991 (LP). Leucheria suaveolens (d’Urv.) Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 538. 1897. Basionym: Perdicium suaveolens d’Urv. TYPE: not localized. Additional specimen examined: Argentina. Islas Malvinas (Falkland Islands): East Falkland, Mt. Tumbledown, W of Port Stanley, D. M. Moore 519 (LP). Leucheria tenuis Less., Syn. gen. Compos.: 401. 1832. TYPE: Chile. Prov. O’Higgins: Rancagua, Monte La Leona, C. Bertero 159 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Tabaco, O. Zöllner s.n. (LP). Leucheria thermarum (Phil.) Phil., Verh. Deutsch. Wiss. Verein Santiago de Chile 2: 203. 1892. Basionym: Chabraea thermarum Phil. TYPE: Chile. Prov. Ñuble: Baños de Chillán, s.d., Philippi s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Río Negro: Dept. Bariloche, Cerro Otto, J. V. Crisci 533 (LP). Leucheria tomentosa (Less.) Crisci, Darwiniana 20: 114. 1976. Basionym: Lasiorrhiza tomentosa Less. TYPE: Chile. Prov. Valparaíso: Valparaíso, E. F. Poeppig 217 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Ligua, E. Barros 7435 (LP). Leucheria viscida (Bertero ex Colla) Crisci, Darwiniana 20: 67. 1976. Basionym: Chabraea viscida Bertero ex Colla. TYPE: Chile. Prov. O’Higgins: Rancagua, Tagua Tagua, C. Bertero 160 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Curicó: Cerro Condell, E. Barros 2619 (LP).

Literature. Crisci (1976b), Katinas et al. (2008a, b)

Leunisia Phil. Anales Univ. Chile 23: 383. 1863; Linnaea 33: 120. 1864. TYPE: Leunisia laeta Phil. 57 of Figs. 55–58.

628

L. Katinas, et al.

Etymology. In honor of the German clergyman and botanist Johannis Leunis (1802– 1873). Shrubs. Leaves alternate; sessile; blades lanceolate to oblong, pinnately veined, margin entire to dentate, pubescent. Capitulescences monocephalous, terminal; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically rounded, crowned by collector hairs. Cypselae attenuate at the apex, pubescent; pappus paleaceous– setaceous, scabrid bristles, unequally fused at the base. Pollen Description (Parra & Marticorena, 1972; Crisci, 1974a). Pollen prolate, medium or large size, tricolporate, exine Trixis type. Habitat and Distribution. Monotypic genus endemic to the Andean region of central Chile. Species list 1. Leunisia laeta Phil., Anales Univ. Chile 23: 383. 1863. TYPE: Chile. Prov. Coquimbo: Huatalame, s.d., H. Volkmann s.n. (holotype, G not seen, G photo F n. 28865 at LP!). Additional specimen examined: Chile Prov. Aconcagua: Cordón Caquis, Cerro Chache, O. Zöllner 429 (LP). Literature. Reiche (1905).

Lophopappus Rusby Bull. Torrey Bot. Club 21: 487. 1894, emend. Cabrera, Bol. Soc. Argent. Bot. 5: 45. 1953. TYPE: Lophopappus foliosus Rusby. 58 of Figs. 55–58. Etymology. From the Greek lophos, crest, and pappos, apical tuft of bristles on the fruit, with reference to the pappus bristles plumose at the apex. Shrubs, stems with brachyblasts and macroblasts. Leaves alternate; sessile; blades linear to elliptic, pinnately veined, margin entire to spinose-dentate, glabrous to pubescent. Capitulescences monocephalous or in loose to dense apical cymes; capitula short-pedunculate, homogamous, discoid or disciform; receptacle epaleate; involucre multiseriate. Florets isomorphic or dimorphic, bisexual, corollas all or partly tubular-funnelform, five-lobed, or all or partly bilabiate; anther apical appendages acute, tails papillose; style bifid, branches rounded at the apex, dorsally papillose. Cypselae attenuate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles, plumose at the apex. Pollen Description. Lophopappus (113 of Figs. 112–116) (Tellería et al., 2003). Pollen subprolate to prolate-spheroidal, medium to large size, tricolporate, exine


629

Trixis type, Proustia type, or both types in a single specimen, with polar caps only in L. berberidifolius Cabrera and L. peruvianus Cabrera (see also Parra & Marticorena, 1972; Crisci, 1974a). Habitat and Distribution. Genus of six species distributed in the Andes from Peru to Argentina. Species list 1. Lophopappus berberidifolius Cabrera, Bol. Soc. Argent. Bot. 5: 48. 1953. TYPE: Peru. Dept. Huancavelica: Sachahuacta, 7 km al SO de Conaica, O. Tovar 961 (holotype, LP!). 2. Lophopappus blakei Cabrera, Bol. Soc. Argent. Bot. 5: 48. 1953, nom. novum. Basionym: Proustia cuneata S. F. Blake. TYPE: Peru. Dept. Cuzco: Ollantaytambo, O. F. Cook & G. B. Gilbert 538 (holotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: desvío km 82, Estancia Piscacucho, A. L. Cabrera & H. A. Fabris 13462 (LP). 3. Lophopappus cuneatus R. E. Fr., Ark. Bot. 5: 29. 1906. TYPE: Bolivia. Quebrada Honda, pr. Tarija, R. E. Fries 1045 (isotype, US not seen, US photo at LP!). Additional specimen examined: Argentina. Jujuy: Dept. Cochinoca, Abra Pampa, Cerro Huancar, A. L. Cabrera 15425 (LP). 4. Lophopappus foliosus Rusby, Bull. Torrey Bot. Club 21: 487. 1894. TYPE: Bolivia. Dept. La Paz: vicinity of La Paz, M. Bang 66 (isosyntype, US not seen, US photo at LP!). Additional specimen examined: Bolivia. Dept. La Paz: Valle de la Luna, A. L. Cabrera 2570 (LP). 5. Lophopappus peruvianus Cabrera, Bol. Soc. Argent. Bot. 5: 50. 1953. TYPE: Peru. Dept. Ancash: Prov. Bolognesi, cerca de Aquia, E. Cerrate 1552 (holotype, LP!). 6. Lophopappus tarapacanus (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 157. 1971. Basionym: Gochnatia tarapacana Phil. TYPE: Chile. Prov. Tarapacá: Sotoca, s.d., Philippi s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Tarapacá: a+- 10 km volviendo de Tranque de Caritaya, M. Ricardi & C. Marticorena 25604 (LP*). Observation. Lophopappus has been also placed in the subtribes Mutisiinae and Gochnatiinae (Cabrera, 1961), because of the tubular florets and rounded, dorsally papillose style branches. The type of exine stratification in pollen of Lophopappus (Crisci, 1974a; Tellería et al., 2003) clearly indicates that this genus belongs to the subtribe Nassauviinae (= tribe Nassauvieae). Style and corolla features link Lophopappus to the genus Proustia, and some authors (Ferreyra, 1995) consider that Lophopappus should be merged with Proustia. Literature. Cabrera (1953a, 1971b).

Macrachaenium Hook. f. Fl. antarct. 2: 321. 1847. TYPE: Macrachaenium gracile Hook. f. 59 of Figs. 59–62.

630

L. Katinas, et al.

Figs. 59–62 59 Macrachaenium gracile Hook. f. A Habit. B Marginal corolla. C Central corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1971b: 324). Scale bars: A=2 cm; B, C, F=2 mm; D, E=1 mm. 60 Marticorenia foliosa (Phil.) Crisci. A Habit (branch). B Palea. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Crisci, 1974b: 41). Scale bars: A=5 cm; B=3.5 mm; C=7 mm; D=4 mm; E=1 mm; F=10 mm. 61 Moscharia pinnatifida Ruiz & Pav. A Habit. B Pseudocephalium. C Marginal capitulum. D Floret of marginal capitulum (without cypsela). E Central capitulum. F Stamen. G Upper part of the style. H Cypsela with pappus. (Redrawn from Crisci, 1974c: 170). Scale bars: A=9 cm; B=10 mm; C, D=4 mm; E, F=2 mm; G=0.6 mm; H=1.5 mm. 62 Nassauvia axillaris (Lag. ex Lindl.) D. Don. A Habit. B Capitulum. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Cabrera, 1978: 633). Scale bars: A=8 cm; B=7 mm; C, F=3 mm; D=1 mm; E=0.6 mm

Etymology. From the Greek makros, big, and achaena or achaenium, the typical fruit of Asteraceae, in reference to the size of the cypselae. Herbs perennial, acaulescent, with one or more scapes. Leaves pinnately veined, pubescent; basal leaves rosulate, base inflated and vaginate; pseudopetiolate; blades broadly elliptic, margin runcinate to pinnately partite; upper leaves similar to the basal ones but reduced, or lacking; sessile to decurrent. Capitulescences mono-


631

cephalous, on scapes; capitula homogamous, discoid or radiate; receptacle epaleate; involucre two- to three-seriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages truncate to acute, tails smooth; style bifid, branches rounded at the apex, dorsally papillose. Cypselae attenuate at the apex, glabrous; pappus of plumose, capillary bristles. Pollen Description. Pollen subprolate, elliptic in equatorial view, circular in polar view, large size, P × E = (55–63×42–49) μm. Tricolporate, colpi long with psilate membrane, mesoaperturate. Exine Trixis type, 6–8 μm at the equator slightly slender at the poles, with polar elevations. Nexine 1.2–2 μm thick. (see also Parra & Marticorena, 1972; Crisci, 1974a; Tellería et al., 2003). Habitat and Distribution. Monotypic genus with two varieties endemic to Nothofagus forests in southern Argentina and Chile. Species list 1. Macrachaenium gracile Hook. f., Fl. antarct. 2: 321. 1847. TYPE: Argentina. Prov. Río Negro: Dept. Bariloche, Lago Nahuel Huapi, Laguna Frías, cerro Riggi, A. L. Cabrera 6048 (holotype of M. gracile var. radiatum, LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: R. Goodall 283 (LP). Observation. Macrachaenium has sometimes been placed in the subtribe Mutisiinae (Cabrera, 1961) because of its rounded, dorsally papillose style branches. The type of exine stratification in the pollen of Macrachaenium (Crisci, 1974a; Tellería et al., 2003) clearly indicates the position of this genus within the Nassauviinae (= tribe Nassauvieae). Literature. Cabrera (1971b), Katinas (1995).

Marticorenia Crisci J. Arnold Arbor. 55: 38. 1974. TYPE: Leuceria foliosa Phil. [= Marticorenia foliosa (Phil.) Crisci]. 60 of Figs. 59–62. Etymology. In honor of the Chilean botanist Clodomiro Marticorena (b. 1929) who made important contributions to the knowledge of the flora of Chile. Shrubs. Leaves alternate; sessile, decurrent to clasping; blades ovate-elliptic, pinnately veined, margin lobulate, pubescent. Capitulescences a dichotomous cyme; capitula pedunculate, homogamous, discoid; receptacle paleaceous, paleae embracing all the florets; involucre one- to two-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of plumose, capillary bristles.

632

L. Katinas, et al.

Pollen Description. Pollen subprolate to prolate, medium size, tricolporate, exine Trixis type (see Crisci, 1974b). Habitat and Distribution. Monotypic genus endemic to the biogeographic region of Central Chile. Species list 1. Marticorenia foliosa (Phil.) Crisci, J. Arnold Arbor. 55: 40. 1974. Basionym: Leuceria foliosa Phil. TYPE: Chile. Prov. Santiago: Las Arañas, Cajón del Mapocho, R. Philippi 1132 (neotype, designated by Crisci, 1974b: 42, LP!). Additional specimen examined: Chile. Prov. Aconcagua, Valle Castro, cerca de Río Blanco, O. Zöllner 2994 (LP). Literature. Crisci (1974b).

Moscharia Ruiz & Pav., nom. cons. Fl. peruv. prodr.: 91. 1794, non Forssk., 1775. TYPE: Moscharia pinnatifida Ruiz & Pav. 61 of Figs. 59–62. Moscaria Pers., Syn. pl. 2: 379. 1807, an orthographic variant of the name Moscharia. Moschifera Molina, Sag. stor. nat. Chili: 294. 1810, an orthographic variant of the name Moscharia. Mosigia Spreng., Syst. veg. 3: 366. 1828, nom. superfluum. Gastrocarpha D. Don, Brit. Fl. Gard. 3: 229. 1827–1829. TYPE: Gastrocarpha runcinata D. Don [= Moscharia pinnatifida Ruiz & Pav.]. Etymology. From the Greek moschos, musky, because of the strong musky fragrance of the plant. Herbs annual. Leaves pinnately veined, glabrous to pubescent; basal leaves subrosulate; petiolate; blades oblanceolate, pinnately parted; upper leaves petiolate to amplexicaul; blades lanceolate, margin entire to lobed. Capitulescences a capituliform pseudocephalium; capitula subsessile, homogamous, discoid; outer capitula of the pseudocephalia with two florets, the involucre of two phyllaries, the outer phyllary keeled, embracing the two florets, the inner phyllary not embracing and placed between the two florets and interpreted as a palea; central capitulum one- to five-flowered, involucre uniseriate; receptacle paleate or epaleate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, pubescent; pappus of very short, crown-like, scabrid bristles, narrowly paleaceous, or pappus absent in the outer cypselae. Pollen Description. Moscharia (Parra & Marticorena, 1972). Pollen spheroidalprolate, medium size, tricolporate, exine Oxyphyllum type (see also Crisci, 1974a).


633

Habitat and Distribution. Genus of two species endemic to the biogeographic region of Central Chile. Species list 1. Moscharia pinnatifida Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 186. 1798. TYPE: Habitat in Regni Chilensis, aridis arenosis et petrosis locis ad Aconcaguae, Quillotae et Rancaguae provincias, et copiosè versus S. Jacobi Chilensis urbem (type, not localized). Additional specimen examined: Chile. Región Metropolitana: Reserva Nacional Río Clarillo, M. Bonifacino 1321 (LP). 2. Moscharia solbrigii Crisci, Contr. Gray Herb. 205: 172. 1974. TYPE: Chile. Prov. Coquimbo: Dept. Ovalle, south end Fray Jorge forest, J. L. Morrison & C. R. Worth 16441 (holotype, GH not seen). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Parque Nacional Fray Jorge, C. Marticorena et al. 521 (LP). Literature. Crisci (1974c), Hellwig (1985), Katinas and Crisci (2000), Katinas et al. (2008a).

Nassauvia Comm. ex Juss. Gen. pl.: 175. 1789. TYPE: Nassauvia magellanica J. F. Gmel. 62 of Figs. 59–62. Acanthophyllum Hook. & Arn., Comp. Bot. Mag.: 37. 1835. TYPE: Acanthophyllum axillare (Lag. ex Lindl.) Hook. & Arn., based on Triptilion axillare Lag. ex Lindl. [= Nassauvia axillaris (Lag. ex Lindl.) D. Don]. Caloptilium Lag., Amen. nat. Españ.: 34. 1811. [= Nassauvia sect. Caloptilium (Lag.) Benth., in Benth. & Hook., Gen. pl. 2: 503. 1873]. TYPE: Caloptilium lagascae (D. Don) Hook. & Arn., based on Sphaerocephalus lagascae D. Don [= Nassauvia lagascae (D. Don) F. Meigen var. lagascae]. Mastigophorus Cass., Dict. sci. nat. 34 222. 1825. [= Nassauvia sect. Mastigophorus (Cass.) DC., Prodr. 7: 50. 1838]. TYPE: Mastigophorus gaudichaudii Cass. [= Nassauvia gaudichaudii (Cass.) Cass. ex Gaudich.]. Nassovia Batsch, Tab. affin. regni veg. 251: 1802, an orthographic variant of the name Nassauvia. Panargyrum DC., Ann. Mus. Natl. Hist. Nat. 19: 9. 1812, an orthographic variant of the name Panargyrus established by de Candolle and used by most researchers, including Cabrera (1982). Panargyrus Lag., Amen. nat. Españ.: 33. 1811. [= Nassauvia sect. Panargyrum (Lag.) Wedd., Chlor. andina 1: 52. 1855]. TYPE: Panargyrum lagascae DC. [= Nassauvia darwinii (Hook. & Arn.) O. Hoffm. & Dusén]. Pentanthus Less., Syn. gen. Compos.: 398. 1832. TYPE: Pentanthus aculeatus Less. [= Nassauvia aculeata (Less.) Poepp. & Endl. var. aculeata]. Piptostemma Spach, Hist. nat. veg. 10: 34. 1841, a name attributed to Don by Spach and considered a synonym of Nassauvia, but never published by Don. The name was retaken by de Candolle as a section of Nassauvia (Crisci, 1974a; Cabrera, 1982).

634

L. Katinas, et al.

Portalesia Meyen, Reise Chile 1: 316. 1834. TYPE: Portalesia procumbens Meyen [= Nassauvia lagascae (D. Don) F. Meigen var. lagascae]. Sphaerocephalus Lag. ex DC., Ann. Mus. Natl. Hist. Nat. 19: 67. 1812. TYPE: Sphaerocephalus lagascae D. Don. [= Nassauvia lagascae (D. Don) F. Meigen var. lagascae]. Strongyloma DC., Prodr. 7: 52. 1838. [= Nassauvia subgen. Strongyloma (DC.) Cabrera, Darwiniana 24: 364. 1982]. TYPE: Strongyloma axillare (Lag. ex Lindl.) DC., based on Triptilion axillare Lag. ex Lindl. [= Nassauvia axillaris (Lag. ex Lindl.) D. Don]. Strongylomopsis Speg., Comun. Mus. Nac. Buenos Aires 1: 135. 1899. TYPE: Strongylomopsis fuegiana Speg. [= Nassauvia fuegiana (Speg.) Cabrera]. Triachne Cass., Bull. Sci. Soc. Philom. Paris: 11. 1818. TYPE: Triachne pygmaea Cass. [= Nassauvia pygmaea (Cass.) Hook. var. pygmaea]. Trianthus Hook. f., Fl. antarct. 2: 320. 1847. TYPE: Trianthus ulicinus Hook. f. [= Nassauvia ulicina (Hook. f.) Macloskie]. Etymology. In honor of the prince of Nassau, who took part in the trip of Bougainville around the world in 1766. Herbs perennial, subshrubs or shrubs, sometimes cushion-like, frequently with brachyblasts and macroblasts. Leaves imbricate; sessile to clasping; blades linearsubulate to ovate, several parallel veins reduced in number in the linear blades, margin entire to partite, spiny to unarmed, sometimes revolute, glabrous to pubescent. Capitulescences monocephalous, racemose, spicate, to glomerulose or pseudocephalium; capitula sessile to pedunculate, homogamous, discoid or radiate, five-flowered, occasionally one- to four-flowered; receptacle epaleate; involucre two-seriate. Florets isomorphic, bisexual, corolla bilabiate, outer lip expanded or not; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus of bristles filiform to narrowly paleaceous, laciniate to plumose, rarely crown-like. Pollen Description. Nassauvia (Parra & Marticorena, 1972; Crisci, 1974a). Pollen spheroidal, prolate to suboblate, small to medium size, tricolporate, exine Oxyphyllum type (see also Markgraf & D’Antoni, 1978; Wingenroth & Heusser, 1983). Habitat and Distribution. Genus of 38 South American species distributed in the Andes and Patagonia, from southern Bolivia to the Islas Malvinas (Falkland Islands). Species list 1. Nassauvia aculeata (Less.) Poepp. & Endl., Nov. gen. sp. pl. 1: 12. 1835. Basionym: Pentanthus aculeatus Less. TYPE: Chile. Andes de Antuco, Sierra Velluda, E. F. Poeppig 227 (lectotype, designated by Cabrera, 1982: 357, P not seen). Additional specimen examined: Argentina. Prov. Neuquén: Pino Hachado, A. L. Cabrera & J. V. Crisci 19113 (LP).


635

2. Nassauvia ameghinoi Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 542. 1897. TYPE: Argentina. Prov. Santa Cruz: San Jorge, Feb 1896, C. Ameghino s.n. (holotype, LP!). 3. Nassauvia argentea Phil., Anales Univ. Chile 87: 82. 1894. TYPE: Chile. Prov. Valdivia: Huahim, Jan 1887, O. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Argentina. Prov. Neuquén: San Martín de los Andes, Cerro Chapelco, A. L. Cabrera 19722 (LP). 4. Nassauvia argyrophylla Speg. ex Cabrera, Notas Mus. La Plata, Bot. 4: 160 1939. TYPE: Argentina. Prov. Chubut: Río Corcovado, 1/15 March 1901, N. Illín s.n. (holotype, LP!). 5. Nassauvia axillaris (Lag. ex Lindl.) D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. Basionym: Triptilion axillare Lag. ex Lindl. TYPE: Chile, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, Quebrada del Paso de la Cruz de Piedra, A. Ruiz Leal 11765 (LP). 6. Nassauvia chubutensis Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 615. 1897. TYPE: Argentina. Prov. Chubut: Teka-choique, 1889, C. Moyano s.n. (holotype, LP!). 7. Nassauvia coronipappa Arroyo & Martic., Brittonia 40: 332. 1988. TYPE: Chile. Ultima Esperanza: Cerro Donoso, Río de las Chinas valley, M. T. K. Arroyo et al. 87314 (holotype, CONC!). 8. Nassauvia cumingii Hook. & Arn., Comp. Bot. Mag. 1: 37. 1835. TYPE: Cordillera, s.d., H. Cuming 237 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Las Heras, Las Cuevas, refugio militar General Lamadrid, O. Boelcke et al. 9818 (LP). 9. Nassauvia darwinii (Hook. & Arn.) O. Hoffm. & Dusén, Wiss. Erg. Schwed. Exp. Magellansl. 3: 112. 1900. Basionym: Panargyrum darwinii Hook. & Arn. TYPE: Argentina. Prov. Santa Cruz: Port Desire, C. Darwin 313 (lectotype, designated by Cabrera, 1982: 351, K not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Punta María flat, between Viamonte and Río Grande, R. N. Goodall 316 (LP). 10. Nassauvia dentata Griseb., Abh. Königl. Ges. Wiss. Göttingen 6: 125. 1854. TYPE: Sine loc., R. Philippi 125 (isotypes, K not seen, P not seen). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Lácar, Hua-Hum, Cerro Malo, J. H. Hunziker 6996 (LP). 11. Nassauvia digitata Wedd., Chlor. andina 1: 48. 1855. TYPE: Chile: s.d., C. Gay s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Ñorquín, Termas de Copahue, A. L. Cabrera 6195 (LP). 12. Nassauvia dusenii O. Hoffm., Rep. Princeton Univ. Exp. Patagonia, Botany 8: 879. 1906. TYPE: Chile. Patagonia occidentalis, Río Aysen, P. K. Dusén 584 (isotype, B not seen, B photo F n. 16032 at LP!). Additional specimen examined: Argentina. Prov. Chubut: Dept. Futaleufú, Esquel, La Hoya, A. L. Cabrera et al. 23132 (LP). 13. Nassauvia fuegiana (Speg.) Cabrera, Physis (Buenos Aires) 10: 285. 1931. Basionym: Strongylomopsis fuegiana Speg. TYPE: Argentina. Prov. Tierra del Fuego: Golfo San Sebastián, Jan 1896, O. Mauri s.n. (holotype, LP!).

636

L. Katinas, et al.

14. Nassauvia gaudichaudii (Cass.) Cass. ex Gaudich., Ann. Sci. Nat. (Paris) 5: 103. 1825. Basionym: Mastigophorus gaudichaudii Cass. TYPE: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), April 1825, C. Gaudichaud s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), East Falkland, Eliza Cove, ca. 1 1/2 miles S of Port Stanley, D. M. Moore 559 (LP). 15. Nassauvia glomerata (D. Don) Wedd., Chlor. andina 1: 53. 1855. Basionym: Panargyrum glomeratum D. Don. TYPE: Chile. Andes of Chile, Valle de los Ciegos, s.d., J. Gillies s.n. (lectotype, designated by Cabrera, 1982: 361, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Malalhue, Alto Valle del Atuel, Quebrada del Arroyo Niehd, A. Ruiz Leal & F. Roig 15677 (LP). 16. Nassauvia glomerulosa (Lag. ex Lindl.) D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. Basionym: Triptilion glomerulosum Lag. ex Lindl. TYPE: Argentina. Cordillera del Portillo, 1813, M. Nee s.n. (isotype, G not seen, G photo F n. 8235 at LP!). Additional specimen examined: Argentina. Prov. Chubut: Tecka, Estancia La Blanche, A. Soriano 2216 (LP). 17. Nassauvia hillii Cabrera, Notas Mus. La Plata, Bot. 22: 161. 1939. TYPE: Argentina. Prov. Neuquén: Dept. Aluminé, Pulmarí, cerca de Piedra Gaucha, H. F. Comber 356 (isotype, LP!). 18. Nassauvia juniperina Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 328. 1916. TYPE: not localized. Additional specimen examined: Argentina. Prov. Chubut: a 35 km de Esquel, A. Ruiz Leal 23964 (LP). 19. Nassauvia lagascae (D. Don) F. Meigen, Bot. Jahrb. Syst. 18: 445. 1894. Basionym: Sphaerocephalus lagascae D. Don. TYPE: Chile. Andes of Chile, ascent to Los Peuquenes, J. Gillies 33 (holotype, E not seen). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Distrito Sosneao, Alto Valle del Atuel, Laguna Atuel, A. Ruiz Leal 16882 (LP). 20. Nassauvia latissima Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 330. 1916. TYPE: Argentina. Prov. Tierra del Fuego: Isla de los Estados, C. Skottsberg 273 (isotype, SI not seen, SI photo at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Dept. Bahía Tetis, Estancia Moat, D. M. Moore 1649 (LP). 21. Nassauvia looseri Cabrera, Darwiniana 24: 363. 1982, nom. novum. Basionym: Panargyrum heterophyllum Phil. TYPE: Chile. Cordillera de Santiago, Las Arañas, Jan 1860, G. Landbeck s.n. (isotype, LP!). Additional specimen examined: Chile. Prov. Santiago: Refugio de La Parva, a 4 km de Farellones, Cordillera de Las Condes, G. Looser 5755 (LP). 22. Nassauvia maeviae Cabrera, Darwiniana 24: 367. 1982. TYPE: Argentina. Prov. Santa Cruz: Dept. Güer Aike, laguna La Leona, 51°31″ S, 69°47″ W, 7 Dec 1975, M. N. Correa s.n. (holotype, SI!). Additional specimen examined: Argentina. Prov. Santa Cruz: Dept. Güer Aike, a 14 km desvío de ruta nacional 3 hacia la Estancia Guakenen Aike, L. Katinas 24 (LP). 23. Nassauvia magellanica J. F. Gmel., Syst. nat.: 1281. 1791. TYPE: Chile or Argentina. Magallania, s.d., D. Commerson s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Usuahia, Laguna Azul, M. N. Correa & R. L. Pérez Moreau 1920 (LP).


637

24. Nassauvia pentacaenoides Speg., Anales Soc. Ci. Argent. 53: 34. 1902. TYPE: Argentina. Prov. Santa Cruz: Karr-aik, Lago Argentino, March 1898, C. Ameghino s.n. (holotype, LP!). 25. Nassauvia pinnigera D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, ascent to Planchón, J. Gillies 36 (holotype, E not seen). Additional specimen examined: Argentina. Prov. Neuquén: Cordillera del Viento, La Corona, F. Pastore 83 (LP). 26. Nassauvia planifolia Wedd., Chlor. andina 1: 48. 1855. TYPE: Argentina. E side of Andes, Bridges 1148 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Valle Hermoso, subida hacia la Vega de los Piuquenes, A. Ruiz Leal 27357 (LP). 27. Nassauvia pulcherrima Cabrera, Notas Mus. La Plata, Bot. 22: 157. 1939. TYPE: Argentina. Prov. Río Negro: Dept. Bariloche, Cerro Catedral, s.d., N. J. J. Neumeyer s.n. (holotype, LP!). 28. Nassauvia pygmaea (Cass.) Hook. f., Fl. antarct. 2: 319. 1847. Basionym: Triachne pygmaea Cass. TYPE: Argentina. Patagonia australis, Cerro Toro, 1826? 1896?, O. Nordenskjold s.n. (isotype, K not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Isla de los Estados, Puerto Cook, M. A. Torres 1199 (LP). 29. Nassauvia pyramidalis Meyen, Reise Chile 1: 356. 1834. TYPE: Chile. Cordillera de Santiago, Valle del Yeso, Jan 1868, R. Philippi s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Minas, Cordillera del Viento, cruzando de Tricao Malal al Cajón de Butaló, O. Boelcke 11598 (LP). 30. Nassauvia ramosissima DC., Prodr. 7: 49. 1838. TYPE: Chile. Prov. Colchagua: Talcaregue, 1831, C. Gay s.n. (holotype, G not seen, G photo at LP!). 31. Nassauvia revoluta D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, ascent to Planchón, J. Gillies 35 (holotype, E not seen, E photo at LP!). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Ñorquín, Termas de Copahue, A. L. Cabrera 6282 (LP). 32. Nassauvia ruizii Cabrera, Notas Mus. La Plata, Bot. 22: 162. 1939. TYPE: Argentina. Prov. Mendoza: Dept. Tunuyán, Rincones del Cerro Morado, A. Ruiz Leal 3183 (holotype, LP!). 33. Nassauvia sceptrum Dusén, Rep. Princeton Univ. Exp. Patagonia, Botany, Suppl.: 286. 1914. TYPE: Argentina. Prov. Santa Cruz: Lago Viedma, P. Dusén 5887 (isotype, SI not seen, SI photo at LP!). 34. Nassauvia serpens d’Urv., Mém. Soc. Linn. Paris 4: 610. 1825. TYPE: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), 1825, d’Urville s.n. (holotype, P not seen; isotype, B not seen, B photo F n. 16039 at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), East Falkland, Mt. Urbourne, D. M. Moore 583 (LP). 35. Nassauvia sprengelioides DC., Prodr. 7: 49. 1838. TYPE: Argentina. Prov. Mendoza: de la Cordillera del Planchón, s.d., M. Neé s.n. (holotype, G not seen). Additional specimen examined: Chile. Prov. Curicó: Teno, La Montaña, E. Barros 3801 (LP).

638

L. Katinas, et al.

36. Nassauvia sublobata Cabrera, Darwiniana 24: 323. 1982. TYPE: Argentina. Prov. Neuquén: cordón del Chapelco, nacientes del arroyo Chapelco Grande, al N del pequeño lago La Kika, M. Gentili & P. Gentili 754 (isotype, LP!). Additional specimen examined: Argentina. Prov. Mendoza: Alto Valle de Calmuco, A. Burkart et al. 14401 (LP). 37. Nassauvia ulicina (Hook. f.) Macloskie, Rep. Princeton Univ. Exp. Patagonia, Botany 8: 885. 1906. Basionym: Trianthus ulicinus Hook. f. TYPE: Argentina. Prov. Santa Cruz: Cape Fairweather, s.d., Capt. King s.n. (holotype, K not seen). Additional specimen examined: Argentina. Prov. Chubut: Dept. Rawson, Rawson, Punta Norte, M. M. Job 3009 (LP). 38. Nassauvia uniflora (D. Don) Hauman, Anales Soc. Ci. Argent. 86: 327. 1918. Basionym: Panargyrum uniflorum D. Don. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, Paramillo de las Cuevas, s.d., J. Gillies s.n. (holotype, E not seen). Additional specimen examined: Argentina. Prov. Mendoza: Las Cuevas, Quebrada Benjamín Matienzo, R. Pérez Moreau 12753 (LP).

Observation. Species of Nassauvia are dominant elements in vast areas of Patagonia. Literature. Cabrera (1982), Kalin Arroyo and Marticorena (1988), Freire et al. (1993), Katinas at al. (2008a).

Oxyphyllum Phil. Fl. atacam.: 28. 1860. TYPE: Oxyphyllum ulicinum Phil. 63 of Figs. 63–66. Etymology. From the Greek oxys, sharp, pungent, and phyllon, leaf, referring to the spiny, sharp-pointed leaves. Shrubs. Leaves imbricate; sessile, clasping; blades linear to obovate, entire, pinnatisect to reduced to three-parted spines, pinnately veined, margin entire, involute, pubescent. Capitulescences densely corymbose; capitula short-pedunculate, homogamous, discoid; receptacle paleate, paleae embracing the marginal florets; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate to slightly constricted at the apex, pubescent; pappus with scabrid to plumose, capillary bristles. Pollen Description. Pollen spheroidal, with polar caps in equatorial view, circular in polar view, medium size, P × E = (28–33×26–30) μm. Tricolporate, colpi long with scarcely microgranulate membrane. Exine Oxyphyllum type, scabrate, 3–4 μm thick, thickened at the poles. Nexine 0.5–1 μm thick (see also Parra & Marticorena, 1972; Crisci, 1974a; Tellería et al., 2003). Monotypic genus endemic to the desert areas of northern Chile.


639

Figs. 63–66 63 Oxyphyllum ulicinum Phil. A Habit (branch). B Palea. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (From Zöllner 3892, LP). Scale bars: A=4 cm; B, C, F=2 mm; D, E= 1 mm. 64 Panphalea missionum Cabrera. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela. (A–C redrawn from Cabrera, 1953b: 234; D, E from Spegazzini s.n., LP). Scale bars: A=4 cm; B, E=2 mm; C=0.5 mm; D=0.3 mm. 65 Perezia multiflora (Humb. & Bonpl.) Less. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Cabrera, 1978: 657). Scale bars: A=5 cm; B=4 mm; C=2.5 mm; D=1.5 mm; E=3 mm. 66 Pleocarphus revolutus D. Don. A Habit (branch). B Palea. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (Redrawn from Muñoz Pizarro, 1966, Figure 73, without page number). Scale bars: A=4 cm; B=2.2 mm; C, D=3 mm; E =1.8 mm; F=4.5 mm

Species list 1. Oxyphyllum ulicinum Phil., Fl. atacam.: 173. 1860. TYPE: Chile. Prov. Antofagasta: región del litoral, en Cachinal de la Costa, Paposo, etc., R. Philippi s.n. (type, probably SGO). Additional specimen examined: Hueso Parado, cerca de Taltal, O. Zöllner 3892 (LP).

640

L. Katinas, et al.

Literature. Reiche (1905).

Panphalea Lag. Amen. nat. Españ.: 34. 1811. TYPE: Panphalea commersonii Cass. 64 of Figs. 63–66. Ceratolepis Cass., Bull. Sci. Soc. Philom. Paris: 111. 1819, nom. nudum. Pamphalea DC. Ann. Mus. Natl. Hist. Nat. 19: 68. 1812, an orthograhic variant of the name Panphalea (see Observation). Etymology. From Greek pan, all, whole, and phalos, shiny, bright, probably in reference to the white florets or to the leaves. Herbs annual or perennial, caulescent. Leaves glabrous to pubescent; basal leaves rosulate; petiolate, sessile to clasping; blades linear, lanceolate, ovate to orbicular, palmately to pinnately veined, margin entire, lyrate to pinnatisect; upper leaves alternate, reduced; clasping. Capitulescences racemose to corymbose; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre one- to threeseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, pubescent; pappus absent. Pollen Description. Exine Oxyphyllum type (see Crisci 1974a). Habitat and Distribution. Genus of nine South American species distributed in humid places of southern Brazil, eastern Paraguay, Uruguay, and northeastern Argentina. Species list 1. Panphalea araucariophila Cabrera, Notas Mus. La Plata, Bot. 16: 232. 1953. TYPE: Brazil. State Rio Grande do Sul: Taimbesinho, p. São Francisco de Paula, B. Rambo 49335 (holotype, LP!). 2. Panphalea bupleurifolia Less., Linnaea 5: 8. 1830. TYPE: Brazil. Brasilia meridionalis, s.d., Sello s.n. (holotype, B not seen, B photo F n. 16094 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: arroyo Juan Blanco-Ruta 11, J. V. Crisci 542 (LP). 3. Panphalea cardaminifolia Less., Linnaea 5: 9. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1553 (holotype B, not seen, B photo F n. 16095 at LP!). Additional specimen examined: Brazil. State Santa Catarina: Mun. Lajes, 6 km W of Capão Alto, S. B. Smith & R. M. Klein 11341 (LP). 4. Panphalea commersonii Cass., Bull. Sci. Soc. Philom. Paris: 111. 1819. TYPE: Uruguay. Près de Montevideo, s.d., s. leg., herb. Jussieu s.n. (holotype, P not seen). Additional specimen examined: Uruguay. Dept. Montevideo: Montevideo, alrededores del cerro de Montevideo, J. Chebataroff 232 (LP). 5. Panphalea heterophylla Less., Linnaea 5: 8. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1328 (holotype, B not seen, B


6.

7.

8.

9.

641

photo F n. 16096 at LP!). Additional specimen examined: Argentina. Prov. Entre Ríos: Dept. Concordia, Salto Grande, A. L. Cabrera & A. Sagástegui 19295 (LP). Panphalea maxima Less., Linnaea 5: 9. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1365 (holotype, B not seen, B photo F n. 16097 at LP!). Additional specimen examined: Uruguay. Dept. Salto: Paysandú, Cueva del Tigre, J. Chebataroff 7001 (LP). Panphalea missionum Cabrera, Notas Mus. La Plata, Bot. 16: 235. 1953. TYPE: Argentina. Prov. Misiones: Dept. Candelaria, Loreto, J. E. Montes 2515 (holotype, LP!). Additional specimen examined: Argentina. Prov. Misiones: Campo de las Cuyas, Jan 1907, C. L. Spegazzini s.n. (LP). Panphalea ramboi Cabrera, Notas Mus. La Plata, Bot. 16: 229. 1953. TYPE: Brazil. State Rio Grande do Sul: Taimbesinho, p. São Francisco de Paula, B. Rambo SJ 49420 (holotype, LP!). Panphalea smithii Cabrera, Bol. Soc. Argent. Bot. 7: 200. 1959. TYPE: Brazil. Santa Catarina: Mun. Bom Retiro, bog, Fazenda Campo dos Padres, Campo dos Padres, S. B. Smith & P. Reitz 10417 (holotype, LP!).

Observation. It can be found in the literature the use of the name Pamphalea instead of the original name Panphalea. The basic problem is that the final letter of “pan” is a dental. In Greek, when a dental (the final “n” of “pan”) comes up against a labial, it changes to a labial (“m”), hence “pam.” The International Code of Botanical Nomenclature (Greuter et al., 2000) in article 60.1 suggests maintaining the original spelling, not imposing linguistically correct spellings. Therefore the original spelling, Panphalea, must be maintained. Those who want to use “pam” must propose conservation, documenting overwhelming usage of “pam” (Dan Nicolson, Smithsonian Institution, personal communication). Literature. Cabrera (1953b, 1959a), Katinas and Crisci (2008).

Perezia Lag. Amen. nat. Españ.: 31. 1811. TYPE: Perdicium magellanicum L. f. [= Perezia magellanica (L. f.) Lag.]. 65 of Figs. 63–66. Calorezia Panero, Phytologia 89: 199. 2007. TYPE: Calorezia nutans (Less.) Panero [= Perezia nutans Less.]. Clarionea Lag. ex DC., non Cass., Ann. Mus. Natl. Hist. Nat. 19: 65. 1812. TYPE: Clarionea magellanica Lag. ex DC. [= Perezia magellanica (L. f) Lag.]. Clarionia D. Don, Trans. Linn. Soc. London 16: 204. 1830, an orthographic variant of the name Clarionea. Clarionema Phil., Linnaea 28: 717. 1858. TYPE: Clarionema humilis Phil. [= Perezia pedicularidifolia Less.]. Drozia Cass., Dict. Sci. Nat. 34: 217. 1825. TYPE: Drozia dicephala Cass. [= Perezia pungens (Humb. & Bonpl.) Less.]. Dumerilia Less., Linnaea 5: 13. 1830, non Lag. ex DC., 1812, nom. illeg.

642

L. Katinas, et al.

Heteranthus Cass., Dict. sci. nat. 21: 110. 1821, an admitted orthographic variant of the name Homoianthus (Simpson Vuilleumier, 1969). Homanthis Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 10. 1818, probably an orthographic variant of the name Homoianthus. Homoeanthus Spreng., Syst. veg. 3: 503. 1826, an orthographic variant of the name Homoianthus. Homoianthus DC., Ann. Mus. Natl. Hist. Nat. 19: 65. 1812. TYPE: Homoianthus pungens (Humb. & Bonpl.) DC., based on Chaetanthera pungens Humb. & Bonpl. [= Perezia pungens (Humb. & Bonpl.) Less.]. Isanthus DC., Prodr. 7: 63. 1838, nom. nudum. Pogonura DC. ex Lindl., Intr. nat. syst. bot.: 263. 1836, nom. nudum. Scolymanthus DC., Prodr. 7: 63. 1838, nom. nudum. Stenophyllum Less., Syn. gen. Compos.: 412. 1832, a name frequently listed as a synonym of Perezia, but it was actually used only as a sectional name by Lessing (Simpson Vuilleumier, 1969). Etymology. In honor of Lázaro (or Laurentio) Pérez, a former pharmacist from Toledo, Spain. Herbs perennial, caulescent or acaulescent. Leaves pinnately veined or parallel veined when sheathing, glabrous to densely pubescent; basal leaves rosulate; pseudopetiolate to clasping; blades linear, oblong to ovate, margin entire to lyrate or pinnately partite; upper leaves alternate, numerous to absent, reduced to bracts to broadly oblong or ovate, margin entire, ciliate to lacerate; clasping. Capitulescences monocephalous, corymbose to paniculate; capitula pedunculate to subsessile, homogamous, discoid or radiate; receptacle epaleate; involucre multiseriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to villose; pappus of scabrid, capillary bristles. Pollen Description. Perezia (114 of Figs. 112–116) (Parra & Marticorena, 1972). Pollen subprolate to spheroidal-oblate, medium size, tricolporate, exine Oxyphyllum type (see also Simpson Vuilleumier, 1969; Crisci, 1974a; Crisci & Marticorena, 1978; Markgraf & D’Antoni, 1978; Wingenroth & Heusser, 1983). Habitat and Distribution. Genus of 32 South American species mainly from the Andes and Patagonia, from Colombia to the Islas Malvinas (Falkland Islands), with a few species in eastern Argentina, Brazil, Paraguay, and Uruguay. Species list 1. Perezia bellidifolia (Phil.) Reiche, Anales Univ. Chile 116: 437. 1905. Basionym: Homoeanthus bellidifolius Phil. TYPE: Chile. Prov. Valdivia: Cordillera de Valdivia, Huahim, O. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Lácar, Cerro Chapelco, A. L. Cabrera et al. 22987 (LP).


643

2. Perezia calophylla (Phil.) Reiche, Anales Univ. Chile 116: 435. 1905. Basionym: Homoeanthus calophyllus Phil. TYPE: Chile. Prov. Valdivia: Pucaullu, O. Philippi 2254 (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquén: San Martín de los Andes, Bridarolli 2229 (LP). 3. Perezia carduncelloides Griseb., Pl. lorentz.: 150. 1874. TYPE: Argentina. Prov. Tucumán: Ciénaga, P. G. Lorentz 320 (holotype, B not seen, B photo F n. 16065 at LP!; isotype, CORD not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Volcán, subida al Cerro Horqueta, A. L. Cabrera et al. 16994 (LP). 4. Perezia carthamoides (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 34. 1835. Basionym: Clarionea carthamoides D. Don. TYPE: Argentina. Prov. Mendoza: Alto de La Laguna, 1821, J. Gillies s.n. (holotype, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Cordillera del Tigre, King 330 (LP). 5. Perezia ciliaris D. Don ex Hook. & Arn., Comp. Bot. Mag. 1: 34. 1835. TYPE: Chile: s.d., J. Dombey s.n. (lectotypes designated by Simpson Vuilleumier, 1969: 95, P, G, not seen, P photo at LP!, G photo F n. 8278 at LP!). Additional specimen examined: Argentina. Prov. Tucumán: Quebradas del Barón, H. A. Fabris 1382 (LP). 6. Perezia ciliosa (Phil.) Reiche, Anales Univ. Chile 116: 426. 1905. Basionym: Clarionea ciliosa Phil. TYPE: Chile. Tarapacá, Cerro de Copacoya, F. Philippi 2250 (isotype, LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Susques, Alto Chorrillos, A. L. Cabrera & H. Schwabe 124 (LP). 7. Perezia coerulescens Wedd., Chlor. andina 1: 39. 1855. TYPE: Peru. Cuzco, Oct 1839-Feb 1840, C. Gay s.n. (holotype, P not seen). Additional specimen examined: Peru. Dept. Puno: Prov. Melgao, Hacienda Huaycho, E. Antazana 51 (LP). 8. Perezia delicata Vuilleum., Contr. Gray Herb. 199: 128. 1969. TYPE: Argentina. Prov. Río Negro: región del Lago Nahuel Huapí, Paso de las Nubes, A. L. Cabrera 5917 (holotype, LP!). Additional specimen examined: Argentina. Prov. Río Negro: Cerro López, H. A. Fabris 2161 (LP). 9. Perezia eryngioides (Cabrera) Crisci & Martic., J. Arnold Arbor. 59: 352. 1978. Basionym: Trixis eryngioides Cabrera. TYPE: Brazil. State Santa Catarina: Mun. Irani, campo de Irani, S. B. Smith & R. M. Klein 7777 (holotype, LP!*). 10. Perezia fonkii (Phil.) Reiche, Anales Univ. Chile 116: 434. 1905. Basionym: Clarionea fonkii Phil. TYPE: Chile. Cordillera del Doce, Fonk 59 (isotype, LP!). Additional specimen examined: Argentina. Prov. Río Negro: Dept. Bariloche, Cerro López, O. Boelcke 1963 (LP). 11. Perezia kingii Baker, Fl. bras. 6: 380. 1884. TYPE: Uruguay. Concepción, Puerto de Salamanca, Oct 1875, P. G. Lorentz s.n. (holotype, B not seen, B photo F n. 16074 at LP!; isotype, GH not seen). Additional specimen examined: Uruguay. Dept. Florida: San Pedro del Timote, arroyo Timote, A. Gallinal et al. PE-5–334 (LP). 12. Perezia lactucoides (Vahl) Less., Linnaea 5: 22. 1830. Basionym: Perdicium lactucoides Vahl. TYPE: Chile. Prov. Magallanes: s.d., D. Commerson s.n. (isotype, C not seen, C photo F n. 22606 at LP!). Additional specimen examined:

644

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

L. Katinas, et al.

Argentina. Prov. Tierra del Fuego: Ushuaia, N. Correa & R. Pérez Moreau 1853 (LP). Perezia linearis Less., Syn. gen. Compos.: 412. 1832. TYPE: Chile. Prov. Bío Bío: near Antuco, E. F. Poeppig 767 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Río Negro: Dept. Bariloche, Cerro López, H. A. Fabris 2179 (LP). Perezia lyrata (J. Rémy) Wedd., Chlor. andina 1: 42. 1855. Basionym: Homoianthus lyratus J. Rémy. TYPE: Chile. Prov. Colchagua: Talcaregue, C. Gay 283 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Talca: Laguna del Maule, O. Boelcke et al. 10430 (LP). Perezia magellanica (L. f.) Lag., Amen. nat. Españ.: 31. 1811. Basionym: Perdicium magellanicum L. f. TYPE: Argentina. Prov. Tierra del Fuego, s.d., Forster s.n. (holotype, LINN 1003.5 not seen). Additional specimen examined: Argentina. Prov. Santa Cruz, Dept. Lago Argentino, Cerro Fitz Roy, P. Agostini B-30 (LP). Perezia mandonii Rusby, Mem. Torrey Bot. Club. 3: 66. 1893. TYPE: Bolivia. Oruro, Capí, M. Bang 777 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Volcán, subida al cerro Horqueta, A. L. Cabrera et al. 16996 (LP). Perezia megalantha Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 540. 1897. TYPE: Argentina. Prov. Santa Cruz: Lago Argentino, 1890, Tonini del Furia s.n. (holotype, LP!). Perezia multiflora (Humb. & Bonpl.) Less., Linnaea 5: 19. 1830. Basionym: Chaetanthera multiflora Humb. & Bonpl. TYPE: Ecuador. Prov. Pichincha: Volcán, Antisana, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Aguada Cerro Abra Morada, A. L. Cabrera et al. 16997 (LP). Perezia nutans Less., Syn. gen. Compos.: 409. 1832. TYPE: Chile. Antuco, meseta de Antuco, E. F. Poeppig 896 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Minas, cajón del arroyo Covunco, O. Boelcke et al. 11460 (LP). Perezia pedicularidifolia Less., Syn. gen. Compos.: 410. 1832. TYPE: Chile. Prov. Bío Bío: Andes de Antuco, Pico de Pilque, s.d., E. F. Poeppig s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Río Negro: Dept. Bariloche, laguna Frías, Cerro Riggi, A. L. Cabrera 6052 (LP). Perezia pilifera (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 34. 1835. Basionym: Clarionea pilifera D. Don. TYPE: Argentina. Prov. Mendoza: Andes de Mendoza, Cerro de Pulcura, s.d., J. Gillies s.n. (holotype, K not seen). Additional specimen examined: Chile. Prov. Ñuble: Termas de Chillán, Pirigallo, A. L. Cabrera 3642 (LP). Perezia pinnatifida (Humb. & Bonpl.) Wedd., Chlor. andina 1: 40. 1855. Basionym: Chaetanthera pinnatifida Humb. & Bonpl. TYPE: Lectotype, designated by Simpson Vuilleumier, 1969: 140, the illustration in Pl. Aequin. 2: 170, pl. 136. 1809. Additional specimen examined: Bolivia. La Paz: Larecaja, vicinity of Sorata, between Pongo and Amlaya, Mandon 20 (US, not seen).


645

23. Perezia poeppigii Less., Syn. gen. Compos.: 411. 1832. TYPE: Chile. Boreal Andes, s.d., E. F. Poeppig s.n. (holotype, P not seen). Additional specimen examined: Chile. Prov. Santiago: Alhué, Monte Cantillana, E. Barros 2025 (LP). 24. Perezia prenanthoides Less., Syn. gen. Compos.: 409. 1832. TYPE: Chile. Prov. Bío Bío: Andes de Antuco, Valle de Quilai, E. F. Poeppig 923 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Río Negro: Cerro López, A. Corte 226 (LP). 25. Perezia pungens (Humb. & Bonpl.) Less., Linnaea 5: 20. 1830. Basionym: Chaetanthera pungens Humb. & Bonpl. TYPE: Ecuador. Prov. Pichincha: Mt. Pichincha near Quito, s.d., F. Humboldt & A. Bonpland s.n. (holotype, P not seen; isotype, B not seen, B photo F n. 16084 at LP!). Additional specimen examined: Peru. Prov. Cajamarca: Dept. Cajamarca, Challuayaco, entre Kumulca y Celendín, I. Sánchez V. & W. Ruiz V. 468 (LP). 26. Perezia purpurata Wedd., Chlor. andina 1: 43. 1855. TYPE: Bolivia. Potosí, d’Orbigny 1420 (holotype, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Salta: Sierra del Cajón, D. Rodríguez 1376 (LP). 27. Perezia pygmaea Wedd., Chlor. andina 1: 40. 1855. TYPE: Bolivia. Prov. La Paz: ravine of Chuquiaguillo, near La Paz, 1851, H. A. Weddell s.n. (holotype, P not seen, P photo F n. 38128 at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, Mina Aguilar, A. L. Cabrera et al. 19789 (LP). 28. Perezia recurvata (Vahl) Less., Linnaea 5: 21. 1830. Basionym: Perdicium recurvatum Vahl. TYPE: Chile. Prov. Magallanes: strait of Magallan, s.d., D. Commerson s.n. (holotype, C not seen, C photo F n. 22607 at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Ushuaia, Río Grande, N. Alboff 374 (LP). 29. Perezia spathulata (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 33. 1835. Basionym: Clarionea spathulata D. Don (see Observations). TYPE: Chile: s. d., H. Ruiz & J. A. Pavon s.n. (type, probably MA). Additional specimen examined: Argentina. Prov. Neuquén: Dept. Lácar, Hua-Hum, camino al Lago Queñi, J. Diem 3252 (LP). 30. Perezia squarrosa (Vahl) Less., Linnaea 5: 15. 1830. Basionym: Perdicium squarrosum Vahl. TYPE: Uruguay. Dept. Canelones: sommet du Morne de Montevideo, 1767, D. Commerson s.n. (holotype C, not seen). Additional specimen examined: Paraguay. Paraguarí: Ybytimi, Montes 12951 (LP). 31. Perezia sublyrata Domke, Notizbl. Bot. Gart. Berlin-Dahlem 13: 248. 1936. TYPE: Peru. Puno, Santa Rosa, Stafford 345 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tilcara, subida al Abra de Remate, H. Sleumer 4075 (LIL). 32. Perezia volcanensis Cabrera, Fl. Prov. Jujuy 10: 660. 1978. TYPE: Argentina. Prov. Jujuy: Dept. Tumbaya, Estación Volcán, L. Castillón 8155 (holotype, LIL, photo at LP!: see Observations).

Observations. Perezia spathulata is cited as a synonym of P. viscosa Less. in the revision of Perezia (Simpson Vuilleumier, 1969). However, as mentioned by Cabrera (1971b), P. spathulata is the correct name by priority.

646

L. Katinas, et al.

The citation of the type specimen of Perezia volcanensis as L. Castillón 7020 in Flora of Jujuy (Cabrera, 1978) is a typographic error (Katinas, 1995; Freire & Iharlegui, 2000). The genus Calorezia, containing two species of Perezia, was recently proposed by Panero (2007) based on a chloroplast DNA coding regions study in preparation by Panero et al. Since there are not clear morphological characters that support this new genus, it is included in the synonymy of Perezia. Literature. Simpson (1973), Simpson Vuilleumier (1969), Crisci and Marticorena (1978), Katinas (1995).

Pleocarphus D. Don Trans. Linn. Soc. London 16: 228. 1830. TYPE: Pleocarphus revolutus D. Don. 66 of Figs. 63–66. Etymology. From the Greek pleos, plenty, full, filled, and karphos, any small dry body, in reference to the receptacle completely paleate. Shrubs, stems with brachyblasts and macroblasts. Leaves alternate; sessile; blades linear to linear-lanceolate, pinnately veined, margin sinuate to dentate, revolute, glabrescent. Capitulescences densely paniculate; capitula sessile to shortly pedunculate, homogamous, discoid; receptacle paleate, paleae embracing all the florets; involucre uniseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of scabrid, capillary bristles. Pollen Description (Parra & Marticorena, 1972). Pollen prolate–subprolate, medium size, tricolporate, exine Trixis type (see also Crisci, 1974a). Habitat and Distribution. Monotypic genus endemic to rocky soils of central and northern Chile. Species list 1. Pleocarphus revolutus D. Don, Trans. Linn. Soc. London 16: 229. 1830. TYPE: Chile. Prov. Coquimbo: s.d., Caldcleugh s.n. (holotype, G not seen, G photo at LP!, US!). Additional specimen examined: Chile. Prov. Coquimbo: alrededores de La Serena, Quebrada Romeral, A. L. Cabrera 11407 (LP). Literature. Gay (1847).

Polyachyrus Lag. Amen. nat. Españ.: 37. 1811. TYPE: Cephaloseris poeppigii Kuntze ex Less. [= Polyachyrus poeppigii (Kunze ex Less.) Less.] 67 of Figs. 67–70.


647

Figs. 67–70 67 Polyachyrus carduoides Phil. A Habit (branch). B Capitulum of two florets. C Floret (without cypsela). D Cypsela of inner floret, with pappus. E Stamen. F Upper part of the style. G Cypsela of outer floret, with pappus. (Redrawn from Muñoz Pizarro, 1966, Figure 67 without page number). Scale bars: A=5 cm; B=4 mm; C, E=2 mm; D, G=2.5 mm; F=1.3 mm. 68 Proustia cuneifolia D. Don. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Cabrera 1978: 614). Scale bars: A=2 cm; B, E=6.5 mm; C=4 mm; D=2 mm. 69 Triptilion achilleae DC. A Habit. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. F Lateral view of palea of the pappus. (Redrawn from Katinas et al., 1992: 102). Scale bars: A=2 cm; B=2 mm; C, D=1 mm; E, F= 3 mm. 70 Trixis grisebachii Kuntze. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Katinas, 1996b: 65). Scale bars: A=5 cm; B=10 mm; C=2 mm; D= 1.5 mm; E=4 mm

Bridgesia Hook., Bot. misc. 2: 228. 1831. TYPE: Bridgesia echinopsoides Hook. [= Polyachyrus sphaerocephalus D. Don]. Cephaloseris Poepp. ex Rchb., Consp. regn. veg. 1: 100. 1828. TYPE: Cephaloseris poeppigii Kunze ex Less., Linnaea 5: 5. 1830 [= Polyachyrus poeppigii Kunze ex Less. var. poeppigii]. Diaphoranthus Meyen, Reise Chile 1: 404. 1834. TYPE: Diaphoranthus fuscus Meyen [= Polyachyrus fuscus (Meyen) Walp.]. Polyachrus Hook. & Arn., Comp. Bot. Mag. 1: 36. 1835, an orthographic variant of the name Polyachyrus. Polyachurus DC., Ann. Mus. Natl. Hist. Nat. 19: 69. 1812, an orthographic variant of the name Polyachyrus. Polyachyris Walp., Repert. Bot. Syst. (Walpers) 2: 681. 1843, an orthographic variant of the name Polyachyrus.

648

L. Katinas, et al.

Etymology. From Greek polys, many, and achyron, chaff, husks, referring to the number of pappus bristles. Subshrubs erect or decumbent. Leaves pinnately veined, pubescent; basal leaves rosulate to sub-rosulate; decurrent to amplexicaul; blades linear to oblong, pinnately lobed to pinnately partite, auriculate at the base, margin spiny to unarmed; upper leaves alternate, similar to the basal leaves but reduced. Capitulescences a corymbose pseudocephalium; capitula shortly-pedunculate, homogamous, discoid, with two florets, rarely three, surrounded by an involucre of five phyllaries: four enclosing one floret and one enclosing the other floret or all florets, the phyllary between florets interpreted as a palea; receptacle paleate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs Cypselae truncate to constricted at the apex, glabrescent to pubescent; pappus dimorphic, scabrid to plumose bristles, shorter in the inner floret, capillary to somewhat paleaceous. Pollen Description. Polyachyrus (Parra & Marticorena, 1972). Pollen subprolatespheroidal, medium size, tricolporate, exine Oxyphyllum type (see also Tellería et al., 2003). Habitat and Distribution. Genus of seven species endemic to the arid regions of northern and central Chile and Peru. Species list 1. Polyachyrus annuus I. M. Johnst., Contr. Gray Herb. 8: 134. 1929. TYPE: Chile. Prov. Antofagasta: Dept. Tocopilla, Tocopilla, ca. 6 km N of port and about opposite Caleta Duendes, I. M. Johnston 3538 (isotype, CONC!). Additional specimen examined: Peru. Dept. Arequipa: Prov. Camaná, R. Ferreyra 2518 (LP). 2. Polyachyrus carduoides Phil., Fl. atacam.: 202. 1860. TYPE: Chile. Prov. Antofagasta: Dept. Taltal, Valle Sandón, s.d., R. A. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Atacama: Encantada, s.d., G. E. Gulig s.n. (LP). 3. Polyachyrus cinereus Ricardi & Weldt, Gayana, Bot. 26: 26. 1974. TYPE: Chile. Prov. Antofagasta: region about Aguada Cachina, waterhole in Quebrada Cachina, ca. 6 km in land from Caleta Esmeralda, I. M. Jhonston 5726 (holotype, GH not seen). Additional specimen examined: Chile. Prov. Copiapó: Pan de Azúcar, S. Roig 19 (LP). 4. Polyachyrus fuscus (Meyen) Walp., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 1: 288. 1843. Basionym: Diaphoranthus fuscus Meyen. TYPE: Chile. Cordillera de Copiapó, 1833, Meyen s.n. (holotype, B not seen, B photo F n. 15992 at LP!). Additional specimen examined: Chile. Prov. Antofagasta: la Chimba, E. Barros 2083 (LP). 5. Polyachyrus gayi J. Rémy, Fl. chil. 3: 372. 1847. TYPE: Chile. Costas de las provincias septentrionales, s.d., C. Gay s.n. (holotype, P not seen, P photo F n. 15993 at LP!; isotype, SGO not seen). Additional specimen examined: Chile. Prov. Antofagasta: la Chimba, E. Barros 2079 (LP).


649

6. Polyachyrus poeppigii (Kunze ex Less.) Less., Linnaea 5: 5. 1830. Basionym: Cephaloseris poeppigii Kunze ex Less. TYPE: Chile. Prov. Valparaíso: Concón, E. F. Poeppig 221 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Valparaíso: Quebrada en ruta 5 y Puente Juan Soldado, M. Bonifacino 1366 (LP). 7. Polyachyrus sphaerocephalus D. Don, Trans. Linn. Soc. London 16: 229. 1830. TYPE: Chile. Prov. Tarapacá: Dept. Iquique, Usmagama, Feb 1885, F. Philippi s.n. (holotype, B not seen, B photo F n. 15996 at LP!). Additional specimen examined: Peru. Dept. Lima: Infiernillo, A. L. Cabrera 10891 (LP).

Literature. Ricardi and Weldt (1974), Katinas and Crisci (2000), Katinas et al. (2008a).

Proustia Lag. Amen. nat. Españ.: 33. 1811. TYPE: Proustia pyrifolia DC. 68 of Figs. 67–70. Etymology. In honor of Ludovico Proust, a chemist from Madrid, Spain, of the early nineteenth century. Shrubs or little trees, erect or scandent, some branches modified in thorns to unarmed, stems with brachyblasts and macroblasts. Leaves alternate; petiolate to subsessile; blades linear, oblong to ovate, pinnately veined, margin entire to spinosedentate, glabrous to pubescent. Capitulescences densely racemose to paniculate; capitula subsessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth to slightly papillose; style bifid, branches rounded at the apex, dorsally papillose. Cypselae truncate to slightly constricted at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles. Pollen Description. Proustia (Tellería et al., 2003). Pollen subprolate, medium size, tricolporate, exine Trixis type and both Proustia and Trixis type in a single specimen, with polar caps only in P. pyrifolia (see also Parra & Marticorena, 1972; Crisci, 1974a). Habitat and Distribution. Genus of three South American species distributed in the Andean regions of southern Peru, Bolivia, central Chile, and Argentina. Species list 1. Proustia cuneifolia D. Don, Trans. Linn. Soc. London 16: 202. 1830. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably G). Additional specimen examined: Argentina. Prov. Salta: Dept. Rosario de Lerma, Santa Rosa de Tastil, A. L. Cabrera 9008 (LP). 2. Proustia ilicifolia Hook. & Arn., Bot. Beechey Voy.: 28. 1840. TYPE: Chile. Coquimbo, s.d., s. leg. (type, probably K). Additional specimen examined:

650

L. Katinas, et al.

Chile. Prov. Atacama: Dept. Copiapó, La Puerta, M. Ricardi & C. Marticorena 3667 (CONC). 3. Proustia pyrifolia DC., Ann. Mus. Natl. Hist. Nat. 19: 70. 1812. TYPE: Hab. in Chili prope Talcahuano, s.d., Lagasca s.n. (type, not localized). Additional specimen examined: Chile. Prov. Valparaíso: Cerro de la Campana, April 1933, Grandjot s.n. (CONC). Observation. Proustia has sometimes been placed in the subtribe Mutisiinae (Cabrera, 1961; Fabris, 1968) because of its rounded, dorsally papillose style branches. The type of exine stratification in the pollen of Proustia (Crisci, 1974a; Tellería et al., 2003) clearly indicates the position of this genus within the subtribe Nassauviinae (= tribe Nassauvieae). Literature. Fabris (1968), Crisci (1974a).

Triptilion Ruiz & Pav. Fl. peruv. prodr.: 102. 1794. TYPE: Triptilion spinosum Ruiz & Pav. 69 of Figs. 67–70. Triptilium DC., Ann. Mus. Natl. Hist. Nat. 19: 59. 1812, an orthographic variant of the name Triptilion. Etymology. From the Greek treis, tria, three, and ptilon, feather, referring to the three scales of the pappus. Herbs annual or perennial, caulescent to acaulescent. Leaves pinnately veined, pubescent; basal leaves rosulate to spread; sessile; blades linear-lanceolate to orbicular, entire to pinnatisect, margin spiny to unarmed; upper leaves alternate, similar to the basal leaves but reduced. Capitulescences corymbose, paniculate to glomerulose or a pseudocephalium; capitula sessile to pedunculate, homogamous, discoid, fiveflowered; receptacle epaleate; involucre two- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus with three, rarely two or four, narrowly paleaceous bristles, longitudinally plicate, apically plumose to laciniate, rarely crown-like. Pollen Description. Triptilion (Parra & Marticorena, 1972). Pollen spheroidal to suboblate, small to medium size, tricolporate, exine Oxyphyllum type (see also Crisci, 1974a). Habitat and Distribution. Genus of seven species endemic to Chile and to the subantarctic forests of southwestern Argentina. Species list 1. Triptilion achilleae DC., Prodr. 7: 51. 1838. TYPE: Chile. Tagua-Tagua, C. Bertero 708 (holotype, P!). Additional specimen examined: Argentina.


2.

3.

4.

5.

6.

7.

651

Prov. Neuquén: Dept. Huiliches, Lago Huechulaufquen, A. L. Cabrera 11286 (LP). Triptilion benaventii J. Rémy, Fl. chil. 3: 357. 1848. TYPE: Chile. Concepción, C. Gay 252 (holotype, P!). Additional specimen examined: Chile. Malleco, camino de Icalma a Liucura, Marimenuco, en las inmediaciones de la confluencia del arroyo Chanchocó con el Bío Bío, Pfister 7382 (CONC). Triptilion berteroi Phil., Anales Univ. Chile 87: 91. 1894. TYPE: Chile. Rancagua, 1818, C. Bertero s.n. (holotype, SGO!). Additional specimen examined: Chile. Santiago, Colina, cerro en la vecindad de Los Baños, s.d., Behn s.n. (CONC). Triptilion capillatum (D. Don) Hook. & Arn., Comp. Bot. Mag. 2: 43. 1836. Basionym: Nassauvia capillata D. Don. TYPE: Chile. Baths of Collina, s.d., Macrae s.n. (phragment of holotype, K!). Additional specimen examined: Chile. Quillota, Cerro La Campana, O. Zöllner 7280 (LP). Triptilion cordifolium Lag. ex Lindl., Bot. Reg. 10: 853. 1824. TYPE: Lectotype, designated by Katinas et al., 1992: 112, the illustration in Bot. Reg. 10: 852. 1824. Additional specimen examined: Chile. Petorca: Zapallar, Kausel 4530 (LP). Triptilion gibbosum J. Rémy, Fl. chil. 3: 356. 1848. TYPE: Chile. Colchagua, s.d., C. Gay s.n. (type, P!). Additional specimen examined: Chile. Elqui, La Serena, E. Barros 2429 (LP). Triptilion spinosum Ruiz & Pav., Syst. veg. fl. peruv. chil. 1: 185. 1798. TYPE: Chile. Concepción, 1782, H. Ruiz & J. A. Pavón s.n. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Cerro Moreno, O. Zöllner 3367 (LP).

Literature. Katinas et al. (1992, 2008a).

Trixis P. Browne Civ. nat. hist. Jamaica: 312. 1756. TYPE: Trixis inula Crantz. 70 of Figs. 67–70. Bowmania Gardner, London J. Bot. 2: 9. 1843. TYPE: Bowmania verbascifolia Gardner [= Trixis nobilis (Vell.) Katinas]. Castra Vell., Fl. flumin.: 343. 1829 (1825). TYPE: Castra praestans Vell. [= Trixis praestans (Vell.) Cabrera]. Prionanthes Schrank, Pl. rar. hort. monac. 2: 51. 1820. TYPE: Prionanthes antimenorrhoea Schrank [= Trixis divaricata (Kunth) Spreng. var. divaricata]. Tenorea Colla, Hortus ripul.: 137. 1824, non Raf., 1814, Gasp., 1845, nec K. Koch, 1869, nom. illeg. TYPE: Tenorea berteri Colla [= Trixis inula Crantz]. Etymology. From the Greek trixos, trissos, three times, in reference to the tridentate outer lip of the corolla. Shrubs, subshrubs, erect to climbing, or herbs perennial, caulescent. Leaves alternate, rarely basal leaves rosulate; petiolate, sessile, decurrent to auriculate;

652

L. Katinas, et al.

blades linear, elliptic to ovate, pinnately veined, margin mucronulate to serrate, sometimes revolute, pubescent. Capitulescences corymbose, paniculate to spicate; capitula sessile to pedunculate, homogamous, discoid or radiate; receptacle epaleate; involucre one- to five-seriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth to papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex to longly rostrate, pubescent; pappus of scabrid, capillary bristles. Pollen Description. Trixis (115 of Figs. 112–116) (Parra & Marticorena, 1972). Pollen prolate, medium to large size, tricolporate, exine Trixis type (see also Crisci, 1974a). Habitat and Distribution. Neotropical genus of 38 species distributed from southern United States to central Chile and Argentina. Species list 1. Trixis aggregata Rusby, Mem. Torrey Bot. Club 6: 71. 1896. TYPE: Bolivia. Bolivian palteau, M. Bang 1150 (isotype, BM!). Additional specimen examined: Bolivia. Dept. Cochabamba, Prov. Cercado, Cerro San Pedro, J. Steinbach 9705 (MO). 2. Trixis alata D. Don, Trans. Linn. Soc. London 16: 192. 1830. TYPE: Mexico. State Morelos: Ayacapixtl (Yecapixtla), s.d., M. Sessé & Moçiño s.n. (holotype, G-DEL not seen). 3. Trixis angustifolia DC., Prodr. 7: 69. 1838. TYPE: Mexico. State San Luis Potosí: pied des monts de Catorce, J. L. Berlandier 1353 (isotype, NY not seen, NY photo at LP!). 4. Trixis cacalioides (Kunth) D. Don, Trans. Linn. Soc. London 16: 187. 1830. Basionym: Perdicium cacalioides Kunth. TYPE: Peru. Andes, s.d., A. Bonpland s.n. (holotype, P!). Additional specimen examined: Peru. Dept. Amazonas, Prov. Chachapoyas, Río Marañón at Balsas, Hutchinson & Wright 5407 (MO). 5. Trixis calcicola B. L. Rob., Proc. Amer. Acad. Arts 49: 516. 1913. TYPE: Mexico. State Guerrero: Iguala Canyon, near Iguala, C. G. Pringle 13921 (isotype, US not seen, US photo at LP!). 6. Trixis californica Kellogg, Proc. Calif. Acad. Sci. 2: 182. 1862. TYPE: Mexico. State Baja California Norte: Cedros Island, Veatch, s.d., s. leg. (holotype, CAS not seen). Additional specimen examined: Mexico. Baja California: Sierra de la Giganta, Pilón de las Parras, W of Loreto, A. Carter & H. Sharsmith 4237 (LP). 7. Trixis calycina D. Don, Trans. Linn. Soc. London 16: 187. 1830. TYPE: Brazil. State Bahia, s.d., C. Martius s.n. (holotype, M not seen, M photo at LP!). Additional specimen examined: Brazil. State Bahía: Mun. Morro da Chapéu, Serra do Tombador, G. Hatschbach & Guimaraes 42354 (LP). 8. Trixis chiapensis C. E. Anderson, Brittonia 23: 347. 1971. TYPE: Mexico. State Chiapas: 24 mi SE of Comitán, W. Anderson & C. Anderson 5494 (holotype, MICH not seen).


653

9. Trixis divaricata (Kunth) Spreng., Syst. veg. 3: 501. 1826. Basionym: Perdicium divaricatum Kunth. TYPE: Peru. Carico, F. Humboldt & A. Bonpland 3650 (holotype, P not seen, P photo F n. 37850 at LP! MO!). Additional specimen examined: Argentina. Prov. Tucumán: Dept. Trancas, El Cadillal, S. Venturi 377 (LP). 10. Trixis glaziovii Baker, Fl. bras. 6: 391. 1884. TYPE: Brazil. State Río de Janeiro: Itatiaia, Campo Feio, A. F. M. Glaziou 6582 (holotype, P!). Additional specimen examined: Brazil. State Minas Gerais: Caldas, 10 Jan 1919, Hoehne s.n. (SP). 11. Trixis glutinosa D. Don, Trans. Linn. Soc. London 16: 189. 1830. TYPE: Brazil. State Minas Gerais: inter Minas Novas et fluvium São Francisci, s.d., C. Martius s.n. (holotype, M not seen, M photo at LP!); State Goiás, rod. br. 153, 10 km W de Anápolis, G. Hatschbach 36682 (LP!). 12. Trixis grandibracteata C. E. Anderson, Brittonia 23: 348. 1971. TYPE: Mexico. State Guerrero: 21 km E of Petatlán, on road to Tlapa, at km 121, W. Anderson & C. Anderson 5729 (isotype, NY not seen, NY photo at LP!). 13. Trixis grisebachii Kuntze, Revis. gen. pl. 3: 183. 1898. TYPE: Argentina. Prov. Jujuy: Jujuy, April 1871, P. G. Lorentz & G. H. E. W. Hieronymus s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Santiago del Estero: Dept. Río Hondo, camino de San Miguel de Tucumán a Río Hondo, L. Katinas et al. 97 (LP!). 14. Trixis haenkei Sch. Bip., Bot. Voy. Herald: 314. 1856. TYPE: Mexico. State Sinaloa: probably from the mountains along the Sinaloa-Durango border, Seemann 2029 (holotype, K not seen). 15. Trixis hassleri Chodat, Bull. Herb. Boissier 3: 782. 1903. TYPE: Paraguay. Dept. Amambay: Río Apa, Hassler 8318 (isotype, GH!). Additional specimen examined: Paraguay. Dept. Amambay: 30 km SE de Bella Vista, A. Krapovickas & A. Schinini 32645 (SI). 16. Trixis hyposericea S. Watson, Proc. Amer. Acad. Arts 25: 157. 1890. TYPE: Mexico. State Jalisco: in the barranca near Guadalajara, C. G. Pringle 1741 (isotype, US not seen, US photo at LP!). 17. Trixis inula Crantz, Inst. rei herb. 1: 329. 1766. TYPE: Jamaica, s.d., s. leg. (holotype, LINN 1000.3 not seen, LINN photo at LP!). Additional specimen examined: Colombia. Dept. Atlántico: Puerto Colombia, Elias 1158 (LP). 18. Trixis lessingii DC., Prodr. 7: 70. 1838. TYPE: Brazil. Brasilia aequinoctialis, F. Sellow 2068 (lectotype, designated by Katinas, 1996b: 72, K!). Additional specimen examined: Brazil. State Santa Catarina: Matos Costa, R. M. Klein 3569 (LP). 19. Trixis megalophylla Greenm., Proc. Amer. Acad. Arts 41: 270. 1905. TYPE: Mexico. State Guerrero: between Sochi and Tlakinsala, E. W. Nelson 2038 (isotype, US not seen, US photo at LP!). 20. Trixis mexicana Lex., Nov. veg. descr. 1: 27. 1824. TYPE: not localized. Additional specimen examined: Mexico. State Michoacán: about 17 mi W of Jiquilpán, A. Cronquist 9757 (LP). 21. Trixis michuacana Lex., Nov. veg. descr. 1. 28. 1824. TYPE: Mexico. State Michoacán: prope Vallisoletum, s.d., LaLlave s.n. (holotype, G-DEL not seen). Additional specimen examined: Mexico. State Jalisco: Barranca du Rio Santiago, Dec 1912, Diguet s.n. (LP).

654

L. Katinas, et al.

22. Trixis nelsonii Greenm., Proc. Amer. Acad. Arts 41: 270. 1905. TYPE: Mexico. State Chiapas: between San Cristóbal and Teopsica, E. W. Nelson 3459 (isotype, US not seen, US photo at LP!). 23. Trixis nobilis (Vell.) Katinas, Darwiniana 34: 74. 1996. Basionym: Castra nobilis Vell. TYPE: Lectotype, designated by Katinas, 1996b: 74, the illustration in Fl. flumin. icon. 8: tab. 82. 1831 (1827). Additional specimen examined: Brazil. State Goiás: Serra dos Pirineus, 20 km of Corumbá de Goiás, on road to Niquelandia, Irwin et al. 18746 (SP). 24. Trixis ophiorhiza Gardner, London J. Bot. 6: 461. 1847. TYPE: Brazil. State Minas Gerais: bewteen the Río de São Francisco and Formigas, G. Gardner 4959 (holotype, BM!). Additional specimen examined: Brazil. Federal district: Fundaçao Zoobotánica, Pires et al. 9583 (LP). 25. Trixis pallida Less., Linnaea 5: 30. 1830. TYPE: Brazil. s.d., Sello 3267 (isotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Chaco: Dept. Sargento Cabral, Colonia La Elisa, Schulz 1050 (LP!). 26. Trixis parviflora C. E. Anderson, Brittonia 23: 351. 1971. TYPE: Mexico. State Oaxaca: ca. 12 km N of Amusgos, on road from Pinotepa Nacional to Tlaxiaco, near km 205, W. Anderson & C. Anderson 5636 (isotype, NY not seen, NY photo at LP!). 27. Trixis peruviana Katinas, Darwiniana 34: 83. 1996. TYPE: Peru. Dept. Lima: Cajatambo, Baños de Churín, Landeman 5425 (holotype, K!). 28. Trixis praestans (Vell.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 61. 1936. Basionym: Castra praestans Vell. TYPE: Lectotype, designated by Katinas, 1996b: 61, the illustration in Fl. flumin. icon. 8: tab. 80. 1831 (1827). Additional specimen examined: Brazil. State Minas Gerais: Grão Mogol, G. Hatschbach 41513 (LP). 29. Trixis pringlei B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 10. 1904. TYPE: Mexico. State Oaxaca: Tomellín Canyon, C. G. Pringle 5894 (holotype, GH not seen). 30. Trixis proustioides Hieron., Bot. Jahrb. Syst. 28: 653. 1901. TYPE: Colombia. Triana 1508 (holotype, B not seen, B photo F n. 16108 at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Santa Bárbara, camino de Santa Clara a El Fuerte, Abra de los Morteros, A. L. Cabrera et al. 27968 (LP). 31. Trixis pruskii D. J. N. Hind, Kew Bull. 55: 382. 2000. TYPE: Brazil. State Bahia: Mun. Río Contas, Carrapato, beira Río da Agua Suja, Ganev 2483 (isotype, US not seen, US photo at LP!). 32. Trixis pterocaulis B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 8. 1904. TYPE: Mexico. State Colima: Manzanillo, E. Palmer 897 (isotype, US not seen, US photo at LP!). 33. Trixis sagasteguii Cabrera, Bol. Soc. Argent. Bot. 10: 42. 1962. TYPE: Peru. Dept. La Libertad: Prov. Bolívar, Catal, ruta Río Marañón-Longotea, A. López & A. Sagástegui 3163 (holotype, LP!). 34. Trixis silvatica B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 12. 1904. TYPE: Mexico. State Oaxaca: unterhalb Totolapam, Seler 1633 (lectotype, designated by Anderson, 1972: 55, GH not seen; isotypes, NY not seen, NY photo at LP!, US not seen, US photo at LP!). Additional specimen examined:


35.

36.

37.

38.

655

Mexico. Oaxaca: 35 mi W of Tehuantepec, along the Pan-American highway, A. Cronquist 9681 (LP). Trixis spicata Gardner, London J. Bot. 6: 462. 1847. TYPE: Brazil. State Minas Gerais: campos near the Río Claro, G. Gardner 4957 (holotype, BM!). Additional specimen examined: Paraguay. Dept. Amambay: Sierra de Amambay, T. Rojas 10251 (LP). Trixis thyrsoidea Dusén ex Malme, Kongl. Svenska Vetensk. Acad. Handl. 12: 116. 1933. TYPE: Brazil. State Paraná: Paraná, Río Branco, P. Dusén 13836 (holotype, S not seen, S photo at LP!). Additional specimen examined: Brazil. State Santa Catarina: Vacas Gordas, Urubici, P. Reitz 6685 (LP). Trixis vauthieri DC., Prodr. 7: 69. 1838. TYPE: Brazil. State Minas Gerais: Tejuco, M. Vauthier 278 (isotype, GH!). Additional specimen examined: Brazil. State Minas Gerais: Serra do Curral, Nova Lima, 5 miles SE of Belo Horizonte, Williams & Assis 8018 (MO). Trixis verbascifolia (Gardner) S. F. Blake, Contr. U. S. Natl. Herb. 22: 655. 1924. Basionym: Bowmania verbascifolia Gardner. TYPE: Brazil. State Río de Janeiro: Organ Mtns., G. Gardner 5797 (holotype, BM!). Additional specimen examined: Brazil. State Río de Janeiro: Serra dos Orgãos, A. L. Cabrera 12255 (LP*).

Literature. Anderson (1972), Katinas (1996b), Hind (2000).

Tribe Stifftieae D. Don D. Don, Trans. Linn. Soc. London 16: 291. 1830. TYPE: Stifftia J. C. Mikan. Stifftioideae (D. Don) Panero, pro parte, Phytologia 89: 356. 2007. TYPE: Stifftia J. C. Mikan. Wunderlichioideae Panero & V. A. Funk, pro parte, Phytologia 89: 357. 2007. Wunderlichieae Panero & V. A. Funk, Phytologia 89: 357. 2007. TYPE: Wunderlichia Riedel ex Benth. Simple to branched subshrubs, shrubs, trees, or vines; stems woolly to glabrous. Leaves simple, alternate, often clustered at stem apex, unarmed; usually petiolate, the petioles, when present, often clasping; blades oblanceolate, less commonly linear, orbicular to elliptic, often thickly coriaceous, glabrous and eglandular, or tomentose to woolly beneath. Capitulescences monocephalous, terminal, less commonly cymose, with few to many capitula. Capitula usually large, homogamous, discoid, with one to ca. 100 flowers; involucre multiseriate, the series graduate; receptacle paleate, partially paleate to epaleate. Florets bisexual, corollas actinomorphic, tubular-funnelform to sub-campanulate, deeply five-lobed, exceptionally bilabiate; anthers mostly exserted, apical appendages acuminate, tails long, smooth to papillose, free to those of adjacent anthers often connate; style cleft into two short, bilobed branches, the branches erect, apically rounded, dorsally rugulose to papillose with the branch apex often smooth and the papillae extending beyond the bifurcation point onto the shaft, occasionally styles bifid, branches recurved and dorsally smooth. Cypselae usually prismatic and glabrous; pappus of scabrid capillary

656

L. Katinas, et al.

bristles, basally connate or not, inner series apically broadened to plumose, unequal to subequal in length. Pollen with exine structure heterogeneous, most genera with compact ectosexine constituted by very densely arranged columellae, and endosexine with stout, ramified columellae; exine surface perforate, microechinate to echinate. Characters that define the tribe are in italics. Stifftieae include six genera and 48 species mostly centered in the Guayana Highlands in Colombia, Guyana, and Venezuela. Some species are distributed in the Andes of Ecuador and Peru, and central-eastern Brazil. Genera: Chimantaea Maguire, Steyerm. & Wurdack, Quelchia N. E. Br., Stenopadus S. F. Blake, Stifftia J. C. Mikan, Stomatochaeta (S. F. Blake) Maguire & Wurdack, Wunderlichia Riedel ex Benth. Key to Genera 1a. Leaves caducous; plants flowering in the leafless condition; pappus connate basally, deciduous as a ring, bristles barbellate to plumose ……. Wunderlichia 1b. Leaves commonly long-persistent; plants thus flowering when leafy; pappus basally free, persistent, bristles scabrid. 2a. Corolla lobes stiffly erect to very slightly recurved at extreme apex; corolla cream-colored, not reddish. 3a. Branched or unbranched subshrubs or shrubs; leaves tomentose to woolly beneath; corolla lobes frequently pilose at tip; anther tails free; cypselae often pilose .…………………………… Chimantaea 3b. Much-branched subshrubs or trees; leaves glabrous or, if pubescent, then generally rapidly glabrate; corolla lobes apically glabrous; tails of adjacent anthers appressed or truly connate; cypselae usually glabrous ……….…………………………………… Stomatochaeta 2b. Corolla lobes strongly coiled to flexuous, never stiffly erect; corolla cream-colored, yellowish, or reddish. 4a. Capitula one-flowered, in axillary to terminal glomerate cymes; anther tails papillose ………..……………………………. Quelchia 4b. Capitula many-flowered in monocephalous or in lax capitulescences; anther tails smooth to occasionally papillose. 5a. Shrubby trees or vines; receptacles epaleate; corolla tube fivenerved; anther tails free ……………………………….. Stifftia 5b. Shrubs to big trees; receptacles sometimes paleate; corolla tube ten- to 15-nerved; anther tails often connate …….. Stenopadus

Chimantaea Maguire, Steyerm. & Wurdack Mem. New York Bot. Gard. 9: 428. 1957. TYPE: Chimantaea mirabilis Maguire, Steyerm. & Wurdack. 71 of Figs. 71–74. Etymology. Named for the Chimantá Massif (state Bolívar, Venezuela), the world’s largest Massif of closely clustered tepuis, and where all the species of the genus occur.


657

Figs. 71–74 71 Chimantaea mirabilis Maguire, Steyerm. & Wurdack. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (Redrawn from Aristeguieta, 1964: 886). Scale bars: A=3 cm; B, E=12 mm; C=4.5 mm; D=1 mm. 72 Quelchia cardonae Steyerm. A Habit (branch). Q. eriocaulis Maguire, Steyerm. & Wurdack. B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (A redrawn from Pruski, 1997: 355; B–E from Huber 11591, US). Scale bars: A=4 cm; B=7 mm; C=3 mm; D=1 mm; E=5 mm. 73 Stenopadus andicola Pruski. A Habit (branch). B Corolla. S. huachamacari Maguire. C Stamen. D Upper part of the style. S. andicola Pruski. E Cypsela with pappus. (A, B, E redrawn from Pruski & Beltrán, 2003: 10; C, D from Maguire et al. 30116, MO). Scale bars: A= 5 cm; B, E=10 mm; C=9 mm; D=1 mm. 74 Stifftia chrysantha J. C. Mikan. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (A, B, E redrawn from Baker, 1884: Figure 94; C, D from Hutchinson 8832, US). Scale bars: A=3 cm; B, C, E=10 mm; D=1.2 mm

Shrubs or subshrubs unbranched to branched. Leaves alternate, densely inserted, commonly clustered at stem apex; sessile to petiolate; blades linear, rhomboidal to obovate, thickly coriaceous, pinnately veined, margin entire, frequently revolute, glabrous above except sometimes along midrib, tomentose to woolly beneath. Capitulescences monocephalous, terminal; capitula sessile, homogamous, discoid; receptacle epaleate or with a few deciduous marginal paleae; involucre multiseriate. Florets isomorphic, bisexual, actinomorphic, tubular-funnelform, deeply five-lobed, lobes stiffly erect to slightly recurved, frequently pilose at the tip; anthers yellow to reddish, generally exserted, apical appendages acute to acuminate, tails smooth, free;

658

L. Katinas, et al.

style yellow to reddish, bilobed, branches dorsally rugulose to papillose much below the bifurcation point. Cypselae often densely pilose; pappus of scabrid bristles, the innermost somewhat longer, apically broadened. Pollen Description. Chimantaea (116 of Figs. 112–116). Pollen subprolate, elliptic in equatorial view, circular in polar view, large size. Tricolporate, long colpi with scabrate–microgranulate membrane. Exine Mutisia type, microechinate. Nexine ca. 2 μm thick (see also Carlquist, 1957b). Chimantaea cinerea (Gleason & S. F. Blake) Maguire, Steyerm. & Wurdack. Pollen size P × E = (64–68×50–54) μm. Exine 6 μm thick. Ratio ectosexine/endosexine: ca. 1:1. Chimantaea huberi. Pollen size P × E = (48–57×37–42) μm. Exine 5.5–6 μm thick, ectosexine slightly thickened at the poles. Ratio ectosexine/endosexine: 1:1; 1.5:1. SEM: tectum very perforate, apertural membrane scabrate (116 of Figs. 112–116). Note: pollen similar to that of Neblinaea promontorium, Stifftia uniflora, Stenopadus campestris, S. colveii and S. huachamacari (see also Tellería, 2008). Habitat and Distribution. Chimantaea is a genus of nine species endemic to the eastern Venezuelan Guayana Highland. All nine species of Chimantaea occur on the Macizo del Chimantá, but subspecies of two species are disjunct, occurring outside of the massif on Auyan-tepuí and Murisipán-tepuí. Species list 1. Chimantaea acopanensis Steyerm., Ann. Missouri Bot. Gard. 71: 335. 1984. TYPE: Venezuela. State Bolívar: Piar, Massif of Chimantá, plateau of Acopantepui, J. A. Steyermark et al. 129932 (isotype, US!). 2. Chimantaea cinerea (Gleason & S. F. Blake) Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 429. 1957. Basionym: Stenopadus cinereus Gleason & S. F. Blake. TYPE: Venezuela. State Bolívar, Auyantepui, G. H. H. Tate 1111 (holotype, NY!). Additional specimen examined: Venezuela. Bolívar, B. K. Host 3745 (MO). 3. Chimantaea eriocephala Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 431. 1957. TYPE: Venezuela. State Bolívar: Chimantá massif Churitepuí (Muru-tepuí), J. J. Wurdack 34225 (holotype, NY!). 4. Chimantaea espeletoidea Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 433. 1957. TYPE: Venezuela. State Bolívar: Chimantá massif, below upper falls of Río Tirica, J. A. Steyermark & J. J. Wurdack 574 (isotype, NY!). 5. Chimantaea huberi Steyerm., Ann. Missouri Bot. Gard. 71: 335. 1984. TYPE: Venezuela. State Bolívar: Piar, J. A. Steyermark et al. 128815 (isotype, US!). Additional specimen examined: Venezuela. State Bolívar: Mun. Gran Sabana, macizo de Chimantá, SE del Apacará-tepui, O. Huber 12034 (LP*). 6. Chimantaea humilis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 432. 1957. TYPE: Venezuela. State Bolívar: Chimantá massif, central section, between W and E branches of headwaters of Río Tirica, J. A. Steyermark & J. J. Wurdack 747 (holotype, NY!). 7. Chimantaea lanocaulis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 430. 1957. TYPE: Venezuela. State Bolívar: Apacara-tepui, J. A. Steyermark 75912 (isotype, US!).


659

8. Chimantaea mirabilis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 432. 1957. TYPE: Venezuela. State Bolívar: Río Tirica, J. A. Steyermark & J. J. Wurdack 821 (isotype, US!). 9. Chimantaea rupicola Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 431. 1957. TYPE: Venezuela. State Bolívar: Río Tirica, J. A. Steyermark & J. J. Wurdack 748 (holotype, US!).

Literature. Maguire et al. (1957b), Aristeguieta (1964), Pruski (1997).

Quelchia N. E. Br. Trans. Linn. Soc. London, Bot. 6: 41. 1901. TYPE: Quelchia conferta N. E. Br. 72 of Figs. 71–74. Etymology. Named in honor of John Quelch who, together with Frederick McConnell, climbed Mount Roraima in 1894 and 1898 and collected the type material. Shrubs or small trees monopodial to few-branched. Leaves alternate, often clustered at the stem apex; petiolate; blades oblanceolate, elliptic to obovate, coriaceous, pinnately veined, margin entire, glabrous to densely woolly beneath. Capitulescences cymose, glomerate, axillary or terminal; capitula long-pedunculate to subsessile, homogamous, one-flowered; receptacle epaleate; involucre multiseriate. Floret bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed to rarely becoming nearly bilabiate, lobes coiled at apex; anthers sometimes redstreaked, partly to mostly exserted, apical appendages acuminate, tails papillose, free; style bifid, sometimes reddish, branches dorsally smooth. Cypselae glabrous to slightly pilose; pappus of scabrid, capillary bristles, the innermost slightly apically broadened and longer. Pollen Description. Quelchia bracteata Maguire, Steyerm. & Wurdack (117 of Figs. 117–118). Pollen subprolate, elliptic-subrectangular in equatorial view, circular in polar view, large size, P × E = (55–62×45–48) μm. Tricolporate, colpi long with margin and microgranulate membrane. Exine Gongylolepis type, 7–10 μm thick in the whole grain, spines ca. 2.5 μm length. Ratio ectosexine/endosexine: 1:2; 1:2.5. Nexine ca. 2 μm thick. SEM: tectum very perforate. Pollen similar to that of Duidaea marahuacensis and Glossarium rhodanthum (see also Carlquist, 1957b; Tellería, 2008). Habitat and Distribution. Four species plus one named hybrid, all endemic to the Guayana Highland in southern Venezuela. Species list 1. Quelchia bracteata Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 435. 1957. TYPE: Venezuela. State Bolívar: Chimantá, J. A. Steyermark & J. J. Wurdack 683 (holotype, NY!). Additional specimens examined: Venezuela.

660

2.

3.

4.

5.

L. Katinas, et al.

State Bolívar: Dist. Piar, macizo del Chimantá, O. Huber 11591 (US); id., J. Pipoly et al. 7217 (US*). Quelchia cardonae Steyerm., Fieldiana, Bot. 28: 670. 1953. TYPE: Venezuela. State Bolívar: cumbre del cerro Acopan, Río Caroni, Guayana, F. Cardona 2288 (holotype, US!). Quelchia conferta N. E. Br., Trans. Linn. Soc. London, Bot. 6: 41. 1901. TYPE: Border of Roraima, Brazil, Guyana, and Bolívar, Venezuela: Mt. Roraima, F. McConnell & J. Quelch 652 (holotype, K!). Quelchia eriocaulis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 435. 1957. TYPE: Venezuela. State Bolívar: Toroni-tepui, J. A. Steyermark & J. J. Wurdack 675 (isotype, US!). Quelchia xgrandifolia Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 436. 1957. TYPE: Venezuela. State Bolívar: Chimantá Massif, Toronotepui, J. A. Steyermark & J. J. Wurdack 756 (isotype, NY!).

Observation. Quelchia combines characteristics of Stifftieae and Mutisieae. Species of the genus have smooth style branches, and capitula with corollas that range from tubular, slightly zygomorphic, to bilabiate. Literature. Maguire et al. (1957a), Aristeguieta (1964), Pruski (1991, 1997).

Stenopadus S. F. Blake Bull. Torrey Bot. Club 58: 489. 1931, emend. Maguire & Wurdack, and collaborators, Mem. New York Bot. Gard. 9: 379. 1957. TYPE: Stenopadus talaumifolius S. F. Blake. 73 of Figs. 71–74. Etymology. From the Greek steno, narrow, and padus, foot, in reference to the narrow paleae at the “foot” of the cypselae as often seen in the type species. Shrubs or big trees. Leaves alternate; petiolate; blades oblanceolate to orbicular, coriaceous, pinnately veined, margin entire, often revolute, usually glabrous. Capitulescences monocephalous or cymes two- to four-headed, terminal; capitula sessile to short-pedunculate, homogamous, discoid; receptacle moderately paleate to epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed, tube ten- to 15-nerved, lobes strongly coiled to less commonly flexuous; anthers sometimes green to purplish, exserted, apical appendages acuminate, tails smooth to papillose, often connate with the tails of adjacent stamens; style sometimes reddish, bilobed, branches dorsally papillose much below the bifurcation point. Cypselae glabrous; pappus of scabrid, capillary bristles to flattened and widened at the apex, the innermost longer. Pollen Description. Stenopadus (118 of Figs. 117–118). Pollen prolate, elliptic or subrectangular in equatorial view, circular–subcircular in polar view, large size.


661

Tricolporate, colpi long or short, with psilate or microgranulate membrane, with or without diffuse mesoaperture. Exine Stenopadus type and Mutisia type, microechinate, equally thickened in the whole grain or with polar caps, scabrate. Ratio ectosexine/endosexine: 1:1; 1:1.5 (see also Carlquist, 1957b; Barroso & Maguire, 1973; Tellería, 2008). Stenopadus chimantensis Maguire, Steyerm. & Wurdack (118A, B of Figs. 117–118). Pollen size P × E = (62–72×36–52) μm. Colpi 45– 50 μm length with membrane densely microgranulate around the endoaperture. Exine Stenopadus type, with polar caps, 3–6 μm thick at the equator and 7–9 μm at the poles. Ratio ectosexine/endosexine: 1:1. Nexine 2–2.5 μm thick. SEM: tectum psilate at the poles and very perforate with striate aspect at the equator. Pollen similar to that of Stenopadus connellii, S. jauaensis, S. sericeus, S. talaumifolius, and Stomatochaeta condensata (see also Tellería, 2008). Stenopadus connellii (N. E. Br.) S. F. Blake (118E of Figs. 117–118). Pollen size P × E = (67–75×42–50) μm. Colpi 40–50 μm length with granulate membrane. Exine Stenopadus type, psilate, 5–5.5 μm thick in the whole grain. Nexine 2 μm thick. SEM: tectum very perforate, with striate aspect and very small microspines. Note: some grains have more than three apertures, and pyramidal or quadrangular shape. Pollen similar to that of Stenopadus chimantensis, S. jauensis, S. sericeus, S. talaumifolius, and Stomatochaeta condensata (see also Tellería, 2008). Stenopadus huachamacari Maguire (118D of Figs. 117–118). Pollen size P × E = (52–64×36–43) μm. Colpi long with psilate margin and scabrate membrane, mesoaperture diffuse. Exine Mutisia type, 4–6 μm thick, slightly thickened at the poles. Nexine 2 μm thick. SEM: tectum very perforate. Pollen similar to that of Chimantaea huberi, C. cinerea, Neblinaea promontorium, Stifftia uniflora, Stenopadus campestris, and S. colveii (see also Tellería, 2008). Stenopadus talaumifolius (118C of Figs. 117–118). Pollen size P × E = (81.5–90×50–61) μm. Colpi 53–64 μm length with membrane psilate or scarcely microgranulate. Exine Stenopadus type, scabrate, 6–8 μm thick in the whole grain. Nexine ca. 2.5 μm thick. Note: pollen similar to that of Stenopadus chimantensis, S. connelli, S. jauensis, S. sericeus, and Stomatochaeta condensata (see also Tellería, 2008). Habitat and Distribution. Stenopadus is a genus of 15 species, 14 of these found in the sandstone areas of the Guayana Highland in Colombia, Venezuela, Guyana, and Brazil, and one species found in the Andes of Ecuador and Peru. Species list 1. Stenopadus andicola Pruski, Novon 8: 67. 1998.. TYPE: Ecuador. ZamoraChinchipe: Cantón Nangaritza, valle del Río Nangaritza, Miazi, W. Palacios 6712 (holotype, US!). 2. Stenopadus aracaënsis Pruski, Ann. Missouri Bot. Gard. 76: 999. 1989. TYPE: Brazil. State Amazonas: plateau of northern massif of Serra Aracá, G. T. Prance et al. 29184 (holotype, IMPA!). 3. Stenopadus campestris Maguire & Wurdack, Mem. New York Bot. Gard. 9: 383. 1957. TYPE: Venezuela. State Amazonas: Puerto Pimichin, B. Maguire & J. J. Wurdack 35578 (holotype, NY!). 4. Stenopadus chimantensis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 427. 1957. TYPE: Venezuela. State Bolívar: Chimantá, Toronó-

662

5.

6.

7.

8.

9.

10. 11.

12.

13.

14.

15.

L. Katinas, et al.

tepui, J. A. Steyermark & J. J. Wurdack 652 (holotype, NY!). Additional specimen examined: Venezuela. State Bolívar: Distr. Piar, macizo del Chimantá, sección centro del Churí-tepui, J. Pipoly et al. 7142 (MO*). Stenopadus colombianus Cuatrec. & Steyerm., Bot. Mus. Leafl. 17: 99. 1955. TYPE: Colombia. Vaupes: Río Kananarí, cerro Isibukurí, R. E. Schultes & I. Cabrera 15031 (holotype, F!). Stenopadus colveei (Steyerm.) Pruski, Ann. Missouri Bot. Gard. 76: 998. 1989. Basionym: Stomatochaeta colveei Steyerm. TYPE: Venezuela. State Bolívar: cumbre Cerro Guaiquinima, J. A. Steyermark et al. 117407 (holotype, VEN!). Stenopadus connellii (N. E. Br.) S. F. Blake, Bull. Torrey Bot. Club 58: 490. 1931. Basionym: Stifftia connellii N. E. Br. TYPE: Border of Roraima, Brazil; Guyana; and Venezuela. Mount Roraima, F. McConnell & J. Quelch 661 (holotype, K!). Additional specimen examined: Venezuela. State Bolívar: Kukenán-tepui, R. Liesner 23109 (MO*). Stenopadus cucullatus Maguire & Wurdack, Mem. New York Bot. Gard. 9: 387. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Yutaje, B. Maguire & C. K. Maguire 35053 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atures, Caño Yutaje, at S base of Serranía de Yutaje, Holst & R. Liesner 3043 (MO). Stenopadus huachamacari Maguire, Mem. New York Bot. Gard. 8: 151. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, B. Maguire et al. 30026 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, Río Cunucunuma, B. Maguire et al. 30116 (MO*). Stenopadus jauensis Aristeg., Acta Bot. Venez. 3: 40. 1968. TYPE: Venezuela. Meseta de Jáua: cerro Jáua, cumbre de Ika, J. A. Steyermark 97867 (isotype, NY!). Stenopadus kunhardtii Maguire, Mem. New York Bot. Gard. 8: 151. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Sinapo, B. Maguire & L. Politi 28304 (holotype, US!). Stenopadus megacephalus Pruski, Ann. Missouri Bot. Gard. 76: 1001. 1989. TYPE: Guayana. Upper Mazaruni River basin, Mt. Ayanganna, S. S. Tillett et al. 45100 (holotype, NY!). Stenopadus obconicus Maguire & Wurdack, Mem. New York Bot. Gard. 9: 381. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37041 (holotype, US!). Stenopadus sericeus Maguire & Aristeg., Mem. New York Bot. Gard. 9: 385. 1957. TYPE: Venezuela. Ilo-tepuí: Gran Sabana, B. Maguire 33490 (holotype, NY!). Stenopadus talaumifolius S. F. Blake, Bull. Torrey Bot. Club 58: 491. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Duida, G. H. H. Tate 759 (holotype US!). Additional specimen examined: Venezuela. Dept. Atapabo, Huachamacari, R. Liesner 18346 (MO*).

Literature. Maguire et al. (1957a, b), Aristeguieta (1964), Pruski (1989b, 1991, 1997, 1998), Díaz-Piedrahita and Vélez-Nauer (1993), Pruski and Beltrán (2003).


663

Stifftia J. C. Mikan Del. fl. faun. bras. 1: t. 1. 1820. TYPE: Stifftia chrysantha J. C. Mikan. 74 of Figs. 71–74. Aristomenia Vell., Fl. flumin.: 8. 1829 (1825). TYPE: Aristomenia fruticosa Vell. [= Stifftia fruticosa (Vell.) D. J. N. Hind & Semir]. Augusta Leandro, Denkschr. Königl. Akad. Wiss. München 7: 235. 1820, non Pohl, 1828, nom. rejic. TYPE: Augusta grandiflora Leandro [= Stifftia chrysantha J. C. Mikan]. Mocinia DC. Prodr. 7: 26. 1838. TYPE: Mocinia mutisioides DC. [= Stifftia chrysantha J. C. Mikan]. Sanhilaria Leandro ex DC., Prodr. 7: 26. 1838, nom. nudum. Etymology. Named for C. E. Stifft, geologist and author of several works from ca. 1807–1832. Trees shrubby or vines. Leaves alternate; petiolate; blades elliptical to oblong, chartaceous or less commonly sub-coriaceous, pinnately veined, margin entire, completely glabrous to weakly arachnoid beneath. Capitulescences corymbose to paniculate, less commonly monocephalous; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular-funnelform, deeply fivelobed, tube five-nerved, the lobes completely coiled or only near the apex; anther apical appendages acuminate, tails smooth to sometimes papillose, free; style bilobed, branches dorsally smooth to slightly rugulose at the apex. Cypselae glabrous to weakly puberulent; pappus of scabrid bristles, capillary to somewhat paleaceous, the innermost slightly apically broadened and longer. Pollen Description. Stifftia (Fig. 119). Pollen spheroidal, elliptic in equatorial view, circular in polar view, large size. Tricolporate, colpi long with margin, and scabrate or psilate membrane. Exine Mutisia type, microechinate, with or without polar caps. Ratio ectosexine/endosexine: 1:1 (see also Carlquist, 1957b; Tellería, 2008). Stifftia chrysantha (Fig. 119). Pollen elliptic, subtriangular in polar view, planaperturate, P × E = (66.5–85×58–65) μm. Exine with polar caps, 8–14 μm thick at the equator and 11–16 μm thick at the poles, ectosexine and internal tectum very thickened at the poles. Ratio ectosexine/endosexine: 2:1; 1:1. Nexine ca. 2 μm thick. SEM: tectum with very small perforations. Stifftia uniflora Ducke. Pollen spheroidal, circular in polar view. P × E = (48–60×42–54) μm. Exine 6– 6.5 μm thick in the whole grain. Nexine ca. 1.5 μm thick. Note: some grains lack the endoaperture. Pollen similar to that of Chimantaea huberi, C. cinerea, Neblinaea promontoriorum, Stenopadus campestris, S. huachamacari and S. colveii (see also Tellería, 2008). Habitat and Distribution. Stifftia is a genus of seven species occurring in Brazil and French Guiana. Stifftia is the only genus of this tribe occurring both north and south of the Amazon River.

664

L. Katinas, et al.

Species list 1. Stifftia cayennensis H. Rob. & B. Kahn, Phytologia 58: 248. 1985. TYPE: French Guiana. Cayenne route de Saint Laurent à Paul Isnard, entre le PK 70 et Citron, au PK 118, G. Cremers 8153 (holotype, US!). 2. Stifftia chrysantha J. C. Mikan, Del. fl. faun. bras. 1: 1. 1820. TYPE: Brazil. Río de Janeiro, s.d., Mikan s.n. (isotype, NY!). Additional specimens examined: Brazil, Hutchinson 8832 (US); Río de Janeiro, Jardín Botánico, Cabrera 12242 (LP*). 3. Stifftia fruticosa (Vell.) D. J. N. Hind & Semir, Kew Bull. 53: 618. 1998. Basionym: Aristomenia fruticosa Vell. Icon. fl. flumin. 8: tab. 84. 1831 (1827). TYPE: Lectotype, designated by Hind & Semir, 1998a: 618, the illustration Icon. fl. flumin. 8: tab. 84. 1831 (1827). 4. Stifftia hatschbachii H. Rob., Syst. Bot. 16: 687. 1991. TYPE: Brazil. Espirito Santo: Mun. Linhares, Reserva Forestal Sooretama, G. Hatschbach et al. 53506 (isotype, US!). 5. Stifftia parviflora (Leandro) D. Don, Trans. Linn. Soc. London 16: 291. 1833. Basionym: Augusta parviflora Leandro. TYPE: Brazil. State Rio de Janeiro: near Rio de Janeiro, s.d., Leandro s.n. (type, probably P). Additional specimens examined: Brazil. State Minas Gerais: Faz. do Brejao, José do Alfredo, Cordesburgo, Heringer 7680 (LP*); State São Paulo: Piassaguéra, Lemos 17110 (LP*). 6. Stifftia racemosa H. Rob., Syst. Bot. 16: 691. 1991. TYPE: Brazil. State Minas Gerais: capoeira Ibirité, municipio de Betim, G. Mendes Magalhães 974 (holotype, US!). 7. Stifftia uniflora Ducke, Arq. Inst. Biol. Veg. 2: 72. 1935. TYPE: Brazil. State Amazonas: Río Negro, Santa Isabel, A. Ducke 24148 (isotype, US!). Additional specimen examined: Brazil. State Amazonas: Río Negro, Santa Isabel, A. Ducke s.n. (LP*). Literature. Maguire et al. (1957a), Robinson and Kahn (1985), Pruski (1991), Robinson (1991), Roque and Pirani (1997), Hind and Semir (1998a).

Stomatochaeta (S. F. Blake) Maguire & Wurdack Mem. New York Bot. Gard. 9: 388. 1957, based on Stenopadus subgen. Stomatochaeta S. F. Blake, Bull. Torrey Bot. Club 58: 490. 1931. TYPE: Stenopadus crassifolius S. F. Blake [= Stomatochaeta crassifolia (S. F. Blake) Maguire & Wurdack]. 75 of Figs. 75–76. Dermatophyllum Sch. Bip. ex M. R. Schomb., Reis. Br.-Guiana 3: 1079. 1848 (1849), non Scheele 1848, nom. nudum. Etymology. From the diminutive of Greek stoma, mouth, and chaite, hair, in reference to the “hairy mouth” (technically the pubescent upper portion of the corolla throat) of the corollas. Subshrubs or trees much branched. Leaves alternate, often clustered at the stem apex; petiolate; blades narrowly spathulate, oblanceolate to obovate, coriaceous,


665

pinnately veined, margins entire, thickened, revolute, glabrous, if pubescent, then generally rapidly glabrate. Capitulescences monocephalous, terminal; capitula sessile to subsessile, homogamous, discoid; receptacle partially paleate to epaleate; involucre multiseriate. Florets isomorphic, bisexual, actinomorphic, tubular-funnelform, very deeply five-lobed, lobes very long, stiffly erect, glabrous; anthers generally exserted, apical appendages acuminate, tails smooth to very slightly papillose, free but appressed to the tails of adjacent stamens to truly connate; style bilobed, branches dorsally papillose much below the bifurcation point. Cypselae usually glabrous; pappus of scabrid, capillary bristles, the innermost apically broadened and longer. Pollen Description. Stomatochaeta condensata (120 of Figs. 120–121). Pollen subprolate, spheroidal-elliptic in equatorial view, circular in polar view, large size, P × E = (52–55×40–45) μm. Tricolporate, colpi long with microgranulate membrane. Exine Stenopadus type 4–5 μm thick, slightly thickened at the poles. Nexine 2 μm thick. Note: pollen similar to that of Stenopadus chimantensis, S. connelli, S. jauensis, S. sericeus and S. talaumifolius (see also Carlquist, 1957b; Tellería, 2008). Habitat and Distribution. A genus of six species, all of which are endemic to the Guayana Highland of southern Venezuela, Guyana, and northern Brazil. Species list 1. Stomatochaeta acuminata Pruski, Brittonia 41: 35. 1989. TYPE: Venezuela. State Bolívar: Piar, summit of Angasima, O. Huber 11374 (holotype, VEN!). 2. Stomatochaeta condensata (Baker) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stifftia condensata Baker. TYPE: Guyana. Roraima, Schomburgk 543 (holotype, K not seen). Additional specimens examined: Venezuela. State Bolívar: R. Liesner 19415 (MO*); cumbre de Cerro Guaiquinima, río Carapo, J. Steyermark & E. Dunsterville 113254 (US).

Figs. 75–76 75 Stomatochaeta condensata (Baker) Maguire & Wurdack. A Habit (branch). B Corolla. C Stamen. D Upper part of the style. E Cypsela with pappus. (A from Steyermark & Dunsterville 113524, US; B–E from Liesner 19415, MO). Scale bars: A=3.5 cm; B=7 mm; C=6.5 mm; D=2 mm; E=6 mm. 76 Wunderlichia mirabilis Riedel ex Baker. A Habit (branch with immature leaves). B Palea. C Corolla. D Stamen. E Upper part of the style. F Cypsela with pappus. (From Macedo 3527, MO). Scale bars: A=5 cm; B, D=10 mm; C=18 mm; E=0.8 mm

666

L. Katinas, et al.

3. Stomatochaeta crassifolia (S. F. Blake) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stenopadus crassifolius S. F. Blake. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida, central camp., G. H. H. Tate 1014 (isotype, US!). 4. Stomatochaeta cylindrica Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. TYPE: Venezuela. State Bolívar: Cerro Guaiquinima, Río Paragua, 1 km E of Cumbre Camp, B. Maguire 32782 (holotype, NY!). 5. Stomatochaeta cymbifolia (S. F. Blake) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stenopadus cymbifolius S. F. Blake. TYPE: Venezuela. State Bolívar: Auyán-tepui, G. H. H. Tate 1322 (holotype, NY!). 6. Stomatochaeta steyermarkii Aristeg., Acta Bot. Venez. 3: 39. 1968. TYPE: Venezuela. Bolívar: Jaua meseta, cerro Jaua, J. A. Steyermark 97917 (isotype, US!).

Literature. Maguire et al. (1957a), Aristeguieta (1964), Pruski (1989a, 1991, 1997).

Wunderlichia Riedel ex Benth. in Benth. & Hook. f., Gen. pl. 2: 489. 1873. TYPE: Wunderlichia mirabilis Riedel ex Baker (lectotype designated by Hind & Semir, 1998b). 76 of Figs. 75–76. Etymology. Named in honor of Mr. Wunderlich, a collector of Russian and Asian flora in the 1830s (his main sets of duplicates are at K & LE), a contemporary of Riedel, and an author of one botanical paper in 1842. Shrubs or small trees very stout, virgate, pyrrophytic. Leaves alternate, deciduous; commonly petiolate; blades ovate to orbicular, coriaceous, less commonly chartaceous, caducous, pinnately veined, margin entire, both surfaces densely tomentose to glabrate. Capitulescences monocephalous or two- to ten-headed, loosely to densely corymbose to racemose; capitula massive, short-pedunculate to sessile, homogamous, discoid; receptacle strongly paleate-setiferous to epaleate, paleae long-linear, flat; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular-campanulate, deeply five-lobed, lobes coiled; anthers exserted, apical appendages short-acuminate, tails smooth, free to often connate with the tails of adjacent stamens; style bilobed, dorsally papillose much below the bifurcation point. Cypselae glabrous, apically tomentose to densely tomentose throughout; pappus of many barbellate to plumose bristles, basally connate, deciduous as a ring, the innermost longer and flattened. Pollen Description. Wunderlichia mirabilis (121 of Figs. 120–121) (Barroso & Maguire, 1973; Tellería et al., 2003). Pollen prolate, large size, tricolporate, exine Wunderlichia type. Pollen similar to that of Wunderlichia azulensis, Eurydochus bracteatus and Salcedoa mirabaliarum (see also Tellería, 2008). Habitat and Distribution. The genus occurs in Brazil (states of Bahia, Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro) and contains six species.


667

Species list 1. Wunderlichia azulensis Maguire & G. M. Barroso, Revista Brasil. Biol. 33: 402. 1973. TYPE: Brazil. State Minas Gerais: Pedra Azul, G. Mendes Magalhães 12039 (holotype, NY!). Additional specimen examined: Brazil. State Minas Gerais: Mun. Pedra Azul, 10 km E da cidade na estrada para Almenara, R. M. Harley et al. 25209 (MO).

Fig. 77 Styles of Mutisioideae (SEM): A, B. Tribe Mutisieae. A. Style branches papillose (Chaptalia nutans, from Job s.n., LP). B. Style branches smooth (Gochnatia patazina, from Macbride & Featherstone 2520, US). C, D. Tribe Stifftieae. C. Style papillose below the branches bifurcation point (Stenopadus kunhardti, from Maguire & Politi 28303, NY). D. Style branches rugulose (Stifftia chrysantha, from Cabrera 12242, LP). E, F. Tribe Nassauvieae. E. Style branches apically crowned by collector hairs. E. Detail of the style branch apex (Leucheria purpurea, from Goodall 4146, GH)

668

L. Katinas, et al.

Fig. 78 Style papillae in genera of Mutisioideae and related subfamilies (SEM). a–c Subfamily Mutisioideae. a Tribe Mutisieae (Chaptalia nutans, from Job s. n., LP). b Tribe Stifftieae (Stenopadus kunhardtii, from Maguire & Politi 28303, NY). c Tribe Nassauvieae (Proustia pyrifolia, from Morrison et al. 17163, GH). d, e Subfamily Cichorioideae, tribe Lactuceae. d Portion of style. e Detail of the papillae (Taraxacum officinale, from Sancho 103, LP). f Subfamily Carduoideae, detail of the papillae (Carduus pycnocephalus, from Cabrera 8559, LP)


669

Fig. 79 Styles of generic groups within the subfamily Mutisioideae, tribe Mutisieae (SEM). a Ainsliaea group, note the rounded papillae at the apex (Ainsliaea latifolia, from Handel & Mazzetti 8501, W). b Hecastocleis schockleyi, detail of the smooth surface (Train 3973, from LP). c Eriachaenium magellanicum, note the rounded papillae (O’Donnell 41079, from LP)

2. Wunderlichia bahiensis Maguire & G. M. Barroso, Revista Brasil. Biol. 33: 404. 1973. TYPE: Brazil. Bahía: entre Lençóis e Itaberaba, E. Pereira 2063 (holotype, RB not seen). Additional specimen examined: Brazil. State Bahía: Río das Contas, estrada para a cachoeira do Fraga, no Río Brumado, a 3 km de Municipio de Río de Contas, A. Furlan et al. 1703 (US). 3. Wunderlichia crulsiana Taub., Bot. Jahrb. Syst. 21: 456. 1896. TYPE: Brazil. State Goiás: Serra dos Veadeiros, Ule 28 (lectotype, designated by Barroso & Maguire, 1973: 403, PR not seen). Additional specimen examined: Brazil. State Goiás: Chapada dos Veadeiros (near Alto Paraíso de Goiás), J. A. Ratter et al. 2615 (MO). 4. Wunderlichia insignis Baill., Bull. Mens. Soc. Linn. Paris 1: 285. 1881. TYPE: Brazil. State Rio de Janeiro: Nova Friburgo, A. F. M. Glaziou 12060 (lectotype, designated by Barroso & Maguire, 1973: 404, PR not seen; isotype, NY!). 5. Wunderlichia mirabilis Riedel ex Baker, Fl. bras. 6: 343. 1884. TYPE: Brazil. State Minas Gerais: Serra da Caraça and Serra da Lapa, Riedel 493 (lectotype, designated

670

L. Katinas, et al.

Figs. 80–85 80 Pollen of Actinoseris angustifolia (Gardner) Cabrera. A SEM equatorial view showing the aperture and the microechinate surface. B, C LM equatorial view. B Lalongate endoaperture. C Optical section showing the exine structure. (From Hatschbach 29986, LP). Scale bars: 10 μm. 81 Pollen of Adenocaulon bicolor Hook. A SEM equatorial view. B LM equatorial view in high focus showing the lalongate endoaperture (arrow). C LM polar view in optical section showing the compact ectosexine and the columellate endosexine. (From Hunziker 4943, LP). Scale bars: A=0.5 μm; B, C=10 μm. 82 Pollen of Ainsliaea DC. A SEM equatorial view in A. uniflora Sch. Bip. showing the colporate aperture (from US 1132181). B LM equatorial view in optical section in A. macroclinidioides Hayata showing the colpate aperture and the two columellatae layers of exine (from Morata 17066, AAU). Scale bars: 10 μm. 83 Pollen of Aphyllocladus sanmartinianus Molfino. SEM polar view showing the exine surface slightly microechinate (from Cabrera et al. 24290, LP). Scale bar: 10 μm. 84 Pollen of Brachyclados lycioides D. Don. A SEM polar view showing the exine few sculpturate. B LM equatorial view in optical section. (From Cabrera & Fabris 14865, LP). Scale bars: 10 μm. 85 Pollen of Chaetanthera Ruiz & Pav. A Pollen type I, LM equatorial view in optical section in C. revoluta (Phil.) Cabrera, showing the elliptic to subrectangular nexine outline (from Cabrera 8874, LP). B Pollen type II, LM equatorial view in optical section in C. brachylepis Phil. showing the dumbbell-shaped nexine outline and the endocingulus (from Barros 7443, LP). Scale bars: 10 μm


671

Figs. 86–91 86 Pollen of Chaptalia nutans (L.) Pol. A SEM equatorial view showing the aperture membrane densely microgranulate. B SEM polar view. (From Joly 145AB3, LP). Scale bars: 10 μm. 87 Pollen of Chucoa ilicifolia Cabrera. A SEM equatorial view showing the smooth apertural membrane. B SEM polar view showing the small microspines. (From López 1090, LP). Scale bars: 10 μm. 88 Pollen of Cnicothamnus lorentzii Griseb. A SEM equatorial view. B LM equatorial view in optical section. (From Maldonado 408, LP). Scale bars: 10 μm. 89 Pollen of Cyclolepis genistoides D. Don. A LM equatorial view in high focus showing the aperture. B Optical section showing the exine structure. (From Forcone 459, CORD). Scale bars: 12 μm. 90 Pollen of Duidaea marahuacensis Steyerm. SEM equatorial view (from Liesner 24644, US). Scale bar: 10 μm. 91 Pollen of Eriachaenium magellanicum Sch. Bip. A SEM equatorial view. B SEM polar view. C LM equatorial view in high focus. (From Sleumer 908, LP). Scale bars: A, B=10 μm; C=13 μm

672

L. Katinas, et al.

Figs. 92–94 92 Pollen of Gerbera hieracioides (Kunth) Zardini. A SEM polar view showing a slightly microechinate surface. B LM equatorial view showing the endoaperture with horns (arrows). (From Sagástegui 106, LP). Scale bars: A=10 μm; B=12 μm. 93 Pollen types I and II of Gochnatia Kunth. A–C Pollen type I. A SEM polar and equatorial view of G. barrosii Cabrera (from Mathes 3, LP). B, C LM equatorial view of G. polymorpha (Less.) Cabrera (from Glaziou 3039, LP). D, E Pollen type II. D SEM equatorial view of G. shaferi (Britton) Jervis & Alain (from Victorin & Clemente 22031, NY). E LM equatorial view in optical section of G. buchii (Urb.) Jiménez showing the exine structure (from Jiménez 3613, LP). Scale bars: 10 μm. 94 Pollen of Gongylolepis huachamacari Maguire. A SEM equatorial view. B LM optical section of equatorial view. (From Maguire et al. 30080, MO). Scale bars: 10 μm


673

Figs. 95–100 95 Pollen of Hecastocleis shockleyi A Gray. A SEM equatorial view. B LM equatorial view in high focus showing the colpate aperture. C LM polar view in optical section. (From Train 3973, LP). Scale bars: 10 μm 96 Pollen of Hyaloseris cinerea (Griseb.) Griseb. SEM polar view (from Ragonese 9335, LP). Scale bar: 10 μm. 97 Pollen of Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind. SEM equatorial view showing a very irregular arrangement of microspines, note the tectum surface very perforate (from Chebataroff 3015, LP). Scale bar: 10 μm. 98 Pollen of Lulia nervosa (Less.) Zardini. A LM equatorial view in optical section showing the exine structure. B LM polar view in high focus. (From Smith & Klein 10576, LP). Scale bars: 10 μm. 99 Pollen of Lycoseris trinervis (Ruiz & Pav. ex D. Don) S. F. Blake. SEM equatorial view (from Diers 1249, LM). Scale bar: 10 μm. 100 Pollen of Macroclinidium rigidulum (Miq.) Makino. A SEM equatorial view showing the short ectocolpus and the very perforate exine surface. B LM equatorial view in optical section. (From Deguchi et al. 6382, MO). Scale bars: 10 μm

674

L. Katinas, et al.

Figs. 101–106 101 Pollen of Mutisia kurtzii R. E. Fr. A SEM equatorial view. B SEM polar view. (From Budin s.n., LP). Scale bars: 10 μm. 102 Pollen of Myripnois dioica Bunge. SEM equatorial view. (From Sheean s.n., NY). Scale bar: 10 μm. 103 Pollen of Neblinaea promontoriorum Maguire & Wurdack. SEM equatorial view. (From Silva & Brazzao 60895, NY). Scale bar: 10 μm. 104 Pollen of Pachylaena atriplicifolia D. Don ex Hook. & Arn. SEM polar view. (From Frenguelli s.n., LP). Scale bar: 10 μm. 105 Pollen of Pertya cordifolia Mattf. SEM equatorial view (from Steward & Cheo 972, NY). Scale bar: 10 μm. 106 Pollen of Plazia daphnoides Wedd. SEM polar view (from Cabrera 9315, LP). Scale bar: 10 μm


675

Figs. 107–111 107 Pollen of Trichocline reptans (Wedd.) B. L. Rob. SEM equatorial view (From Cabrera et al. 24099, LP). Scale bar: 10 μm. 108 Pollen of Urmenetea atacamensis Phil. A SEM polar view. B LM equatorial view in optical section. (From Cabrera & Schwabe 80, LP). Scale bars: 10 μm. 109 Pollen of Ameghinoa patagonica Speg. A SEM equatorial view showing the very small microspines and the microgranulate membrane. B LM equatorial vew showing the exine estructure. (From Soriano 3208, LP). Scale bars: 10 μm. 110 Pollen of Cephalopappus sonchifolius Nees & Mart. SEM polar view (from Cabrera 12243, LP). Scale bar: 10 μm. 111 Pollen of Criscia stricta (Spreng.) Katinas. SEM equatorial view (from Crisci 545, LP). Scale bar: 10 μm

by Hind & Semir, 1998b: 1011, K not seen). Additional specimens examined: Brazil. State Minas Gerais: Mun. Gouveia, Parauna, G. Hatschbach 26992 (LP*); State Goias: Municipio Goias, Morro D. Francisco, A. Macedo 3527 (MO). 6. Wunderlichia senaeii Glaz. ex Maguire & G. M. Barroso, Revista Brasil. Biol. 33: 405. 1973. TYPE: Brazil. State Minas Gerais: 3–5 km E of Serra, along road from Conceiçao to Diamantina, B. Maguire et al. 49125 (lectotype, designated by Barroso & Maguire, 1973: 405, NY!). Literature. Barroso and Maguire (1973), Roque and Pirani (1997), Roque (2001b), Freire et al. (2002).

676

L. Katinas, et al.

Figs. 112–116 112 Pollen of Leucheria floribunda DC. A SEM equatorial view. B Detail of the exine structure. (From Grandjot 3353, GH). Scale bars: 10 μm. 113 Pollen of Lophopappus tarapacanus (Phil.) Cabrera. SEM equatorial view showing small microspines (from Ricardi & Marticorena 25604, LP). Scale bar: 10 μm. 114 Pollen of Perezia eryngioides (Cabrera) Crisci & Martic. SEM polar view (From Smith & Klein 7777, LP). Scale bar: 10 μm. 115 Pollen of Trixis verbascifolia (Gardner) S. F. Blake. SEM equatorial view (From Cabrera 12255, LP). Scale bar: 10 μm. 116 Pollen of Chimantaea huberi Steyerm. SEM equatorial view (from Huber 12034, LP). Scale bar: 10 μm


677

Figs. 117–118 117 Pollen of Quelchia bracteata Maguire, Steyerm. & Wurdack. SEM equatorial view (from Pipoly et al. 7217, US). Scale bar: 10 μm. 118 Pollen of Stenopadus. A SEM equatorial view in S. chimantensis Maguire, Steyerm. & Wurdack. B LM equatorial view in S. chimantensis Maguire, Steyerm. & Wurdack, note the polar caps (arrows). (From Pipoly et al. 7142, MO). C LM equatorial view in S. talaumifolius S. F. Blake (from Liesner 18346, MO). D SEM equatorial view in S. huachamacari Maguire (from Maguire et al. 30116, MO). E SEM equatorial view in S. connelli (N. E. Br.) S. F. Blake (from Liesner 23109, MO). Scale bars: A, B, D=10 μm; C=12 μm; E=9 μm

Excluded Taxa Barnadesioideae K. Bremer & R. K. Jansen Ann. Missouri Bot. Gard. 79: 415. 1992. Barnadesiinae Benth., in Benth. & Hook. f., Gen. pl. 2: 168. 1873. Herbs, shrubs, subshrubs or little trees, usually with axillary spines, unique indumentum of long, two-celled, barnadesioid hairs on vegetative and reproductive organs. Corollas villose, central florets tubular-funnelform with equal to unequal segments, marginal florets bilabiate to sub-ligulate; anther apical appendages obtuse, tails long to short; style bifid to bilobed, branches smooth to dorsally papillose above or above and below the branch bifurcation point; pappus generally of plumose bristles, the bristles constituted by barnadesioid hairs, scaly, or rarely absent. The Barnadesioideae comprise a small subfamily with 88 species in nine genera restricted to South America, ranging from the northern Andes in Venezuela south into Argentinian Patagonia. Observation. Barnadesioideae with the genera Arnaldoa, Barnadesia, Chuquiraga, Dasyphyllum, Doniophyton, Fulcaldea, Huarpea Cabrera, and Schlechtendalia,

678

L. Katinas, et al.

Fig. 119 Pollen of Stifftia chrysantha J. C. Mikan. a SEM equatorial view. b SEM polar view. c LM equatorial view in optical section showing the polar thickened exine. d LM polar view in optical section, note the thick margin of the apertures (arrow). (From Cabrera 12242, LP). Scale bars: 10 μm

were considered as a subtribe of Mutisieae by Cabrera (1961, 1977), following the ideas of Bentham. Later the genus Duseniella was added to this group of genera (see Duseniella below). Jansen and Palmer (1987) reported the presence of a 22 kb cpDNA inversion in several Asteraceae. The inversion was found to be absent in three genera of the, by that time, Mutisieae-Barnadesiinae and in other families close to Asteraceae suggesting a basal dichotomy between the Barnadesiinae and the rest of the family. The result was later confirmed by chloroplast DNA sequence data (Kim et al., 1992) and its taxonomic rank was subsequently elevated to subfamily by Bremer and Jansen (1992). From a morphological point of view, the members of the Barnadesioideae are characterized by unique, axillary spines (excepting Duseniella, Huarpea, Schlechtendalia, and some species of Chuquiraga and Dasyphyllum), and by a unique indumentum in vegetative and reproductive organs, the barnadesioid hairs (Bremer & Jansen, 1992; Katinas & Stuessy, 1997). In addition, villose corollas are very


679

Figs. 120–121 120 Pollen of Stomatochaeta condensata (Baker) Maguire & Wurdack. SEM equatorial view (from Liesner 19415, MO). Scale bar: 10 μm. 121 Pollen of Wunderlichia mirabilis Riedel ex Baker. LM equatorial view in optical section showing the exine structure (from Hatschbach 26992, LP). Scale bar: 12 μm

common in the subfamily. The pollen of the Barnadesioideae differs from that of Mutisioideae because the exine is spongy, granulate, granulate–columellate or scarcely columellate and in some genera it is caveate (Urtubey & Tellería, 1998; Zao et al., 2000). Our results show that most members of Mutisioideae (tribes Mutisieae and Stifftieae) share the same type of style with those of Barnadesioideae, providing evidence of a strong morphological link between both subfamilies.

Brachylaena R. BR. Trans. Linn. Soc. London 12: 115. 1817. TYPE: Brachylaena neriifolia (L.) R. Br., based on Baccharis neriifolia L. Oligocarpha Cass., Bull. Sci. Soc. Philom. Paris: 151. 1817. TYPE: Oligocarpha nereifolia Cass., Dict. sci. nat. 34: 22. 1823 [= Brachylaena neriifolia (L.) R. Br.]. Trees or shrubs dioecious. Leaves alternate; petiolate to subsessile; blades elliptic to obovate, margin entire to dentate-spinose, glabrous to subglabrous above, tomentose beneath. Capitulescences racemose to paniculate; capitula unisexual, subsessile to pedunculate; receptacle epaleate; involucre multiseriate, outermost phyllaries often extending into the stalk. Florets in male capitula with corollas tubular sub-campanulate, deeply five-lobed, lobes coiled; anther apical appendages apiculate, tails smooth; style bifid, widened below the branch bifurcation point, branches appressed, collector hairs above and somewhat below the bifurcation point,

680

L. Katinas, et al.

the hairs long and acute; florets in female capitula with corollas tubular-filiform, shallowly five-lobed, lobes erect; anthers absent or reduced to staminodes; style not widened, branches smooth. Cypselae pubescent; pappus of barbellate bristles, widened and plumose at the apex. Habitat and Distribution. Genus of ca. 15 species of Africa and Madagascar. Observation. Brachylaena and its sister genus Tarchonanthus have been variously placed in the tribes Anthemideae, Astereae, Inuleae, Mutisieae, Vernonieae, and Tarchonantheae. The molecular study of Keeley and Jansen (1991) indicated that the two genera form a distinct clade at or near the base of Cichorioideae. Recent molecular studies indicate that Brachylaena and Tarchonanthus are sister to the African taxa Dicoma and Pasaccardoa (Kim et al., 2002) or sister to Cardueae and Oldenburgia (Funk et al., 2005), and thus are included in the Carduoideae. Our study shows that the widened styles covered by acute hairs is another character that supports the exclusion of Brachylaena from the Mutisioideae. Literature. Paiva (1972), Pope (1992), Cilliers (1993), Kimball et al. (2004).

Cloiselia S. Moore J. Bot. 44: 148. 1906. TYPE: Cloiselia carbonaria S. Moore. Shrubs or little trees. Leaves alternate to fasciculate; short-pseudopetiolate; blades oblanceolate to spathulate. Capitulescences monocephalous or in cymes, terminal; capitula short-petiolate, homogamous, discoid; receptacle epaleate, alveolate; involucre multiseriate. Florets bisexual, zygomorphic, transitional between actinomorphic and ligulate, often villous; anther apical appendages apiculate to caudate, tails laciniate, long, acute, tails connate; style bilobed, branches appressed, dorsally pubescent reaching the bifurcation point, hairs acute. Cypselae villose; pappus of scabrid, capillary bristles. Habitat and Distribution. Genus of four species endemic to Madagascar. Observation. A cladistic analysis of the genus Dicoma based on morphological data (Ortiz, 2000) showed that some of its species constitute a well-supported monophyletic group, segregated from the remaining species of Dicoma. This group of species constitute the genus Cloiselia (Ortiz, 2006a), characterized by corollas much longer than the involucre, disc floret epidermal cell cuticle ornamentation “intestine-like,” long simple (not twin) hairs on disc florets corolla, stamen insertion near the corolla base, and pollen echinate, among other characters. According to the cladistic analysis (Ortiz, 2000), Cloiselia is sister to a monophyletic group comprising the genera Macledium, Pasaccardoa and Dicoma s. str. An interesting aspect of Cloiselia is the presence of some characters that appear to be archaic within Asteraceae, as they are present in Barnadesioideae, such as zygomorphic corollas, marginal veins of the adjacent lobes separate from the corolla base, long eglandular


681

hairs in corollas (but not barnadesioid hairs) (Ortiz, 2006a). However, molecular phylogenetic analyses (Kim et al., 2002) do not support a close relationship between Cloiselia and the Barnadesioideae. Literature. Humbert (1963), Ortiz (2000, 2001, 2006a).

Dicoma Cass. Bull. Sci. Soc. Philom. Paris: 12. 1817. TYPE: Dicoma tomentosa Cass. Hochstetteria DC., Prodr. 7: 287. 1838. TYPE: Hochstetteria schimperi DC. [= Dicoma schimperi (DC.) Baill. ex O. Hoffm.]. Herbs annual or perennial, shrubs, or rarely small trees. Leaves alternate; sessile to subsessile, sometimes pseudopetiolate or clasping; blades usually ovate, margin entire to serrulate, pubescent in both surfaces. Capitulescences monocephalous or in cymes; capitula sessile to pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets bisexual, corolla actinomorphic, deeply five-lobed, tube narrow, gradually dilated into a limb to subcampanulate, lobes coiled; anther apical appendages acute to acuminate, tails papillose; style bifid, branches appressed, apically pubescent and also hairs in a subapical ring, hairs acute. Cypselae villose; pappus usually of scabrid bristles. Habitat and Distribution. Genus of ca. 35 species most from central and southern Africa and Madagascar, two species from Asia (Arabian Peninsula, India, and Pakistan). Observation. Dicoma and its related genera Cloiselia, Erythrocephalum, Gladiopappus, Macledium, Pasaccardoa, Pleiotaxis, and the unpublished genus Dicomopsis (Ortiz, 2006b) have been included in the tribe Mutisieae (Jeffrey, 1967; Cabrera, 1977; Grau, 1980; Karis et al., 1992; Bremer, 1994; Ortiz, 2000, 2006a). Bremer (1994) called this complex, with the exception of the recently resurrected genera Cloiselia and Macledium, and Dicomopsis, the Dicoma group. The genera of the Dicoma group have been characterized by the presence of mostly actinomorphic corollas with narrow tubes and wide limbs, acuminate or apiculate anther appendages, papillose style branches with a ring of subapical hairs, and echinate pollen in some genera (Cloiselia, Erythrocephalum, Pleiotaxis). Hansen (1991a) suggested, based on a set of features in common (e.g., petal epidermis pattern, style branches with a tuft of hairs, corollas campanulate, cypselae turbinate–oblong) that Dicoma and relatives are more related to the tribe Cardueae or Arctoteae than to Mutisieae. The phylogenetic relationships between the Dicoma group and Cardueae have been supported by the molecular studies of Kim et al. (2002) and Funk et al. (2005). Based on molecular data, the genera of the Dicoma group have been segregated in the new tribe Dicomeae (Panero & Funk, 2002). According to Panero and Funk (2002), the Dicomeae is the sister group of the tribes Cardueae and Tarchonantheae, and the three tribes together constitute the subfamily Carduoideae. Further, Funk et al. (2005) included also Oldenburgia in this monophyletic group. Our observations of the styles of

682

L. Katinas, et al.

specimens of Dicoma show similarities to the carduoid style of the Cardueae, supporting the exclusion of this genus from Mutisioideae. Literature. Wilson (1923), Jeffrey (1967), Pope (1991, 1992), Ortiz et al. (1998), Ortiz (2000), Ortiz and Pulgar (2002), Ortiz and Netnou (2005).

Duseniella K. Schum. Just’s Bot. Jahresber. 28: 475. 1902. TYPE: Duseniella patagonica (O. Hoffm.) K. Schum., based on Dusenia patagonica O. Hoffm. Dusenia O. Hoffm., Wiss. Erg. Schwed. Exp. Magellansl. 3: 246. 1900, non C. A. Mull., 1897, nom. illeg. TYPE: Dusenia patagonica O. Hoffm. [= Duseniella patagonica (O. Hoffm.) K. Schum.]. Herbs dwarf, unarmed, annual, with barnadesioid hairs. Leaves apically alternate and basally opposite; sessile; blades linear-oblong, entire, pubescent to glabrous. Capitulescences monocephalous, terminal; capitula sessile, heterogamous, discoid; receptacle epaleate; involucre multiseriate. Florets with corolla actinomorphic, tubular-funnelform, five-lobed, lobes erect; marginal florets female; central florets bisexual; anther apical appendages acute, tails smooth; style bifid, branches dorsally papillose above and below the bifurcation point. Cypselae truncate at the apex, villose; pappus of ten scabrid, lanceolate scales. Habitat and Distribution. Monotypic genus endemic to Patagonia in Argentina. Observation. Although Duseniella had been already referred to the subtribe Barnadesiinae by Jeffrey (1967), Cabrera (1977) placed it in the Mutisieae– Gochnatiinae, probably because some of its characters deviate from those typical of Barnadesiinae, e.g., unarmed branches, non villose florets, and scaly pappus. Morphology-based cladistic studies (Bremer, 1994; Stuessy et al., 1996) supported relationships between Duseniella and Doniophyton, a member of the subfamily Barnadesioideae, based on similar habit (short-lived herbs); basally opposite, linear leaves; female marginal florets with tubular corolla; and collapsed testa epidermis. In addition, we observed barnadesioid hairs in Duseniella, which are exclusive to the Barnadesioideae. Literature. Cabrera (1971b).

Erythrocephalum Benth. in Benth. & Hook. f., Gen. pl. 2: 488. 1873. TYPE: Erythrocephalum zambesianum Oliv. & Hiern, Fl. Trop. Afr. 3: 441. 1877. Achyrothalamus O. Hoffm., Bot. Jahrb. Syst. 15: 541. 1893. TYPE: Achyrothalamus marginatus O. Hoffm. [= Erythrocephalum marginatum (O. Hoffm.) S. Ortiz & A. P. Cout.].


683

Herbs annual or subshrubs. Leaves alternate, often rosulate; sessile, often auriculate at the base; blades linear to elliptic, margin entire to denticulate, usually tomentose on the lower surface. Capitulescences monocephalous, terminal; capitula pedunculate, homogamous, radiate or discoid; receptacle paleate; involucre multiseriate. Florets bisexual, radiate capitula with marginal corollas bilabiate, outer lip expanded; central corollas actinomorphic, tubular-campanulate, deeply five-lobed, tube narrow, abruptly dilated into a limb, lobes erect; discoid capitula with florets actinomorphic, corollas tubular-campanulate; anther apical appendages acuminate to apiculate, tails papillose, long, subrounded; style bifid, branches separate, dorsally pubescent and also hairs in subapical ring, hairs acute. Cypselae glabrous to sparcely villose; pappus of scabrid, linear scales, or absent. Habitat and Distribution. Genus of ca. 13 species most from eastern Tropical Africa. Observation. The genus Erythrocephalum was included in the Dicoma group (Bremer, 1994) (see the Observation in Dicoma). The exine with an Helianthoid pattern in the pollen of Erythrocephalum is completely absent in Mutisioideae (Ortíz & Coutinho, 2001; our own observations). Also, we observed that the style of Erythrocephalum has similarities to the carduoid style of the Cardueae, supporting the exclusion of this genus from Mutisioideae. Literature. Jeffrey (1967), Pope (1992), Ortiz (2000), Ortiz and Coutinho (2001).

Gladiopappus Humbert Bull. Soc. Bot. France 95: 181. 1948. TYPE: Gladiopappus vernonioides Humbert. Shrubs. Leaves alternate; petiolate; blades oblanceolate to obovate, entire. Capitulescences monocephalous, terminal to axillar; capitula sessile, homogamous, radiate; receptacle fimbriate; involucre multiseriate. Florets dimorphic, bisexual, marginal florets with corollas bilabiate, outer lip expanded; central florets with corollas actinomorphic, tubular-campanulate, deeply five-lobed, tube narrow, abruptly dilated into a limb, lobes coiled; anther apical appendages apiculate, tails papillose; styles bilobed, branches appressed, dorsally sparcely papillose, papillae reaching the bifurcation point. Cypselae papillose; pappus dimorphic, outer row of setae, inner row of scabrid scales and longer. Habitat and Distribution. Monotypic genus endemic to Madagascar. Observation. The genus Gladiopappus was included in the Dicoma group by Bremer (1994). It shares the typical corolla abruptly dilated into a tube with the genera of this group, but, according to the literature (Humbert, 1948, 1963), the style is sparsely papillose and lacks the typical subapical tuft of collector hairs (see the Observation in Dicoma), a feature that would approach Gladiopappus to Mutisieae. Until we can analyze the morphology of this rare genus, we maintain Gladiopappus excluded from Mutisioideae.

684

L. Katinas, et al.

Literature. Humbert (1948, 1963).

Hesperomannia A. Gray Proc. Amer. Acad. Arts 6: 554. 1864–65. TYPE: Hesperomannia arborescens A. Gray. Trees to 5 m tall. Leaves alternate; sessile to petiolate; blades linear-elliptic to ovate, entire, glabrous to pubescent. Capitulescences monocephalous or clusters of two to ten capitula; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual; corolla actinomorphic, tubularfunnelform, deeply five-lobed; anther apical appendages acute, tails smooth; style bifid, branches with collector hairs above and much below the bifurcation point, acute hairs. Cypselae trigonous, truncate, glabrescent; pappus of rigid, scabrid bristles, plumose to scabrid at the apex. Habitat and Distribution. Genus of three species endemic to the Hawaiian Islands. Observation. Hesperomannia has longly been placed in the tribe Mutisieae. Phylogenetic comparisons within the Vernonieae using sequences of both ndhF and the internal transcribed spacer regions of nuclear ribosomal DNA (Kim et al., 1998) revealed that Hesperomannia belongs to the tribe Vernonieae and is sister to African species of Vernonia Schreb. In addition, we observed vernonioid styles, although the branches are short. With respect to pollen morphology, Hesperomannia cannot be included in the Mutisioideae. The exine section in LM micrographs (Selling, 1947; Marticorena & Parra, 1975; our own observations) is similar to the modified Anthemoid pattern of pollen of Vernonieae and Liabeae (Skvarla et al., 1977). This pattern is characterized by having much broader columellae than noted for the Anthemoid pattern present in Mutisioideae (Skvarla et al., 1977; see also Wortley et al., 2007). Literature. Degener (1946), Carlquist (1957a), Wagner et al. (1990).

Macledium Cass. Dict. sci. nat. 34: 39. 1825. [= Dicoma subgen. Macledium (Cass.) Less., Syn. gen. Compos.: 169. 1832]. TYPE: Macledium burmanni Cass., nom. illeg. Cullumiopsis Drake, Bull. Mus. Hist. Nat. (Paris) 5: 103. 1899. TYPE: Cullumiopsis grandidieri Drake [= Macledium grandidieri (Drake) S. Ortiz]. Nitelium Cass., Dict. sci. nat. 35: 11. 1825. TYPE: Nitelium rubescens Cass. [= Macledium spinosum (L.) S. Ortiz]. Herbs or shrubs. Leaves alternate, sometimes rosulate; sessile to subsessile, subtending; blades ovate to oblanceolate, margin serrulate to entire, tomentose especially beneath. Capitulescences monocephalous, cymes terminal to axillar, or corymbose to paniculate; capitula subsessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corollas actinomorphic, tubular-campanulate, deeply five-lobed, tube narrow, gradually dilated into a limb to sub-campanulate, lobes erect; anther apical appendages acuminate,


685

tails papillose; style bifid, branches appressed, dorsally pubescent and also with a basal ring of slightly longer hairs, almost reaching the bifurcation point, hairs acute. Cypselae pubescent; pappus of plumose, capillary bristles, often scales. Habitat and distribution. Genus of 20 species of Central and South Africa. Observation. The genus Macledium has been lately resurrected (Ortiz, 2001) to include some distinct species that constitute a monophyletic group within the genus Dicoma (Ortiz, 2000). Macledium is different from Dicoma by its phyllaries with inconspicuous midrib, central corollas with straight lobes, and dorsally pubescent style branches with a basal ring of short hairs reaching the bifurcation point (Ortiz, 2001) (see the Observation in Dicoma). Our observations confirm that the styles resemble to the carduoid style of the Cardueae, supporting the exclusion of this genus from Mutisioideae. Literature. Ortiz (2000, 2001).

Moquinia DC., Nom. Cons. Prodr. 7: 22. 1838, non Spreng. 1828, emend. Gamerro, Darwiniana 30: 132. 1990. TYPE: Moquinia racemosa (Spreng.) DC., based on Conyza racemosa Spreng. Spadonia Less., Syn. gen. Compos.: 99. 1832. TYPE: Spadonia racemosa (Spreng.) Less., based on Conyza racemosa Spreng. [= Moquinia racemosa (Spreng.) DC.]. Trees or shrubs monoecious or gynodioecious. Leaves alternate; petiolate; blades obovate, cuneate, entire, glabrescent to tomentose beneath. Capitulescences racemose, corymbose, spicate to paniculate; capitula unisexual or bisexual, sessile to subsessile, discoid; receptacle epaleate; involucre multiseriate. Florets actinomorphic, tubular sub-campanulate, deeply five-lobed, lobes coiled to erect. Bisexual capitula: anthers with acute apical appendages, tails short, smooth, style bifid, widened below the branch bifurcation point, branches with collector hairs above and below the bifurcation point; female capitula: florets without anthers or with staminodes; male capitula: florets with rudimentary ovaries. Cypselae villose; pappus of long and short, somewhat paleaceous, scabrid bristles. Habitat and distribution. Genus of two species endemic to the states of Bahia and Minas Gerais in Brazil. Observation. Moquinia was included in the synonymy of Gochnatia by Cabrera (1950) and further separated from this genus by the same author (Cabrera, 1969). Moquinia remained in the tribe Mutisieae until Gamerro (1990) performed a morphological and palynological study of the genus. In his work, the genus Pseudostifftia H. Rob. of the tribe Vernonieae (Robinson, 1979) was considered a synonym of Moquinia, and Moquinia was transferred to the Vernonieae. The main characters to justify the transference are: the short apical anther appendage; the glandular hairs in corolla, leaves, and fruits; the pollen morphology; the welldeveloped stylopodium; the widened style; and the endothecium characters

686

L. Katinas, et al.

(Gamerro, 1990). Robinson (1994) separated the genera Pseudostifftia and Moquinia and placed them in their own tribe, the Moquinieae, because of differences in the style and pollen characters with the Vernonieae (see also Wortley et al., 2007). Literature. Cabrera (1969), Gamerro (1990). Pasaccardoa Kuntze Revis. gen. pl. 1: 354. 1891. TYPE: Pasaccardoa grantii Kuntze. Phyllactinia Benth., in Benth. & Hook. f., Gen. pl. 2: 488. 1873, non Lév., 1851, nom. illeg. TYPE: Phyllactinia grantii Benth. Herbs or subshrubs. Leaves alternate; sessile to subsessile, subtending; blades elliptic to oblanceolate, usually denticulate, gland-dotted, pubescent in the lower surface, sometimes discolor. Capitulescences monocephalous, terminal to axillar; capitula subsessile, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Marginal florets neuter, corollas true ray, limb three-lobed; central florets bisexual, corollas actinomorphic, tubular-campanulate, deeply five-lobed, tube narrow, abruptly dilated into a limb, lobes usually coiled; anther apical appendages acuminate, tails papillose, long, acute; style bifid, branches appressed, dorsally pubescent and also hairs in subapical ring, hairs acute. Cypselae dimorphic; marginal cypselae rudimentary, rostrate at the apex, glabrous or pubescent; central cypselae ribbed, very hairy at the base or pubescent throughout; pappus of numerous, persistent scales. Habitat and Distribution. Genus of four species of Tropical Asia. Observation. The genus Pasaccardoa was included in the Dicoma group (Bremer, 1994) (see the Observation in Dicoma). Our observations of the styles of specimens of Pasaccardoa show similarities to the carduoid style of the Cardueae, supporting the exclusion of this genus from Mutisioideae. Literature. Jeffrey (1967), Pope (1992), Ortiz (2000).

Pleiotaxis Steetz in Peters, Naturw. Reise Mossambique 6: 499. 1864. TYPE: Pleiotaxis pulcherrima Steetz. Herbs or subshrubs. Leaves alternate to sometimes the basal rosulate; sessile to subsessile, often basally auriculate; blades linear to elliptic, margin denticulate, glabrous to pubescent. Capitulescences monocephalous or in cymes; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets bisexual, corolla actinomorphic, tubular-campanulate, deeply five-lobed, tube narrow, abruptly dilated into a limb, lobes coiled, sometimes patent; anther apical appendages apiculate-bulbous, tails papillose, long, subrounded; style bilobed to bifid, branches separate, pubescent at the apex and also hairs in a subapical ring, hairs acute. Cypselae glabrous to pubescent; pappus of scabrid, capillary bristles.


687

Habitat and Distribution. Genus of ca. 26 species of Tropical Africa. Observation. The genus Pleiotaxis was included in the Dicoma group (Bremer, 1994) (see the Observation in Dicoma). Our observations of the styles of specimens of Pleiotaxis show similarities to the carduoid style of the Cardueae, supporting the exclusion of this genus from Mutisioideae. In addition, conspicuous, well-developed spines, different from those of Mutisioideae, were observed in the pollen of species of Pleiotaxis. Literature. Moore (1925), Jeffrey (1967), Pope (1992), Ortiz and Rodríguez-Oubiña (1998), Ortiz (2000), Freire et al. (2002).

Tarchonanthus L. Sp. pl.: 842. 1753. TYPE: Tarchonanthus camphoratus L. Trees or shrubs dioecious, often aromatic. Leaves alternate; short-petiolate to subsessile; blades ovate, elliptic to obovate, margin entire to lobed, usually tomentose beneath. Capitulescences paniculate, terminal; capitula unisexual, pedunculate, discoid; receptacle epaleate, glabrous or with silky hairs; involucre one- to two-seriate, phyllaries free to connate. Florets actinomorphic, tubularfunnelform, villose. Male capitula with actinomorphic, five-lobed corollas, anthers exserted, apical appendages acute, short, tails smooth; style bifid, branches dorsally papillose above and below the bifurcation point; female capitula with corollas actinomorphic, shorter than the male corollas, four- to five-lobed; anthers absent or reduced to staminodes; style bilobed, widened below the branch bifurcation point, branches flat, smooth. Cypselae flattened, pubescent; pappus absent. Habitat and Distribution. Genus of five species ranging from Saudi Arabia through East Africa to the Cape Province of South Africa. Observation. Tarchonanthus together with Brachylaena are currently placed in the tribe Tarchonantheae (Keeley & Jansen, 1991; Kim et al., 2002; Funk et al., 2005) (see Observation in Brachylaena). The pollen morphology of Tarchonanthus positions this genus close to Anthemideae, although more studies are needed (Skvarla et al., 1977). Literature. Paiva (1972), Beentje (1998), Herman (2002). Warionia Benth. & Coss. Bull. Soc. Bot. France 19: 165. 1872. TYPE: Warionia saharae Benth. & Coss. Shrubs aromatic, with latex. Leaves alternate; sessile; blades oblong to obovate, pinnately partite, margin sinuate to undulate, fleshy, glabrous. Capitulescences monocephalous or two- to five-headed, corymbose to paniculate; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular, deeply five-lobed, lobes coiled; anther apical appendages acute, tails smooth; style bifid, branches with collector

688

L. Katinas, et al.

hairs above and somewhat below the branch bifurcation point, hairs acute. Cypselae truncate at the apex, villose; pappus of paleaceous-setose, barbellate bristles. Habitat and Distribution. Monotypic genus, endemic to the African Sahara desert. Observation. Warionia was regarded as a member of the Mutisieae by Cabrera (1977) and excluded from the tribe by Hansen (1991a). In the morphology-based cladogram of Karis et al. (1992), Warionia appears as an independent branch between the Cardueae and the Vernonioid complex. Bremer (1994) placed it among the genera unassigned to a tribe in Cichorioideae. In a recent study of the tribe Arctoteae (Funk et al., 2004), Warionia is sister to the genus Gundelia L., and both are members of the tribe Gundelieae at the base of the Lactuceae. The style and pollen morphology of Warionia differs from that of the members of Mutisioideae (see also Wortley et al., 2007). Literature. Ozenda (1991), Audissou (1999), Katinas et al. (in press). Acknowledgments We are grateful to Jorge Crisci and Bob Kowal for critical reading of the manuscript, and to Peter Hoch and Vicki Funk for their assistance and constant support. Jorge Crisci also provided some unpublished SEM photographs of Nassauvieae. Anonymous reviewers are also acknowledged for their helpful comments on an earlier version of this paper. Thanks are given to Susana Freire for helping with the Ainsliaea group, to Santiago Ortiz for comments and material on the African taxa, to Nicholas Hind for helping with material at Kew, to Melica Muñoz Schick for helping with literature and specimens of Chilean taxa, and to Dan Nicolson and Daniel Giuliano for nomenclatural advice. We also thank the curators of hebaria for loans of specimens, all the persons and institutions that granted permission to redraw their illustrations, and Hugo Calvetti for the designing of the SEM plates. The John Simon Guggenheim Memorial Foundation (LK), the Latin American, Caribbean, and Iberian Studies (LACIS) program (LK), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina, Agencia de Promoción Científica y Tecnológica, Argentina, the Botany Department of the University of Wisconsin-Madison, and the Missouri Botanical Garden are also acknowledged for their generous support.

Literature Cited Anderberg, A. A. & S. E. Freire. 1990. Luciliopsis perpusilla Weddell is a species of Chaetanthera Ruiz & Pavon (Asteraceae, Mutisieae). Taxon 39: 430–432. Anderson, C. 1972. A monograph of the Mexican and Central American species of Trixis (Compositae). Memoirs of the New York Botanical Garden 22: 1–68. Aristeguieta, L. 1964. Compositae. Pp. 485–941. In: T. Lasser (ed.), Flora de Venezuela 10. Instituto Botánico, Dirección de Recursos Naturales Renovables, Ministerio de Agricultura, Caracas, Venezuela. ———. 1967. Compositae. Pp. 345–370. In: J. A. Steyermark (ed.), Flora of the Auyan-tepui. Acta Botánica Venezuelica 2. Instituto Botánico, Dirección de Recursos Naturales Renovables, Ministerio de Agricultura, Caracas, Venezuela. Ariza Espinar, L. 1973. Revisión del género Hyaloseris (Compositae). Kurtziana 7: 195–211. Arroyo, M. T. K., A. M. R. Davies & I. Till-Bottraud. 2004. Chaetanthera acheno-hirsuta (Tombesi) Arroyo, A. M. R. Davies & Till-Bottraud elevated to species, new for the flora of Chile. Gayana, Botánica 61: 27–31. Audissou, J.-A. 1999. Warionia saharae. British Cactus and Succulent Journal 3: 124–126. Baagøe, J. 1977. Microcharacters in the ligules of the Compositae. Pp. 119–139. In: V. H. Heywood, J. B. Harborne & B. L. Turner (eds.), The Biology and Chemistry of the Compositae, Vol. 1. Academic, London. Bacigalupi, R. 1931. A monograph of the genus Perezia, section Acourtia, with a provisional key to the section Euperezia. Contributions of the Gray Herbarium of Harvard University 97: 1–81. Badillo, V. M. 1988. Sobre la vigencia del género Cardonaea Aristeg. et al. (Compositae–Mutisieae). Ernstia 46: 8–10.


689

Baker, J. G. 1884. Compositae III. Mutisiaceae. Pp. 339–398. In: C. F. P. Martius (ed.), Flora Brasiliensis 6(3). Monachii in typographia regia C. Wolf et fil. et in offic. lithograph. Keller, Munich. Barreda, V. D. 1993. Late Oligocene?—Miocene pollen of the families Compositae, Malvaceae, and Polygonaceae from the Chenque Formation, Golfo San Jorge Basin, southeastern Argentina. Palynology 17: 169–186. ———, V. Barreda, L. Palazzesi & M. C. Tellería. 2008. Fossil pollen grains of Asteraceae from the Miocene of Patagonia: Nassauviinae affinity. Review of Palaeobotany and Palynology 151: 51–58. Barrie, F. R. 2006. Report of the General Committee: 9. Taxon 55: 795–800. Barroso, G. M. & B. Maguire. 1973. A review of the genus Wunderlichia (Mutisieae, Compositae). Revista Brasileira de Biologia 33: 379–406. Beentje, H. J. 1998. The genus Tarchonanthus (Compositae–Mutisieae). Kew Bulletin 54: 81–95. Bentham, G. & J. D. Hooker. 1873. Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definite, volume 2. Lovell, Reeve and Co., London. Bittmann, M. 1990a. Die Gattung Adenocaulon (Compositae): I. Morphologie. Candollea 45: 389–420. ———. 1990b. Die Gattung Adenocaulon (Compositae): II. Ökologie, Verbreitung und Systematik. Candollea 45: 493–518. Bolick, M. R. 1978. Taxonomic, evolutionary, and functional considerations of Compositae pollen ultrastructure and sculpture. Plant Systematics and Evolution 130: 209–218. Bond, P. 1987. A revision of Oldenburgia. South African Journal of Botany 53: 493–500. Bremer, K. 1987. Tribal interrelationships of the Asteraceae. Cladistics 3: 210–253. ———. 1992. Ancestral areas: A cladistic reinterpretation of the center of origin concept. Systematic Biology 41: 436–445. ———. 1993. Intercontinental relationships of African and South American Asteraceae: A cladistic biogeographic analysis. Pp. 104–135. In: P. Goldblatt (ed.), Biological Relationships Between Africa and South America. Yale University Press, New Haven. ———. 1994. Asteraceae: Cladistics and Classification. Timber, Portland, Oregon. ———. 1996. Major clades and grades of the Asteraceae. Pp. 1–7. In: D. J. N. Hind & H. J. Beentje (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994, Vol. 1. Royal Botanic Gardens, Kew. ——— & M. H. G. Gustafsson. 1997. East Gondwana ancestry of the sunflower alliance of families. Proceedings of the National Academy of Sciences of the United States of America 94: 9188–9190. ——— & R. K. Jansen. 1992. A new subfamily of the Asteraceae. Annals of the Missouri Botanical Garden 79: 414–415. Brummitt, R. K. 2005. Report of the Committee for Spermatophyta: 57. Taxon 54: 1093–1103. Burkart, A. 1944. Estudio del género de Compuestas Chaptalia con especial referencia a las especies argentinas. Darwiniana 6: 505–594. Cabrera, A. L. 1936. Las especies argentinas y uruguayas del género Trixis. Revista del Museo de La Plata, Sección Botánica 2: 31–86. ———. 1937. Revisión del género Chaetanthera (Compositae). Revista del Museo de La Plata, Sección Botánica 2: 87–210. ———. 1944. Vernonieas argentinas (Compositae). Darwiniana 6: 265–379. ———. 1950. Observaciones sobre los géneros Gochnatia y Moquinia. Notas del Museo de La Plata, Botánica 15: 37–48. ———. 1951. Notas sobre compuestas de América Austral. Darwiniana 9: 363–386. ———. 1953a. Compuestas peruanas nuevas o críticas. Boletín de la Sociedad Argentina de Botánica 5: 37–50. ———. 1953b. Las especies del género Pamphalea (Compositae). Notas del Museo de La Plata, Botánica 16: 225–237. ———. 1954. Compuestas sudamericanas nuevas o críticas. II. Notas del Museo de La Plata, Botánica 17: 71–80. ———. 1959a. Compositae catarinenses novae. Boletín de la Sociedad Argentina de Botánica 7: 187–200. ———. 1959b. Revisión del género Dasyphyllum (Compositae). Revista del Museo de La Plata, Sección Botánica 9: 21–100. ———. 1961. Compuestas argentinas. Clave para la determinación de los géneros. Revista del Museo Argentino de Ciencias Naturales, “Bernardino Rivadavia” Instituto Nacional de Investigación de las Ciencias Naturales, Botánica 2: 291–362. ———. 1965. Revisión del género Mutisia (Compositae). Opera Lilloana 13: 5–227. ———. 1968. Rehabilitación del género Holocheilus Cassini (Compositae). Revista del Museo de La Plata, Sección Botánica 11: 1–15. ———. 1969. El género Moquinia (Compositae). Boletín de la Sociedad Argentina de Botánica 11: 255–261.

690

L. Katinas, et al.

———. 1970. Actinoseris, nuevo género de compuestas. Boletín de la Sociedad Argentina de Botánica 13: 45–52. ———. 1971a. Revisión del género Gochnatia (Compositae). Revista del Museo de La Plata, Sección Botánica 12: 1–160. ———. 1971b. Compositae. Pp. 1–451. In: M. N. Correa (ed.), Flora Patagónica. Colección Científica del INTA, Buenos Aires, Argentina. ———. 1974. Tres Compositae nuevas de Minas Gerais (Brasil). Boletim do Museu Botanico Municipal 15: 1–4. ———. 1975. Una nueva especie de Onoseris (Compositae) de Brasil. Boletim do Museu Botanico Municipal 19: 1–3. ———. 1976. Una nueva especie de Mutisia (Compositae) de Patagonia. Darwiniana 20: 210–212. ———. 1977. Mutisieae—Systematic review. Pp. 1039–1066. In: V. H. Heywood, J. B. Harborne & B. L. Turner (eds.), The Biology and Chemistry of the Compositae, Vol. 2. Academic, London. ———. 1978. Compositae. Pp. 9–726. In: A. L. Cabrera (ed.), Flora de la Provincia de Jujuy, República Argentina. Colección Científica del INTA, Buenos Aires, Argentina. ———. 1982. Revisión del género Nassauvia (Compositae). Darwiniana 24: 283–379. ——— & A. Willink. 1973. Biogeografía de América Latina. O.E.A. Serie de Biología, Monografía 13, Washington, DC. Cabrera-Rodríguez, L. 1992. Systematic Study of the Genus Rzedowskiela R. Cabrera (Mutisieae, Compositae). Doctoral Thesis, The University of Texas, Austin. ———. 2001. Six new species of Acourtia (Asteraceae) and a historical account of Acourtia mexicana. Brittonia 53: 416–429. ——— & G. Dieringer. 2003. Pollen structure of Acourtia (Asteraceae): New findings and some taxonomic considerations. International Journal of Plant Sciences 164: 287–294. Candia, R. & A. D. Dalmasso. 1995. Dieta del guanaco (Lama guanicoe) y productividad del pastizal en la Reserva La Payunia, Mendoza (Argentina). Multequina 4: 5–15. Carlquist, S. 1957a. Systematic anatomy of Hesperomannia. Pacific Science 11: 207–215. ———. 1957b. Anatomy of Guayana Mutisieae. Memoirs of the New York Botanical Garden 9: 441–475. ———. 1957c. Wood anatomy of Mutisieae (Compositae). Tropical Woods 106: 29–45. ———. 1958. Anatomy of the Guayana Mutisieae. Part II. Memoirs of the New York Botanical Garden 10: 157–184. ———. 1961. Comparative Plant Anatomy. Holt, Rinehart & Winston, New York. ———. 1976. Tribal interrelationships and phylogeny of the Asteraceae. Aliso 8: 465–492. Cassini, H. 1816. Quatriéme memoire sur le famille des synantherées, contenant l’analyse de l’ovaire et de ses accesoires. Journal de Physique, de Chimie, d’Histoire Naturelle et des Arts 85: 5–21. ———. 1819. Suit du sixiéme mémoire sur la familie des Synanthérées, contenat les caractères des tribus. Journal de Physique, de Chimie, d’Histoire Naturelle et des Arts 88: 189–204. ———. 1824. Mutisiées. Pp. 462–465. In: G. Cuvier (ed.), Dictionnaire des Sciences Naturelles, Vol. 33. Le Normant, Paris. ———. 1825. Nassauviées. Pp. 204–208. In: G. Cuvier (ed.), Dictionnaire des Sciences Naturelles, Vol. 34. Le Normant, Paris. Cilliers, S. S. 1993. Synopsis of the genus Brachylaena (Asteraceae) in southern Africa. Bothalia 23: 175–184. Crisci, J. V. 1971. Sobre una especie de Jungia (Compositae) del Perú. Boletín de la Sociedad Argentina de Botánica 13: 341–346. ———. 1974a. A numerical–taxonomic study of the subtribe Nassauviinae (Compositae, Mutisieae). Journal of the Arnold Arboretum 55: 568–610. ———. 1974b. Marticorenia: A new genus of Mutisieae (Compositae). Journal of the Arnold Arboretum 55: 38–45. ———. 1974c. Revision of the genus Moscharia (Compositae, Mutisieae) and a reinterpretation of its inflorescence. Contributions of the Gray Herbarium of Harvard University 205: 163–173. ———. 1976a. Burkartia: Nuevo género de Mutisieae (Compositae). Boletín de la Sociedad Argentina de Botánica 17: 241–246. ———. 1976b. Revisión del género Leucheria (Compositae: Mutisieae). Darwiniana 20: 9–126. ———. 1980. Evolution in the subtribe Nassauviinae (Compositae, Mutisieae): A phylogenetic reconstruction. Taxon 29: 213–224. ——— & S. E. Freire. 1986. El género Calopappus (Compositae, Mutisieae). Caldasia 15: 57–69. ——— & C. Marticorena. 1978. Transfer of the Brazilian Trixis eryngioides to Perezia (Compositae, Mutisieae). Journal of the Arnold Arboretum 59: 352–359. Cristóbal, C. L. & A. L. Cabrera. 1982. Novedades en Hyaloseris Griseb. (Compositae). Hickenia 1: 255–257.


691

Davies, A. M. R. 2006. Chaetanthera kalinae (Mutisieae, Asteraceae), a new species from Chile. Novon 16: 51–55. de Candolle, A. P. 1838. Prodromus systematis naturalis regni vegetabilis 7. Treuttel and Würtz, Paris. de Candolle, M. 1812. Observations sur les plantes Composées, ou Syngenèses, troisième mémoire: Sur les Composées a corolles labiées ou Labiatiflores. Annales du Muséum National d’Histoire Naturelle 19: 59–72. Degener, O. 1946. Hesperomannia. In: Flora Hawaiiensis. New Illustrated Flora of the Hawaiian Islands. Published by the author. Díaz-Piedrahita, S. 2001. Tostimontia, un nuevo género de Asteraceae (Mutisieae) originario de Colombia. Revista de la Academia Colombiana de Ciencias Exactas 25: 7–10. ——— & C. Vélez-Nauer. 1993. Revisión de las tribus Barnadesieae y Mutisieae (Asteraceae) para la Flora Colombiana. Monografías del Jardín Botánico Jose Celestino Mutis 1: 1–162. Dimon, M. T. 1971. Problémes généraux soulevés par l’étude pollinique de Composés Mediterranéennes. Naturalia Monspeleliensia, Série Botanique 22: 129–144. Don, D. 1830. Descriptions of the new genera and species of the class Compositae belonging to the floras of Peru, Mexico, and Chile. Transactions of the Linnean Society of London 16: 171–297. Dormer, K. J. 1962. The fibrous layer in the anthers of the Compositae. New Phytologist 61: 150–163. Egeröd, K. & B. Stähl. 1991. Revision of Lycoseris (Compositae–Mutisieae). Nordic Journal of Botany 11: 549–574. Elsik, W. C. & T. E. Yancey. 2000. Palynomorph biozones in the context of changing paleoclimate, middle Eocene to lower Oligocene of the northwest Gulf of Mexico. Palynology 24: 177–186. Erbar, C. & P. Leins. 1995. Portioned pollen release and the syndromes of secondary pollen presentation in the Campanulales–Asterales complex. Flora 190: 323–338. Erdtman, G. 1960. The acetolysis method, a revised description. Svensk Botanisk Tidskrift utgifven uf Svenska Botaniska Föreningen (Stockholm) 54: 561–564. ———. 1966. Pollen Morphology and Plant Taxonomy. Hafner, New York and London. ———. 1969. Handbook of Palynology: An Introduction to the Study of Pollen Grains and Spores. Hafner, Munksgaard, Copenhagen and New York. Fabris, H. A. 1968. Revisión del género Proustia (Compositae). Revista del Museo de La Plata, Sección Botánica 11: 23–49. Ferreyra, R. 1944. Revisión del género Onoseris. Journal of the Arnold Arboretum 25: 349–395. ———. 1949. Una nueva especie de Onoseris procedente del Perú. Publicaciones del Museo de Historia Natural “Javier Prado,” Serie B, Botánica 2: 1–4. ———. 1953. Las especies peruanas del género Chaetanthera (Compositae). Publicaciones del Museo de Historia Natural “Javier Prado,” Serie B, Botánica 6: 1–8. ———. 1959. Dos especies nuevas de Onoseris (Compositae) para la flora peruana. Publicaciones del Museo de Historia Natural “Javier Prado,” Serie B, Botánica 11: 1–6. ———. 1980. Especies nuevas de Compuestas peruanas. Boletín de la Sociedad Peruana de Botánica 8: 75–82. ———. 1995. Family Asteraceae: Part VI. Fieldiana, Botany 35: 1–101. Franchet, M. A. 1888. Mutisiacées du Yun-nan. Journal de Botanique (Morot) 2: 1–7. Freire, S. E. 1996. Asteraceae, Tribu XII. Mutisieae, Subtribu 2. Gochnatiinae (excepto Gochnatia). Pp. 1–8, 12–17. In: A. T. Hunkiker (ed.), Flora Fanerogámica Argentina, Fascículo 28. PROFLORA, Córdoba, Argentina. ———. 2002. Two new species of Ainsliaea (Asteraceae, Mutisieae) from China and Vietnam. Novon 12: 451–455. ———. 2007. Systematic revision and phylogeny of Ainsliaea DC. (Asteraceae, Mutisieae). Annals of the Missouri Botanical Garden 94: 79–191. ——— & L. Iharlegui. 2000. Ejemplares tipo de Asteraceae (= Compositae) de A. L. Cabrera. Darwiniana 38: 307–364. ——— & L. Katinas. 1995. Morphology and ontogeny of the cypsela hairs of Nassauviinae (Asteraceae, Mutisieae). Pp. 107–143. In: D. J. N. Hind, C. Jeffrey & G. V. Pope (eds.), Advances in Compositae Systematics. Royal Botanic Gardens, Kew. ———, J. V. Crisci & L. Katinas. 1993. A cladistic analysis of Nassauvia Comm. ex Juss. (Asteraceae, Mutisieae) and related genera. Botanical Journal of the Linnean Society 112: 293–309. ———, L. Katinas & G. Sancho. 2002. Gochnatia (Asteraceae: Mutisieae) and the Gochnatia complex: Taxonomic implications from morphology. Annals of the Missouri Botanical Garden 89: 525–550. Funk, V. A., R. Chan & S. C. Keeley. 2004. Insights into the evolution of the tribe Arctoteae (Compositae: subfamily Cichorioideae s.s.) using trnL-F, ndhF, and ITS. Taxon 53: 637–655.

692

L. Katinas, et al.

———, R. J. Bayer, S. Keeley, R. Chan, L. Watson, B. Gemeinholzer, E. Schilling, J. L. Panero, B. G. Baldwin, N. Garcia-Jacas, A. Susanna & R. K. Jansen. 2005. Everywhere but Antarctica: Using a supertree to understand the diversity and distribution of the Compositae. Biologiske Skrifter 55: 343–374. Gamerro, J. C. 1990. Identidad de Pseudostifftia con Moquinia (Compositae) y consideraciones sobre la ubicación tribal del taxón. Darwiniana 30: 123–136. Garnatje, T. & J. Martin. 2007. Pollen studies in the genus Echinops L. and Xeranthemum group (Asteraceae). Botanical Journal of the Linnean Society 154: 549–557. Gay, C. 1847. Historia física y política de Chile. Tomo 3, Museo de Historia Natural de Santiago, Santiago de Chile, Chile. Gleason, H. A. 1931. The botanical results of the Tyler-Duida expedition. Bulletin of the Torrey Botanical Club 58: 277–501. González-Medrano, J., J. L. Villaseñor & R. Medina. 2004. A new species of Gochnatia (Asteraceae, Mutisieae) from the desert scrubland of the state of Hidalgo, Mexico. Novon 14: 434–436. Graham, A. 1996. A contribution to the geologic history of the Compositae. Pp. 123–140. In: D. J. N. Hind & H. J. Beentje (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994, Vol. 1. Royal Botanic Gardens, Kew. Grau, J. 1980. Die testa der Mutisieae und ihre systematische Bedeutung. Mitteilungen aus der Botanischen Staatssammlung München 16: 269–332. Greuter, W., J. McNeill, F. R. Barrie, H.-M. Burdet, V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & D. L. Hawksworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Code). Regnum Vegetabile 138. Gustafsson, M. H. G., A. S.-R. Pepper, V. A. Albert & M. Källersjö. 2001. Molecular phylogeny of the Barnadesioideae (Asteraceae). Nordic Journal of Botany 21: 149–160. Hansen, H. V. 1985a. A taxonomic revision of the genus Gerbera (Compositae–Mutisieae) sections Gerbera, Parva, Piloselloides (in Africa) and Lasiopus. Opera Botanica a Societate Botanica Lundensi 78: 5–36. ———. 1985b. Notes on Gerbera sect. Pseudoseris (Compositae–Mutisieae). Nordic Journal of Botany 5: 451–453. ———. 1985c. A taxonomic revision of the genus Perdicium (Compositae–Mutisieae). Nordic Journal of Botany 5: 543–546. ———. 1988. A taxonomic revision of the genera Gerbera sect. Isanthus, Leibnitzia (in Asia), and Uechtritzia (Compositae, Mutisieae). Nordic Journal of Botany 8: 61–76. ———. 1990. Phylogenetic studies in the Gerbera-complex (Compositae, tribe Mutisieae, subtribe Mutisiinae). Nordic Journal of Botany 9: 469–485. ———. 1991a. Phylogenetic studies in Compositae tribe Mutisieae. Opera Botanica a Societate Botanica Lundensi 109: 1–50. ———. 1991b. SEM-studies and general comments on pollen in the tribe Mutisieae (Compositae) sensu Cabrera. Nordic Journal of Botany 10: 607–623. Harling, G. 1991. Compositae–Mutisieae. Pp. 1–106. In: G. Harling & L. Andersson (eds.), Flora of Ecuador, Vol. 42. Department of Systematic Botany, University of Göteborg, Section of Botany, Riksmuseum, Stockholm, Pontificia Universidad Católica del Ecuador, Quito. ———. 1995. The genus Jungia L. fil. (Compositae–Mutisieae). Acta Regiae Societatis Scientiarum et Litterarium Gothoburgensis, Botanica 4: 5–133. ———. 1997. A new species of Jungia (Compositae) from Bolivia. Novon 7: 246–248. Harris, J. G. & M. Wolf Harris. 1994. Plant Identification Terminology. An Illustrated Glossary. Spring Lake, Spring Lake, Utah. Hellwig, P. 1985. Bau der blüten und köpfchen von Moscharia und verwandten gattungen der Compositae–Mutisieae. Mitteilungen aus der Botanischen Staatssammlung München 21: 1–47. Heng, L. 1995. Discussion on floristic features of the rare and endangered Compositae plant of China: Nouelia insignis. Acta Botanica Yunnanica 17: 401–404. (In Chinese). Herman, P. P. J. 2002. Revision of Tarchonanthus camphoratus complex (Asteraceae–Tarchonantheae) in southern Africa. Bothalia 32: 21–28. Hershkovitz, M. A., M. T. K. Arroyo, C. Bell & L. F. Hinojosa. 2006. Phylogeny of Chaetanthera (Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages. Molecular Phylogenetics and Evolution 41: 594–605. Heusser, C. J. 1971. Pollen and Spores of Chile: Modern Types of the Pteridophyta, Gymnospermae and Angiospermae. The University of Arizona Press, Tucson. Hickman, J. C. (editor) 1993. The Jepson Manual: Higher Plants of California. University of California Press, Berkeley and Los Angeles. Hind, D. J. N. 2000. A new species and a commentary on the genus Trixis (Compositae: Mutisieae) in Bahia, Brazil. Kew Bulletin 55: 381–386.


693

———. 2001a. Problems in Pertya Sch.Bip. Compositae Newsletter 36: 9–13. ———. 2001b. A new combination in Amblysperma (Compositae: Mutisieae). Kew Bulletin 56: 711–713. ———. 2004. A new species of Jungia (Compositae: Mutisieae: Nassauviinae) from Bolivia. Kew Bulletin 59: 311–314. ———. 2007. Tribe Mutisieae. Pp. 90–123. In: K. Kubitzki (ed.), The Families and Genera of Vascular Plants, Vol. 8. Springer, Berlin and Heidelberg. ——— & J. Semir. 1998a. A new combination in Stifftia (Compositae: Mutisieae). Kew Bulletin 53: 617–622. ——— & ———. 1998b. Typification of Wunderlichia (Compositae: Mutisieae). Kew Bulletin 53: 1011–1012. Hoffmann, O. 1894. Compositae. Pp. 87–391. In: A. Engler & K. Prantl (eds.), Die Natürlichen Pflanzenfamilien, Vol. 4(5). von Wilhelm Engelmann, Leipzig. Humbert, H. 1948. Un genre nouveau remarquable de Mutisiées à Madagascar. Bulletin de la Société Botanique de France 95: 181–183. ———. 1963. 189e Famille Composées. Flore de Madagascar et des Comores. 3. Museum d’Histoire Naturelle, Paris. Jansen, R. K. & K. -J. Kim. 1996. Implications of chloroplast DNA data for the classification and phylogeny of the Asteraceae. Pp. 317–339. In: D. J. N. Hind & H. Beentje (eds.), Compositae: Systematics. Proceedings of the International CompositaeConference, Kew, 1994, Vol. 1. Royal Botanic Gardens, Kew. ——— & J. D. Palmer. 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proceedings of the National Academy of Sciences of the United States of America 84: 5818–5822. ——— & ———. 1988. Phylogenetic implications of chloroplast DNA restriction site variation in the Mutisieae (Asteraceae). American Journal of Botany 75: 753–766. Jarvis, C. E. & N. J. Turland (editors). 1998. Typification of Linnean specific and varietal names in the Compositae (Asteraceae). Taxon 47: 347–370. Jeffrey, C. 1967. Notes on Compositae II. The Mutisieae in east tropical Africa. Kew Bulletin 21: 177–223. ———. 1977. Corolla forms in Compositae—Some evolutionary and taxonomic speculations. Pp. 111– 118. In: V. H. Heywood, J. B. Harborne & B. L. Turner (eds.), The Biology and Chemistry of the Compositae, Vol. 1. Academic, London. ———. 2004. Systema Compositarium (Asteraceum) nova. Botanical Journal 89: 1817–1822. ———. 2007. Introduction with key to tribes. Pp. 61–87. In: K. Kubitzki (ed.), The Families and Genera of Vascular Plants, Vol. 8. Springer, Berlin, Heidelberg. Jervis, R. N. 1954. A Summary of the Genus Gochnatia Including a Revision of the West Indian Species which Comprise the Section Anastraphioides. Ph.D. Dissertation, University of Michigan. Jiménez Rodríguez, F., L. Katinas, M. C. Tellería & J. V. Crisci. 2004. Salcedoa gen. nov., a biogeographic enigma in the Caribbean Mutisieae (Asteraceae). Systematic Botany 29: 987–1002. Johnston, I. M. 1929. Undescribed species from the Cordillera of Atacama. Contributions of the Gray Herbarium of Harvard University 85: 164–172. Kalin Arroyo, M. T. & C. Marticorena. 1988. A new species of the South American genus Nassauvia (Compositae: Mutisieae) from Chilean Patagonia. Brittonia 40: 332–334. Karis, P. O., M. Källersjö & K. Bremer. 1992. Phylogenetic analysis of the Cichorioideae (Asteraceae), with emphasis on the Mutisieae. Annals of the Missouri Botanical Garden 79: 416–427. Katinas, L. 1994. Un nuevo género de Nassauviinae (Asteraceae, Mutisieae) y sus relaciones cladísticas con los géneros afines de la subtribu. Boletín de la Sociedad Argentina de Botánica 30: 59–70. ———. 1995. Tribu XII. Mutisieae. Subtribu 4. Nassauviinae. Pp. 5–58. In: A. T. Hunziker (ed.), Flora Fanerogámica Argentina, Fascículo 13. PROFLORA, Córdoba, Argentina. ———. 1996a. Tribu XII. Mutisieae. Subtribu 3. Mutisiinae. Pp. 1–40. In: A. T. Hunziker (ed.), Flora Fanerogámica Argentina, Fascículo 4. PROFLORA, Córdoba, Argentina. ———. 1996b. Revisión de las especies sudamericanas del género Trixis (Asteraceae, Mutisieae). Darwiniana 34: 27–108. ———. 1998. The Mexican Chaptalia hintonii is a Gerbera (Asteraceae, Mutisieae). Novon 8: 380–385. ———. 2000. Implications of morphological phylogenetics for the placement of the genera Adenocaulon and Eriachaenium (Asteraceae). Plant Systematics and Evolution 223: 229–250. ———. 2004a. The Gerbera-complex (Asteraceae, Mutisieae): To split or not to split. Sida 21: 935–940. ———. 2004b. Amblysperma should be retained under Trichocline. Taxon 53: 108–112. ———. 2008. The genus Pachylaena (Asteraceae, Mutisieae). Botanical Journal of the Linnean Society 157: 373–380. ——— & J. V. Crisci. 2000. Cladistic and biogeographic analyses of the genera Moscharia and Polyachyrus (Asteraceae, Mutisieae). Systematic Botany 25: 33–46.

694

L. Katinas, et al.

——— & J. V. Crisci. 2008. Reconstructing the biogeographical history of two plant genera with different dispersion capabilities. Journal of Biogeography 35: 1374–1384. ——— & T. F. Stuessy. 1997. Revision of Doniophyton (Compositae, Barnadesioideae). Plant Systematics and Evolution 206: 33–45. ———, J. V. Crisci & S. E. Freire. 1992. Revisión sistemática y análisis cladístico del género Triptilion Ruiz et Pavón (Asteraceae, Mutisieae). Boletín de la Sociedad de Biología de Concepción 63: 101–132. ———, J. V. Crisci, M. C. Tellería, V. Barreda & L. Palazzesi. 2007. Early history of Asteraceae in Patagonia: Evidence from fossil pollen grains. New Zealand Journal of Botany 45: 605–610. ———, J. V. Crisci, R. Schmidt Jabaily, C. Williams, J. Walker, B. Drew, J. M. Bonifacino & K. J. Sytsma. 2008a. Evolution of secondary heads in Nassauviinae (Asteraceae, Mutisieae). American Journal of Botany 95: 229–240. ———, M. C. Tellería & J. V. Crisci. 2008b. A new species of Leucheria (Asteraceae, Mutisieae) from Chile. Novon 18: 366–369. Katinas, L., M. C. Tellería, A. Susanna & S. Ortiz. Warionia (Asteraceae): A relict genus of Cichorioideae? Anales del Jardín Botánico de Madrid 65. In press. Keeley, S. C. & R. K. Jansen. 1991. Evidence from chloroplast DNA for the recognition of a new tribe, the Tarchonantheae, and the tribal placement of Pluchea (Asteraceae). Systematic Botany 16: 173–181. Keighery, G. J. 2005. A new species of Amblysperma (Asteraceae: Mutisieae). Compositae Newsletter 42: 26–31. Kim, H.-G., S. C. Keeley, P. S. Vroom & R. K. Jansen. 1998. Molecular evidence for an African origin of the Hawaiian endemic Hesperomannia (Asteraceae). Proceedings of the National Academy of Sciences of the United States of America 95: 15440–15445. Kim, K.-J., R. K. Jansen, R. S. Wallace, H. J. Michaels & J. D. Palmer. 1992. Phylogenetic implications of rbcL sequence variation in the Asteraceae. Annals of the Missouri Botanical Garden 79: 428–445. Kim, K.-J., D. J. Loockerman & R. K. Jansen. 2002. Systematic implications of ndhF sequence variation in the Mutisieae (Asteraceae). Systematic Botany 27: 598–609. ———, K.-S. Choi & R. K. Jansen. 2005. Two chloroplast DNA inversions originated simultaneously during the early evolution of the sunflower family (Asteraceae). Molecular Biology and Evolution 22: 1783–1792. Kimball, R. T., D. J. Crawford & T. K. Lowrey. 2004. A molecular study of Brachylaena (Asteraceae): phylogenetic and biogeographic implications. Abstracts, Botany 2004, Snowbird, Utah. Kitamura, S. 1937. Tribe Mutisieae. Memoirs of the College of Science, Kyoto Imperial University, Series B, Biology 13: 292–316. Koch, M. F. 1930. Studies in the anatomy and morphology of the Compositae flower. II. The corollas of the Heliantheae and the Mutisieae. American Journal of Botany 17: 995–1010. Koster, J. T. 1945. Chaetanthera. Blumea 5: 673–674. Koyama, H. 1973. On the taxonomic characters of the genus Pertya. Proceedings of the Japanese Society of Plant Taxonomists 3: 20–21. ———. 1975. Notes on Pertya hossei and its allies. Bulletin of the National Science Museum, Tokyo 1: 49–58. Lagasca, M. 1811. Disertación sobre un nuevo orden de plantas de la clase de las Compuestas. Amenidades Naturales de las Españas: 26–44. Leins, P. 1968. Versuch einer Gliederung der Inulinae und Buphtalminae nach den Pollenkorn typen. Berichte der Deutschen Botanischen Gesellschaft, Berlin 81: 498–504. Lessing, C. F. 1832. Synopsis Generum Compositarum. Duncker and Humblot, Berlin. Lin, N. N., H. Wang, D. Li & S. Blackmore. 2005. Pollen morphology of eight genera of the subtribe Mutisiinae Less. sensu Bremer (Compositae) from Asia. Journal of Integrative Plant Biology 47: 1036– 1046. Lindley, J. 1829. Mutisioideae. Pp. 1072–1074. In: J. Loudon (ed.), An Encyclopaedia of Plants. Longman, Rees, Orme, Brown and Green, London. Luan, S., T.-Y. Chiang & X. Gong. 2006. High genetic diversity vs. low genetic differentiation in Nouelia insignis (Asteraceae), a narrowly distributed endemic species in China, revealed by ISSR fingerprinting. Annals of Botany 98: 583–589. Maguire, B. 1956. Distribution, endemicity, and evolution patterns among Compositae of the Guayana Highland of Venezuela. Proceedings of the American Philosophical Society 100: 467–475. ———. 1967. Compositae. The botany of the Guayana Highland—Part VII. Memoirs of the New York Botanical Garden 17: 437–439. ——— & J. J. Wurdack. 1958. New Guayana Compositae. Boletín de la Sociedad Venezolana de Ciencias Naturales, Caracas 20: 54–59.


695

———, ——— & collaborators. 1957a. The botany of the Guayana Highland—Part II. Memoirs of the New York Botanical Garden 9: 235–392. ———, J. A. Steyermark, J. J. Wurdack & collaborators. 1957b. Botany of the Chimantá Massif-I. Gran Sabana, Venezuela. Memoirs of the New York Botanical Garden 9: 393–439. Markgraf, V. & H. L. D’Antoni. 1978. Pollen Flora of Argentina. Arizona, Tucson. Marticorena, C. & O. Parra. 1975. Morfología de los granos de polen de Hesperomannia Gray y Moquinia DC. (Compositae–Mutisieae). Estudio comparativo con géneros afines. Gayana, Botánica 29: 1–22. ——— & M. Quezada. 1974. Compuestas nuevas o interesantes para Chile. Boletín de la Sociedad de Biología de Concepción 48: 99–108. Mattfeld, J. 1934. Compositae Novae Sinenses. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 11: 103–110. Mitsui, Y., S.-T. Chen, Z.-K. Zhou, C.-I.-Peng, Y. F. Deng & H. Segoguchi. 2007. Phylogeny and biogeography of the genus Ainsliaea (Asteraceae) in the Sino-Japanese region based on nuclear rDNA and plastid DNA sequence data. Annals of Botany 101: 111–124. Molfino, J. F. 1953. Una nueva especie del género Aphyllocladus. Boletín de la Sociedad Argentina de Botánica 5: 30–34. Mondin, C. A. 1995. Holocheilus monocephalus (Asteraceae–Mutisieae), nova espécie do sul do Brasil. Napaea 11: 31–34. ——— & C. L. Vasques. 2004. O gênero Holocheilus Cass. (Asteraceae–Mutisieae–Nassauviinae) no Rio Grande do Sul, Brasil. Iheringia, Botanica 59: 161–172. Moore, D. M. 1983. Flora of Tierra del Fuego. Anthony Nelson, Shropshire, England. Moore, S. 1925. The genus Pleiotaxis Steetz (Compositae). Journal of Botany 63: 43–50. Muñoz Pizarro, C. 1966. Sinopsis de la Flora Chilena: Claves para la Identificación de Familias y Géneros. Ediciones de la Universidad de Chile, Santiago. Nayar, M. P. & M. Ahmedullah. 1985. Catamixis baccharoides Thomson—An endemic chasmophyte of the W. Himalaya under threat. Bulletin of the Botanical Survey of India 27: 248–250. Nesom, G. L. 1983. Biology and taxonomy of American Leibnitzia (Asteraceae: Mutisieae). Brittonia 35: 126–139. ———. 1984. Taxonomy and distribution of Chaptalia dentata and C. albicans (Asteraceae: Mutisieae). Brittonia 36: 396–401. ———. 1995. Revision of Chaptalia (Asteraceae: Mutisieae) from North America and continental Central America. Phytologia 78: 153–188. Nordenstam, B. & E. El-Ghazaly. 1977. Floral micromorphology and pollen ultrastructure in some Centaureinae (Compositae) mainly from Egypt. Publicatins of the Cairo University Herbarium N° 7 & 8. Ohwi, J. 1965. Flora of Japan. National Science Museum, Tokyo; Smithsonian Institution, Washington, DC. Ortiz, S. 2000. A phylogenetic analysis of Dicoma Cass. and related genera (Asteraceae: Cichorioideae: Mutisieae) based on morphological and anatomic characters. Annals of the Missouri Botanical Garden 87: 459–481. ———. 2001. Reinstatement of the genus Macledium Cass. (Asteraceae, Mutisieae): Morphological and phylogenetic arguments. Taxon 50: 733–744. ———. 2006a. Systematics of Cloiselia (Asteraceae, Mutisieae s. l.), a reinstated Madagascan genus. Systematic Botany 31: 421–431. ———. 2006b. African Mutisieae. P. 14. In: Book of Abstracts. The International Compositae Alliance (TICA–Deep Achene), Barcelona, Spain. ——— & A. P. Coutinho. 2001. Achyrothalamus reduced to Erythrocephalum (Asteraceae, Mutisieae). Taxon 50: 389–403. ——— & N. C. Netnou. 2005. A new species of Dicoma (Asteraceae, Mutisieae) from South Africa. Botanical Journal of the Linnean Society 147: 509–513. ——— & I. Pulgar. 2002. A new species of Dicoma Cass. (Asteraceae: Mutisieae) from Namibia. Botanical Journal of the Linnean Society 139: 317–322. ——— & J. Rodríguez-Oubiña. 1998. The infraspecific taxa of Pleiotaxis huillensis (Mutisieae, Asteraceae). Nordic Journal of Botany 19: 23–26. ———, J. Rodríguez-Oubiña & M. Tadesse. 1998. A taxonomic revision of Dicoma (Asteraceae: Cichorioideae: Mutisieae) for the horn of Africa. Annals of the Missouri Botanical Garden 85: 440–459. Ozenda, P. 1991. Flore et Végétation du Sahara, 3rd ed. Centre National de la Recherche Scientifique, Paris. Paiva, J. A. R. 1972. New and little known species from the flora zambesiaca area. Boletim da Sociedade Broteriana 46: 355–384. Panero, J. L. 2007. Calorezia, a new genus of tribe Nassauvieae (Asteraceae, Mutisioideae). Phytologia 89: 198–201. ——— & V. A. Funk. 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proceedings of the Biological Society of Washington 115: 909–922.

696

L. Katinas, et al.

——— & ———. 2007. New infrafamilial taxa in Asteraceae. Phytologia 89: 356–360. ——— & ———. 2008. The value of sampling anomalous taxa in phylogenetic studies: Major clades of the Asteraceae revealed. Molecular Phylogenetics and Evolution 47: 757–782. Parra, O. & C. Marticorena. 1972. Granos de polen de plantas chilenas, II. Compositae–Mutisieae. Chile. Gayana, Botánica 21: 1–107. Pehlivan, S. 1995. Pollen morphology of some Turkish endemic Centaurea. Grana Palynologica 34: 29–38. Poljakov, P. P. 1967. Systematika i proischozdenie sloznocvetnych. Alma-Ata, Nauka. Pope, G. V. 1991. Notes on Dicoma Cass. (Compositae). Kew Bulletin 46: 699–709. ———. 1992. Flora Zambesiaca. Managing Commitee on Behalf of the Contributors to Flora Zambesiaca. Royal Botanic Gardens, Kew. Pruski, J. F. 1989a. Notes on the Compositae of the Guayana Highland—I. A new species of Stomatochaeta and the reduction of Guaicaia to Glossarion (Compositae: Mutisieae). Brittonia 41: 35–40. ———. 1989b. Compositae of the Guayana Highland—II. Novelties in Gongylolepis and Stenopadus (Mutisieae). Annals of the Missouri Botanical Garden 76: 993–1003. ———. 1991. Compositae of the Guayana Highland—V. The Mutisieae of the Lost World of Brazil, Colombia, and Guyana. Boletim do Museo Paraense de Historia Natural 7: 335–392. ———. 1997. Asteraceae. Pp. 177–393. In: J. A. Steyermark, P. E. Berry & B. K. Holst (eds.), Flora of the Venezuelan Guayana, Vol. 3. Missouri Botanical Garden, St. Louis. ———. 1998. Stenopadus andicola sp. nov. (Asteraceae: Mutisieae), a new generic record for Ecuador. Novon 8: 67–69. ———. 2004. Missouri Botanical Garden, Research: Asteraceae (Compositae). The Missouri Botanical Garden, Poster, St. Louis. ——— & H. Beltrán. 2003. Stenopadus andicola (Compositae: Mutisieae), a new generic record for Peru. Compositae Newsletter 39: 2–12. ——— & G. Sancho. 2004. Asteraceae or Compositae (Aster or sunflower family). Pp. 33–39. In: N. P. Smith, S. V. Heald, A. Anderson, S. A. Mori & D. W. Stevenson (eds.), Flowering Plants of the Neotropics. Princeton University Press, Princeton. Punt, W., S. Blackmore, S. Nilsson & A. Le Thomas. 1994. Glossary of pollen and spores terminology. Laboratory of Palaeobotany and Palynology. Contributions Series of the University of Utrech, ser. 1, 1: 1–71. Rao, R. R. 1995. Tribe Mutisieae Cass. Pp. 163–186. In: P. K. Hajra, R. R. Rao, D. K. Singh & B. Uniyal (eds.), Flora of India, Bulletin of the Botanical Survey of India. Vol. 13. Botanical Survey of India, India. Reiche, C. 1905. Mutisieas. Pp. 288–460. In: Flora de Chile, Vol. 4. Imprenta Cervantes, Santiago de Chile, Chile. Reveal, J. L. & R. M. King. 1973. Re-establishment of Acourtia D. Don (Asteraceae). Phytologia 27: 228–232. Ricardi, M. & E. Weldt. 1974. Revisión del género Polyachyrus (Compositae). Gayana, Botánica 26: 3–34. Robinson, H. 1979. Two new genera of Vernonieae (Asteraceae) from Brazil: Heterocypsela and Pseudostifftia. Phytologia 44: 442–450. ———. 1980. Harthamnus, a new genus of Mutisieae from Bolivia. Phytologia 45: 451–455. ———. 1983. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contributions to Botany 54: 1–69. ———. 1987. Some suggestions regarding the significance of chloroplast DNA variation in the Asteraceae. Phytologia 63: 316–324. ———. 1988. A new species of Trichocline from northern Peru. Phytologia 65: 47–49. ———. 1991. Two new species of Stifftia with notes on relationships of the genus (Asteraceae: Mutisieae). Systematic Botany 16: 685–692. ———. 1994. Notes on the tribes Eremothamneae, Gundeliae, and Moquinieae, with comparisons of their pollen. Taxon 43: 33–44. ———. 1999. Generic and subtribal classification of American Vernonieae. Smithsonian Contributions to Botany 89: 1–116. ——— & R. D. Brettell. 1973. Tribal revisions in Asteraceae. VI. The relationships of Eriachaenium. Phytologia 26: 71–72. ——— & B. Kahn. 1985. A new species of Stifftia from Cayenne. Phytologia 58: 248–251. ——— & C. Marticorena. 1986. A palynological study of the Liabeae (Asteraceae). Smithsonian Contributions to Botany 64: 1–50. Roque, N. 1997. A reassessment of Actinoseris polymorpha (Less.) Cabrera (Mutisieae–Compositae), with a new combination. Kew Bulletin 52: 197–204. ———. 2001a. Five new species of Richterago (Compositae, Mutisieae): A genus endemic to Brazil. Novon 11: 341–349.


697

———. 2001b. Fenología de Wunderlichia mirabilis Riedel ex Baker (Compositae–Mutisieae) na Serra do Cipó, Minas Gerais, Brasil. Sitientibus, Série Ciencias Biologicas 1: 108–111. ——— & D. J. N. Hind. 2001. Ianthopappus, a new genus of the tribe Mutisieae (Compositae). Novon 11: 97–101. ——— & N. Nakajima. 2001. Two new species of Richterago Kuntze (Asteraceae, Mutisieae) from Minas Gerais. Kew Bulletin 56: 697–703. ——— & J. R. Pirani. 1997. Flora da Serra do Cipó, Minas Gerais: Compositae–Barnadesieae e Mutisieae. Boletim de Botânica da Universidade de São Paulo 16: 151–185. ——— & ———. 2001. Reinstatement of the name Richterago Kuntze and recircumscription of the genus to include species formerly treated as Actinoseris (Endl.) Cabrera (Compositae, Mutisieae). Taxon 50: 1155–1160. ——— & M. S. F. Silvestre-Capelato. 2001. Generic delimitation of Gochnatia, Richterago and Ianthopappus (Compositae–Mutisieae) based on pollen morphology. Grana Palynologica 40: 197– 204. Rydberg, P. A. 1917. Flora of the Rocky Mountains and Adjacent Plains. Published by the author, New York. Rzedowski, J. 1983. Algunas adiciones al género Acourtia (Compositae–Mutisieae). Boletín de la Sociedad Botánica de México 45: 97–109. Sagástegui Alva, A. & M. O. Dillon. 1985. Four new species of Asteraceae from Peru. Brittonia 37: 6–13. Salgado-Labouriau, M. L. 1982. Pollen morphology of the Compositae of the northern Andes. Pollen and Spores 24: 397–452. Sancho, G. 1996. Asteraceae, Tribu XII. Mutisieae, Subtribu 2. Gochnatia. Pp. 9–11. In: A. T. Hunziker (ed.), Flora Fanerogámica Argentina, Fascículo 28. PROFLORA, Córdoba, Argentina. ———. 1999. Gochnatia calophylla: A synonym of Gochnatia oligocephala (Gardner) Cabrera (Asteraceae, Mutisieae). Willdenowia 29: 235–237. ———. 2000. Revisión y filogenia de la sección Moquiniastrum Cabrera del género Gochnatia Kunth (Asteraceae, Mutisieae). Fontqueria 54: 61–122. ———. 2004. Phylogenetic relationships within the genus Onoseris (Asteraceae, Mutisieae) inferred form morphology. Systematic Botany 29: 432–447. ———, S. E. Freire, L. Katinas & M. C. Tellería. 2005. A new species and a new combination in Andean Mutisieae (Asteraceae). Taxon 54: 85–90. Scott, L., A. Cadman & I. McMillan. 2006. Early history of Cainozoic Asteraceae along the southern African west coast. Review of Palaeobotany and Palynology 142: 47–52. Selling, O. H. 1947. Studies in Hawaiian Pollen Statistics. Part III. The Pollen of the Hawaiian Phanerogams. Bernice P. Bishop Museum Special Publication 38: 1–430. Simpson, B. B. 1973. Contrasting modes of evolution in two groups of Perezia (Mutisieae, Compositae) of southern South America. Taxon 22: 525–536. ———. 1978. Compositae, tribe Mutisieae (except Trixis). Pp. 1–10. In: C. T. Rogerson ed.), North American Flora 2. The New York Botanical Garden, New York. Simpson Vuilleumier, B. 1969. The systematics and evolution of Perezia section Perezia (Compositae). Contributions of the Gray Herbarium of Harvard University 199: 1–163. Skvarla, J. J. & B. L. Turner. 1966. Systematic implications from electron microscopic studies of Compositae pollen: A review. Annals of the Missouri Botanical Garden 53: 220–256. ———, B. L. Turner, V. C. Patel & A. S. Tomb. 1977. Pollen morphology in the Compositae and in morphologically related families. Pp. 141–248. In: V. H. Heywood, J. B. Harborne & B. L. Turner (eds.), The Biology and Chemistry of the Compositae, Vol. 1. Academic, London. Small, J. 1919. The origin and development of the Compositae. New Phytologist 11: 1–333. Southworth, D. 1983. Exine development in Gerbera jamesonii (Asteraceae: Mutisieae). American Journal of Botany 70: 1038–1047. Stearn, W. T. 1996. Botanical Latin: History, Grammar, Syntax, Terminology and Vocabulary. Timber, Portland, Oregon. Steyermark, J. A., B. Maguire & collaborators. 1972. The Flora of Meseta del Cerro Jáua. Memoirs of the New York Botanical Garden 23: 833–892. Stix, E. 1960. Pollenmorphologische Untersuchungen an Compositen. Grana Palynologica 2: 41–107. Stuessy, T. F., T. Sang & M. L. Devore. 1996. Phylogeny and biogeography of the subfamily Barnadesioideae with implications for early evolution of the Compositae. Pp. 463–490. In: D. J. N. Hind & H. J. Beentje (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994, Vol. 1. Royal Botanic Gardens, Kew. Teeri, T. H., P. Elomaa, M. Kotilainen & V. A. Albert. 2006. Mining plant diversity: Gerbera as a model system for plant developmental and biosynthetic research. BioEssays 28: 756–767. Tellería, M. C. 2008. Taxonomic significance of pollen types in the Guyana Highland-centred composite genera of Mutisioideae (Asteraceae). Botanical Journal of the Linnean Society 156: 327–340.

698

L. Katinas, et al.

——— & A. Forcone. 2002. Morfología del polen de las mieles del valle de Río Negro, valle inferior del río Chubut y llanura del río Senguerr (Patagonia Argentina). Boletín de la Sociedad Argentina de Botánica 37: 235–250. ——— & L. Katinas. 2004. A Comparative palynologic study of Chaetanthera (Asteraceae, Mutisieae) and allied genera. Systematic Botany 29: 752–773. ——— & ———. 2005. The unusual occurrence of tricolpate pollen within Mutisieae (Asteraceae). Grana Palynologica 44: 91–97. Tellería, M. C. & L. Katinas. New insights into the pollen morphology of the genus Mutisia (Asteraceae, Mutisieae). Plant Systematics and Evolution. In press. ———, E. Urtubey & L. Katinas. 2003. Proustia and Lophopappus (Asteraceae, Mutisieae): Generic and subtribal relationships based on pollen morphology. Review of Paleobotany and Palynology 123: 237–246. Thomson, T. 1865. On two new genera of Compositae Mutisiaceae from India. Journal of the Linnean Society, Botany 9: 342–344. Tormo-Molina, R. & J. L. Ubera-Jiménez. 1995. Tipos polínicos de la Tribu Cardueae en la península Ibérica. Monografías del Jardín Botánico de Córdoba 2: 1–52. Tseng, Y. C. 1985. Materials for Chinese Pertya (Compositae). Guihaia 5: 327–335. Turner, B. L. 1986. A new species of Hyaloseris (Asteraceae–Mutisieae) from Bolivia. Phytologia 59: 317–318. ———. 1993a. New taxa, new combinations, and nomenclatural comments on the genus Acourtia (Asteraceae: Mutisieae). Phytologia 74: 385–413. ———. 1993b. Berylsimpsonia (Asteraceae: Mutisieae), a new genus of the Greater Antilles. Phytologia 74: 349–355. Ulloa Ulloa, C. & P. M. Jørgensen. 1996. A new species of Mutisia (Compositae–Mutisieae) from Ecuador. Novon 6: 131–133. Urtubey, E. 1999. Revisión del género Barnadesia (Asteraceae: Barnadesioideae, Barnadesieae). Annals of the Missouri Botanical Garden 86: 57–117. ——— & T. F. Stuessy. 2001. New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae) based on morphology. Taxon 50: 1043–1063. ——— & M. C. Tellería. 1998. Pollen morphology of the subfamily Barnadesioideae (Asteraceae) and its phylogenetic and taxonomic implications. Review of Paleobotany and Palynology 104: 19–37. Vezey, E. L., L. E. Watson, J. J. Skvarla & R. Estes. 1994. Plesiomorphic and apomorphic pollen structure characteristics of Anthemideae (Asteroideae: Asteraceae). Amererican Journal of Botany 81: 648–657. Von Bunge, A. A. 1835. Myripnois. Mémoires présentés a l’ Académie Impériale des Sciences de St.Pétersbourg par Divers Savans et Lus dans ses Assemblées 2: 112. Wagenitz, G. 1955. Pollenmorphologie and Systematick in der Gattung Centaurea L. s. l. Flora 142: 213–279. Wagner, W. L., D. R. Herbst & S. H. Sohmer. 1990. Manual of the Flowering Plants of Hawaii. University of Hawaii Press, Honolulu. Wilson, F. C. 1923. Revision of the genus Dicoma. Bulletin of Miscellaneous Information 1923: 377–388. Wingenroth, M. & C. J. Heusser. 1983. Pollen of the high Andean Flora. Instituto Argentino de Nivología y Glaciología, IANIGLA, CONICET, Buenos Aires, Argentina. Wodehouse, R. P. 1929a. Pollen grains in the identification and classification of plants, III. The Nassauviinae. Bulletin of the Torrey Botanical Club 56: 123–138. ———. 1929b. Pollen grains in the identification and classification of plants, IV. The Mutisieae. American Journal of Botany 16: 297–313. ———. 1935. Pollen Grains. McGraw-Hill, New York. Wortley, A. H., V. A. Funk, H. Robinson, J. J. Skvarla & S. Blackmore. 2007. A search for pollen morphological synapomorphies to classify rogue genera in Compositae (Asteraceae). Review of Palaeobotany and Palynology 146: 169–181. Zardini, E. M. 1974. Sobre la presencia del género Gerbera en América. Boletín de la Sociedad Argentina de Botánica 16: 103–108. ———. 1975. Revisión del género Trichocline (Compositae). Darwiniana 19: 618–733. ———. 1980. Lulia: Un nuevo género de Compuestas. Boletín de la Sociedad Argentina de Botánica 19: 255–258. Zao, Z., J. J. Skvarla, R. K. Jansen & M. De Vore. 2000. Phylogenetic implications of pollen morphology and ultrastructure in the Barnadesioideae(Asteraceae). Lundellia 3: 26–40. ———, ——— & ———. 2006. Mutisieae (Asteraceae) pollen ultrastructure atlas. Lundellia 9: 57–76. Zavada, M. S. & S. E. De Villiers. 2000. Pollen of the Asteraceae from the Paleocene–Eocene of South Africa. Grana Palynologica 39: 39–45.


Index to Names Accepted scientific names appear in boldface; other names are seen in italics. Acanthophyllum Acanthophyllum axillare Achnopogon Achnopogon steyermarkii Achnopogon virgatus Achyrothalamus Achyrothalamus marginatus Acourtia Acourtia acevedoi Acourtia aspera Acourtia belizeana Acourtia bravohollisiana Acourtia butandae Acourtia caltepecana Acourtia carpholepis Acourtia carranzae Acourtia ciprianoi Acourtia collina Acourtia cordata Acourtia coulteri Acourtia cuernavacana Acourtia dieringeri Acourtia discolor Acourtia dissiticeps Acourtia dugesii Acourtia durangensis Acourtia elizabethiae Acourtia erioloma Acourtia formosa Acourtia fragrans Acourtia fruticosa Acourtia gentryi Acourtia glandulifera Acourtia glomeriflora Acourtia gracilis Acourtia grandifolia Acourtia guatemalensis Acourtia hidalgoana Acourtia hondurana Acourtia hooveri Acourtia huajuapana Acourtia huajuapana var.

699

actinomorpha Acourtia humboldtii Acourtia intermedia Acourtia joaquinensis Acourtia lepidopoda Acourtia lobulata Acourtia longifolia Acourtia lozanii Acourtia macvaughii Acourtia mexiae Acourtia mexicana Acourtia michoacana Acourtia microcephala Acourtia moctezumae Acourtia molinana Acourtia moschata Acourtia nana Acourtia nelsonii Acourtia nudicaulis Acourtia nudiuscula Acourtia oaxacana Acourtia ovatifolia Acourtia oxylepis Acourtia palmeri Acourtia parryi Acourtia patens Acourtia pilulosa Acourtia pinetorum Acourtia platyphylla Acourtia platyptera Acourtia potosina Acourtia pringlei Acourtia pulchella Acourtia purpusii Acourtia queretarana Acourtia reticulata Acourtia runcinata Acourtia rzedowskii Acourtia scapiformis Acourtia scaposa Acourtia simulata Acourtia sinaloana Acourtia souleana Acourtia tenoriensis Acourtia tomentosa Acourtia turbinata

700

Acourtia umbratalis Acourtia venturae Acourtia veracruzana Acourtia wislizenii Acourtia wrightii Acourtia zacatecana Actinoseris Actinoseris angustifolia Actinoseris arenaria Actinoseris hatschbachii Actinoseris polymorpha Actinoseris polyphylla Actinoseris radiata Actinoseris stenophylla Adenocauleae Adenocaulinae Adenocaulon Adenocaulon bicolor Adenocaulon chilense Adenocaulon himalaicum Adenocaulon lyratum Adenocaulon nepalense Aglaodendron Aglaodendron cheiranthifolium Ainsliaea Ainsliaea acerifolia Ainsliaea angustata Ainsliaea apiculata Ainsliaea aptera Ainsliaea apteroides Ainsliaea bonatii Ainsliaea brandisiana Ainsliaea caesia Ainsliaea cavaleriei Ainsliaea chapaensis Ainsliaea cordifolia Ainsliaea crassifolia Ainsliaea dissecta Ainsliaea elegans Ainsliaea foliosa Ainsliaea fragrans Ainsliaea fulvipes Ainsliaea glabra Ainsliaea glumacea Ainsliaea gracilis Ainsliaea grossedentata Ainsliaea henryi

L. Katinas, et al.

Ainsliaea hypoleuca Ainsliaea kawakamii Ainsliaea lancangensis Ainsliaea latifolia Ainsliaea linearis Ainsliaea macrocephala Ainsliaea macroclinioides Ainsliaea mairei Ainsliaea nana Ainsliaea nervosa Ainsliaea oblonga Ainsliaea parvifolia Ainsliaea paucicapitata Ainsliaea pentaflora Ainsliaea pertyoides Ainsliaea pingbianensis Ainsliaea pteropoda Ainsliaea pteropoda var. apteroides Ainsliaea pteropoda var. macrocephala Ainsliaea qianiana Ainsliaea ramosa Ainsliaea reflexa Ainsliaea rubrinervis Ainsliaea smithii Ainsliaea spanocephala Ainsliaea spicata Ainsliaea triloba Ainsliaea trinervis Ainsliaea uniflora Ainsliaea walkeri Ainsliaea yunnanensis Aldunantea Aldunantea chilensis Aldunantea gnaphalioides Aldunantea lycopodioides Amblysperma Amblysperma minor Amblysperma scapigera Ameghinoa Ameghinoa patagonica Amphoricarpus Anandria Anastraphia Anastraphia attenuata Anastraphia buchii Anastraphia calcicola Anastraphia cowellii


Anastraphia crassifolia Anastraphia cubensis Anastraphia ekmanii Anastraphia elliptica Anastraphia enneantha Anastraphia gomezii Anastraphia ilicifolia Anastraphia inertexta Anastraphia maisiana Anastraphia mantuensis Anastraphia microcephala Anastraphia montana Anastraphia obtusifolia Anastraphia oligantha Anastraphia parvifolia Anastraphia pauciflosculosa Anastraphia picardae Anastraphia recurva Anastraphia shaferi Anastraphia tortuensis Anastraphia wilsonii Anthemideae Aphyllocaulon Aphyllocladus Aphyllocladus decussatus Aphyllocladus denticulatus Aphyllocladus ephedroides Aphyllocladus sanmartinianus Aphyllocladus spartioides Aplophyllum Archetypae Arctoteae Aristomenia Aristomenia fruticosa Arnaldoa Arnica cordata Arnica crocea Arnica gerbera Arnica grandis Arnica piloselloides Arnica serrata Artemisia verlotorum Aster crocatus Asteroideae Atasites Atractylis mexicana Atractylis purpurea

701

Augusta Augusta grandiflora Augusta parviflora Baccharis blanchetiana Baccharis neriifolia Barnadesia Barnadesieae Barnadesiinae Barnadeseae Barnadesioideae Berniera Berniera nepalensis Bertolonia Berylsimpsonia Berylsimpsonia crassinervis Berylsimpsonia vanillosma Bichenia Bichenia aurea Bichenia auriculata Bichenia cineraria Bichenia reptans Bichenia sinuata Bowmania Bowmania verbascifolia Brachyclados Brachyclados caespitosus Brachyclados lycioides Brachyclados megalanthus Brachylaena Brachylaena neriifolia Bridgesia Bridgesia echinopsoides Burkartia Burkartia lanigera Calopappus Calopappus acerosus Caloptilium Caloptilium lagascae Calorezia Caloseris Caloseris rupestris Calyceraceae Campanulaceae Cardonaea Cardonaea jauaensis Cardueae Carduoideae

702

Carduus pycnocephalus Carlininae Carmelita Carmelita formosa Carthamus tinctorius Cassiopea Castra Castra illustris Castra nobilis Castra praestans Cataleuca Catamixis Catamixis baccharoides Celmisia spathulata Centaurea Centroclinium Centroclinium albicans Cephalopappus Cephalopappus sonchifolius Cephaloseris Cephaloseris poeppigii Ceratolepis Chabraea Chabraea gayana Chabraea glabriuscula Chabraea integrifolia Chabraea landbeckii Chabraea nutans Chabraea polyclados Chabraea purpurea Chabraea salina Chabraea thermarum Chabraea viscida Chaenanthophorae Chaetachlaena Chaetachlaena odorata Chaetanthera Chaetanthera acerosa Chaetanthera acheno-hirsuta Chaetanthera apiculata Chaetanthera australis Chaetanthera aymarae Chaetanthera boliviensis Chaetanthera brachylepis Chaetanthera chilensis Chaetanthera chiquianensis Chaetanthera ciliata

L. Katinas, et al.

Chaetanthera cochlearifolia Chaetanthera dealbata Chaetanthera dioica Chaetanthera elegans Chaetanthera euphrasioides Chaetanthera flabellata Chaetanthera flabellifolia Chaetanthera glabrata Chaetanthera glandulosa Chaetanthera gnaphalioides Chaetanthera incana Chaetanthera kalinae Chaetanthera lanata Chaetanthera leptocephala Chaetanthera limbata Chaetanthera linearis Chaetanthera lycopodioides Chaetanthera microphylla Chaetanthera minuta Chaetanthera moenchioides Chaetanthera multiflora Chaetanthera pentacaenoides Chaetanthera perpusilla Chaetanthera peruviana Chaetanthera pinnatifida Chaetanthera planiseta Chaetanthera pulvinata Chaetanthera pulvinata var. achenohirsuta Chaetanthera pungens Chaetanthera pusilla Chaetanthera renifolia Chaetanthera revoluta Chaetanthera serrata Chaetanthera spathulifolia Chaetanthera sphaeroidalis Chaetanthera splendens Chaetanthera stuebelii Chaetanthera subgen. Carmelita Chaetanthera subgen. Egania Chaetanthera subgen. Oriastrum Chaetanthera subgen. Proselia Chaetanthera subgen. Tylloma Chaetanthera tenella Chaetanthera valdiviana Chaetanthera villosa Chaetantherae


Chaptalia Chaptalia albicans Chaptalia angustata Chaptalia anisobasis Chaptalia arechavaletae Chaptalia azuensis Chaptalia callacallensis Chaptalia chapadensis Chaptalia cipoensis Chaptalia comptonioides Chaptalia cordata Chaptalia cordifolia Chaptalia crassiuscula Chaptalia crispata Chaptalia dentata Chaptalia denticellata Chaptalia denticulata Chaptalia dolichopoda Chaptalia eggersii Chaptalia ekmanii Chaptalia estribensis Chaptalia exscapa Chaptalia flavicans Chaptalia gossypina Chaptalia graminifolia Chaptalia hermogensis Chaptalia hidalgoensis Chaptalia hintoni Chaptalia hololeuca Chaptalia ignota Chaptalia incana Chaptalia integerrima Chaptalia isernina Chaptalia latipes Chaptalia leiocarpa Chaptalia leptophylla Chaptalia lyratifolia Chaptalia madrensis Chaptalia malcabalensis Chaptalia mandonii Chaptalia maxima Chaptalia media Chaptalia membranacea Chaptalia meridensis Chaptalia modesta Chaptalia montana Chaptalia mornicola

703

Chaptalia nipensis Chaptalia nutans Chaptalia oblonga Chaptalia obovata Chaptalia ovalis Chaptalia paramensis Chaptalia piloselloides Chaptalia piloselloides var. graminifolia Chaptalia primulacea Chaptalia pringlei Chaptalia pumila Chaptalia rocana Chaptalia rotundifolia Chaptalia runcinata Chaptalia sect. Leria Chaptalia sect. Lieberkuhna Chaptalia sect. Loxodon Chaptalia seemannii Chaptalia shaferi Chaptalia similis Chaptalia sinuata Chaptalia spathulata Chaptalia stenocephala Chaptalia stuebelii Chaptalia texana Chaptalia tomentosa Chaptalia transiliens Chaptalia turquinensis Chaptalia undulata Chaptalia vegaensis Chardinia Cherina Cherina microphylla Chimantaea Chimantaea acopanensis Chimantaea cinerea Chimantaea eriocephala Chimantaea espeletoidea Chimantaea huberi Chimantaea humilis Chimantaea lanocaulis Chimantaea mirabilis Chimantaea rupicola Chionoptera Chionoptera gayophyta Chondrochilus Chondrochilus crenatus

704

Chucoa Chucoa ilicifolia Chucoa lanceolata Chuquiraga Cichorioideae Cladoseris Cladoseris annua Clarionea Clarionea carthamoides Clarionea ciliosa Clarionea fonkii Clarionea magellanica Clarionea pilifera Clarionea spathulata Clarionema Clarionema humilis Clarionia Clarionia runcinata Cleanthes Cleanthes hieracioides Cleistanthium Cleistanthium nepalense Cloiselia Cloiselia carbonaria Clybatis Clybatis volkmanni Cnicothamnus Cnicothamnus azafran Cnicothamnus lorentzii Cnicothamnus lorentzii var. azafran Conyza racemosa Copernicia Criscia Criscia stricta Cullumiopsis Cullumiopsis grandidieri Cursonia Cursonia peruviana Cyclolepis Cyclolepis denticulata Cyclolepis genistoides Cyclopis Dasyphyllum Dermatophyllum Diaphoranthus Diaphoranthus fuscus Diaspananthus

L. Katinas, et al.

Diaspananthus uniflorus Diaspanthus Diazeuxeae Diazeuxis Diazeuxis mutisiana Diazeuxis trinervis Dicoma Dicoma schimperi Dicoma subgen. Macledium Dicoma tomentosa Dicomeae Dicomopsis Dinoseris Dinoseris salicifolia Dipterocypsela Dolichlasium Dolichlasium lagascae Doniophyton Doronicum incanum Drozia Drozia dicephala Duidaea Duidaea marahuacensis Duidaea pinifolia Duidaea rubriceps Duidaea tatei Dumerilia Dumerilia axillaris Dumerilia humboldtii Dumerilia paniculata Dusenia Dusenia patagonica Duseniella Duseniella patagonica Egania Egania acerosa Egania apiculata Egania dioica Egania revoluta Eizaguirrea Eizaguirrea candollei Elachia Elachia euphrasioides Elachia renifolia Elachia splendens Epiclinastrum Epiclinastrum candollei


Eriachaenium Eriachaenium magellanicum Erigeron scandens Erythrocephalum Erythrocephalum marginatum Erythrocephalum zambesianum Eupatorium rigidulum Eurydochus Eurydochus bracteatus Eurydochus cortesii Euthrixia Euthrixia salsoloides Facelidae Frageria Fulcaldea Gastrocarpha Gastrocarpha runcinata Gerbera Gerbera ambigua Gerbera aurantiaca Gerbera bojeri Gerbera cordata Gerbera crocea Gerbera delavayi Gerbera diversifolia Gerbera elliptica Gerbera emirnensis Gerbera galpinii Gerbera gossypina Gerbera henryi Gerbera hieracioides Gerbera hintonii Gerbera hybrida Gerbera hypochaeridioides Gerbera jamesonii Gerbera knorringiana Gerbera kokanica Gerbera lacei Gerbera leandrii Gerbera linnaei Gerbera maxima Gerbera nivea Gerbera parva Gerbera perrieri Gerbera petasitifolia Gerbera piloselloides Gerbera raphanifolia

705

Gerbera ruficoma Gerbera sect. Lasiopus Gerbera serrata Gerbera tomentosa Gerbera viridifolia Gerbera wrightii Gerbereae Gerberia Gerberieae Gerberinae Gladiopappus Gladiopappus vernonioides Glossarion Glossarion bilabiatum Glossarion rhodanthum Gochnatia Gochnatia amplexifolia Gochnatia angustifolia Gochnatia arborescens Gochnatia arequipensis Gochnatia argentina Gochnatia argyrea Gochnatia attenuata Gochnatia barrosoae Gochnatia blanchetiana Gochnatia boliviana Gochnatia buchii Gochnatia calcicola Gochnatia cardenasii Gochnatia cinerea Gochnatia cordata Gochnatia corymbosa Gochnatia cowellii Gochnatia crassifolia Gochnatia cubensis Gochnatia curviflora Gochnatia decora Gochnatia densicephala Gochnatia discoidea Gochnatia discolor Gochnatia ekmanii Gochnatia elliptica Gochnatia enneantha Gochnatia floribunda Gochnatia foliolosa Gochnatia gardneri Gochnatia glomeriflora

706

Gochnatia glutinosa Gochnatia gomezii Gochnatia hatschbachii Gochnatia haumaniana Gochnatia hiriartiana Gochnatia hypoleuca Gochnatia ilicifolia Gochnatia intertexta Gochnatia lanceolata Gochnatia magna Gochnatia maisiana Gochnatia mantuensis Gochnatia microcephala Gochnatia mollisima Gochnatia montana Gochnatia obtusifolia Gochnatia oligantha Gochnatia oligocephala Gochnatia orbiculata Gochnatia palosanto Gochnatia paniculata Gochnatia paniculata var. densicephala Gochnatia parvifolia Gochnatia patazina Gochnatia pauciflosculosa Gochnatia picardae Gochnatia polymorpha Gochnatia pulchra Gochnatia purpusii Gochnatia ramboi Gochnatia recurva Gochnatia rotundifolia Gochnatia rusbyana Gochnatia sagraeana Gochnatia sect. Discoseris Gochnatia sect. Leucomeris Gochnatia sect. Pentaphorus Gochnatia shaferi Gochnatia smithii Gochnatia sordida Gochnatia spectabilis Gochnatia suffrutescens Gochnatia tarapacana Gochnatia tortuensis Gochnatia vargasii Gochnatia velutina Gochnatia vernonioides

L. Katinas, et al.

Gochnatia wilsonii Gochnatieae Gochnatinae Gochnatiinae Gochnatioideae Gongylolepis Gongylolepis benthamiana Gongylolepis bracteata Gongylolepis colombiana Gongylolepis cortesii Gongylolepis erioclada Gongylolepis fruticosa Gongylolepis glaberrima Gongylolepis huachamacari Gongylolepis jauaensis Gongylolepis martiana Gongylolepis oblanceolata Gongylolepis paniculata Gongylolepis paruana Gongylolepis pedunculata Goodeniaceae Guaicaia Guaicaia bilabiata Guariruma Gundelia Gundelieae Gypothamnium Gypothamnium pinifolium Harthamnus Harthamnus boliviensis Hecastocleideae Hecastocleidoideae Hecastocleis Hecastocleis shockleyi Hesperomannia Hesperomannia arborescens Heteranthus Heterolepis Hieracium glumaceum Hilairia Hipposeris Hochstetteria Hochstetteria schimperi Holocheilus Holocheilus brasiliensis Holocheilus fabrisii Holocheilus hieracioides


Holocheilus illustris Holocheilus monocephalus Holocheilus pinnatifidus Holocheilus schulzii Homanthis Homoeanthus Homoeanthus bellidifolius Homoeanthus calophyllus Homoianthus Homoianthus lyratus Homoianthus pungens Huarpea Hyalis Hyalis argentea Hyalis lancifolia Hyaloseris Hyaloseris andrade-limae Hyaloseris boliviensis Hyaloseris camataquiensis Hyaloseris cinerea Hyaloseris longicephala Hyaloseris quadriflora Hyaloseris rubicunda Ianthopappus Ianthopappus corymbosus Idicium Ingenhusia Ingenhusia radiata Isanthus Isotypus Isotypus onoseroides Jobaphes Jobaphes virgatus Jungeae Jungia Jungia axillaris Jungia beckii Jungia brachyphylla Jungia calyculata Jungia coarctata Jungia crenatifolia Jungia ferruginea Jungia fistulosa Jungia floribunda Jungia glandulifera Jungia gracilis Jungia hirsuta

707

Jungia karstenii Jungia mitis Jungia ovata Jungia paniculata Jungia pauciflora Jungia polita Jungia pringlei Jungia rugosa Jungia schuerae Jungia sect. Dumerilia Jungia sellowii Jungia sordida Jungia spectabilis Jungia stuebelii Jungia vitocensis Jungia weberbaueri Jungia woodii Jungieae Labiatiflorae Lactuceae Lasiopus Lasiopus ambiguus Lasiopus bojeri Lasiopus viridifolius var. hirsutus Lasiorrhiza Lasiorrhiza glacialis Lasiorrhiza leontopodioides Lasiorrhiza lithospermifolia Lasiorrhiza tomentosa Lavidia Lavidia caespitosa Lefrovia Lefrovia rhaponticoides Leibnitzia Leibnitzia anandria Leibnitzia knorringiana Leibnitzia nepalensis Leibnitzia occimadrensis Leibnitzia ruficoma Leibnitzia seemanii Leptica Leptica ciliata Leria Leria leiocarpa Leria media Leria nutans Leria nutans var. sinuata

708

Leria spathulata Leria stenocephala Lerieae Leucaeria Leuceria Leuceria anthemidifolia Leuceria foliosa Leuceria nivea Leuceria thrincioides Leuchaeria Leucheria Leucheria achillaeifolia Leucheria amoena Leucheria apiifolia Leucheria bridgesii Leucheria candidissima Leucheria cerberoana Leucheria coerulescens Leucheria congesta Leucheria cumingii Leucheria daucifolia Leucheria diemii Leucheria eriocephala Leucheria floribunda Leucheria garciana Leucheria gayana Leucheria gilliesii Leucheria glabriuscula Leucheria glacialis Leucheria glandulosa Leucheria hahnii Leucheria hieracioides Leucheria integrifolia Leucheria landbeckii Leucheria leontopodioides Leucheria lithospermifolia Leucheria magna Leucheria menana Leucheria millefolium Leucheria multiflora Leucheria nutans Leucheria oligocephala Leucheria paniculata Leucheria papillosa Leucheria polyclados Leucheria pteropogon Leucheria purpurea

L. Katinas, et al.

Leucheria rosea Leucheria runcinata Leucheria salina Leucheria scrobiculata Leucheria senecioides Leucheria stuebelii Leucheria suaveolens Leucheria tenuis Leucheria thermarum Leucheria tomentosa Leucheria viscida Leucomeris Leucomeris decora Leucomeris spectabilis Leukeria Leunisia Leunisia laeta Liabeae Liatris latifolia Lieberkuhna Lieberkuhna bracteata Lophopappus Lophopappus berberidifolius Lophopappus blakei Lophopappus cuneatus Lophopappus foliosus Lophopappus peruvianus Lophopappus tarapacanus Loxodon Loxodon brevipes Luciliopsis perpusilla Lulia Lulia nervosa Lycoseris Lycoseris boliviana Lycoseris colombiana Lycoseris crocata Lycoseris eggersii Lycoseris grandis Lycoseris mexicana Lycoseris minor Lycoseris peruviana Lycoseris retroflexa Lycoseris trinervis Lycoseris triplinervia Macledium Macledium burmanni


Macledium grandidieri Macledium spinosum Maclovia Macrachaenium Macrachaenium gracile Macrachaenium gracile var. radiatum Macroclinidium Macroclinidium hybridum Macroclinidium koribeanum Macroclinidium rigidulum Macroclinidium robustum Macroclinidium suzuki Macroclinidium trilobum Marticorenia Marticorenia foliosa Martrasia Martrasia auriculata Martrasia crenata Martrasia pubescente Martrasia sericea Mastigophorus Mastigophorus gaudichaudii Mimela Mimela pedicularifolia Minythodes Mocinia Mocinia mutisioides Moquinia Moquinia argentina Moquinia argyrea Moquinia curviflora Moquinia gardneri Moquinia hypoleuca Moquinia mollissima Moquinia oligocephala Moquinia orbiculata Moquinia racemosa Moquinia velutina Moscaria Moscharia Moscharia pinnatifida Moscharia solbrigii Moschifera Mosigia Mutisia Mutisia acerosa Mutisia acuminata

709

Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia Mutisia

alata andersonii araucana arequipensis brachyantha burkartii campanulata cana clematis coccinea cochabambensis comptoniaefolia decurrens discoidea friesiana grandiflora hamata hastata hieronymii homoeantha ilicifolia intermedia involucrata kurtzii lanata lanigera latifolia ledifolia lehmannii linearifolia linifolia lutzii macrophylla magnifica mandoniana mathewsii microcephala microneura microphylla ochroleuca oligodon orbignyana pulcherrima rauhii retrorsa rimbachii rosea

710

Mutisia saltensis Mutisia sect. Guariruma Mutisia sinuata Mutisia sodiroi Mutisia speciosa Mutisia spectabilis Mutisia spinosa Mutisia splendens Mutisia stuebelii Mutisia subspinosa Mutisia subulata Mutisia tridens Mutisia venusta Mutisia vicia Mutisia wurdackii Mutisiaceae Mutisieae Mutisinae Mutisiinae Mutisioideae Myripnois Myripnois dioica Myripnois uniflora Nassauvia Nassauvia aculeata Nassauvia aculeata var. aculeata Nassauvia ameghinoi Nassauvia argentea Nassauvia argyrophylla Nassauvia axillaris Nassauvia capillata Nassauvia chubutensis Nassauvia coronipappa Nassauvia cumingii Nassauvia darwinii Nassauvia dentata Nassauvia digitata Nassauvia dusenii Nassauvia fuegiana Nassauvia gaudichaudii Nassauvia glomerata Nassauvia glomerulosa Nassauvia hillii Nassauvia juniperina Nassauvia lagascae Nassauvia lagascae var. lagascae Nassauvia latissima

L. Katinas, et al.

Nassauvia looseri Nassauvia maeviae Nassauvia magellanica Nassauvia pentacaenoides Nassauvia pinnigera Nassauvia planifolia Nassauvia pulcherrima Nassauvia pygmaea Nassauvia pygmaea var. pygmaea Nassauvia pyramidalis Nassauvia ramosissima Nassauvia revoluta Nassauvia ruizii Nassauvia sceptrum Nassauvia sect. Caloptilium Nassauvia sect. Mastigophorus Nassauvia sect. Panargyrum Nassauvia serpens Nassauvia sprengelioides Nassauvia subgen. Strongyloma Nassauvia sublobata Nassauvia ulicina Nassauvia uniflora Nassauviaceae Nassauvieae Nassauvinae Nassauviinae Nassauvioideae Nassovia Neblinaea Neblinaea promontoriorum Neocaldasia Neocaldasia colombiana Neoshinneria Nitelium Nitelium rubescens Nothofagus Nouelia Nouelia insignis Oldenburgia Oldenburgia grandis Oldenburgia intermedia Oldenburgia papionum Oldenburgia paradoxa Oleaceae Oligocarpha Oligocarpha nereifolia


Onoseridae Onoserideae Onoseridinae Onoseris Onoseris acerifolia Onoseris alata Onoseris albicans Onoseris amplexicaulis Onoseris annua Onoseris brasiliensis Onoseris cabrerae Onoseris castelnaeana Onoseris chrysactinioides Onoseris costaricensis Onoseris cumingii Onoseris donnell-smithii Onoseris drakeana Onoseris fraterna Onoseris gnaphalioides Onoseris hastata Onoseris heterophylla Onoseris hieracioides Onoseris humboldtiana Onoseris hyssopifolia Onoseris linearifolia Onoseris lopezii Onoseris macbridei Onoseris minima Onoseris odorata Onoseris onoseroides Onoseris parva Onoseris peruviana Onoseris purpurea Onoseris sagittata Onoseris salicifolia Onoseris silvatica Onoseris subgen. Hipposeris Onoseris spathulata Onoseris speciosa Onoseris stricta Onoseris weberbaueri Oreoseris Oreoseris lanuginosa Oreoseris nivea Oriastrum Oriastrum cochlearifolium Oriastrum pentacaenoides

711

Oriastrum pulvinatum Oriastrum pusillum Oriastrum sphaeroidale Oxydon Oxydon bicolor Oxyphyllum Oxyphyllum ulicinum Pachylaena Pachylaena atriplicifolia Pachylaena elegans Pamphalea Panargyrum Panargyrum darwinii Panargyrum glomeratum Panargyrum heterophyllum Panargyrum lagascae Panargyrum uniflorum Panargyrus Panphalea Panphalea araucariophila Panphalea bupleurifolia Panphalea cardaminifolia Panphalea commersonii Panphalea heterophylla Panphalea maxima Panphalea missionum Panphalea ramboi Panphalea smithii Pardisium Pasaccardoa Pasaccardoa grantii Pentanthus Pentanthus aculeatus Pentaphorus Pentaphorus foliolosus Pentaphorus glutinosus Perdiceae Perdiciaceae Perdicieae Perdicium Perdicium brasiliense Perdicium cacalioides Perdicium capense Perdicium chilense Perdicium cordatum Perdicium divaricatum Perdicium lactucoides

712

Perdicium leiocarpum Perdicium magellanicum Perdicium piloselloides Perdicium purpureum Perdicium recurvatum Perdicium squarrosum Perdicium suaveolens Perezia Perezia aspera Perezia bellidifolia Perezia calophylla Perezia carduncelloides Perezia carpholepis Perezia carthamoides Perezia ciliaris Perezia ciliosa Perezia coerulescens Perezia collina Perezia coulteri Perezia cuernavacana Perezia delicata Perezia dissiticeps Perezia dugesii Perezia dugesii var. pilulosa Perezia erioloma Perezia eryngioides Perezia fonkii Perezia fruticosa Perezia glandulifera Perezia grandifolia Perezia hooveri Perezia kingii Perezia lactucoides Perezia lanigera Perezia lepidopoda Perezia linearis Perezia lobulata Perezia longifolia Perezia lozanii Perezia lyrata Perezia magellanica Perezia mandonii Perezia megalantha Perezia michoacana Perezia moschata Perezia multiflora Perezia nana

L. Katinas, et al.

Perezia nelsonii Perezia nudicaulis Perezia nudiuscula Perezia nutans Perezia oxylepis Perezia palmeri Perezia parryi Perezia patens Perezia pedicularidifolia Perezia pilifera Perezia pinetorum Perezia pinnatifida Perezia platyphylla Perezia platyptera Perezia poeppigii Perezia prenanthoides Perezia pringlei Perezia pungens Perezia purpurata Perezia purpusii Perezia pygmaea Perezia recurvata Perezia scapiformis Perezia scaposa Perezia simulata Perezia spathulata Perezia squarrosa Perezia sublyrata Perezia tomentosa Perezia turbinata Perezia umbratalis Perezia viscosa Perezia volcanensis Perezia wislizenii Perezia wrightii Pereziopsis Pereziopsis donnell-smithii Pertya Pertya aitchisoni Pertya angustifolia Pertya berberidioides Pertya bodinieri Pertya bodinieri var. berberidioides Pertya cordifolia Pertya corymbosa Pertya desmocephala Pertya discolor


Pertya glabrescens Pertya henanensis Pertya hossei Pertya hybrida Pertya macrophylla Pertya mattfeldii Pertya monocephala Pertya ovata Pertya phylicoides Pertya pungens Pertya scandens Pertya scandens f. schultziana Pertya sect. Macroclinidium Pertya simozawai Pertya sinensis Pertya tsoongiana Pertya uniflora Pertya yakjushimensis Pertyeae Pertyoideae Phyllactinia Phyllactinia grantii Piloselloides Piptostemma Platycheilus Platycheilus ochroleucus Plazia Plazia cheiranthifolia Plazia conferta Plazia daphnoides Pleiotaxis Pleiotaxis pulcherrima Pleocarphus Pleocarphus revolutus Poaceae Pogonura Polyachrus Polyachurus Polyachyreae Polyachyrinae Polyachyris Polyachyrus annuus Polyachyrus carduoides Polyachyrus cinereus Polyachyrus fuscus Polyachyrus gayi Polyachyrus poeppigii

713

Polyachyrus poeppigii var. poeppigii Polyachyrus sphaerocephalus Portalesia Portalesia procumbens Prionanthes Prionanthes antimenorrhoea Proselia Proselia serrata Proustia Proustia crassinervis Proustia cuneata Proustia cuneifolia Proustia ilicifolia Proustia pyrifolia Proustia reticulata Proustia vanillosma Pseudostifftia Ptilurus Ptilurus daucifolius Quelchia Quelchia bracteata Quelchia cardonae Quelchia conferta Quelchia eriocaulis Quelchia xgrandifolia Rhinactina Rhodoseris Rhodoseris conspicua Richterago Richterago campestris Richterago caulescens Richterago conduplicata Richterago elegans Richterago lanata Richterago petiolata Richterago riparia Salcedoa Salcedoa mirabaliarum Sanhilaria Schaetzellia Schaetzellia deckeri Schinopsis Schlechtendalia Scolymanthus Senecio pteropogon Seris

714

Seris amplexifolia Seris angustifolia Seris discoidea Seris onoseroides Seris polymorpha Seris polyphylla Seris sagittatus Spadonia Spadonia paniculata Spadonia polymorpha Spadonia polymorpha var. sordida Spadonia racemosa Sphaerocephalus Sphaerocephalus lagascae Sphagnum Stenopadus Stenopadus andicola Stenopadus aracaënsis Stenopadus campestris Stenopadus chimantensis Stenopadus cinereus Stenopadus colombianus Stenopadus colveei Stenopadus connellii Stenopadus crassifolius Stenopadus cucullatus Stenopadus cymbifolius Stenopadus huachamacari Stenopadus jauensis Stenopadus kunhardtii Stenopadus megacephalus Stenopadus obconicus Stenopadus sericeus Stenopadus subgen. Stomatochaeta Stenopadus talaumifolius Stenophyllum Stifftia Stifftia cayennensis Stifftia chrysantha Stifftia condensata Stifftia connellii Stifftia fruticosa Stifftia hatschbachii Stifftia martiana Stifftia parviflora Stifftia racemosa

L. Katinas, et al.

Stifftia uniflora Stifftieae Stomatochaeta Stomatochaeta acuminata Stomatochaeta colveei Stomatochaeta condensata Stomatochaeta crassifolia Stomatochaeta cylindrica Stomatochaeta cymbifolia Stomatochaeta steyermarkii Strongyloma Strongyloma axillare Strongylomopsis Strongylomopsis fuegiana Taraxacum officinale Tarchonantheae Tarchonanthus Tenorea Tenorea berteri Thyrsanthema Tiltilia Tostimontia Tostimontia gunnerifolia Triachne Triachne pygmaea Trianthus Trianthus ulicinus Trichocline Trichocline arenaria Trichocline aurea Trichocline auriculata Trichocline beckeri Trichocline boecheri Trichocline catharinensis Trichocline caulescens Trichocline cineraria Trichocline cordifolia Trichocline dealbata Trichocline denticulata Trichocline deserticola Trichocline exscapa Trichocline heterophylla Trichocline humilis Trichocline incana Trichocline linearifolia Trichocline macrocephala


Trichocline macrorhiza Trichocline maxima Trichocline nervosa Trichocline plicata Trichocline reptans Trichocline sect. Amblysperma Trichocline sinuata Trichocline spathulata Trichocline speciosa Trinacte Triptilion Triptilion achilleae Triptilion axillare Triptilion benaventii Triptilion berteroi Triptilion capillatum Triptilion cordifolium Triptilion gibbosum Triptilion glomerulosum Triptilion spinosum Triptilium Trixidea Trixideae Trixidinae Trixis Trixis aggregata Trixis alata Trixis angustifolia Trixis cacalioides Trixis calcicola Trixis californica Trixis calycina Trixis chiapensis Trixis divaricata Trixis divaricata var. divaricata Trixis eryngioides Trixis glaziovii Trixis glutinosa Trixis grandibracteata Trixis grisebachii Trixis haenkei Trixis hassleri Trixis hyposericea Trixis inula Trixis lessingii Trixis megalophylla

715

Trixis mexicana Trixis michuacana Trixis nelsonii Trixis nobilis Trixis ophiorhiza Trixis pallida Trixis parviflora Trixis peruviana Trixis pinnatifida Trixis praestans Trixis pringlei Trixis proustioides Trixis pruskii Trixis pterocaulis Trixis rugulosa Trixis sagasteguii Trixis schulzii Trixis silvatica Trixis spicata Trixis thyrsoidea Trixis vauthieri Trixis verbascifolia Tugarinovia Tussilago albicans Tussilago anandria Tussilago dentata Tussilago exscapa Tussilago integerrima Tussilago nutans Tussilago pumila Tylloma Tylloma glabratum Tylloma lanatum Tylloma limbatum Tylloma minutum Tylloma pusillum Uechtritzia Uechtritzia armena Uechtritzia kokanica Uechtritzia lacei Urmenetea Urmenetea atacamensis Vernonia Vernonieae Warionia Warionia saharae

716

Weberbauerella Weberbaueriella Weberbaueriella johnstoniana Wunderlichia Wunderlichia azulensis

L. Katinas, et al.

Wunderlichia Wunderlichia Wunderlichia Wunderlichia Wunderlichia

bahiensis crulsiana insignis mirabilis senaeii