New combinations in South American Myricaceae

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des, ~t 2900 m d'alt," Triana 825 p.p. (LECTO-. TYeE: here designated, P; ISOLECTOTYPES: MO,. NY, US). Myrica parvifolia vat. major A. Chev., Mem. Soc. Sci.
New combinations in South American Myricaceae CARLOS P A R R A - O .

Parra-O., C. (Instituto de Ciencias Naturales--Universidad Nacional de Colombia, Apartado Aereo 7495, Bogota, Colombia; email: [email protected]. edu.co). New combinations in South American Myricaceae. Brittonia 54: 322326. 2002.--New combinations and lectotypification are made of the South American species formerly of Myrica in the genus Morella. A key and a list of synonyms are presented. Key words: Myricaceae, Morella, Myrica, South America. Parra-O., C. (Instituto de Ciencias Naturales---Universidad Nacional de Colombia, Apartado Aereo 7495, Bogot~i, Colombia; email: [email protected]. edu.co). New combinations in South American Myricaceae. Brittonia 54: 322326. 2002.--Se presentan las nuevas combinaciones y las lectotipificaciones necesarias para transferir a Morella las especies suramericanas anteriormente asignadas a Myrica. Adem~is se incluyen una clave y una lista de sinrnimos.

Various problems o f nomenclature and typification o f the Myricaceae have arisen in recent years. In this article, I present a synthesis o f the nomenclature that includes the core genus of Myricaceae, Myrica L. (sensu Chevalier, 1901). The p r o b l e m began when the strongly different morphology led to the genus Myrica being separated into three m a j o r groups, which are well differentiated f r o m each other and are supported by different kinds of evidence (Baird, 1968; Macdonald, 1989; Wilbur, 1994). As Wilbur pointed out, the level assigned to these groups (i.e., whether three genera or one genus with three subgroups) is a decision that depends on " . . . both tradition and the prevailing philosophy o f the period." Traditionally, one of these groups (represented by M. asplenifolia L.) was separated long ago as the genus Comptonia L'Hrr. ex Aiton because it had pinnatifid leaves, stipules, and a burrlike fruit with 6 8 persistent bracteoles (Bornstein, 1997; Chevalier, 1901; Elias, 1971; Wilbur, 1994). The other groups have been treated as distinct taxonomic categories throughout their history. Authors such as Elias (1971) proposed two subgenera: subgen. Myrica (with two species, M. gale and M. hartwegii

S. Watson) and subgen. Morella (with ca. 40 species). Early authors such as Spach (1841) and Chevalier (1901) recognized three genera: Myrica L. ( 4 0 - 5 0 species), Gale D u h a m e l (two to three species), and Comptonia L'Hrr. ex Aiton (one species). After Chevalier (1901) published his " M o n o g r a p h i e des M y r i c a c r e s , " other authors (Greene, 1910a, 1910b; Nieuwland, 1910) tried to resolve the ranking of these taxa, but some changes were not formalized or the proposals were not adopted. Recent authors (Baird, 1968; Wilbur, 1994) listed several characters that support Spach's and Chevalier's criteria for considering the three groups as different genera. The characters that separate Myrica and Gale are principally the terminal buds (present vs. lacking), the vessel element distribution (diffuse porous vs. ring porous), the insertion o f aments (on old w o o d mainly below the leaves vs. at the s u m m i t of the branchelets o f the preceding year), the stamen n u m b e r (highly variable, to 22 per flower vs. consistently reduced, mostly 5 or less), the adnation of bracteoles to the fruit wall (none vs. strongly adnate), the fruit (drupelike with hard wall and covered with

Brittonia, 54(4), 2002, pp. 322-326. 9 2002, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

ISSUED: 16 April 2003

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papillae vs. nutlike with thin wall and without papillae), the wax production of the fruit (present vs. absent), and the chromos o m e number (n = 8 vs. n = 24, 48) (Baird, 1968; Wilbur, 1994). The crux of the nomenclatural p r o b l e m (which originated with the acceptance o f Myrica subgen. Myrica and Myrica subgen. Morella as two separate genera) has been the lectotypification o f the genus Myrica. I f M. cerifera were designated as the lectotype o f Myrica, then Myrica could be treated in the sense of Chevalier (1901), and M. gale and M. hartwegii should be segregated into Gale. However, if M. gale were designated as the lectotype of Myrica, then all the species belonging to subgen. Morella (most o f the species occurring in the Americas, Africa, eastern Asia, the Philippines, and Malaysia) would be included in Morella Lour. Small (1903) adopted Morella Lout. as the correct name for all the sSutheastern United States species of Myrica. Later criticism b y Greene (1910b) discouraged the use of this genus. Baird (1968) analyzed the p r o b l e m and proposed the suitable nomenclatural combinations with typification o f each one, supporting the use o f Morella (to accommodate all the species o f Myrica sensu Chevalier), but he never published these new combinations. Wilbur (1994) independently c a m e to the same conclusion as Baird and published the nomenclatural transferrals o f four species of Myrica f r o m North A m e r i c a and Canada to Morella. He proposed new names and combinations to define subgenera and series within Morella. This proposal was not followed by Bornstein (1997) in his Flora of North America. Verdcourt and Polhill (1997), unaware of Wilbur's paper, summarized the m o s t important points o f the problem and proposed the conservation of Myrica and Gale following the criteria o f Hylander (1945) and Rehder (1949), w h o considered Myrica cerifera to be the lectotype of Myrica and m a d e the transfer of Myrica gale and Myrica hartwegii to

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Gale. This proposal required that Myrica gale b e c o m e Gale belgica Dumort. to avoid tautonomy. Although some recent papers (e.g., Ferguson, 1998) supported the lectotypification of Myrica by M. cerifera, the proposal was rejected b y the Committee for Spermatophyta. The only species growing in Sweden (M. gale) was well known to Linnaeus and it seemed only right that that species should serve as the generic type (Killick et al., 1998; Brummitt, 1999). After this rejection, Killick et al. (1998) transferred the African species previously accepted by K_illick (1969) and White (1993) to Morella, making 13 new specific c o m binations and five infraspecific ones. It is evident that the remaining species o f Myrica should be transferred to Morella, which I do in this publication. A key to the species of Morella f r o m South America and their s y n o n y m y are included. At present m a n y publications continue using names that were already in s y n o n o m y since Chevalier (1901) [e.g., M. arguta, M. caracas-

sana, M. polycarpa (M. pubescens)]. I agree that all the South American species of Morella belong to subgen. Cerothamnus (Tidestr.) W i l b u r ser. Cerothanmus (Tidestr.) Wilbur (Wilbur, 1994). Morella rotundata (Steyermark & Maguire) C. Parra-O. is excluded f r o m this key because the species' description was based only on sterile specimens and I have seen no fertile specimens. This species has been poorly collected in the Chimant~i Massif in Venezuelan Guayana. Nevertheless, vegetative characteristics o f the isotype and paratype indicate that the species is distinct within Morella.

Morella Lour., F1. Cochinch. 458. 1790. TYPE: Morella rubra Lour.

Cerophora Raf., Alsog. Americana 11. 1838 (p:p.). Myrica L. subgen. Morella (Lour.) Engler, Nat. Pflanzenfam. III 1: 27. 1893.

Myrica sensu Chevalier, non L., Mem. Soc. Sci. Nat. Cherbourg 32: 286. 1901.

CerothamnusTidestr., Elys. Marian., Ferns. 41. 1910.

Key to the Species of Morella of South America 1. Plants monoecious; leaf blades lanceolate, pubescent beneath (generally), the base attenuate, the main, secondary, and/or tertiary veins beneath densely or sparsely pubescent; aments with staminate

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flowers on the lowest part of the rachis, pistillate flowers on the upper part; main bract of the pistillate flowers lanceolate or short-lanceolate, densely pubescent at apex, margin, and dorsal portion; fruits pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morella pubescens Plants dioecious (rarely monoecious); leaf blades variously shaped, glabrous or minutely and sparsely pubescent beneath, the base acute or cuneate, the main vein (and sometimes secondary veins beneath) densely or sparsely pubescent; aments with only staminate flowers or pistillate flowers; main bract o f pistillate flowers rhomboid, s o m e t i m e s lanceolate, sparsely pubescent at apex and margin (with shorter hairs s o m e t i m e s present on dorsal portion); fruits usually glabrous. 2. Leaves lanceolate-linear, 5 - 7 times longer than wide; staminate inflorescence with c o m p o u n d male flowers; in the Andes o f southern Peru and northern Chile .......................... Morella pavonis 2. Leaves of various shapes, 2 - 4 times longer than wide; staminate inflorescence with single male flowers; elsewhere in A n d e a n South America. 3. L e a v e s 3 - 4 t i m e s l o n g e r t h a n wide, elliptic-oblong or oblong; fruits ovoid . . . . . . . . . . . ............................................................................................................... Morella funckii 3. Leaves in a different proportion and shape; fruits spherical. 4. Young twigs usually pubescent; leaf blades obovate or elliptic-obovate, 3.1-9.1 • 1.33.8 cm, the apex obtuse or subobtuse, the margin dentate or dentate-serrate; s e c o n d a r y bracts in the pistillate flowers 4 - 6 ; s t a m i n a l c o l u m n 0 . 5 - 2 . 5 m m long ............ .............................................................................. Morella singularis 4. Young twigs pubescent or glabrous; leaf blades o f various shapes, 1.1-6.4 • 0 . 4 - 2 . 2 cm, the apex acute or subacute, the margin entire or serrate; secondary bracts in the pistillate flowers 2 - 4 ; staminal column 0.2-1.2 m m long 5. Leaf blades elliptic or narrowly elliptic, the margin entire or s o m e t i m e s serrate and then only in the distal 1/3, glabrous; staminate flowers with 2 ovoid or rhomboid secondary bracts; secondary bracts in the pistillate flowers 3 - 4 ; A n d e a n Venezuela, Colombia, Ecuador, and Peru, 2 0 0 0 - 4 0 0 0 m ........................... Morella parvifolia 5. Leaf blades obovate-lanceolate or obovate-oblong, the margin serrate (at least in the distal 2/3), s o m e t i m e s p u b e m l e n t in and/or between teeth; staminate flowers with or without s e c o n d a r y bracts, if present the bracts linear; pistillate flowers with 2 3(4) s e c o n d a r y bracts; A n d e a n B o l i v i a and A r g e n t i n a , 1 4 0 0 - 2 7 0 0 m ............ .................................................................................. Morella chevalieri

M o r e l l a funckii (A. Chev.) C. Parra-O., comb. nov. Myrica funckii A. Chev., M e m . Soc. Sci. Nat. Cherbourg 32: 286. 1901. TYPE: V E N E Z U E L A . M6rida: Culata, 8000 ft, Apr 1847, Funck & Schlim 1541 ( L E C T O T Y P E I here designated, P; ISOLECTOTYPE" G ) .

M o r e l l a p a r v i f o l i a (Benth.) C. Parra-O., comb. nov. Myrica parvifolia Benth., P1. Hartweg. 251. 1846. TYPE: E C U A D O R . Monte de Tiopullo dicto prope Quito, H a r t w e g 1378 [HOLOTYPE: K; photos: COL,' M O , US (F negative no. 11552)]. Myrica parvifolia var. obtusa Benth., Pl. Hartweg 25l. 1846. TYPE: C O L O M B I A : prope Bogota, Hartweg 1379 (HOLOTYPE:K; photo: COL). Myrica parvifolia var. longipedunculata A. Chev., M e m . Soc. Sci. Nat. Cherbourg 32: 284. 1901. TYPE: C O L O M B I A . Boyac~i: "entre Tunja et Paypa, ~ 1400 poises d'alt," M a r 1843, Linden 1305 [LECTOTVPE: here designated, P; photo: M O (F negative no. 11581)].

In the original description o f Myrica parvar. longipedunculata, Chevalier

vifolia

(1901) cited two specimens o f Linden 1305, one o f t h e m deposited in " H . M u s . " (P), the other in " H . Bruxelles" (BR). The duplicate at B R has not been found there (R Stoffelen, pers. comm.). Myrica parvifolia vat. lucens A. Chev., M e m . Soc. Sci. Nat. Cherbourg 32: 285. 1901. TYPE: COL O M B I A . Prov. Bogot~i, 1844, Goudot s.n. (LEC-

TOTYPE: here designated, P). Myrica parvifolia vat. macrostachya A. Chev., M e m . Soc. Sci. Nat. Cherbourg 32: 284. 1901. TYPE: C O L O M B I A . "Prov. Bogot~i: dans les Andes, ~t 2900 m d'alt," Triana 825 p.p. (LECTOTYeE: here designated, P; ISOLECTOTYPES: MO, NY, US). Myrica parvifolia vat. major A. Chev., Mem. Soc. Sci. Nat. Cherbourg 32: 286. 1901. TYPE: ECUADOR. "Andes," 1861, Spruce 5135 (rlOLOTYPE: P).

M o r e l l a pavonis (C. DC.) C. Parra-O., comb. nov. Myrica pavonis C. DC., Prodr. 26 (2): 151. 1868. TYPE: PERU: prope Huayaquil, Ruiz & Pavdn s.n. [LECTOTYPE: here designated, G; photos: COL, MO, US (F negative no. 8538)]. Myrica pavonis var. glandulosa A. Chev., Mem. Soc. Sci. Nat. Cherbourg. 32: 288. 1901. TYPE:

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PERU: Ruiz & Pavdn s . n . [LECTOTYPE:here designated, P; ISOLECTOTYPE:BM (not found)]. In the o r i g i n a l d e s c r i p t i o n o f M y r i c a p a v o n i s , de C a n d o l l e (1864) c it e d t w o s p e c i m e n s in the p r o t o l o g u e : " I n P e r u v i ~ p r o p e H u a y a q u i l (herb. P a v o n in h. B o i s s . ) " a n d " S a n t a p r o p e L i m a (herb. K e w . S i m n. 8 8 ) . " I d e s i g n a t e th e s p e c i m e n at G as th e lectotype of M. p a v o n i s for a few reasons: the s p e c i m e n " S i m n. 8 8 " w a s n o t f o u n d at K (S. O w e n s , pers. c o m m . ) ; the sheet d e p o s i t e d in G is an e x c e l l e n t s a m p l e s h o w i n g all the d i a g n o s t i c c h a r a c t e r s o f the s p e c i e s ; a n d G is the d e p o s i t o r y o f de C a n d o l l e ' s herbarium. T h e label o f the l e c t o t y p e e v i d e n t l y c o n tains a g e o g r a p h i c a l error. T h e o r i g i n a l labe l says: " M y r i c a s a l i c i f o l i a de H u a y a q u i l . 1785. Peru. F l o r M a s c u l i n a . " H o w e v e r , it is l i k e l y that the l a b e l i n c o r r e c t l y identifies " H u a y a q u i l " as the c o l l e c t i o n l o c a l e a n d that the l e c t o t y p e c o m e s f r o m P e r u and n o t f r o m Ecuador. T h i s is s u p p o r t e d by the f o l l o w i n g facts. T h o u g h R u i z and P a v r n h a d w a n t e d to c o l l e c t in G u a y a q u i l a n d n e a r b y p r o v i n c e s i n E c u a d o r , v a r i o u s factors p r e vented them from visiting these localities (Steele, 1964). F u r t h e r m o r e , at the t i m e this s p e c i m e n was c o l l e c t e d , R u i z and P a v r n w e r e in Peru, n e a r H m i n u c o and a d j a c e n t areas (Steele, 1964). B a s e d on a v a i l a b l e e v i d e n c e , there are n o h e r b a r i u m s p e c i m e n s o f M o r e l l a f r o m G u a y a q u i l , E c u a d o r (inc l u d i n g n e a r b y areas), or f r o m z o n e s n e a r sea l e v e l ( L a e g a a r d , 1999). M o r e l l a p a v o n i s is k n o w n to g r o w o n l y in c e n tr a l and southern P e r u and in n o r t h e r n Chile, b e t w e e n 1000 and 3 2 0 0 m.

Morella

pubescens

(Humb. & Bonpl. ex

W i l l d . ) Wilbur, R h o d o r a 103: 121. 2001. Myrica pubescens Humboldt & Bonpland ex Will-

denow, Sp. P1. 4: 746. 1806. TYPE: "Habitat in Nova Granada," Humboldt & Bonpland s.n. [HOLOTYPE: herb. Willdenow in B; photos: NY, US (F negative no. 11553)]. Myrica arguta Kunth, Nov. Gen. Sp., 2: 17, t. 98. 1817. TYPE: COLOMBIA. "Crescit in regno Novogranatensi in Paramo de San Fortunato, juxta viam Fusagasugensem, alt. 1480 hexap. Floret Septembri," Humboldt & Bonpland s.n. [HOLOTYPE: P (n.v.); microfiche: NY; photo: NY (F negative no. 39603)]. Myrica caracasana Kunth, Nov. Gen. Sp., 2: 18. 1817. TYPE: VENEZUELA. "Crescit in alsis Pro-

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vinciae Venezuelae, in monte Silla de Caracas, alt. 1200 hex., Floret Januario," Humboldt & Bonpland s.n. [HOLOTYPE:P (n.v.); photo: US (F negative no. 39605); microfiche: NY]. Myrica pubescens var. caracasana (Kunth) A. Chev., Mem. Soc. Sci. Nat. Cherbourg 32: 292. 1901. Myrica macrocarpa Kunth, Nov. Gen. Sp. 2: 16. 1817. TYPE: PERU. "Crescit in temperatis regni Peruviani prope Ayavaca, alt. 1370 hexap." Humboldt & Bonpland s . n . (HOLOTYPE:P; microfiche: NY). Myrica polycarpa Kunth, Nov. Gen. Sp. 2:18.1817. TYPE: "Crescit cum Myrica arguta in regno Novogranatensi in Paramo de San Fortunato, juxta viam Fusagasugensem, alt. 1480 hex, floret Septembri," Humboldt & Bonpland s.n. [HOLOTYPE: P (n.v.); microfiche: NY]. Myrica interrupta Benth., P1. Hartweg. 251. 1846. TYPE: COLOMBIA: "Juxta cataratam Tequindama, prov. Bogota," Hartweg 1380 (HOLOTYPE:K; photos: COL, NY, US (F negative no. 11554)]. Myrica pubescens var. interrupta (Benth.) A. Chev., Mem. Soc. Sci. Nat. Cherbourg 32: 294.

1901. Myrica arguta var. peruviana C. DC., Prodr. 16(2): 153. 1868. TYPE: PERU. Pavon s . n . (HOLOTYPE:

G; photo: COL). Myrica arguta var. tinctoria C. DC., Prodr. 16(2): 153. 1868. TYPE: PERU. Pavon s.n. (LECTOTYPE:

here designated, G; photo: COL; ISOLECTOTYPE: P). Myrica pubescens vat. glandulosa A. Chev., Mem.

Soc. Sci. Nat. Cherbourg. 32: 293. 1901. TYPE: PERU: Hartweg 832 (LECTOTYPE: here designated, P; ISOLECTOTYPE:NY). Myrica pubescens var. tomentosa A. Chev., Mem. Soc. Sci. Nat. Cherbourg 32: 294. 1901. TYPE: PERU: "Rivrro-Canta" (LECTOTVPE: here des-

ignated, P). Myrica costata Rusby, Descr. S. Amer. P1.8. 1920.

TYPE: BOLIVIA. La Paz. Prov. Noryungas: Unduavi, Buchtien 2810 (HOLOTYPE:NY; ISOTYPES: NY, US).

Morella rotundata (Steyermark & Maguire) C. Parra-O., co m b . nov. Myrica rotundata Steyermark & Maguire, Mem.

New York Bot. Gard. 17 (1): 441. 1967. TYPE: VENEZUELA. Bolfvar: Apacar~i-tepui, Chimantzl Massif, Steyermark 75084 [HOLOTYPE: VEN (n.v.); tSOTYPE:NY].

M o r e l l a s i n g u l a r i s (C. P a r r a - O . ) C. ParraO., c o m b . nov. Alchornea cerifera Croizat, Caldasia 2: 128. 1943.

TYPE: COLOMBIA. Putumayo: Lado sur de la Laguna de la Cocha, Cuatrecasas 11824 (HOLOTYPE: US; ISOTYPE: C O L ) . Myrica singularis C. Parra-O., Caldasia 21: 5. 1999. nom. nov. for A1chornea cerifera Croizat, non Myrica cerifera L., Sp. P1. 2: 1024. 1753.

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Bornstein, A, 1997. Myricaceae. Pages 429-435. In: M o r e l l a c h e v a l i e r i C. Parra-O.,

Brittonia 5 2 : 3 2 1 . 2 0 0 0 . TYPE: B O L I V I A . Santa Cruz: Prov. Florida, 4 km W of center of Samaipata, 18~ 63~ 20002 1 0 0 m , 31 D e c 1 9 9 2 (fr), M . N e e & L V a r g a s 4 3 4 3 6 [HOLOTYPE: N Y ; ISOTYPES: AD (n.v.), MO, LPB (n.v.), SI (n.v,),

USZ (n.v.)]. Myrica pubescens var. glabra A. Chev., Mere. Soc. Sci. Nat. Cherbourg 32: 293. 1901. TYPE: PERU: "Andes de l'interior," 1836-1837, Mathews 939 (HOLOTYPE: P).

Acknowledgments I thank the fellowship program o f the Andrew W. Mellon Foundation for financing my research as a visiting student at The N e w York Botanical Garden from July 1999 to January 2000. I thank the staff at The N e w York Botanical Garden for allowing access to the herbarium and the library, and for providing the necessary facilities for completing this investigation. Thanks to D. Lentz ( N Y ) and M. N e e ( N Y ) for their c o n t i n u o u s s u p p o r t i n a l l a s p e c t s r e l a t e d to t h i s s t u d y . T h a n k s t o S. O w e n s a n d C. B a r k e r ( K ) , P. M o r a t a n d M . P i g n a l (P), M . Baessler (B), E Jacquemoud ( G ) , P. S t o f felen and E. Robbrecht (BR), and C. Jarvis and R. Vickery (BM) for loans, photographs, and information concerning the types. Thanks to the curators of the herbaria at COL, MO, and US for providing access to their collections, and to Instituto de Ciencias Naturales of the Universidad Nacional de Colombia for administrative support. Thanks to D. Verdcourt (K) for sending me a copy of his article and commentaries on the nomenclature changes of the African species of M o r e l l a . I am also indebted to A. Macdonald, Lakehead University, Can a d a ; A'. B o r n s t e i n ( S E M O ) ; a n d a n a n o n ymous reviewer for their valuable comments on the manuscript.

Literature Cited Baird, J. R. 1968. A taxonomic revision of the plant family Myricaceae of North America, north of Mexico. Unpubl. Ph.D. dissertation. University of North Carolina, Chapel Hill.

Flora of North America Editorial Committee, editors. Flora of North America, North of Mexico. Vol. 3. Oxford University Press, New York. Brummitt, R. K. 1999. Proposals to conserve or reject. Report of the Committee for Spermatophyta. Taxon 48: 367. Candolle, A. C. de. 1864. Myricaceae. Pages 147155. In: Prodromus Systematis Naturalis Regni Vegetabilis. Vol. 16: 147-155. Chevalier, A. 1901. Monographie des Myricacres. Mrm. Soc. Sci. Nat. Cherbourg 32: 85-341. Elias, T. 1971. The genera of Myricaceae in the Southeastern United States. J. Arnold Arbor. 52: 305-318. Ferguson, D. K. 1998. The contribution of micromorphology to the taxonomy and fossil record of the Myricaceae. Taxon 47: 333-335. Greene, E. L. 1910a. Nomenclature of the bayberries. Lear. Bot. Observ. Crit. 2: 37-40. - - . 1910b. A new name for the bayberries. Lear. Bot. Observ. Crit. 2: 101-104. Hylander, N. 1945. Nomenklatorische und systematische Studien fiber Nordische Gef~isspflanzen. Uppsala Univ. Arsskr. 7: 337. Killick, D. J. B. 1969. The South African species of Myrica. Bothalia 10: 5-17. - R. M. Polhill & B. Verdcourt. 1998. New combinations in African Myricaceae. Kew Bull. 53: 993-995. Laegaard, S. 1999. Myricaceae. Page 611. In: P. M. Jorgensen & S. Le6n-Yafiez, editors. Catalogue of the Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard., Vol. 75. Missouri Botanical Garden Press, St. Louis, Missouri. Macdonald, A. 1989. The morphology and relationships of the Myricaceae. Pages 147-165. In: P. R. Crane & S. Blackmore, editors. Evolution, Systematics, and Fossil History of the Hamamelidae. Vol. 2: "Higher" Hamamelidae. Systematics Association Special Volume No. 40 B. The Systematics Association, Claredon Press, Oxford. Nieuwland, J. A. 1910. The name of our American wax bayberries. Amer. Midl. Naturalist 1: 238-243. Rehder, A. 1949. Bibliography of cultivated trees and shrubs. Arnold Arboretum, Harvard University, Jamaica Plain, Massachusetts. Small, J. 1903. Flora of the Southeastern United States. Published by the author, New York. Spach, E. 1841. Hi stoire Naturelle des Vrgrtaux. Phanrrogames. Vol. 11. Parig. Steele, A. R. 1964. Flowers for the King; the Expedition of Ruiz and Pavdn and the Flora of Peru. Duke University Press, Durham, NC. Verdeourt, B. & R. Polhill. 1997. Proposals to conserve the names Myrica and Gale (Myricaceae) with conserved types. Taxon 46: 347-348. White, F. 1993. African Myricaceae and the history of the Afromontane flora. Opera Bot. 121: 173-188. Wilbur, R. L 1994. The Myricaceae of the United States and Canada: genera, subgenera and series. Sida 16: 93-107.