ASEF 4-2008.indb - Laboratoire de zoologie

Ann. soc. entomol. Fr. (n.s.), 2008, 44 (4) : 459-470

ARTICLE

Paramesopsocidae, a new Mesozoic psocid family (Insecta: Psocodea “Psocoptera”: Psocomorpha) Dany Azar (1)*, Lara Hajar (2), Chadi Indary (3) & André Nel (4)* (1)

(2)

Lebanese University, Faculty of Sciences II, Department of Biology, Fanar - Matn - P. O. box 26110217, Lebanon Equipe Paléoenvironnements et Paléoclimats, Institut des Sciences de l’Evolution de Montpellier, Université Montpellier II, UMR 5554, Place Eugène Bataillon, F-34095 Montpellier Cedex 5, France (3) Jbeil, Kartaboun St., Diab blg, 4th flour, Lebanon (4) UMR 5202, Entomologie, CP 50, Muséum national d’Histoire naturelle, 45 rue Buffon. F-75005 Paris, France * Authors for correspondence

Abstract. Paramesopsocus lu n. gen., n. sp. and Paramesopsocus adibi n. sp. are respectively described from the Early Cretaceous amber of Lebanon and from the Late Jurassic limestone of Karatau (Kazakhstan). They are placed within the suborder Psocomorpha, and in the Mesozoic extinct family Paramesopsocidae n. fam. A cladistic phylogeny for Psocomorpha is given including our fossil taxa. The discovery of these new taxa demonstrates the necessity of a deep cladistic redefinition of the currently admitted major subdivisions of this suborder. Résumé. Paramesopsocidae, a new Mesozoic psocid family (Insecta : Psocodea “Psocoptera” : Psocomorpha). Paramesopsocus lu n. gen., n. sp. et Paramesopsocus adibi n. sp. sont décrits respectivement de l’ambre crétacé inférieur du Liban et du paléolac jurassique supérieur de Karatau (Kazakhstan). Ils sont placés dans le sous-ordre Psocomorpha, et dans la famille mésozoïque Paramesopsocidae n. fam. Une analyse phylogénétique cladistique des Psocomorpha est proposée, incluant nos taxons fossiles. La découverte de ces nouveaux taxons confirme la nécessitée d’une redéfinition phylogénétique approfondie des subdivisions majeures actuellement admises pour ce sous-ordre.

Keywords: Amber, Early Cretaceous, Late Jurassic, Lebanon, Kazakhstan.

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ecent cladistic analyses demonstrate the polyphyletic nature of several groups of insects, one of the most remarkable being the lice (Phthiraptera), which is now included within the Psocoptera to constitute the order Psocodea (Grimaldi & Engel 2005, 2006a, Yoshizawa & Johnson 2006, Johnson et al. 2004). Although we totally agree with the new data of the phylogeny of “Psocoptera”, we are considering for convenience in the following the “Psocoptera” (= psocids) in the traditional sense, i.e. without including the “Phthiraptera” (= lice), except when using the term Psocodea. Psocodea are a relatively small order with about 5,500 described recent psocid species (Lienhard & Smithers 2002; Lienhard 2003), and about 5,000 lice species (Grimaldi & Engel 2005). The earliest definitive phtirapteran is from the Eocene oil shales of Eckfeld maar near Manderscheid, Germany (Wappler et al. 2004). The oldest records of psocid-like Paraneoptera

E-mail: [email protected], [email protected] Accepté le 29 mai 2008

are as old as the Permian of Kansas (USA), C.I.S. and New South Wales (Australia) but also the Triassic of Issikkul (C.I.S.) (Carpenter 1992). The oldest definite psocids are recorded from the Middle Jurassic of China (Huang et al. 2008), the Upper Jurassic of Germany and Kazakhstan, the Late Jurassic (or Early Cretaceous?) of Western Liaoning in China (Lin 1976), and the Cretaceous amber from Lebanon (D. Azar, unpub. doctoral thesis, Univ. Paris-Sud, 2000; Poinar & Milki 2001; Perrichot et al. 2003; Azar & Nel 2004), France (Perrichot et al. 2003), Myanmar (Cockerell 1916, 1919; Nel & Waller 2007), Canada (Spahr 1992), Siberia (Taymir Peninsula) (Vishnyakova 1975), USA (New Jersey) (Gelhaus & Johnson 1996), and Spain (Alava) (Baz & Ortuño 2000, 2001a,b). Recently six new taxa of Psocoptera from the Lebanese amber have been studied (Perrichot et al., 2003 Azar & Nel 2004, Grimaldi & Engel 2006b). The taxa described from the amber are sufficiently well preserved in detail for correct comparisons with the recent psocids. Unfortunately the study of psocids preserved in limestone is mainly based on the wing venations, and when possible on some details preserved on legs and head.

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D. Azar, L. Hajar, C. Indary & A. Nel

There are very few attempts of phylogenetic analyses of the Psocoptera. Smithers (1972) dedicated a large part of his work “The classification and Phylogeny of Psocoptera” to the study of the phylogenetic relationships within Psocoptera, but his work is typological. Smithers (1972) proposed dendrograms with several lineages based on homoplastic or pleisiomorphic characters. Several years afterward the same author (Smithers 1991) admitted that his phylogeny needed revision. Perrichot et al. (2003) presented a tentative of cladistic phylogeny for Trogiomorpha, nevertheless this later phylogeny is incomplete because based on few characters. Johnson et al. (2004), Grimaldi & Engel (2006a), and Yoshizawa (2002) proposed molecular

and morphological phylogenetic analyses; the last one concerned the Psocomorpha only. We describe one new genus with two new species, viz. Paramesopsocus lu from the Early Cretaceous amber of Lebanon, and Paramesopsocus adibi from the Jurassic limestone of Karatau (Kazakhstan). Both are placed into the Psocomorpha, and in the Mesozoic new family Paramesopsocidae. This new fossil is of interest because it presents several characters that are currently considered as ‘characteristic’ of the Troctomorpha. Material and methods We follow the works of Smithers (1972, 1990) and Mockford (1993) as essential tools for the systematic of the psocids. We follow the wing venation nomenclature and body structures of

Figures 1–4 Paramesopsocus lu n. gen., n. sp. 1, Photograph of the holotype n. 746 C, male, dorsal view; 2, idem, ventral view; 3, drawing of the same specimen, dorsal view, scale bar = 1 mm; 4, photograph of the paratype n. 422, male, dorsal view.

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A new Mesozoic psocid family

Smithers (1972), and Lienhard (1998). The fossils in amber were carefully prepared in Canada balsam medium, following the method described by Azar et al. (2003), in order to observe as many characters as possible. Thus the ‘absences’ of structures are accurate, which is different of structures that are ‘not visible’ but maybe present.

Suborder Psocomorpha Family Paramesopsocidae n. fam.

and two anal veins. Hind wing with closed middle cell and M forked into M1 and M2.

Genus Paramesopsocus n. gen. Type species. Paramesopsocus lu n. gen., n. sp.

Other species. Paramesopsocus adibi n. sp. Etymology. Named after the close similarities in forewing venation of the recent genus Mesopsocus. Feminine gender. Diagnosis. As for the family.

Type species. Paramesopsocus lu n. gen., n. sp.

Etymology. Named after the close affinities with the extant family Mesopsocidae. Diagnosis. Antenna with 13 flagellomeres with fine secondary annulations. Legs with three-segmented tarsi, tarsal claws with one preapical tooth, no pulvillus. Wings glabrous and hyaline. Forewing with thickened and sclerotized pterostigma bearing a nodus at its base; free areola postica; hooked nodulus present;

Paramesopsocus lu n. sp. (Figs 1–19) Material. Holotype specimen n. 746 C (male) (figs 1–3), paratypes specimens n.: 422 (fig. 4), 427 A (figs 5, 6) and 886 (fig. 7, 8) (all males), coll. Azar, provisionally deposited in the Muséum National d’Histoire Naturelle, Paris.

Figures 5–8 Paramesopsocus lu n. gen., n. sp. 5, Photograph of the paratype n. 427 A, male, dorsal view; 6, idem, ventral view; 7, Photograph of the paratype n. 886, male, dorsal view; 8, idem, ventral view.

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Figures 9–14 Paramesopsocus lu n. gen., n. sp. 9, Photograph of the head of the holotype n. 746 C, lateral view, arrow indicates the lacinia; 10, drawing of the same specimen, lateral view, scale bar = 0.3 mm; 11, photograph of the head of the same specimen, frontal view; 12, drawing of the head of the same specimen, frontal view, scale bar = 0.3 mm; 13, photograph of the head of the paratype n. 427 A, dorsal view; 14, drawing of the antenna of the holotype n. 746 C, scale bar = 0.1 mm.

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Type locality and horizon. Early Cretaceous, Barremian – Lower Aptian, Hammana/Mdeirij, Caza Baada, Mouhafazit Jabal Libnen, Lebanon. Etymology. Named after L. U. initials of Lebanese University, the place where this study was conducted. Diagnosis. The only visible difference with P. adibi is the forewing with Rs and M fused for a short distance near their bases. Description. Head elongated, presenting color ornamentation (figs 9–13). Antenna with 15 segments (13 flagellomeres),

flagellomeres finely secondary annulated, all filiform, elongate, shortest flagellomere 0.046 mm long, longest 0.1 mm long (fig. 14); pedicel 0.064 mm long, 0.032 mm wide; scape 0.057 mm long, 0.038 mm wide. Anteclypeus 0.025 mm long, 0.22 mm wide; postclypeus 0.14 mm long, 0.25 mm wide (figs 9–12). Compound eyes slightly flattened with a large diameter of 0.2 mm and a small diameter of 0.16 mm. Three large ocelli disposed in triangle between compound eyes. Lacinia with a small tooth in inner side at basal third, and a round apex with two bulbs in inner side and a bulb in outer side (fig. 15). Maxillary palps four-segmented, second maxillary palpomere

Figures 15–19 Paramesopsocus lu n. gen., n. sp. 15, drawing of the lacinia of the holotype n. 746 C, scale bar = 0.05 mm; 16, drawing of the forewing of the same specimen, scale bar = 0.5 mm; 17, drawing of the hindwing of the same specimen, scale bar = 0.5 mm; 18, photograph of the male genitalia of the paratype n. 886; 19, drawing of the male genitalia of the holotype n. 746 C, scale bar = 0.3 mm.

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mx2 longer than mx4; mx1 0.019 mm long, 0.0032 mm wide; mx2 0.096 mm long, 0.0032 mm wide; mx3 0.0038 mm long, 0.002 mm wide; mx4 0.089 mm long, 0.0032 mm wide (fig. 10). Labial palps two-segmented. Thorax 0.55 mm long, 0.52 mm wide; mesothorax nearly triangular. Legs (measurements made on one of the holotype midlegs) with three-segmented tarsi, first tarsomere is much longer than others, 0.3 mm long, second tarsomere 0.046 mm long, and third one 0.053 mm long. Distal tarsomeres bearing claws with one preapical tooth. Forewing glabrous and hyaline, 2.37 mm long, 0.79 mm wide (fig. 16). Pterostigma thickened and sclerotized. Sc short, distally curved, and reaching R, then fused with it for 0.74 mm till 1.19 mm from wing base. Distal free part of Sc (Sc’ sensu Lienhard 1998) making a sharp angle with costal margin and reaching it 1.29 mm from wing base; R1 simple, reaching costal margin with a sharp angle, 1.81 mm from wing base. Rs basally oblique, branching slightly more distally with M; common portion Rs + M 0.038 mm long, at 1.15 mm from wing base; fork of R2+3 and R4+5 1.62 mm distal of wing base; R2+3 and R4+5 nearly straight; reaching wing margin respectively at 2.09 and 2.27 mm from wing base. Radial cell closed, longer than broad, five-angled. M basally fused with Cu1; fork between M1+2 and M3 1.71 mm distal of wing base; M1 and M2 separating 1.87 mm distal of wing base; M1 straight, reaching wing apex, 0.45 mm long; M2 straight, 0.34 mm long; M3 nearly straight, 0.31 mm long. Fork of Cu1 in Cu1a and Cu1b 1.38 mm from wing base; Cu1a strongly curved and longer than Cu1b. Areola postica (AP) free; no cross-vein between AP cell and M. Cu2 weaker than other veins, 0.62 mm long; a distinct nodulus. 1A straight, 0.90 mm long; 2A visible, 0.31 mm long. Hind wing glabrous and hyaline, smaller than forewing, 1.73 mm long, 0.54 mm wide (fig. 17). Sc short, 0.18 mm long, not reaching anterior wing margin; Rs distally fused with M for 0.12 mm. R1 reaching anterior wing margin 1.14 mm from wing base. Fork of Rs into R2+3 and R4+5 1.33 mm from wing base. Basi-radial cell four-angled; R2+3 0.21 mm long; R4+5 0.42 mm long. Fork of M into M1 and M2 1.13 mm from wing base; M1 0.42 mm long; M2 0.23 mm long; M1 nearly twice as long as M2. Cu1 and Cu2 reaching posterior wing margin respectively at 0.97 mm and 0.73 mm from wing base; anal vein (1A) forking into 1Aa and 1Ab 0.14 mm from wing base. Abdomen 1.22 mm long, 0.64 mm wide. Male with triangular hypandrium 0.51 mm long, 0.48 mm wide at base; parameres (visible on specimen 886) triangular-shaped, elongated and curved dorsally, 0.17 mm long, 0.07 mm large basally (figs 18, 19).

Comment. According to the keys to the families of Lienhard (1998), Paramesopsocus lu n. gen., n. sp. would fall in the Psocomorpha because of the following characters: adult with tarsi three-segmented; forewing with thickened and sclerotized pterostigma; no scales. After the keys of Mockford (1993), P. lu has some psocomorphan characters such as ‘forewing with thickened and sclerotized pterostigma’. But it has also some characters of the suborder Troctomorpha, viz. antenna with 13 flagellomeres; adult with tarsi three-

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segmented; secondary annulations on flagellomeres. Thus its position is problematic. The character ‘secondary annulations on flagellomeres’ only present in recent Troctomorpha, is also present in the very inclusive Jurassic Archipsyllidae. It is very probably a plesiomorphy (Huang et al. 2008). Recent Psocomorpha have 11 flagellomeres or fewer, while Troctomorpha usually have 13 but sometime fewer (Mockford 1993). The presence of only 11 flagellomeres is probably a plesiomorphy, as the Archipsyllidae have 11 flagellomeres and are in a very inclusive position in the Psocodea (Huang et al. 2008). The number of flagellomeres is not sufficient to assign this fossil taxon to one or the other of the two suborders, because this character seems to be homoplastic. The character ‘adult with tarsi three-segmented’ is present in psocids of the two suborders. The absence of pulvillus is a character of Troctomorpha but also of some Psocomorpha. Yoshizawa (2002) considered it as highly homoplastic. The ‘pretarsal claws with preapical tooth’ is also a character present in Troctomorpha, but plesiomorphic after Yoshizawa (2002). Other characters present in Troctomorpha are also shared by some Psocomorpha, i.e. ‘hind wing with M vein two-branched’ (probably a plesiomorphy as it is present in the Archipsyllidae and the Hemipsocidae, psocomorphan family having an inclusive position in Yoshizawa’s phylogeny); ‘forewing with nodulus and two anal veins’; ‘wings glabrous and hyaline’. If we neglect the character ‘thickened and sclerotized pterostigma’ (characteristic of the Psocomorpha) in the key to recent families of Smithers (1990), P. lu would fall near the electrentomoid family Compsocidae Mockford 1967 for the hind wing M forked and the forewing with nodulus (see also Mockford 1967; Lienhard & Smithers 2002). This family comprises two recent genera Compsocus Banks 1930 and Electrentomopsis Mockford 1967, and one fossil genus Burmacompsocus Nel & Waller 2007. Compsocus has 14 flagellomeres while Electrentomopsis and Burmacompsocus have only 13 as in P. lu. Electrentomopsis and Burmacompsocus differ from P. lu in the absence of first segment of Rs in hind wing. Nevertheless all Compsocidae have hyaline and not sclerotized pterostigma, unlike P. lu. The characters ‘thickened and sclerotized pterostigma’, ‘pterostigma with nodus’, and ‘hooked nodulus’ strongly support an attribution to the Psocomorpha, as these characters are considered as synapomorphies of this suborder (Mockford 1967; Yoshizawa 2002). According to Lienhard (1998), P. lu would fall in the psocomorphan family Mesopsocidae because of


Figures 20–25 Paramesopsocus adibi n. sp. 20, Photograph of the holotype specimen n. 2784/2096; 21, drawing of habitus of the holotype specimen n. 2784/2096, scale bar = 1 mm; 22, photograph of the paratype specimen n. 2066/3589; 23, photograph of the paratype specimen n. 2066/3602; 24, photograph of the paratype specimen n. 2066/3605; 25, photograph of the paratype specimen n. 2384/286.

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the following features: macropterous insect with threesegmented tarsi; thick pterostigma; areola postica free; wings glabrous; nodulus present. After the keys of the recent psocids of Smithers (1990), P. lu falls in the same family for the following characters: macropterous; legs with tarsi three-segmented; absence of scales; wing venation complex (venation not reduced to some parallel veins); sclerotized pterostigma; free areola postica with Cu1a and Cu1b separating near costal margin; wings glabrous; nodulus present. However, P. lu has some characters not shared by Mesopsocidae such as: absence of pulvilli; flagellomeres with secondary annulations (this character is normally proper to the Troctomorpha); presence of two anal veins; fork of M into M1 and M2 in hind wing (this character is present in many Troctomorpha and Trogiomorpha). Because of these characters we cannot attribute P. lu to the Mesopsocidae but to a new extinct family very close to the Mesopsocidae that has ‘kept’ some characters considered as ‘basal’ by Smithers (1972).

Paramesopsocus adibi n. sp. (Figs 20–29) Material. Holotype specimen PIN 2784/2096 (figs 20, 21), paratypes: specimens PIN 1782/135, 2066/1591, 2066/3494 (a) and (b) (part and counterpart), 2066/3526 (a) and (b) (part and counterpart), 2066/3586, 2066/3589 (fig. 22), 2066/3593 (a) and (b) (part and counterpart), 2066/3602 (fig. 23), 2066/3605 (fig. 24), 2239/535, 2239/547, 2239/622, 2384/286 (fig. 25), 2384/291, 2465/993, 2554/388, 2784/1998, 2784/2001, 2784/2340, 2997/3706, 2997/4011, 2997/3973, 2997/4018, and PIN 2997/5142, all deposited in the Palaeontological Institute (PIN RAS) Russian Academy of Sciences, Moscow. Type locality and horizon. Late Jurassic (Oxfordian or Kimmeridgian) (Kirichkova & Doludenko 1996), Karatau, South Kazakhstan. Etymology. Named after Mr Adib Indary, father of one of us. Diagnosis. The only visible difference with P. lu is the forewing with Rs and M joined by a small cross-vein. Description. Head 0.64 mm large, eyes rounded, diameter 0.23 mm, 0.34 mm apart (figs 9-13). Number of antennal segments unknown, flagellomeres finely secondary annulated, all filiform, elongate, pedicel 0.06 mm long, 0.03 mm wide; scape 0.09 mm long, 0.04 mm wide.

Figures 26–29 Paramesopsocus adibi n. sp. 26, Drawing of forewing of the paratype specimen n. 2066/3602, scale bar = 1 mm; 27, drawing of forewing of the paratype specimen n. 2066/3602, scale bar = 1 mm; 28, photograph of the paratype specimen n. 2384/286, hind wing; 29, drawing of hind wing of the holotype specimen n. 2784/2096, scale bar = 1 mm.

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Thorax 0.78 mm wide, mesothorax nearly triangular. Forewing glabrous and hyaline, 2.15 mm long, 0.75 mm wide (holotype) (fig. 21), for paratype PIN 2066/3586 2.46 mm long, 1 mm wide (fig. 27), and for paratype PIN 2066/3602 2.24 mm long, 0.77 wide (fig. 26). Pterostigma thickened and sclerotized. Sc short, distally curved and reaching R, then fused with it for 0.6 mm till nearly 1.11 mm from wing base. Distal free part of Sc (Sc’ sensu Lienhard, 1998) forming a sharp angle with costal margin. R1 simple, reaching costal margin at about 1.64 mm from wing base. Rs basally oblique and joined to M by a small cross-vein, fork of R2+3 and R4+5 1.4 mm distal of wing base; R2+3 and R4+5 curved; reaching wing margin respectively at 1.93 and 2.13 mm from wing base. Radial cell closed, longer than broad. M basally fused with Cu1; fork between M1+2 and M3 1.44 mm distal of wing base; M1 and M2 reaching wing apex and separating 1.85 mm distal of wing base. Fork of Cu1 into Cu1a and Cu1b 1.31 mm from wing base; Cu1a curved and longer than Cu1b. Areola postica (AP) free; no cross-vein between AP cell and M. Cu2 weaker than other veins; a distinct nodulus. 1A straight; 2A present. Hind wing glabrous and hyaline, smaller than forewing, about 1.83 mm long, 0.52 mm wide (figs 28, 29), but basal part of hind wing not visible. R1 reaching anterior wing margin 1.19 mm from wing base. fork of Rs into R2+3 and R4+5 1.34 mm from wing base. Basi-radial cell four-angled. R2+3 0.27 mm long; R4+5 0.45 mm long. Fork of M into M1 and M2 1.18 mm from wing base; M1 0.59 mm long; M2 0.36 mm long. Cu1 reaching posterior wing margin at same level as fork of M into M1 and M2. Anal area not visible. Abdomen 0.62 mm wide. Genitalia not well preserved.

Comment. After the keys to the families of Lienhard (1998), P. adibi n. sp. falls in the Psocomorpha for the same characters as above for P. lu: adult with threesegmented tarsi; thickened and sclerotized pterostigma; no scales. This last character is unclear in our fossils because some scale-like structures could be seen, but we think that this is rather related to artifact of observation, and/or due to the impression of the insect organic material on the grain of the holding rock. As for P. lu, the thickened and sclerotized pterostigma is currently considered as being an apomorphy of Psocomorpha (Yoshizawa 2002). But the same difficulty as above for P. lu occurs due to the presence of secondary annulations on flagellomeres, which is a troctomorphan character after the keys of Mockford (1993). According to the keys of recent psocids of Smithers (1990) and Lienhard (1998), P. adibi would fall in the Mesopsocidae for the same characters as in P. lu. However, P. adibi has the same characters as P. lu that exclude an attribution to the Mesopsocidae. The great similarity of tarsal and antennae structure, and wing venations between these two species allows the attribution of P. adibi to the new family Paramesopsocidae. But as some characters (genitalia, lacinia, pulvilli, etc.) are missing in the Karatau fossils we prefer to consider this attribution as only tentative.

Remark. Rasnitsyn (2002: 129) stated that all the psocid material of the Late Jurassic of Karatau studied herein are attributable to the electrentomoid Amphientomidae: Electrentominae. He also (loc. cit.: 136, caption of fig. 163) attributed one of these fossils to an ‘undescribed booklouse of the family Mesopsocidae from the Late Jurassic of Karatau…’. Unfortunately we did not examine this last particular figured fossil, but after the illustration of Rasnitsyn, it clearly belongs to P. adibi. In fact a rapid examination of the psocid material of Karatau could overlook some details and thus would orient workers to place them into Mesopsocidae after taking into account the thickened pterostigma. But the same characters for P. adibi as for P. lu exclude an attribution to this last family, viz. a Rs-M cross-vein and two anal veins in the forewing, and a forked median vein in the hind wing. It is true that some features (antennae with secondary annulation, forewing with short Sc ending in R, Rs and M joined by a cross-vein, presence of two anal veins, long areola postica with sharp apex directed towards wing base, and hind wing with M forked), would direct the attribution of P. adibi to the Amphientomidae and more precisely to the Electrentominae. But in the Amphientomidae, there is no closed basi-radial cell in the hind wing (absence of first segment of Rs), the forewing median vein is three-branched with branches close together, and the pterostigma is not thickened and not sclerotized, unlike in P. adibi.

Phylogenetic position in Psocomorpha Traditionally Psocomorpha were regarded as including four infraorders (Psocetae, Homilopsocidea, Epipsocetae, and Caeciliusetae), but Yoshizawa (2002) based on a cladistic phylogeny added two more, Archipsocetae comprising Archipsocidae, and Hemipsocetae including Hemipsocidae, these two families were assigned previously to Homilopsocidea and Psocetae respectively. Our new family Paramesopsocidae has to be placed very basally in the suborder Psocomorpha, if we admit that the three characters ‘wings with pterostigma thickened and sclerotized’, ‘hooked nodulus formed by truncated spines fused a their base’, and ‘presence of nodus’, are not homoplasic and are really apomorphies of Psocomorpha, as proposed by Yoshizawa (2002). In order to verify the position of the Paramesopsocidae we made an attempt of cladistic analysis based on all the 68 characters and 50 taxa used by Yoshizawa (2002, table 1) and available on the internet page (http://insect3.agr. hokudai.ac.jp/psoco-web/data/index.html), to which we added our genus Paramesopsocus and the character

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number 69: ‘antenna secondary annulation present (0), or absent (1)’. Based on this matrix most parsimonious and consensus trees were found using PAUP* 4 beta 10 (Swofford 2001). A heuristic search was performed using PAUP* 4 beta 10 (Swofford 2001). It yielded 10989 most parsimonious trees with a length of 201, consistency index (CI) = 0.4179; homoplasy index

(HI) = 0.5821; retention index (RI) = 0.8010. A strict consensus tree was also obtained (fig. 30) showing that the phylogenetic relationships near the basal node are unresolved, but nearly all the results of Yoshizawa (2002) are maintained. The Archipsocidae are regarded as the basalmost clade of Psocomorpha, followed by the Hemipsocidae, then by a large polytomy including

Figure 30 Strict consensus cladogram of the most parsimonious trees of Psocomorpha plus Paramesopsocus.

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all the remaining psocomorphan families including our fossil family. This later was inserted in this polytomy without however affecting the topology of the strict consensus tree obtained by Yoshizawa (2002). When looking at the obtained most parsimonious trees, the Paramesopsocidae generally fall in a very ‘basal’ position amongst the group forming the polytomy in the strict consensus tree. Among the great clades (infraorders) proposed by Yoshizawa (2002), only the Homilopsocidea and the Caeciliusetae (in part) are not maintained as clades in our strict consensus cladogram. Polytomies are usually regarded as problems. Nevertheless it is preferred in our case for some nodes because of the lack of information to resolve the evolutionary relationship between these taxa. This indicates that a lot of work is still to be done on Psocodea. More morphological or/and molecular characters must be found for future studies of phylogeny of Psocodea including the fossil taxa in order to improve and determine the history and scenarios of evolution of this group. Acknowledgments. We are grateful to an anonymous referee for the important and useful remarks made on the first version of this paper. This paper is a contribution to the ANR project: AMBRACE (BLAN 07-1_184190).

References Azar D. 2000. Les ambres mésozoïques du Liban. Thèse de Doctorat de l’Université Paris XI, Paris, 164 + 144 p. Azar D., Nel A. 2004. Four new Psocoptera from Lebanese amber (Insecta: Psocomorpha: Trogiomorpha). Annales de la Société Entomologique de France (N.S.) 40: 185-192. Azar D., Perrichot V., Neraudeau D., Nel A. 2003. New psychodid flies from the Cretaceous ambers of Lebanon and France, with a discussion about Eophlebotomus connectens Cockerell, 1920 (Diptera, Psychodidae). Annals of the American Society of America 96: 117-127. Badonnel A. 1943. Psocoptères. Faune de France 42, Paris, 164 p. Baz A., Ortũno V. M. 2000. Archaeatropidae, a new family of Psocoptera from the Cretaceous amber of Alava, Northern Spain. Annals of the Entomological Society of America 93: 367-373. Baz A., Ortũno V. M. 2001a. New genera and species of empheriids (Psocoptera: Empheriidae) from the Cretaceous amber of Alava, northern Spain. Cretaceous Research 22: 575-584. Baz A., Ortuño V. M. 2001b. A new electrentomoid psocid (Psocoptera) from the Cretaceous amber of Alava (Northern Spain). Deutsche Entomologische Zeitschrift 48: 27-32. Carpenter F. M. 1992. Superclass Hexapoda. Treatise on Invertebrate Paleontology, Arthropoda 4 (3-4): 1-655. Cockerell T. D. A. 1916. Insects in Burmese amber. American Journal of Science 42: 135-138. Cockerell T. D. A. 1919. Insects in Burmese amber. The Entomologist 52: 241-243. Garcia-Aldrete A. N. 1984. The Trogiomorpha (Pscoptera), of Chamela, Jalisco, Mexico. Folia Entomologica Mexicana 59: 25-69. Gelhaus J. K., Johnson R. 1996. First record of crane flies (Tipulidae: Limoniinae) in Upper Cretaceous Amber from New Jersey, U.S.A. Transactions of the American Entomological Society 122: 55-65. Grimaldi D., Engel M. S. 2005. Evolution of the insects. Cambridge University Press, Cambridge, 755 p.

Grimaldi D., Engel M. S. 2006a. Fossil Liposcelididae and the lice ages (Insecta: Psocodea). Proceedings of the Royal Society (B) 273: 625-633. Grimaldi D., Engel M. S. 2006b. Extralimital Fossils of the ‘‘Gondwanan’’ Family. Sphaeropsocidae (Insecta: Psocodea). American Museum Novitate 3523: 1-18 Gurney A. B. 1943. A synopsis of the psocids of the tribe Psyllipsocini, including the description of an unusual new genus from Arizona. Annals of the Entomological Society of America 36: 195-220. Huang D.-Y., Nel A., Azar D., Nel P. 2008. Phylogenetic relationships of the Mesozoic paraneopteran family Archipsyllidae (Insecta: Psocodea). Geobios 41(4): 461-464. Johnson K. P., Yoshizawa K., Smith V. S. 2004. Multiple origins of parasitism in lice. Proceedings of the Royal Society of London 271: 17711776. Kirichkova A. I., Doludenko M. P. 1996. New data on the Jurassic phytostratigraphy in Kazakstan. Stratigrafiya. Geologocheskaya Korrelatsiya 4: 35-52 (in Russian, English translation in Stratigraphy and Geology Correlation 4: 440-449. Lienhard C. 1998. Psocoptères euro-méditerranéens. Faune de France 83, Paris, 20 + 517 p. Lienhard C. 2000. A new genus of Prionoglarididae from a Namibian cave (Insecta: Psocoptera). Revue Suisse de Zoologie 107: 871-882. Lienhard C. 2003. Nomenclatural amendments concerning Chinese Psocoptera (Insecta), with remarks on species richness. Revue Suisse de Zoologie 110: 695-721. Lienhard C., Smithers C. N. 2002. Psocoptera world catalogue and bibliography. Instrumenta Biodiversitatis, Muséum d’Histoire Naturelle de Genève, 5, 41 + 745 p. Lin Q.-B. 1976. The Jurassic fossil insects from western Liaoning. Acta Palaeontologica Sinica 15, 97–121. [In Chinese with abstract in English.] Mockford E. L. 1967. The electrentomoid psocids (Psocoptera). Psyche 74: 118-165. Mockford E. L. 1984. Two new species of Speleketor from southern California with comments on the taxonomic position of the genus (Psocoptera: Prionoglarididae). Southwestern Naturalist 29: 169-179. Mockford E. L. 1993. North American Psocoptera (Insecta). Flora & Fauna Handbook, 10, CRC press,Boca Raton, 455 p. Nel A., Waller A. 2007. The first fossil Compsocidae from the Cretaceous Burmese amber (Insecta, Psocoptera, Troctomorpha). Cretaceous Research 28: 1039-1041. Neraudeau D., Perrichot V., Dejax J., Masure E., Nel A., Philippe M., Moreau P., Guillocheau F., Guyot T. 2002. Un nouveau gisement à ambre insectifère et à végétaux (Albien terminal probable): Archingeay (Charente-Maritime, France). Geobios 35: 233-240. Perrichot V., Azar D., Néraudeau D., Nel A. 2003. New Psocoptera in the Lower Cretaceous ambers of southwestern France and Lebanon (Insecta: Psocoptera: Trogiomorpha). Geological Magazine 140: 669-683. Poinar G. O. Jr., Milki R. 2001. Lebanese amber. The oldest ecosystem in fossilized resin. Oregon State University Press, Corvallis, 96 p. Rasnitsyn A. P. 2002. 2.2.1.2.4. Superorder Psocidea Leach, 1815. p. 125-133. In: Rasnitsyn A. P., Quicke D. L. J. (Eds.). History of insects. Kluwer Academic Publishers, Dordrecht: 544 p. Smithers C. N. 1972. The classification and phylogeny of Psocoptera. Australian Museum Memoir 14: 1-351. Smithers C. N. 1990. Keys to the families and genera of Psocoptera (Arthropoda, Insecta). Technical Reports of the Australian Museum 1990: 1-82. Smithers C. N. 1991. Psocoptera (Psocids, booklice), p. 412-420 in: Naumann I. D., Carne P. B., Lawrence J. F., Nielsen E. S., Spradbery J. P., Taylor R. W., Whitten M. J., Littlejohn M. J. (Eds). The insects of Australia: a textbook for students and research workers, 2nd edition. CSIRO, Melbourne University Press, Melbourne.

469


Spahr U. 1992. Ergänzungen und Berichtigungen zu R. Keilbachs Bibliographie und Liste der Bernsteinfossilien. Klasse Insecta (Ausgenommen: Apterygota, Hemipteroidea, Coleoptera, Hymenoptera, Mecopteroidea). Stuttgarter Beiträge zur Naturkunde (B) 182: 1-102. Swofford D. L. 2001. PAUP*. Phylogenetic Analysis using Parsimony (*and other methods), Version 4. Sinauer Associates, Massachusetts. Vishnyakova V. N. 1975. Psocoptera in Late Cretaceous insect-bearing resins from the Taimyr. Entomological Review 54: 63-75.

470

Wappler T., Smith V. S., Dalgleish R. 2004. Scratching an ancient itch: an Eocene bird louse fossil. Proceedings of the Royal Society (B) 271 (Suppl. 5): 255-258. Yoshizawa K. 2002. Phylogeny and higher classification of suborder Psocomorpha (Insecta: Psocodea: Psocoptera). Zoological Journal of the Linnean Society 136: 371-400. Yoshizawa K., Johnson K. P. 2006. Morphology of male genitalia in lice and their relatives and phylogenetic implications. Systematic Entomology 31: 350-361