Insecta, Psocodea, Psocomorpha) Dany Azar, André Nel, Didier Neraudeau

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A new cretaceous psocodean family from the Charente-Maritime amber (France) (Insecta, Psocodea, Psocomorpha) Dany Azar, André Nel, Didier Neraudeau

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Dany Azar, André Nel, Didier Neraudeau. A new cretaceous psocodean family from the Charente- Maritime amber (France) (Insecta, Psocodea, Psocomorpha). Geodiversitas, Museum National d’Histoire Naturelle Paris, 2009, 31 (1), pp.117-127. insu-00392801

HAL Id: insu-00392801 https://hal-insu.archives-ouvertes.fr/insu-00392801 Submitted on 3 Mar 2021

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. A new Cretaceous psocodean family from the Charente-Maritime amber (France) (Insecta, Psocodea, Psocomorpha)

Dany AZAR Lebanese University, Faculty of Sciences II, Department of Biology, Fanar-Matn P.O. box 26110217 (Lebanon) [email protected]

André NEL Muséum national d’Histoire naturelle, CNRS UMR 7205, Département Systématique et Évolution, case postale 50, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected]

Didier NÉRAUDEAU Université Rennes I, UMR CNRS 6118 Géosciences, campus de Beaulieu bât. 15, 263 avenue du Général Leclerc, F-35042 Rennes cedex (France) [email protected]

Azar D., Nel A. & Néraudeau D. 2009. — A new Cretaceous psocodean family from the Charente-Maritime amber (France) (Insecta, Psocodea, Psocomorpha). Geodiversitas 31 (1) : 117-127.

ABSTRACT Arcantipsocus courvillei n. gen., n. sp. is described from the Cretaceous amber of KEY WORDS Archingeay (France). It is placed within the suborder Psocomorpha, and in the Insecta, Mesozoic extinct family Arcantipsocidae n. fam. characterized by 14-segmented Psocodea, Psocoptera, antenna; legs with tarsi 3-segmented; forewing setose with evanescent veins; Psocomorpha, pterostigma dark, thickened and setose; M 2-branched; areola postica free; Arcantipsocidae n. fam., amber, nodulus present; hind wing with M bifurcated, without basi-radial cell; claws Cretaceous, with a preapical tooth. A cladistic phylogeny for Psocomorpha is given includ- France, ing the new fossil taxon. Th e discovery of this new taxon demonstrates the new family, new genus, necessity of a deep phylogenetic redeﬁ nition of the currently admitted major new species. subdivisions of this suborder.

GEODIVERSITAS • 2009 • 31 (1) © Publications Scientiﬁ ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 117 Azar D. et al.

RÉSUMÉ Une nouvelle famille de Psocodae de l’ambre crétacé de Charente-Maritime (France) (Insecta, Psocodea, Psocomorpha). Arcantipsocus courvillei n. gen., n. sp. est décrit de l’ambre crétacé d’Archingeay MOTS CLÉS (France). Il est placé dans le sous-ordre Psocomorpha et dans la famille méso- Insecta, zoïque éteinte Arcantipsocidae n. fam., caractérisée par : antennes avec 14 seg- Psocodea, Psocoptera, ments ; pattes avec trois segments tarsaux ; aile antérieure cétose avec des nervures Psocomorpha, évanescentes ; ptérostigma épais et cétose ; M avec deux branches ; areola postica Arcantipsocidae n. fam., ambre, libre ; nodulus présent ; aile postérieure avec M bifurquée, sans cellule basi-radiale ; Crétacé, griﬀ es avec une dent préapicale. Une analyse phylogénétique cladistique pour France, les Psocomorpha est donnée en incluant notre taxon fossile. La découverte de ce famille nouvelle, genre nouveau, nouveau taxon démontre la nécessité d’une redéﬁ nition phylogénétique profonde espèce nouvelle. des subdivisions majeures actuellement admises au sein de ce sous-ordre.

INTRODUCTION Recently two new taxa of Psocodea from the Charente-Maritime (Archingeay) French amber Recent cladistic analyses reveal the paraphyletic have been studied by Perrichot et al. (2003). nature of several orders of insects. Th e most sig- We describe herein Arcantipsocus courvillei n. gen., nifi cant being the lice (order Phthiraptera), which n. sp., from the Archingeay Cretaceous amber of is now included within the Psocoptera to form France. It is placed into the suborder Psocomorpha, the order Psocodea (Yoshizawa & Johnson 2003a, and in the Cretaceous extinct family Arcantipso- b, 2006; Johnson et al. 2004; Grimaldi & Engel cidae n. fam. 2005, 2006a). Th e Psocodea is a relatively small order with ABBREVIATIONS about 10 000 valid extant species. Th eir earliest Cu cubital vein; record is from the Permian of Kansas (USA), Com- M median vein; mx1-4 fi rst to fourth maxillary palpomere; monwealth of Independent States (C.I.S.) and R radial vein; New South Wales (Australia) (Carpenter 1992); Rs radial sector. although Mockford (1993: 2) considered that the earliest unquestionable fossil psocids known are from Cretaceous amber. If the pre-Late Jurassic SYSTEMATIC PALAEONTOLOGY fossils are currently considered as representatives of the paraneopteran stem group, but resembling We follow in part the catalogue of Lienhard & Psocoptera, Huang et al. (2008) demonstrated Smithers (2002), and the works of Smithers (1972, that the Middle Jurassic Chinese Archipsyllidae 1990) and Mockford (1993) as essential tools for Handlirsch, 1906 are Psocodea. Amber Cretaceous the systematic of the order. We follow the nomen- Psocodea are recorded from Lebanon (Azar 2000; clature of wing venation and body structures of Poinar & Milki 2001; Perrichot et al. 2003; Azar & Smithers (1972), and Lienhard (1998). Th e fossil Nel 2004; Grimaldi & Engel 2006b), France (Per- was carefully prepared in Canada balsam medium, richot et al. 2003), Canada (Spahr 1992), Siberia following the method described by Azar et al. (Taymir Peninsula) (Vishnyakova 1975), USA (2003), in order to observe as many characters as (New Jersey) (Gelhaus & Johnson 1996), Spain possible. Th us the “absences” of structures are ac- (Alava) (Baz & Ortuño 2000, 2001), and Myanmar curate, which is diff erent of structures that are “not (Cockerell 1916, 1919). visible” but may be present.

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FIG. 1. — Photograph of Arcantipsocus courvillei n. gen., n. sp., FIG. 2. — Photograph of Arcantipsocus courvillei n. gen., n. sp., holotype no. ARC 10.2, male, dorsal view. holotype no. ARC 10.2, male, ventral view.

Suborder PSOCOMORPHA Roesler, 1944 ETYMOLOGY. — After “Arcanti” from “Arcantiatum” old name of Archingeay and “psocus”; gender masculine.

Family ARCANTIPSOCIDAE n. fam. DIAGNOSIS. — In addition to the family diagnosis: anten nal fl agellomeres decreasing progressively in length TYPE GENUS. — Arcantipsocus n. gen. by present des- forwards apex. Lacinia with two shoulders each made ignation. of two smooth teeth, the fi rst being in the inner middle of visible part of lacinia and the second situated slightly DIAGNOSIS. — Antenna with 12 fi liform fl agellomeres. La- before the tip. Maxillary palpus 4-segmented, with mx4 cinia present. Legs with tarsi 3-segmented, distal segment the longest and cylindrical. Forewing patterned. Most of tarsi bearing claws with one preapical tooth. Forewing veins evanescent except in their terminal parts. Paraproct membrane setose, veins basally evanescent; pterostigma with bilobed process. thickened and setose; M 2-branched; areola postica free; nodulus present. Hind wing with M bifurcate, without basi-radial cell. Paraproct with lobed process. Arcantipsocus courvillei n. sp. (Figs 1-10) Genus Arcantipsocus n. gen. MATERIAL. — Holotype specimen no. ARC 10.2 (male), TYPE SPECIES. — Arcantipsocus courvillei n. gen., n. sp. deposited in the palaeontology collections of the Muséum by present designation. national d’Histoire naturelle, Paris.

GEODIVERSITAS • 2009 • 31 (1) 119 Azar D. et al.

FIG. 3. — Drawing of the habitus of Arcantipsocus courvillei n. gen., n. sp., holotype no. ARC 10.2, male, dorsal view. Scale bar: 1 mm.

(Fig. 4), with mx4 the longest and cylindrical; mx1 0.04 mm long and 0.03 mm wide; mx2 0.15 mm long and 0.03 mm wide; mx3 0.05 mm long and 0.03 mm wide; mx4 0.18 mm long and 0.03 mm

FIG. 4. — Drawing of the maxillary palpus of Arcantipsocus courvillei wide. Labial palpus not visible. Visible part of lac- n. gen., n. sp., holotype no. ARC 10.2, male. Scale bar: 0.1 mm. inia 0.09 mm long (Figs 5; 6), with two shoulders made of two smooth teeth each, the fi rst being in TYPE LOCALITY AND HORIZON. — Archingeay-Les the inner middle and the second situated slightly Nouil lers, Charente-Maritime, France; Lower Creta- before the tip, the apex formed of two smooth teeth, ceous, uppermost Albian. one of them being very small. Th orax 0.58 mm wide; mesothorax nearly tri- ETYMOLOGY. — After Dr Philippe Courville, palaeon- tologist who helped us in collecting fossil insects in angular. this amber. Legs with tarsi 3-segmented, distal segment bearing claws with one preapical tooth (Fig. 7). DIAGNOSIS. — As for the genus. Forewing patterned and setose, 1.97 mm long and 0.57 mm wide (Fig. 3). Marginal setae crossing. DESCRIPTION Two rows of setae on veins. Apex slightly acuminate. Total body length 2.11 mm (Figs 1-3). Head nearly Most veins evanescent except in their terminal parts. triangular. Antenna with 14 segments (12 fl agel- Pterostigma dark, thickened and setose, convex lomeres) 2.42 mm long, fl agellomeres fi liform, and not connected to Rs by a cross-vein. Sc diff use elongate, and decreasing progressively in length. and evanescent. R1 simple reaching costal margin Th e fi rst fl agellomere being the longest 0.3 mm in at 1.55 mm from wing base. Rs evanescent and length, the shortest the last one 0.1 mm. Pedicel hardly visible; fork of R2 + 3 and R4 + 5 1.41 mm and scape nearly cylindrical, respectively 0.11 and distal of wing base; R2 + 3 and R4 + 5 strongly 0.1 mm in length, and 0.05 and 0.03 mm wide. curved; reaching wing margin respectively at 1.7 Compound eyes nearly rounded with 0.22 mm of and 1.88 mm from wing base. M 2-branched. M1 diameter. Th ree large ocelli disposed in triangle be- and M2 separating 1.58 mm distal of wing base; M1 tween compound eyes. Maxillary palpus 4-segmented reaching wing margin, 0.38 mm long; M2 nearly

120 GEODIVERSITAS • 2009 • 31 (1) New Psocodea (Insecta) from Cretaceous French amber

straight, 0.27 mm long. Fork of Cu1 in Cu1a and Cu1b 1.31 mm from wing base; Cu1a strongly curved and longer than Cu1b. Areola postica (AP) free; no cross-vein between AP cell and M. Cu2 very weak and hardly visible. A distinct nodulus present. Anal vein (A) very poorly preserved. Hind wing hyaline, smaller than forewing, with setose margin, 1.55 mm long and 0.5 mm wide (Fig. 8). Sc not visible. R fused basally with M and Cu. R1 0.23 mm long, not reaching anterior wing margin. No basi-radial cell. Bifurcation of Rs into R2 + 3 and R4 + 5 1.1 mm from wing base. M bifurcated. Bifurcation of M into M1 and M2 1 mm from wing base; M1 0.45 mm long; M2 0.32 mm long. Cu1 reaching posterior wing margin at 1.16 mm. Remaining veins hidden. Abdomen 1 mm long and 0.45 mm wide. Male appendages relatively well preserved (Figs 9; 10), hypandrium transparent and not well sclerotized, with a ﬂ attened posterior extremity; paraproct with sharply bilobed process; phallosome visible by transparency, Y-shaped.

DISCUSSION

Th e new fossil possesses characters shared by both suborders Psocomorpha and Troctomorpha, viz. adult with tarsi 3-segmented, hind wing with M vein FIG. 5. — Photograph of the apical part of the lacinia of Arcanti- 2-branched (probably a plesiomorphy as it is present psocus courvillei n. gen., n. sp., holotype no. ARC 10.2, male. in the Archipsyllidae and the Hemipsocidae Pearman, 1936, psocomorphan families having an inclusive position in Yoshizawa’s phylogeny), and forewing with nodulus. According to the keys proposed by Mockford (1993) and Lienhard (1998), Arcanti- psocus courvillei n. gen., n. sp. falls in the suborder

Psocomorpha because of the characters “forewing FIG. 6. — Drawing of the apical part of the lacinia of Arcantipso- with thickened and sclerotized pterostigma”, and “no cus courvillei n. gen., n. sp., holotype no. ARC 10.2, male. Scale scales on wings and body”. Th e character “thickened bar: 0.05 mm. and sclerotized pterostigma” is considered as being apomorphic of Psocomorpha by Yoshizawa (2002) and Mockford (1967). But with antennae with 12 ﬂ agellomeres, the new fossil has also features of the Troctomorpha. Th us its position is problematic and we have to discuss more precisely these characters. FIG. 7. — Drawing of distal part of fore tarsus of Arcantipsocus Recent Psocomorpha have 11 ﬂ agellomeres or courvillei n. gen., n. sp., holotype no. ARC 10.2, male. Scale bar: fewer, while Troctomorpha usually have 13 but 0.1 mm.

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Th erefore, the new fossil cannot be assigned to any of the known families. Because of the unique combination of characters mentioned above and the thickened and sclerotized pterostigma that is apomorphic for the Psocomorpha, we attribute Arcantipsocus n. gen. to a new extinct family within this suborder. FIG. 8. — Drawing of the hindwing of Arcantipsocus courvillei n. gen., n. sp., holotype no. ARC 10.2, male. Scale bar: 0.5 mm. PHYLOGENY sometimes fewer (Mockford 1993). Th e presence Th ere are very few attempts of phylogenetic analy- of only 11 fl agellomeres is likely plesiomorphic, as ses of the Psocoptera. Smithers (1972) dedicated a the Archipsyllidae have 11 fl agellomeres and are in large part of his work Th e Classifi cation and Phyl- a very inclusive position in the Psocodea (Huang ogeny of Psocoptera to the study of the phylogenetic et al. 2008). Th us the number of fl agellomeres is relationships within Psocoptera, but his work is not suffi cient to assign this fossil taxon to one of typological. Smithers (1972) proposed dendro- the two suborders since this character seems to be grams with several lineages based on homoplastic homoplastic. or plesiomorphic characters. Several years later If we neglect the character “thickened and he admitted that his phylogeny needed revision sclerotized pterostigma” that is diagnostic of the (Smithers 1991). Perrichot et al. (2003) presented a Psocomorpha in the key to recent families of Smith- tentative of cladistic phylogeny for Trogio morpha, ers (1990), Arcantipsocus n. gen. falls in the trocto- nevertheless this later phylogeny is incomplete morphan Amphientometae family Compsocidae because based on few characters. Yoshizawa et Mockford, 1967 for its hind wing vein M forked al. (2006) proposed another phylogeny of the and the forewing with a nodulus. But all Compso- same group, based on molecular data. Yoshizawa cidae have hyaline and unsclerotized pterostigma, (2002), Johnson et al. (2004) and Grimaldi & Engel unlike Arcantipsocus n. gen. (2006a), proposed molecular and morphological If we consider this character, Arcantipsocus n. gen. phylogenetic analyses; the last one concerned the falls near the psocomorphan family Bryopsocidae Psocomorpha only. Mockford, 1984 after the following combina- Our new family Arcantipsocidae n. fam. should tion of characters: macropterous insect; legs with fall within the Psocomorpha Homilopsocidea (sensu trimerous tarsi; absence of scales; complex wing Yoshizawa 2002), if we admit that the character venation, in contrast to some psocids with vena- “pterostigma thickened and sclerotized” is not tion reduced to some parallel veins; sclerotized homoplastic, and is a real apomorphy of the Psoco- pterostigma; head not elongate; free areola postica morpha, as proposed by Yoshizawa (2002). Th e status with Cu1a and Cu1b separating near posterior of this character is debatable as Yoshizawa (2002) margin; forewing margin and membrane setose; noted that “although a thickened pterostigma is hind wing with some setae on margin in addition observed in Archipsocidae, it appears to be much to setae between arms of radial fork. However, thinner than in other families of Psocomorpha (...) Arcantipsocus n. gen. diff ers from all Bryopso- diff erent degrees of thickness of the pterostigma cidae by the following characters: antennae with may provide further evidence for the phylogenetic 12 fl agellomeres instead of 11; hind wing without placement of Archipsocidae as the basalmost clade basi-radial cell; bifurcation of M in hind wing into of Psocomorpha”. Also it is shared by the Mesozoic M1 and M2 (this last character is shared by the family Archipsyllidae that is supposed to be a more majority of psocids belonging to the suborders inclusive group than all recent Psocodea (Huang Troctomorpha and Trogiomorpha). et al. 2008).

122 GEODIVERSITAS • 2009 • 31 (1) New Psocodea (Insecta) from Cretaceous French amber

FIG. 9. — Photograph of the male genitalia of Arcantipsocus courvillei n. gen., n. sp., holotype no. ARC 10.2.

In order to verify the position of the Arcanti- psocidae n. fam., we made an attempt of cladistic analysis based on all the 68 characters and 50 taxa used by Yoshizawa (2002) (see Appendix for matrix) and available on the internet (http://www.psocodea. org/kazu/data/psocomorpha/psocomorpha.nexus), to which we added our genus Arcantipsocus n. gen. and another fossil troctomorphan genus Electrento- mum Enderlein, 1911. A heuristic search was per- formed using PAUP* 4 beta 10 (Swoﬀ ord 2001). It yielded 165 most parsimonious trees with a length of 199, consistency index (CI) = 0.4171; homo- plasy index (HI) = 0.5829; retention index (RI) = 0.8076. A strict consensus tree was also obtained showing that phylogenetic relationships near the basal node are unresolved. Our consensus tree is congruent with the results obtained by Yoshizawa (2002). His six infraorders are maintained in our results. Archipsocidae Enderlein, 1903 (Archi- psocetae) are regarded as the most inclusive clade of Psocomorpha, followed by the Hemipsocidae (Hemipsocetae), then by the remaining four infraorders of Epipsocetae, Caeciliusetae, Psocetae, FIG. 10. — Drawing of the male genitalia of Arcantipsocus courvillei and Homilopsocidea that include our fossil family, n. gen., n. sp., holotype no. ARC 10.2. Scale bar: 0.2 mm.

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Echmepteryx TROGIOMORPHA Tapinella Troctopsocidae* Electrentomum TROCTOMORPHA Tineomorpha Paraphientomum Archipsocus ARCHIPSOCETAE Pararchipsocus Archipsocidae Hemipsocus Hemipsocidae HEMIPSOCETAE Psilopsocus Psilopsocidae Psocus Psocidae Stigmatoneura PSOCETAE Myopsocus Myopsocidae Lichenomima Elipsocus Elipsocidae Hemineura Trichopsocus Trichopsocidae Calopsocus Calopsocidae Heterocaecilius Pseudocaecilius Pseudocaeciliidae Ophiodopelma Philotarsus Philotarsidae Arionella Arcantipsocus n. gen. Arcantipsocidae n. fam. HOMILO- Bryopsocus Bryopsocidae PSOCIDEA Kaestneriella Peripsocidae Peripsocus Idatenopsocus Mesopsocus Mesopsocidae Ectopsocus Ectopsocidae Nanolachesilla PSOCOMORPHA Lachesilla (Vietnam) Lachesilla Lachesillidae Eolachesilla Epipsocus Epipsocidae Epipsocopsis Dolabellopsocus Dolabellopsocidae Isthmopsocus EPIPSOCETAE Cladiopsocus Cladiopsocidae Spurostigma Triplocania Ptiloneuridae Ptiloneura Notiopsocus Asiopsocidae Asiopsocus Stenopsocus Stenopsocidae Graphocaecilius Amphipsocus CAECILIUSETAE Kolbea Amphipsocidae Matsumuraiella Dasydemellidae Kodamaius Caecilius Caeciliusidae Dasypsocus

FIG. 11. — Strict consensus cladogram of the Psocomorpha including Arcantipsocus n. gen. and Electrentomum Enderlein, 1911. See Appendix for the matrix of character state used. *, all the genera belonging to the family. together with a group of four families Philotarsidae, clear synapomorphy. Th is later was inserted in the Pseudocaeciliidae, Calopsocidae, and Trichopsocidae cladogram without however aﬀ ecting the topology but this clade of ﬁ ve families is not supported by any of the strict consensus tree obtained by Yoshizawa

124 GEODIVERSITAS • 2009 • 31 (1) New Psocodea (Insecta) from Cretaceous French amber

(2002). Traditionally Psocomorpha were regarded as CARPENTER F. M. 1992. — Superclass Hexapoda, in including four infraorders (Psocetae, Homilopsoci- MOORE R. C. & KAESLER R. L. (eds), Treatise on dea, Epipsocetae, and Caeciliusetae) but Yoshizawa Invertebrate Paleontology, Arthropoda 4 (3/4), Geo- logical Society of America and University of Kansas, (2002), based on a cladistic phylogeny added two Boulder, Colorado and Lawrence: 1-655. more, Archipsocetae comprising Archipsocidae, and COCKERELL T. D. A. 1916. — Insects in Burmese amber. Hemipsocetae including Hemipsocidae, these two American Journal of Science 42: 135-138. families were previously assigned to Homilopsocidea COCKERELL T. D. A. 1919. — Insects in Burmese amber. and Psocetae respectively. More morphological or/ Entomologist 52: 241-243. ENDERLEIN G. 1903. — Die Copeognathen des Indo- and molecular characters are needed for future stud- australischen Faunengebietes. Annales Historico- ies of phylogeny of Psocodea including the fossil Naturales Musei Nationalis Hungarici 1: 179-344. taxa in order to precise the history and scenarios GELHAUS J. K. & JOHNSON R. 1996. — First record of of evolution of this group. crane fl ies (Tipulidae: Limoniinae) in Upper Creta- ceous Amber from New Jersey, U.S.A. Transactions of the American Entomological Society 122: 55-65. GRIMALDI D. & ENGEL M. S. 2005. — Evolution of Acknowledgements the Insects. Cambridge University Press, Cambridge, We are grateful to Dr Vincent Perrichot for making 755 p. the material available for study. We thank also the GRIMALDI D. & ENGEL M. S. 2006a. — Fossil Liposcelidi- anonymous reviewers for their helpful remarks on dae and the lice ages (Insecta: Psocodea). Proceedings of the Royal Society, serie B, 273: 625-633. an earlier version of the manuscript. Th is paper is a GRIMALDI D. & ENGEL M. S. 2006b. — Extralimital contribution to the Agence nationale de la Recherche fossils of the ‘‘Gondwanan’’ Family. Sphaeropsocidae (ANR) project: AMBRACE no. BLAN 07-1-184190 (Insecta: Psocodea). American Museum Novitates and to the scientifi c project “Th e Study of the Fossil 3523: 1-18. Insects in Lebanon and their Outcrops: Geology of HUANG D.-Y., NEL A., AZAR D. & NEL P. 2008. — Phylo- genetic relationships of the Mesozoic paraneopteran the Outcrops – Historical and Biodiversity Evolu- family Archipsyllidae (Insecta: Psocodea). Geobios tion” fi nanced by the Lebanese University. 41: 461-464. JOHNSON K. P., YOSHIZAWA K. & SMITH V. S. 2004. — Multiple origins of parasitism in lice. Proceedings of REFERENCES the Royal Society of London 271: 1771-1776. LIENHARD C. 1998. — Psocoptères euro-méditerranéens. Faune de France. Fédération française des Sociétés de AZAR D. 2000. — Les ambres mésozoïques du Liban. Sciences naturelles, Paris, 83: i-xx + 517 p. Th èse de Doctorat de l’Université Paris XI, France, LIENHARD C. & SMITHERS C. N. 2002. — Psocoptera 164 p. + 144 p., annexes. (Insecta): World catalogue and bibliography. Instru- AZAR D. & NEL A. 2004. — Four new Psocoptera from menta Biodiversitatis 5: xli + 745 p. Lebanese amber (Insecta: Psocomorpha: Trogiomor- MOCKFORD E. L. 1967. — Th e electrentomoid psocids pha). Annales de la Société entomologique de France (Psocoptera). Psyche 74: 118-165. 40 (2): 185-192. MOCKFORD E. L. 1984. — Relationships among philo- AZAR D., PERRICHOT V., NÉRAUDEAU D. & NEL A. tarsid and pseudocaeciliid genera and a proposed 2003. — New psychodid fl ies from the Cretaceous new family Bryopsocidae (Psocoptera). Psyche 91: ambers of Lebanon and France, with a discussion about 309-318. Eophlebotomus connectens Cockerell, 1920 (Diptera, MOCKFORD E. L. 1993. — North American Psocoptera Psychodidae). Annals of the Entomological Society of (Insecta). Flora & Fauna Handbook 10. Sandhill Crane America 96 (2): 117-127. Press, Gainsville, Florida, 455 p. BAZ A. & ORTU˜ NO V. M. 2000. — Archaeatropidae, a PERRICHOT V., AZAR D., NÉRAUDEAU D. & NEL A. new family of Psocoptera from the Cretaceous amber 2003. — New Psocoptera in the Lower Cretaceous of Alava, Northern Spain. Annals of the Entomological ambers of southwestern France and Lebanon (Insecta: Society of America 93: 367-373. Psocoptera: Trogiomorpha). Geological Magazine 140 BAZ A. & ORTU˜ NO V. M. 2001. — New genera and (6): 669-683. species of empheriids (Psocoptera: Empheriidae) POINAR G. O. JR & MILKI R. 2001. — Lebanese Amber. from the Cretaceous amber of Alava, northern Spain. Th e Oldest Ecosystem in Fossilized Resin. Oregon State Cretaceous Research 22: 575-584. University Press, Corvallis, Oregon, 96 p.

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Submitted on 12 October 2007; accepted on 18 December 2008.

126 GEODIVERSITAS • 2009 • 31 (1) New Psocodea (Insecta) from Cretaceous French amber

APPENDIX

Matrix of characters states used for the cladistic analysis (Fig. 11). The list of characters is available on the internet (http://www. psocodea.org/kazu/data/psocomorpha/psocomorpha.nexus) (Yoshizawa 2002). *, all the genera belonging to the family.

11111111112222222222333333333344444444445555555555666666666 12345678901234567890123456789012345678901234567890123456789012345678 Echmepteryx 00000000000000000000020?00000000000000000000000?10000000011000000000 Tapinella 0000000000000?000000000?00000000002000000000000100000000000000000000 Troctopsocidae* 00000000000000000000100?00000000002000000000000100000000000000000000 Tineomorpha 00000000000000000000020?00000000000000000000000?10000000000000000000 Paramphientomum 00000000000000000000020?00000010000000000000000?10000000000000000000 Archipsocus 10000000100002100100021?11000011110000011100000102000000001?20000000 Pararchipsocus 10000000100002100100021?11000011110000011100000102000000001?00000000 Hemipsocus 10110000100010000200000010000110110000011001000?10001000000000000000 Psilopsocus 10100000100010010111100010000010112000101001000102001001?10000100000 Psocus 10101000100010010111100010000210112000110001100002001001010000100001 Sigmatoneura 10101000100010010111100010000210112000110001100002001001010000100001 Myopsocus 10100000100010010111100010000210112000101001000?10001101000000000000 Lichenomima 10100000100010010111100010000210112010101001000000001101000000000000 Elipsocus 10000000100011110111000010000010111000000000000000000010000001000000 Hemineura 10000000100011110111000010000010111000000000000000000010000001000100 Ectopsocus 10100000100111110111000010001010112010011100000000100020000001000100 Eolachesilla 10100000100111110111000010000010112000001000000000100000000001000100 Nanolachesilla 10111000100111110111000010000010112000011000000000100030001?1??00100 Lachesilla (Vietnam) 10101000100111110111000010000010112000010000000?12000000001?1??00100 Lachesilla 10101000100111110111000010000010112000011000000?12000000001?2??00000 Trichopsocus 10101000100011110211000110000010110001011110000000100000010001011000 Pseudocaecilius 10101000100011110211001110000010110100011110111000000020010101011000 Ophiodopelma 10101000100011110211001110000010110100011100110000100020010101011000 Heterocaecilius 1010100010?011110211001110000010110100011110111000100020010101011000 Calopsocus 00101011100011110211011110000010110100011010111000100020010101011000 Bryopsocus ??????????0????????101001000001?1?0????0?0?0000000110030?100010?2000 Kaestneriella 10110000100011110111000010001010112000010000000001110030010001012000 Peripsocus 10100000100011111111100010001010110000010000000001110030010001012000 Philotarsus 10101100100011110111001110000011110101000000100100010030010001012000 Aaroniella 10101000100011110111001110000011110101000000000100110030010001012000 Idatenopsocus 10000000101011110111100010000010111001000000000000010030010201012000 Mesopsocus 10000000101011110111100010000010111000000000000100000030010201012000 Epipsocus 10111111101011010111000110010011110200010000000000000000001?00000120 Epipsocopsis 10111111101011010111000110010011110200010000000000000000001?00000120 Dolabellopsocus 10101111101011010111000110010000110200011000000000000000001?00000120 Isthmopsocus 10101111101011010111000110010000110200011000000000000000001?00000120 Cladiopsocus 10101111101011010111000110010000110200010000000??0000000000000000120 Spurostigma 10101111101011010111000110000000110200010000000000000000000000000110 Triplocania 10111111100011010111000110010001110200000000001?00000000000000000110 Ptiloneura 1011111110?01101011100011001000111020000000????????00000000000000110 Notiopsocus 10100011111111111211000010001010110000012100000000000000000010000200 Asiopsocus 1?10?111?11???1????11000100000101?2000?101?0??0000000000000?100002?0 Stenopsocus 100000111011111111110000101001101110000111100000000000001000000002?0 Graphocaecilius 10000011101111111111000010100110112000011110000000000000100000000200 Amphipsocus 110000111011111111110001100000111102000111100000000000000000000002?0 Kolbea 110000111011111111110001100000111102000111100000000000000000000002?0 Matsumuraiella 11000011101111111111010110000011111000011110000000000000000000000200 Caecilius 100000111011111111110000100000111100000111100000000000000000000002?0 Kodamaius 110000111011111111110001100001111100000111100000000000000000000002?0 Dasypsocus 000000111011111111110000100000111100000111100000000000000000000002?0 Arcantipsocus 10100000?10?1?1101000111100?001?0?00000020?1???00?1????????????????? Electrentomum 00000000000000000000100000010000?020–000000?????????????????????????

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