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The first fossil Compsocidae from Cretaceous Burmese amber (Insecta, Psocoptera, Troctomorpha)
Cretaceous Research 28 (2007) 1039e1041 www.elsevier.com/locate/CretRes
The first fossil Compsocidae from Cretaceous Burmese amber (Insecta, Psocoptera, Troctomorpha) A. Nel*, A. Waller CNRS UMR 5202, Muse´um National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005 Paris, France Received 12 September 2006; accepted in revised form 5 February 2007 Available online 25 July 2007
Abstract The first fossil record of the Compsocidae, Burmacompsocus perreaui gen. et sp. nov., is described from Late Albian Burmese amber. Its strong similarity to the two extant compsocid genera suggests a remarkable morphological stability within this group of 100 Ma. This family, now known only in Central America, was certainly more widespread in the past. Ó 2007 Elsevier Ltd. All rights reserved. Keywords: Insecta; Psocomorpha; Troctomorpha; Compsocidae; First fossil record; Cretaceous; Burmese amber
1. Introduction
Derivation of name. After Burma and Compsocus.
The Psocoptera: Trogiomorpha are rather well represented in the Cretaceous (Perrichot et al., 2003; Azar and Nel, 2004), but Mesozoic fossils of the suborder Troctomorpha are much scarcer (Baz and Ortu~ no, 2001; Lienhard, 2002; Grimaldi and Engel, 2006). Within this group, there is only one Mesozoic record of electrentomoid psocids, namely Manicapsocidus enigmaticus Baz and Ortu~ no, 2001. The small family Compsocidae Mockford, 1967 has been unknown in the fossil record hitherto. Furthermore, it is only known from two species from Central America. Thus the discovery of this group in the Late Albian Burmese amber is of great interest for its age and biogeography. In our description below we follow the body and wing venation terminology of Yoshizawa (2005).
Type species. Burmacompsocus perreaui sp. nov.
2. Systematic palaeontology Family: Compsocidae Mockford, 1967 Genus Burmacompsocus gen. nov. * Corresponding author. E-mail address: [email protected] (A. Nel). 0195-6671/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2007.02.002
Diagnosis. Body structure and wing venation nearly identical to those of the recent Compsocidae, 13-segmented antenna; first segment of Rs absent in hind wing; micro-vestitute of fore wing membrane in form of short points; vein 2A not joining 1A. Burmacompsocus perreaui sp. nov. Figs. 1e2 Derivation of name. After our colleague Dr Michel Perreau. Material. Holotype specimen Burm1; paratype specimen Burm2; Michel Perreau coll., provisionally housed in the collection of fossil insects, MNHN, Paris. Diagnosis. As for the genus. Age and outcrop. Late Albian, Burmese amber, Myanmar. Description. Both the holotype and paratype are complete but of unknown sex. Body and wings without flattened scales;
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head 0.3 mm long, 0.42 mm wide; labrum without ridges from base to margin; lacinia tip with lateral cusp curved outward, bearing a few, indistinct rounded denticles, no secondary tooth visible on cusp; maxillary palp with P4 0.1 mm long, P3 0.04 mm long, P2 0.14 mm long; three ocelli present, arranged in a triangle; antenna with 13 flagellomeres, at least the fourth and fifth being annulate; scape 0.08 mm long, pedicel 0.04 mm long; first flagellar segment f1 0.12 mm long, second f2 0.1 mm long; several long setae on frons; compound eyes large, reaching hind margin of head capsule; frontal suture present. Fore wing hyaline, glabrous, micro-vestitute of membrane in form of short points, wing 2 mm long, 0.64 mm wide; pterostigma closed basally, not sclerotized, 0.38 mm long, 0.10 mm wide; areola postica 0.46 mm long, 0.12 mm wide, not joined to M by a cross-vein; nodulus present; two anal veins not fused distally. Hind wing hyaline, 1.8 mm long, 0.5 mm wide; first segment of Rs absent; vein M two-branched. Fore femora 0.32 mm long, a row of about 10 denticles present on anterior carina of fore femur; tibia 0.28 mm long, tarsi 0.24 mm long; mid femora 0.4 mm long, tibia 0.4 mm long, tarsi 0.28 mm long; hind femora 0.4 mm long, tibia 0.78 mm long, tarsi 0.52 mm long, t1 0.40 mm long, t2 0.06 mm long, t3 0.06 mm long; all tarsi three-segmented; pretarsal claw with two preapical denticles; pulvillus not visible. Abdomen 0.5 mm long; genital structure poorly visible. Discussion. Following Mockford (1993), Burmacompsocus gen. nov. falls within the suborder Troctomorpha because of the following characters: antenna with 13 flagellomeres; at least the fourth and fifth being annulate; tarsi three-segmented; fore wing without sclerotized pterostigma. Following the key
Fig. 1. Burmacompsocus perreaui gen. et sp. nov., holotype Burm1. A, C, photograph and drawing respectively of dorsal view. B, photograph of ventral view. D, drawing of wings. Scale bars represent 1 mm.
Fig. 2. Burmacompsocus perreaui gen. et sp. nov., paratype Burm2, photograph. Scale bar represents 1 mm.
A. Nel, A. Waller / Cretaceous Research 28 (2007) 1039e1041
to modern families of Smithers (1990), Burmacompsocus would fall in the Compsocidae Mockford, 1967 because of the following characters: macropterous; tarsi three-segmented; body and wings without flattened scales; fore wing venation complex; pterostigmal area not more opaque than rest of the membrane; fore wing with nodulus; fore wing with two anal veins; hind wing with M forked. Mockford (1967) gave the following diagnostic characters for this electrentomoid family: antenna with 13e14 segments (13 in Burmacompsocus); frontal suture present, as in Burmacompsocus; lacinia tip with lateral cusp curved outward (present in Burmacompsocus), bearing few indistinct rounded denticles (no secondary tooth visible on cusp in Burmacompsocus); row of denticles present on anterior carina of fore femur (visible in Burmacompsocus); tarsi three-segmented (as in Burmacompsocus); pretarsal claw with two preapical denticles (visible in Burmacompsocus); mesothoracic tergal lobes not protruding above a line drawn from vertex of head to dorsal surface of abdomen (not applicable to our fossils); pterostigma closed basally (as in Burmacompsocus); vein M in hind wing one-branched (in fact two-branched, as in Burmacompsocus); first segment of Rs in hind wing present or absent (absent in Burmacompsocus); third ovipositor valvula deeply bilobed apically (not visible in Burmacompsocus). The last diagnostic character ‘vein 2A of fore wing joining vein 1A’, is not shared by Burmacompsocus. The Compsocidae comprises the two modern genera Electrentomopsis Mockford, 1967 and Compsocus Banks, 1930, both from the Central America (Panama, Mexico). Burmacompsocus shares with Electrentomopsis the 13-segmented antenna (14 in Compsocus); absence of first segment of Rs in hind wing; micro-vestitute of fore wing membrane in form of short points (Mockford, 1967; Smithers, 1990).
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Burmacompsocus differs from both Electrentomopsis and Compsocus in that vein 2A does not join 1A. Acknowledgements We are very grateful to our colleague, Dr. Michel Perreau, for allowing us to work on the type material of Burmacompsocus. References Azar, D., Nel, A., 2004. Four new Psocoptera from Lebanese amber (Insecta: Psocomorpha: Trogiomorpha). Annales de la Socie´te´ Entomologique de France 40, 183e190. Baz, A., Ortu~no, V.M., 2001. A new electrentomoid psocid (Psocoptera) from the Cretaceous amber of Alava (northern Spain). Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin, Deutsche Entomologische Zeitschrift 48, 27e32. Grimaldi, D.A., Engel, M.S., 2006. Fossil Liposcelididae and the lice ages (Insecta: Psocodea). Proceedings of the Royal Society of London, B 273, 625e633. Lienhard, C., 2002. Classification of Psocoptera with checklist of genera (including Cenozoic and Cretaceous fossils, in amber) compiled by Charles Lienhard. Natural History Museum, Geneva, Switzerland. http://www. ville-ge.ch/musinfo/mhng/page-e/artowho.htm#lienhard. Mockford, E.L., 1967. The electrentomoid psocids (Psocoptera). Psyche 74, 109e165. Mockford, E.L., 1993. North American Psocoptera (Insecta). Flora; Fauna Handbook 10. Sandhill Crane Press, Gainesville, Florida, xviii þ 455 pp. Perrichot, V., Azar, D., Neraudeau, D., Nel, A., 2003. New Psocoptera in the Lower Cretaceous ambers of southwestern France and Lebanon (Insecta: Psocoptera: Trogiomorpha). Geological Magazine 140, 669e683. Smithers, C.N., 1990. Keys to the families and genera of Psocoptera (Arthropoda, Insecta). Technical Reports of the Australian Museum 1990 (2), 1e82. Yoshizawa, Y., 2005. Morphology of Psocomorpha (Psocodea: ‘Psocoptera’). Insecta Matsumarana, New Series 62, 1e44.
The first fossil Compsocidae from Cretaceous Burmese amber (Insecta, Psocoptera, Troctomorpha) A. Nel*, A. Waller CNRS UMR 5202, Muse´um National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005 Paris, France Received 12 September 2006; accepted in revised form 5 February 2007 Available online 25 July 2007
Abstract The first fossil record of the Compsocidae, Burmacompsocus perreaui gen. et sp. nov., is described from Late Albian Burmese amber. Its strong similarity to the two extant compsocid genera suggests a remarkable morphological stability within this group of 100 Ma. This family, now known only in Central America, was certainly more widespread in the past. Ó 2007 Elsevier Ltd. All rights reserved. Keywords: Insecta; Psocomorpha; Troctomorpha; Compsocidae; First fossil record; Cretaceous; Burmese amber
1. Introduction
Derivation of name. After Burma and Compsocus.
The Psocoptera: Trogiomorpha are rather well represented in the Cretaceous (Perrichot et al., 2003; Azar and Nel, 2004), but Mesozoic fossils of the suborder Troctomorpha are much scarcer (Baz and Ortu~ no, 2001; Lienhard, 2002; Grimaldi and Engel, 2006). Within this group, there is only one Mesozoic record of electrentomoid psocids, namely Manicapsocidus enigmaticus Baz and Ortu~ no, 2001. The small family Compsocidae Mockford, 1967 has been unknown in the fossil record hitherto. Furthermore, it is only known from two species from Central America. Thus the discovery of this group in the Late Albian Burmese amber is of great interest for its age and biogeography. In our description below we follow the body and wing venation terminology of Yoshizawa (2005).
Type species. Burmacompsocus perreaui sp. nov.
2. Systematic palaeontology Family: Compsocidae Mockford, 1967 Genus Burmacompsocus gen. nov. * Corresponding author. E-mail address: [email protected] (A. Nel). 0195-6671/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2007.02.002
Diagnosis. Body structure and wing venation nearly identical to those of the recent Compsocidae, 13-segmented antenna; first segment of Rs absent in hind wing; micro-vestitute of fore wing membrane in form of short points; vein 2A not joining 1A. Burmacompsocus perreaui sp. nov. Figs. 1e2 Derivation of name. After our colleague Dr Michel Perreau. Material. Holotype specimen Burm1; paratype specimen Burm2; Michel Perreau coll., provisionally housed in the collection of fossil insects, MNHN, Paris. Diagnosis. As for the genus. Age and outcrop. Late Albian, Burmese amber, Myanmar. Description. Both the holotype and paratype are complete but of unknown sex. Body and wings without flattened scales;
1040
A. Nel, A. Waller / Cretaceous Research 28 (2007) 1039e1041
head 0.3 mm long, 0.42 mm wide; labrum without ridges from base to margin; lacinia tip with lateral cusp curved outward, bearing a few, indistinct rounded denticles, no secondary tooth visible on cusp; maxillary palp with P4 0.1 mm long, P3 0.04 mm long, P2 0.14 mm long; three ocelli present, arranged in a triangle; antenna with 13 flagellomeres, at least the fourth and fifth being annulate; scape 0.08 mm long, pedicel 0.04 mm long; first flagellar segment f1 0.12 mm long, second f2 0.1 mm long; several long setae on frons; compound eyes large, reaching hind margin of head capsule; frontal suture present. Fore wing hyaline, glabrous, micro-vestitute of membrane in form of short points, wing 2 mm long, 0.64 mm wide; pterostigma closed basally, not sclerotized, 0.38 mm long, 0.10 mm wide; areola postica 0.46 mm long, 0.12 mm wide, not joined to M by a cross-vein; nodulus present; two anal veins not fused distally. Hind wing hyaline, 1.8 mm long, 0.5 mm wide; first segment of Rs absent; vein M two-branched. Fore femora 0.32 mm long, a row of about 10 denticles present on anterior carina of fore femur; tibia 0.28 mm long, tarsi 0.24 mm long; mid femora 0.4 mm long, tibia 0.4 mm long, tarsi 0.28 mm long; hind femora 0.4 mm long, tibia 0.78 mm long, tarsi 0.52 mm long, t1 0.40 mm long, t2 0.06 mm long, t3 0.06 mm long; all tarsi three-segmented; pretarsal claw with two preapical denticles; pulvillus not visible. Abdomen 0.5 mm long; genital structure poorly visible. Discussion. Following Mockford (1993), Burmacompsocus gen. nov. falls within the suborder Troctomorpha because of the following characters: antenna with 13 flagellomeres; at least the fourth and fifth being annulate; tarsi three-segmented; fore wing without sclerotized pterostigma. Following the key
Fig. 1. Burmacompsocus perreaui gen. et sp. nov., holotype Burm1. A, C, photograph and drawing respectively of dorsal view. B, photograph of ventral view. D, drawing of wings. Scale bars represent 1 mm.
Fig. 2. Burmacompsocus perreaui gen. et sp. nov., paratype Burm2, photograph. Scale bar represents 1 mm.
A. Nel, A. Waller / Cretaceous Research 28 (2007) 1039e1041
to modern families of Smithers (1990), Burmacompsocus would fall in the Compsocidae Mockford, 1967 because of the following characters: macropterous; tarsi three-segmented; body and wings without flattened scales; fore wing venation complex; pterostigmal area not more opaque than rest of the membrane; fore wing with nodulus; fore wing with two anal veins; hind wing with M forked. Mockford (1967) gave the following diagnostic characters for this electrentomoid family: antenna with 13e14 segments (13 in Burmacompsocus); frontal suture present, as in Burmacompsocus; lacinia tip with lateral cusp curved outward (present in Burmacompsocus), bearing few indistinct rounded denticles (no secondary tooth visible on cusp in Burmacompsocus); row of denticles present on anterior carina of fore femur (visible in Burmacompsocus); tarsi three-segmented (as in Burmacompsocus); pretarsal claw with two preapical denticles (visible in Burmacompsocus); mesothoracic tergal lobes not protruding above a line drawn from vertex of head to dorsal surface of abdomen (not applicable to our fossils); pterostigma closed basally (as in Burmacompsocus); vein M in hind wing one-branched (in fact two-branched, as in Burmacompsocus); first segment of Rs in hind wing present or absent (absent in Burmacompsocus); third ovipositor valvula deeply bilobed apically (not visible in Burmacompsocus). The last diagnostic character ‘vein 2A of fore wing joining vein 1A’, is not shared by Burmacompsocus. The Compsocidae comprises the two modern genera Electrentomopsis Mockford, 1967 and Compsocus Banks, 1930, both from the Central America (Panama, Mexico). Burmacompsocus shares with Electrentomopsis the 13-segmented antenna (14 in Compsocus); absence of first segment of Rs in hind wing; micro-vestitute of fore wing membrane in form of short points (Mockford, 1967; Smithers, 1990).
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Burmacompsocus differs from both Electrentomopsis and Compsocus in that vein 2A does not join 1A. Acknowledgements We are very grateful to our colleague, Dr. Michel Perreau, for allowing us to work on the type material of Burmacompsocus. References Azar, D., Nel, A., 2004. Four new Psocoptera from Lebanese amber (Insecta: Psocomorpha: Trogiomorpha). Annales de la Socie´te´ Entomologique de France 40, 183e190. Baz, A., Ortu~no, V.M., 2001. A new electrentomoid psocid (Psocoptera) from the Cretaceous amber of Alava (northern Spain). Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin, Deutsche Entomologische Zeitschrift 48, 27e32. Grimaldi, D.A., Engel, M.S., 2006. Fossil Liposcelididae and the lice ages (Insecta: Psocodea). Proceedings of the Royal Society of London, B 273, 625e633. Lienhard, C., 2002. Classification of Psocoptera with checklist of genera (including Cenozoic and Cretaceous fossils, in amber) compiled by Charles Lienhard. Natural History Museum, Geneva, Switzerland. http://www. ville-ge.ch/musinfo/mhng/page-e/artowho.htm#lienhard. Mockford, E.L., 1967. The electrentomoid psocids (Psocoptera). Psyche 74, 109e165. Mockford, E.L., 1993. North American Psocoptera (Insecta). Flora; Fauna Handbook 10. Sandhill Crane Press, Gainesville, Florida, xviii þ 455 pp. Perrichot, V., Azar, D., Neraudeau, D., Nel, A., 2003. New Psocoptera in the Lower Cretaceous ambers of southwestern France and Lebanon (Insecta: Psocoptera: Trogiomorpha). Geological Magazine 140, 669e683. Smithers, C.N., 1990. Keys to the families and genera of Psocoptera (Arthropoda, Insecta). Technical Reports of the Australian Museum 1990 (2), 1e82. Yoshizawa, Y., 2005. Morphology of Psocomorpha (Psocodea: ‘Psocoptera’). Insecta Matsumarana, New Series 62, 1e44.