© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at A new pygmy mole cricket in Cretaceous amber from Burma (Orthoptera: Tridactylidae) Sam W. HEADS Abstract: A new genus and species of Tridactylidae (Orthoptera: Caelifera: Tridactyloidea) is described from mid-Cretaceous Burmese amber. Burmadactylus grimaldiigen. et sp.nov. is the first tridactylid to be formally described from a Cretaceous amber and is assigned to the extant subfamily Dentridactylinae. The new genus is distinguished from all other Dentridactylinae by unusual- ly small male paraproctal lobes and represents the first record of an extant tridactylid subfamily from the Mesozoic. A key to the genera of Dentridactylinae is also provided. Key words: Tridactyloidea, Tridactylidae, Dentridactylinae, burmite, Myanmar. Santrauka:Remiantisvidurin(cid:0)s kreidos Birmos (Myanmaro) gintaru aprašoma nauja šeimos Tridactylidae (Orthoptera: Caelifera: Tridactyloidea) gentis ir nauja ru–šis. Burmadactylus grimaldii gen. et sp.nov. yra pirmoji Tridactylidae ru–šis apib(cid:0)dinama pagal Krei- dos gintaru˛. Ji priskiriama dabartiniu metu egzistuojan(cid:0)iam Dentridactylinae pošeimiui. Kartu tai pirmas atvejis, kai dabartinio Tri- dactylidae pošeimio r(cid:0)šis randama mezozojuje. Nauja gentis nuo visu˛ kitu˛ Dentridactylinae skiriasi ne(cid:0)prastai mažomis patino pa- raproktalin(cid:0)mis skiaut(cid:0)mis (paraproctal lobes). Taip pat pateikiamas Dentridactylinae gencˇiu˛ apib(cid:0)dinimo raktas. Raktiniai žodžiai:Tridactyloidea, Tridactylidae, Dentridactylinae, birmitas, Myanmaras. Introduction an skin (GRIMALDIet al. 2002). This incredibly diverse microbiota and a probable Turonian-Cenomanian age In terms of its biotic diversity, Burmese amber is by combine to make Burmese amber one of the most signi- far the richest Cretaceous amber in the world. The ficant Cretaceous ambers in the world (GRIMALDI & abundance, diversity and exquisite preservation of its ENGEL2005), though study of this extraordinary resour- biological inclusions have made Burmese amber the ce is still in its infancy. subject of intense scientific scrutiny in recent years1 Orthoptera are rare in amber, so it is perhaps not (GRIMALDI1996; ZHERIKHIN& ROSS2000; GRIMALDIet surprising that the study of orthopteran inclusions has al. 2002; GRIMALDI& ENGEL2005). To date, over 300 lagged far behind that of other insect groups. Only species representing some 130 families in 27 hexapod eight specimens were recorded from Burmese amber by orders have been recorded, amongst which are the ol- dest Strepsiptera (GRIMALDI& ENGEL2005) as well as GRIMALDIet al.(2002), all of which are small ensiferan nymphs. Indeed, adult orthopterans are extremely scar- the only Mesozoic Zoraptera (ENGEL & GRIMALDI ce in amber as they are usually large enough to free 2002) and Embiodea (ENGEL& GRIMALDI2006). In ad- themselves from resin traps. This complicates the study dition to hexapods, the amber has also yielded other or- of amber Orthoptera considerably as little can be glea- ganisms including bryophytes and higher plants (an an- ned from immature specimens beyond basic identifica- giosperm flower), nematode worms, pulmonate gastro- tion to family-group. Nevertheless, POINAR et al. pods, a remarkable peripatid onychophoran, myriapods, (2007) recently described Longioculus burmensis (El- numerous arachnids, a feather and fragments of reptili- canidae) from a small adult specimen previously figured by POINAR et al. (2005). Here, I describe a new genus 1Burmese amber, or ”Burmite“, was exported to China as early as the and species of Tridactylidae from Burmese amber, repre- first century AD and was highly sought after for use in jewellery or as a medium for sculptors (GRIMALDIet al. 2002). However, mining ceased senting the first Mesozoic record of the extant subfam- abruptly in the mid-twentieth century and it was not until comparati- vely recently (1999/2000) that interest from the Leeward Capital Cor- ily Dentridactylinae. AZAR& NEL(2008) described the Denisia 26, poration led to renewed mining and exploitation efforts. Important dentridactyline Guntheridactylus from Eocene French zugleich Kataloge der oberösterreichischen collections of Burmese amber inclusions are held at the American Mu- amber and additional tridactylids are known from both Landesmuseen seum of Natural History in New York and the Natural History Museum in London, and were summarised by GRIMALDIet al. (2002). Dominican (HEADS 2009, this volume) and Arch in - Neue Serie 86(27050–98)2: © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at geay ambers (PERRICHOT 2004). However, the current Tridactyloidea are an ancient group, though fossil paper constitutes the first record of an extant tridactylid representatives are extremely rare. The earliest definiti- subfamily from the Mesozoic. ve tridactyloids are known from the Early Cretaceous of Central Asia (Transbaikalia and Mongolia), Brazil and The specimen described here is deposited in the am- southern England (SHAROV1968; MARTINS-NETO1990; ber collection of the Division of Invertebrate Zoology GOROCHOV 1992; GOROCHOV et al. 2006) though the (Entomology), American Museum of Natural History, precise relationships of these primitive taxa remains un- New York (AMNH) and was studied using a Zeiss certain. GOROCHOVet al.(2006) place all the Cretace- stereo microscope. Drawings were made with the aid of a ous genera in the extinct tridactylid subfamily Mongo- camera lucida and photomicrographs produced using a scope-mounted Olympus digital camera. The age and loxyinae GOROCHOV, 1992. However, this subfamily is defined solely on the basis of plesiomorphic tegminal biotic diversity of Burmese amber are reviewed in detail venation and is almost certainly paraphyletic. More - by ZHERIKHIN & ROSS (2000), ROSS & YORK (2000), over, it is still unclear as to whether the Cretaceous ge- RASNITSYN& ROSS(2000) and GRIMALDIet al. (2002). nera represent stem-group Tridactylidae sensu stricto or are the stem-group to a clade comprising Tridactylidae Systematic account and Ripipterygidae. Tridactylidae were also reported from the Early Cretaceous Archingeay amber of south- Superfamily Tridactyloidea west France by PERRICHOT(2004) though they were not The superfamily Tridactyloidea comprises three fami- figured and remain undescribed. Guntheridactylus gri- lies: Tridactylidae BLANCHARD, 1845, Ripipterygidae AN- maultiAZAR & NELis the only tridactylid described so DER, 1939 and Cylindrachetidae GIGLIO-TOS, 1914. The far from the Cenozoic, though undescribed species of El- Ripipterygidae2 are very closely related to Tridactylidae lipes (Tridactylidae) and Ripipteryx(Ripipterygidae) are and have at times been considered a subfamily of the lat- present in Miocene amber from the Dominican Repu- ter (e.g. CHOPARD1949; RAGGE1955). They are never- blic and will be described elsewhere (HEADS in prep.). theless distinct from the tridactylids in having elongate Cylindrachetidae are at least as old as the Tridactylidae- mesotibia with parallel or near-parallel margins and ma- Ripipterygidae clade and although they are entirely un- les with one-segmented cerci. Cylindrachetidae (sand known as fossils, their disjunct austral distribution sup- gropers) are a peculiar group of fossorial insects known ports a Mesozoic origin and suggests that the family ra- only from Patagonia, Australia and New Guinea. Mono- diated prior to the break up of Gondwana. phyly of Tridactyloidea is supported by a number of mor- phological characters: [1] the prosternum connected di- Family Tridactylidae BRULLÉ, 1835 rectly to the pronotum by means of a precoxal bridge; [2] protarsus with two tarsomeres, inserted on the inner sur- Tridactyles BRULLÉ, 1835: Histoire naturelle des Insectes 9: 192. face of the protibia [3] mesotarsus with two tarsomeres; Tridactylites BLANCHARD, 1845: Histoire des Insectes, 246, [4] metatarsus always reduced to only one tarsomere; [5] 255–257. arolia entirely absent; [6] abdomen with nine fully sclero- Tridactylites SAUSSURE, 1874: Bulletin de Société Vaudoise des tised sterna in both sexes, the ninth forming a simple sub- Sciences Naturelles 13: 466–467. genital plate lacking styli; [7] abdominal repugnatorial Tridactylidae BRUNNER VON WATTENWYL, 1882: Prodromus glands; and [8] male paraproct with distinctive sclerotised der europäischen Orthopteren, 453. hooks. Members of the superfamily superficially resemble Type genus: TridactylusOLIVIER, 1789. certain Grylloidea (Ensifera) with tridactylids and ripip- terygids approximating true crickets (Gryllidae) and Nomenclatural note: Priority for family-group cylindrachetids remarkably convergent on mole crickets names based on the type genus TridactylusOLIVIER, 1789 (Gryllotalpidae). Indeed, the tridactyloid families have in dates from Tridactyles BRULLÉ, 1835 and not Tri- the past been classified along with the Grylloidea (TIN- dactylites SAUSSURE, 1874 as widely purported in the DALE1928) though their caeliferan identity has now been literature. Moreover, the name Tridactylites was first confirmed beyond doubt (ANDER1934). used by BLANCHARD (1845), almost 30 years before SAUSSURE(op. cit.). 2Ripipterygidae is commonly misspelled as ”Rhipipterygidae“. This is Diagnosis: Tridactylidae are distinguished from the due to an early misspelling of the genus-group name Ripipteryx NEWMAN, 1834 by AUDINET-SERVILLE(1838). This error was repeated Cylindrachetidae by their saltatorial metathoracic legs by SAUSSURE(1859) and gained wide usage during the first half of the with markedly enlarged femora and long, slender tibiae. 20th century. Indeed, ANDER(1939) used this incorrect spelling when forming the combining stem of the family-group name (i.e. Rhipipte- As stated previously, Tridactylidae are most closely re- ryg-). GÜNTHER(1994) subsequently recognised RhypipteryxAUDINET- lated to Ripipterygidae, but can be distinguished from SERVILLE, 1838as a junior synonym of RipipteryxNEWMAN, 1834 and the latter by the male cerci two-segmented. The family was followed by OTTE(1997), though misspelling of both the genus- group and family-group names is still widespread. is characterised by their very small size (a feature shared 76 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at with Ripipterygidae) and significantly reduced tegminal Diagnosis: As for the genus (vide supra). venation comprising between two and four longitudinal Description: =: body 3.77 mm long measured from veins. Members of the family are generally a glossy black the head to the abdominal apex; the specimen has un- and white with occassional brown/off-white patches and dergone some lateral compaction with its ventral and usually have fossorial prothoracic legs, often with a right lateral sides embedded in a large and turbid bubble number of dactylar processes present on the protibia. (Figs 1A-B, 2). Head: hypognathous, capsulate; vertex somewhat inflated anteriorly; frons broadly rectangular; Subfamily Dentridactylinae GÜNTHER, 1979 frontoclypeal sulcus prominent; clypeus quadrate, nar- Dentridactylinae GÜNTHER, 1979: Deutsche Entomologische rowing distally; labrum markedly smaller than clypeus, Zeitschrift 26, 258, 263. almost triangular; ocelli apparently absent; compound Type genus: DentridactylusGÜNTHER, 1974. eyes large and suboval; antennae moniliform, with ten articles; scape and pedical identical in size and shape to Diagnosis: The subfamily Dentridactylinae com- the flagellomeres; mandible largely obscured by tur- prises three extant genera, Bruntridactylus GÜNTHER, bitidy of the amber; maxillary and labial palpi long, with 1979, DentridactylusGÜNTHER, 1974 and Paratridactylus clavate terminal palpomeres bearing numerous small se- EBNER, 1943 and the fossil genus GuntheridactylusAZAR tae. Thorax: pronotum large, shield-like, extending pos- & NEL, 2008 from Eocene French amber, which are teriorly to entirely cover the mesonotum; margins of united based on the presence of a distinctive subapical pronotum with a prominent sulcus (Figs 1A-B, 2); denticular process on the metatarsus. prosternum largely obscured by the turbidity of the am- ber; tegmina strongly sclerotised though heavily distort- Genus Burmadactylusgen.nov. ed by compaction, with at least two longitudinal veins; Type species: Burmadactylus grimaldiisp.nov. by mo- hindwing absent. Legs: profemur robust, slightly curved notypy. distally, with a strongly concave ventral surface and very long external ventral setae (Fig. 3); protibia broad, Etymology: The genus-group name is formed from a strongly inflated distally with a dense covering of strong combination of ”Burma“ and ”dactylus“, a suffix com- setae (Fig. 3); basitarsus short, with two pairs of pulvil- monly used in the genus-group names of tridactylids. li; apical tarsomere elongate, slightly curved distally; Diagnosis: Burmadactylus is distinguished from all pretarsus with two prominent ungues; mesofemur other Dentridactylinae by the unusually small paraproc- markedly longer and narrower than the profemur (see tal lobes which are much longer (as long as the cerci) in Fig. 2); mesotibia similar in shape to the protibia but all other members of the subfamily. Moreover, Bur- more pointed apically; mesotarsus almost identical to madactylus can be separated from Bruntridactylus and the protarsus; metathoracic leg saltatorial; metafemur Guntheridactylusby the absence of metatibial swimming very large, as long as the abdomen and greatly inflated plates, and from Paratridactylus by the presence of along its entire length, with prominent dorsal carina; tegmina. The new genus is similar to Dentridactylus, par- external genicular lobe prominent, smooth, with sever- ticularly in the morphology of the metathoracic legs, al irregularly spaced setae (Figs 1A, 2); metatibia almost but can be distinguished from the latter by the small as long as metafemur, very slender, subquadrate in sec- paraproctal lobes and cerci with long setae. tion, with two parallel rows of 12 small and evenly spaced spines on the dorsal margins; two small subapical Burmadactylus grimaldiisp.nov.(Figs 1-4) spurs and two much longer apical spurs (Figs 1C, 4); Holotype: Adult = in Upper Cretaceous amber metatarsus a little over twice as long as the apical spurs, from Burma (Union of Myanmar); Katchin, Tanai Vil- with prominent apical (dent1) and subapical (dent2) lage, on Ledo Rd. 105 km NW Myitkyna, leg. R. denticular processes (Figs 1C, 4). Abdomen: terga and CRUICKSHANK, Leeward Capital Corp., 2000. AMNH sterna equally well sclerotised, narrowing posteriorly to- number Bu1452. Syninclusions: in addition to the afo- ward abdominal apex and somewhat distorted by lateral rementioned tridactylid, this piece of amber also con- compaction (Fig. 2); subgenital plate broad, without tains a small hemipteran nymph and several collembo- styli, bearing a marginal fringe of long setae; cerci long, two-segmented with numerous long setae; apical seg- lans representing at least two species. ment smaller than basal segment; paraproctal hooklets Etymology: The specific epithet is a noun in the entirely obscured by a large bubble; paraproctal lobe genitive case and is named in honour of Dr David cone-shaped, markedly smaller than the basal segment GRIMALDI (AMNH, New York) in recognition of his of the cercus, with numerous short apical setae. significant contribution to our knowledge of Burmese amber and its inclusions. 77 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Fig. 1: Holotype of Burmadactylus grimaldiigen. et sp.nov. (AMNH Bu1452); (A) left lateral view of habitus; optical distortion of the metatibial apices is caused by curvature of the amber surface; scale bar 1.0 mm; (B) right dorsolateral view of the head, thorax and prothoracic leg; scale bar 0.5 mm; (C) left metatibia and apex of metafemur; scale bar 0.5 mm. 78 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Figs 2-4: Burmadactylus grimaldiigen. et sp.nov. (AMNH Bu1452). (2) general habitus of the specimen; scale bar 0.5 mm; (3) right prothoracic leg showing the arrangement of setae; scale bar 0.25 mm; (4) apex of left metatibia (oblique dorsal view) showing details of the metatarsus, apical and subapical spurs; scale bar 0.25 mm. Abbreviations: asp, apical spurs of metatibia; dent1, apical denticular process of metatarsus; dent2, subapical denticular process of metatarsus; sasp, subapical spurs of metatibia. Fig. 5: Geographical distribution of Dentridactylinae. 79 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Key to the genera of Dentridactylinae wanan continents during the Cretaceous; a situation not dissimilar to that of the closely related tridactyloid fami- 1 Tegmina present . . . . . . . . . . . . . . . . . . . . . . . .2 – Tegmina absent . . . . . . . . .ParatridactylusEBNER ly Cylindrachetidae (GÜNTHER 1992; GRIMALDI & EN- 2 Metatibia with swimming plates . . . . . . . . . . .3 GEL 2005). The monotypic genus Paratridactylus is known only from Equatorial Guinea and is characterised – Metatibia without swimming plates . . . . . . . . .4 3 Antennae with only nine segments; tegmina with by the absence of tegmina and hindwings (GÜNTHER 1979, 1995). To date, the holotype male of P. eidmanni short longitudinal vein between costal margin and Sc and a straight longitudinal vein posterior EBNER, 1943 remains the only specimen of this aberrant genus and little is known of its affinities other than its of A1; metatibia with four pairs of swimming pla- placement in Dentridactylinae. The genus Bruntridacty- tes [Eocene French amber] . . . . . . . . . . . . . . . . . lus is more geographically widespread than the other . . . . . . . . . . . . . .GuntheridactylusAZAR& NEL genera, occurring in central and southeastern Africa, – Paraproctal lobes of = dorsoventrally compressed Turkey, Uzbekistan, India, Bangladesh, Burma and Tai- with numerous long setae, broad basally and narro- wan (Fig. 5). The fossil genus Guntheridactylusis known wing to a blunt apex; Ysubgenital plate not pro- only from the Eocene French amber (AZAR& NEL2008) jecting posteriorly . . . .BruntridactylusGÜNTHER and confirms the presence of Dentridactylinae in Europe 4 Paraproctal lobes of =near equal in length to the during the Cenozoic. Although the distribution of Den- cerci, cigar-shaped; both cerci and paraproctal lo- tridactylus suggests a gondwanan origin for this genus, bes without long setae; Ysubgenital plate gently there is no evidence to suggest that the subfamily was re- rounded posteriorly . . .DentridactylusGÜNTHER stricted entirely to Gondwana. On the contrary, thedis- – Paraproctal lobes of =much shorter than the cer- covery of Burmadactylusconfirms the presence of Dentri- ci, cone-shaped, with short apical setae; cerci dactylinae in Laurasia during the mid-Cretaceous. Mo- with numerous long setae; = subgenital plate reover, the patchy distribution of Bruntridactylus across with a fringe of long setae around the posterior Africa and Eurasia suggests that large-scale Cenozoic ex- margin [Cretaceous Burmese amber] . . . . . . . . .. tinctions might have contributed significantly to the . . . . . . . . . . . . . . . . . . . . .BurmadactylusHEADS present distributions of Old World Tridactylidae. In summary, Burmadactylus represents the first re- Discussion cord of an essentially modern tridactylid from the Cre- Tridactylidae are undoubtedly an ancient group taceous and confirms the great antiquity of this group. with a number of genera known from the Cretaceous Moreover, the presence of Burmadactylus in Laurasia (Cratodactylus MARTINS-NETO, 1990; Cretoxya GORO- challenges the supposed Gondwanan origin for the CHOVet al., 2006; MongoloxyaGorochov, 1992; Mono- Dentridactylinae as suggested by the austral disjunction dactyloides SHAROV, 1968; and Monodactylus SHAROV, of Dentridactylusand implies a formerly more widespre- 1968). However, all of these genera are characterised by ad distribution for the subfamily. GRIMALDI(1992) and plesiomorphic tegminal venation, comprising numerous GRIMALDI& ENGEL(2006) showed that fossil represen- longitudinal veins and a rich crossvenation. Based on tatives of crown group taxa can be used successfully to this feature, GOROCHOV (1992) and GOROCHOV et al. interpret the formerly more widespread distributions of (2006) united the Cretaceous genera in the subfamily Recent lineages. Nevertheless, a more detailed under- Mongoloxyinae, though its definition based on a plesio- standing of tridactyloid relationships is required in order to interpret fully the interesting distribution of these morphy renders the monophyly of this group highly fascinating and unusual orthopterans. questionable. Indeed, Mongoloxyinae most probably re- presents a paraphyletic assemblage of stem-group taxa that gave rise to the two modern tridactylid subfamilies Zusammenfassung (and possibly the Ripipterygidae) sometime in the late Eine neue Gattung und Art der Familie Tridactyli- Mesozoic. The discovery of B. grimaldiiconfirms the an- dae (Orthoptera: Caelifera: Tridactyloidea, Grabschre- tiquity of the Dentridactylinae and supports the asser - cken) wird aus burmesischem Bernstein (mittlere Krei- tion that they, along with their likely sister-group the de) eingeführt und der Unterfamilie Dentridactylinae Tridactylinae, diversified before the mid-Cretaceous. zugeordnet. Burmadactylus grimaldii gen. et sp.nov. ist Burmadactylus appears to be most closely related to der erste Tridactylide, der formal aus kreidezeitlichem Dentridactylus, with both genera having a very similar ar- Bernstein beschrieben wird. Die neue Gattung unter- rangement of spines and spurs on the metatibiae. The scheidet sich von allen anderen Dentridactylinae durch geographical distribution of Dentridactylus presents a ungewöhnlich kleine, paraproctale Loben bei Männ- classic example of austral disjunction (Fig. 5) and sug- chen. Ein Bestimmungsschlüssel zu den Gattungen der gests that the genus was effected by the drifting of Gond- Dentridactylinae wird präsentiert. 80 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Acknowledgements GRIMALDID.A. (1992): Vicariance biogeography, geographic ex- tinctions, and the North American Oligocene tsetse flies. — Many thanks go to Dr David GRIMALDI (AMNH, In: NOVACEKM.J. & Q.D. WHEELER(Eds), Extinction and Phylo- New York) for his encouragement and hospitality dur- geny. Columbia University Press, New York: 178-205. ing my visit to the AMNH in September 2007. I am al- GRIMALDID.A. (1996): Amber: Window to the Past. —Abrams so very grateful to Dr Michael ENGEL (University of and the American Museum of Natural History, New York. Kansas, Lawrence) and Dr David MARTILL (University GRIMALDID.A. & M.S. ENGEL(2005): Evolution of the Insects. — of Portsmouth) for comments on the manuscript, and Cambridge University Press, Cambridge and New York. Mr Robert LOVERIDGE (University of Portsmouth) for GRIMALDI D.A. & M.S. ENGEL (2006): Extralimital fossils of the assistance with photography. Thanks also go to Dr Dany “Gondwanan” family Sphaeropsocidae (Insecta: Psocodea). —American Mus. Nov. 3523: 1-18. AZAR (Lebanese University) and Dr André NEL (Muséum National d’Histoire Naturelle, Paris) for pro- GRIMALDID.A., M.S. ENGEL& P.C. NASCIMBENE(2002): Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, viding a copy of their manuscript on Guntheridactylus, biotic diversity and paleontological significance. —Ameri- and to Dr Andrej GOROCHOV (Zoological Institute, can Mus. Nov. 3361: 1-71. Russian Academy of Sciences, St Petersburg) for useful GÜNTHERK.K. (1974): Über die Tridactyloidea in den Sammlun- debate. Support was provided by NERC grant gen des Museums f. Naturgeschichte der Stadt Genf. — NER/S/A/2004/14238. Rev. Suisse Zool. 81: 1027-1074. GÜNTHERK.K. (1979): Einige Bemerkungen über die Gattungen References der Familie Tridactylidae BRUNNERund Klassifikation der Tri- dactylodea. —Deutsche Entomol. Z. 26: 225-264. ANDERK. (1934): Über die Gattung Cylindrachetaund ihre syste- GÜNTHERK.K. (1992): Revision der Familie Cylindrachetidae GI- matische Stellung (Orthoptera Saltatoria). — Arkiv Zool. GLIO-TOS, 1914 (Orthoptera, Tridactyloidea). —Deutsche En- 26A(21): 1-16. tomol. Z. 39: 233-291. ANDER K. (1939): Vergleichend-Anatomische und Phylogeneti- GÜNTHERK.K. (1994): Die Tridactyloidea-Fauna Kolumbiens (Or- sche Studien über die Ensifera (Saltatoria). —Opusc. Ento- thoptera, Caelifera). —Deutsche Entomol. Z. 41: 1-56. mol. Suppl. 2: 1-306. GÜNTHERK.K. (1995): Die Tridactyloidea des südlichen Afrika (Or- AUDINET-SERVILLE J.G. (1838): Histoire naturelle des Insectes Or- thoptera, Caelifera). —Deutsche Entomol. Z. 42: 213-286. thoptéres. —Librarie Encyclopédique de Roret, Collection HEADSS.W. (2009): New pygmy grasshoppers in Miocene amber des suites a Buffon, Roret, Paris. from the Dominican Republic (Orthoptera: Tetrigidae). — AZAR D. & A. NEL (2008): First Tridactylidae from the Eocene Denisia 26: 69-74. French amber (Insecta: Orthoptera). —Alavesia 2: 169-175. MARTINS-NETOR. G. (1990): Um novo gênero e duas novas espé- BLANCHARD E. (1845): Histoire des Insectes, leurs moeurs, leurs cies de Tridactylidae (Insecta, Orthopteroidea) na Forma- métamorphoses et leur classification. Volume 2. —Didot, ção Santana (Cretáceo Inferior do nordeste do Brasil). — Paris. Ann. Acad. Brasileira Ciên. 62: 51-59. BRULLÉA. (1835): Cinquième Ordre. Orthoptères [in Orthoptères NEWMAN E. (1834): Entomological Notes. — Entomol. Mag. 2: et Hemiptères]. —In: AUDOUINJ.V. & A. BRULLÉ(Eds), Histoi- 200-205. re naturelle des Insectes, 9. Pillot, Paris. OLIVIERA.G.(1789–1832): Encyclopédie Méthodique ou par ord- BRUNNERVONWATTENWYLK. (1882): Prodromus der europäischen re de matières; Histoire naturelle, 10 volumes. —Panckou- Orthopteren. —Engelmann Verlag, Leipzig. cke, Paris. CHOPARDL. (1949): Ordre des Orthoptères. —In: GRASSÉP.P. (Ed.), OTTED. (1997): Orthoptera Species File 6. Tetrigoidea and Tri- Traité de Zoologie 9: 617-722. dactyloidea and addenda to OSF vols 1-5. —Orthopterists’ EBNERR. (1943): Über einige Orthoptera Saltatoria von Fernan- Society and Academy of Natural Sciences of Philadelphia. do Poo. —Ann. Mus. Civ. Stor. Nat. Genova 6: 1-21. PERRICHOT V. (2004): Early Cretaceous amber from south-west ENGELM.S. & D.A. GRIMALDI(2002): The first Mesozoic Zorapter France: insight into the Mesozoic litter fauna. —Geol. Acta (Insecta). —American Mus. Nov. 3362: 1-20. 2: 9-22. ENGELM.S. & D.A. GRIMALDI(2006): The earliest webspinners (In- POINARG.O., R. BUCKLEY& A. BROWN(2005): The secrets of Burme- secta: Embiodea). —American Mus. Nov. 3514: 1-15. se amber. —Mid-America Paleontol. Soc. 29: 20-29. GIGLIO-TOSE.(1914): Sulla posizione sistematica del gen. Cylin- POINARG.O., A.V. GOROCHOV& R. BUCKLEY(2007): Longioculus bur- drachetaKIRBY. —Ann. Mus. Civ. Stor. Nat. Genova 6: 81- mensis, n.gen., n.sp. (Orthoptera: Elcanidae) in Burmese 101. amber. —Proc. Entomol. Soc. Washington 109: 649-655. GOROCHOVA.V. (1992): New fossil Orthoptera and Phasmoptera RASNITSYNA.P. & A.J. ROSS(2000): A preliminary list of arthropod from the Mesozoic and Cainozoic of Mongolia. —Sovmest- families present in the Burmese amber collection at The naya Sovetsko-Mongol’skaya Paleontol. Eksped. Trudy 41: Natural History Museum, London. —Bull. Nat. Hist. Mus., 117-121. [in Russian] London (Geol.) 56: 21-24. GOROCHOV A.V., E.A. JARZEMBOWSKI & R.A. CORAM (2006): Gras- ROSSA.J. & P.V. YORK(2000): A list of type and figured specimens shoppers and crickets (Insecta: Orthoptera) from the Lower of insects and other inclusions in Burmese amber. —Bull. Cretaceous of southern England. —Cret. Res. 27: 641-662. Nat. Hist. Mus., London (Geol.) 56: 11-20. 81 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at SAUSSUREH. DE(1859): Orthoptera Nova Americana (Diagnoses Praeliminaires). II. — Rev. Mag. Zool. 11: 59-63, 201-212, 315-317, 390-394. SAUSSUREH. DE(1874): Notice sur un Orthoptère du Pays de Gex, le Tridactyle (Tridactylus). — Bull. Soc. Vaudoise Sci. Nat. 13: 466-467. SHAROVA.G. (1968): Phylogeny of the Orthopteroidea. —Trudy Paleontol. Inst. Nauk, 251 pp. [in Russian] TINDALE N.B. (1928): Australasian mole-crickets of the family Gryllotalpidae (Orthoptera). —Rec. South Australian Mus. 4: 1-42. ZHERIKHINV.V. & A.J. ROSS(2000): A review of the history, geolo- gy and age of Burmese amber (Burmite). —Bull. Nat. Hist. Mus., London (Geol.) 56: 3-10. Address of author: Sam W. HEADS Palaeobiology Research Group School of Earth and Environmental Sciences University of Portsmouth, UK E-Mail: [email protected] 82