A New Collection of Insects from the Upper Triassic of South Africa
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1 Ann. Natal Mur. Yol. 22(3) Pages 791-820 Pietermaritiburg Novenibcr I976 1 A new collection of insects from the Upper Triassic of South Africa E. F. Riek (Division of Entomology, CSIRO, Canberra City, Australia) SYNOPSIS A second collection of insect fossils fro111 Bird's River and a single specimen from Mount Fletcher are described and illustrated. Meganisoptera, Odonata, Plecoptera and Lcpidoptera are recorded for the fauna, bringing the number of orders known to 11. Meganisoptcra arc rarely recorded from strata as late as the Triassic. Lcpidoptera have not previously been recognized in strata older than the Cretaceous. Large cicada-like Homoptera were a feature of thc fauna. A cercopoid honioptcran has unique lobcs on thc margin of the clavus that would producc a cryptic pattern when the wings are folded. The known fauna of 33 species from the Molteno Formation is listed. Fourteen species and a nymph have been described froin the Molteno Forniation (Zeuner 1939; Riek 1974, 1975). A new collection of 38 specilneiis from Bird's River (C-Dt 11) and one specimen from Mount Fletcher (C-MFI) includes 18 new species, and provides additional data on two described species. Mount Fletcher is a new locality in the Molteno for insect fossils. Insects are now known from nine localities in the Formation. Meganisoptera, Odonata, Plecoptera and Lepidoptera are recorded for the fauna, bringing tlie number of orders known to I 1. The dragonfly-like Meganisoptera are rarely recorded froin strata as late as the Triassic. The specinien is fragmentary, but this is not unusual with insect wings of large size. Large cicada-like Homoptera were diverse: they are represented by five species referred to three families. A dysmorphoptilid cercopoid has a unique lobed margin to the clavus that would produce a cryptic pattern when the wings were folded but the function of the lobes is not known. Lepidoptera have not previously been recognized in strata older than the Cretaceous. The Mesopsychidae, also known from the Upper Triassic of Australia, which are now referred to the Lepidoptera, have had a chequered career. They have previously been referred to a separate order Paratrichoptera, to tlie Trichoptera, to the Mecoptera, or considered closely related to Diptera. Most insects froin the fauna are of moderate to large size but there are also a few small Homoptera, Mecoptera and Coleoptera. The fauna is closely similar to that of the Upper Triassic of Australia although there are some notable additions especially Epheineroptera, Meganisoptera and Protozygopterous Odonata. The Plecoptera, Rlattodea, Orthoptera, Homoptera, Coleoptera and Lepidoptera are so similar to those of the Australian Upper Triassic that, unless informed of the country of origin, one would readily accept them as part of the Australian fauna of the period. 792 ANNALS OF 711E NATAL MUSEUM, VOI . 22(3), 1976 INSECTS KNOWN FROM THE MOI,TENO FORMATlON Ephemeroptera: Xenophlebioidca Xenophlebiidae X(~noj1h1cbiaoptatn Riek in pres~ Meganisoptera Paralogidae Triu,,rsologus hisrriutus gen. et sp. nov. Odonata: Protozygoptera Triassoneuridae nov. Tt,iassoncum crnck2rroni gen. ct sp. nov. T. heidiar sp. nov. T. rcgtrlariv sp. nov. Plecoptera ?Gripopterygidae Euxenoperlinae Euxenopcrla clur-a sp. nov. Blattodea Poroblattinidae Trinrsoblcrtta nntnlensis (Zeuner) 196 1 T. robz4,rtu Riek I974 T. ~i~nple.~sp. nov. Mesoblattinidae Snmcrroblattn par~wlusp. nov. Sun~aroblattellai.(~~lutcr gen. et sp. nov. ?Mantodea Mesorthopteridae Mesorthoptrron sin1ili.v Riek 1974 Orthoptera Haglidae Hqyln tontortu Riek 1974 Proparagryllacrididae Dorclrcthtin roblato gen. ci sp. nov. Geinitziidae Flr~tc~lzitziaj~ict~ir~tn gen. ct sp. nov. Hemiptera: Ho~noptera Mesogereonidae Triurrogcrron cli\ti~~ctnnzgen. et ~p.nov. Dunstaniidae Flc~tchcrianan~rigtln sp. nov. Dunstaniu pctropl~ilnsp. nov. Cicadoprosbolidae Pro t bolornorj~haclara Riek 1974 L~~pto~~rosbolclq,iilu gen. et sp. nov. Scy tinopteridae Srj~tinoj?t~rrxdrforta sp. nov. Dysmorphoptilidae Tenrzctittin gen nov. protubrrar~.\ Riek 1974 Protopsyllidiidae nymph Megaloptera Euchauliodidae Ec~cknuliock~sclistir~ct~i,~ Riek 1974 Mecoptera Mesopanorpodidae Afiistcllr~ck~licatula Riek 1974 RIEK: NEW UPPFR TRIASSIC INSECTS Lepidoptera : Paratrichoptera Mesoset~daenov. Mrsoses optata gen. el sp. nov. M. n~agnasp. nov. Coleoptera Cuped~dae Moltc~noc~lpestocsnrowi Zeuner 1961 Permosynidae Ademo yne .cl~cciosaRiek 1 974 A. prirca Riek 1974 A. reductn sp. nov. U~~~Xomau~riadt'prc.5 ,a Zeuner 1961 ?S~lphidae P,ccvu'o.sill,lzitc~~natu1cnsi.s Zeuner 1961 SYSTbMATICS ORDER MEGANISOPTERA The order 1s recorded froin the Upper C'1rboniferou5 of Europe and North Amer~ca, Lower PerinIan of North Anierlca 'ind the Urals, the Per~nianof the Archangel area, tlie Upper Perrn~dnof Australia and the Triass~cof France, and posc~blyfrom the Tr~ass~cof Germany although assignincnt of the very ~r~conipletewing of Relsla g~l~rcu(Rels) reinalns In doubt (Carpenter 1931). A cpecies ~c recorded from the Upper Trlassic oS soutl~ernAfrlca. The largest known insects were Meganiaoptera. Some speaes, however, were of a nioderate slzc comparable with liecent Odonata. The Megan~sopterahave a rather pr~m~tivehaw wlng ven'ltlon dlthough there are basal fusions between the veiny MP and CuA, and Cu and anal stem. The dom~nanttwo fam~l~cs,Paralog~dae and Meganeundae, are drctinguished on the branclung of Re and MA. The Tr~adotypldae have unu\ual brancli~ngof the cub~to-analfield The new species resemblec Paralog~daeIn the open venation, In whlch the armc of the first foiking of RE are d~vergentand MA 1s not expanded into a large sector before the middle of ~t\length. The precostal field I< short wlth Sc endlng about level with or before the first forhiog of Rs, as In Paralogidac but also 111 Typh~nac(Mega- neur~dae) The preco~t~ilfield 15 long In Meganeurinae. Fanlily Paralogidae Dirzgnocic. Venation open. Armc of tlie first forklng of Rs divergent. MA not expanded Into a large sector before thc middle of the wing. Precostal field short, w~thSc ending at or before the level of the firlt forking of Rs. Genils Triassologus &en. nov. Type .specic.s. Trias~ologusDiscrirrt~is sp. nov. Ditrgnoci\. S~lnilarto Ol~got~~p~i,and Puralogut a\ far as can be ascertdined from the ~re\ervedportion of the wing but w~tlia dlstlnct double row of cells between the uppernioct two branches oS Rs, lower branch of Rs dist~nctlyforked, and MA and MP ,uhl7,1rallel nntll nell after the first forking of Rs. The genus re>embles Olrgo~jy~l~rInore than Prrrnloguc. 794 ANNALS OF THE NATAL MUSEUM, VOL. 22(3), 1976 Triassologus biseriatus sp. nov. (Fig. 1 ; Plate 1, fig. 5) Type. C-Dt. 11 764, in Bernard Price Institute. Description. Apical half of wing but very incomplete caudally. Length as preserved c. 40 mm, indicating a wing length of c. 95 mm. Costal margin thickened, with a distinct median channel. Subcostal space narrowing apical with R very close to margin. Rs preserved only just distally of the first forking, the lower branch forking apparently before its middle and with the two branches divergent. Area between intercalated vein and upper branch of Rs with a double row of cells and area below with a single row. One cross-vein from R to the upper branch of Rs very much more oblique than the others. Cross-veins between MA and MP simple at least until the level of the forking of the lower branch of Rs. Cubital field only fraglnentar~lypreserved but branches curving strongly to posterior margin. ORDER ODONATA Suborder Protozygoptera The Protozygoptera differ from Zygoptera in not having a conlpletely formed nodus, the discoidal cell is not closed basally and the subnodus is either not developed or not fully aligned with the incipient nodus. The discoidal cell is not closed basally in the fore wing of the Recent species H[>rni- phlebia rnirabili.~Selys and Chorisnragrion risi Morton but the nodus and subnodus are fully formed. The d~scoidalcell is not closed basally in Perrnagriorz falltlarzdicz~m (Tillyard 1928) from the Upper Permian of the Falkland Islands and as the nodus is not completely formed it is preferable to refer the Perniagrionidae to the Proto- zygoptera although Tillyard had considered it as the oldest representative of the Zygoptera. Species from the Upper Triassic of southern Africa resemble Pennagrion,fulhla~~di- cunr in Inany characters, including the open disco~dalcell, but the nodus and sub- nodus are even less defined. However, the basal venation is even more reduced than in many Recent species. The species are referred to a separate famlly of the Protozygoptera. Family Triassoneuridae fam. nov. Protozygoptera with Sc extending allnost to the middle of the wing. The two ailtenodals situated towards the base of the costal space. Postnodals not in line with the cross-veins behind them. Subnodus not developed, the cross-vein more or less transverse. Cu+A free from MP only froin the lower corner of the discoidal cell. It would appear that the stem of Cu-i-A is anastornoscd with MP, to the discoidal cell, and not with the hind margin of the wing. This is similar to the condition in typical Protozygoptera and in Pcrmcrgrio~ except that the anastomosis is more extensive distally. The family differs from the Permagrionidae in the more distal position of the incipient node, the non-development of a subnodus, the non-alignment of the post- nodals with the cross-veins behind them and the anastomosis of Cu-+A with MP to the lower corner of the discoidal cell. RIEK: NEW UPPER TRIASSIC INSECTS Figs 1 -5. (1) fiicr.s.ro1o~rr.shiscr-icrtrrs gcn. et sp. nov. Holoty ~e C-Dt 11 764; (2) Tr-icrssoncrrrarmrk.rsof~i yen. et sp. nov. 1-lolotypc C-Dt 11 763 h; (3) Trrtr\.ronc~rrrtrl~c,it/icrc> sp.