Pl83485 Pl83·l86 Invertebrates, mainly insects, from the freshwater,

Pl83487 Lower Cretaceous, Koonwarra Fossil Bed PI83f88 Pl83189 ( Group), South , Pl83il90 PETER A. JELL AND PETER M. DUNCAN Pl83~91 Pl83t92 Pl83493 JELL, P.A. & DUNCAN. P .M., 1986:7: 10. Invertebrates, mainly insects, from the freshwater, Lower Cretaceous, Koonwarra Fossil Bed (Korumburra Group), , Victoria. PI8lt94 Mem. Ass. Australas. Palaeontols 3, 111-205. ISSN 08!0-8889. Pl83495 The Lower Cretaceous Koonwarra Fossil Bed, encountered in a road cutting south of on the , yielded more than 80 species of invertebrates. More than 70 are insects, associated with an ostracode, a syncarid (Koonaspides indistinctus gen. et sp. nov.), an anostracan and a cladoceran among the Crustacea, two spiders and a harvestman among the arachnids, possible earthworms, bryozoan statoblasts and a bivalved mollusc. The insects represent 12 orders dominated by the Hemiptera, Coleoptera and Diptera in terms of diversity but dominated numerically by aquatic immature Ephemeroptera and Diptera; Odonata, Blattodea, Plecoptera, Orthoptera, Psocoptera, Mecoptera, Trichoptera, Hymenoptera and probably Siphonaptera are lesser components. Insects, identified specifically and described taxonomically, are PromirarQ cephalota gen. et sp. nov., Australurus plexus gen. et sp. nov., Dulcim4nna sculptor gen. et sp. nov., Peraphlebill tetrastich/a gen. et sp. ·nov., Eodinotoperla duncanae gen. et sp. nov., Edgariekill una gen. et sp. nov., Duncanove/ia extensa gen. et sp. nov., Choristopanorpa drinnani sp. nov., Neoparachorista cfarkae sp. nov., Prochoristella /eongatha sp. nov., Cretacochorista parva gen. et sp. nov ., Tarwinia austral is gen. et sp. nov., Niwratla elongata gen. et sp. nov., Chironomaptera collessi sp. nov., Pseuda/ysiinia fragmenta sp. nov., Atherimorpha festuca sp. nov., Eoichneumon duncanae gen. et sp. nov., Westratia nana gen. et sp. nov., and Cretaco/ormica explicata gen. et sp. nov. However, the majority of the fauna is left in open nomenclature because features of generic or specific importance among living relatives are not clearly preserved on the fossils. The emphasis is on immature stages of the insects; some of the fossils are more or less in life position: in several instances larval and adult stages may be referred to the same species. Most significantly, many of the insects are referred. to living families; closer relationship to the living insects may be anticipated when more features used in modern taxonomy become available on the fossils. This fauna provides the first real insight into the antiquity of the living insect fauna of . The palaeoenvironment indicated by the invertebrate fauna is a shallow, semi-isolated body of water marginal to a shallow freshwater lake with periodic, probably seasonal, replenishing of the fauna from the lake and periodic mass-mortality of the isolated fauna; the mechanism for this mortality remains uncertain. A present-day environment in Lake Muirhead ~ust east of the Grampians in western Victoria is suggested as possibly comparable to the fossil setting. P.A. Jell, Queensland Museum, P.O. Box 3QO, South Brl.sbane, Queensland, Australia 4101; P.M. Duncan, Glenburnie Road, via Whittlesea, Victoria, Australia 3757; received 4 September, 1985.

FOSSIL INSECTS have been known from the Queensland the best known. The only Cretaceous Australian continent since I 870 but it was not until form described to date is .a dragonfly wing from this century that extensive studies by Tillyard, the marine Lower Cretaceous of nonh Queensland Riek, Evans, Dunstan and others made known the (Woodward, 1884). Riek (1970b) and Willmann diverse Permian and Triassic insect faunas of (in Hennig, 1981, p.83, note 98) briefly alluded to southern Queensland and New South Wales. the Koonwarra fauna described below. .Vinually all of the living orders of insects are The Lower Cretaceous Koonwarra Fossil Bed represented among these Permian and Triassic was discovered in 1961 by Country Roads Board {aunas but because no representatives of living of Victoria workmen straightening and widening families are present, they provide minimal a section of the South Gippsland Highway 3km information on the antiquity of the living east of Koonwarra and 142km from ; Australian fauna. Discovery of post-Triassic the site appears as NMVPL425 on the Museum of faunas is necessary to shed any light on this Victoria Palaeontological Locality Register and all question. A brief review of Australian fossil insect material described below comes from this one faunas (Riek, 1970b) indicated a relatively poor locality, although from several different levels record, with those from the Tertiary of southern within the Sm bed. The sediments have been dated 112 P.A. JELL AND P.M. DUNCAN

as late Aptian in age using macro and micro plant relationship may be even closer than thal fossils (Dettmann, 1963, this memoir; Douglas, has been ·suggested by their tentative chlSSi 1969, 1973; Douglas in Douglas et al., 1976). if more of the diagnostic characters used la Accounts of the fish fauna, the geological setting, accepted classification of living species had the age, and the palaeoecology as well as a available from the fossils. In this res measured section and detailed locality maps of the indicates that the modern Australian insect f. site were provided by Waldman (1971) who is at least 120 million years old and that referred to the fossiliferous horizon as the transition from or demise of the distinctive T · 'Koonwarra fish-bed'. We prefer the more general fauna probably occurred in the Jurassic; di 'Koonwarra Fossil Bed' in so far as the fish are of Jurassic insect faunas in Australia win not dominant either in numbers of species or in necessary to further define the event. In a nu numbers of individuals; they may have provided of the fossil species it is possible to associate the greatest amount of biomass but even in that adult insect with immature aquatic stages (e.1.- ·. regard vegetable matter may have been more some of the midges there are pupae and adulu · '' significant. in the beetles larvae and adults) which is a Although discovered more than 20 years ago, without precedent in the fossil history or the extensive Koonwarra insect fauna has received insects. This relatively complete fauna or only passing comment to date (Carroll, 1962; insects of a shallow lake enables comparison Talent, 1965; Talent et al., 1966; Riek, 1970a, rates of evolution, of a number of different i 1970b; Waldman, 1971). Insect specimens from orders, over a considerable time span; 120 mi Koonwarra in the collections of the Museum of years is approximately one-third of the total Victoria numbering several thousand, thanks to span of insect life and half the duration the painstaking preparation by P.M.D. over many stabilized insect evolution. ae, pupa1 years, have been considered the most interesting The fauna contains several invertebrates 01 rrai::an, th\ part of the fauna from 'one of the great fossil than insects that are of considerable signific '·.·. stai.. oblast~ localities of the Mesozoic Era' (Dettmann & with regard to the evolution of their respect ·"

under the action of directional winds. The lake along stream margins and in slow-flowing pools, ssifid1tioo . inhabitants should be most numerous under these while the other two have been compared with sed iii 11le .conditions with the stream and terrestrial Coloburiscoides Lestage 1935 that clings to rocks had lieca components much rarer; this is found to be the in fast riffles), and of the simuliid larvae. The resptj:t it case (Table I). simuliid was probably a sand-dwelling form sect fiiuna similar to living ones that occur in the sandy l thai the sections of some streams just before they enter a ve Triassic The fauna of the lake was apparently very similar lake, or during drier periods, where the now discdvery to that of present-day, shallow, cold, freshwater disappears into the sand. a wih be lakes of southeastern Australia including Tas­ a nufilbcr mania. This component includes both benthos and The terrestrial fauna :ociat~ Ille plankton and both may be represented by dead Only a small part of what must have been an . es (e.i!., in bodies er moulted cuticles. For example the extensive terrestrial insect fauna is preserved. The adult~and complete pupae of chironomids are preserved in semiaquatic gerroid waterbugs probably lived in is allnosa lateral aspect with the abdomen curved (Fig. 57E­ marginal aquatic sites or on plant-covered water ry of the l)whereas cast pupal cuticles from which the adult surfaces around the margins of the lake in habitats 1a of lhe emerged on the surface of the lake arc preserved similar to those of their modern relatives Jarisdn of in dorsoventral aspect with the abdomen stretched (Andersen, 1982, p.264). The rhagionid dipteran, rent ihsea. out straight and all intersegmental membranes the dragonfly, and the adult scorpiontlies all lived 20 million fully extended (Fig. 57A-D). It would be easy to on the wing above the lake or its feeder stream, or total.lime in1erpret these two forms as separate species. in the surrounding vegetation. The carabid and 1ratioh of The plankton included chaoborid larvae and staphylinid beetles probably scavenged in the litter 12upae, pupae of chironomid midges, the along the water's edge. Among the parasitic wasp inostracan, the conchostracan, the cladoceran and fossils are some whose living relatives attack wood­ che statoblasts. boring beetle larvae; they are active fliers or small The bottom fauna in the shallows would have enough to be carried by the wind. The pulicid fleas i\icluded the immature stages of the mayfly tend to indicate the presence of marsupials and/ Auslralurus plexus gen. et sp. nov., the dragonfly, or some other terrestrial vertebrates; to date the lhe scorpionfly Nannochorista sp., the stonefly, site has yielded only a few bird feathers (Talent et the caddisfly and the dytiscid and hydrophilid al., 1966; Waldman, 1970) to represent that group beetles. In the weeds, probably in slightly deeper but a substantial contemporary reptilean fauna is water, would have been the coenagrionid being discovered in Victoria (Molnar et al., 1981) .. damselfly larvae, some hydrophilid beetles, ~tracodes and possibly some Australurus. The ehironomid larvae, which are rare in the collection Site palaeoecology although the free-swimming pupae are common, ~1·ould have burrowed in the mud, especially in the Waldman (1971) discussed the palaeoecology of kss consolidated zone some distance from shore. this site at great length, stating in his abstract that this may also have been the home of the syncarid it was 'lacustrine or paludal, and mass mortality d of the xiphosuran if indeed the latter was a occurred periodically due to anoxic winter ptrmanent inhabitant of the lake. conditions beneath an ice-cover; the phenomenon known as 'winter kill'.' The transported stream fauna We agree with Waldman's conclusions that:- Species from the inflow stream were represented 1. The palaeo-environment formed a shallow part only by moulted cuticles which were carried either of, or was periodically linked with, a larger body in the body of the stream or on the surface, of water (e.g. a lake) from which it received a ~y depending on the method of ecdysis and the periodic repopulation of its aquatic fauna. 2. ount of agitation through the course of the Conditions within this environment were of the ream. Those cuticles that floated were more still-water type, or nearly so, for most of the period ely to be preserved, and to be preserved intact of sedimentary deposition. 3. Mass-mortality of an those that travelled submerged and tended to fish fauna occurred periodically. His further tie to the lake floor as the flow rate dropped, conclusion that a surface film caused the death of tling into situations not conducive to good the fauna and caused the environment to become ossilization. The fauna from the inflow stream is anaerobic, as well as his preferred option of snow resented by moulted cuticles of the immature covered ice forming the surface film, is quite ages of the mayflies Promirara cephalota gen. et plausible. Indeed the siphlonurid mayflies live nov., Dulcimanna sculptor gen. et sp. nov. and today in cool mountain streams and lakes in hlonuridae? gen. nov. (the former having central Tasmania and the cantharid beetle being finity with Mirawara Harker 1954 that lives similar to a species that swarms, at times, in the 114 P.A. JELL AND P.M. DUNCAN

A B c EPHEMEROPTERA SIPHONAPTERA Promirara cephalota c n Tarwinia australis u Auslra/urus plexus c n s Niwralia elongala .u Dulcimanna sculptor r n Pulicid indet. 1 u Siphlonuridae? gen. nov. r n Pulicid indet. 2 u ODONATA DIPTERA Peraphlebia tetrastk·hia r a,n Tipulid larva indet. u Coenagrionid indet. c n w Tipulid pupa indet. I u BLATTODEA Tipulid pupa indet. 2 u fl Methana sp. c a,n Tipulid? indet. u a Limoniinid indet. 1 r a PLECOPTERA unaf list for i Limoniinid indet. 2 u a :c = comml Eodinotoperla duncanae c a,n s • t = larva; pl Dixid indet. u a and preserv ORTHOPTERA Chironomaplera collessi c l,p = infauna; 1: Gryllacridoid? indet. u a Tanypodine indet. c a Tridactylid indet. u n Tanypodine pupa indet. l c p PSOCOPTERA Tanypodine pupa indet. 2 c p Edgariekia una u a Simuliid indet. I c a,I Simuliid indet. 2 u a ever, the o\ HEMIPTERA Pseudalysiinia fragmenta u a e, ad!.jlt inse~ Cixiid indet. u a Mycetophilidae gen. nov. u a wattrbugs: Cicadelloid indet. 1 c a erans f¥eopan A I herimorpha f estuca u a Cicadclloid indet. 2 u a islellq Jeong! Acalyptrate wings indet. r a Psyllid indet. u a nid, some ofi Brachyceran larvae indet. r I Anthocorid indet. u a ant) in~the sal Veliid indet. r a TRICHOPTERA the quesiio~ ofi line clay \amma1 Duncanove/ia extensa r a Calamoceratid pupa indet. u p inconi~eivabl~ Gelastocorid indet. r n of the wat, Homopteran indet. u a Oecetis? sp. u I rated ln the f! COLEOPTERA HYMENOPTERA into thq fine s! w back\vater ot Dytiscid indet. 1 r a,l s Eoichneumon duncanae u a and Aulacid indet. u a un~rowdei Dytiscid indet. 2 c a,! s Is, partiyularly • Dytiscid indet. 3 u a s Westratia nana r a dead anill'jals eitr Gyrinid? indet. a b Proctotrupid indet. u a , tend~, to su4 Hydraenid indet. u a Bethyloid? indet. u a t that affected ~ Hydrophilid indet. c a w Pemphredonine? indet. r a y not h11ve b Hydrophilid larva indet. r I w Cretacoformica explicata u a ebrate~. I~ thi! insects arid fish i Staphylinid indet. 1 r a ii bed wi'th the l Staphylinid indet. 2 u a CONCHOSTRACA n...... ,,." I Pselaphid indet. u a Cyzicus ? banchocarus rser laminae h Helodid indet. c I w Talent 1965 c a ambers, this m Cantharid indet. u a CLADOCERA Waldman · (197 I interpretati9ns on ~ Mordellid indet. u a Daphniid indet. c e,a p Coleoptera indet. c a,I tvidence we:mightJ ANOSTRACA However, we are MECOPTERA Anostracan indet. c a p possibility r}rovidel Choristopanorpa drinnani u a for the mixiit~ of'.~ Neoparachorista c/arkae u a OSTRACODA that explan\ttton 1~ Cypridoid indet. u Prochoriste/la leongatha u a a b Waldma'\1 (1981 Cretacochorista parva u a SYNCARIDA varves impl\ed traq Nannochorista sp. c I s Koonaspides indistinctus u a b dee~water yarves)I laminae in" sprin~

~w{ EARLY CRETACEOUS INVERTEBRATES, KOONWARRA 115

ARACHNIDA Oligochacte inde1. 2 c a u a Oligochaete indet. 3 u a Araneae indet. I u a BRYOZOA Araneae inde1. 2 u a Victalimulus macqueeni Phylactolaematid Riek & Gill 1971 u a b statoblasls c e p OLIGOCHAETA BIVALVIA Oliaochactc indct. I a f Unionoid indct. u a

Table/. Faunal list for Koonwarra Fossil Bed showing in Column A specimen numbers (u = unique specimen; r rare i.e. 2-5; c = common i.e. more than 5); in Column B growth stages reprcscmed (a adull; c = cphippium; 11 = nymph; I larva; p = pupa); in Column C ecological setting (i inllow; p = plankton; s = resident bent hos in shallows and preserved in place; w resident bemhos in weed bed or further offshore; b = u11diffcrcn1ia1~'tl bemhos; f infauna; t = 1crrcs1rial).

snow gum country tend to support Waldman's melting ice and the finer laminae laid down in theory of an ice cover on the lake during winter. autumn-winter. However, if they arc shallow However, the occurrence of well-preserved, water varves (Waldman, 1971, 1984), as seems complete, adult insects (e.g. the cockroaches, the most likely, then the coarser laminae will have gerroid waterbugs, the large beetles, the been deposited during and immediately after the mecopterans Neoparachorisla clarkae sp. nov. and greatest rainfall period. This means that the fine Prochoristefla leongatha sp. nov., the fleas, the fosil-bearing layers may not have been deposited rhagionid, some of the parnsitic wasps and the during autumn-winter as suggested by Waldman flying ant) in the same beds as the aquatic fauna (1984, fig. I) if the same rainfall pattern existed begs the question of how they came to be there if then as now. Predominantly winter rains would the fine clay laminae were deposited under ice. It mean fossil laminae deposition in summer-autumn seems inconceivable that they floated on the and make a surface ice cover less likely. Until the surface of the water before it froze, were not Early Cretaceous rainfall regime is belier incorporated in the frozen layer and then gradually understood the seasonality of the varves remains sank into the fine sediments on the floor of the in doubt. shallow backwater of the lake. Moreover, the more Another ecological observation worth noting is even and uncrowded distribution of invertebrate the extreme rarity (I valve only) of ostracodes. In fossils, particularly the aquatic nymphs preserved present-day lakes of southeastern Australia where as dead animals either on or through the fine clay flourish invertebrate faunas comparable to that layers, tends to suggest that the mass mortality from Koonwarra, ostracodes arc invariably a event that affected the fish fauna (Waldman, 1971) common element of the fauna. All indicators may not have been as catastrophic for the suggest that the lake at Koonwarra should have invertebrates. In this regard it should be noted that had a similarly rich ostracode fauna. The virtual the insects and fish are largely separated within the absence of ostracodes in the Fossil Bed suggests fossil bed with the insects and plants dominant in that the body of water in which those sediments the finer laminae near the base and the fish in were laid down was isolated from the main lake coarser laminae higher in the Bed (Drinnan & by dry land. This suggests possibly a levee along p Chambers, this memoir). In the same way that the mouth of an inflow stream isolating a pool that Waldman (1971) dismissed alternative received surface waters from the lake only when p interpretations on the basis of apparently contrary strong winds blew across the lake to form waves evidence we might now ignore his ice film theory. that carried the flotsam, plankton and some However, we are not prepared to dismiss it as a benthos over into the shallow isolated backwater. p possibility provided an explanation can be found As ostracodc shells sink after death they would for the mixing of the terrestrial and aquatic insects; have remained on the floor of the main lake. 1hat explanation is not apparent to us now. Clearly there is more work to be done on the b Waldman (1984) indicated that occurrence of intcrprctalion of palcoecology or the site; ii is not varves implied traditional seasonal control (i.e. for within the scope of this paper to begin that work b deepwater varves) on sedimentation with coarser as a considerable amount or detailed sedi­ laminae in spring-summer due to run-off from mentological study is necessary. 116 P.A. JELL AND P.M. DUNCAN

Modern analogy and is outside the scope of the present study. However, possible analogy is mentioned tO Lake Muirhead, a clay-pan just east of the promote study of that modern environment in tht, Grampians in western Victoria, has been suggested hope that it will shed further light on the.·· as having an ecological setting today that closely Koonwarra Fossil Bed. In particular it seems thll resembles that of the Lower Cretaceous 'The Clay-Settling Hypothesis' mentioned bf. Koonwarra Fossil Bed (E.F. Riek, pers. comm.). Waldman (1971, p.107) might well apply in rhil The Grampians area is an isolated mountain region marginal basin at Lake Muirhead, but faunal . reaching elevations of around IOOOm with steep content of the sediments in that basin needs study. eastern slopes descending abruptly to a badly drained peneplain at an elevation of 200-300m. The few small short streams have beds on the Systematic palaeontology slopes of boulders and pebbles of metamorphosed sandstone and siltstone but on the plain they have Throughout this paper we have, as far as possible, open shallow beds of sand size and smaller applied morphological terminology and particles entering large claypan regions within 3- classification used in 'The Insects of Australia' Skm. (CSIRO, 1970). Specimens described below all come from Lake Muirhead is one such shallow claypan that NMVPL425 in the Koonwarra Fossil Bed and are is fed by a single small broad shallow sandy housed in the following collections indicated by · stream, that is almost dry in summer and that has the prefixes (in brackets) attached to the no outlet stream. During the wet season sheet registration numbers:- (NMVP) Palaeontological·. nooding is general on the peneplain and the inflow Collection, Museum of Victoria, Melbourne; stream brings a component of fine sand and silt. (MUGD) Palaeontological Collection of the Most of the sand is deposited close to the inlet. Department of Geology, University of Melbourne: The lake is of such size and depth that wind action (AMF) Palaeontological Collection, Australian can produce obvious waves and cause water to Museum, Sydney. The bulk of the material back up on the lee shore, with periodic overnow described was collected by P.M.D. between 1962 into shallow marginal basins, which then retain and 1967. Over 300 specimens from the toral some water when the winds abate and lake levels collection of several thousand formed a numbered return to normal. These marginal basins develop (PDl-338) set containing most of the illustrated slight bars on their lakeward side. material. With the donation of the Duncan The sediment in at least one of these marginal Collection to the Museum of Victoria those basins has a varve-like character with a number of numbered specimens have been registered in the microlamellae distinguishable within the darker range NMVPI03001-103338 so that original ·· bands. These bands in the sediment are numbers are retained as the last three digits. Other undoubtedly due to changes in water level, but it specimens previously registered in collections of is not known whether these are due mainly to the Geological Survey of Victoria and of the inflow' into the basin or to changes produced Zoology Department at Monash University arc through wind action. Sedimentation rates in the now in the Museum of Victoria collections and re· lake are not known but it appears that wider bands registered accordingly. Suffixes 'A' and 'B' with ·, of coarser material represent wet-season inflow; a registration number indicate part and the microlamellae within the narrower fine zones counterpart of the same specimen. are correlated with smaller changes in level due to combination of overbank flooding and wind Phylum ARTHROPODA action. Prevailing winds would be uniform close to mountains so development of marginal basins Class INSECTA in which microlamellae could develop wo1.1ld be Order EPHEMEROPTERA (Mayflies) restricted to a small part of the lake margin. Superfamily BAETOIDEA Klapalck 1909 The fauna of Lake Muirhead is similar to that found in the Koonwarra Fossil Bed but it has not Family SIPHLONURIDAE Banks 1900 been possible to determine whether the PROM!RARA gen. nov. microlamellae at Lake Muirhead enclose any animals or not. Moreover, no data are available Etymology. From Latin prefix pro - before and Fig. I. P,.qmirara slight variation on generic name Mirawara. chlorjde NM' on the chemical environment of this marginal x4.(:,0,doi basin to determine whether it has all the necessary. Type species. Promirara cephalota sp. nov. E, Mrsal viei features to qualify as a direct analogy for the of ill(S:omple! Koonwarra situation. Assembling such data may Diagnosis. Based on nymph only. Similar to living require considerable study over a number of years Mirawara but with mouthparts less highly adapted EARLY CRETACEOUS INVERTEBRATES, KOONWARRA 117

yflie.•.) dck 1909

.S 19()0 F o - bcforr /. Promirara ceplwlotu gen. cl sp. nov. A , dorsal view o f almos1 comple1e holotypc whi1encd wi1h ammonium /firawura. d1loride NMVPI02472B, x 3.5. 13, dorsal view of almosl complete holotype in natural state NMVP102472A, x 4. C,D, dorsal views of counterparts of incomplete paratypc N:VIVP 102-t68B and A, x 6 and x 7, respectively. [/ sp. nm-. [,dorsal view of incomplete pnratype whitened with ammonium chloride NMVP 102473A, x 5. F, dorsal view of imcornplcte paratype counterpart in natural state NMVP 10247Jll, x 5. Similar 10 Ii\ ; highly ada 11 8 P.A. JELL AND P.M. DUNCAN

c

1. AJI. Si phi '<~IVP 1032• S MVPIOJ21 )mph MU• If, d,1rsal v ~l\'1'2250c EARLY CRETACEOUS INVERTEBRATES, KOONW ARRA 119

c

• J. A-0, Promirara cephalota gen. et sp. nov. A,B, camera lucida sketches from Fig. 21 and 2H, respectively. :; C,D, camera lucida sketches of very immature and mature nymphs from Fig. 2F and 20, respectively. E,F, , $iphlonuridae? gen. nov., camera lucida sketches of nymphs from Fig. 2A and 28, respectively. G,H, Dulcimanna sculptor gen. et sp. nov., camera lucida sketches of nymphs from Fig. 7E and 70, respectively. ,J.A,B, Siphlonuridae'? gen. nov. A, lateral view of mature nymph NMVP103210, x 9. B, lateral view of nymph NMVPI03209, x 7. C-1, Promirara cephalota gen. et sp. nov. C,D, dorsal view of mature paratype nymph NMVPI03208, x Sand x 3.$, respectively. E, paratype nymph NMVP27038, x 4. F, very immature paratype nymph MUG037S7A, x 10. G, dorsal view of paratype posterior abdominal segment NMVPI024678, x 7. H, dorsal view of anterior half of paratype adult NMVP22400B, x4, I, dorsal view of paratype adult NMVP22SOOA, x 3. 120 P.A. JELL AND P.M. DUNCAN

for predation. Abdomen with seven pairs of of a broad lamella with basai fibrilliform t subequal gills, first pair not reduced. Cerci with extending almost entire length of anterior gills hairs on the lateral as well as median margin but (possibly protruding at caudal margin) reduced hairs on lateral margin shorter and not extending posterior gills; caudal filaments short, less th to the base of the cercus. half abdominal length, with short articles; med' filament and inner margins of lateral ones Remarks. Promirara is so similar to Mirawara that dense long hairs; lateral filaments with d it may be considered ancestral to the recent genus. fringe of short hairs on outer margin over d Promirara was probably a less active swimmer portion but without hairs towards base. with less developed predatory habits than Mirawara, if one may judge from structure of the Remarks. The fossil nymph is similar to that abdominal gills and of the mouth-parts. living Mirawara (see Riek, l 955b) though there. Mirawara, which has a wide but discontinuous small but distinct differences. The mouthparts' distribution over much of the higher regions of highly modified, as in Mirawara, but the te · eastern Australia, is cold-adapted and needs well spines on the maxilla are shorter and the ter aerated water for survival. As there is no reason tooth on the mandible is shorter than in Mira to consider that these are recently acquired In Mirawara the first gill is reduced in size, t characters, it is reasonable to assume that in form it is not unlike that of the fossil s Promirara was also cold-adapted. Promirara has a broadly expanded area ant to the longitudinal ridge of the gill wh PROMIRARA CEPHALOTA sp. nov. (Figs I. 2C-l, Mirawara has a strongly sderotized ant 3A-D) margin developed apparently as a swimming Lateral caudal filaments of Promirara retain Etymology. Greek Kephalotos - with a head. primitive feature of hairs on the lateral marlf' · Material. Holotype NMVPI02472 an almost though the hairs are reduced in length and do . mature nymph 30mm long. Paratypes NMVP- extend to the base of the filament. Hairs ar0:.:· 22400, 22500, 27038, 102467, 102468, 102473, present on the lateral margin in Mirawara, th'" 102480, 102599, 102683, 103208, MUGD3753, they do occur to a slight extent in Tasmanoph 3757. Tillyard 1921, another Australian siphlonurid, in a few other siphlonurid nymphs. Description. Mature nymph more than 30mm long. Abdomen with distinct pigmentation AusTRALURus gen. nov. pattern, preserved only in very immature nymphs, consisting of a central irregularly pigmented area Etymology. Latin australis - southern and joined to a dark lateral area by an oblique arm on - water-bird. each side of each segment. Head distinctly wider than pronotum, with Type species. Australurus plexus sp. nov. large but not bulging eyes, appearing a little wider Diagnosis. Nymphs only. Head elo than long (possibly due to distortion upon mandibles large; antenna long; face compaction); labrum with anterior margin antennal insertions ridged. Legs long and,· appearing emarginate at meson; maxilla with long tibia shorter than tarsus; tarsal claw app .. ~ styliform terminal processes; mandible with long. Mesonotum raised and produced ca . heavily sclerotized terminal tooth; pronotum pronotum large. Abdomen with seven widening posteriorly but not strongly produced; unmodified lamellar abdominal gills; gill st wing pads extending to posterior of second or third imperfectly known but each gill apparently abdominal segment. Legs relatively long; femur strengthened longitudinal ridge at or close much wider than tibia-tarsus; tibia about equal in margin and possibly with a fibrillar tuft length to tarsus, with a slight transverse groove segments I to 9 strongly produced posterol close to its base. Abdominal segments with into spines. Caudal filaments with dense posterolateral corners produced into pointed of fine hairs except on outer margins o carinate tips, eighth and ninth segments having lateral filament. strongly produced almost spinose tips, with prominent median carina; gills on each side of first Discussion. This genus differs from the' seven segments, broadly lamellar, with strong Tasmanophlebia mainly in the gene curved ridge some distance behind anterior margin abdominal gills which occur, as is general (closer to margin on first than on other gills), and family, on the first seven abdominal scgme all subequal not reduced in size on posterior the first gill is not modified into a c segments; anterior gill sub-elytroid, more heavily succeeding gills as in Tasmanophlebia. This sderotized than posterior gills; each gill consisting seem to indicate that the fossil nymphs L/\RI.Y C RET1\ CTOUS INVERTUlRATES, KOO NWA RRt\ 121

lliform tuf1 -ior gills and ) redt.iced o -t, le~s than .. · .. . ' _-:, r '. :!cs; rnedi a11 . .. .. ,,_.. ,,,,,. . ~ 1 ones with with denst ove1 distal e.

r to that of gh thNeart 1th parts arc he terminal he tei"m inaJ 1 1\1irall'ara. .ize, though >si I species. ca anteri r ill whereas d anterior 1rn1ing aid. 1 retain the 11 margi!ll, and do nOI 1irs are noc •ra, though A 111opli!ebitl 1nurid, and

1 and ll/V

v.

• 4. A11stral11rus plexus gen. ct sp. nov. All in dorsal view. A, pararypc NMVP I02475, x7. n,c, part and counterpart of pararype NMVP22398, x 8 and x 9 , respectively. D, paratypc M UGD37488, x 7. E, paratype N:-.'I VP52550D, x 8.5. F,G, counterpart and parl o f holo1ype N\1VPI02450B a nd 1024501\, respectively, x 8. 122 P.A. JELL AND P.M. DUNCAN

partly bury themselves in sand or silt as is the habit 102457, 102465, !02475, 102476, !02478, 102479, of nymphs of Tasmanophlebia. Tasmanophlebia 102486, 102492, 102538, 102619, 102893, 103120,. lives either at the margins of lakes or in sandy 103222, MUGD3747, 3748, 3754. This is an backwaters and some silty areas of small, well­ extremely common species and a further 100 or. aerated streams; the species are all cold-adapted. more specimens are housed in the Museum of Although the mesonotum is extended in the Victoria. fossil there is no suggestion of dorsal median spines on the basal abdominal segments as in Diagnosis. As for genus. nymphs of Tasmanophlebia. There is no Description. Length of mature nymph (with dark. suggestion of a dense marginal fringe on the outer wing-pads) 9- IOmm. Abdomen with distinct margin of the cercus in the fossil species; such a pigmentation pattern consisting of a median and fringe is present in Tasmanophlebia. Although the a pair of lateral longitudinal bands joined at t fossil species is similar to Tasmanophlebia it is anterior margin of each segment. Medi probably not directly ancestral because, although longitudinal band indistinct on two caud most of the specialized features of Tas­ segments. Lateral regions unpigmented. Vertex O manophlebia occur in more generalized form in head apparently deeply pigmented or heavil Australurus, the cerci, lacking a lateral fringe, are sclerotized. Wing-pads on almost mature nym more specialized and have developed to the with rour main pigmentation lines correspondi condition that is of almost uniform occurrence to R, M, CuA and A. throughout the Siphlonuridae. Head elongated, often preserved wl The fossil nymph can also be compared with mandibles displaced laterally; mandibles I the nymphs of Baetidae, especially Centropti/um with transverse groove or ridge about mid Eaton 1869 which occurs in similar habitats along distinctly longer than wide, tapering only slig the margins of present day lakes, but the fossil to apex; antennae widely separated, first species is markedly larger, has well-developed possibly second segment enlarged, flagel pleural spines, relatively large gills, a broad distinctly longer than distance between ant clypeus and labrum, and the labrum is apparently insertions; face, above antenna! insertions, not deeply emarginate at the apex. an oblique curved ridge from just mesal Jurassic siphlonurids discussed by antenna! insertions to inner margin of eye; Sinitshenkova (1985) from Siberia and Mongolia large, oval, extending to caudal margin of h bear certain similarities to A ustralurus but each vertex slightly produced caudally at me may be distinguished by the combination of its clypeus relatively long; labrum apparently features. Australurus differs from Mesobaelis deeply emarginate at meson. Brauer, Redtenbacher & Ganglbauer 1889 by its Pronotum, seen better in lateral compress! spinose pleural tips, its more slender femora, its long, apparently slightly raised at meson cau elongate mandibles only slightly tapering and its posterolateral margin not strongly produ abdominal pigmentation pattern and from mesonotum large, posteriorly obviously raised,. Mogzonurella Sinitshenkova 1985 in its spinose produced at apex almost to a point; m pleural tips, more elongate head, relatively longer rimmed, with a ridge close to and convergf and more slender legs and relatively shorter caudal meson to the apical rim so that the apical filaments. The specimens figured by Hong (1982a, of the not um appears to have. two grooves on pl. 18, figs 2,3) as Ephemeropsis trisetalis Eich­ side converging to apex; wing-pads not ext wald from Upper Jurassic strata in the Neimenggol beyond apex of third abdominal se Autonomous Region of China also bear converging slightly to apex; inner margins st resemblance to Australurus particularly in the apex almost pointed. Legs long and t short caudal filaments but their pleural tipsdo not expanded; trochanter small; femur not appear to be spinose, their gills appear to be wide as tarsus; combined tibia-tarsus ( strongly supported and features of the head are claw) longer than femur; tarsus of fore and .·· not evident. legs about 1.5 times as long as tibia; ti~ 1 . tarsus tapering slightly; tibia with a deep I · AUSTRALURUS PLEXUS sp. nov. (Figs 4-6) groove on upper surface close to joint with .. ; Etymology. From Latin plexus - braided or tarsal claw stout, tapering regularly, lensti;, interwoven. fully preserved though apparently long. ;'' \ Abdomen with basal 9 segments subeqwil.; A ustrQlurus Pl x 6. B,~, part~ Material. Holotype NMVPl02450, a mature segment as long as second; segments 2-9 canal) !iMVPlq female nymph almost complete, except for apex posterolateral margin produced to a Iona, NMVP1'02478, i of caudal filaments and apical parts of hind legs. spine; lateral margins of abdomen .. NMVP!02455,,, Paratypes NMVP22398, 486!0, 52550, 102455- abdomen widest at segments 4 and 5: EAllLY C RETACEOUS INVERTLBRATLS, KOONWARRA 123

102.i i 1031. IS IS r 100 scum

rcssiol'I ·aud31l oducC'd .ised and

:rging II rcgi

Fig. 5. Australurus plr.rns gen. et sp. nov. All in dorsal view. A, 1wo paratypcs Ni'v!VP102457 (left) and NMVP 102456, .1al, fo x 6. B,C, part and counterpart of paratype showing dark central organ through abdomen (probably alimentary -9 \\idi canal) NMVP 102492A and 10249213, respcclively, x 6. D, incompi.:tc paratype N:\1VP 103222, x 10. E, para type g, sh NMVPI02478, x 7. F, paratypc N1vlVPI02538B, x 8. G, paratypc NMVPI02486A, x8. H , paratype con\ c l , NMVPI02455, x 10. i; 1cn 124 P.A. JELL AND P.M. DUNCAN

segment about 0.5 as Jong as ninth, its apex fine long hairs along lateral rounded though slightly produced; sternum of outer edges of cerci. ninth segment not produced over tenth segment; gills on segments I-7, lamellar, with strengthened Discussion. This genus, based only on the nymp fore margin, held laterally and slightly dorsal, and is assigned to the Siphlonuridae on its size, t either with a gill tuft at base or with a branched structure of the labrum· exposed anteriorly, tracheal trunk; caudal filaments only about 0.5 as abdominal gills and the distribution of hairs long as abdomen, with very short articles, with the caudal filaments. Posterolateral corners dense marginal fringes on median filament and abdominal segments are not spinose as in ot inner margin of each lateral filament; outer margin members of th~ family but this may be interpret of lateral filaments without obvious fringe even at as the ancestral state for the family. T apex. unsegmented tarsi distinguish it from Mirawa Immature nymph with only slightly raised the lack of hairs on lateral margins of the mesonotum and small wing-pads, and nymph separate it from Tasmanophlebia and usually preserved in lateral view. buriscoides is distinguished by its distinctive Comments on type specimen. Holotype (Fig. DULCI MANNA SCULPTOR sp. nov. (Figs 30,H, 4F,G) a mature female nymph, almost complete, except for apex of caudal filaments and apical Etymology. Latin sculptor- a carver. A noun portions of hind legs. Numerous paratype apposition. specimens, some preserving structures more distinctly than holotype. NMVP22398 (Fig. 48,C) Material. Holotype NMVPI02507. shows dark wing-pads of the mature nymph and NMVP103341, AMF66758. general body form. MUGD3748 (Fig. 4D) is a complete nymph that preserves the gills on both Diagnosis. As for genus. sides of the abdomen though they are not easily discernible on one side where they overlie the thin Description, Available specimens each 20mm I lateral nange of the segments. The displaced (without caudal filaments) and 2mm wide mandible and structure of the legs are also clear. thorax; head subrounded, a little longer than NMVP48610 (Fig. 6D) a lateral compression of an separated from pronotum by markedly const immature nymph showing segmentation of the neck; eyes large, bulging, situated dorsolat thorax. The head is particularly well preserved. at posterior of head; labrum with entire, NMVP102619 (Fig.'6G) shows abdominal gills and anterior margin; maxilla gently curved, head structure. spinose tip; mandible broad, with scleroti anteromedially. Pronotum with gently C() DULC!MANNA gen. nov. lateral margins; wing pads not extending first abdominal segment; legs stout, · Etymology. An anagram for Mrs llma Duncan, sclerotized longitudinal ridges on fe for her support during many years of collecting. becoming only slightly longer from fore to leg; femora all approximately same lengtll:·' Type species. Dulcimanna sculptor sp. nov. width; tibiae becoming longer and slightly st' Diagnosis. Nymph only known, large, markedly to .hind leg; tarsi unsegmented, becoming elongate body 10 times as long as wide. Head to hind leg: tarsal claw sclerotized, curved? rounded, relatively small, with large bulging dorsal relatively long. ,,, eyes, with entire anterior margin on labrum. Legs Abdomen long and slender, with sort;"' with femora of approximately equal length from evidenced by the many crinkles prestf · fore to hind legs, with tibiae and unsegmented tarsi formed by compression; abdominal gil · - becoming longer to hind leg, with strong curved segments I to 7, flat, lobate, with prominent relatively long tarsal claw. Abdomen with flat running down gill close to anterior mar,..1: lo bate gills on segments I to 7, with gently convex markedly sderotized ring around point of j internal margins not spinose posterolaterally. with abdomen; three caudal filaments Caudal filaments equal, bearing dense comb of without obvious segmentation, with fringe Cl( ~' :'.~~~

Fig. 6. Australurusplexus gen. et sp. nov. A, dorsal view of paratype MUGD3754B, x 11. B, dorsal view of NMVPI02893, x 7. C, dorsal view of paratype NMVP22398, x 9. D. lateral view of paratype NM~P x8. E, lateral view of paratype NMVP103120, x6. F, dorsal view of paratype NMVPI0246SA. II dorsal view of incomplete paratype NMVP102619A, x 8. H, dorsal view of para type NMVP 1024768, 1' dorsal view of incomplete paratype MUGD3747B, x 8. · · EARLY CRETACEOUS INVERTEIJRATES, KOONWARRA 125

margins except

only c•n the nym lae 011 its size, 1 sed anteriorly, r butioh of hairs )lateral corners ol ;pinose as in orhcr may be interpreted the . family. Tht it from Mirawaro, argin~ of the cer hfebia and Colo. its di ;tinctivc gilt .

>v. (Figs 30,H, 7)

l carv•:r. A noun 1

0250·1. Paraty

ns each 20mm Ion nd 2h1m wide c longer than \1 ick arkedly constricr ated dorsolater:ill vith entire, con\c 1tly •:urved, " ii 'lith sderotized hp ith g·~ntly com extending beyo legs stout, "ii ges on femor from fore to hi same length ind slightly stouter , becoming Ion tized, curved

:r, wlth soft 1 kl cs presuma jominal gills h prominent ri .erior margin j point of junc:f filaments eq with fringe or Ii 126 P.A. JELL AND P.M. DUNCAN

long hairs on each margin except on the outer the available specimens but this feature is not margins of lateral filaments. conclusive. Remarks. The three available nymphs are not sufficient to provide a complete picture of this Order ODONAT A (Dragonflies and damselflies». species because features such as antennae, some Suborder ANISOPTERA Selys 1840 mouthparts and most thoracic features are not clear. However, sufficient detail is available to be Superfamily LlBELLULOIDEA Tillyard 1926 confident it does not belong to any previously Family MESOPHLEBllDAE Tillyard 1916 described genus. The principal diagnostic feature of th S1rHLONURIDAE? gen. nov. (Figs 2A,B, 3E,F) Mesaphlebiidae is development of a wide spa with at least two rows of cells between R2 and R · Material. NMVPI03209 and NMVPI032IO both Although it may not be advisable to define a fami mature nymphs. on one character the fact that adult wing venatio Description. Nymph. Length to 6.5mm plus is known in each of the assigned genera only fro cerci of 2.5mm. Head rounded, mouthparts distal fragments and the venation that is kno conspicuous, occupying half area of head in lateral bears little n;semblance to other families in matt view; eye relatively small; clypeus and labrum of fine detail leads us to agree with Tillyard t small and narrow; labrum rounded at apex, about this group of Australian Mesozoic taxa constitut as long as wide; frons convex in lateral profile; a separate family. mandible large, subrounded except for tapered Tillyard (1916, p.24) made tentative assignm apex; maxilla, labium, and labial palps of this family (as a subfamily) to the· Anisopt conspicuous below mandible in lateral view, but later (1922, p.452) removed it to together occupying twice area of mandibles. Anisozygoptera although the heading of t Pronotum long, with prominent transverse carina section indicated the Archizygoptera. This may towards anterior and posterior margins, not assumed to have been a typographical error strongly produced posterolaterally, with transverse perhaps an indication of the uncertainty or posterior margin; mesonotum large, strongly placement. Whatever the intention of Tillyard extended posteriorly to reach as far as apex of fifth consider that his first opinion was correct. abdominal segment caudally; wing pads open venation resembles that of the Libelluloi conspicuous, apparently separated from one and Gomphidae; the brace vein continues the another (i.e. left from right) by extended of the base of the pterostigma as in Gomphi mesonotum, with eight clearly defined, radiating and in Synthemidae; the postnodals are incom veins; legs and sternum of abdominal segment 1 as in Synthemidae and Gomphidae. The nym not well defined on available material; sternum of mask is almost identical with that of many Ii abdominal segment I not attached to Libelluloidea. metasternum; segments 2 to 9 increasing regularly However the fossil species differs from in length so that 9 is twice as long as 2; without Synthemidae and Gomphidae in the cell prominent posterolateral pleural spines except development between R3 and IR3. There are t perhaps on segment 9 although some heavier rows of cells towards the margin and four r chitinization is evident in the lateral midline; the wing margin in the fossil species. T segment 10 short, less than half segment 9; caudal usually a single row in most living anisopt filaments moderately long, slightly more than half but there is a double row and occasional! length of abdomen. Gills not preserved. irregular third row at the wing margin in Libellulidae and Corduliidae. The fossil f Discussion. The body form of this species differs also in not having developed a sec resembles the living Baetisca Walsh 1862 and intercalary stem between R3 and R4 w Coloburiscoides of the Siphlonuridae. Since the present in Libellulidae and Corduliidae but latter occurs in Australia whereas the former is Synthemidae. On the basis of these comp ·. only known in cool temperate North America the Mesophlebiidae is tentatively placed witbid (Edmunds el al., 1976) it would seem much more Libelluloidea. likely that the similarities with Coloburiscoides would prove the more significant. PERAPHLEBIA gen. nov. The lateral midline darkening could reflect pleural spines because the same effect is seen in Etymology. Greek pera - beyond or very( lateral compressions of Australurus plexus where phlebos - vein. dorsoventral compressions confirm the presence of pleural spines. The gills may all have fallen from Type species. Peraphlebia tetraslichia sp. EARLY C RETACEOUS INVERTE£lRA TES, KOON\V ARRA 127 T. feature is

.nd damsclnicu 340 Tilly

milie'I in mancn ith Tlllyard I : taxa constitutes ative assig the A.nisop1rr1 ivcd it to 1 1eadirig of I era. This ma)' aphic ~ l error 1cert:fi nty of on of illyard •as correct. :he Libellulo· ontinlles the .sin Gomph· Js are incom 1e. Tile nym 1t of many Ii\·

ing anisopt occasional!) margin in 'he fossil f oped a scco nd R4 whl

md or ve ry; 7. Dulcimanna sculptor gen. et sp. nov. All dorsal views whitened with ammonium chloride. A, paratype r-;MVP 103341, x 4. B- D, pa rt and counterpart of holotypc NMVP 102507A (C) and 1025078 (B,D}, x 8, x 6 and x 4, respectively. E-G, part and counterpart of paratype AMF66758A (E,F} and 667588 (G}, x 4, x 5 •ichia sp. nm·. and x 4, respectively. Plants and invertebrates from the Lower Cretaceous Koonwarra Fossil Bed, South Gippsland, Victoria

P.A. Jell & J. Roberts editors

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I. i Lf ( ' / . _)

()O, 3 ,~;<_, {{!_ l/'J( ,,

Published by the Association of Australasian Palaeontologists Sydney 1986 R00177 85417 Cqnteii

FIQra o~ Sopth q AND~

Methods i Vicioriani The Otwd The GiPP.t Tal(onoml No(able ui Coinparisi Paffieoeni Sys.·tematil Dh1ision I JUNGERMj PLl}GIOC~ JU~GERMj R1qcARDl DE!-!DROd Memoir 3 of the Association of Australasian HErAnc~ Palaeontologists TH~.' LLIT~ Dh,Jsion ~ sPqRoPHJ ©AAP SPqROPHj GA¥ETO~ National Library of Australia card numbers GA~ETO~ ISBN 0 949466 02 6 BRVOPHYJ ISSN 0810-8889 Division~ LYCOPOq Isoj3TES rj Division$ I PH'(LLOT! EQ\JISETU Div.Jsion ~ SriiENOPJ GL\!ICHEtj AojANTni ONfCHIOi CL~DOP~ Typesetting by Press Etching Pty Ltd, 18 Little Edward Street, Spring Hill, Qld 4006. ~~.::.R~~tElDi"'.ision PH¥LLOP THfNNFEn TAl!NIOPl SMlNIA ~ Div.jsion q Printed by Gt~KGO u Watson Ferguson & Co., 35 Hamilton Road, Moorooka, Qld. 4105.