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Discovery and Analysis of the Oldest Mayflies (Insecta, Ephemeroptera) Known from Amber

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Discovery and Analysis of the Oldest Mayflies (Insecta, Ephemeroptera) Known from Amber THE OLDEST MAYFLIES FROM AMBER BulL Soc. His. Nut.. Toulouse, 133, 1997,77-82 Discovery and Analysis of the Oldest Mayflies (Insecta, Ephemeroptera) Known from Amber by W.P. McCafferty Department of Entomology, Purdue University, West Lufayette, IN (USA) 47907 Conovirilus poinuri (McCafferty n. gen. et n. sp. (family Leptophlebiidae) and Baetidae sp. 1 are described from adult mayfly fossils taken in Lebanese amber from the Lower Cretaceous. These mayflies represent the oldest mayflies known from amber and are significant in that they also represent the oldest corroborated fossils of their respective families. Baetidae sp. 1 cannot be resolved beyond family (obvious from its tarsal formula) because of the condition of the fossil specimen ;however, genitalia, leg and hindwing characterization available on the fossil of C. poinari allows the taxon to be placed to a relatively ancient and plesiotypic Southern Hemisphere clade of Atalophlebiinae genera that also includes Adenophlebia, and Aprionyx and the Atalophlebioides complex. The distribution and age of Conovirilus is consistent with that of the clade as can be deduced from phylogeny and extant distributions. The study exemplifies the predictive value of phylogeny and how it can be tested with paleontological data. Keywords : fossil mayflies, Leptophlebiidae, Baetidae, Lebanese amber, Conovirilus. D6maveri.e et analyse des plus densEphCm?ms (Insecta, Ephemeroptera) connus dam I'Ambre. Conoviriluspoinuri McCafferty n. gen. et n. sp. (famille Leptophlebiidae) et Baetidae sp. 1 sont dhits l partir d'CphCm&res adultes pris dans l'ambre Libanais, date du Cdtad InfCrieur. Ce matkriel repdsente les plus anciens Eph6mCropt2res connus dam l'ambre ;sa port& ap- par& aussi dam le fait qu'il s'agit Cgalement des plus anciens fossiles confirm% dans leurs familles respectives. Baetidae sp. 1 ne peut etre r6- solu au dell de la famille (Cvidente par la formule tarsale) en raison de l'ktat du sp6cimen. En revanche, la caractCrisation possible des genita- lia, pattes et ailes postkrieures du spkirnen de C. poinari autorise A placer ce taxon dans un clade, relativement ancien et plCsiotypique de l'HCmisph&reSud, de genres d'Atalophlebiinae qui inclut aussi Adenophlebia, Aprionyx et le complexe Atalophlebioides. La &partition et 1'8ge de Conovirilus sont compatibles avec ceux du clade ainsi qu'on peut le d6duire de la phylogknie et des rCpartitions existantes. Cette Ctu- de met en Cvidence la valeur pddictive de la phylogknie et la fa~ondont cette dernih peut Ctre test& par les donnks palkontologiques. Mots clCs : kphkmkes fossiles, Leptophlebiidae, Baetidae, ambre du Liban, Conovirilus. INTRODUC~ON ginally ascribed to the Oligocene may actually be Eocene Previous to this report, the known mayfly fauna from fos- (Schliiter 1975). Baltic amber genera include *Baetis Leach silized resins has consisted of 31 nominal species, represen- (Baetidae), Baltameletus Demoulin (Siphlonuridae s.l.), tative of 22 genera and 12 families. All of the families except Balticophlebia Demoulin (Arneletopsidae), Blastumphlebia Australiphemeridae are extant, and 10 of the genera pre- Demoulin (?Leptophlebiidae), Brevitibia Demoulin viously known from amber are also extant (see asterisked (?Ametropodidae), ?*Choroterpes Eaton (?Leptophlebii- genera, below). Other than one family, three genera, and five dae), *Cinygma Eaton (Heptageniidae), Cronicus Eaton species, this fauna has been described from Baltic amber, (Coloburiscidae), Denina McCafferty (Ephemeridae), which dates to the Oligocene and Eocene epochs [25-60 mil- Electmgenia Demoulin (Arthropleidae), *Heptagenia Walsh lion years ago (mya)] although some of the Baltic amber ori- (Heptageniidae), *Metretopus Eaton (Metretopodidae), 78 W.P. McCAFFERTY (2' *Paraleptophlebia Lestage (Leptophlebiidae), *Rhithrogena mya) (SCHLEEand DIETRICH1970). The new data are sign& Eaton (Heptageniidae), *Siphlonurus Eaton (Siphlonuri- cant because of the dearth of paleontological inforrnatim dae), *Siphloplecton Clemens (Metretopodidae), Succino- from this period as explained above, and because the leptw genia Demoulin (Heptageniidae), *Timpanoga Needham phlebiid fossil is in good enough condition to allow it to k (Ephemerellidae), and Xenophlebia Demoulin (?Siphlonu- precisely placed in the generic phylogeny of ridae s.1.). Leptophlebiidae. Furthermore, the evolutionary data haw Philolimnias sinica was described by HONG(1979) from important biogeographic ramifications, as will be shown be- Chinese amber dating to the Eocene (ca 50-60 mya). The low. single specimen upon which its description was based can ostensibly be assigned to the Ephemerellidae based on wing venation. However, it should be noted that other interpreta- 1. ConoviriZus McCafferty n. gen. (fam. Leptophlebii- tions of its morphology (e.g., mouthparts and abdominal ter- dae) :description minalia) are clearly not consistent with Ephemeroptera and Male adult are misrepresentations of more obscure portions of the fos- Compound eyes (Figs. 1-3) large, meeting on meson of sil, as can be seen from a whole-specimen photograph. head, dorsal three-fourths missing (Fig. 3). Species of Cretoneta Tshernova and Paleoanthus Kluge are known from Taymyr retinite (Siberian) from the Upper Frons with apical margin not deeply scalloped, shaped as Cretaceous (ca 70-100 mya). Kluge (1993) removed Creto- in Figure 3. neta from the family Leptophlebiidae and placed it in Forelegs and midlegs incomplete. Hindlegs (Figs. 1, 2) Siphlonuridae s.1. (viz., Pisciforma), and he placed Paleoan- with 4-segmented tarsus (Fig. 4) ; claws (Fig. 5) similar, ea- thus in a new family Paleoanthidae (Ephemeroidea). From ch apically hooked and with smaller opposing hook. its characterization, however, Paleoanthus actually falls wi- Forewing venation indeterminate. Hindwing costal margin thin the context of the previously known extinct family (Fig. 6) convex, with apex of convexity located slightly Australiphemeridae (see MCCAFFERTY1990). There may be beyond one-third distance from base ;venation indetermina- some other mayflies from Upper Cretaceous Siberian resins te. not accounted for here. Ninth abdominal sternum (Fig. 7) not enlarged posteriorly One of the least represented time periods in geologic his- and not cleft medially. tory with respect to mayflies is the Lower Cretaceous (ca Genitalia (Figs. 1,7) with forceps 3-segmented ; segment 120-135 mya). As pointed out by MCCAFFERTY(1900), this 1 elongate, slightly thickened at base ; segments 2 and 3 period is critical for assessing the biogeographic history of short, together approximately one-sixth length of segment 1. mayflies because it was prior to the final breakup of major Penes fused and conical ;apices obscured by exudate (Fig. 7, landmasses. It is also critical for assessing the fate of the pointers). Mesozoic fauna and origin of the Cenozoic fauna because it was just prior to the major biotic extinctions that took place Cerci and median caudal filament broken (median caudal by the end of the Cretaceous. Previous to this report only 13 filament represented by broken truncate stub (not tapered nominal species in 12 genera, representative of at least seven and rudimentary). families, were known from the Lower Cretaceous. Exempli- Larva and egg stages : Unknown. fying the transitional nature of this fauna is the fact that all of 2. 'Qpe species the known Lower Cretaceous genera had disappeared by the Cenozoic, but three of the seven families presently known Conoviriluspoinari McCafferty n. sp. from the Lower Cretaceous record are present today. 3. Generic etymology Australiphemeridae, Epeoromimidae, Hexagenitidae, and The generic nomen (formed from a combination of the Pristiplociidae did not survive to the Cenozoic, but Lepto- Latin words conus and virilis) is an allusion to the unusual phlebiidae, Oligoneuriidae and Siphlonuridae did. Among cone shaped penes. other families known from the Mesozoic (Jurassic but not yet Cretaceous) several did not continue to the Cenozoic, but 4. ConoviriZus poinari McCafferty n. sp. :description Coloburiscidae, Behningiidae, and Ephemerellidae survived Male adult to the present. Some other extant families not yet confirmed Body (Figs. 1,2) 4.3 mm long. Color patterns indetermi- from the Mesozoic record can be assumed to have been pre- nate, except where noted below. sent by the Lower Cretaceous based on their hypothesized phylogenetic positions relative to families known from the Compound eyes 0.6 mm diameter at base. fossil record. Antennae 0.7 mm long. In this report, I describe and analyze the first mayflies (one Forelegs incomplete ; femur 0.75 rnm long. Midlegs in- leptophlebiid and one baetid) taken from Lebanese amber. complete ; femur 0.75 mm long. Hindlegs with femur Lebanese amber dates from the Lower Cretaceous (120- 135 0.75 mm, tibiae 0.65 rnrn long, tarsi 0.38 mrn long. THE OLDEST MAYFLIES FROM AMBER Figs. 1-2. Conoviriluspoi~riMCCAPFERTY, n. gen., n. sp. Fig. 1 : Male adult (pbtographed in amber), dorsal view. Fig. 2 : Male adult (photographed in amber), ventral view. Figs 1-2. Conovirilus poi~riMCCAPFERTY, n. gen., n. sp. Fig. 1 : Adulte mgle (photographic5 dam l'ambre), wedorsale. Fig. 2 : Adulte mgle (photographit dans l'ambre), weventrale. W.P. McCAFFERTY exudate Figs. 3-7. Conoviriluspoinari McCm,n. gen., n. sp. Fig. 3 : Fmntal view of head (frons indicated). Fig. 4 : Hindleg tarsus. Fig. 5 : Tarsal claws (spread in amber). Fig. 6 : Outline of hindwing (dotted line pumed). Fig. 7 : Tenninalia, ventral view (obscuring white exudate
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