Leptophlebiidae

Atalophlebia australis, copyright Keith Martin-Smith.

Belongs within: Furcatergalia.

The Leptophlebiidae are a group of mayflies in which the hind wings are relatively small or absent (Peters & Campbell 1991).

When mayflies last for millions of years
Published 10 March 2016
Mayfly Paraleptophlebia prisca preserved in amber, from Penney & Jepsen (2014).

Of all the media available for the preservation of fossils, none approaches perfection anywhere near as close as amber. There is little structural that amber does not preserve: external apperance, soft tissues, even cellular structure may potentially be examined. Amber offers us a window into the past unlike any other. The first amber deposits to go on the record (going back as far as the ancient Greeks), and the largest deposit yet discovered, was the Baltic amber of northern Europe, formed in the Eocene or Oligocene epoch* from the sap of a relative of the modern pines.

*Calculating the age of amber deposits is not easy. The amber itself cannot be dated directly (it is too old to be carbon-dated, of course, and it usually contains no mineral sediments that can be dated by other means) so aging it depends on indirect methods such as comparison of enclosed fossils with other aged samples, or dating of the deposits in which the amber is buried. Fossil comparisons suffer from the fact that the types of organisms that tend to be found preserved in amber are usually different from those preserved by other means (so we may know that two amber deposits are of similar age as each other, but we may still not know what that age actually is). Dating of surrounding deposits may be more straightforward, but is complicated by the fact that amber’s relative buoyancy makes it prone to reworking (when a geological specimen becomes eroded out of its original formation and reburied in a younger one, thus making it appear younger than it is). Of course, if an amber deposit was produced over a long period of time, it may be impossible to tell if a particular piece comes from early or late in that period. Current consensus seems to indicate an Eocene age for the Baltic amber, but older references may refer to it as Oligocene or even Miocene.

Most of the insects we find preserved in Baltic amber are similar to those we find today (though differences in climate between now and then mean that the amber contains a number of groups that we would not expect to find so far north today). The specimen shown at the top of this post is assigned to a genus of mayfly (of the diverse family Leptophlebiidae) that remains widespread in Europe and North America today, of which it represents the earliest record. Like other members of the genus, it is a relatively small mayfly with a wingspan of less than 15 millimetres (I came across this fly-fishing website referring to the frustration of anglers trying to handle such small flies). Several specimens of this species are known from the Baltic amber, representing the subimaginal and imaginal stages of both sexes (mayflies are unique among living insects in that their wings become functional before they are fully mature; when a mayfly first emerges from the water it is as a near-mature subimago, subsequently moulting to a fully mature imago). Presumably, like modern mayflies, Paraleptophlebia prisca emerged as synchronised swarms, many individuals of which may have found themselves landing on an unwisely chosen tree-trunk and trapped within weeping sap.

This species was first described in 1856 by F. J. Pictet-Baraban along with other species of ‘Neuroptera’ from Baltic amber (at the time, the order Neuroptera included a wide range of insects with relatively unspecialised wings such as mayflies, dragonflies, bark-lice and lacewings). Pictet assigned it to the genus Potamanthus, another Recent mayfly genus albeit one in a different family Potamanthidae. He did express some uncertainty about this placement, an uncertainty that was later borne out by Demoulin (1968) who re-identified it as Paraleptophlebia on the basis of wing venation and male genital characters. Subsequent authors have agreed with Demoulin’s assessment, regarding the mayflies of today as little different from those you might have found flying about over 50 million years ago.

Systematics of Leptophlebiidae

Characters (from Peters & Campbell 1991): Adult fore wings with vein MA forked, no marginal intercalaries present in interspace between veins MP2 and CuA; genital forceps of male with two short, terminal segments, segmentation often indistinct. Nymph with clypeus apparently fused to frons; maxillary and labial palps three-segmented, although segmentation often indistinct; dorsal and ventral portions of gills similar in shape and structure (ventral portion occasionally lost).

<==Leptophlebiidae [Leptophlebioidea]
|--LeptophlebiinaeGS15
| |--LeptophlebiaGS15
| | |--L. marginata (Linnaeus 1767)GS15
| | `--L. vespertina (Linnaeus 1758)M86
| `--Paraleptophlebia Lestage 1917GS15, D70 [incl. Oligophlebia Demoulin 1965GS15]
| |--P. prisca (Pictet-Baraban & Hagen 1856)D70, GS15 (see below for synonymy)
| |--P. spinosa Ueno 1931TYM08
| `--P. submarginata (Stephens 1835)GS15
`--AtalophleboadentataGS15
|--Calliarcys Eaton 1881 [Calliarcyinae]GS15
| |--*C. humilis Eaton 1881GS15
| `--C. van Godunko, Sroka et al. 2015GS15
`--+--HabrophlebiinaeGS15
| |--Habrophlebia lauta Eaton 1884GS15
| `--Habroleptoides confusa Sartori & Jacob 1986GS15
`--+--Terpides [Terpidinae]GS15
`--AtalophlebomaxillataGS15
|--Castanophlebia [Castanophlebiinae]GS15
`--AtalophlebiinaeGS15
|--AustrophlebioidesPC91
|--JappaPC91
|--UlmerophlebiaPC91
|--Kirrara proceraPC91
|--Nousia (Australonousia) fusculaPC91
|--Koorrnonga inconspicuaPC91
|--Garinjuga maryannaePC91
|--Neboissophlebia occidentalisPC91, ASD05
|--Nyungara bunniPC91, ASD05
|--Bibulmena kadjinaPC91, ASD05
|--AtalomicriaPC91
|--ThraulophlebiaPC91
|--Thraulus Eaton 1881GS15
| |--T. bellus Eaton 1881GS15
| `--T. thraker Jacob 1988GS15
`--AtalophlebiaPC91
|--A. australasicaPC91
|--A. australisPC91
`--A. yuganaPC91

Leptophlebiidae incertae sedis:
ConovirillusKS02
Lepismophlebia platymeraS02
Leptoneta Sinitshenkova 1989 non Simon 1872KS02
MassartellaMD03
Deleatidium myzobranchiaCQ04, PC66
Acanthophlebia cruentataCQ04
ZephlebiaPC66
|--Z. cruentataPC66
|--Z. dentataPC66
`--Z. versicolorPC66
Atalophlebioides aucklandensis Peters 1971PC91, H79
Choroterpes Eaton 1881D68
Blasturophlebia Demoulin 1968D68
`--*B. hirsuta Demoulin 1968D68
Acentrella Bengtsson 1912GS15
`--A. joosti [=Baetis joosti]SBG11
AtalonellaR74
MasharikellaR74
CareospinaP92
BorinquenaP92

Paraleptophlebia prisca (Pictet-Baraban & Hagen 1856)D70, GS15 [=Potamanthus priscusD70, Leptophlebia priscaD68; incl. Oligophlebia calliarcys Demoulin 1965D70, O. longiceps Demoulin 1965D70]

*Type species of generic name indicated

References

[ASD05] Armstrong, K. N., A. W. Storey & P. M. Davies. 2005. Effects of catchment clearing and sedimentation on macroinvertebrate communities of cobble habitat in freshwater streams of southwestern Australia. Journal of the Royal Society of Western Australia 88 (1): 1–11.

[CQ04] Collier, K. J., & J. M. Quinn. 2004. Factors affecting distribution and abundance of the mayfly Acanthophlebia cruentata (Leptophlebiidae) in North Island, New Zealand, streams. New Zealand Entomologist 27: 17–28.

[D68] Demoulin, G. 1968. Deuxième contribution à la connaissance des ephéméroptères de l’ambre oligocène de la Baltique. Deutsche Entomologische Zeitschrift (N. F.) 15 (1–3): 233–276.

[D70] Demoulin, G. 1970. Troisieme contribution a la connaisance des ephemeropteres d l’ambre Oligocene de la Baltique. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 46 (2): 1–11.

[GS15] Godunko, R. J., P. Sroka, T. Soldán & J. Bojková. 2015. The higher phylogeny of Leptophlebiidae (Insecta: Ephemeroptera), with description of a new species of Calliarcys Eaton, 1881. Arthropod Systematics and Phylogeny 73 (2): 259–279.

[H79] Hubbard, M. D. 1979. On the origin of flightlessness in Ephemeroptera. Systematic Zoology 28 (2): 227.

[KS02] Kluge, N. Yu., & N. D. Sinitshenkova. 2002. Order Ephemerida Latreille, 1810. The true mayflies (=Ephemeroptera Hyatt et Arms, 1891 (s. l.); =Euephemeroptera Kluge, 2000. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 89–97. Kluwer Academic Publishers: Dordrecht.

[M86] Marten, M. 1986. Drei für Deutschland neue und weitere, selten gefundene Eintagsfliegen aus der Fulda (Insecta, Ephemeroptera). Spixiana 9 (2): 169–173.

[MD03] Molineri, C., & E. Domínguez. 2003. Nymph and egg of Melanemerella brasiliana (Ephemeroptera: Ephemerelloidea: Melanemerellidae), with comments on its systematic position and the higher classification of Ephemerelloidea. Journal of the North American Benthological Society 22 (2): 263–275.

[PC66] Pendergrast, J. G., & D. R. Cowley. 1966. An Introduction to New Zealand Freshwater Insects. Collins: Auckland.

Penney, D., & J. E. Jepson. 2014. Fossil Insects: An introduction to palaeoentomology. Siri Scientific Press: Manchester.

[PC91] Peters, W. L., & I. C. Campbell. 1991. Ephemeroptera (mayflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 1 pp. 279–293. Melbourne University Press: Carlton (Victoria).

Pictet-Baraban, F. J., & H. Hagen. 1856. Die im Bernstein befindlichen Neuropteren der Vorwelt. In: Berendt, G. C. Die im Bernstein befindlichen organischen Reste der Vorwelt vol. 2 pp. 41–126. Nicolaischen Buchhandlung: Berlin.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[R74] Riek, E. F. 1974. Ephemeroptera (mayflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 33–34.Melbourne University Press.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

[SBG11] Staniczek, A. H., G. Bechly & R. J. Godunko. 2011. Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera). Insect Systematics and Evolution 42: 101–138.

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

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