Cambridge Entomological Club, 1874
PSYCHE

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This is the CEC archive of Psyche through 2000. Psyche is now published by Hindawi Publishing.

Frank M. Carpenter.
Fossil Insects from Antarctica.
Psyche 76(4):418-425, 1969.

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FOSSIL INSECTS FROM ANTARCTICA*
BY FRANK M. CARPENTER
Harvard University
The only fossil insects from Antarctica that have been formally described are two beetles, Grahamelytron crofti Zeuner and Ade- mosynoides antarctica Zeuner, both from a Jurassic deposit on Mount Flora, Hope Bay, Grahamland, at the northern tip of the Antarctic peninsula (Zeuner, 1959). Since these are known by isolated elytra, their family positions are obscure and conjectural at best.
Two other fossil insects from Antarctica have been reported in the literature but not named. One of these, a well preserved wing of "Permo-Carboniferous" age, was found in the Theron Mountains, near the Filchner Ice Shelf, during the Trans-Antarctic Expedition of 195 5-58 ( Plumstead, 1962). Unfortunately, the specimen appears to have been lost in the mail after it was consigned to Dr. Zeuner for study (personal communication, Dr. James Schopf), the only record of it being the photograph published by Plumstead. Although overlain by plant fragments, the wing was apparently well preserved and its venation could have been worked out satisfactorily from 'the specimen. Even the small, published photograph is sufficient to show that the insect was homopterous, although venational details are not clear enough to permit determination of family affinitie~.~ Hornoptera of this general type are not uncommon in Permian de- posits in the Soviet Union, United States and Australia. The other specimen, a wing fragment of Permian age, was found in the Polar- star Formation of the Sentinel Mountains of Antarctica on the east slope of Polarstar Peak (Tasch and Riek, 1969). Despite the obscure nature of this fossil, Riek was led to conclude that it was a part of a homopterous fore wing, with a venation reminiscent of the family Stenoviciidae, known from the Permian and Triassic of eastern Australia and the Permian of Russia. My own, subsequent study of this specimen, made with the aid of ammonium chloride and under several different types of illumination, has revealed the presence of two additional longitudinal veins and numerous cross *This research has been supported in part by Grant No. GB 7308 from the National Science Foundation, F. M. Carpenter, Principal Investigator. 'Gressitt's suggestion (1967) that the fossil might be neuropterous is not really supportable.




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19691 Carpenter - Fossil insects 419
veins, as well as a coarse rugosity of the wing membrane, not men- tioned or shown by Riek. However, the fossil is still very fragmentary and although it might well be homopterous, its family position is most obscure.
Two additional fossil insects from Antarctica, one Jurassic and the other Permian, have been sent to me for study. Both are suffi- ciently well preserved to justify formal description and naming. The Jurassic specimen is an odonate, collected from a pond deposit within the so-called Mawson Tillite on Carapace Nunatak, South Victoria Land.2 It belongs to the suborder Anisozygoptera, which was a major one in the Jurassic Period, and to that complex of families which includes the Liassophlebiidae. The general vena- tional pattern, the nature of the arculus, nodus and pterostigrna, as well as the curvature of CUP and IA, are very similar to those of Limsophlebia. There are some differences in the nature of the antenodal cross veins but, all details considered, it seems advisable to broaden our concept of the family Liassophlebiidae to include the antenodal structure of the Antarctic species, for reasons given below, rather than to establish another family. The new specimen does clearly represent an undescribed genus and species. Caraphlebia Carpenter, new genus
This is related to Liasophleb'ia, but the hind wing has several weak antenodals in addition to the two strong, primary ones. The venation is much like that of Liasso'phlebia (see figures I and 3) but the cross veins between R2 and R3, proximally, are long and apparently not interrupted by transverse connections; the space be- tween MP and CuA is very narrow; IR2 apparently arises more distally than in Liassophlebia; and the anal area of the wing is small. The shapes of the discoidal cell, subdiscoidal cell, CUP and rA are very much as in Liassophlebia.
Type species: Caraphlebia antorctica, n. sp. Caraphlebia antarctica Carpenter, n. sp. Figure I
Hind wing: length of wing, 40 mm; width, at level of arculus, 8 mm. Primary antenodals very well developed, the costal and sub- costal elements aligned; the other antenodals weak and indistinct, but under glycerin-alcohol eleven are visible in the costal area and seven in the subcostal area, none aligned; pterostigma long and 'Basaltic lavas enclosing the fossiliferous pond deposit are presently being dated by the Potassium/Argon method.




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Psyche
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Figure 1. Caraphlebia antarct.'ca n.sp. Drawing based on holotype, collected on Carapace Nunatak, Antarctica. Sc, subcosta; Rl, radius; R2, R3, IR3, R4-+5, branches of radial sector; MA, anterior media ; CUP, pos- terior cubitus; dc, discoidal cell. Length of wing, 40 mm. Figure 2. Turanothemis nodalis Pritykina. Drawing based on holotype (hind wing), Jurassic of Karatau, USSR (from Pritykina, 1968). Figure 3. Liassophlebia mirabilis Tillyard. Drawing based on holo- type (hind wing), Jurassic of England (from Tillyard, 1925).



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19691 Carpenter - Fossil insects 42 1
slender; R4+ 5 arising slightly nearer to the arculus than to the nodus; other venational details are shown in figure I. Holotype : No. I 65874, U. S. National Museum, Smithsonian Institution, Washington; found in rock forming part of the "Maw- son Tillite" and collected on Carapace Nunatak, Antarctica, by H. W. Borns, Jr. and B. A. Halls3 The rock matrix is part of a volcanic mudflow and it includes many remains of Conchostraca, Ostracoda, and other Crustacea, as well as numerous body frag- ments of insects; the latter are unidentifiable even to ordinal level, though some appear to be parts of aquatic nymphs (possibly mayflies or stoneflies).
The plants of the Carapace Nunatak tillite are cycads and conifers and they are indicative of Jurassic age (Town- row, 1967). This is consistent with the occurrence of the family Liassophlebiidae, which is known only from Jurassic deposits. The specimen consists of a single wing, quite clearly but peculiarly preserved. The wing has been torn just beyond the nodus in such a way as to make it difficult to trace the subnodal vein, although its approximate position is obvious in the fossil. As a result of tearing along the posterior margin, the distal portions of the veins in the posterior half of the wing are not perfectly aligned with the basal portions, though the amount of shift is not uniform. Nevertheless, the use of large photographs has enabled the preparation of a drawing of the wing, shown in figure I ; this drawing includes only those strucrtures that are visible in the fossil, except for the very apex, which is indicated by dotted lines. One of the peculiar features of this specimen is the preservation of the veins on the two counterparts: apparently, the convex veins are well preserved on one half and the concave veins on the other. Such a
separation of the convex and concave veins can be duplicated in Recent insect wings by separating the two membranes just after the adult has developed its wings or by the use of caustic potash. Some wing veins in the fossil are only faintly indicated on the rock but they become very clear if the specimen is moistened with alcohol or glycerin-alcohol. The thin antenodals, for example, cannot be discerned unless the fossil is treated in this way. It is possible that the use of glycerin-alcohol on the specimens of Liassophlebia, which are in the British Museum, might also reveal the presence of faint antenodals in the costal area, since they do occur in the subcos,tal area (Tillyard, 1925). Even if this should prove not to be the case, the similarities between the wings of Liassophlebia and Cara- 'For the location of this Nunatak, see Borns and Hall, 1969, p. 871, fig. 1.



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422 Psyche [December
phlebiff are so marked that family separation of these genera seems unwarranted.
A related genus, Turanathemh (see figure 2)) has recently been described by Pritykina from the Jurassic of Karatau (Kazakhstan) in the Soviet Union and has been assigned to a separate family, Turanothemisridae (Pritykinz, 1968). In her account of this family, the author makes no comparisons with any other specific family, simply stating that it differs sharply from all other families of this series of Aniswygoptera by the presence of only 'the two primary antenodds in the costal area and by the f~m of the dk- coidal cell in the hind wings. However, in Lkopitlebia, as already noted, only the two primary antenodais have been reported {Tillyard, 1925) and its discoidal cell (hind wing) has precisely the same form as that of the fossil on which Tsiranothem'fs is based. Since no other distinguishing characteristics of the Turanothemistidae are dis- cernible, I consider the family Turanothemistidae inseparable from the Liassophlebiidae, which, on this basis, is known from Jurassic deposits of England, Siberia and Antarctica. The new Permian insect, found in conchostracan-bearing beds of the Mount Glossopten's Formation, Ohio Range, is a small but well preserved nymph. Since very little is known of nymphal forms of Paleozoic insects and especially since no venational pattern is discern- ible in the wing pads, the ordinal affinities of the fossil cannot be determined with any degree of certainty. However, the specimen is very close to a Permian nymph, Uralonympha Zaiessky, described from Tchekarda, in the Ural Mountains of the USSR, and similar to another, Permoleuctropsis Martynov, from a Permian deposit near Orenburg, USSR. The similarity of the new nymph to Urah- nympha is especially strong in the form of the prothorax and the position of the wing pads (see figure 6). The Antarctic species is accordingly being assigned to the genus Urabnympha, in preference to making another separate genus that could not be satisfactorily distinguished from Uralonympha at the present state of our knowl- edge.
The ordinal position of these nymphs is conjectural. Urala- nympha has generally been considered an immature form of a stone- fly (Zaiessky,
1935; Sharov, 1962) but there is an equally good possibility that it belongs to the Protorthoptera. Until a series of such nymphs has been found in association with numerous adults, the affinities of Urdonympha and other little-known nymphs of the Paleozoic will remain obscure. For the present, Ura/onY7nfha is best considered a member, incertae sedu, of the order Perlaria.



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Carpenter - Fossil insects
Figure 4. Uralonympha schopfi nsp. Drawing based on holotype, showing dorsal view of insect; collected at Leaia Led,ge, Ohio Range, Antarctica.
Figure 5. Uralonymfha schopfi n.sp. Drawing of holotype, showing ventral view.
Figure 6. Uralonymph var'ica Zalessky. Drawing based on holotype, Permian of Tchekarda, Ural Mountains, USSR (from Zalessky, 1939).
The following is a description of the nymph from the Ohio Range :
Uralonympha schopfi Carpenter, n. sp.
Figures 4 and 5
Based on a single specimen of a nymph, complete except for the last four or five abdominal segments. Length of specimen as pre- served, from the clypeus to the posterior edge of the sixth abdominal segment, 8 mm; antennae slender, 2.5 mm. long, showing 21 segments, though the basal few segments are not discernible; eyes prominent, bulging, width across the eyes 0.7 mm; clypeus prominent and mandi- bles large ; pronotum oval, 2.2 mm. wide, I .4 mm. long; mesothorax I .7 mm. long, metathorax I .3 mm. long. Fore wing pad about 3 mm. long; hind wing pad 2.5 mm. long.
The pronotum resembles that of Uralonympha in being distinctly oval. The legs are considerably more robust than those of Uralo- nympha. The fore legs are little known but the meso- and meta- thoracic legs are preserved as far as the femora; they show a distinct and rather large coxa, a small trochanter and well developed femur. The dimensions of the legs and their segments are as follows: mesocoxa, I. I mm. ; mesotrochanter, .3 mm. ; meso femur, I .7 mm. ;



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424 Psyche [December
metafemur, 2 mm.
The abdominal segments are of equal size, 1.5 mm. long. Since the terminal portion of the abdomen is not pre- served, the cerci are not included in the fossil. Holotype: No. 165875, U. S. National Museum; collected by Dr. James Schopf, for whom the species is named (field no. ANT 67-I-a&b) ;
it was found in a small piece of carbonaceous shale, bleached to white by weathering (Leaia Zone), west face, Mercer Ridge, Ohio Range. S~O~O'S, I 13O45'W; conchostracans and a typical Glossopteris flora occur in the same shale, which is considered to be late or middle Permian in age (Doumani and Tasch, 1965). The fossil is extraordinarily well preserved and if the specimen were an adult insect, the venational details would have allowed precise deter- mination of its systematic position.
Both of the new fossils described herein are indicative of the presence of productive insect-bearing deposits in Antarctica and the excellent preservation of these particular specimens justifies further exploration of the deposits concerned, with special reference to insect remains.
ACKNOWLEDGMENTS
I am indebted to Dr. J. M. Schopf of the USGS Coal Geology Laboratory for the opportunity of studying the specimen of Ura- Zonyrnpha; to Professors Borns and Hall of the University of Maine for making available the specimen of Caraphlebia ; to Professor Paul Tasch of the University of Wichita for certain collecting data; and to Mr. Jessa Merida of the U. S. National Museum for the loan of the specimen from the Polarstar Formation. REFERENCES
BORNS, H. W., JR. AND B. A. HALL
1969. Mawson "Tillite" in Antarctica: Preliminary Report of a Vol- canic Deposit of Jurassic Age. Science, 166: 870-872. DOUMANI, G. A. AND PAUL TASCH
1965. A leaiid conchostracan zone (Permian) in the Ohio Range, Holick Mountains, Antarctica. Amer. Geophys. Union Antarctic Res. Ser., 6: 229-239.
GRESSITT, J. LINSLEY
1967. Entomology of Antarctica. Antarctic Res. Series, 10 (No. 1574) : 25.
PLUMSTEAD, E. P.
1962. Fossil floras of Antarctica. Trans-Antarctic Exped., 1955-1958. Sci. Ref., 9: 66 and pi. 11, fig. 2.
PRITYKINA, L. N.
1968. Odonata. In: Jurassic insects from Kara-tau. Acad. Sci. USSR, 3 8 : 26-54. (Russian).




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19691 Carpenter - Fossil insects 425
SHAROV, A. G.
1962. Principles of Paleontology (Osnovy). Insecta, p. 138. (Russian). TASCH, P. AND EDGAR E. RIEK
1969. Permian insect wing from Antarctic Sentinel Mountains. Science, 166: 1529-1530.
TILLYARD, R. J.
1925. The British Liassic Dragonflies (Odonata). British Museum, Fossil Insects, 1: 1-38.
TOWNROW, J. A.
1967. Fossil Plants from Allan and Carapace Nunataks and from the Upper Mill and Shackleton Glaciers, Antarctica. New Zeal. Journ. Geol. Geophys., 10 : 456-73.
ZALESSKY, G.
1939. Etudes des insectes permiens du basin de la Sylva et problemes de l'kvolution dans la classe des insectes, Problems of Paleont., 5: 33-91.
ZEUNER, F.
1959. Jurassic beetles from Grahamland, Antarctica. Palaeontology, l(4) : 407-409.




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