A. P. Rasnitsyn.
Review of the fossil Tiphiidae, with description of a new species (Hymenoptera).
Psyche 93(1-2):91-101, 1986.

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REVIEW OF THE FOSSIL TIPHIIDAE,
WITH DESCRIPTION OF A
NEW SPECIES (HYMENOPTERA)*
Paleontological Institute,
Academy of Sciences of the USSR,
Profusoyuznaja, 113, 117868,
Moscow USSR
Through the courtesy of Professor Frank M. Carpenter (Harvard University, Cambridge, Mass.) and Dr. Paul E. S. Whalley (British Museum, Natural History, London, U.K.) I have been able to study the type specimens (good photographs of the specimen in one case) of all described extinct species ever attributed to the Tiphiidae. Five of them have been described as members of the subfamily Antho- boscinae by Cockerell: in 1906 (Lithotiphia scudderi, Geotiphiafox- iana), 1910 (G. sternbergi, G. halictina) and 1927 (G. pachysoma); while Hoplisidea kohliana was described originally as a member of the Sphecidae (Cockerell, 1906) and later transferred to the Antho- boscinae by Evans (1966).
From my study of these specimens I have found that the latter species most probably belongs to the Sceliphronini (Sphecidae) and I will treat it elsewhere. The five other species are discussed below and one new species is described. All the species described by Cockerel1 are from the Lower Oligocene of Florissant, Colorado; the new one is from the ?Upper Oligocene of the Sikhote-Ah Mts., Maritime Province of the USSR. Only the holotypes are known for all these species and each specimen is a female, suggesting a female biased tiphiid population during the Oligocene. Only two other fossil specimens of Tiphiidae have been men- tioned in the literature; both were found in Baltic amber collected by A. Menge and both were identified by Brische (1886) as "Tiphia (?)". Unfortunately, Menge's collection is apparently lost (Heie, 1967, p. 119).
*Manuscript received by the editor August 3, 1985 91




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92 Psyche [VOI. 93
The species treated here (figures 1-7) can be assigned to the Tiphiidae on the basis of the strongly fossorial nature of the legs (mid and hind tibiae thick and spiny), combined with the pleisio- morphic wing venation; the latter differs distinctly from that of the Scoliidae, which do have similar fossorial adaptations. In one case (Fig. 2) this indirect evidence is confirmed by the structure of the mesosternum, which shows the pair of lamellae that characteristi- cally partly cover the midcoxae.
The fossil species show a habitus and female wing venation typi- cal for the Anthoboscinae. Nevertheless, they do not belong to that subfamily, mainly because their antennal sockets are overlain with tubercles, clearly seen in one case (Fig. 6) and less clear in another (Fig. 7). There are additional features distinguishing the fossils from Anthoboscinae, viz., flagellum straight or variously bent (Figs. 1,2, 4, 6) instead of tightly curled (as in all female Anthoboscinae stu- died), femora lacking genual plates (Figs. 1-3, 7) or propodeum with longitudinal lines (Figs. 4, 5).
All Tiphiidae with the antennal sockets partly covered by frontal tubercles or ridges belong to the Myzininae and Methochinae. The latter subfamily is not involved here, since its members have thin tibiae bearing only weak spines. [I follow V. Gorbatovsky (personal communication) in treating Pterombrus Smith as a member of the subfamily Methochinae]. Therefore, the Myzininae is the only sub- family with the characters of the fossils and in particular with those of Geotiphia. [Lithotiphia is poorly known but I consider it similar enough to the former genus to classify them together and not to reject Lithotiphia as a tiphiid incertae sedis]. Within the Myzininae the fossils take an isolated position because of the very primitive, male-like wing venation of the females.
Both of these extinct genera can be identified by the following diagnoses. Lithotiphia (Fig. I): forewing with cu-a cross-vein ante- furcal; head capsule with a short oral cavity, distant from occipital carina; hind tibiae very strongly swollen. Geotiphia (Figs. 2-7): fore wing with cu-a interstitial or postfurcal; oral cavity longer, with hypostomae reaching occipital carina; hind tibiae less swollen. The latter genus possibly deserves to be divided into two genera, since sternbergi and pachysoma, in contrast to other species, show mid and/ or hind femora with the genual plates, and the propodeum with longitudinal lines. The propodeal structure is unknown in any other



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Rasnitsyn- Fossil Tiphiidae
Figure 1.
Lithotiphia scudderi Cockerell, holotype, no. 2022, Museum of Com- parative Zoology, Harvard University. Wing cells are lettered. Scale line in all fig- ures, 3 mm.
species and I hesitate to create another new genus on a sole charac- ter. The following is a descriptive account of the species in these two genera. The details shown in the figures are generally not described below.
Lithotiphia scudderi Cockerel1
Figure 1
Lithotiphia scudderi Cockerell, 1906, p. 51 Body length, 12.3 mm; fore wing length, about 5 mm (Length is measured here from base to apex of cell 3r). Gastral terga with light spots. Integumental sculpture not discernible because of covering by Canada balsam. Holotype: M.C.Z. no. 2022.



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[Vol. 93
Figure 2.
Geotiphia foxiana Cockerell, holotype, no. 2021, Museum of Compar- ative Zoology, Harvard University.
Geotiphia foxiana Cockerel1
Figure 2
Geotiphia foxiana Cockerell, 1906, p. 52 Body length, as preserved, 11 mm (probably originally 12 mm.); fore wing length, 6.2 mm. Integumental sculpture not discernible. Ground color moderately dark, the flagellum, tibiae, tarsi, veins, and pterostigma less dark; metasomal sterna with light spots sub- laterally, 2nd sternum having the spots large and contiguous. Color pattern of terga unknown. Wing membrane not infumate. Holo- type: M.C.Z. no. 2021.
Geotiphia halictina Cockerel1
Figure 3
Geotiphia halictina Cockerell, 1910, p. 279 Body length, 18 mm; fore wing length, 3.5 mm. Venation similar to that of foxiana, but differing in smaller size and the position of cell



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Rasnitsyn- Fossil Tiphiidae
Figure 3.
Geotiphia halictina Cockerell, drawing based on photograph of holo- type, no. 18619, Museum of the University of Colorado. 3r remote from wing margin apically. Integumental sculpture and color pattern unknown. (Description based on photograph of holotype).
Geotiphia sternbergi Cockerel1
Figure 4
Geotiphia sternbergi Cockerell, 19 10, p. 277 Body length, 8 mm; fore wing length, 12 mm. Head with posterior surface punctate dorsally and laterally, finely punctatorugose medially. Thorax with distinct, moderately large punctures dor- sally; lateral adscutellar depression, metanotum and propodeurn finely reticulate. Gastral terga with sculpture fine and sparse, not clear in detail. Ground color dark (not known for fore and mid



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Figure 4. Geot@hia sternbergi Cockerelly holotype, no. 18868, American Museum of Natural Historyy New York, Wing veins are lettered. legs), anterior metasomal segments with small light spots laterally. Fore wing apex infumate. Differs from the above species by its large size, modified antenna1 segments, and in having the hind femur with genual plate; fore wing with cell 2rm very long, and possibly in having the propodeum with longitudinal lines. Holotype: A.M.N.H., no. 18868.
Geotiphia pachysoma Cockerel1
Figures 5 and 6
Geotbhia pachysoma Cockerelly 1927å p. 432. Body length, 9.2 mm; fore wing length, 6.0 mm. Head punctato- rugose dorsomedially in part, thorax smooth, with distinct but weak



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Rasnitsyn- Fossil Tiphiidae
Figure 5.
Geotiphia pachysoma Cockerell, holotype, no. In. 26929, British Museum (N.H.), London. Dorsal view.
punctures dorsally; lateral parts of metanotum striate longitudi- nally. Body with ground color dark, without obvious light spots; wing membrane infumate in apical two-fifths. Similar to sternbergi in having genual plates and dissected propodeum, differing in small size and in having cell 2 rm shorter; genual plates longer. Holotype: B.M. (N.H.), no. In 26929.
Geotiphia orientalis, new species
Figure 7
Fore wing length about 6 mm. Pterostigma rather long, with 2r-rs arising halfway before apex; cell 3r rounded at costal margin; RS between RS-kM and 2r-rs almost straight; cells lr, 2rm and 3rm all relatively short; 2rm and 3rm of subequal length; lm-cu just before the middle of 2rm; 2m-cu at the middle of 3rm, which has the



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Figure 6. Same as Fig. 5, ventral view.
posterior side very short and the distal side (3r-m) strongly arched; crossvein cu-a at the fork of M-kCu; posterior genual plates absent on mid and hind femora. Surface sculpturing indistinct. Body struc- ture as preserved lacks taxonomically important features, the details in part difficult to interpret. Ground color moderately dark; tibiae, tarsi, venation, pterostigma, and metasomal segments 2 and 3 less dark and without light spots (subsequent segments not preserved). Wing membrane not infumate.
Holotype (only specimen known): no. 34291 100, Paleontological Institute, Moscow, USSR; collected at Bolshya Svetlovodnaya River, Pozharsky District, Maritime Province, USSR: ?Upper Oligocene.




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Figure 7.
GeotMhia orientalis, holotype, no. 34291 100. Paleontol. Inst. Acad. Sciences, USSR, Moscow.
Comparison. As preserved this species is very similar to foxi- ana, differing in having a longer pterostigma, the posterior side of cell 3rm shorter, and the metasomal segments without light spots [The latter difference may be meaningless because the color pattern is known only for the metasonal sterna in foxiana and possibly only for terga in orientalis].
The above data show considerable taxonomic and anagenetic evolution of the subfamily Myzininae since the early Oligocene, an interval of about 35 million years. Both fossil genera have been replaced with a wide array of living genera, and even the most primitive modern genus, Myzinum Latreille, is probably further away from its Oligocene predecessors than these predecessors are from their anthoboscine ancestor. A paleontological history is not



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100 Psyche [VOI. 93
known for any living myzinine genera, probably because of their preference for environments unfavorable to fossilization (xeric bio- topes or, in the case of Hylornesa, tropical forests), but all of them can be easily derived from Geotiphia morphologically (but not from Lithotiphia, because of the apomorphic position of the cu-a cross- vein). Geotiphia can be characterized in short as an anthoboscine with supraantennal tubercles, a position not consistent with the current phylogenetic scheme showing synapomorphies for a11 Tiphiidae other than Anthoboscinae and additional synapomor- phies for all Tiphiidae except Anthotoscinae and Thynninae (Brothers, 1975). An alternative scheme with Myzininae independ- ent of other subfamilies (excluding Anthoboscinae and probably Metochinae) seems to me more realistic.
The paleontological records indicate the minimal age of the Myzininae as Early Oligocene. The records seem too scanty, how- ever, to help in identifying the geographic area where the subfamily arose.
Types of the previously described fossil Tiphiidae are studied. Two genera and six species are recognized, each species known only from the holotype: Lithotiphia Cockerell, with only one species, scudderi Cockerell; and Geotiphia Cockerell, with foxiana Cocke- re11 (type-species), halictina Cockerell, orientalis, n.sp., sternbergi Cockerell, and pachysoma Cockerell. The fossils are found to represent the most primitive members of the subfamily Myzininae, indicating that the subfamily originated from the Anthoboscinae independently of the Thynninae, Tiphiinae, and Brachycistidinae. Hoplisidea kohliana Cockerel1 is now determined as belonging to the Sceliphronini of the family Sphecidae and will be treated else- where. All species mentioned are from the Lower Oligocene of Flor- issant, Colorado, except the new one, G. orientalis, which is from the ?Upper Oligocene of Sikhote- Alin Mts., Maritime Province of USSR.




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Rasnitsyn- Fossil Tiphiidae
BRISCHKE, D.
1886. Die Hymenopteren des Bernsteins. Schrift. naturf. Gesellsch. Danzig, n.f., 6: 278-279.
BROTHERS, D. J.
1975.
Phylogeny and classification of the Aculeate Hymenoptera, with special reference to Mutillidae. Univ. Kansas Sci. Bull., 50(11): 483-648. COCKERELL, T. D. A.
1906. Fossil Hymenoptera from Florissant, Colorado. Bull. Mus. Comp. Zoology, Harvard Univ., 50: 33-58.
1910.
Fossil insects and a crustacean from Florissant, Colorado. Bull. Amer. Mus. Nat. Hist., 28: 275-288.
1927.
Hymenoptera and a caddis larva from the Miocene of Colorado. Ann. Mag. Nat. Hist., (9)20: 429-435.
EVANS, H. E.
1966. The comparative ethology and evolution of the sand wasps. Harvard Univ. Press, Cambridge, Mass., p. 1-526. HEIE, 0.
1967.
Studies on fossil aphids (Homoptera, Aphidodea). Spol. Zool. Mus. Hauniensis, 26: 1-274.




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