F. M. Carpenter.
The Affinities of Eomerope and Dinopanorpa (Mecoptera).
Psyche 79:79-87, 1972.

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THE AFFINITIES OF EOMEROPE AND
DINOPANORPA (MECOPTERA)*
BY F. M. CARPENTER
Harvard University
The two fossil Mecoptera discussed below were originally de- scribed by T. D. A. Cockerell many years ago. One of them,
Eomerope tortriciformis, was obtained in the Oligocene shales at Florissant, Colorado; and the other, Dinopanorpa megarche, was collected in a Miocene deposit near the Amagu River in eastern Siberia. Study of the type specimens for the present paper was made possible by the courtesy of the authorities of the Peabody Mu- ~eum at Yale University, for Eomerope, and of the U. S. National Museum, for Dinopanorpa.
Eomerope was assigned by Cockerel1 (1909) to the mecopterous family Meropeidae, which, at that time, was a monotypic family, represented by Merope tuber Newman, a little-known species infrequently collected in the eastern part of the United States. However, a second species, hstromerope poultoni from Western Australia, was described by Killington in 1933. These two genera, although having obvious differences in facies, are closely related, as indicated by the similar structure of the male genitalia. In his account of Eomerope, Cockerell made no reference to another mecopteron, Notiothauma reed MacLachlan (1877), which occurs in part of Chile and which is the only known representative of the family Notiothaumidae. In all probability, Cockerel1 was not aware of this insect, since its existence was not generally made known until the publication of Esben-Petersen's monograph of the Mecoptera in 1921. From my study of the type of Eomerope and comparisons with specimens of Mer~ope and Notiothauma, I am convinced that Eomerope belongs to the family Notiothaumidae instead of the Meropeidae. The reasons for this conclusion are given below, follow- ing the account of the genus and species. Family Notiothaumidae Esben-Petersen
Genus Elomerope Cockerel1
Eornerope Cockerel!, 1909, p. 381
Wing venation as in Notiothauma but with fewer cross veins, the *Partial financial support of this research is acknowledged to the National Science Foundation (Grant no. GB 27333, F. M. Carpenter, Harvard Uni- versity, principal investigator).
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Psyche [March-June




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19721
Carpenter - Eonzerope and Dinopanorpa
number of cellules being about half that in Notioth,auma. Type species: Eomerope tortricifonnis Cockerell. E omerope tortricif ornzis Cockerell
Figs. I and 5
Eomerope tortriciformis Cockerell, 1909, p. 381 Length of fore wing, 14 mm. ; length of body, 13 mm. The wing expanse of the insect was about 32 mm., some 10 mm. less than N. reedi.
Type: No. 26176, Peabody Museum, Yale University; collected at Florissant, Colorado, in 1907.
The specimen shows the whole insect (see figure 5). The wings
are almost symmetrically arranged, with a pair on each side slightly overlapped; the veins in the apical and posterior regions of both pairs are not discernible. The legs are long and unusually spinose, as in Notiothauma. The specimen, obviously a male, has the 10th abdominal segment forming a characteristic genital bulb, comparable to that in Notiothauma (see Crampton, I 93 I ) . The preserved part of the venation of the fore wing is shown in figure I. At the base of the wing is a cluster of heavy setae, as in Notiothauma. The costal area is abruptly narrowed basally. Sc is a distinct vein, as in Notiothamza, with a series of irregular veinlets arising anteriorly from its basal branch. RI arises from Rs, as in Notiothauma, by diverging anteriorly, Rs continuing the straight line of R; only a few of the basal branches of Rs are preserved; IA and 2A are represented only by their curved basal portions that strongly resemble the curved bases of Notiothauma. The venation of the anterior-basal part of the hind wing is like that of the fore wing.
The similiarity of the venation of Eomerope to that of Notio- thaw is at once obvious by comparing figure I with figure 2, which shows the basal part of the wing of Notiothauma. The venational pattern is essentially the same, the only notable difference being the smaller number of cross veins and cellules in Eomerope. The differ- Figure 1. Eomerope tortriciformis Cockerel1 ; drawing of preserved part of fore wing, based on holotype
(original).
Figure 2. Notiothauma recdi MacLachlan; drawing of proximal part of fore wing (after Crampton, 1930).
Figure 3. Meropc tuber Newman; drawing of proximal part of fore wing (original).
Figure 4.
Austromcropc poultoni Killington; drawing of proximal part of fore wing (after Killington, 1933).




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82 Psyche [March-June
Figure 5. Eomerope fortriciformil Cockerel!; photograph of holotype; length of fore wing, 14 mm.




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19721 Carpenter - Eomerope and Dinopanorpa 83 ences between Eomerope and the two genera of Meropeidae are readily apparent by considering figures 3 (Merope) and 4 (Austro- merope). In Merope the costal area is relatively narrow and lacks cellules; Rs diverges posteriorly from RI. In Austromerope the costal area is also narrow, though there are a few cellules; RI, not Rs, continues the straight line of R, Rs diverging posteriorly from RI ; and the anal veins do not have distinctly curved bases. From the evidence of the wing venation, it is clear that Eomerope is more closely related to Notiotkaunza than to the Meropeidae, but stronger evidence is furnished by the structure of the abdomen in the male. In the Meropeidae the males lack the genital bulb charac- teristic of most Mecoptera but possess a pair of long claspers, which extend far beyond the end of the abdomen, especially in Austromerope. In Eomerope, the abdomen has the small bulb like that of Notio- thaum, with very short forceps that do not extend beyond the abdomen.
Family Din~~anorpidae, new family
Large Mecoptera, related to the Orthophlebiidae and Pano'rpidae. Hind wing : several strong cross veins between Sc and costal margin ; RI extending almost to wing apex, curving posteriorly near its termi- nation; Rs with at least 8 terminal branches, all directed posteriorly near the wing margin; M with at least 5 branches; stem of Cu free at base; 'CuA coalesced with M for a short distance basally and CUP coalesced with IA; first basal cross vein (m-cu) between CuA and MP very long and oblique; cross veins more numerous than in Panorpidae and Orthophlebiidae. Fore wing and body unknown.
Genus Dinopanorpa Cockerel1
Dinopanorpa Cockerel!, 1924, p. 2
Hind wing: costal space relatively broad (for a hind wing), with 5 strong veinlets to margin; stem of Cu somewhat closer to IA than to M.
Type species : Dinopanorpa megarche Cockerell. Dinopanorpa megarche Cockerel1
Figs. 6 and 9
Dinopanorpa megarche Cockerell, 1924, p. 2 Length of hind wing, 30 mm.; maximum width, 10 mm.; esti- mated wing expanse, 65 mm. Other specific characteristics are diffi- cult to designate, in the absence of the fore wing and body; however, the number and arrangement of cross veins would almost certainly



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Figure 6. Dbzopanor$a megarche Cockerell; drawing of hind wing, based on holotype (original).
Figure 7. Panorpa nebulosa Westwood; drawing of hind wing (ori- (ginal).
Figure 8. Ort/zo$hlebia liassica Mantell; drawing of hind wing (after Tillyard, 1933).




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86 Psyche [March-June
be in this category. Details of the venation are shown in figure 6. As preserved, the wing is dark brown) with several white spots and bands (figure 9)) as frequently seen in existing panorpids. Type: No. 69173, U. S. National hfuseum; collected by A. Kuznetzov, '(on the Amagu River) Maritime Province, coast of Siberia, opposite the southern end of Sakhalin Island." Rohdendorf (1957) records the 1cGcality as in the Lower Arnul-sk Region of the USSR, on the bank of the Kudya River, a tributary of the Amagu River, and indicates its age as Lower Miocene, Cockerel1 states ( 1924) that the flora of the deposit) including such genera as Ginko, Cornus, Taxodium, and Qum-cus, indicates a warm-temperate climate. The type specimen consists of a very well-preserved hind wing, lacking only a part of the apical-posterior region (figure 9). Cockerell placed Dinopanorpa in the family Panorpidae, Tillyard ( 1933, p, 26) transferred it to the extinct family Orthophlebiidae (otherwise known only from the Triassic and Jurassic periods) and Martynova (1962, p* 291) considered it a synonym of Orthophlebia in the same family. Actually, as noted by Cockerell, Dinopanorpa presents) in the hind wing, a remarkable combination of characters. The presence of 5 strong veinlets between Sc and the costal margin is a feature that does not occur in the Orthophlebiidae or Panorpidae, although it is seen in some of the Permian and Triassic genera of other families. The form of RI, extending nearly to the wing apex and directed poste~iorly in the apical region, is unique in the known Mecoptera, extinct and Recent) as noted by Cockerell; in other members of the order, RI is much shorter and is curved anteriorly at its termination. Cross veins are at least twice as numerous in Dinlopanorpa as in the Panorpidae and Qrthophlebiidae. Another peculiar feature) also noted by Cockerell, is the long and oblique m-cu cl-oss vein (figure 61, although it could be an abnormality in this particulai- wing. In contrast, the structure of Cu, including its stem, the nature of its branching and the coalescence of CuA and CUP (with M and IA respectively), is virtually identical with that in the Panorpidae (figure 7) but, incidentally, quite unlike that of Orthophlebiu (figu-e $)* The extensive branching of Rs, with at least 8 terminal branches) is totally unlike the condition in the Panorpidae, with 5 branches ts Rs. Dinopanorpa also has a 5- branched M, although that vein is rarely more than 4-branched in Panorpidae.
In view of the differexes and peculiarities noted above, assignment of Dinopanorpa to either Orthophlebiidae or Panorpidae seems un-



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19721 Carpenter - Eomerope and Dinopanorpa 87 justified. The position of the Dinopanorpidae in the mecopteran phylogeny will remain obscure until the fore wing and body struc- tures are known. However, it can ha-~dly be cons!dered intermediate between the Panorpidae and Orthophlebiidae because of the peculiar form of RI and Rs and the presence of costal veidets. More likely, it is a specialized derivative of some early AJesozoic st~ck) close to the Orthophlebiidae.
REFERENCES
COCKERELL, T. D. A,
1909. Descriptions of Tertiary Insects) VI. Am. Journ. of Sci (4) 27: 381-387.
1924. Foss~il Insects in the United States National Museum. Proc. U.S.N.M. 64: 1-15.
CRAMPTON, G. C.
1930. The Wings of the Remarkable Archaic Mecoptera Notiothauma reedi MacLachlan with Remarks on their Protoblattoid Affinities. Psyche 37: 83-103.
1931. The Genitalia and Terminal Structures of the Archaic Mecop- teron Notiothauma reedi; Compared with Related Holometoba from the Standpoint of Phylogeny. Psyche 38: 1-21. KILLINGTON, F. J.
1933. A New Genus and Species of Meropeidae (Mecoptera) from Australia. Ent. Mo. Mag. 69: 1-4.
MARTYNOVA) 0.
1962. Mecoptera, in Osnovy (Insects), pp. 283-294, ROHDENDORF, B. B,
1957. Paleoentomological Investigation in the USSR. Trudy Paleont. Inst. 66: 1-100.
TILLYARD, R. J.
1933. The Panorpoid Complex in the British Rhaeic and Lias. Brit. Mus. Fossil Insects, No. 3: 1-79.




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NESTING BEHAVIOR AND DESCRIPTION OF THE
LARVA OF BO THYATOSTETHUS DISTINCTUS FOX (HYMENOPTERA: SPHECIDAE)
Bothynostethus is a predominantly neotropical genus having a single described species, distinctus Fox, in America north of Mexico. The genus has been little studied, and its placement in the sphecid classifica,tion has been the subject of some dispute. Kohl (1896) and Handlirsch ( 1925) put Bothj~n~ostethus in the Nyssoninae and related it to Alysson. In the Synoptic Catalog of Hymenoptera of America North of Mexico, ICmmbein (1951) placed it in the tribe Misco- phini of the subfamily Larrinae. More recently, Menke (1968) grouped B~othynostethus, Scaflheutes, WiZZinkieZZa, and Bohartella in the tribe Bothynostethini of the subfamily Larrinae. However) Evans (1964) has questioned, on the basis of larval structure, whether the Larrinae and Crabroninae deserve the status of separate sub- families, and he has cited Bothynostethus as a genus in which the adults resemble certain Crabroninae. Despite Menke's ( I 968) in- clusion of the Bothynostethini in the Larrinae, he informs us (correspondence) that Bothynostethus is not a "typical" member of the subfamily, and that "it would not be difficult to derive a crabronid from a Bothynostethus-like ancestor". It is clearly of interest to discover what light a study of nesting behavior and larval structure may shed on these problems. Cazier and Mortenson (1965) found B. distinctus nesting in the ground and preying upon a chrysomelid beetle, but othei-wise the genus has not been studied in the field. Recently we discovered a small aggrega- tion of B. distinctus nesting near Albany, N. Y., and a single indi- vidual nesting in Bedford, Mass., and we are able to present considerable new information on nesting behavior as well as a description of the mature larva. Our studies represent the first records of the species from those two states, as it has not previously been reported from north of New Jersey. In the Discussion we reconsider the proper phylogenetic placement of Bothynostethus. The specimens from our studies bear our note numbers and have 'Department of Forest Entomology, State University College of Forestry, Syracuse, New York 13210.
2Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138.
Manu~cript receiaed by the editor July 5, 1972.



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