F. M. Carpenter.
The Lower Permian Insects of Kansas. Part 3. The Protohymenoptera.
Psyche 37:343-374, 1930.

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19301 Permian Insects of Kansas
THE LOWER PERMIAN INSECTS OF KANSAS
PART 3. THE PROTOHYMENOPTERA
Museum of Comparative Zoology, Cambridge, Mass. The order Protohymenoptera was established by Tillyard (1924) to include a series of fossil wings contained in the Yale collection from the Lower Permian of Kansas. As indicated by the name, Tillyard believed that this group was ancestral to the Hymenoptera and was sufficient to prove that the Hymenoptera originated independently from the rest of the holometabolous insects (1926b). While it is true that certain morphological features of the Hymen- optera, such as the reduced wing venation, and the poly- nephric Malpighian system, remove the group from the true panorpoid orders ; the results of morphological studies (Crampton, 1919, 1927) indicate that the Hymenoptera arose from the same stem as all the other insects with a similar metamorph~sis.~ It is therefore advisable for us to submit any paleontological evidence which seems inconsistent with this view to the carefullest examination. Aside from phylogenetic consideratons, the question of the affinities of the Protohymenoptera has close bearing on the interpretation of the wing venation in the Hymen- optera; for the specialized condition of the wings in the Hymenoptera has prevented entomologists from satisfac- torily homologizing the veins with those of other insects. As a result, a number of different systems of venational * National Research Fellow, Harvard University. These studies have also been aided by grant No. 280 of the Bache Fund, National Academy of Sciences, and a Sheldon Traveling Fellowship from Harvard University. Part 1 (Introduction and the Order Mecoptera) was published in the Bulletin of the Museum of Comparative Zoology 70 (2), 1930; Part 2 (Paleodictyoptera, Protodonata, and Odonata) will appear in the American Journal of Science, 21 (2), 1931.
Handlirsch, however, has held the opposite view (1906-08).



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344 Psyche [December
nomenclature are being used in the Hymenoptera, each more or less limited to a certain group of families. Com- stock, MacGillivary, and a few others have proposed sys- tems intended to homologize the venation with that of other groups, but hymenopterists in general have not accepted these schemes. More recently Tillyard has ad- vanced a new and radically different system (1924) based on the assumption that the Protohymenoptera exhibit the primitive condition of the venation in the hymenopterous line of descent; and he believes that through this interpre- tation the whole of the homologies of the veins at once "becomes clear and of the utmost simplicity" (1926~~ p. 256).
Let us consider briefly the opinions expressed by other entomologists on the affinities of the Protohymenoptera, and on the new system of venation in the Hymenoptera, based on these fossils. Lameere (1927) accepts the insects as representatives of the group ancestral to the Hymen- optera, and also adopts the venational interpretation, with a few slight modifications. Handlirsch (l927), however, has briefly (without discussion) suggested that the fossils are more closely related to the Megasecoptera than to the Hymenoptera, and may actually belong to that order. Cockerel1 (1927) appears to accept the fossils as of hymen- opterous nature, but suggests some changes in the inter- pretation of the veins. Crampton (1927) agrees that the fossils are hymenopterous and he favors the venational system.
Martynov has studied the question more thoroughly. In 1928, while collecting insects in the Permian beds (Kazan) of Northwest Russia, he discovered a single wing which resembles both the Kansan Protohymenoptera and certain Carboniferous Megasecoptera. From his study of this fossil and the Kansan specimens in the Yale collection, he concluded that the Protohymenoptera were really close to the Megasecoptera, and not related to the Hymenoptera. "They belong to my subdivision Paleoptera containing all orders, chiefly extinct, in which the wings were outspread when at rest. Indeed, both Protohymen perkanus Till. and Permohymen schucherti Till. were found in the out- spread condition of the wings, and even Tillyard thinks



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19301 Permian Insects of Kansas 345
that such a discovery 'suggests very strongly that the in- sects of this family rested with outspread wings, as did the Palseodictyoptera and Megasecoptera.' It is true that among Asthenohymenidse specimens 'occur, in which one wing is laid exactly over the other,' but the same takes place in the case of the Zygoptera, and this manner of resting in Zygoptera in no way proves that they are allied to the Hymenoptera or to any other Holometabola or even Neoptera. When an insect of the division Neoptera, for instance a Neuropteron, puts its wings on the dorsum in roof-shaped manner, the upper surface of the wings is exposed upwards and outwards, with costal borders placed beneath, along the support. On the contrary, in Zygoptera, as also in some Agnatha, when at rest the upper surfaces of the wings are turned inwards, with the dorsal borders looking upwards, i. e., the manner of folding the wings at rest is very different in the Zygoptera and in the Neoptera. Perhaps some forms of Protohymenoptera could 'fold' their wings back on the dorsum, but such 'folding,' in all proba- bility, recalled that of the Zygoptera or of the Agnatha, but not of the Hymenoptera or Copeognatha, or in general, of the Neoptera.
Further, such facts as that the fore and hind wings in the 'Protohymenoptera' were of almost equal size, had the same wing venation, were not linked together in flight by booklets, or by any hairs, and had some veins, at least the costa and the radius 'serrated along its outer edge, in exactly the same manner as that in Odonata,' clearly manifests that the whole order belongs to the divi- sion Paleoptera and is allied partly to the Megasecoptera, partly to the Odonata and Protodonata. The membrane of the wings was glassy, as in Agnatha or in Megasecop- tera or in Odonata. Many Neuroptera, Mecoptera and Hymenoptera and even some Trichoptera have also a glassy membrane, but in these groups the wing membrane is furnished not only with chetoids . . . but also with numer- ous true hairs.
In the wings of the Hymenoptera both
chetoids and hairs are present everywhere, though the hairs sometimes become very small, minute. The wing membrane in the Odonata and Agnatha, as well as, proba- bly, in Megasecoptera, is really glassy, i. e., perfectly de- prived of both hairs and chetoids. One should suppose



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346 Psyche [ December
that in the group ancestral to the Hymenoptera we must inevitably find some remains of hairs, similar to those which exist, for example, in the wings of the fossil Mecop- tera. Thanks to the kindness of Professor Schuchert . . . I examined the excellent remains of both Protohymen and Permohymen, but I could not perceive any hairs on their wings. The wing of the new genus Aspidohymen from Tikhie Gory also does not show any traces of hairs. Thus, such a
character of the wing membrane manifests also that this order is very far removed from the order Hymen- optera, as well as from the remaining Holometabola, and is allied rather to the order Odonata, etc., i. e., belongs to the division Paleoptera."
(1930, p. 79.)
In 1927, one year previous to Martynov's discovery of the Russian specimens, I was fortunate enough to secure fifty-five representatives of the Protohymenoptera in the limestone at Elmo, Kansas. At that time it was already clear that a knowledge of the structure of the body of these insects would help us immensely in determining their affini- ties. Since the Yale specimens consisted only of wings, the chief aim of our collecting trip was to obtain specimens of Protohymenoptera with the body preserved. In this respect we were most successful; several fossils show por- tions of the bodies, and one specimen includes nearly the whole body, with the minutest details perfectly preserved. Through the kindness of Professor Dunbar I have been able to examine the twenty-three specimens of Protohymen- optera in the Yale collection, including Tillyard's types; and I have also found four additional fossils in Dr. Sel- lards' collection. From my study of all these representa- tives of the group, totaling eighty-two, I am convinced that Handlirsch and Martynov were correct in their as- sumption that the Protohymenoptera were megasecop- terous. The evidence which leads me to this conclusion will be presented after the description of the fossils. It will be observed in these descriptions that I have employed an interpretation of the venation which differs from that used by Tillyard, as well as from that offered by Martynov. These changes in the nomenclature of the veins are necessary because both of these authors have been mislead by an erroneous conception of the convexities



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19301 Permian Insects of Kansas 347
and concavities of the veins.
The evidence for this will
also be considered after the descriptions. The Kansan Protohymenoptera were separated by Till- yard into two families, Protohymenidae and Astheno- hymenidae, based largely on the degree of development of the pterostigma, the presence or absence of the radial sec- tor, and the relative lengths of the second anal vein. This classification is a convenient one and is used here, but the most striking difference between the families lies in the width of the costal space. The second of the characteristics mentioned by Tillyard must be omitted, for the radial sector is present in all forms. One change in the terminology of the families must be made. In 1906 Sellards described one of the Kansan fossils in his collection as Doter minor; this specimen consisted of the thorax, abdomen, two long cerci, and the two front wings, but the latter were so twisted and folded that he was unable to determine satis- factorily the nature of the venation. He was not sure of the taxonomic position of the fossil, but believed that it was related to the Protephemeroidea. Handlirsch (1919) placed it within a separate family, Doteridse, and doubt- fully considered it to be a Paleodictyopteran. When I ex- amined this type specimen in 1927 at Austin, Texas, I observed at once from the nature of the wing membrane and the venation that it belonged to the genus Astheno- hymen, which Tillyard had just described. From the photo- graph, drawings, and notes which I made at that time, I am now able to recognize it as the same species which Tillyard called A. ddari, the commonest of all the Proto- hymenoptera in the Elmo limestone. Of course this iden- tity is not obvious from Sellards' figure of Doter minor, for the distortion of his fossil and the fact that the insect possessed an unusual type of venation prevented Sellards from obtaining a correct idea of the venation. This syn- onymy requires us to change the name of the genus from Asthenohymen to Doter, and the corresponding family name to Doteridse. It is interesting to note that Dr. Martynov has observed (1930, p. 85) a similarity between the vena- tion of Doter, as figured by Sellards, and the venation of Asthenohymen, and he has suggested that Doter may be a Protohymenopteron. Dr. Martynov deserves to be con-



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348 Psyche [December
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Carpenter-Permian Insects.




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19301 Permian Insects of Kansas 349
gratulated on the keenness of his observation and the accuracy of his conclusion.
Fore wing: costal margin very slightly convex, nearly straight; costal space present at very base of wing only; Sc rather long, extending a short distance beyond the origin of Rs; R close to the costal margin, except at the very base; Rl diverging downward distally, away from the pterostigma ; pterostigma very slender ; Cu straight at base of wing; Cul and Cu2 diverging near the base; 9-14 cross-veins.
Hind wing: a little shorter and broader than fore wing, and differently shaped; venation similar. Protohymen Tillyard
Protohymen Tillyard, Amer. Journ. Sci. (5) 8 (44) : 113, 1924.
Fore wing; very slender, subpetiolate; R remaining in contact with the costa to the pterostigma and joined to the costal margin by a heavy cross-vein distad of the ptero- stigma ; Cul diverging backward directly after its origin ; 13-14 cross-veins. The positions of the cross-veins, desig- nated by Tillyard as generic features, are subject to great individual variation; but the cross-vein between 1A and the hind margin is always remote from the base of the wing.
Hind wing: more petiolate than fore wing; Cu more remote from R+M at the base than in the fore wing. Genotype : Proto hymen permianus Tillyard. Protohymen permianus Tillyard
Figs. 1, 11
Protohymen permianus Tillyard, Amer. Journ. Sci. (5) 8 (44) : 114, 1924.




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Carpenter~Permian Insects.




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193 01 Permian Insects of Kansas 351
Protohymen permianus Tillyard, Amer. Journ. Sci. (5) 11 (61) : 58, 1926.
Fore wing: length, 10-13 mm.; width, 3 mm.; at the base the costa bends outward slightly, but there is no pre- costal area; R comes in contact with the costa before the divergence of Cul and Cu2, and at the middle of the ptero- stigma it diverges downward toward the apex; Cu2 strongly undulated. Cross-veins: one between Rl and R2+3 ; one between R2^3 and R4+5 ; two between R4+5 and MA; two or three between MA and MP; one between MP and Cul; two or three between Cul and Cu2; two between Cu2 and 1A; one or two between 1A and the hind margin; the cross-veins between 1A and the hind margin are always attached to the distal half of the anal vein. Hind wing: length, 9-11 mm.; width, 3.5 mm.; vena- tion nearly identical with that of the fore wing; the length of the oblique vein between MP and Rs+MA is less than in the fore wing.
Holotype: No. 5001 (hind wing), Peabody Museum, Yale University; paratype: No. 5002 (fore wing), Pea- body Museum ; No. 1702, Tillyard collection. In the Harvard collection there are ten specimens of this species, as follows: No. 3060ab, fore and hind wings complete, with fragments of head and thorax; No. 3061ab, fore and hind wings, folded together, but well preserved; No. 3062ab, hind wing, complete; No. 3063ab, fore wing, nearly complete; No. 3064ab, one pair of fore and hind wings (J. W. Wilson, collector) ; No. 3065ab, hind wing, complete except for the base; No. 3066ab, basal third of wing; No. 3067, middle third of wing; No. 3068ab, distal third of wing; No. 3070ab, basal two-thirds of wing. All specimens collected by F. M. Carpenter, unless otherwise indicated. In the Sellards collection there are two speci- mens : No. 1068, distal two-thirds of fore wing; No. 1558, distal third of wing.
The holotype specimen of this species is well preserved, but the base of the wing is missing. Since none of the other Yale specimens have this part preserved, Tillyard's figure is incomplete at the base. The wing is longer and more nearly petiolate than he supposed. The subcosta is



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not well preserved in the Yale material? but in several of the new fossils it is very clear; it does not extend to the pterostigma as thought by Tillyard.
The pterostigma
is also differently formed than shown in his figure; the photograph accompanying his paper (plate 4? fig. I) dem- onstrates clearly the true nature of this structure. The
costa and Rl widen slightly in the region of the ptero- stigma? so that there is no space formed between these veins until Rl diverges towards the apex. Rl appears to
fork here, but in reality? as I shall show later, the upper "branch" which connects Rl to the margin is a modified cross-vein. There is considerable variation in the distri- bution of the cross-veins, and sometimes a cross-vein may be absent altogether or an extra one added. The holotype specimen has two cross-veins between 1A and the hind margin, but all the other specimens I have seen possess only one here. The short vein, resembling a cross-vein? between the base of Cul and R+M is apparently the basal part of MP.
Protohymen elongatus n. spa
Fig. 2
Fore wing: length, 20 mm. ; width? 3.5 mm. ; extremely slender and pointed ; Rl separating from the costa at the beginning of the pterostigma, so that there is a distinct space between these veins in the region of the pterostigma (text fig. I) ; Cul diverges upward after its separation from Cu2, and touches R+M; Cu2 only very slightly undu- lated; cross-veins distributed essentially as in permianus, but the one between IA and the hind margin is attached to the middle part of the anal vein.
Holotype : No. 3069ab9 Museum of Comparative Zoology ; F. M. Carpenter? collector.
This species is based on a well-preserved and nearly complete fore wing. The only part missing includes the area between the terminations of MP and RI? along the posterior border. Elongatus can readily be distinguished from the previous species by its more slender habitus and the corresponding tenuity of the veins.




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Permohymen Tillyard
Permohymen Tillyard, Amer. Journ. Sci. (5) 8 (115), 1924.
Fore wing: very broad; Rl remaining in contact with costal vein only a little beyond the origin of Rs; ptero- stigrnal space wider than in Protohymen, narrowly tri- angular; Rl without a definite cross-vein between it and the costal margin in the region of the pterostigma; Cul remains parallel to R+M and in contact with it for some distance after its origin ; 9-10 cross-veins ; cross-vein be- tween IA and the hind margin is attached close to the base of the wing.
Hind wing: much broader than the fore wing, especially at the base; venation similar.
Genotype : Permohgmen schucherti Tillyard. Permohymen schwherti Tillyard
Permohymen sch,ucherti Tillyard, Amer. Journ. Sci. (5) 8 (44) : 116, 1924.
Permohymen schucherti Tillyard, Amer. Journ. Sci. (5) 11 (61) : 61, 1926.
Fore wing: length, 13 mm.; width, 3.5 mm.; costal margin bends outward at base; Rl comes in contact with costa just before termination of Sc; Cu2 straight; cross- veins: one between Rl and R2+3; one between R2+3 and R4+5; one between R4+5 and MA; two between MA and MP; one between MP and Cul; two between Cul and .
Cu2; one between Cu2 and IA; one between IA and mar- gin of wing.
Hind wing : length, 12 mm. ; width, 4 mm. ; pterostigma a little broader apically than in fore wing; venation similar.
Holotype : No. 5003, Peabody Museum ; counterpart, No. 1704b, Tillyard collection.
In the Harvard collection there are four specimens of this species, as follows: No. 3071, fore wing complete; No.



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19301 Permian Ir,?sects of Kansas 355
3072a, apical half of wing (fore?) ; No. 3073, hind wing, complete and splendidly preserved ; 3074ab, base of hind wing. In the Sellards collection I find one specimen, No. 308 (reverse, No. 309), fore wing, complete except for apex.
The holotype specimen consists of a very nearly com- plete fore wing and hind wing, with parts of the corre- sponding pairs. The bases of the fore wings are obscured by an abrupt bend in the surface of the rock, and the bases of the hind wings are missing entirely. Tillyard was therefore not able to obtain a clear vision of th.e venation at the base of the wings, and his figure of this part is in- correct. Cul is undoubtedly united with Cu2 at the base of the wing, but it runs along in contact with RiM for quite a distance before diverging backwards; this is essentially the same structure as in Protohymen, only in that genus Cul diverges from R+M much nearer to the base of the wing. The subcosta cannot, of course, be seen in tKe holotype, but it is clearly preserved in several of the Harvard specimens. It is a strongly developed vein, ex- tending only a short distance beyond the posterior diver- gence of Cul. The apex of the fore wing is not pointed as figured by Tillyard. The apex of the left wing, on which Tillyard apparently based his figure, is distorted by an irregularity in the surface of the rock. The true shape of the wing is clearly shown in the right wing of the holo- type, as can be seen in the photograph accompanying Till- yard's paper (plate 4, fig. 2).
Fore wing: anterior margin slightly concave, the maxi- mum curvature near the base of the wing; costal margin serrated; costal space broad basally, present along the en- tire anterior margin of the wing; Sc short, not extending as far distad as the origin of Rs; R remote from the costal margin at the base, approaching nearer to it at the middle of the wing; Rl starts to diverge from the costa near the middle of the wing ; pterostigma weakly developed, slen- derly oval; frequently the pterostigma is bordered distally by a well-developed cross-vein between Rl and the anterior



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