F. M. Carpenter.
Studies on North American Carboniferous Insects. 3. A Spilapterid from the Vicinity of Mazon Creek, Illinois (Palaeodictyoptera).
Psyche 71(3):117-124, 1964.

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I-f~ntdonnira d/ibasfnsitasi, n.sp. Photograph of halotype (ori~inai). X 2. The fore wing ia the lower of the two wings. Ps,nå´fl 71:ll7-126 [1%4). h~p:ffp!yche.iå´nlcluh~~B/71/7l-l1 htnd



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19461 Carpenter - Carboniferous insects
ORDER PALAEODICTYOPTERA
Family Spilapteridae Handlirsch
Spilapteridae Handlirsch, 1906, Die fossilen Insekten p. 101. The insects included in this family have distinct cross veins, which usually form definite rows; a true archedictyon is absent. The anterior margin of the wings (fore and hind) tends to be concave, often strongly so; the shapes of the fore and hind wings may be very similar (e.g., FIomaloneuru) but more often the hind wing has a broader cubital-anal region (e.g., Dunbaria); in such cases, although the cubitus and anal veins may have a few more branches than in the fore wing, the venation is essentially alike in the two wings. The radial sector has at least six terminal branches, both MA and MP have at least two branches, and CuA has several branches. The wings are commonly marked with bands or spots. The family is known from Upper Carboniferous strata to the Upper Permian. I consider the families Dunbariidae Handlirsch ( 1937); Doropteridae Zalessky ( 1946) and Neuburgiidae Rohdendorf ( I 96 I ) to be inseparable from the Spilapteridae. Genus Homaloneura Brongniart
Homaloneura Brongniart, 1885, Bull. Soc. Amis Sci. Natur. Rouen, 1885 :50- 68; 1893, Recherches pour servir a l'histoire des insectes fossiles des temps primaires, p. 316.
~omaloneukna Handlirsch, 1906, Die fossilen Insekten, p. 46. Homaloneurites Handlirsch, 1906, ibid., p. 47. Homaloneura belongs to that series of genera of the Spilapteridae in which the subcosta etxends nearly to the apex of the wings and in which there is no anastomosis of MA with Rs (or R) or of
CuA with MP (or M). It is related to Doropteron (Permian, USSR), Permiakovia Martynov (Permian, USSR), Neuburgia Martynov (U. Carb., USSR) and Dunbaria Tillyard (L. Permian, Kansas). Unfortunately, our knowledge of even the wings of most of these genera is unsatisfactory; Permidovia is known only from -
the hind wing and Doropteron and Neuburgia are known from the fore wings or parts of the fore wings. Some synonymy may eventu- ally be necessary when more is known about thes,e genera, but
Explanation of Plate 16
Homaloneura dabasinskasi, nsp. Fore and hind wings; original drawing of holotype by Dr. J. Kukalovi. C, costa (+), Sc, subcosta (-), Rl, radius (+), Rs, radial sector (-), MA, anterior media (+ ), MP, posterior media (-), CuA, anterior cubitus (+), CUP, posterior cubitus (-), lA, first anal (+ ).




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19641 Carpenter - Carboniferous insects 121 Homaloneura, being the first of this series to be described, will
remain valid. It is distinguished from the other genera of the Spilapteridae in having the concavity of the front margin of both fore and hind wings very slight and in having the anal region of the hind wing much less extensively developed than in the other genera in which the hind wing is known. Homaloneura elegans Brongniart was designated the type-species of the genus Homaloneura by Handlirsch in 1922 ; a photograph of the type specimen of elegans is given on plate 17 of the present paper. Homaloneura dabasinskasi, n. sp.
Plates 15 and 16
Fore wing: length, 55 mm.; maximum width, 15 mm. Costal margin very slightly concave, apex of wing rounded, hind margin with two slight indentations, one near the posterior branch of MA and the other near the anterior termination of MP. Costal space broad in the basal half of the wing but narrowing abruptly towards the middle of the wing and continuing of uniform width for the rest of the wing until the termination of Sc, just before the apex; R and RI nearly parallel to Sc; Rs arising about a quarter of the wing length from the base, dividing to form five main branches, about half of which fork once again; the branches of Rs tending to arise more dichotomously than pectinately; M arising from the base of the wing as an independent vein, forming MA and MP slightly distally of the origin of Rs; MA with three terminal branches; MP with four; Cu apparently arising independently from the wing base, forking very shortly into CuA (with four branches) and CUP (forked). The basal connections of the anal veins are not clearly preserved; the first of these terminates in three branches, the next three are forked. Cross veins distinct, arranged as shown in plate 16. The wing is distinctly marked with transverse bands and spots.
Hind wing:
length, 50 mm. ; maximum width, 18 mm. This is shaped like the fore wing except that the concavity of the costal margin is slightly more pronounced and the anal area is somewhat more rounded. The number of anal veins appears to be the same -- - -- - -
Explanation of Plate 17
Homaloneura elegans Brongniart, U. Carboniferous, Commentry, France, Photograph of type in Laboratoire de Paleontologie, Paris. (Original). X 4. Note the prothoracic lobes and the similarity of the wing venation to that of H. dabasinskasi. The posterior margin of the hind wing is in- completely preserved.




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19641 Carpenter - Carboniferous insects 123 in both wings. The venational pattern is basically as in the fore wing, with only slight differences in the positions of the branches) mainly those of the radial sector. The wing markings and cross veins are essentially as in the fore wing. Holotype: collected by Mr. Walter Dabasinskas in June, 1960, in an ironstone nodule, found at the Greer Earthmoving and Mechanics Schoo'l, Will County) Illinois, 4 miles north of raid- wood. The type is contained in Mr. Dabasinskas' collection and the species is named for him. This fossil is the most spectacularly preserved fossil insect which I have ever seen in a nodule from the Mazon Creek region and it is indeed one of the striking Carboni- ferous insects known to me. As can be observed in the photograph (plate 15) the fore wing is posterior in position to the hind wing and the apex of the fore wing rests near the base of the hind wing. That these two wings were derived from one side of the same specimen of an insect seems almost certain. The subcosta is concave in one wing and convex in the )other; presumabIy, the wings broke away from the body as the insect rested in water and one of the wings turned over as well as rotating through 180å before coming to rest in the mud. The counterpart of the half of the specimen shown in plate 15 has the apex of the fore wing and basal region of the hind wing completely preserved.
A comparison of the wings of dabasinshmi with those of H. elegans and the other Commentry species shows enough differences to require a distinct species for the new f,oIssil but not enough in my opinion to justify generic separation. The hind margin of the fore wing of dabminskmi is somewhat more sharply curved basally than in the Commentry species. The wing markings of dabasins4asi are distinctly different from those of elegans, which has a dark, longitudinal stripe along RI distally; but they are similar to those of another Commentry species, H. ornata Brongniart. In addition to the features mentioned above as specific in nature, several other structures preserved in this new fossil deserve mention, since they may turn out to be common to all Spilapteridae and perhaps to other families of the Palaeodictyoptera. One of these is the series of fine ridges at the base of the costal area of both fore and hind wings (see plate 16). These do not appear to be veins in Explanation of Plate 18
Homaloneura bonnieri Brongniart, U. Carboniferous, Commentry, France. Photograph of type in Laboratoire de Paleontologic, Paris. (Original). X 2. Note the prothoracic lobes and the similarity of the hind wing to that of H. dabasinskasi




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124 Psyche [September
this species and the more basal area cannot be considered as a precostal area) as it occurs in the Orthoptera or Protodonata. Since similar ridges or actual veinlets have been observed (Kukalovi, 1960) in other Palaeodictyoptera (Breyeria and Ostrava) they may turn out to be characteristic of most Palaeodictyoptera. Another interesting structure present in dabasins4asi is the thickened basal stem of R. Examined under certain angles of illumination) the posterior edge of this stem appears as an independent vein extending almost from the base of M to the origin of Rs. Careful examination of the base of the wing, under various types of illumination) however) has con- vinced me that this apparent vein is actually the posterior edge of the stem of R. Similar widening of R can be seen in the Commentry specimens of HomaZonew-a.
To enable further comparisons of dabminskmi with the described species of HomaZoneuraJ I am including here (plates 17 and 18) phofiographs of the types of H. elegans Brongniart and of H. bon- nieri Brongniart) which are contained in the Laboratoire de Paleontologic of the Museum National d9Histoire Naturelle) Paris. It is regrettable that the Illinois fossil does not show such body structures as the prothoracic lobes) which are visible in both of the Commentry fossils depicted.
BRONGNIART, c.
1885. Les insectes fossiles des terrains prirnaires. Bull, Soci6ti Arnis des Sciences Naturelles de Rouen, 1885: 50-68. 1893. Recherche8 pour servir 5 l'histoire des insectes fossiles des temps primaires. Saint-Etienne, pp. 1-493.
HANDLIRSCH, A.
1906. Die fossilen Insekten, Leipzig, p. 108. 1922. Fossilium Catalogus, (1) 16: 47.
KUKALOVL, J.
1960. New Palaeodictyoptera (Insecta) of the Carboniferous and Permian of Czechoslovakia. Sbornik UUG, paleon. 25 ~239-250. ROHDENDORF, B. B.
1961. Palaeozoic insects of the Kuznetsk Basin, Trudy Paleont. Inst., 85 : 72.
ZALESSKY, G.
1946. Sur un reprksentant d'un nouveau ordre des Hemiodonates de Perimien de 1'0ural. Bull. SOC, Nat. Moscou, 51 : 63-70.



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