C. Riley Nelson and William D. Tidwell.
Brodioptera stricklandi n. sp. (Megasecoptera: Brodiopteridae), a new fossil insect from the Upper Manning Canyon Shale Formation, Utah (Lowermost Namurian B).
Psyche 94(3-4):309-316, 1987.

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BRODIOPTERA STRICKLANIV. SP.
(MEGASECOPTERA: BRODIOPTERIDAE),
A NEW FOSSIL INSECT FROM THE UPPER
MANNING CANYON SHALE FORMATION, UTAH
(LOWERMOST NAMURIAN B)
BY C. RILEY NELSON' AND WILLIAM D. TI DWELL^ The insect described in this report was collected from the upper- most units of the time-transgressive unit-the Manning Canyon Shale Formation, in central Utah. This formation of Late Missis- sippian to Early Pennsylvanian (Namurian A and B) age consists predominately of shales with interbedded limestones, orthoquartz- ites and some siltstones. A flora has been described from the upper portion of this formation at and near the collecting site for this insect. The flora from the upper Manning Canyon Shale, as presently defined, contains 43 genera and 103 species (Tidwell, 1967; Tidwell et al., 1974; Webster et al., 1984). Thus, it represents the most diversified flora of Carboniferous age presently known in western North America. Plant fossils from this formation consist of fern or fern-like foliage, lycopods, species with calamitean affinities, various seed types, cordaitean taxa and several forms related to microsporangiate structures. These plants indicate that the area was a swampy, moist lowland with perpetual summer-like conditions (Tidwell, 1975).
The Manning Canyon Shale is the oldest horizon reported to be insect-bearing in western North America (Durden, 1984). The age of the uppermost Manning Canyon Shale Formation remains uncer- tain. Many paleontologists and geologists consider the formation to be entirely Mississippian (Upper Namurian A: Bissell, 1959; Gordon and Duncan, 1970; and Webster et al., 1984). Others (Chamberlain and Clark, 1973; Sando, 1985) place the base of the formation in Upper Mississippian and the upper portion in the Lower Pennsyl- vanian (Namurian B and C). The majority of the fossil plants from the upper shales of this formation are encountered only in the Penn- Department of Zoology and 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
Manuscript received by the editor June 10, 1987.



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3 10 Psyche [vo~. 94
sylvanian, whereas a small number of species are typically Missis- sippian. Therefore, we consider this upper flora, as well as the insect, to be lowermost Namurian B in age. The insect specimen described in this paper is a compression fossil consisting of the body, four wings, cerci, legs, and antennae. It is assigned to Brodioptera of the paleopterous order Megasecoptera, which is related to the orders Diaphaneropterodea and Paleodicty- optera. All of these Paleozoic orders had sucking beaks and proba- bly fed by sucking the contents of fructifications and cones of lycopods, cordaiteans, and pteridosperms (Kukalova-Peck, 1983, 1985).
The oldest North American pterygote insect known so far is Met- ropator pusillus Handlirsch collected near "Altamount Colliery" in the Anthracite coal region of Pennsylvania (Carpenter, 1965). It is of historical note that there were two Altamount Collieries both near Frackville, Pennsylvania. Coal mined at the Altamount Col- liery #I was in the Tumbling Run Member of the Pottsville Forma- tion which is Morrowan (Late Namurian) in age and coal mined at the Altamount Colliery #2 was in The Sharp Mountain Member of this same formation. The latter member is of Atokan (partly West- phalian B and partly Westphalian C) age. Since the coals for the Altamount Colliery are not older than the Tumbling Run Member, then the oldest this insect can be is Namurian B. A rich Namurian entomofauna has been described from Europe, including Poland, Czechoslovakia, Belgium, and numerous speci- mens from the Ruhr Valley of West Germany. Further, two very well preserved and almost complete wings of Protodonata were reported from Namurian strata of Argentina in South America by Riek and Kukalova-Peck (1984).
The order Megasecoptera occurs from Upper Carboniferous to Upper Permian (Brues, Melander, and Carpenter, 1954). The numerous families composing this order have been separated mainly on differences in their venation. Most members of this order have been named on the basis of single wings. Because of its complete- ness, the specimen from the Manning Canyon Shale is a significant contribution.




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19871 Nelson & Tidwell- Brodioptera
Fig. 1.
A. Habitus of Brodioptera stricklani n. sp., holotype. B. Drawing of Brodioptera stricklani n. sp. BYU 3243, dorsal view of the male terminalia.



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Psyche
[Vol. 94
Megasecoptera
Brodiopteridae Carpenter, 1963
Brodioptera Copeland, 1957
Brodioptera stricklani n. sp.
Figs. 1-2
HOLOTYPE. Male, UTAH, Utah Co., 1.5 miles East of highway 73 at point 4.3 miles west of Lehi (Sec. 9 T5S R1 W, Utah County, Utah in Jordan Narrows Quadrangle; 40å¡23'30" 11 1å¡57'30"W in clay pits of the Manning Canyon Shale Formation of lowermost Pennsylvanian (lowermost Namurian B) age. Deposited in Brigham Young University paleontological collection as specimen #3 160. DESCRIPTION. Male, length from head to distal portion of geni- talia 26 mm; expanse measured between apices of forewings of complete specimen 57 mm; head with vaguely preserved mouth- parts; antennae filiform, incomplete right remnant 9 mm in length, left remnant 40 mm; legs poorly preserved, three on left side, two on right; length of each forewing 28 mm, width of each forewing meas- ured at apex of Sc 8 mm; length of right hindwing 25 mm, not flat during preservation, length of left hindwing 28 mm, width of each hindwing 8 mm measured at apex of Sc. Wings as in Figs. 2a-d; Sc joining C near midlength of each wing; Ri and Rs forking in basal quarter, Rs with four branches; MA and MP forking at 113 wing length, MA arching anteriorly near fork with MP to nearly contact Rs; CuA and CUP forking in basal 113; anal veins composed of three branches with As seen only in right hind wing (Fig. 2d); cross- veins few but straight and irregularly placed. Abdomen with faint indication of segmentation; genitalia 4.8 mm in length, composed of lateral claspers and medial gonapophyses, neither claspers nor gonapophyses with annulations; cerci long and filiform more than 54 mm in length.
PRESERVATION. The specimen of B. stricklani consists of a part and counterpart with characters better preserved on one rather than the other of the faces. The specimen is lying ventral side up. In this position, the basal portions of the wings are covered from view with the legs lying over parts of the specimen. Details of the head as well as the thorax including attachments of the legs and wings cannot be observed. The specimen is a compression fossil with concavities and convexities of the wings not apparent. Many of the structures of the specimen are indicated by hematitic material; however, the legs and



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19871 Nelson & Tidwell- Brodioptera
Fig. 2. A-D. Wings of Brodioptera stricklani n. sp., holotype; A. left forewing;
B. right forewing. C. left hindwing; D. right hindwing.



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some of the veins are raised areas or impressions depending on which face is viewed. Vague outlines of the head and sucking beak are indicated near the basal end of the antennae. DIAGNOSIS.
Brodioptera stricklani is closely related to B. cum- berlandensis Copeland 1957. Comparison of the wing of B. cumber- landensis (Carpenter, 1963) reveals that it more closely resembles the forewing of B. stricklani than the hindwing. The two may be separated on the basis of details of forking patterns of the veins. The shorter subcosta of B. stricklani joins the costa at about the mid- point of the wing while that of B. cumberlandensis joins the costa well beyond the midpoint. The radial sector, media, and cubitus all fork more distally in B. stricklani than in B. cumberlandensis. Both the fore and hind wings of B. stricklani have a relatively broader anal region than does B. cumberlandensis. Further, the MA in B. stricklani does not curve forward as sharply as in B. cumber- landensis.
ETYMOLOGY. This species is named in honor of Mr. Dave Strick- lan of Provo, Utah, who donated the holotype specimen for this study.
DISCUSSION. The well-spread wings of B. stricklani give supple- mentary support to the concept that Megasecoptera were unable to fold their wings over the dorsum of the abdomen and, therefore, that Carpenter (1963) was correct in placing Brodioptera in Mega- secoptera. Further, the adequately preserved genitalia of this speci- men closely resemble those of extant Ephemeroptera (Edmunds, et al., 1976) and should provide valuable information for future phy- logenetic and evolutionary studies. It is interesting to note, as did Carpenter (1963), that such apparently apomorphic reduction in venation is present in an insect found so near the age (Namurian) in which the oldest winged insects have been discovered. ADDITIONAL MATERIAL. A second specimen consisting of genita- lia and cerci has been collected near the same locality as the holo- type. The specimen is considerably smaller than the holotype but has a similar genitalic structure. This specimen consists of both the part and counterpart that reveal the dorsum of the specimen including the dorsal attachment of the cerci to the abdomen. This second specimen is tentatively assigned to B. stricklani and is depos- ited as figured specimen #3243 in BYU paleontological collection.



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19871 Nelson & Tidwell-Brodioptera 315
We thank Dave Stricklan for the generous donation of the holotype specimen as well as Victor Call for the second specimen. Many thanks are given to Dr. J. Kukalova-Peck of Carleton University for valuable information and numerous suggestions that greatly improved this paper and to Dr. A. T. Cross of Michigan State University for his assistance with stratigraphic problems. Additional thanks are extended to J. Chandler for help with literature searches and to N. Hebbert for inking the illustrations. A fossil megasecopteran of the family Brodiopteridae, Brodi- optera stricklani n. sp., is described. This is the first report of a nearly complete specimen of the family and is the first insect recorded from the transitional Mississippian-Pennsylvanian (Nam- urian A and B) Manning Canyon Shale Formation of Utah in west- ern North America.
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