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Chapter 8. Raphidioptera John D CHAPTER 8. RAPHIDIOPTERA JOHN D. OSWALD' ABSTRACT Three adult snakefly specimens are recorded These are the first authenticated records of Ra- from the Lower Cretaceous Santana Formation, phidioptera, fossil or extant, from South America from near Nova Olinda, Ceara, Brazil. Two species and the Southern Hemisphere. A catalog of de- are described and figured: Baissoptera brasiliensis scribed Raphidioptera fossils (34 named species n. sp. (Baissopteridae) and an unnamed species. in 15 genera and 10 families) is presented. INTRODUCTION Three specimens representing two species AMNH 44400/44407 (paratype, male?, com- of fossil snakeflies from the Lower Creta- plementary halves). ceous Santana Formation of the state of DESCRIPrION: Head: eyes hemispherical; Ceara, Brazil, were recently found in a col- postocular region elongate, abruptly nar- lection of fossil insects belonging to the rowed posteriorly; three ocelli present, ar- American Museum of Natural History ranged in a triangular pattern; antennae fili- (AMNH). These species are described and form, scape enlarged; mouthparts concealed. figured below. One species is assigned to the Thorax: pronotum elongate, enclosing pro- genus Baissoptera, originally described by thorax laterally; legs robust, tarsi 5-segment- Martynova (1961) from the Upper Jurassic ed; forewing as in figure 3; hindwing as in ofthe USSR; the other species is poorly pre- figure 4; wing bases mostly obscured; costal served and is left unnamed. These specimens crossveins simple; remigial crossveins nu- authenticate the past presence ofRaphidiop- merous, arranged posteriorly in well-devel- tera in South America, and may have im- oped gradate series; trichosores absent; Sc portant zoogeographic implications for the joining Cjust proximal to pterostigma; ptero- order. For general accounts of the Santana stigma large and opaque (reddish-brown as Formation, its fossil biota and stratigraphy, preserved). Abdomen: shorter than wings; see papers by Grimaldi and Maisey in this parts of an elongate ovipositor are preserved volume. in specimen AMNH 43287. MEASUREMENTS (mm): AMNH 43287. ACKNOWLEDGMENTS Body length (excluding ovipositor) = ca. 19.5; I thank David A. Grimaldi of the Ameri- forewing = ca. 14.5; hind wing = ca. 12.5. can Museum for the opportunity to examine Body length ofAMNH 44400/44407 also ca. the material treated here, and H. Asp6ck, F. 19.5. M. Carpenter, and J. K. Liebherr for review- DIscussIoN: I tentatively refer these two ing the manuscript. specimens to the genus Baissoptera Martyn- ova based on the following venational char- TAXONOMY acters: in the forewing and hind wing-re- migial crossveins numerous and posteriorly FAMILY BAISSOPTERIDAE arranged in well-defined series, Sc and C Baissoptera brasiliensis, n. sp. anastomosing just proximal to pterostigma, Figures 1, 3, 4 pterostigma long and opaque; in the hind wing only -MA inserting on Rs near divergence MATERIAL: Two specimens; AMNH 43287 of Rs from R. The first of these four char- (holotype female, exposed dorsally) and acters is particularly diagnostic. The venation of B. brasiliensis agrees extremely well with Martynova's figures of the venation of B. ' Department ofEntomology, Comstock Hall, Cornell martinsoni (Martynova, 1961: fig. 6, fore- University, Ithaca, New York 14853. wing) and B. kolosnitsynae (Martynova, 1961: 154 1 990 OSWALD: RAPHIDIOPTERA 155 AWW lo^ "Mft" 2, ...i4 X w _giPs _S 4 . t p ...... r .1 4 2 Figs. 1, 2. Raphidioptera spp. 1. Baissoptera brasiliensis, holotype female, dorsal habitus (AMNH 43287). 2. Raphidiopterawsp., female,,dorsal4habitus (AMNH 44406). Scale lines = 10 mm. fig. 7, hind wing). With the exception of the ally, covered sparsely with small tubercules prominent ocelli observable on AMNH (setal prominences?); legs represented only by 43287, the visible body characters of the fragments or impressions; wings all but lost, present material are also consistent with a single small venational fragment (right fore- Martynova's (1961, fig. 8) diagrammatic rep- wing?) is just visible, other wings each rep- resentation of the head and partial thorax of resented only by a short segment and collin- a Baissoptera sp. Martynova's figure differs ear impression of a single prominent by the lack of ocelli. longitudinal vein (Sc?). Abdomen: position marked by a clear ventral impression, in- Raphidioptera sp. cluding the impression of the proximal por- tion ofan elongate ovipositor; sclerites large- Figure 2 ly lost. MATERLAL: AMNH 44406 (female, ex- MEAsuREMENTs: Body length (excluding posed dorsally). ovipositor) = ca. 12.4 mm. DESCRIPTION: Head: postocular region DIscussIoN: The form of the head and elongate, but proportionally shorter than in pronotum, together with the impression of Baissoptera brasiliensis; ocelli apparently ab- an elongate ovipositor, clearly place this in- sent, but possibly lost with pieces ofthe frac- sect in the order Raphidioptera. The single tured head capsule; antennae lost; labrum visible fragment ofvenation is consistent with rectangular, mandibles large, other mouth- this determination, but is insufficient to allow parts concealed. Thorax: pronotum trape- more detailed taxonomic placement. The zoidal, elongate, enclosing prothorax later- smaller size, shorter postocular head capsule 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 195 '0 .- 1.4 '00 cn '0 o * z. C.) C< rA0_ , 'o o o i Q Ct 04 ._ 000 0 U 1990 OSWALD: RAPHIDIOPTERA 157 length, and apparent lack of ocelli differen- The discovery of fossil snakeflies in Brazil tiate this specimen from Baissoptera brasi- (at ca. 7°S current latitude; ca. 9°S early Cre- liensis described above. taceous latitude [Smith et al., 1981, map 30]) establishes the past presence of Raphidiop- GENERAL DISCUSSION tera in what is now South America. This oc- currence extends the southern distributional The insect order Raphidioptera contains limit of the order Raphidioptera into the approximately 170 living species placed in Southern Hemisphere, approximately 250 two families, Raphidiidae [ca. 150 spp.] and farther south than any previously known fos- Inocelliidae [ca. 20 spp.] (H. Aspock, 1986). sil or extant snakefly. In addition, if the pa- This small extant fauna is generally regarded leoclimatic conditions under which the San- as the relict of a once more diverse fauna tana Formation formed were warm and arid (Carpenter, 1956), based on the relatively (as suggested by Grimaldi and Maisey, this large number of described fossil Raphidiop- volume), the present find suggests that not all tera (presently 34 species placed in 15 genera snakefly lineages have required cold temper- and 10 families, see Appendix 1). atures to complete development. If so, other It has long been noted that living snakeflies factors (e.g., low vagility, subsequent extinc- are geographically restricted to the Northern tion) may be necessary to explain the appar- Hemisphere. Raphidia herbsti Esben-Peter- ent absence of snakeflies from Southern sen (1912: 273), the only snakefly ever re- Hemisphere regions today. ported from the Southern Hemisphere [Chile], The presence offossil species ofthe family was treated by U. Aspock and H. Asp6ck Baissopteridae in Asia and South America (1969) as conspecific with Agulla occidentis suggests that this family was broadly distrib- Carpenter, a species of the western United uted throughout Laurasia and northern States and southwestern Canada (U. Asp6ck Gondwanaland during the Upper Jurassic and 1975). These authors regarded Esben-Peter- Lower Cretaceous. During this period South sen's record as erroneous or adventitious. America and Africa were contiguous (Smith Though the possibility ofthe existence ofliv- et al., 1981). Baissoptera brasiliensis likely ing snakeflies in South America cannot be represents a component ofthis ancestral fau- entirely ruled out (H. Aspock, personal com- na whose fossils have been displaced west- mun.), extant New World Raphidioptera are ward by the fragmentation of Gondwana- presently known to occur south only to ap- land. proximately 19°N in southern Mexico (H. Baissoptera brasiliensis does not appear Asp6ck, 1986: figs. 4 and 5), where they ap- closely related to extant North American pear to occur at high elevations. Raphidioptera (H. Aspock, personal com- Until now, all fossil snakeflies had also been mun.). This observation tends to refute the recovered from Northern Hemisphere sites idea that North American snakeflies may have (North America-British Columbia, Colo- been derived from South American ances- rado, Kansas and Labrador; Eurasia-Baltic tors. But, it does not discriminate between Amber, China, England, and several locali- two other hypotheses ofthe origin ofthe North ties in the USSR), supporting the hypothesis American snakefly fauna: (1) that this fauna that Raphidioptera have always been restrict- was derived solely from eastern Asian snak- ed to northern latitudes. Immatures ofliving eflies which crossed the Bering land bridge snakeflies appear to require a period of cold sometime after the Miocene (Carpenter, 1936: temperature to complete adult development 154), or (2) that this fauna was derived from (H. Aspock, 1986). If one accepts a northern the ancestors ofa static remnant ofan earlier center of origin for the Raphidioptera, this Laurasian snakefly fauna (H. Asp6ck, in requirement could explain (1) the absence of press). recent Raphidioptera from tropical regions, In conclusion, the discovery ofRaphidiop- (2) the apparent restriction of Raphidioptera tera in South America is intriguing
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