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Enigmocarnus Chloropiformis Gen. Et Sp. Nov., and Parallel Evolution of Protandrial Symmetry in Carnidae (Diptera)

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Enigmocarnus Chloropiformis Gen. Et Sp. Nov., and Parallel Evolution of Protandrial Symmetry in Carnidae (Diptera) SYSTEMATICS Enigmocarnus chloropiformis gen. et sp. nov., and Parallel Evolution of Protandrial Symmetry in Carnidae (Diptera) 1 MATTHIAS BUCK AND STEPHEN A. MARSHALL Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada, N1G 2W1 Downloaded from https://academic.oup.com/aesa/article-abstract/100/1/9/8317 by guest on 15 November 2018 Ann. Entomol. Soc. Am. 100(1): 9Ð18 (2007) ABSTRACT Enigmocarnus chloropiformis Buck gen. et sp. nov. is described on the basis of a single male from east central Texas. The phylogeny of Carnidae is analyzed based on a matrix of 25 morphological characters, and a key to the World genera of Carnidae is presented. The new genus is characterized by a reduced head chaetotaxy (bristles short and mostly pale), notable gray pruinosity of the body, and a unique conÞguration of pregenital sclerites (the protandrium), which precludes placement of this species in any previously described genus of Carnidae. Enigmocarnus possesses a nearly symmetrical protandrium like Carnus and Meoneura, but the position of sternite 7 indicates that symmetry evolved independently and on a different path. The protandrial peculiarities exhibited by Enigmocarnus are discussed in the framework of Carnoidea relationships. KEY WORDS Carnidae, new genus, phylogeny, protandrium, symmetry In the present article, we describe a new genus and segments 7 and 6 is responsible for protandrial asym- species of Carnidae from North America. Enigmocar- metry. The protandrium is further modiÞed through nus gen. nov. is only the Þfth extant genus to be fusion and loss of sclerites and spiracles, which takes described in this small family after Carnus Nitzsch, place in various ways and to a different extent in Meoneura Rondani, Hemeromyia Coquillett, and different taxa. Such fusion and loss is usually irrevers- Neomeoneurites Hennig (Meoneurites Hennig is known ible and therefore of great value for phylogenetic from Baltic amber only). The new genus exhibits an reconstruction (Hennig 1958). unusual conÞguration of sclerites in the male protan- drium (pregenital segments) and provides a good ex- ample for the usefulness of protandrial characters in Materials and Methods phylogenetic reconstruction. All Cyclorrhapha pos- sess a so-called circumverted hypopygium (with a 360Њ Morphology and Terminology. Abdominal mor- clockwise rotation of the genitalia) (Crampton 1944), phology was studied on specimens that were cleared which is unique in Diptera and is assumed to have in hot 10% KOH and neutralized in glacial acetic acid; evolved only once in the stem species of the Cyclor- cleared parts are stored in glycerin and kept in a rhapha from a postulated inverted (rotated 180Њ clock- microvial with the specimen. The terminology largely wise) ancestral condition (GrifÞths 1972). Rotation of follows McAlpine (1981) except for structures of the the male genitalia to a variable degree has occurred phallic complex, which were taken from Andersson several times independently within Diptera as cou- (1977). The term “protandrium” was coined by Stey- pling and mating positions changed during the evo- skal (1957) and refers to the postabdominal segments lutionary history of the order (e.g., McAlpine 1981). before the hypopygium (segments 6Ð8 or 7 and 8). Ontogenetically, rotation is either completed within The segments and sclerites of the protandrium are also the puparium before emergence (Schizophora: e.g., referred to as “pregenital segments” and “pregenital Schra¨der 1927, Gleichauf 1936) or takes place in part sclerites”. Costal sector 1 is measured from humeral before and after emergence (“Aschiza,” as far as crossvein to subcostal break, Cs2 from subcostal break known; e.g., Kessel 1968). The rotation not only affects to apex of R2 ϩ 3, Cs3 between apices of R2 ϩ 3 and Њ the genitalia proper (which are rotated by 360 ) but R4 ϩ 5. also segment 8 (rotated by 180Њ) and usually segments Photography. Photographs were taken with a Mi- 7 and 6, which are rotated up to 90Њ. Sternites 6 and 7 croptics Digital Lab XLT imaging system using a usually have the appearance of having been dragged Canon EOS 1 Ds camera and Microptics ML-1000 ßash clockwise to a degree dependent on their proximity to Þber optic illumination system. Each image was as- the completely inverted segment, and this rotation of sembled from a series of photographs (with different focal planes) using the computer freeware CombineZ 1 Corresponding author, e-mail: [email protected]. (Hadley 2005). 0013-8746/07/0009Ð0018$04.00/0 ᭧ 2007 Entomological Society of America 10 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 100, no. 1 Downloaded from https://academic.oup.com/aesa/article-abstract/100/1/9/8317 by guest on 15 November 2018 Figs. 1–3. E. chloropiformis Buck sp. nov., habitus. (1) Head and thorax, lateral. (2) Head, frontal. (3) Head and thorax, dorsal. Scale bar ϭ 0.1 mm. Phylogenetic Analysis. Parsimony analysis of the interfrontals, short vibrissa and two pairs of bristles on character matrix was performed with PAUP* 4.0b10 facial ridge above vibrissa. Frons with distinct ocellar for Windows (Swofford 2001) by using the “branch triangle. Antennae in deep foveae; foveae delimited and bound” algorithm. medially and ventrally by sharply deÞned carina. Eye Acronym of Depository. DEBU, Department of distinctly transverse, gena high. Environmental Biology, University of Guelph, Thorax. Thorax (Figs. 1 and 3) with bristles rela- Guelph, Ontario, Canada. tively short and mostly pale including one pair of prescutellar dorsocentrals, two notopleurals, one post- Systematics pronotal, one postalar, two pairs of scutellars, two anepisternals (one reclinate at anterior margin, one dorsoclinate near middle of ventral portion), and one Enigmocarnus Buck gen. nov. katepisternal. No distinct supra-alar bristles. Acros- Type species: Enigmocarnus chloropiformis sp. nov. tichals in approximately four poorly deÞned rows. Diagnosis. Very small (body length Ϸ1.3 mm), Legs simple. Fore and mid-tibiae with ventroapical gray-pruinose ßies (Figs. 1Ð3). Female unknown. bristles, legs otherwise without conspicuous bristles. Head. Head with short, inconspicuous, pale bristles Wing (Fig. 4) as in the genus Meoneura. Costa extend- (Figs. 1Ð3), including four pairs of orbital bristles (Þrst ing to R4 ϩ 5, with humeral and subcostal breaks; sub- and second orbitals inclinate, third exclinate and costa reduced in apical half, last sector of M very faint, fourth reclinate), ocellars, slightly divergent postver- crossveins r-m and dm-cu closely approximated and ticals, inner and outer verticals, small inner occipital shifted toward base of wing, bm-cu, CuA2 and A1 bristles just inside inner verticals, two pairs of small absent, anal lobe well developed. January 2007 BUCK AND MARSHALL: Enigmocarnus GEN. NOV.11 2. Face below antennal foveae high (Fig. 2). Head bristles inconspicuous and pale (Figs. 1Ð3). Height of gena at narrowest point about half eye height (Fig. 1). Supra-alar bristles not differen- tiated (Fig. 3). Thorax heavily gray-pruinose (Nearctic) ........Enigmocarnus gen. nov. Ð Face below antennal foveae reduced, linear. Head bristles well developed and dark. Genal height usually distinctly less than half eye height. Pre- and postsutural supra-alar bristles Fig. 4. E. chloropiformis Buck sp. nov., right wing. Scale differentiated. Thorax shining to dull, never ϭ bar 0.1 mm. gray-pruinose .....................3 Downloaded from https://academic.oup.com/aesa/article-abstract/100/1/9/8317 by guest on 15 November 2018 3. Wing lacking crossvein dm, usually broken off. Abdomen. Preabdomen with four large tergites Postvertical bristles absent. Katepisternal bris- (syntergite 1 ϩ 2, tergites 3Ð5). Protandrium (fused tle small, hair-like. Female sternites 2Ð5 absent. tergite 6 ϩ sternites 6, 7, 8) nearly symmetrical; tergite Abdomen physogastric. Ectobiontic on birds and sternite 6 transverse, forming a complete ring; (widespread) .................Carnus tergite 6 fused with sternite 8 laterally; sternite 7 in Ð Crossvein dm present (as in Fig. 4); wing not mid-ventral position, fused to sternite 6 but free from dehiscent. Postvertical bristles present. Katepis- sternite 8; tergite 7 absent. Spiracles 7 placed slightly ternal bristle well developed. Female sternites asymmetrically. Epandrium saddle L-shaped; sursty- 2Ð5 present. Not ectobiontic on birds (wide- lus simple, setulose. Cerci rudimentary. Subepandrial spread) ....................Meoneura sclerite well-developed, free from epandrium poste- 4. Second orbital bristle reclinate. Facial carina riorly. Hypandrium with broad and short anterior broad, plateau-like, sclerotized throughout. apodeme, fused to phallapodemic plate. Phallapo- Proboscis somewhat elongate, prementum deme small. Postgonites simple. Phallus extremely longer than wide, not bulbous. Anepisternum short, distal portion membranous and bare. Ejacula- bare; two or three subequal katepisternal bris- tory apodeme moderately developed. tles (Argentina, Chile) ......Neomeoneurites Etymology. The name refers to the unusual and Ð Second orbital bristle inclinate. Facial carina nar- initially puzzling protandrial morphology of the male. row, if slightly expanded then median portion pale Recognition. Enigmocarnus is separated from other and desclerotized. Proboscis short, prementum carnid genera in the key below. The genus is very not longer than wide, bulbous. Anepisternum with distinctive because of its short, predominantly pale hairs and bristles; only one katepisternal bristle
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