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Novltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICANt MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3190, 30 pp., 16 figures, 1 table March 3, 1997 The Bird Flies, Genus Carnus: Species Revision, Generic Relationships, and a Fossil Meoneura in Amber (Diptera: Carnidae) DAVID GRIMALDI' ABSTRACT Variation in the male genitalia of Carnus and are exclusively nest associates of birds, the adults other features reveal that specimens from eastern probably being hematophagous parasites of the and northern North America are the same as the nestlings. Larvae of Carnus are described for the European species, C. hemapterus Nitzsch, 1818. first time. They are most distinctive for the single, A Carnus from western North America is newly midventral row of eight fleshy protuberances. described, C. occidentalis, n.sp., as are two new A new species of carnid is described in 20 mil- species from Florida (C. floridensis, n.sp.) and lion-year-old (Miocene) amber from the Dornini- central Mexico (C. mexicana, n.sp.). Specimens can Republic, Meoneura vieja, n.sp., belongiag to of two probable additional species from southern the sister genus of Carnus. This is the first carnid Mexico are also discussed, for which only female known from the Caribbean. The only other fossil specimens are available. Single records of the ge- camid is in Eocene Baltic amber. Ages of the two nus from southeast Asia and Africa indicate that fossil species are consistent with their phyloge- the genus is probably global, but yet uncovered netic rank, and suggest that carnids probably orig- in most areas because of the specialized collecting inated in the Paleocene, when many other families required to find them. All host records of Carnus of "acalyptrate" Cyclorrhaphan flies were also ra- are summarized and new ones presented. The flies diating. INTRODUCTION In the nests of various birds, primarily in its wings: Carnus. Europeans have been the Holarctic Region, a curious tiny fly sheds studying Carnus hemapterus Nitzsch since ' Curator and Chairman, Department of Entomology, American Museum of Natural History. Copyright © American Museum of Natural History 1997 ISSN 0003-0082 / Price $3.60 2 AMERICAN MUSEUM NOVITATES NO. 3190 the early 19th century (fig. 1), but the genus Mary LeCroy, Dept. of Ornithology wasn't even reported in North America until (AMNH) kindly checked and updated the 1942 (Bequaert, 1942), despite the fact that taxonomic names and classification of the list insect inhabitants of North American bird of bird hosts. Development of the museum's nests had been surveyed before then (e.g., amber collection is made possible by the Dobroscky, 1925). Adults of the fly are usu- generosity of Chairman Emeritus of the mu- ally found attached in small groups to bare seum, Robert G. Goelet. My field and labo- areas of the axillary region on nestlings. ratory research on Dominican amber has Whether they are feeding on blood or skin been sponsored by NSF grant BSR 9020102. secretions has been controversial, as will Pinned material was received on loan from briefly be reviewed below. This report is a the following institutions and their respective detailed morphological study of adult Carnus curators: from throughout the range, originally aimed AM Australian Museum, Sydney (D.K. at examining intraspecific and possible inter- McAlpine) specific variation, and which uncovered sev- CAS California Academy of Sciences (PH. eral new species. The whole study was in- Arnaud, Jr.) spired by the finding of some specimens in CNCI Canadian National Collection of In- the sister genus to Carnus, Meoneura Ron- sects, Ottawa (J. Cumming) dani, in Oligo-Miocene amber (20 million HNHM Hungarian Natural History Museum (L. years old) from the Dominican Republic- Papp) only the second fossil species in the family. HUMB Zoologische Institut, Humboldt Muse- Description of an interesting Meoneura fossil um, Berlin (H. Schumann) INHS Illinois Natural History Survey (Kath- led to a consideration of the generic relation- leen Methven) ships of Carnus, with the possibility of per- MHNP Musee d'Histoire Naturelle, Paris (L. haps identifying the ancestral habit of such Tsacas) specialized flies, and the origins of the car- NHRS Naturhistoriska Riksmuseet, Stockholm nids. (B. Viklund) NMNH U.S. National Museum of Natural Histo- MATERIALS, METHODS, ACKNOWLEDGMENTS ry, Smithsonian Institution (W Mathis) UTSU Utah State University (W. Hanson) Dissection of pinned specimens followed techniques described elsewhere (Grimaldi, SYSTEMATICS 1987). Preparation and study of amber spec- GENUS CARNUS NITZSCH imens is given in Grimaldi (1993), and a reassessment of the age of Dominican amber TYPE SPECIES: Carnus hemapterus Nitzsch, is presented by Grimaldi (1995) and, most 1818. recently, by Ituralde-Vinent and MacPhee Carnus Nitzsch, 1818: 305; Collin, 1911 (synon- (1996). It is a pleasure to acknowledge the ymy of Cenchridobia). help of Roy Larimer, who has allowed me to Cenchridobia Schiner, 1862: 435. sort through his caches of Dominican amber DIAGNOSIS: Tiny flies living primarily in for scientifically valuable specimens, and to bird nests, undoubtedly monophyletic by the Jake Brodzinsky, a constant source of the following suite of apomorphies: notum hav- rare and unusual in Dominican amber. Car- ing one pair of dorsocentral setae; pleural oline Chaboo helped with literature searches membrane of abdomen with dense, setiferous and prevented various distractions from de- spots (more than in other genera of carnids) scending on me. A review and unpublished (figs. 2d; 4a, b; 13); females with abdominal information on the African Carnus were sternites lost (reduced in males) (fig. 13); ab- kindly provided by Eliane DeConinck (Royal dominal tergites reduced in males and fe- Central African Museum, Tervuren). I am males; wings with vein A1 and crossvein extremely grateful for the detailed review dm-cu lost (fig. 11); wings dehiscent (fig. provided by Terry Wheeler (McGill Univer- 2c); adults physogastric, females particularly sity), whose knowledge of acalyptrate geni- so. talia was an important source of information. GENERAL DESCRIPTION: Eye light, cream- 1997 GRIMALDI: BIRD FLIES (CARNUS) 3 I ' l "Ie ,I," -Iga ~ ,~V A;4S'W/tJ-p1117?,^fRt^/2st'W'SgX -----I a I ('d V/ J c .af - a N N Ky If. Fig. 1. The earliest accounts of Carnus. Top, male and female of C. hemapterus (from Germar, 1822); bottom, a normal and engorged female of C. hemapterus (from Egger [1854], Verh. Zool. Bot. Ges. Wien 4, p. 3, fig. 2). The genus was not reported in North America until 1942. 4 AMERICAN MUSEUM NOVITATES NO. 3190 Fig. 2. A more modem view of Carnus: scanning electronmicrographs of C. occidentalis, n.sp. a. Habitus, physogastric female (ventral view). b. Head, profile. c. Base of dehisced wing, which is broken along the line from the subcostal break through the anterior crossvein. d. Setulae in the abdominal, pleural membrane. Each seta is anchored in a sclerotized, conical base. colored, slightly ovoid, bare (no interfacetal ta about same length as orbital setae, with setulae); 1 pair anterior, inclinate orbital setae short pubescence; 2 pairs vibrissae present, and 2 pairs posterior, lateroclinate orbital se- ventral one slightly thinner, setae ventral to tae present; postocellar setae lost; face vir- these upturned; proboscis with labellum small, tually nonexistent, merely a deep, vertical labium heavily sclerotized, broad, sometimes sulcus (closure of sulcus varies greatly); aris- bulbous; antennal bases lying in deep fossae. Fig. 3. Third instar larva of Carnus occidentalis, n.sp. a. Entire larva (ventral). b. Cephalic region (head is collapsed into the oral cavity). c. Ventral protuberance on abdominal segment 5. d. Ventral protuberance on abdominal segment 7. e. Posterior end (dorsal surface is up). f. Cavity into which posterior spiracle is recessed. 1997 GRIMALDI: BIRD FLIES (CARNUS) 5 q Q-ies .,,i. ,,,. A v .!&g-.s,k:,S 00, 6 AMERICAN MUSEUM NOVITATES NO. 3190 Notum with following setae (per side): 1 ekbol, Kelt VIII//13/70, Papp, 16; Alcsiit, postpronotal, 1 presutural, 2 notopleurals, 1 VI/20/63, Csokof6sze K bol Kelt, Warga, 36 dorsocentral, 1 postsutural supra-alar, 1 pres- (diss. no. 18); Szentendre, III-V/70, Papp, cutellar acrostichal; acrostichal setulae in no 3d (diss. no. 17). Sweden: Sk. Malmo. Bel- noticeable rows, most numerous at anterior leviegarden, Klackt ur holk nr. Sturnus vul- end; 2 pairs of scutellar setae. Legs with cox- garis, 28/IV/86, 106' (diss. nos. 19, 20), 4? ae, femora, and most of each tibia brown, (NRS). Switzerland (all in HNHM): Kauf- tarsi yellowish. Female tergites reduced to dorf, Sur poussin de Chereche, 3NVI/79, Bau- narrow, rectangular shape, tergite 1 and 5 de, 1' (diss. no. 16), 29Y; Aniere (GE), nest heavily sclerotized, others much more lightly of Falco tinnunculus, 9/VI/79, Baude, 36. so. USA: Colorado: Colorado Springs, Male genitalia with epandrium complete; V1118/70, R.M. Stabler, ex: Sparrow Hawk, cerci small, unsclerotized, without setae or 1 (diss. no. 7), 29 (NMNH); Indiana: setulae; hypandrium narrow, U shaped; ae- Wayne Co., Centerville, 5/VIII/61, G.L. deagus (phallus) short, bulbous, membra- Ward, Sturnus vulgarus nestling, 16 (diss. nous; paraphysis (paramere) triangular. no. 4) (NMNH). Maryland: Howard Co., 4/VI/85, ex: young kestrels in nest box; Low- Carnus hemapterus ry, Martin, and Wallace, 26' (diss. nos. 1, Figures 4a; 5a, b; 6-8; 13 11), 29 (NMNH). Massachusetts: Middle- hemapterus Nitzsch, 1818: 305; Germar, 1822: pl. sex Co., Cambridge, 7/VII/81, N.E. Woodley, 24, 25; Hennig, 1937: 72.; Sabrosky, 1965 (cat- 26' (diss. no. 12) (NMNH). New Jersey: alog): 729. Cape May Co., Avalon, VI/5/79, W.G. Rob- Cenchridobia eggeri Schiner, 1862: 436. ichaud, ex: osprey nestling, 1 6 (diss. no. 2), Carnus setosus Stobbe, 1913: 193. 19 (NMNH). New York: Albany, IV/4/72, J.A. Wilcox, 26 DIAGNOSIS: Found throughout the Palearc- (NMNH); Tompkins Co., tic region and eastern and northern North Ithaca, ex: falcon (no date or coll.), 16 , 1 9 America, and distinguished from occidental- (INHS); Jamesville, J.R.
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