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 origin 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

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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

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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. Philips, VI/24/76, is, n.sp., by the relatively longer surstylus ex: nest of American kestrel, 26 (CNC). (apical width/total length 0.40) (fig. 6), and Rhode Island: Richmond, VI/1/71, A. Lav- by the long paraphysis in lateral view (0.54 allee, 3d (diss. nos. 3, 10), 3? (NMNH). greatest width/length) (fig. 7). Distinguished Utah: Cache Co., Smithfield, 27/V/74, TL. from orientalis as given below. Comparisons Whitworth, magpie nest in willow, 76' (diss. between eastern North American and Euro- no. 21), 8? (UTSU, 1 of each sex in pean specimens revealed no consistent dif- AMNH); West Nibley Lumber Mill, 16-18/ ferences, although the ventral lobe of the IV/74, T.L. Whitworth, magpie nest, Nos. epandrium in some European Carnus was 1758, 1788, 36 (diss. no. 22), 2? (UTSU). longer and thinner. HOSTS: Bequaert (1942) summarized the MATERIAL EXAMINED: AFRICA: Egypt, 25 hosts and localities for European specimens km W. Marsa' Matruih, W. Desert, IV/28/59, and records of Carnus hemapterus, which in- M.N. Kaiser, nest of Corvus corax, 3 d (diss. cluded 29 species of birds in 15 families. He no. 9), 2? (NMNH); CANADA: British noted that there was a slight predilection for Columbia: Osoyoos L., N. end, 21/IV/85, birds nesting in sheltered areas or cavities, R.J. Cannings, in saw whet owl nest, 1 6 but never a species that was a ground or (diss.), 2? (CNC). New Brunswick: F'ton swamp nester. Capelle and Whitworth (1973) [Fredericton], VI/27/5 1, N.R. Brown, "nest reviewed all the North American records, and young of Sphyrapicus varius", 2? which included 15 species of raptors, passer- (CNC); Priceville, near Docktown, VII/6/68, ines, and woodpeckers, but, again, no ground ex: nest of yellow-bellied sapsucker, 2? nesters. Fitzner and Woodley (1983), Main (CNC). Ontario: no specific information. and Wallis (1974), and Wilson (1977) added EUROPE: England: Oxford, 111/54, J.R. new records of raptor and woodpecker hosts. Vockeroth, 19 (CNC). Hungary (all in Additional records are added here from mu- HNHM): Aranyosgandany, gyurgyalag fesz- seum specimens that have been studied. Ap- 1997 GRIMALDI: BIRD FLIES (CARNUS) 7

Carnus floridensis

Carnus sp. A Carnus sp. B Carnus mexicana Fig. 4. Heads and abdomens of Carnus occidentalis, floridensis, mexicana, and two species from Chiapas, Mexico known only from females. 8 AMERICAN MUSEUM NOVITATES NO. 3190

C. hemapterus (New Jersey, 62) (Egypt, 68)

C. mexicana

C. floridensis (holotype) Fig. 5. Male terminalia of Carnus. Epandrium, surstyli, and genitalia of C. hemapterus, from New Jersey and Egypt, and of C. floridensis, n.sp. (holotype). Genitalia and surstyli of C. mexicana, n.sp. 1997 GRIMALDI: BIRD FLIES (CARNUS) 9

NEARCTIC PALEARCTIC EASTERN WESTERN hemapterus I I occidentalis l

hemapterus

Indiana (64) J

British Columbia (d29) Fig. 6. Variation in and comparison between surstyli in C. hemapterus and C. occidentalis. pendix 1 is a complete summary of all host the laboratory and found that they readily records of Carnus. fed, and were attracted to, blood from FEEDING: Whether the adults feed on the scratches. Engel (1919) reported that the pro- blood of birds (primarily nestlings) or skin boscis is usually attached to where the quill secretions has been controversial. Noller inserts (feeding, perhaps, on oily secretions), (1920) transferred Carnus adults to birds in and that no blood was seen through abdom- 10 AMERICAN MUSEUM NOVITATES NO. 3190 inal membranes in freshly collected flies. Be- phalic region is entirely covered with simple quaert (1942) noted that the mouthparts were (unbranched) spinules (fig. 3b). The posterior not similar to other higher flies that had rasp- end has five pairs of lobes (fig. 3e), and the ing structures on the labellum, such as Sto- posterior spiracles are retractable enough to moxys (Muscidae) and tsetse flies (Glossini- be entirely recessed (fig. 3f). dae), and so were probably unable to pene- DISTRIBUTION: European records of Carnus trate the skin. Capelle and Whitworth (1973) hemapterus were summarized by Bequaert found, as did Fitzner and Woodley (1983), (1942), which included the following coun- small groups of 3-8 adults feeding in bare tries: Austria, Finland, Germany, Italy, Lith- axillary regions of nestling Swainson's uania, the Netherlands, Romania, Switzer- hawks and red-shafted flickers. Of most in- land, and Yugoslavia. Additional records are terest was that Fitzner and Woodley observed Spain (Carles-Tolra [1993]), Hungary (Papp, dried blood spots in the axillary region where 1984; this study), eastern Russia (Papp, the flies were found. Evidence does not pre- 1984), and Sweden (Papp, 1984; this study). clude, and in fact suggests, that the flies are Papp (1984) also listed "Nearctic and Afro- blood feeders. This could be simply ad- tropical Regions" under the distribution of dressed by first starving the flies, letting them C. hemapterus; some of these records, as feed, and then examining their gut contents shown below, refer to other species. soon after their removal from the host. Sim- American records of Carnus were most re- ply removing the flies from a host in nature cently reviewed by Capelle and Whitworth would not guarantee that the fly has fed re- (1973), with new records cited by Fitzner cently. and Woodley (1983) and Cannings (1986). REPRODUCTION: Reports conflict as to States and provinces previously reported are: whether Carnus is ovoviviparous. De Mei- Arizona (Bequaert, 1951: species identity un- jere (1913) cited Brauer (1880) as stating that certain, specimens were possibly occidental- first-instar larvae were extracted from the ab- is, n.sp.), Baja California (Bequaert, 1951; domen of a gravid female [presumably based Sabrosky, 1965: probably occidentalis, on specimens from Germany]. Capelle and n.sp.), British Columbia (Bequaert, 1951; Whitworth (1973) mentioned that a female Cannings, 1986), Florida (Bequaert, 1942: from Utah laid eggs. probably C. floridensis, n.sp., see below), In- LARVAE: Until now, the larvae have been diana (Wilson, 1977), Massachusetts (Main undescribed, although de Meijere (1913), and Wallis, 1974), New Brunswick (Sabros- Hennig (1937), and Capelle and Whitworth ky, 1965), New Jersey (Kirkpatrick and Col- (1973) illustrated the puparium. The pupar- vin, 1989), New York (Bequaert, 1942), Utah ium is distinctive for its produced posterior (Lloyd and Philip, 1966; Capelle and Whit- spiracles and pronounced, evenly spaced an- worth, 1973: included hemapterus and occi- nuli (numbering about 16-22). Using mate- dentalis, but these species were not distin- rial from the Capelle and Whitworth study, guished at the time), Washington state (Fitz- the third-instar larva is most distinctive for ner and Woodley, 1983, specimens of which the midventral row of eight fleshy protuber- could have been occidentalis or hemapterus). ances, one per abdominal segment (figs. 3a, Additional records, based on material ex- c, d). This is apparently unique for the Cy- amined for this study, include Colorado, clorrhapha. The protuberances are not exten- Maryland, and Rhode Island. Fitzner and sions of the creeping welts, since there are Woodley remarked that the species was un- no curved spinules on them (figs. 3c, d), as known from the midwestern (Great Plains) is seen in the assorted cyclorrhaphan larvae states (e.g., Dakotas, Minnesota, Iowa, Ne- with paired "prolegs" (e.g., Ephydra braska, Kansas, Texas, New Mexico, Wyo- [Ephydridae], Cladochaeta [Drosophilidae]). ming, and Montana, although it has been Function(s) of the protuberances are un- found in Colorado). Two slide-mounted known, but unlikely to serve in locomotion specimens were examined from the CAS, la- because there are no spinules. Each protu- beled as "Rocky Mt. Laboratory, 19 June, berance has either a single, central slit or a 1963, G. D. Lloyd, Nest 21-2N," which is pair of paramedian slits (figs. 3c, d). The ce- in Hamilton, Montana (P. H. Arnaud, Jr., per- 1997 GRIMALDI: BIRD FLIES (CARNUS) I1I

sonal commun.) (orientation of the mount Malaysia ("Selangor, Rantau Panjang, 8 km prevents a definitive identification, but the N. Klang, ex: juvenile fish owl"). Holotype male specimen appears to be hemapterus). is in the Bishop Museum, Honolulu (no. Previous authors maintained that this might 7604), paratypes in the Natural History Mu- represent a disjunction of eastern and west- seum, London; and the National Museum of ern populations. Carnus hemapterus clearly Natural History, Smithsonian Institution. occupies the eastern half of North America, It should be noted that an intensive survey and northern latitudes west to northern Utah of nearly 70 bird nests, from 23 species and and southern British Columbia (fig. 9). Car- eight families of birds, done in Hokkaido, nus occidentalis, n.sp., is the fly former au- northern Japan, revealed no Carnus (Iwasa thors called hemapterus from southwestern et al., 1995). North America. An apparent absence of Carnus in mid- Carnus occidentalis, new species western North America may be attributable Figures 2, 3, 6, 7 to the poor sampling for these flies, but the distinct identity of the western North Amer- DIAGNOSIS: Very similar to C. hemapterus ican species (reported below) suggests the but consistently distinguished from it by disjunction to be real. Indeed, an examina- male genitalia in occidentalis having a short- tion of bird nests from western Montana re- er, deeper surstylus that is almost square (api- vealed no Carnus (Jellison and Philip, 1933). cal width/total length approximately 0.57, vs. Prior to the present study, virtually no com- 0.30 to 0.45 in hemapterus [fig. 6]); paraph- parisons had been made among individuals ysis (paramere) in lateral view much shorter, of Carnus from various localities. Bequaert almost triangular (greatest width/greatest (1942) mentioned that he was unable to com- length = 0.75, vs. 0.55 in hemapterus) (fig. pare European and American specimens, but 7). did note (on the basis of old descriptions) Head, notum, pleura light, shiny brown to that the setation in the abdominal pleural black; proclinate interfrontal setae cruciate; membrane was slightly different in the ocellar setae almost parallel; antenna with American specimens, and the "[external] pedicel tan, flagellomere 1 dusty gray. No- male terminalia [were] virtually identical." tum with apical scutellar setae cruciate, leg Given that apparent Holarctic species are of- coloration typical for genus. Setulae on ab- ten two species, Nearctic and Palearctic (al- dominal pleural membrane short, longer ones beit subtly different), separate identities are dorsad. Male genitalia: synsternite VII + not surprising. VIII, a vestigial pair of crescentic sclerites flanking epandrium; paraphysis (paramere) Carnus orientalis longer than wide; surstylus with blunt teeth on medial edge, trapezoidal in shape, longer Carnus orientalis Maa, 1968: 33. than wide. DIAGNOSIS: Separated from hemapterus TYPES: Holotype (male): UTAH: Box El- and occidentalis by orientalis having a nar- der Co., Mantua, Devil's Gate, ex: flicker rower interocular distance; labium shorter [species], 14/VII/68, T.L. Whitworth. In and slightly longer; cheek not as deep; ab- AMNH. Paratypes: 14d (diss. no. 23), 19? dominal sternites (males) almost square from same series as holotype (in AMNH and (rectangular in other species); abdominal UTSU). pleural membrane with setulae sparser in OTHER MATERIAL: California: Contra both sexes; female tergites much larger than Costa Co., Walnut Creek, V/12/50, S. Kent any other species in the genus. Species status Carnie, "Host: Falco peregrinus anatum, un- is unquestionable based on somatic features dersides of wing at elbow jt., among pin alone, but diagnostic features of male geni- feathers and down," 1 d (diss. no. 5), 1 ? talia could not be deciphered from Maa's de- (NMNH). Idaho: Boise Co., Lucky Pk. Nur. scriptions or illustrations. 14/V1184, R. Meadows, "Malaise trap," 1 6 SPECIMENS: Known only from the type se- (diss. no. 15). Utah: Box Elder Co.: Devil's ries of 8 dealate males and 11 females from Gate, VII/69, TL. Whitworth, flicker nest, 12 AMERICAN MUSEUM NOVITATES NO. 3190

NEARCTIC PALEARCTIC EASTERN WIESTERN

hemapterus oci

'.-0VI Massachusetts (c12) / dz

Rhode Island (610) U1

New Jersey (62)

Ca Maryland (611)

(X ) \ heemapterus

Colorado (67) / floridensis LUtah (621)

Briti ish Columbia (629)

Fig. 7. Variation in and comparison between paraphyses and other parts of the male genitalia in C hemapterus, C. occidentalis, and C. floridensis.

9G, 2? (diss. nos. 13, 14); 5 mi NW Brig- ETYMOLOGY: "Western," referring to its ham City, 25/V/93, C.D. Marti, "from ab- location in North America. domen of Tyto alba chicks", 2 ?; Davis Co.: DIscusSION: Distribution of this species 3 mi W. Kaysville, 25/V/93, C.D. Marti, 3 Y, overlaps that of C. hemapterus, at least in "paras. of Tyto alba chicks"; Summit Co., northern Utah. It should be noted that the Heiner's Canyon, nr. Coalville, 25/VI/64, brief survey by Lee and Rychman (1954) of G.D. Lloyd, ex: hawk nest, 12-3N (1G, 1 : owl nests in California revealed no Carnus. CAS); nr. Coalville, 6500 ft, G.D. Lloyd, 25/VI/64, on nestling sparrow hawk, 3 G Carnus floridensis, new species (diss. no. 6). Willard BSN, 9/VII/69, Capelle Figures 4b, 5c, 8 and Whitworth, on flicker nestling, 33G, DIAGNOSIS: Eye light, cream-colored, con- 24? (in UTSU and AMNH). trasting with dark blackish-brown body 1997 GRIMALDI: BIRD FLIES (CARNUS) 13

(head, all of thorax, labium, femora, most of ETYMOLOGY: In reference to Mexico, the each tibia). Antenna with pedicel and flagel- only known locality. lomere I dark, black-brown. Ocellar setae divergent; proclinate interfrontal setae conver- Carnus sp. A gent but not cruciate; labium bulbous. Setu- Figures 4c; 8c lae on pleural membrane of abdomen black, about twice the length of setulae in other spe- FEATURES: Larger black species; ocellar se- cies. Female tergites narrow, except tIV tae divergent; proclinate interfrontal setae (W/L = 2.2 [vs. 2.4-3.6 in other species]); upright, almost parallel, another pair poste- tergites with long setae on posterior margin, riad and nearly cruciate. Proboscis slightly about twice the length of setae in other spe- longer and thinner than in other species (ex- cies. Male genitalia with epandrium having cept Carnus sp. B), labellum small and nar- long, stout setae, about twice the length of row. Abdominal setae (pleural setulae and other species; synsternite VII + VIII com- tergal setae) black, long. Male unknown. plete dorsally; aedeagal apodeme short (ca. SPECIMENS: MEXICO: Chiapas, 8 mi E. 0.7X length in other species); paraphysis a San Cristobal, 19NV/69, B. Peterson, ex: owl simple, triangular shape; surstylus rectangu- nest. 2? . In CNC. lar, not trapezoidal, with blunt teeth on the DIScuSSION: Both this species and Carnus apical edge. sp. B appear to be distinct species on the TYPES: Holotype (male): FLORIDA: Wak- basis of somatic features only. However, ulla Co., 15/V/92, on woodpecker nestling since the other species of Carnus in this (diss. no. 24). Paratype: female, same data. study are most consistently diagnosed on the Both specimens in AMNH. basis of male genitalia, I hesitate to describe ETYMOLOGY: In reference to the type and these as new species without additional ma- only known locality, in Florida, USA. terial. Carnus sp. A and B are sympatric at least in southern Mexico, but were reared from different hosts. Carnus mexicana, new species Figures 4e, 5d-e, 8b Carnus sp. B DIAGNOSIS: Coloration light brown to tan, Figure 4d all tarsi yellowish white. Head: Proclinate in- FEATURES: Body entirely dark, black- terfrontal setae convergent, but not cruciate; brown, except for eyes and light yellowish ocellar setae parallel to slightly divergent; vi- tarsi. Proboscis slightly longer and thinner brissae with large distance between two ip- than in other species (except Carnus sp. A), silateral ones (distance approx. 5X the di- labellum narrow. Proclinate interfrontal setae ameter of the seta). Thorax: scutellar setae virtually upright, convergent to almost par- with apical pair parallel to convergent (in allel. Eye longer than high, cheek very deep; very replete females they are cruciate from vertex flat in profile. Apical scutellar setae pressure of the abdomen). Abdomen: pleural with tips virtually touching. Tergites not ob- setae short; female tergites narrower than in served well, since specimens considerably other species, especially tIl, tIll (W/L = 3.2 shriveled, but pleural and tergal setae long, [vs. 3.8-4.0 in other species]); apical sternite as in floridensis. in female less sclerotized and smaller (by ap- SPECIMENS: MEXICO: Chiapas, 8 mi E. prox. one-half the size) in other species. San Cristobal, 19/V/69, B. Peterson, ex: Male genitalia: synsternite VII + VIII com- Sparrow hawk nest. 10? (in CNC). plete; surstylus with just 2 blunt apical teeth, roughly triangular in shape; paraphysis tri- African Carnus angular, longer than wide. sp. TYPES: Holotype (male): MEXICO: Du- DeConinck (1986), cited by Barraclough rango, 10 mi W. El Salto, ex: flicker young. (1994), reported the first Carnus from sub- Paratypes: 15?, 7d, same data. Holotype saharan Africa. Locality: ZAIRE: near Mit- and 18 paratypes in CNC; 4 paratypes in waba, 08°51'S, 26°43'E, Kaswabilenga, AMNH. 14/X/47, Rene Verheyen, on nestlings of the 14 AMERICAN MUSEUM NOVITATES NO. 3190

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sp. B floridensis Fig. 8. Female tergites in Carnus. Distension of the abdomen is trivial variation. 1997 GRIMALDI: BIRD FLIES (CARNUS) 15 kingfisher, Halcyon albiventris. Since this ism rates of birds may be high, the flies have kingfisher is widespread through South Af- little noticeable effect on the birds. Carnus rica, Gabon, Zaire, Kenya, and Somalia, the hemapterus was found on 91 of 103 (88%) fly is thought to be similarly widespread. If nestling barn owls (Tyto alba) in southwest- this Carnus is as polyphagous as the Euro- ern New Jersey (Kirkpatrick and Colvin, pean and North American species, it proba- 1989). The flies were on the chicks only until bly feeds on many birds. A description of the fifth week. In Germany, Carnus hemap- this species will be published by DeConinck. terus infested nestlings of the starling, Stur- nus vulgaris, from hatching, and the heaviest GENERAL DISCUSSION infestation was when the nestlings were six and seven days old (Walter and Hudde, All but one of the more than 65 records 1987). Parasitism apparently did not reduce of Carnus associate the fly with nests of tree- breeding success of the starlings. nesting birds (appendix 1) (the one exception Adults of Carnus probably do not disperse is the mention by Maa [1968] of Carnus he- with their avian hosts. No reports have ever mapterus on the gray egret, Ardea cinerea, been made of banded, mist-netted, wild-cap- in The Netherlands). More than one-third of tured or even domestic birds having Carnus the records report Carnus in raptor nests, 10 on them. The flies may just be much easier records in woodpecker nests, 9 with corvids, to detect in a nest or nest hole and on nest- and the remaining records being one to four lings and fledglings. However, Carnus lacks reports each in various families of smaller, modifications seen in fleas and other ecto- perching birds (passeriformes). The apparent parasitic Diptera (such as hippoboscids, stre- predilection for raptors may be due to the blids, and nycteribiids), which allow the par- conspicuous size of the nests, and the carrion asite to grip the hairs or feathers of its flying debris that falls into the bottom of the nest, or moving host. These include an extremely in which larval carnids would breed. Later I flattened body (for moving among hair and discuss why tree-nesting birds are parasit- feathers) and combs (ctenidia) or strong ized. claws for gripping. Carnus also lacks the dis- Behavior of the flies and field observations tinctive ectoparasite feature of reduced eyes. indicate that the flies specifically feed on If Carnus must disperse to nests simply by blood and/or oily and other secretions of its own flight, instead of by phoresy, then the nestling birds. This is probably their sole broad range of hosts seen in C. hemapterus way of life, given the distinct adaptations of must be directly related to this method of Carnus for avian ectoparasitism, such as de- host colonization. hiscent wings; short, stiff setae over much of Three of the five species of Carnus are the body; and physogastry. Presumably, an each known from only a single series or pair enormous clutch of eggs or one or more large of specimens from one nest (e.g., orientalis, larvae are produced with the volume of mexicanus, floridensis). This applies as well blood/oil meal that distends the female ab- to three additional, undescribed species from domen two to three times the size of the rest Mexico and Africa. No records exist from of the body. Physogastry is also seen in the South America, or from India, southern Chi- hippoboscid fly Melophagus, a hematopha- na, Indochina, or even from central and east- gous ectoparasite of ungulates, although the ern Russia; and one doubts if appropriate ef- physogastry is not as pronounced as in Car- forts to sample flies like Carnus have ever nus. Interestingly, additional protein for vi- been made in these regions. Given the spotty tellogenesis in Carnus may be derived from records of Carnus from Africa and Central autolysis of the flight muscles, which was America, and the poor sampling of them shown to occur in histological studies by from an area as intensively studied as North Mercier (1928). A feature that preadapts America, it would not be surprising if Car- Carnus well to ectoparasitism are antennae nus were found on all continents except Aus- deeply recessed in fossae (which exists in the tralia (and Antarctica, of course). In fact, other genera of carnids). based on land areas, and the diversity of trop- A few studies indicate that, while parasit- ical bird faunas, Carnus is likely to be sev- 16 AMERICAN MUSEUM NOVITATES NO. 3190 eral times more diverse in the African, In- length of flagellomere I. Vibrissae stout, sub- dopacific, and New World tropics than are tended by 4 pr. of subvibrissal setae: pair im- presently known to exist in northern temper- mediately ventral to vibrissae smallest, next ate regions. Fifty world species of Carnus pair similar in size and orientation to vibris- would not be an unreasonable number to ex- sae; setae of large lateral pair upturned and pect, but this would require an extremely am- divergent; lateralmost pair convergent. Pro- bitious program of field work to decipher. boscis with small palps and labellum; pre- mentum only slightly bulbous, considerably GENUS MEONEURA smaller than in Carnus. Face width/head Figures lb, 12b, 14, 15 width = 0.41; cheek depth/eye depth = 0.34. Meoneura Rondani, 1856: 128; Hennig, 1937: 59 Thorax: Notum with flat dorsal surface; (subsequent citations and synonyms). acrostichal setulae well developed, not in rows. Setae: 1 large postpronotal; 1 presu- Meoneura vieja, new species tural (lateral); 2 notopleurals (anterior one slightly stouter); 1 pr. large prescutellar ac- DIAGNOSIS: Easily distinguished from Car- rostichals; 2 supra-alars; only 1 (large) pair nus and other Meoneura with abdominal of dorsocentrals; 2 pr. scutellars. No color spots on the basis of the distinctive male gen- patterns distinct on thorax or abdomen. italia, with surstyli that are deeply forked, for Wings: completely hyaline. Section of costal virtually one-half their length (similar to that vein proximal to sc break with row of 5 in the living species Meoneura anceps Frey, stout, black, spinelike setae (no indication of M. californica Sabrosky, and M. milleri Gre- a line of weakness in this area). R2+3 short. gor); surstyli without brushes of long setae; Costal vein ending midway between apices ventral lobe beneath the cerci bifurcate, with of R2+3 and R4+5. Crossveins dm-cu and r-m several fine setae at tips of the lobes. Ab- present, separated by a distance approxi- dominal segments 3-5 with tergites virtually mately 1.5 times length of crossveins. Vein devoid of setae; pleural membrane with setu- Ml very light, barely noticeable. Vein A, lae more numerous than in most Meoneura. largely absent. Wing length = mm. Legs: DESCRIPTION: Head: Unicolorous brown. Forefemur with dorsal row of 3 and ventral Eyes bare, without fine pile, much lighter in row of 4 long setae; midtibia with 2 large color than rest of head. Orbital setae: 2 pos- apical setae; hind femur with ventral row of terior pairs strongly lateroclinate; 2 anterior 5-6 stiff short setae on distal half, one seta pairs inclinate, all approximately equal in twice the size of others. size (anteriormost orbitals slightly shorter); a Abdomen: Male and female tergites 2-4 much smaller, finer setula lying between each virtually devoid of setae, with only 2-3 setae orbital seta; ipsilateral setae and setulae in near lateral margins. Tergite 5 with normal line. Pair of small mediofrontal setae present; setation, including row of larger setae along convergent, with ends touching. Two pairs posterior margin. Tergite 6 minute, with sev- small inclinate setae bordering ocellar trian- eral long setae, segment 7 with 2 lateral gle. Ocellar triangle extended to ca. 0.6X plates. Pleural membrane of segments 1 and length of frons. Ocellar setae thick (slightly 2 without setulae; segment 3 with 4-5 setu- thicker than other setae on head), strongly lae, each one arising from dark spot at its divergent. One pair of fine, parallel post- base; segments 4 and 5 with more setulae ocellar setae present. Inner vertical setae lon- than normally found in Meoneura. Female gest on head; pair of outer vertical setae di- sternites small. Ovipositor telescoping (seg- vergent, ca. 0.7X length of inner verticals. ments 6, 7, 8), apically membranous. Surstyli Face with narrow carina lying between 2 of male genitalia without brushes of long se- deep fossae; base of carina forming membra- tae; ventral lobe beneath cerci bifurcate, with nous triangle at oral margin. Fossae have several fine setae at tips of lobes. steep, definite edges extended laterally to in- TYPES: Holotype: Male: AMNH DR-14- ner margin of eye and ventrally to just be- 600, in amber from the Miocene of the Do- tween vibrissae. Antenna with pedicel with 1 minican Republic (Grimaldi, 1995), although stout seta on anterior surface; arista ca. 2.5X specific provenance is unknown. Genitalia 1997 GRIMALDI: BIRD FLIES (CARNUS) 17

Fig. 9. Records of Carnus in the New World.

are well preserved and exposed. Paratypes: phylinid beetle. Area around carnid was re- female, in excellent condition, AMNH no. moved and polished into a small piece so as 11 842A. Originally in large, clear, deeply to better view the face. A fine crack along colored piece of amber also containing a sta- the midventral line of the abdomen obscures 18 AMERICAN MUSEUM NOVITATES NO. 3190

Fig. 10. Global distribution and records of Carnus. this area somewhat: the piece must be care- 2. Apomorphic: One pair of dorsocentral fully positioned and lit to view the sternites. setae present. Plesiomorphic: Two pairs. Paratype, female, AMNH DR-14-599, spe- 3. Apomorphic: Pair of (proclinate) inner cific locality in Dominican Republic un- frontal setae present. Plesiomorphic: known. None present. ETYMOLOGY: Spanish for "old man," re- 4. Apomorphic: Aedeagus (phallus) long, ferring to age of the specimen. coiled, with extensive spicules. Plesiom- orphic: short, bare. GENERAL DISCUSSION 5. Apomorphic: Female cerci fused (McAlpine, 1989). Plesiomorphic: Cerci GENERIC RELATIONSHIPS: Below is a list of not fused. apomorphic characters used in constructing 6. Apomorphic: Bases of antennae (pedi- the cladogram of genera and other groups of cel, flagellomere recessed into carnids The were I) deep (fig. 16). characters polar- fossae. Fossae not ized based on comparison to the outgroup Plesiomorphic: pres- family Milichiidae, which has been generally ent. recognized as the 7. Apomorphic: Facial carina broad, at sister group to the Carni- least width of antenna. dae. Since the characters are completely con- Plesiomorphic: sistent with each other (fig. 16), the clado- Carina very narrow. gram was not analyzed with computerized al- 8. Apomorphic: Male tergite VI reduced or gorithms. Character numbers given below lost. Plesiomorphic: not reduced or lost. appear on the cladogram, and dates of the 9. Apomorphic: Abdominal pleural mem- two amber fossils are superimposed for es- brane with sparse to many fine setulae, timates of the minimum age of several each with a sclerotized base. Plesiom- clades. orphic: Pleural membrane bare. 10. Apomorphic: Apical half of vein Ml 1. Apomorphic: Upper fronto-orbital setae very weak. Plesiomorphic: Vein M, as directed outward/laterad; lower (anteri- strong as other veins. or) fronto-orbitals directed inward/me- 11. Apomorphic: Labium enlarged to bul- diad. Plesiomorphic: All fronto-orbitals bous. Plesiomorphic: Width of labium directed mediad or posteriad. no more than rest of proboscis. 1997 GRIMALDI: BIRD FLIES (CARNUS) 19

Meoneura vieja

Meoneura pteropleuralis Fig. 11. Wings of Carnus and Meoneura: Carnus occidentalis, n.sp.; the amber fossil, Meoneura vieja, n.sp.; and Meoneura pteropleuralis Sabrosky. All to the same scale. 20 AMERICAN MUSEUM NOVITATES NO. 3190

12. Apomorphic: Anal vein very reduced or lost. Plesiomorphic: Anal vein distinctly present, extended almost to margin of wing. 13. Apomorphic: Small basal wing cell (bm- cu) not closed. Plesiomorphic: Cell closed. 14. Apomorphic: Vein Sc incomplete or lost. Plesiomorphic: Vein Sc present, strong, often extending to costal vein. 15. Apomorphic: Rs vein meets vein R slightly proximal to inner end of the basal cell. Plesiomorphic: Rs meets R distal to end of basal cell. 16. Apomorphic: Male tergite VI lost. Ple- siomorphic: Tergite VI reduced. 17. Apomorphic: Portion of costal vein proximal to subcostal break with 2 rows of large, spinelike setae. Lengths of setae about 2.5x width of costal vein. Ple- siomorphic: Setae shorter and thinner, lengths no more than 1.5 X width of costal vein. 18. Apomorphic: Crossveins very close to- gether, separated by distance equal to or slightly greater than length of crossvein. Meoneura digitata Plesiomorphic: Crossvein distance much greater, at least several times the length of either crossvein. 19. Apomorphic: Postvertical setae cruciate. Plesiomorphic: Setae convergent or parallel. 20. Apomorphic: Setulae in pleural membrane of abdomen long and thick, with large, heavily sclerotized base to each seta (McAlpine [1989] erroneously indicated that this feature was restricted to males). Plesiomorphic: Setae much smaller, without such heavily sclerotized bases. 21. Apomorphic: Surstylus (male genitalia) deeply bilobed. Plesiomorphic: Not lobed; a simple, undivided lobe. 22. Apomorphic: Broad lobe or pair of lobes on surstylus with fringe of long setae. Plesiomorphic: Without fringe. 23. Apomorphic: Setulae in pleural mem- Meoneura vieja brane of abdomen very numerous (sev- Fig. 12. Female abdomens of two Meoneura eral hundred per side). Plesiomorphic: (ventral view): M. digitata Sabrosky (extant) and Much sparser, 50 or less. M. vieja, n.sp. (fossil). Setation in vieja is actually 24. Apomorphic: Abdominal sternites of fe- typical for most of the living Meoneura, that in male lost. Plesiomorphic: Present. digitata is highly derived. 1997 GRIMALDI: BIRD FLIES (CARNUS) 21

." :..1.-t r T.,...I 1r* r 't I;' r r .

Carnus hemapterus Meoneura sp. Fig. 13. Female abdomens of Carnus hemapterus and Meoneura sp. (from Yellowstone National Park, Wyoming). Telescoping terminal segments of the Meoneura are everted. 22 AMERICAN MUSEUM NOVITATES NO. 3190

Fig. 14. Paratype female of Meoneura vieja, n.sp. (AMNH 11842A) in Dominican amber. Habitus and frontal view of the head. 1997 GRIMALDI: BIRD FLIES (CARNUS) 23

*Ai

Fig. 15. Photomicrographs of Meoneura vieja, n.sp., in Dominican amber. Above, female paratype (DR-14-599); below: male holotype (DR 14-600), with rendered detail of genitalia in terminal view. 24 AMERICAN MUSEUM NOVITATES NO. 3190

Meoneura

Miocene- - l_ __ _ |______28 1 9 28 27 21 26 25 AEocene 18424 .17 23 16 15 o 14 Eocene 13 12 10

4 .3

Fig. 16. Cladogram of generic relationships in the Carmidae. No attempt was made here to define monophyly of Australimyza, Meoneurites, Neomeoneurites, and Hemeromyia.

25. Apomorphic: Physogastry. Plesiomorph- duced, especially in females. Plesiom- ic: No physogastry. orphic: Tergites extend laterad. 26. Apomorphic: Crossvein dm-cu lost. Ple- siomorphic: Crossvein present. The inclusion of Australimyza Harrison, as 27. Apomorphic: Wing dehiscent. Plesiom- a subfamily in the Carnidae, has been pro- orphic: Not dehiscent. posed elsewhere (Colless and D.K. Mc- 28. Apomorphic: Aedeagus (phallus) short, Alpine, 1991; J.F. McAlpine, 1989), although abbreviated, without vestiture. Plesiom- Griffiths (1972) indicated there was no close orphic: Long, coiled, with spicules. relationship between this genus and carnids. 29. Apomorphic: Abdominal tergites re- Australimyza is composed of 6 described and 1997 GRIMALDI: BIRD FLIES (CARNUS) 25 many undescribed species from New Zealand Male genitalia of Meoneura offer a pleth- and Australia, where they breed in beach ora of features for cladistic analysis of the wrack. The clade of Neomeoneurites (two genus. However, for purposes here only cer- species in Chile [Hennig, 1972; Wheeler, tain obvious features were selected to group 1994]) and the Baltic amber fossil, Meoneu- some species. One feature, character 20, rites Hennig (1965), is defined on the basis groups the North American species Meoneu- of a broad facial carina (character 7). Whee- ra pteropleuralis Sabrosky and M. digitata ler (1994) discussed several apomorphic Sabrosky (fig. 12, top). At least four North male genitalic features of living Neomeoneu- American species (californica, lamellata, rites, the existence of which are unknown for flavifacies, and wirthi) and one European the fossil Meoneurites. It is interesting that, species (milleri Gregor) are grouped on the without the Baltic amber fossil, one would basis of characters 21 and 22. The fossil have assumed that the early history of the Meoneura appears to be a basal member of carnids was entirely an austral one (as based this clade, since it possesses character 21 but on Australimyza and Neomeoneurites). The not 22. One should not expect this prelimi- extinction of present-day austral taxa from nary attempt at cladistics of Meoneura to northern latitudes has actually been discussed necessarily make much biogeographic sense, elsewhere, specifically with respect to organ- particularly in light of the very poor Neo- isms in Baltic amber (Ander, 1942). The next tropical sampling of this genus. However, it taxon on the cladogram is Hemeromyia, a ge- can be assumed on the basis of these rela- nus of four Palearctic, two Nearctic, and one tionships that the genus Meoneura possibly African species. Little is known of its biol- originated in the lower Miocene to late Oli- ogy, but the report by Cole (1969), of spec- gocene. imens in New Mexico reared from the nest With an Oligo-Miocene origin of Meoneu- of a mouse (Peromyscus truei), is consistent ra, and appearance of the more plesiomorph- with what is known of the habits of other ic clade, Meoneurites + Neomeoneurites, by carnids. the time of the Eocene Baltic amber, the Car- The close relationship between Meoneura nidae can be assumed to appear by the Pa- and Carnus has never been questioned and leocene. Given the virtual lack of higher cy- rests on substantial morphological criteria: clorrhaphan fossils from the Cretaceous, car- characters 10-15, above. It is possible, in nid origins in the Paleocene would be entire- fact, that Carnus may eventually be found to ly consistent with radiations of the modern be paraphyletic with respect to Meoneura, families of Cyclorrhapha in the earliest Ter- since only two characters of Meoneura are tiary. known thus far to be apomorphic with re- EVOLUTION OF HOST USE: Like most car- spect to Carnus: the spinose base of the cos- nids, Meoneura is closely associated with fe- tal vein in Meoneura (character 17), and the ces of mammals and birds. In fact, there are close proximity of the crossveins on the wing several records of Meoneura in close asso- of Meoneura (character 18) (which is impos- ciation with birds. Meoneura lamellata was sible to evaluate for Carnus, since the distal reported about the "openings of sand mar- crossvein is lost). Thus, effectively only one tin's burrows" in England (Collin, 1930), apomorphy establishes the monophyly of and in the nests of bank swallows in Alaska Meoneura with respect to Carnus. The ques- (Sabrosky, 1959). Two records associate tion of Meoneura paraphyly could probably Meoneura with marine birds: Collin reported be answered with a complete, worldwide re- M. seducta as being probably associated with vision of the genus. It should be noted that gannetts on Grasholm Island. (Collin, 1937) characters 26 (loss of crossvein dm-cu) and and an undetermined species of the fly was 10 (atrophy of vein M1) in Carnus may be associated with a penguin rookery on Gough developmentally very closely associated: Island (in the south Atlantic, in the Tristan striking convergence of both features appears de Cunha group) (Barraclough, 1994). There in Paramyia nitens (Milichiidae). Paramyia are two records of Meoneura reared from old has even lost vein A1 as in Carnus, but the bird nests: M. neottiophila, in England (Col- basal cells remain. lin, 1930); and M. hungarica, from a Juncus 26 AMERICAN MUSEUM NOVITATES NO. 3190 nest in Hungary (Papp, 1977b). Clearly, the terous ectoparasites, the preening and ruf- ancestal Carnus also bred in bird nests, like fling of stiff, adult contour feathers would some Meoneura. probably prevent the adult flies from reach- The almost exclusive use by Carnus of ing the skin. Whatever the mode of host evo- tree-nesting birds, vs. ground nesters, may lution, the parasitic lifestyle of Carnus adults relate to the development of the young birds. is derived from behavioral and morphologi- Most ground nesters have precocial young cal specializations that further used the con- that leave the nest quickly after hatching; tree tents of the nest: its inhabitants. nesters have predominantly altricial young The Meoneura in amber were preserved (nestlings), which live in the nest for several almost certainly when flying around trunks weeks. For the flies to parasitize young pre- of the amber tree (Hymenaea). It is plausible cocial birds, emergence of adult flies would that they were breeding in or searching for need to be perfectly synchronized with the the nests of birds in Hymenaea trees, since hatching of the birds. Nestlings are parasit- feathers are also preserved in Dominican am- ized instead of adult birds probably because ber. One feather, in fact, has been identified their skin is more accessible, with at best a as from a woodpecker (family Picidae), prob- layer of down that the flies must penetrate. ably closely related to the extant Antillean Since adult Carnus lack the extremely flat- Piculet (Nesoctites micromegas) (Laybourne tened and hardened bodies seen in other dip- et al. 1994).

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an einigen anderen Trypanosomen. 1987. Carnidae. In J. F McAlpine (ed.), Man- Arch. Protistenkd. 41: 149-168. ual of Nearctic Diptera 2: 909-912. Papp, L. Agric. Canada Monogr. 28. 1977a. Notes on some Becker's types. Ann. Schiner, J. R. Hist.-Nat. Mus. Natl. Hung. 69: 185- 1862. Vorlaufiger Commentar zum dipterolo- 189. gischen Theile der "Fauna austriaca." 1977b. New species and records of Hungarian Odiniidae, Milichiidae and Camidae V. Wien Entomol. Monschr. 6: 428- (Diptera). Acta Zool. Acad. Sci. Hung. 436. 23: 171-181. Stobbe, R. 1984. Family Carnidae. In A. Soos and L. 1913. Zur kenntnis der Gattung Carnus Papp (eds.), Catalogue of Palaearctic Nitzsch (= Cenchridobia Schiner) mit Diptera 10: 118-124. Amsterdam: El- 1 nov. sp. (Dipt.). Dtsch. Entomol. Z. sevier. 1913: 192-194. Rondani, C. Walter, G., and H. Hudde 1856. Dipterologiae Italicae prodromus. Vol. 1987. Carnus hemapterus (Milichiidae, Dip- 1: Genera Italica ordinis dipterorum or- tera), an ectoparasite of nestlings. J. Or- dinatim disposita et distincta et in fam- nithol. 128: 251-255. ilias et stirpes aggregata. Parma. 228 Wheeler, T. A. PP. 1994. A new species of Neomeoneurites Sabrosky, C. W. 1959. The Nearctic species of the filth fly ge- (Diptera: Carnidae) from Argentina, nus Meoneura (Diptera, Milichiidae). with a discussion of male postabdom- Ann. Entomol. Soc. America 52: 17- inal structure in the genus. Can. Ento- 26. mol. 126: 435-441. 1965. Carnidae. In A. Stone et al. (eds.), A Wilson, N. catalogue of the Diptera of American 1977. Ectoparasites found in the nest cavities North of Mexico. U.S. Dep. Agric. of pileated woodpeckers in Oregon. Agric. Handb. 276: 728-729. Bird-Banding 48: 171-173. 1997 GRIMALDI: BIRD FLIES (CARNUS) 29

APPENDIX

Nest (N)/ Carnus Bird Host Family/Species Location Bird (B) Species Ref. ARDEIDAE: Ardea cinerea Netherlands B hemap f PANDIONIDAE: Pandion haliaetus New Jersey B hemap ACCIPITRIIDAE: Haliaetus albicilla Finland N hemap a Aquila heliaca Yugoslavia B hemap a Buteo swainsoni Washington B h/occ b Buteo jamaicensis calurus Utah B h/occ c FALCONIDAE: Falco peregrinus California B occid Finland N hemap a Germany B hemap k Falco tinnunculus Austria N hemap a Germany N hemap a, k Switzerland N hemap i Falco cherrug Romania B hemap a Falco mexicanus Utah B h/occ c Falco sparverius Colorado B hemap i Maryland B hemap New York B hemap Utah B h/occ c, d Mexico N sp. B i Falco sp. Italy B hemap a COLUMBIDAE: Columba livia Finland N hemap a Columba oenas Finland N hemap a Germany B hemap k TYTONIDAE: Tyto alba Germany N, B hemap a, k New Jersey B hemap j Utah B occid i STRIGIDAE: Otus asio Florida N/B flori? a Arizona B occid? e Bubo ketupu Malaya B orient f Aegolius acadicus Brit. Columbia B hemap i "owl" Mexico B sp. A i Aegolius funereus Finland N hemap a Athene noctua Germany B hemap k PICIDAE: Picoides major Austria N hemap a Finland B hemap a Picus viridis Germany B hemap a Dryocopus martius Finland N hemap a Colaptes auratus Mexico N mexic i New York B hemap i Utah B occid d, i Williamson's sapsucker Utah B h/occ d 30 AMERICAN MUSEUM NOVITATES NO. 3190

APPENDIX Continued

Nest (N)! Carnus Bird Host Family/Species Location Bird (B) Species Ref. Sphyrapicus thyroideus Utah B h/occ d New Brunswick B, N hemap pileated woodpecker Oregon N h/occ i "woodpecker" Florida B flori JYNGINAE: Jynx torquilla Germany B hemap a Switzerland B hemap a HIRUNDINIDAE: Delichon urbica Finland N hemap a TURDIDAE: Turdus philomelos Finland N hemap a Turdus iliacus Finland N hemap a Phoenicurus phoenicurus Finland N hemap a SYLVIIDAE: Regulus regulus Finland N hemap a Sylvia atricapilla Germany N hemap a PARIDAE: Parus atricapillus Finland N hemap a Parus ater Finland N hemap a Parus major Germany B hemap k CERTHIDAE: Certhia familiaris Finland N hemap a FRINGILLIDAE: Fringilla coelebs Finland N hemap a PLOCEIDAE: Passer domesticus Switzerland N hemap a STURNIDAE: Sturnus vulgaris Finland, Germany, Italy, Sweden Switzerland N/B hemap a, i, k Indiana B hemap g, i Wisconsin N/B hemap? d CORVIDAE: Pica pica Finland N hemap a Utah B hemap a, d, i Corvus corone cornix Finland N hemap a Egypt N hemap i Corvus corone Lithuania B hemap a Germany B hemap a, k Corvus monedula Netherlands N, B hemap a Finland B hemap a Germany B hemap a, k References: a, Bequaert (1942); b, Fitzner & Woodley (1983); c, Lloyd & Philip (1966); d, Capelle & Whitworth (1973); e, Bequaert (1951); f, Maa (1968); g, Wilson (1977); h, Guigan et al. (1983); i, this study (museum specimens); j, Kirkpatrick & Colvin (1989); k, Walter & Hudde (1987).