PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON
ADULT DIPTERA FROM OZARK CAVES
JEFFREY K. BARNES, MICHAEL E. SLAY, AND STEVEN J. TAYLOR
(JKB) Department of Entomology, University of Arkansas, Fayetteville, AR 72701, U.S.A. (e-mail: [email protected]); (MES) The Nature Conservancy, Ozark Highlands Office, 675 North Lollar Lane, Fayetteville, AR 72701, U.S.A. (e-mail: [email protected]); (SJT) Illinois Natural History Survey, 1816 South Oak Street, Champaign, IL 61820, U.S.A. (e-mail: [email protected]) PROC. ENTOMOL. soc. WASH. 111(2), 2009, pp. 335-353
ADULT DIPTERA FROM OZARK CAVES
JEFFREY K. BARNES, MICHAEL E. SLAY, AND STEVEN J. TAYLOR
(JKB) Department of Entomology, University of Arkansas, Fayetteville, AR 72701, U.S.A. (e-mail: [email protected]); (MES) The Nature Conservancy, Ozark Highlands Office, 675 North Lollar Lane, Fayetteville, AR 72701, U.S.A. (e-mail: [email protected]); (SJT) Illinois Natural History Survey, 1816 South Oak Street, Champaign, IL 61820, U.S.A. (e-mail: [email protected])
Abstract.-The Diptera fauna of Ozark caves is fairly typical of that found elsewhere in the United States east of the Great Plains. The sphaerocerid Spelobia tenebrarum is the only troglobitic dipteran found in Ozark caves. Probable troglophiles include the mycetophilid Macrocera nobilis, the psychodids Psychoda spp., and the phorid Megaselia cavernicola. The tipulids commonly found in Ozark caves, such as Dactylolabis montana, Dolichopeza spp. and Limonia spp. are probably all trogloxenes, as are the Trichoceridae and Heleomyzidae. The mycetophilids, Exechiopsis umbratica and Rymosia sp., and the culicids are among the most common cave dipterans in the study area. The culicids overwinter in caves and other protected places; they are properly classified as trogloxenes. The main sources of food for cave Diptera are other insects, carrion, guano, and allochthanous plant debris. Key Words: cavernicolous, Tipulidae, Mycetophilidae, Sciaridae, Cecidomyiidae, Psychodidae, Trichoceridae, Culicidae, Empididae, Drosophilidae, Dolichopodidae, Phoridae, Heleomyzidae, Sphaeroceridae, Antho myiidae, Bibionidae, Calliphoridae
Until recently, little research has been Oklahoma and included records for 23 published on cavernicolous Ozark Dip Diptera, but most identifications were tera. Most papers that have appeared list not beyond family level. McDaniel and species found in individual caves, giving Smith (1976), McDaniel et al. (1979), few data on collection dates or sexes and Dunivan et al. (1982) reported on encountered and little indication of the life forms found in Arkansas Ozark ecological status of the species or their caves, including 16 species of Diptera, geographical distributions. Hubricht most of which are also reported here. (1950) listed only species that appear to Peck and Lewis (1978) reported on the be able to live and breed in the total invertebrate fauna of caves in Illinois darkness of caves, including the Diptera and eastern Missouri Ozarks. They had species Megaselia cavernicola (Brues) very few Missouri Diptera records. Peck and Spelobia tenebrarum (Aldrich). Black and Peck (1982) found five species of (1971) summarized the cave life of flies in the sandstone crevice caves at Devils Den State Park, Arkansas, but * Accepted by David R. Smith they identified only the mycetophilid 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Macrocera nobilis Johnson to species large percentage of the total biomass in a level. Gardner (1986) inventoried organ given cave. The following records add to isms from more than 400 caves in 38 the growing body of evidence that might Missouri counties. Many of the 25 one day allow better judgments about Diptera reported were identified only to how Diptera use caves. family or genus level. Sutton (1993) published lists of cave species from a MATERIALS AND METHODS mineral prospecting area in southern Names of major collectors are abbre Missouri. Again, many of the Diptera viated in the text: C. J. Bitting (CJB), C. were identified only to family or genus Brickey (CB), G. Graening (GG), D. level. Graening et al. (2003, 2006) listed Fenolio (DF), M. E. Slay (MES), and S. some Diptera from caves in the Sylamore J. Taylor (SJT). Specimens were collect Ranger District of Ozark National For ed opportunistically during visual sur est and the Buffalo National River in veys of caves and by using baited pitfall North Central Arkansas, and they sug traps. Pitfall traps were baited with gested ecological classifications for some rancid Limburger cheese spread and left of the taxa. Barnes (2004) reported on in place for one to six days. overwintering cave mosquitoes of the Identifications were made by the Arkansas and Missouri Ozarks. Elliott senior author except as noted in the (2007) reported on the zoogeography acknowledgments. Specimens were iden and biodiversity of Missouri caves and tified to genus using the keys found in karst and included Spelobia tenebrarum the Manual ofNearctic Diptera, volumes in his list of troglobites and phreatobites 1 and 2 (McAlpine et al. 1981, 1987). in Missouri but otherwise mentioned Species level identifications were made Diptera only briefly. using keys cited in the Manual or more Here we report on additional collec recent literature as appropriate. Speci tions made by M. E. Slay, G. O. mens are preserved in 70% ethanol in Graening, and D. Fenolio in Arkansas vials. Arkansas and Oklahoma speci and Oklahoma and by S. J. Taylor and mens are deposited at the University of M. E. Slay at Fort Leonard Wood Arkansas Arthropod Museum, while (FLW), Pulaski County, Missouri as Missouri specimens will be deposited in part of a study of the cave fauna of that the Illinois Natural History Survey. In military installation (Proffitt 2005). addition to the taxa recorded here, adult It is difficult to assess the true ecolog and immature Chironomidae were also ical classifications of many of the taxa found in some caves. No attempt has yet discussed in this paper. Most Ozark cave been made to identify these specimens. Diptera can probably be regarded as trogloxenes. These are species that enter RESULTS AND DISCUSSION caves, perhaps for shelter, but do not Tipulidae reproduce in caves and are not entirely dependent on the cave environment. Austrolimnophila sp.-ARKANSAS: Troglophiles, on the other hand, com Madison Co.: Hunter's Cave, 28-Apr plete their life cycles in caves, but they 01, GG, J. Gunter, R. Honebrink, B. can also occur in certain environments Wagner, D. Wagner (1 <2). outside the cave. Troglobites must live Dactylolabis montana (Osten Sacken). their entire life underground (Jackson ARKANSAS: Stone Co.: Bud Wallis 1997). Troglobitic and even troglophilic Cave, 28-Apr-02, GG, MES, CB, M. Diptera species are rare (Vandel 1965), Covington (2 rS, 2 <2). MISSOURI: although a few species may represent a Pulaski Co.: FLW: cave no. 23PU280,
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11-May-04, SIT, MES (1 MES, CJB (1 ~). MISSOURI: Pulaski Washington Co.: Synder Cave, 16-Sep Co.: FLW: Chuck's Virgin Cave, 20-Apr 01, MES, R. Honebrink, B. Potter (2 J, 03, SJT, MES (1 ~). 2 ~). MISSOURI: Pulaski Co.: FLW: Boletina sp.-ARKANSAS: Stone Profitt Cave, 24-Mar-03, SJT, K. Loh Co.: Janus Pit, I-Feb-03, GG, MES, D. raff, V. Block (sex unknown); Wolf Den Kampwerth, E. Corfey (1 J). Cave, 16-Jun-04, SJT, MES (3 J, 1 ~). Brevicornu sp.-ARKANSAS: Stone Exechiopsis is a widespread genus, Co.: Blanchard Springs Caverns, 27 occurring throughout most of the North Apr-02, GG, MES, K. Tinkle (I J). ern hemisphere. It was, in terms of both Cerotelion johannseni (Fisher).-MIS species and individuals, the most abun SOURI: Pulaski Co.: FLW: Camp Vista dant mycetophilid genus found hibernat Cave, 13-May-04, SJT, MES (1 J). ing in Estonian caves (Kurina 1996). An Exechia sp.-ARKANSAS: Stone unidentified species was previously re Co.: Blanchard Springs Caverns, 27 ported from a Sharp County, Arkansas Apr-02, GG, MES, K. Tinkle (I J). cave by McDaniel and Smith (1976). Exechiopsis umbratica (Aldrich).- Exechiopsis umbratica was described ARKANSAS: Marion Co.: Forest Trail from a female specimen collected in Ridge Cave, 6-Jun-02, MES (3 J, 3 ~), Shilo Cave, Indiana in July (Johannsen 5-Jun-04, MES, E. Frank, J. Terry (10 1912). It is one of the most abundant fly ~); Overlook(ed) Cave, 5-Jun-04, MES, species collected during the course of the E. Frank, J. Terry (8 ~), 8-Jun-04, MES, present study, and specimens were often CJB (1 J, 3 ~); Toney Bend Mine #2, collected from bedrock walls and ceil 24-May-04, MES, CJB (4 ~, I J); ings, sometimes in association with Newton Co.: Jaw Bone Cave, 16-Sep spider webs. Both males and females 04, MES, CJB (1 ~); Stone Co.: Blan were collected from March to October. chard Springs Caverns, 27-Apr-02, GG, Only one specimen was taken during the MES, K. Tinkle (8 J); Rowland Cave, winter, so it is doubtful that E. umbratica 12-Jun-01, GG, CB (I J, 2 ~). MIS hibernates in the adult stage in Ozarks SOURI: Pulaski Co.: FLW: Cave caves. It seems likely that it is either 23PU493, IO-May-2004, SJT, MES (3 breeding or aestivating in caves during J, 9 ~); Breeden Cave, 25-0ct-03, SJT, the warmer months. It is probably either MES (1 J, I ~); David's Crawl Cave, a troglophile or a trogloxene. 12-Apr-04, SJT, MES (2 J, I ~); Eight Keroplatus carbonarius Bosc.-OK Cave, 19-Apr-03, SJT, MES (2 J, 4 ~); LAHOMA: Delaware Co.: Junkyard Jasper Cave, ll-Apr-04, SJT, MES, J. Cave, 26-Jul-2005, GG, MES (1 ~). Jacoby (3 J, 4 ~); Joy Cave, 15-Jun-04, Leia sp.-ARKANSAS: Marion Co.: SJT, MES (I J); King Cave, 28-Mar-03, Toney Bend Mine #2, 24-May-04, SJT, V. Block (1 J); McCann Cave, 12 MES, CJB (1 J, 1 ~). MISSOURI: May-04, SJT, MES (1 J); small cave near Pulaski Co.: FLW: cave no. 23PU248, Lohraff Cave, 28-Mar-03, SJT, MES, V. 23-0ct-03, SJT, MES (1 ~); New Arch Block (2 ~); Turtle Trap Cave, 6-Sep-04, Cave, 5-Sep-04, SJT, MES, C. Me1hart MES, C. Melhart (2 ~ ); Wolf Den Cave, (1 ~). 16-Jun-04, SJT, MES (3 J, 3 ~). OKLA The Missouri specimens of Leia were HOMA: Delaware Co.: Surprise Cave, 27 taken from bedrock ceilings of caves. Jul-05, GG, DF, W. Puckette (1 ~). Evenhuis (1986) reported that two unde Exechiopsis sp.-ARKANSAS: New scribed Leia males had been found in ton Co.: Sherfield Cave, 2-Jun-01, CB, L. Pigeon Cave on Hawaii's big island. Moritz (1 ~); Tom Watson's Bear Cave, Macrocera nobilis Johnson.-AR 26-Jan-02, MES, CB, M. Ross (1 J); KANSAS: Marion Co.: Forest Trail 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ridge Cave, 6-Jun-02, MES (1 6, 1 2), Cave, 25-0ct-03, SJT, MES (1 6, 4 2); 5-Jun-04, MES, E. Frank, J. Terry (2 6); Little Freeman Cave, 9-Apr-04, SJT, Morning Star Mine #5, 26-May-04, MES, J. Jacoby (1 6). MES, CJB (4 6); Overlook(ed) Cave, Mycomya sp.-MISSOURI: Pulaski 6-Jun-02, MES (1 2), 5-Jun-04, MES, E. Co.: FLW: Henshaw Cave, l8-Apr-03, Frank, J. Terry (3 2); Pretty Junkyard SJT, V. Block (1 2). Spring Cave, 6-Jun-02, MES (1 6); Neuratelia sp.-ARKANSAS: Marion Toney Bend Mine #2, 24-May-04, Co.: Forest Trail Ridge Cave, 5-Jun-04, MES, CJB (1 6). MISSOURI: Pulaski MES, E. Frank, J. Terry (1 2); Toney Co.: FLW: Davis Cave #2, l7-Jun-04, Bend Mine #2, 24-May-04, MES, CJB SJT, MES (1 2); Wolf Den Cave, 16 (1 2); Sharp Co.: Eckel Cave, l4-Dec-Ol, Jun-04, SJT, MES (1 6). GG, DF, B. Sasse (1 2). Macrocera nobilis is known as an McDaniel and Smith (1976) reported inhabitant of forests in Alberta, Ontario, finding Neuratelia sp. near an opening to New England, and North Carolina. It is the surface of Needles Cave, Izard also known to occur in caves in Arkan County, Arkansas. They classified it as sas, Kentucky, Missouri, Oklahoma, a trogloxene. Tennessee, and West Virginia (Peck and Pseudobrachypeza bulbosa (Johans Russell 1976; Vockeroth 1976; Gardner sen).-OKLAHOMA: Delaware Co.: 1986; Graening et al. 2003, 2006). Long's Cave, 26-Jul-05, GG, MES, S. Individual adults of cave populations Hensley, J. Pruett (1 2). are short-lived. They mate year-round at Rymosia sp.-ARKANSAS: Marion cave entrances and oviposit in the dark Co.: Reed Cave, 9-Mar-02, GG, S. zone of caves where larval development McGinnis (1 2); Morning Star Mine and pupation occur. Larvae are more #6, 26-May-04, MES, CJB (2 6, 3 2); often observed than adults. They con Saltpeter Cave, 6-Jun-04, MES, W. struct extensive webs which they can Baker (1 6); Toney Bend Mine #2, 24 travel in pursuit of captured insect prey, May-04 (2 6, 2 2); Newton Co.: mostly other Diptera and especially the Saltpeter Cave, l7-Mar-02, MES, CJB heleomyzid Amoebaleria defessa (Osten (1 6,2 2); Stone Co.: Bud Wallis Cave, Sacken). The larval stadia last 9-10 28-Apr-02, GG, MES, C.B, M. Coving months, and the pupal stage lasts about ton (4 2); Hammer Springs Cave, 17 2 weeks, but the life cycle is not seasonal. Mar-02, GG, CB (1 6). MISSOURI: Peck and Russell (1976) hypothesized Pulaski Co.: FLW: Big Freeman Cave, 9 that during the Wisconsinan glaciation, Jan-04, SJT, MES (3 6, 1 2); David's the species occurred throughout the Crawl Cave, l2-Apr-04, SJT, MES (1 2); forests of the southern-central and east Deadman Cave, l6-Apr-03, SJT, V. ern United States. With retreat of the Block (1 2); Jasper Cave, ll-Apr-04, glaciers, this and other species that are SJT, MES, J. Jacoby (2 2); Phreatic adapted to cool, moist conditions found Cave, 24-0ct-03, SJT, MES (1 6, 1 2); refuge in caves in warming southern Ramsey Cave, 9-Jan-04, SJT, MES (2 6). lowland areas. All specimens reported OKLAHOMA: Cherokee Co.: Crystal here were adults collected in May and Cave, 29-Jan-02 (1 2); Delaware Co.: June by hand in entrance, twilight, and Rock Quarry Cave, 25-Jul-05, GG, MES dark zones. (26,12). Mycetophila sp.-ARKANSAS: Stone Peck and Lewis (1978) found Rymosia Co.: Blanchard Springs Caverns, 27 sp. in three southern Illinois caves and Apr-02, GG, MES, K. Tinkle (1 6). characterized it as a trogloxene, but they MISSOURI: Pulaski Co.: FLW: Breeden did not record collection dates. Reeves et VOLUME Ill, NUMBER 2 341 al. (2000) collected Rymosia sp. from a Cave, 25-Jul-04, CJB (I 2); Toney Bend Georgia cave in September and classified Mine #2, 27-May-04, MES, CJB (1 2); it as a possible troglophile. Moseley Newton Co.: Toney Barnes Cave, 16-Sep (2007) characterized Rymosia sp. as a 04, MES, CJB (1 d'); Tom Barnes Cave, habitual trogloxene wintering on cave ll-Dec-04, N. Youngsteadt, J. Young walls in the Canadian Maritime Prov steadt (2 2); Tom Barnes Cave, 18-Feb inces. The Ozark cave specimens, both 05, N. Youngsteadt, J. Youngsteadt (l d'); males and females, were collected in Walnut Cave, 29-Dec-04, MES, CJB (I January, March through July, and Oc 2); Searcy Co.: Hurricane River Cave, 14 tober, suggesting that Rymosia here is in Apr-02, MES, CB (1 2); 15-Jan-06, W. caves throughout the year. They were Baker (1 d'); Stone Co.: Blanchard collected by hand from entrance, twi Springs Caverns, 27-Apr-02, GG, MES, light, and dark zones on bedrock ceiling K. Tinkle (~102 2); Gustafson Cave, 6 and walls. It is perhaps a troglophile. Oct-02, GG, S. McGinnis, CB (I 2); Hell The mycetophilids E. umbratica and Creek Cave, 16-Sep-00, GG, MES (1 2); Rymosia sp. are among the most com Saltpeter Cave, 31-Mar-02, GG, DF, CB mon cavernicolous dipterans in the (1 2). MISSOURI: Pulaski Co.: FLW: Ozarks. Similarly, Kjaerandsen (1993) Andy's Cave #1, II-Apr-04, SJT, J. found that the mycetophilids Exechia Jacoby (1 2); Andy's Cave #2, ll-Apr exigua Lundstroem, E. festiva Winnertz, 04, SIT, J. Jacoby (4 2); Brooks Cave, 18 Exechiopsis fumbriata (Lundstroem), E. Jun-04, MES (5 2); Chuck's Virgin Cave, subulata (Winnertz), Pseudexechia auri 20-Apr-03, SJT, MES (3 d', II 2); vernica Chandler, and Rymosia fasciata David's Cave, 28-Mar-03, SIT, V. Block (Meigen) accounted for 95% of the (1 2); Davis Cave #1, 27-Mar-03, SJT, V. 20,000 Diptera specimens collected from Block (1 d', 2 2); Davis Cave #2, 17-Jun mines and caves in southern Norway. 04, SJT, MES (1 d'); Eight Cave, 24-0ct Trichonta sp.-MISSOURI: Pulaski 03, SIT, MES (2 d'); Folly Cave, 9-Apr Co.: FLW: Paleospring Cave, 17-Apr 04, SJT, MES, J. Jacoby (2 d'); Jasper 03, SJT, MES, V. Block (1 d'). Cave, II-Apr-04, SJT, MES, J. Jacoby (1 This specimen was taken from Neotoma d'); Joint East Cave, 10-Apr-04, SIT, J. floridana (Ord) scat in a packrat midden. Jacoby (I 2); Joy Cave, 18-Jun-04, MES (I 2); Kilman Cave, 25-Mar-03, SIT, V. Sciaridae Block (2 2); King Cave, 28-Mar-03, SIT, Sciaridae.-ARKANSAS: Benton Co.: V. Block (4 2); Little Freeman Cave, 9 Cave Springs Cave, 6-Mar-02, GG, MES, Apr-04, SJT, MES, J. Jacoby (1 2); B. Wagner (2 2); Pigeon Roost Cave, 28 Martin Cave, 29-Mar-03, MES, SJT, V. Jun-02, GG, B. Sasse (1 d'); Rainy Day Block (1 d', 17 2), IO-Jan-04, SJT, MES Cave, 17-Mar-06, MES, C. Slay (I 2); (4 2), 14-May-04, SIT, MES, K. Lohraff Tom Danforth Cave, 28-Jan-06, W. Baker (1 2), 6-Sep-04, MES, C. Melhart (I 2); (2 2); War Eagle Cavern, ll-May-Ol, A. Phreatic Cave, 24-0ct-03, SJT, MES (2 d'); Brown, C. Brown (1 2); Marion Co.: Ramsey Cave, 9-Jan-04, SJT, MES (1 d'); Forest Trail Ridge Cave, 5-Jun-04, MES, Saltpeter Cave, 9-Jan-04, SIT, MES (1 d', E. Frank, J. Terry (1 2); 8-Jun-04, MES, 37 2); Wilson Cave, 8-Jan-04, SJT, MES CJB (1 2); Morning Star Mine #5, 29 (5 2); Wolf Den Cave, 18-Jun-04, MES May-04, MES, CJB (2 2); Morning Star (1 2). OKLAHOMA: Adair Co.: Dun Mine #6, 29-May-04, MES, CJB (2 2); can Field Cave, 29-Apr-06, S. Hensley, Overlook(ed) Cave, 6-Jun-02, MES (2 2); W. Baker (1 2); Cherokee Co.: Blue 5-Jun-04, MES, E. Frank, J. Terry (I d', I Moon Cave, 29-Jan-02, GG, DF, S. 2); 8-Jun-04, MES, CJB (13 2); Summer Hensley (1 2). 342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Sciaridae, the dark-winged fungus Porricondyla sp.-ARKANSAS: Mar gnats, are among the most commonly ion Co.: Morning Star Mine #5, 26 collected Ozark cave Diptera. Unfortu May-04, MES, CJB (l ------VOLUME 111, NUMBER 2 343 Davis Cave #3, 23-0ct-2003, SJT, MES (Linnaeus), P. satchelli, Psychoda um (1 6); Wolf Den Cave, 16-Jun-2004, bracola Quate, and an unidentified Psy SJT, MES (1 6, 1 S2). choda species in Ontario caves in May, Psychodinae.-ARKANSAS: Marion June, and August. Gardner (1986) re Co.: Morning Star Mine #7, 7-Jun-04, ported unidentified species of Pericoma MES (1 S2); Overlook(ed) Cave, 8-Jun and Psychoda from several Missouri 04, MES, CJB (1 S2); Toney Bend Mine caves and classified them as trogloxenes. #2, 27-May-04, MES, CJB (1 S2); Searcy We suggest that at least P. satchelli could Co.: Square Cave, 25-Jul-04, MES, CB be considered a troglophile. (1 S2). MISSOURI: Pulaski Co.: FLW: Breeden Cave, 25-0ct-03, SJT, MES (1 Trichoceridae 6), Wolf Den Cave, 18-Jun-04, MES Trichocera brevicornis Alexander. (2 S2). ARKANSAS: Madison Co.: Whippoor Both sexes of psychodine psychodids will Cave, 29-Sep-02, GG, S. McGinnis were collected from Ozark caves during (1 6); Washington Co.: Joyce Cemetery all seasons of the year, indicating that Cave, I-Mar-04, MES (1 6), 26-Feb-05, they are probably true troglophiles. M. Ross (2 6). MISSOURI: Pulaski Specimens were taken by hand or in Co.: FLW: Breeden Cave, 25-0ct-03, pitfall traps in the entrance or dark zone SJT, MES (1 6). from scat on bedrock floors or from Trichocera sp. S2 (not identifiable to bedrock ceiling at air temperatures rang species level by morphological means). ing from 11.9 to 25.2°C and relative ARKANSAS: Marion Co.: Reed Cave, humilities ranging from 86.7 to 92.8%. 15-Nov-Ol, GG, T. Snell, P. Shurgar (1 Psychoda satchelli was the most com S2); Searcy Co.: USFS cave on Road monly encountered species and is the No. 91030A, 12-Dec-01 (1 S2); Stone most common species of Psychoda in Co.: Gunner Cave, 27-Jan-Ol, MES, America north of Mexico (Quate 1955). CB, J. Zawislak (1 S2); Washington Co.: Adult Psychodidae, which are usually Jones Cemetery Cave, I-Mar-04, MES (4 nocturnal, are often found in moist, S2); Joyce Cemetery Cave, 26-Feb-05, M. shaded habitats. Larvae of most species Ross (1 S2). OKLAHOMA: Cherokee are saprophagous, consuming decaying Co.: Crystal Cave, 29-Jan-02, GG, DF, organic matter. Larvae of Psychodinae S. Hensley (1 S2). live in various moist or subaquatic Winter crane flies of the genus Tricho habitats. Larvae of P. satchelli have been cera are often found on sunny days in found in compost (Quate 1960). Larvae autumn, winter, and spring. Even on of Psychoda reevesi Quate were collected cold winter days when temperatures are in Georgia caves using sterilized human above freezing, large swarms of males excrement as bait, and adults were reared may be seen dancing above the snow. from the larvae (Quate 2000). Larvae of Females are rarely found in swarms, Psychoda pussilla Tonnoir were collected although our results show they are more on chicken liver bait in Georgia caves, readily collected in Ozark caves than are and adults were reared in the laboratory. males. Trichocera brevicornis apparently Both P. reevesi and P. pussilla were has only one generation per year, with classified as troglophiles by Reeves et adult emergence commencing in Septem al. (2000). Adult Psychoda sp. have been ber. This species was previously recorded found associated with dung in caves in from Vermont, Maryland, North Caro southern Canada. Moseley (2007) classi lina, South Carolina, Georgia, Florida, fied them as habitual trogloxenes. Peck and Kansas. Specimens were collected (1988) reported Psychoda phalaenoides from October to February. Our cave 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimens were collected in Arkansas, SJT, MES, V. Block (l ~); Phreatic Missouri, and Oklahoma from Septem Cave, 24-0ct-03, SJT, MES (1 ~); Profitt ber to March. Females were found in Cave, 24-Mar-03, SJT, V. Block (2 ~); caves during the same season as when Wilson Cave, 8-Jan-04, SJT, MES (8 ~). males are flying (Pratt 1992, Pratt and OKLAHOMA: Delaware Co.: Junkyard Pratt 1984). They are probably utilizing Cave, 26-Jul-05, GG, MES (l ~); Rock the caves as shelter and are assumed to Quarry Cave, 25-Jul-05, GG, MES (1 ~). be trogloxenes. Culex pipiens Linnaeus or C quinque fasciatus Say.-ARKANSAS: Benton Culicidae Co.: Rainy Day Cave, l7-Mar-06, MES, Anopheles punctipennis (Say).-AR C. Slay (2 ~); Newton Co.: Pentrance KANSAS: Izard Co.: Needles Cave, 2 Cave, 4-Mar-06, W. Baker (5 ~). MIS Feb-03, GG, MES, D. Kampwerth, E. SOURI: Pulaski Co.: FLW: Camp Vista Corfey (l ~); Newton Co.: Jaw Bone Cave, 13-May-04, SJT, MES (l ~); small Cave, l6-Sep-04, MES, CJB (4 ~); Willis cave near LohraffCave, 28-Mar-03, MES, Cave, l5-0ct-05, N. Youngsteadt, J. SJT, V. Block (4 ~); McCann Cave, 12 Youngsteadt (l ~). MISSOURI: Pulaski May-04, SJT, MES (1 ~). Co.: FLW: Big Freeman Cave, 9-Jan-04, Uranotaenia sapphirina (Osten Sacken). SJT, MES (4 ~); Breeden Cave, 25-0ct MISSOURI: Pulaski Co.: FLW: 23PU248, 03, SJT, MES (3 ~); Eight Cave, 24-0ct 23-0ct-03, SJT, MES (2 ~); small cave near 03, SJT, MES (l ~); Flat Roof Cave, Lohraff Cave, 28-Mar-03, MES, SJT, V. 10-May-04, SJT, MES (1 ~); Lower Block (2 ~); Phreatic Cave, 24-0ct-03, SJT, Hooten Cave, ll-May-04, SJT, MES, MES (3 ~). K. Lohraff (1 ~); New Arch Cave, 5 These records are in addition to those Sep-04, MES, C. Melhart, SJT (1 ~ ); reported by Barnes (2004). All mosqui Phreatic Cave, 24-0ct-03, SJT, MES (l0 toes reported from Ozark caves are adult ~); Ramsey Cave, 9-Jan-04, SJT, MES females, and most were collected during (4 ~); Wilson Cave, 8-Jan-04, SJT, MES late fall, winter, or early spring. All (l ~). reported species are thought to winter Anopheles quadrimaculatus Say.-MIS as fertilized females. They seek shelter SOURI: Pulaski Co.: FLW: 23PU248, from freezing temperatures in caves and 23-0ct-03, SJT, MES (1 ~). other moist, protected habitats. They are Culex erraticus (Dyar and Knab). properly classified as facultative troglox ARKANSAS: Izard Co.: Needles Cave, enes. Anopheles quadrimaculatus was not 2-Feb-03, GG, MES, D. Kampwerth, E. previously reported wintering in an Corfey (5 ~); Newton Co.: Hutchinson Ozark cave, although it has been collect Waterfall Cave, 29-Dec-04, MES, CJB, ed from caves elsewhere (Hess and C. Melhart (6 ~); Jaw Bone Cave, 16 Crowell 1949, Ives 1938, Peck 1988). In Sep-04, MES, CJB (3 ~); Tom Barnes the southern part of this species' range, Cave, ll-Dec-04, N. Youngsteadt, J. females usually winter near human hab Youngsteadt (l ~), l6-Sep-04, MES, itations in cellars, crawl spaces, or CJB (3 ~); Willis Cave, l5-0ct-05, N. domestic animal quarters in barns. In Youngsteadt, J. Youngsteadt (2 ~), 18 the north, they usually winter in more Nov-05 (7 ~). MISSOURI: Pulaski Co.: isolated areas, such a storm drains, FLW: 23PU248, 23-0ct-03, SJT, MES (2 caves, and hollow trees (Horsfall 1955). ~); 23PU309, l3-May-04, SJT, MES (l Culex erraticus and A. punctipennis are ~); Eight Cave, 24-0ct-03, SJT, MES (l the most common species found winter ~); Little Crevice Cave, 24-0ct-03, SJT, ing in Ozark caves during the course of MES (1 ~); Miller Cave, 28-Mar-03, this study. Several local species that are VOLUME Ill, NUMBER 2 345 thought to winter as fertilized females Reeves et al. (2000) found adults of L. have not been collected from Ozark genualis on the roof and walls of caves, including Culex peccator Dyar Yarborough Cave, Bartow County, and Knab, Culex restuans Theobald, Georgia on 7 September 1998. They Culex salinarius Coquillett, Culex tarsalis surmised that the flies were avoiding Coquillett, Culex territans Walker, and the hot summer day outside. The Ozark Culiseta inornata (Williston). records reported here are for all seasons of the year. In addition, there are six Empididae pinned specimens in the collection of the Platypalpus sp.-ARKANSAS: Bax University of Arkansas Arthropod Mu ter Co.: Bonanza Cave, 7-0ct-02, GG, S. seum. Two males and three females were McGinnis, D. Kampwerth (1 ~). collected at a cave entrance in Newton Tachypeza brachialis (Melander). County, Arkansas on 26 July 1970. One ARKANSAS: Newton Co.: Fitton Cave, male was collected at Roaring River, 13-May-OI, GG, CJB, C. Bitting, S. Missouri on 29 July 1941. No other McGinnis (1 6); Jaw Bone Cave, 16 biological data are available for this Sep-04, MES, CJB (l 6); Stone Co.: Bud species. However, Corpus (1986) reports Wallis Cave, 28-Apr-02, GG, MES, CB, that a related species, L. similis Aldrich, M. Covington (I 6). was collected from drainage culverts and Tachypeza sp.-ARKANSAS: New rock crevices near seeps and waterfalls, ton Co.: Fitton Cave, 13-May-OI, GG, usually from shaded, moist, vertical rock surfaces. Adults reared from larvae in CJB, C. Bitting, S. McGinnis (l 6); Jaw the laboratory lived only three days. Bone Cave, 16-Sep-04, MES, CJB (I ~). Larvae were found only on algal mats Drosophilidae growing on rock below a waterfall. Neurigonella sp.-ARKANSAS: Wash Drosophila melanogaster.-ARKAN ington Co.: Farmer's Cave, 9-Jul-01 SAS: Stone Co.: Blanchard Springs (1 ~). Caverns, 27-Apr-02, GG, MES, K. Tinkle (4 ~). Phoridae Drosophila sp.-ARKANSAS: Mar Mallochphora orphnephiloides (Mal ion Co.: Overlook(ed) Cave, 8-Jun-04, loch).-ARKANSAS: Marion Co.: Reed ~); MES, CJB (1 Square Cave, 25-Jul Cave, 15-Nov-OI, GG, T. Snell, P. Shurgar 04, MES, CB (l ~). MISSOURI: Pulaski (l ~). Co.: FLW: Henshaw Cave, 18-Apr-03, Megaselia cavernicola (Brues).-AR ~). SJT, V. Block (l KANSAS: Benton Co.: Rainy Day Cave, 17-Mar-06, MES, C. Slay (l ~); Dolichopodidae Boone Co.: Big Hole Cave, 10-Nov-01, Liancalus genualis Loew.-ARKAN MES, M. Covington, CB (l ~); Newton SAS: Madison Co.: Wagonshed Cave, Co.: Dear Buster Cave, 28-May-06, W. 24-Jan-01 (l 6); Wounded Knee Cave, Baker (1 ~); Stone Co.: Biology Cave, 27-May-01, GG, CB (1 6, I ~); Marion 17-Sep-00, DF, CB, S. Longing (1 6); Co.: Saltpeter Cave, 6-Jun-04, MES, W. Hammer Springs Cave, 17-Mar-02, GG, Baker (2 6, 2 ~); Newton Co.: Wolf CB (1 ~); Hell Creek Cave, 16-Sep-00, Creek Cave, II-Nov-OO, GG, CB, J. GG, MES (1 ~); Rowland Cave, 12-Jun Gunter (l 6, I ~). 01, GG, CB, J. Disler (l ~, I 6); Twin Liancalus sp.--ARKANSAS: Marion Cave, 28-0ct-05, N. Youngsteadt, J. Co.: Long Ear Mine, 25-May-04, MES, Youngsteadt (2 ~); Washington Co.: CJB (l ~). Jones Cemetery Cave, I-Mar-04, MES ------346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (4 S2). OKLAHOMA: Delaware Co.: Cave, 6-Sep-04, MES, C. Melhart (l J); Long's Cave, 26-Jul-05, GG, MES, S. Wolf Den Cave, 16-Jun-04, SJT, MES Hensley, J. Pruett (l S2); Rock Quarry (1 J); Kilman Cave, 25-Mar-03, SJT, V. Cave, 25-Jul-05, GG, MES (l S2). Block (1 J). OKLAHOMA: Delaware Megaselia cavernicola is commonly Co.: Rock Quarry Cave, 25-JuI-05, GG, found in caves in the eastern United MES (1 J). States, where both sexes occur through Oecothea sp.-ARKANSAS: Marion all seasons. It was previously reported Co.: Coon Cave, 6-Jun-02, MES (l S2); from caves in Independence, Marion, Toney Bend Mine #2, 24-May-04, MES, Newton, and Stone Counties, Arkansas CJB (1 S2); Newton Co.: Jaw Bone Cave, (McDaniel and Smith 1976, Welbourn I6-Sep-04, MES, CJB (1 S2); Toney 1980, Welbourn and Lindsley 1979). Barnes Cave, I6-Sep-04, MES, CJB (l Reeves and Disney (1999) reported M. S2); Willis Cave, I8-Nov-05, N. Young cavernicola larvae on bat carrion and steadt, J. Youngsteadt (1 S2); Stone Co.: human excrement in the field, and the Blanchard Springs, 27-Apr-02, GG, species was commonly collected in carri MES, K. Tinkle (1 S2); Hidden Spring on traps. Larvae consume carrion from Cave, 5-0ct-02, CB, T. Breland, C. the bottom up, and they burrow into Breland (l S2); Twin Cave, 28-0ct-05, mud to pupate. Although commonly N. Youngsteadt, J. Youngsteadt (1 S2); found in caves, surface populations also Washington Co.: Fincher Cave, 9-Jul-OI, have been found (Brues 1950). The GG, J. Gunter, A. Gunter (1 S2). MIS species is classified as a troglophile. A SOURI: Pulaski Co.: FLW: Brooks similar cavernicolous species, Megaselia Cave, I8-Jun-04, MES (1 S2); Cave spelunciphila Disney, was not identified 23PU3IO, I8-Jun-04 (l S2); Chuck's during the course of this study. Virgin Cave, 20-Apr-03, SJT, MES (1 S2); Davis Cave #2, I7-Jun-04, SJT, Heleomyzidae MES (1 S2); Deadman Cave, I9-Apr-03, Oecothea specus (Aldrich).-ARKAN SJT, V. Block (2 S2); Kilman Cave, 25 SAS: Marion Co.: Long Ear Mine, 25 Mar-03, SJT, V. Block (1 S2); New Arch May-04, MES, CJB (l J); Morning Star Cave, 5-Sep-04, MES, C. Melhart, SJT Mine #5, 26-May-04, MES, CJB (1 J); (1 S2); Wolf Den Cave, I6-Jun-04, SJT, Morning Star Mine #7, 4-Jun-04, MES, MES (2 S2). OKLAHOMA: Delaware E. Frank (l J); Pretty Junkyard Spring Co.: Surprise Cave, 27-Jul-05, GG, DF, Cave, 6-Jun-02, MES (1 J); Saltpeter W. Puckette (1 S2). Cave, 6-Jun-04, MES, W. Baker (2 J); Amoebaleria defessa (Osten Sacken). Square Cave, 22-Jul-04, MES, CJB (1 J); ARKANSAS: Benton Co.: Civil War Newton Co.: Jaw Bone Cave, I6-Sep-04, Cave, 29-0ct-00, MES, GG, A. Brown (2 MES, CJB (2 J); Toney Barnes Cave, J); Pigeon Roost Cave, 28-Jun-02, GG, I6-Sep-04, MES, CJB (1 J); Willis Cave, B. Sasse (l J); Rainy Day Cave, 17 15-0ct-05, N. Youngsteadt, J. Young Mar-06, MES, C. Slay (2 J); Carroll steadt (1 J); Stone Co.: Blanchard Co.: Cave on Miner's Rock Trail, 12 Springs Caverns, 27-Apr-02, GG, MES, Aug-OO, GG (1 J); Marion Co.: Back'o K. Tinkle (1 J); Hidden Spring Cave, 5 Beyond Cave, 23-Jun-OI, GG, CJB (1 J); Oct-02, CB, T. Breland, C. Breland Forest Trail Ridge Cave, 5-Jun-2004, (2 J). MISSOURI: Pulaski Co.: FLW: MES, E. Frank, J. Terry (3 J); Marble Big Freeman Cave, 9-Jan-04, SJT, MES Falls Cave, 7-Sep-OI, GG, MES (1 J); (1 J); Camp Vista Cave, 13-May-04, Morning Star Mine #5, 26-May-04. SJT, MES (l J); Deadman Cave,I9 MES, CJB (1 J); Overlook(ed) Cave, 8 Apr-03, SJT, V. Block (l J); Turtle Trap Jun-04, MES, CJB (2 J); Saltpeter Cave, VOLUME 111, NUMBER 2 347 24-May-04, MES (1 6), 6-Jun-04, MES, May-04, MES, CJB (1 S2); Pretty Junk W. Baker (1 6); Newton Co.: Fitton yard Spring Cave, 6-Jun-02, MES (1 S2); Cave, 13-May-OI, GG, CJB, C. Bitting, S. Reed Cave, I5-Nov-OI, GG, T. Snell, P. McGinnis (1 6); Toney Barnes Cave, 16 Shurgar (1 S2); Square Cave, 22-Jul-04, Sep-04, MES, CJB (2 6); Willis Cave, 15 MES, CJB (1 S2); Toney Bend Mine #2, Oct-05, N. Youngsteadt, J. Youngsteadt 24-May-04, MES, CJB (1 S2); Newton (1 6), I8-Nov-05, N. Youngsteadt, J. Co.: Jaw Bone Cave, I6-Sep-04, MES, Youngsteadt (1 6); Stone Co.: Blanchard CJB (1 S2); Saltpeter Cave, I7-Mar-02, Springs Caverns, 27-Apr-02, GG, MES, MES, M. Covington (1 S2); Sherfield K. Tinkle (1 6); Hammer Springs Cave, Cave, 6-Jun-OI, CB, L. Moritz (1 S2); I7-Mar-02, GG, CB (1 6); Hell Creek Sprite Cave, I6-Mar-02, MES, M. Cov Cave, I6-Sep-00, GG (2 6); Rowland ington (1 S2); Tom Watson's Bear Cave, Cave, I2-Jun-OI, GG, CB, J. Disler (1 6); 26-Jan-02, MES, CB, M. Ross (1 S2); Twin Cave, 28-0ct-05, N. Youngsteadt, J. Toney Barnes Cave, I6-Sep-04, MES, Youngsteadt (1 6); Von Wadding's Me CJB (1 S2); Tweet's Cave, 26-0ct-OI, morial Cave, I7-Aug-02, GG, R. Schroe MES, M. Covington (1 S2); Searcy Co.: der, D. Kampwerth, E. Corfey (1 6); Hurricane River Cave, I4-Apr-02, MES, Washington Co.: Snyder Cave, I6-Sep-OI, CB (1 S2); Sharp Co.: Eckel Cave, 14 MES, R. Honebrink, B. Potter (2 6). Dec-Ol, GG, DF, B. Sasse (1 S2); Stone MISSOURI: Pulaski Co.: FLW: Andy's Co.: Blanchard Springs Caverns, 27 Cave #2, II-Apr-04, SJT, MES, J. Jacoby Apr-02, GG, MES, K. Tinkle (1 S2); (1 6); Andy's Cave #1, ll-Apr-04, SJT, Bud Wallis Cave, 28-Apr-02, GG, MES, MES, J. Jacoby (1 6); Breeden Cave, 25 CB, M. Covington (1 S2); Janus Pit, 1 Oct-03, SJT, MES (1 6); Joy Cave, 15 Feb-03, GG, MES, D. Kampwerth, E. Jun-04, SJT, MES (1 S2); Turtle Trap Corfey (1 S2); Martin Hollow Cave, 14 Cave, 6-Sep-04, MES, C. Melhart (1 6); Oct-OO, GG, MES, M. Covington, CB, J. Wolf Den Cave, I6-Jun-04, SJT, MES (1 Gunter (1 S2); Saltpeter Cave, 3I-Mar 6). OKLAHOMA: Delaware Co.: Janu 02, GG, DF, CB (1 S2); Washington Co.: ary-Stansbury Cave, Summer 2003, DF (2 Fincher Cave, 9-Jul-OI, GG, J. Gunter, 6); Murray Co.: Little Crystal Cave, 2 A. Gunter (1 S2); Joyce Cemetery Cave, Aug-05, GG, DF, A. Harris, K. Thomas 26-Feb-05, MES (1 S2). MISSOURI: (1 6). Pulaski Co.: FLW: Andy's Cave #1, Amoebaleria sackeni Garrett.-AR ll-Apr-04, SJT, MES, J. Jacoby (6 S2); KANSAS: Newton Co.: cave no. NW4Ibb, Andy's Cave #2, II-Apr-04, SJT, MES, 4-Mar-06, W. Baker (1 6). MISSOURI: J. Jacoby (1 S2); Davis Cave #3, 23-0ct Pulaski Co.: FLW: Davis No.3 Cave, 23 03, SJT, MES (1 S2); Henshaw Cave, 18 Oct-03, SJT, MES (1 6); Ramsey Cave, 9 Apr-03, SJT, V. Block (1 S2); Lohraff Jan-04, SJT, MES (1 6); Wilson Cave, 8 Cave,.30-Mar-03, SJT, V. Block (1 S2); Jan-04, SIT, MES (1 6). OKLAHOMA: Wilson Cave, 8-Jan-04, SJT, MES (1 S2). Cherokee Co.: Crystal Cave, 29-Jan-02, OKLAHOMA: Delaware Co.: Peachtree GG, DF, S. Hensley (1 6). Cave, 27-Jul-05, GG, MES, W. Puckette Amoebaleria sp.-ARKANSAS: Ben (1 S2); Surprise Cave, 27-Jul-05, GG, DF, ton Co.: James Ditto Cave, 6-Dec-00, W. Puckette (1 S2). GG, CB (1 S2); Boone Co.: Big Hole Heleomyza brachypterna (Loew).-AR Cave, lO-Nov-OI, MES, M. Covington, KANSAS: Benton Co.: Congo Crawl, 4 CB (1 S2); Marion Co.: Elm Cave, 16 May-OI, MES, A. Brown (1 6); Stone Nov-OI, GG, B. Sasse (1 S2); Forest Trail Co.: Von Wadding's Memorial Cave, 17 Ridge Cave, 5-Jun-04, MES, E. Frank, J. Aug-02, GG, R. Schroeder, D. Kamp Terry (4 S2); Morning Star Mine #6, 26 werth, E. Corfey (1 S2). MISSOURI: 348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pulaski Co.: FLW: Joy Cave, 15-Jun-04, baited with fish meal, fermented syrup, SJT, MES (2 S2); Wolf Den Cave, 16-Jun and beef liver. Amoebaleria defessa and 04, SJT, MES (l S2). A. sackeni have been observed at tree sap Most of these collections were taken in wooded areas, but probably most by hand and occasionally from pitfall specimens have been taken deep within traps. Heleomyzids were taken mainly on caves where they occur throughout the bedrock ceilings and walls, but some year (Gill 1962). Busacca (1975) found collections were associated with rock, A. defessa, H brachypterna, H serrata clay, soil, and gravel floors of caves. (Linnaeus), and 0. specus most heavily Unidentified female specimens of Oe concentrated 40-50 m from the entrance cothea sp. were collected under rocks and of an Indiana cave in October, Novem on floors or on clay or soil floors. ber, and January. He examined bat Recorded temperatures ranged from guano, decaying organic matter, dead 12.4 to 15.9°C for air and 11.8 to bats, and meat bait for heleomyzid 14.1°C for soil, with humidity of 92.1 larvae but found none. Reeves and to 99.4%. Specimens of 0. specus were McCreadie (2001) found no heleomyzid recorded from under rock, on bedrock immature in hundreds of carrion baits walls, and on clay or gravel floors in the placed in two caves in Georgia. Larvae twilight and darks zones of caves. of the widespread North American and Recorded temperatures ranged from European species H brachypterna have 12.2 to 12.8°C for air and 11.1 to been found in bird nests in Great Britain 12.2°C for soil, with humidity of 91. 7 (Blair 1931). Collin (1939) reported to 94.0%. Arnoebalaria defessa and A. having seen specimens bred from the sackeni were recorded primarily from nest of a brown owl, from the debris of bedrock ceilings and walls in the twilight an owl's nest in a tree hollow, and from and dark zones of the caves. nests of kestrels, carrion crows, and Each of the species of heleomyzid flies rooks. Both male and female heleorny reported here was found previously in zids are present in Ozark caves through Ozark caves (Barnett 1970, Black 1971, out the year. Most A. sackeni specimens Peck and Lewis 1978, McDaniel et al. were taken in winter, and most H 1979, Welbourn and Lindsley 1979, brachypterna specimens were taken in Gardner 1986). Heleomyzid flies are late spring. frequently found on walls and ceilings of caves, and they are usually classified Sphaeroceridae as troglophiles or trogloxenes (Gill 1962, Leptocera caenosa (Rondani).~AR Holsinger and Culver 1988, Peck and KANSAS: Newton Co.: John Eddings Lewis 1978, Sutton 1993). We know little Cave, 19-0ct-Ol, MES, CJB (1 S2); Stone about their biology or why they enter Co.: Blanchard Springs Caverns, 27 caves in such great numbers. The few Apr-02, GG, MES, K. Tinkle (1 S2). heleomyzine species for which larval food OKLAHOMA: Delaware Co.: East Hol habits are known are thought to feed on low Cave, 26-Jul-05, GG, MES, S. dung, carrion, and other decaying mate Hensley, J. Pruett (1 6, 1 S2). rials. Oecothea specus (previously known Limomyza cavernicola Marshall.~ as Aecothea specus [International Com ARKANSAS: Stone Co.: Hammer mission on Zoological Nomenclature Springs Cave, 17-Mar-02, GG, CB (l 1995]) occurs throughout North America, S2). OKLAHOMA: Delaware Co.: Long's and it is common in collections from caves Cave, 26-Jul-05, GG, MES, S. Hensley, J. and mammal burrows (Garnett and Foote Pruett (1 6); Rock Quarry Cave, 25-Jul 1966). Adults have been collected at traps 05, GG, MES (1 6). VOLUME 111, NUMBER 2 349 Spelobia clunipes (Meigen).-OKLA cheese, and dung baits in a cave in HOMA: Delaware Co.: Rock Quarry Georgia. Limomy:a cavernicola was re Cave, 25-Jul-05, GG, MES (1 6). cently described from in or near caves in Spelobia sp. prob. clunipes (Meigen). Missouri, Kentucky, Oklahoma, and ARKANSAS: Washington Co.: Granny Colorado. It is regarded as troglophile Parker's Cave, l6-Jun-00, GG, S. Long (Marshall 1997). ing (2 S!). OKLAHOMA: Delaware Co.: Spelobia tenebrarum, known only from Long's Cave, 26-Jul-05, GG, MES, S. caves, is the most abundant of the Hensley, J. Pruett (3 S!). Nearctic cave sphaerocerids. It occurs Spelobia tenebrarum (Aldrich).-AR in the south-central United States, south KANSAS: Baxter Co.: Bonanza Cave, 7 of the southern limits of the Wisconsinan Oct-02, GG, D. Kampwerth, S. McGin glacial ice sheet. Like some other cave nis (1 6); Benton Co.: Pigeon Roost flies, it has reduced eyes and an elongat Cave, 28-Jun-02, GG, B. Sasse (1 6); ed antennal arista. It seems to have War Eagle Cavern, Il-May-Ol, A. reduced powers of flight, leaping rather Brown, C. Brown (1 S!); Boone Co.: than flying when disturbed. The species Big Hole Cave, 10-Nov-Ol, MES, M. is commonly associated with fecal mate Covington, CB (2 S!); Marion Co.: rial, especially bat guano, and decaying Marble Falls Cave, 7-Sep-Ol, GG, MES organic matter (Marshall and Peck 1984, (2 6); Pretty Junkyard Spring Cave, 5 1985; Marshall 1985). It is considered a Jun-02, MES (2 S!);Reed Cave, 9-Mar-02, true troglobite (Lewis et al. 2003, Mar GG, S. McGinnis (1 6); Stone Co.: shall and Peck 1984, Marshall and Blanchard Springs Caverns, 27-Apr-02, Fitzgerald 1997) or possibly a troglophile GG, MES, K. Tinkle (3 6 ,1S!); Hammer (Holsinger and Culver 1988). Marshall Springs Cave, 17-Mar-02, GG, CB (1 6); and Reeves (2003) reported adults and Rowland Cave, l2-Jun-Ol, GG, CB, J. immatures of the facultative cavernico Disler (1 6, 2 S!); Twin Cave, 28-0ct-05, lous species Spelobia pickeringi Marshall N. Youngsteadt, J. Youngsteadt (4 S!). to be associated with bat guano in caves MISSOURI: Jasper Co.: Sarcoxie Cave, in South Carolina. Adults were also 8-Feb-02, GG, T. Snell, J. Foresman, J. collected in epigean Malaise and mush Hinsey (1 S!). OKLAHOMA: Cherokee room-baited traps in Georgia and Mary Co.: Crystal Cave, 29-Jan-02, GG, DF, S. land. Hensley (3 6, 2 S!); Delaware Co.: East According to Marshall and Peck Hollow Cave, 26-Jul-05, GG, MES, (1984), the entire genus Telomerina is S. Hensley, J. Pruett (1 S!); January characterized by apparently hypogean Stansbury Cave, Aug. 2002, DF (1 S!). characters, such as reduced eye size. It Telomerina jlavipes (Meigen).-AR is not surprising that some species are KANSAS: Newton Co.: John Eddings found in caves. They characterized T Cave, 19-0ct-Ol, MES, CJB (1 6). jlavipes as a cosmopolitan, necropha Sphaerocerids have been reared from gous, synanthropic species that is not dead bats and meat bait placed in caves predominantly troglophilic. It also has (Busacca 1975). Leptocera caenosa is been collected from caves in Europe, associated primarily with plant detritus Africa, South America, Australia, and in caves and is opportunistically associ New Zealand and from Devil's Den ated with many cavelike environments. It Cave, Washington County, Arkansas. also invades cellars, .mines, urinals, ab It is commonly collected in carrion attoirs, and septic tanks (Conn and traps, dung, decaying fungi, and mam Marshall 1991). Reeves et al. (2000) mal burrows (Marshall and Rohacek found it in association with carrion, 1984). 350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Anthomyiidae saprophagous. Cynomya cadaverina is Pegomya sp.-MISSOURI: Pulaski attracted to excrement and putrid carri Co.: FLW: David's Crawl Cave, 12 on. Adults are also found in kitchens, pa~tries, and cellars of homes in early Apr-04, SJT, MES (1 C;»; Flat Roof spnng and late fall. The larvae have been Cave, 1O-May-04, SJT, MES (1 C;»; Joint implicated in cases of subdermal myiasis East Cave, 1O-Apr-04, SJT, MES (1 C;»; Kilman Cave, 25-Mar-03, SJT, V. Block in warm-blooded animals (Hall 1948). (1 6); Little Crevice Cave, 24-0ct-03, ACKNOWLEDGMENTS SJT, MES (1 C;». OKLAHOMA: Dela ware Co.: Long's Cave, 26-Jul-05, GG, Dr. Raymond J. Gagne, Systematic Entomology Laboratory, Agricultural MES, S. Hensley, J. Pruett (1 C;». Missouri specimens of Pegomya were Research Service, United States Depart found on bedrock ceilings or on packrat ment of Agriculture, identified the Ceci feces in habitats characterized subjective domyiidae. Dr. Steven A. Marshall, De par~me~t ly as having dry to normal moisture of Environmental Biology, University of Guelph assisted with the condition. Larvae of Pegomya are typi identifications of Sphaeroceridae. We cally leaf and stem miners. Pegomya thank the individuals listed in the collec affinis Stein and P. dissecta Meigen have tion records for their assistance during been found in caves in Ontario during fieldwork; without their help, this project summer and fall (Peck 1988). would not have been accomplished. Spe Bibionidae cial thanks go to G. Graening and D. Fenolio for their contribution ofArkansas Bibio albipennis Say.-ARKANSAS: and Oklahoma data and fieldwork. Help Stone Co.: Blanchard Springs Caverns, ful reviews of an earlier draft of this 27-Apr-02, GG, MES, K. Tinkle, (1 C;». manuscript were provided by W. K. Calliphoridae Reeves and J. J. Lewis. Funding for this study was provided by the following Calliphora livida Hall.-ARKANSAS: sources: Arkansas Game & Fish Commis Newton Co.: Squirrel Pit, 2-Feb-02, GG, sion, Arkansas Natural Heritage Com MES (1 C;». mission, Fort Leonard Wood Military Calliphora vicina Robineau-Desvoidy. Installation, Illinois Natural History Sur ARKANSAS: Stone Co.: Blanchard vey, Oklahoma Biological Survey, The Springs Caverns, 27-Apr-02, GG, MES, K. Nature Conservancy (Arkansas Field Tinkle (2 6, 1 C;». MISSOURI: Pulaski Co.: Office, Oklahoma Field Office), Universi FLW: Joy Cave, 18-Jun-04, SIT, MES (l C;». ty of Arkansas, U.S. Fish & Wildlife Cynomya cadaverina Robineau-Des Service (Arkansas, Oklahoma), U.S. For voidy.-OKLAHOMA: Delaware Co.: est Service (Ozark National Forest), and Twin Cave, 5-Mar-Ol, GG, MES, S. U.S. National Park Service (Buffalo Feirer, DF (l 6). National River). Peck and Lewis (1977) found C. vicina in a cave in Illinois. Reeves and Disney LITERATURE CITED (1999) reported finding Calliphora sp. Alexander, C. P. 1919. The crane-flies of New larvae competing with M. cavernicola York. Part I. Distribution and taxonomy of larvae on carrion in caves. Reeves et al. the adult flies. Cornell University Agricultural (2000) reported C. vicina and Calliphora Experiment Station Memoir 25: 771~993. -. 1920. The crane-flies of New York. Part II. vomitoria (Linnaeus) associated with Biology and phylogeny. Cornell University chicken liver bait placed in caves in Agricultural Experiment Station Memoir 38: Georgia. All species of Calliphora are 699~1133. ------._----_. VOLUME III, NUMBER 2 351 1966 (reprint of 1942 edition, with Gardner, J. E. 1986. Invertebrate Fauna from addenda). Family Tipulidae, pp. 196-486. In Missouri Caves and Springs. Missouri Depart Guide to the Insects of Connecticut. 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