Zootaxa 1357: 1–19 (2006) ISSN 1175-5326 (Print Edition) ZOOTAXA 1357 Copyright © 2006 Magnolia Press ISSN 1175-5334 (Online Edition)

Spongillaflies (Neuroptera: Sisyridae) of North America with a key to the larvae and adults

DAVID E. BOWLES National Park Service, Heartland Network Inventory and Monitoring Program, c/o Department of Biology, Missouri State University, 901 South National Avenue, Springfield MO 65897, U.S.A. E-mail: [email protected]

Abstract

The distributions of the two genera (Climacia and Sisyra) and six species of spongillaflies known to occur in Canada, Mexico and the United States are reviewed and summarized. Several new and significant distribution records are presented for states or provinces in these countries. Species diagnoses and a revised, illustrated key to the third instar larvae and adults of all species are presented.

Key words: Spongillaflies, Neuroptera, Sisyridae, Climacia, Sisyra, taxonomy, distribution

Introduction

Two genera and six species of Sisyridae, or spongillaflies, are known to occur in North America (Penny et al. 1997). Parfin and Gurney (1956) published a treatment of the spongillaflies of the western hemisphere that addressed these species and their known distributions at that time. Subsequently, several state or regional studies of spongillaflies in the United States have been published (Isom 1968, Poirrier 1969, Throne 1971, Poirrier & Arceneaux 1972, Brown 1974, Grigarick 1975, White 1976, Huggins 1980, Poirrier & Holzenthal 1980, Lago 1981, Stoaks et al. 1983, Lawson & McCafferty 1984, Clark 1985). Additionally, Penny et al. (1997) addressed the general distribution of sisyrids occurring in North America exclusive of Mexico, but their treatment did not include all of the previously published distribution records for the respective species. Oswald et al. (2002) listed the states of Mexico in which spongillaflies had been collected, but they did not provide specific locality data. Because of these various studies, spongillafly distributions are generally well documented from eastern North America north of Mexico. By comparison, the distributions of sisyrids in the western United States, Canada, and

Accepted by S. Winterton: 11 Oct. 2006; published: 13 Nov. 2006 1 ZOOTAXA Mexico are not well known (Parfin & Gurney 1956, Brown 1974, Penny et al. 1997, 1357 Oswald et al. 2002). Larvae of sisyrid species are poorly known (Pupedis 1980). Additionally, larvae are seldom represented in collections, and, if present, they are often poorly preserved, making identification difficult. The relative rarity of the larvae is further confounded by there being few strong characters to distinguish among the species. However, Parfin and Gurney (1956) were able to provide taxonomic keys to the third instar larvae and adults of most of the species found in North America. Later studies, including those of Poirrier and Arcenaux (1972), Pupedis (1980) and Lago (1981), have provided additional characters for separating some of these species. The accumulated data of these works makes it possible to develop a comprehensive key to both the larvae and adults of all sisyrid species found in North America. The purpose of this paper is to present a revised key to the larvae and adults and summarize new and existing distribution records for the six known species of North American sisyrid species.

Methods

In addition to my own collection (DEB), and previously published records, I have examined and recorded data from more than 1,200 spongillaflies in the following museum collections: Academy of Natural Sciences of Philadelphia (ANSP); American Museum of Natural History (AMNH); Brigham Young University Collection (BYUC); Essig Museum of Entomology, University of California, Berkeley (EMEC); Florida State Collection of Arthropods (FSCA); Illinois Natural History Collection (INHS); Michigan State University Entomology Collection (MSUC); Mississippi Entomological Museum (MEM); Oklahoma Natural History Survey (ONHS); Snow Entomological Museum, University of Kansas (SEMC); Texas A&M University Insect Collection (TAMU); National Museum of Natural History, Washington, DC (USNM); University of Arkansas Arthropod Museum (UAAM); University of Missouri, Columbia (UMC); University of North Texas (UNT); Utah State University Entomological Museum (EMUS); and Bill Shepard Collection (BSC). Terminalia of adult specimens were removed using forceps under a dissecting stereomicroscope. Terminalia were then soaked in 10% potassium hydroxide for approximately 24 hours and then rinsed in 70% isopropyl alcohol to remove any remaining residue. Cleared terminalia were then examined in glycerin using a dissecting stereomicroscope and stored in glycerin-filled genitalia vials attached to the specimen pins or placed in the vial of alcohol as appropriate. Right forewings were removed and photographed with a digital camera fitted to a dissecting stereomicrocope. Whole larvae were also cleared for examination using the same procedure as above. Below I summarize the previously published distributions for spongillaflies for states and provinces in North America, and these records are supplemented with county and

2 © 2006 Magnolia Press BOWLES parish data. Only first published records are presented. New state and province records are ZOOTAXA given with complete specimen label data. County and parish information inferred from 1357 incomplete published records and label data (i.e., where state and other identifying collection information was provided) is placed within brackets. A complete listing of label data for all specimens examined for this study is available from the author upon request. Certain illustrations were redrawn and modified as necessary from their original published sources and these are noted where appropriate. The terminology used herein follows Flint (1998).

Keys to third instar larvae and adult Sisyridae of North America

The following keys are adapted and modified from information in Parfin and Gurney (1956), Poirrier and Arceneaux (1972), Lago (1981), Pupedis (1980) and Weissmar and Waringer (1994). Third instar larvae of S. nigra and S. vicaria usually cannot be reliably separated. However, Pupedis (1980) presented some meristic characters related to the spacing of the paired dorsal thoracic tubercles that may be useful in separating high quality, recently captured specimens.

Larvae 1 Eighth abdominal tergite with a pair of setae; 6th abdominal tergite with middle tubercle longest and bearing a terminal seta that is longer and thinner than that of other tubercles (Fig. 1); small acute projections present on the tubercle adjacent to the bases of thoracic setae...... Climacia, 2 - Eighth abdominal tergite without setae; 6th abdominal tergite with lateral-most tubercle longest and bearing a terminal seta that is much longer and thinner than that of other tubercles (Fig. 2); thoracic tubercles without small acute projections adjacent to the bases of their setae ...... Sisyra, 4 2 Meso- and metathoracic sclerites with five setae, four set on prominent tubercles, one minute and sessile (Fig. 3) ...... 3 - Meso- and metathoracic sclerites with four setae, three set on prominent tubercles, one minute and sessile (Fig. 4) ...... Climacia californica Chandler 3 Sixth abdominal tergite with median tubercle of dorsal plates having seta extending beyond the apex of 10th abdominal segment, and at least 3 times the length of the seta of the lateral tubercle; lateral tubercle with prominent blade-like projections at apex (Fig. 5)...... Climacia areolaris (Hagen) - Sixth abdominal tergite with median tubercle of dorsal plates having seta not extending to 10th abdominal segment, and less than 3 times the length of the seta of the lateral tubercle; lateral tubercle with small, never blade-like projections at apex (Fig. 6) ...... Climacia chapini Parfin and Gurney 4 First abdominal gills with distinct basal hook-like projections (Fig. 7). Fifth lateral

SPONGILLAFLIES © 2006 Magnolia Press 3 ZOOTAXA abdominal tubercle with three setae arranged on two projections (Fig. 9) ...... 1357 ...... Sisyra nigra (Retzius), Sisyra vicaria (Walker) - First abdominal gills without basal hook-like projection (Fig. 8). Fifth lateral abdominal tubercle with two sessile setae and one seta on a projection (Fig. 10)...... Sisyra apicalis Banks

Adults 1. Forewings with a series of outer gradate cross-veins (Fig. 11); radial sector (Rs) with one branch; distinctly maculate with dark brown, reddish and yellow markings or hyaline with only radiomedial pigment streaks. Antennae brown to blackish, occasionally slightly pale apically. Terminal segment of labial palp less than twice as broad as long and scimitar-shaped (Fig. 13)...... Climacia, 2 - Forewings without a series of outer gradate cross-veins (Fig. 12); Rs with two branches; uniformly dark brown and with or without darker brown pigment streaks. Antennae distinctly bicolored, brown and yellow. Terminal segment of labial palp twice as broad as long and hatchet-shaped (Fig. 14)...... Sisyra, 4 2. Forewings mostly hyaline, but with a narrow, conspicuous radiomedial pigment streak (Fig. 15) and with some intervenal streaking present (some specimens have additional pigmentation located posteriad to the radiomedial pigmentation). Terminalia as in Figs. 18–20...... Climacia chapini Parfin and Gurney - Forewings distinctly maculate and with prominent intervenational streaking. Male and female terminalia not as above...... 3 3. Forewings usually 5 mm or longer; pale colored area of pterostigma covering six or more veins (Fig. 16). Antennae dark brown to black. Postocciput dark brown and distinctly darker than rest of head. Terminalia as in Figs. 21–23 ...... Climacia areolaris (Hagen) - Forewings usually 4 mm long or less; pale colored area of pterostigma covering only four veins (Fig. 17). Antennae pale to dark brown. Postocciput pale brown and not distinct in color from rest of head. Terminalia as in Figs. 24–26 ...... Climacia californica Chandler 4. Length 5 mm or less. Forewings with prominent intervenational streaking. Antennae with basal half and apical 4 or 5 segments dark brown, intervening segments pale yellow. Terminalia as in Figs. 27–29 ...... Sisyra apicalis Banks - Length 5 mm or greater. Forewing usually uniformly brown, or with intervenational streaking poorly defined. Antennae uniformly dark in color or with only a few basal segments dark and remaining segments paler in color. Terminalia not as above...... 5 5. Forewing with R2 vein usually forked basal to the junction of Sc and R1 veins. Anten- nae with only 2–3 basal segments dark brown with remaining segments yellow, rarely basal third dark and apical two thirds paler in color. Head with vertex usually pale brown to yellow. Terminalia as in Figs. 30–32 ...... Sisyra vicaria (Walker)

4 © 2006 Magnolia Press BOWLES - Forewing with R2 vein usually forked equal or apical to the junction of Sc and R1 ZOOTAXA veins. Antennae, head and thorax dark brown to black. Terminalia as in Figs 33–35.... 1357 ...... Sisyra nigra (Retzius)

Diagnoses and distributions

Climacia areolaris (Hagen) Figures 3, 5, 16, 21–23

This species is most similar in appearance to C. californica. The forewings are usually 5 mm or longer and the pale colored area of pterostigma covers six or more veins (Fig. 16). The antennae are dark brown to black, and the postocciput is dark brown and distinctly darker than the rest of the head. Male terminalia have the ventral lobe of the ectoproct narrow and finger-like and located posterior to the dorsal portion (Fig. 21). The gonarcus complex in dorsal view has mildly sinuate internal spines that are broadest anteriorly and gradually tapering posteriorly to a fine point (Fig. 22). Female terminalia have tergite 9 as broad as long in lateral view (Fig. 23). The apex of the gonapophyses curve posteriad at an angle of approximately 80o. Climacia areolaris is widespread in the United States and Canada east of the Rocky Mountains, but I have not collected nor examined any material from Mexico. Brown (1974) stated that the record of C. areolaris from Lago de Xochimilco in central Mexico reported by Navás (1928) was probably inaccurate and that those specimens were probably either Climacia chapini or an undescribed species. However, Brown (1974) indicated that the Climacia larvae he collected in nearby Durango State, Mexico were apparently undescribed and possibly represented the same species reported by Navás (1928). Brown (1974) also suggested that the Climacia cocoons he collected in sponge in southern Baja California, Mexico likely represented an undescribed species, and Pupedis (1980) arrived at a similar conclusion. Oswald et al. (2002) reported C. areolaris from San Luis Potosi and Tamaulipas, Mexico based on information originally provided by this author. However, I have reexamined those specimens and determined that the former is C. chapini and the latter C. californica. The known distribution of C. areolaris in the United States, and my examination of material from Mexico, leaves me to conclude that this species likely does not occur in Mexico and that previously published records of its occurrence there are dubious. In this study, new distributional records for C. areolaris are reported from Nova Scotia, Canada, and Kentucky, South Carolina, and West Virginia in the United States. The flight period of C. areolaris ranges from March through December. Adults of C. areolaris are occasionally co-collected with Sisyra vicaria throughout their respective ranges although they normally are more locally abundant than the latter species.

FIGURES 1–10. Structures of larval Sisyridae. 1–2. Abdominal tergite 6, tubercle group, left side, dorsal (redrawn from Pupedis 1980): 1. Climacia. 2. Sisyra. 3–4. Mesothoracic sclerite, right side, dorsal: Climacia spp: 3. C. areolaris and C. chapini. 4. C. californica, (redrawn from Pupedis 1980). 5–6. Abdominal segment 6, lateral turbercle (redrawn from Pupedis 1980): 5. C. areolaris. 6. C. chapini. 7–8. Abdominal segment 1, ventral gill: 7. S. nigra and S. vicaria (redrawn from Weissmair & Waringer 1994). 8. S. apicalis. 9–10. Abdominal segment 5, lateral tubercle (redrawn from Poirrier & Arcenaux 1972): 9. S. nigra and S. vicaria. 10. S. apicalis.

6 © 2006 Magnolia Press BOWLES Published and Supplemental State, County and Parish Records. CANADA. Ontario: ZOOTAXA (Carpenter 1940. Quebec: (Carpenter 1940). MEXICO: [Distrito Federal] (Navás 1928). 1357 San Luis Potosi: (Oswald et al. 2002). Tamaulipas: (Oswald et al. 2002). UNITED STATES. Alabama: county unknown (Parfin & Gurney 1956), [Lawrence, Limestone] (Isom 1968); Baldwin, Jefferson (MEM). Arkansas: county unknown (Parfin & Gurney 1956); Clark, Garland, Johnson (UAAM), Lafayette (UNT), Logan (UAAM), Ouachita (INHS), Perry, Pike (UAAM), Pope (UNT), Stone (INHS, UAAM) Logan (EMEC). Colorado: county unknown (Parfin & Gurney 1956). Connecticut: county unknown (Carpenter 1940), [Hartford, Middlesex] (Parfin & Gurney 1956), [Tolland] (Parfin & Gurney 1956). Delaware: county unknown (Parfin & Gurney 1956). Florida: county unknown (Carpenter 1940), Alachua, [Charlotte, Hendry, Palm Beach, Polk, Volusia] (Parfin & Gurney 1956); Baker, Dade, [Escambia], Highlands, Liberty, Levy, Marion, Oklaloosa, Putnam, Santa Rosa, Suwannee (BYUC), [Hendry], Wakulla (AMNH). Georgia: [Charlton] (Carpenter 1940), [Charlton, Crawford, Houston, Pierce, Richmond, Whitfield] (Parfin & Gurney 1956); [Lanier] (INHS). Illinois: county unknown (Carpenter 1940); [Carroll, Iroquois, Kankakee, Jersey, Saline, Stephenson] (Parfin & Gurney 1956), Champaign, Fayette, Gallatin, Johnson, McLean, Platt, Pope, Vermilion (INHS). Indiana: [Marshall] (Parfin & Gurney 1956); Monroe (FSCA). Kansas: [Riley] (Parfin & Gurney 1956); Chautauqua, Cherokee, Cowley, Crawford, Douglas, Neosho, Johnson, and Woodson (Huggins 1980). Louisiana: [Caddo, Rapides] (Parfin & Gurney 1956); Bossier, Concordia, Jefferson, St. Charles, St. Tammany, Washington (Poirrier 1969), East Feliciana (Poirrier & Arceneaux 1972); Baton Rouge (BYUC), Natchitoches (TAMU). Maine: county unknown (Carpenter 1940), [Hancock, Kennebec] (Parfin & Gurney 1956), [Penobscot] (Parfin & Gurney 1956). Maryland: county unknown (Carpenter 1940), [Worcester] (Parfin & Gurney 1956); Calvert (BYUC). Massachusetts: county unknown (Carpenter 1940) (AMNH), [Middlesex, Worcester] (Parfin & Gurney 1956). Michigan: [Emmet, Ingham] (Carpenter 1940), [Cheboygan, Houghton] (Parfin & Gurney 1956); Chippewa, Kalamazoo (MSU). Minnesota: [St. Louis] (Carpenter 1940), Houston, Lake, Washington (Parfin 1952); Cook, [Ramsey], [Wabasha] (Parfin & Gurney 1956). Mississippi: [Adams] (Carpenter 1940); Amite, Lincoln, Pearl River, Perry, Simpson, Stone (Poirrier & Holzenthal 1980); Franklin, Hancock, Jackson, Madison, Marshall, Quitman, Tishomingo (Lago 1981); Lauderdale (BYUC), Grenada, Pontotoc, Smith, Oktibbeha (MEM). Missouri: Taney (Froeschner 1947), [Taney] (Parfin & Gurney 1956); Boone (INHS, UMC), Pulaski, Shannon (UAAM), Wayne (UMC). New Hampshire: county unknown (Carpenter 1940). New Jersey: county unknown (Carpenter 1940), [Burlington] (Parfin & Gurney 1956). New Mexico: Eddy (Carpenter 1940). New York: county unknown (Carpenter 1940) [Columbia, Erie, Essex, Herkimer, Jefferson, Livingston, Montgomery, Queens, St. Lawrence, Warren, Washington], Westchester (Parfin & Gurney 1956); [Clinton] (INHS), [Columbia] (FSCA). North Carolina: [Moore] (Carpenter 1940); McDowell (INHS). North Dakota: Grand Forks (Stoaks et al. 1983). Ohio: [Ottawa] (Carpenter 1940); Ashtabula (INHS), Hocking (BYUC).

SPONGILLAFLIES © 2006 Magnolia Press 7 ZOOTAXA Oklahoma: county unknown (Carpenter 1940), Adair (BSC), Beckham, Grant, McCurtain 1357 (Parfin & Gurney 1956), Grayson (White 1976), Johnston (Brown 1974), LeFlore (EMEC), Marshall (Brown 1974), McCurtain (Parfin & Gurney 1956), Pushmataha (Parfin & Gurney 1956); Johnston (ONHS), Latimer (DEB), Marshall (INHS, UOBS, FSCA), Ouachita (INHS). Pennsylvania: county unknown (Carpenter 1940). Tennessee: county unknown (Isom 1968). Texas: Bosque (Carpenter 1940), Dallas, Goliad (Parfin & Gurney 1956), Sutton (Carpenter 1940); Anderson (TAMU); Bastrop (INHS, MEM), Bexar (DEB), Bosque (USNM), Brazos, Bowie, Burleson (TAMU), Dimmit (TAMU), [Goliad] (USNM); Hardin (USNM, TAMU), Gonzales (DEB), Hemphill (USNM), Leon, Jasper, Limestone, Montgomery, Newton (TAMU), Robertson (DEB, TAMU), San Jacinto (TAMU, MEM, BYUC), Titus (DEB), Travis, Tyler, Walker, Williamson (TAMU), Wood (DEB). Vermont: county unknown (Carpenter 1940). Virginia: county unknown (Carpenter 1940), [Fairfax] (Parfin & Gurney 1956), (BYUC); Louisa (BYUC). Wisconsin: [Lincoln, Jefferson] (Parfin & Gurney 1956), Barron, Green Lake, Florence, Langlade, Marinette, Marquette, Oconto, Vilas, Waupaca (Throne 1971); [LaCrosse] (FSCA), Oneida (AMNH). New State Records. CANADA. Nova Scotia: Shelburne Co., West Branch Roseway River, hwy 203, Lower Ohio, 24-VI-1993, Baumann & Kondratieff, 1 male (BYUC); Yarmouth Co., Clyde River, hwy. 203, Flintsone Rock, 24-VI-1993, Baumann and Kondratieff, 1 male (BYUC). UNITED STATES. Kentucky: Hart Co., 4.8 mi NNW Horse Cave, 23-V-1987, E. A. Lisowski, BL trap on Green River, 1 female (INHS); McCreary Co., Cumberland River, Cumberland Falls State Park, hwy 90, 11-V-1988, Baumann, Kircher, Kondratieff & Nelson, 1 male (BYUC). South Carolina: Aiken Co., Upper Three Runs, 4-VI-1984, B. C. Kondratieff, 1 male (BYUC); [Darlington Co.], Society Hill, Cedar Creek, 4-V-1944, Frison & Ross, 1 male (INHS); Greenwood Co., Saluda River at SC hwy 34 bridge downstream from dam, 25-VII-1974, Sanderson & Unzicker, blacklight trap, 5 females (INHS). West Virginia: Hampshire Co., North River Mill, 30-VI-1990, C. R. Nelson, R. F. Whitcomb, 1 male (BYUC); Ohio Co., Wheeling, 4- VI-2002, Baumann & Kondratieff, 1 male, 2 females (BYUC).

Climacia californica Chandler Figures 4, 17, 24–26

This species is most similar in appearance to C. areolaris, but it is usually smaller in size and the pale colored area of pterostigma covers only four veins (Fig. 17). The antennae are pale to dark brown, and the postocciput is pale brown and not distinct in color from the rest of the head. Male terminalia in lateral view have the ectoproct nearly parallel-sided (Fig. 24) and its dorsal surface bears numerous, prominent dentigerous-based setae. The internal spines and vertical wall of the gonarcus complex together appear H-shaped in dorsal view (Fig. 25). Female terminalia have tergite 9 longer than broad in lateral view

FIGURES 11–14. Forewings and labial palps of adult Sisyridae (modified from Parfin & Gurney 1956). 11–12. Forewings: 11. Climacia. 12. Sisyra. 13–14. Terminal segment of labial palp: 13. Climacia. 14. Sisyra.

SPONGILLAFLIES © 2006 Magnolia Press 9 ZOOTAXA Climacia californica previously was reported from northern California (Chandler 1357 1953), Oregon (Parfin and Gurney 1956), and Idaho (Clark 1985). New distribution records for this species are reported here from Durango, Mexico, and Texas and Utah in the United States which greatly expands its known range. Moreover, the updated distribution of this species suggests that it is distributed throughout the southwestern United States and Mexico. Climacia californica is occasionally co-collected with C. chapini where their respective ranges overlap. Specimens were collected in May, August and November. Published and Supplemental State and County Records. MEXICO. Sonora (Oswald et al. 2002). UNITED STATES. California: Lake (Chandler 1956) (USNM); Imperial (BYUC). Idaho: Minidoka (Clark 1985); Twin Falls (BYUC). Oregon: [Deschutes] (Parfin & Gurney 1956). New State and Country Records. MEXICO. [Durango]: 2 mi NW Nombre de Dios, Dgo., 24-VII-1969, J. Doyen & J. Haddock, 1 female (EMEC). Michoacan: Patzquero, 31-VIII-1938, L. J. Lipovsky, 1 female (SEMC). Tamaulipas: vic. of Punta Piedra, 25-IX- 1981, W. H. Cross, blacklight trap, 1male, 1 female (MEM). UNITED STATES. Texas: Bandera Co., Camp Verde, Verde Creek, 8-VIII-1992, D. E. Bowles, blacklight, 1 male [genitalia damaged] (DEB); Brewster Co., county park (The Post), 8 km S Marathon, elev. 1090 m, 11-V-1993, J. Gelhaus & D Koenig, 1 female (ANSP); Val Verde Co., Devils River at Dolan Falls, 4-7-VIII- 1994, C. R. Nelson, R. S. Hansen, A. W. Hook, O. Hernandez, S. K. Jasper, R. Gibson, UV-light, 1 female (BYUC); same, but Dolan Falls Preserve, N 30o 25’ 8.9”, W 97o 48’ 46.9”, 3-VI-1997, D. E. Bowles, UV- & MV-light, 1 adult [abdomen missing] (DEB); same, but at Dolan Falls, 15-V- 1999, light trap, 1 male (DEB); same, but Rio Grande at Langtry, 21-VI-1990, E. G. Riley & C. S. Wolfe, UV- light, 1 female (TAMU). Utah: Washington Co., Red Cliffs campground near Leeds, 5-X- 1979, J. A. Stanger, 1 female (BYUC).

Climacia chapini Parfin & Gurney Figures 3, 6, 15, 18–20

The mostly hyaline forewings with narrow, conspicuous radiomedial pigment streaks (Fig. 15) distinguish this species from C. areolaris and C. californica. In lateral view, male terminalia have the posterior border of the ectoproct straight and bearing numerous, coarse, dentigerous-based setae (Fig. 18). The ventral lobe of the ectoproct is triangular in shape and much darker than the dorsal portion and its outer margin also has coarse dentigerous-based setae. Internal spines of the gonarcus complex are broadest anteriorly and taper posteriorly in dorsal view (Fig. 19). Female terminalia have tergite 9 longer than broad in lateral view (Fig. 20). The apex of the gonapophyses curve abruptly posteriad at an angle of approximately 65o with fine serrations dorso-apically.

FIGURES 15–17. Forewings of Climacia. 15. Climacia chapini. 16. Climacia areolaris. 17. Climacia californica

Parfin and Gurney (1956) described this species from specimens collected in Texas and New Mexico. Subsequently, Penny et al. (1997) reported C. chapini from Oklahoma, and Flint (1998) reported the species from Chiapas, Mexico. In this study, new distribution records for C. chapini were recorded from Durango and San Luis Potosi, Mexico suggesting it occurs throughout much of Mexico. The flight period ranges from April through December. Adults are often co-collected with C. californica where their respective ranges overlap, but they are typically more abundant than the latter species.

FIGURES 18–26. Male and female terminalia of Climacia (modified from Parfin & Gurney 1956). 18–20. Climacia chapini: 18. Male, lateral, with gonarcus complex beneath; 19. Male, dorsal; 20. Female, lateral. 21–23. Climacia areolaris: 21. Male, lateral, with gonarcus complex beneath; 22. Male dorsal; 23. Female, lateral. 24–26. Climacia californica: 24. Male, lateral, with gonarcus complex shown beneath; 25. Male, dorsal; 26. Female, lateral.

12 © 2006 Magnolia Press BOWLES Published and Supplemental Records. MEXICO. Chiapas: (Flint 1998);. ZOOTAXA Chihuahua: (Oswald et al. 2002). Veracruz: (Oswald et al. 2002). UNITED STATES. 1357 Oklahoma: county unknown (Penny et al. 1997); Marshall (BSC). New Mexico: Eddy (Parfin & Gurney 1956). Texas: Bosque, [Colorado], Guadalupe, [Live Oak], Sutton (Parfin & Gurney 1956), Victoria (Parfin & Gurney 1956); Bandera (DEB), Bastrop (BYUC), Blanco (TAMU, INHS), Brazos, Burnet, Cameron, Dallas (DEB), Gillespie (TAMU), Gonzales (DEB, TAMU), Hidalgo (TAMU, EMEC), Jasper (TAMU), Kerr (INHS), Kimble (UMC), Mason, Robertson (TAMU), Starr (DEB), Tarrant (EMEC), Travis (BYUC), Val Verde (ANSP, BYUC, DEB, EMEC, INHS, TAMU). New State Records. MEXICO. [Durango]: 2 mi NW Nombre de Dios, Dgo., 24-VIII- [19]69, J. Doyen & J. Haddock, 2 females (EMEC). San Luis Potosi: 3 mi W El Naranjo, elev. 1100’, Platanus lined stream, 31-V-1971, blacklight, A. Newton, 1 female (INHS).

Sisyra apicalis Banks Figures 8, 10, 27–29

This small species usually is 5 mm or less in length and can be distinguished from S. nigra and S. vicaria in having the antennae with the basal half and apical 4 or 5 segments dark brown and the intervening segments pale yellow. The forewings have prominent intervenational streaking. The ectoproct of the male terminalia is elongate in lateral view, the distal two-thirds parallel-sided, and the apex rounded (Fig. 27). The parameres are divided into two distinct portions (Fig. 28), but only the distal portion is visible in lateral view. The female terminalia in lateral view show tergite 9 nearly round in shape and with only a posterior articulation ridge (Fig. 29). The gonapophyses are rounded apically and gradually arch posteriad. Parfin and Gurney (1956) reported S. apicalis from the southern United States and Cuba. In this study, new distributional records for S. apicalis are recorded from Alabama, North Carolina and Texas in the United States. In Mexico and the United States, this species appears to be restricted to the Gulf of Mexico coastal plain (Parfin & Gurney 1956, Poirrier 1969, Poirrier & Arceneaux 1972, Lago 1981, Oswald et al. 2002, this study). I have seen other specimens from the Bahamas, Costa Rica, and Panama. An additional record of S. apicalis from the Amazon River Basin (Penny 1981) suggests this species is widely distributed throughout the Neotropics. Adult specimens have been collected during April, November and December. This species has been collected in February in Costa Rica (Penny et al. 2002) and is present throughout the year in central Amazonia, but more prevalent in June and July (Penny 1981). Although this species has a wide distribution, it is seldom collected and is not well represented in most museums.

FIGURES 27–35. Male and female terminalia of Sisyra (modified from Parfin & Gurney 1956). 27–29. Sisyra apicalis: 27. Male, lateral; 28. Male, dorsal; 29. Female, lateral. 30–32. Sisyra vicaria: 30. Male, lateral; 31. Male, dorsal; 32. Female, lateral. 33–35. Sisyra nigra: 33. Male, lateral; 34. Male, dorsal; 35. Female, lateral.

14 © 2006 Magnolia Press BOWLES Published and Supplemental State, County and Parish Records. MEXICO. Quintana ZOOTAXA Roo: (Oswald et al. 2002). UNITED STATES. Florida: Hardee, Marion (Parfin & 1357 Gurney 1956), Paradise Key (Parfin & Gurney 1956); Baker, Dade, Monroe (FSCA), Nassau (SEMC); Paradise Key, Putnam (FSCA), Sarasota (SEMC). Georgia: [Chatham] (Parfin & Gurney 1956). Louisiana: St. Tammany (Poirrier 1969); East Feliciana (Poirrier & Arceneaux 1972). Mississippi: Lafayette (Lago 1981); Jackson, Oktibbeha (MEM). New State Records. UNITED STATES. Alabama: Baldwin Co., Bon Secour N. W. Refuge, T95, R32 E, Sec 24, 5-6-IX-1988, R. L. Brown, J. MacGown, 1 male; same, but Gulf State Park, 26-VII-1985, R. L. & B. B. Brown, 1male (MEM); North Carolina: [Camden Co.], South Mills, IX-1-1962, A. B. Gurney, 3 adults (sex not determined) (USNM); Texas: Brazos Co., 9.5 mi ENE Kurten, N 30o 48’ 39” , W 96o 10’ 30”, 25-IX- 1998, J. D. Oswald, MV-light, Navasota River bottomland, 1 Ε (TAMU); Sabine Co., 9 mi E Hemphill, “beech bottom”, 24-IX-7-X-1989, R. Anderson & E. Morris, Malaise trap, beech magnolia forest, 1 male, 2 females (TAMU); San Jacinto Co., Big Creek, at TX hwy 150, 2 mi W Shepard, 23-X-1992, UV-light, Moulton & Alexander, 2 males (TAMU).

Sisyra nigra (Retzius) Figures 7, 9, 33–35

This species is closely related to and difficult to distinguish from S. vicaria. The R2 vein in the forewing of S. nigra is forked equal to or apical of the junction of the Sc and R1 veins. The length of this fork varied among the S. nigra specimens I examined, but it usually was about one-half the distance to the junction of Sc and R. The head, antennae, and thorax are dark brown to black in color. In lateral view, the ectoprocts of the male terminalia are twice as long as wide (Fig. 33). In dorsal view, the ectoprocts are roughly parallel-sided for the basal three-fourths, and with the apical one-fourth abruptly arching mesad and tapering to a point (Fig. 34). The parameres of the gonarcus complex are roughly L-shaped and less than one-half the length of ectoprocts. Female terminalia have tergite 9 broadly triangular in lateral view and with only a posterior articulation ridge (Fig. 35). Sisyra nigra has a Holarctic distribution and is widely distributed in Canada, Europe and the northern United States. Most previously published North American records for this species have been reported under the name S. fuscata (Fabricius) which Leraut (1980) indicates is a junior synonym of S. nigra. New distributional records for S. nigra are reported here from Newfoundland and Nova Scotia, Canada, and Montana and Vermont in the United States. The adult flight period ranges from June to August. Adults of S. nigra and S. vicaria are occasionally collected in the same habitats (Parfin & Gurney 1956), although the former is not as common as the latter. Published and Supplemental State and County Records. CANADA. British Columbia: (Walker 1853). Ontario: (Walker 1853). Quebec: (Walker 1853). UNITED

SPONGILLAFLIES © 2006 Magnolia Press 15 ZOOTAXA STATES. Alaska: (Walker 1853). Indiana: county unknown (Smith 1925). Maine: 1357 county unknown (Walker 1853), [Kennebec, Hancock] (Parfin & Gurney 1956); Penobscot (BYUC), [Aroostook] (USNM). Massachusetts: county unknown (Parfin & Gurney 1956). Michigan: county unknown (Walker 1853), [Allegan], Houghton, Livingston (Parfin & Gurney 1956), [Wayne] (Parfin & Gurney 1956); Kalamazoo (AMNH), Cheboygan (SEMC). Minnesota: county unknown (Walker 1853), Cass, [Itasca] (Parfin & Gurney 1956). New York: county unknown (Walker 1853), [Hamilton] (Parfin & Gurney 1956). Wisconsin: county unknown (Walker 1853), [Jefferson, Washburn] (Parfin & Gurney 1956), Barron, Florence, Jefferson, Marquette, Oneida, Sheboygan, Walworth, Washburn, Waukesha, Waupaca, Wood (Throne 1971). New State Records. CANADA. New Foundland: Terra Nova Natl. Park, Junction Pond, Notre Dame Cp., 12-VII-[19]61, 3 adults (sex not determined) (USNM). Nova Scotia: Juniper Park, W of Beechville, 7-VIII-1965, R. A. Evers, 2 females (INHS). UNITES STATES. Montana: Missoula Co., Salmon Lake, 7.5 mi. N of jct 200 & 83, 23- VII-1988, P. Skelley, UV-light, 6 males, 11 females (DEB); same, but 1 male (FSCA). Vermont: Orleans Co., 2 mi N of East Charleston on north shore of Echo lake, 18-VIII- 1967, Sweeping, M. A. Deyrup, 1 adult (sex not determined) (AMNH).

Sisyra vicaria (Walker) Figures 7, 9, 30–32

This species is closely related to and difficult to distinguish from S. nigra. The forewing usually has the R2 deeply forked and basal to junction of the Sc and R1 veins. The antennae have only 2–3 basal segments dark brown in color with the remaining segments yellow or rarely the basal third of the antennae are dark and apical two thirds paler in color. The vertex of the head is pale brown to yellow. The ectoprocts of the male terminalia are as long as wide in lateral view (Fig. 30) and broadly C-shaped in dorsal view (Fig. 31). The gonarcus complex has prominent parameres that are more than one-half as long as the ectoprocts. Female terminalia have tergite 9 broadly oval-shaped in lateral view and with anterior and posterior articulation ridges (Fig. 32). This species is widely distributed in the U.S. and Canada (Parfin & Gurney 1956, Penny et al. 1997). New distributional records are reported here from Alabama, Arkansas, New Hampshire, New Jersey, Ohio and Vermont in the United States. Given the distribution of S. vicaria in southern California, Arizona and Texas, this species likely occurs in northern Mexico as well, but it has not yet been collected there. Adults have been taken in January and April through October. This species is occasionally collected with C. areolaris and S. nigra where their ranges overlap. Published and Supplemental State, County and Parish Records. CANADA. British Columbia: (Currie 1904). Nova Scotia: (Carpenter 1940). Ontario: (Carpenter 1940).

16 © 2006 Magnolia Press BOWLES Quebec: (Carpenter 1940). UNITED STATES. Arizona: county unknown (Parfin & ZOOTAXA Gurney 1956). California: Lake (Grigarick 1975); Mendocino Co. (EMEC). 1357 Connecticut: county unknown (Carpenter 1940); Fairfield, Litchfield, New London, Tolland, Windham (PRC). District of Columbia: (Carpenter 1940). Florida: Alachua, [Duval], Paradise Key (Parfin & Gurney 1956). Georgia: county unknown (Walker 1853); [Charlton, Whitfield, Wilkinson] (Parfin & Gurney 1956), Thomas (FSCA), [Gordon] (INHS). Florida: county unknown (Carpenter 1940), Leon (AMNH), Highlands, Liberty, Jackson, Marion, Okaloosa, Putnam, Sarasota, Suwannee (FSCA). Illinois: county unknown (Carpenter 1940), Mason, [Alexander, Lake, DuPage, Vermilion] (Parfin & Gurney 1956), Champaign, [Clark], Clay, [Cook], Iroquois, [Kankakee], Marion, Schuler, Union (INHS), [Pope] (MEM). Indiana: [Lake] (Parfin & Gurney 1956), Steuben (INHS). Kansas: Hodgeman (Parfin & Gurney 1956), Bourbon, Butler, Chase, Crawford, Douglas, Elk, Kiowa, Johnson, Montgomery, Nemaha, Pottawatomie, Pratt and Woodson (Huggins 1980). Kentucky: Carter, Harlan (Parfin & Gurney 1956), Bell (Brown 1974). Louisiana: Jefferson, Pointe Coupee, St. Charles, St. Tammany (Poirrier 1969); Beauregard, East Feliciana (Poirrier & Arceneaux 1972). Maine: [Aroostook, Kennebec, Penobscot, Kennebec, Washington] (Parfin & Gurney 1956); Waldo Co. (BYUC). Massachusetts: county unknown (Carpenter 1940), [Middlesex, Norfolk] (Parfin & Gurney 1956). Maryland: county unknown (Carpenter 1940), [Montgomery] (Parfin & Gurney 1956). Michigan: Cheboygan, [Emmet], Wayne (Carpenter 1940), Livingston, [Washtenaw] (Parfin & Gurney 1956), Kalamazoo (INHS), Allegan, Ingham, Kalamazoo, Schoolcraft (MSU). Minnesota: county unknown (Walker 1853), Cass, Clearwater, Kittson, Marshall, Polk, Traverse (Parfin 1952), [Itasca, Millelacs] (Parfin & Gurney 1956). Mississippi: Stone, Wilkinson (Poirrier & Holzenthal 1980); Lafayette (Lago 1981), Grenada, Oktibbeha, Smith (MEM), [Marshall] (INHS). Missouri: Dunklin (Froeschner 1947), Boone (INHS, UMC), Cape Girardeau (INHS). New York: county unknown (Carpenter 1940), Tompkins, Westchester counties, [Erie, Essex, Hamilton, Herkimer, Montgomery, Ostego, Suffolk] (Parfin & Gurney 1956). North Carolina: [Wake] (Carpenter 1940). North Dakota: Foster (Stoakes et al. 1983), Stark (BYUC). Oklahoma: Grayson, Marshall (White 1976), Delware, (ONHS), Latimer (DEB, TAMU). Oregon: [Jackson] (Carpenter 1940). Pennsylvania: county unknown (Carpenter 1940). Rhode Island: [Washington] (Parfin & Gurney 1956). Tennessee: [Montgomery] (Parfin & Gurney 1956). Texas: Bexar (Parfin & Gurney 1956) (DEB), Colorado (Carpenter 1940); Aransas (USNM), Brazos (TAMU), Coleman (BYUC), Fort Bend (TAMU), Gonzales (INHS), Grimes (TAMU), Guadalupe (USNM), Jackson (SEMC), Jasper (TAMU), Montgomery (TAMU), [Tarrant] (INHS). Virginia: [Fairfax] (Parfin & Gurney 1956). Washington: [Spokane] (Parfin & Gurney 1956), Giles (SEMC). Wisconsin: county unknown (Carpenter 1940), [Vilas] (Parfin & Gurney 1956), Barron, Grant, Florence, Fond du Lac, Marquette, Oneida, Ozaukee, Walworth, Waukesha, Wood, Vilas (Throne 1971). New State Records. UNITED STATES. Alabama: Baldwin Co., Bon Secour N. W.

SPONGILLAFLIES © 2006 Magnolia Press 17 ZOOTAXA Refuge, T95, R2E, Sec 24, 5-6-IX-1988, R. L. Brown, J. MacGown, 1 male, 2 females 1357 (MEM). Arkansas: Logan Co., Cove Lake, 23-V-1991, blacklight, J. Powell, 1 female (E MEC); [Poinsett Co.], Marked Tree, Little River, 22-V-1957, Ross & Stannard, 1 female (INHS). New Hampshire: no additional data (USNM). New Jersey: Cumberland Co., Parvin St. Park, nr Vineland, 8-VIII-1971, 1 adult (sex not determined) (AMNH). Ohio: Delaware Co., jct Deer Run & Scioto Rivers, 27-VII-1985, S. M. Clark, 1 male (BYUC): same, but O’Shaughnessy Res. nr Shawnee Hills, 7-IX-1985, S. M. Clark, 1 female (BYUC).

Acknowledgements

I thank the many curators, collection managers, and other colleagues who graciously provided me with specimens to study. I especially thank Richard Baumann for his support and interest in this project, and Oliver S. Flint, Jr., Atilano Contreras-Ramos, John D. Oswald, and an anonymous reviewer for critical discussions and comments on an earlier version of this work.

References

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