9.0 kg), and prey items were grouped into four categories, that is, shrimps, crabs, teleost fishes, and elasmobranch fishes. Percent iri’ s were calculated as a percent of total iri within each cobia size class. Acetate impressions of cobia scales were difficult to interpret, therefore, sagittal otoliths of cobia were used to estimate specimen age. Sagittae of cobia were removed, washed in distilled water, and stored dry in individually labeled envelopes. Sagittae were embedded in 14x6x3-mm epoxy molds. Casts were affixed to a microscope slide with a drop of cyanoacrylate glue, then clamped to the arm of a circular low-speed saw. A 0.5-mm transverse section was made through the sagittal focus using a diamond-edge circular blade. The resulting wafer was permanently mounted to a microscope slide with a fixative. Sagittal sections were viewed on a dissecting microscope (16x) with transmitted, polarized light. Cross-sectioned sagittae had an opaque central core, followed by alternating translucent and opaque zones (Fig. 2). Although marginal increment analyses were precluded because specimens were unavailable throughout the year, most sagittae had an opaque edge, or an opaque zone in close proximity to the sagittal edge. Moreover, research in the northern Gulf of Mexico (Franks et al. 1991, Thompson et al. 1991) confirmed the validity of the formation of one translucent and one opaque zone on cobia sagittae each year. Thus, I assumed that one translucent and one opaque zone was deposited each year, and that opaque zones could be used to estimate cobia ages. Opaque zones along the ventral medial axis were counted as apparent annuli; estimated fish ages were based on opaque zone counts. I used the SAS NLIN procedure with the Marquardt option (SAS Institute, Inc. 1987) to estimate von Bertalanffy growth parameters based on individual fork lengths. Lengths referred to in the text are fork lengths. 6 Joseph W. Smith Fig. 2. Cross-sections (0.5 mm thick) of cobia sagittae: a) sagitta from age 3 fish (89 cm FL male, 18x magnification), b) sagitta from age 8 fish (125 cm FL female, 18x magnification). Note that spheres are artifacts of fixative. Life History of Cobia 7 RESULTS Size and Age Composition Four hundred sixteen cobia were collected. Most ( n = 366) were acquired from recreational hook-and-line fishermen, while others came from trawls ( n = 34), gill nets ( n = 7), pound nets (n = 4), stop nets ( n - 2), long hauls ( n = 2), and purse seine ( n = 1). A majority (n = 356) of the specimens came from North Carolina waters, mostly from inlet areas. A few specimens were from the Virginia portion of Chesapeake Bay (n = 17), and others were collected by port agents in South Carolina ( n = 11), and northeast Florida ( n = 15). Research trawls (75-ft high-rise mongoose net) from Daytona Beach, Florida, to Cape Fookout, North Carolina captured 17 specimens at ocean stations in depths 7-17 m. Using pooled data from all gear types, 174 male cobia ranged from 39 to 136 cm and 0.47 to 32.0 kg, and 182 females ranged from 44 to 142 cm and 0.66 to 32.2 kg (Fig. 3). Only 27 of 152 (17.8%) males, taken by hook-and-line, measured greater than 100 cm; conversely 91 of 174 (52.3%) of the females caught by the same gear were greater than 100 cm (Fig. 3). North Carolina enacted bag (2 fish/angler/day) and minimum size limits (33 inches [84 cm] FL) for cobia in 1991, thus bringing the state in line with corresponding cobia regulations in other south Atlantic states and the Federal Fisheries Conservation Zone (3-200 miles from shore). Between 1983 and 1990, 261 cobia caught by hook-and-line were examined, and 65 (24.9%) were less than 84 cm. Between 1991 and 1994, only five (5.3%) of 93 fish caught by hook-and-line were less than 84 cm, and four of these were 82-83 cm. Sectioned sagittae from 326 specimens were examined for opaque zone counts (Fig. 2). Mean observed fork length of cobia increased with opaque zone count (Table 2). Otoliths with no opaque zones distal to the sagittal core presumably came from young-of-the-year cobia that averaged 31-cm ( n = 17, range = 21-46-cm). Age 1 cobia, or those with one opaque zone distal to the core, averaged 51 cm ( n = 9, range = 39-64-cm). Mean length of females was larger than mean length for males at a given estimated age (Table 2). Maximum estimated age was 14 for males, and 13 for females. The von Bertalanffy growth coefficient, k, was greater for males than females, although mean asymptotic size was larger for females (Table 3). Seasonality and Distribution Initial catches of cobia by North Carolina anglers usually occurred in March or April 50-65 km offshore over rocky outcroppings and Table 2. Sample size ( n ), fork length range, mean observed fork lengths (±1 SE), mean observed mass, and von Bertalanffy estimates of fork lengths (VB FL) for each sex of cobia from North Carolina and adjacent waters, 1983-94, by estimated age. Results of Richards’ (1967 and 1977) studies are presented for comparison. All lengths are in cm. 8 Joseph W. Smith o ON C/3 -a t-« cd J3 O 5 XXXOOOnOnOO NO rH (N ON 04 CO NO 00 ON t— I o ■xt- X rH Tt Tt rH I/O t"- 00 On O O O On On O rH 00 00 l/o -d- NO iohaOH O NO O CO (N CN CO CO 5*. T3 3 GO C gh ix S3 oo w 00 t-H +1 IX "O « Gg C/3 X o 4) 00 ^ cd OH(N)ONd’^'Or-ICOCONtd‘,tl/) int^OOOOONONOOOOOOOO t-h Ox On NO fx 00 htohH^nooooH OHHNNNNCOCO O CO CO Tt T— i C4 T— ( (n) MO IO "t COONhCOd’OIO'OcOtN t— icoNdodoNOT-HcboocN HtC4rHr-lrHCslC4C4XrH in 1 'ct CO X X 00 (X ON o\ 00 On o O TCj- X r- 00 Os On t-H t-H t-H t-H CO ^H(NcoHtmxtx-oooNO C/3 5-6 m deep. These sites were characterized by long, straight troughs or embayments (up to several kilometers long and/or wide), often with adjacent feeder creeks or channels, e.g., Bogue Sound, Newport River, and Wallace and Blair channels of Ocracoke Inlet. Peak catches of cobia in the North Carolina sounds occurred during June, and declined thereafter (Table 4). The latest record for an adult cobia taken by hook-and-line in the Carolina sounds during this study was 18 August 1988. Cobia were captured during summer in the nearshore ocean adjacent to buoys and fishing piers, and over artificial reefs and live bottom areas. Catches were often incidental to bottom fishing or live-bait fishing for other species. During May 1988 and June 1991, catches were poor in the sounds following the passage of unseasonable cold fronts that quickly chilled estuarine water temperatures from 26 C to 19 C and 28 C to 22 C, respectively. Juvenile cobia also occurred in North Carolina sounds during summer. Young-of-the-year (based on length frequency distributions Life History of Cobia 11 Table 4. Number of cobia processed that were and caught by hook-and-line in North Carolina by month and date, 1983-90 (date intervals arbitrarily chosen). Dates May June July August September 1-7 1 45 8 0 0 8-15 5 100 5 0 1 16-22 27 6 3 1 0 23-31 20 33 5 0 0 Totals 53 184 21 1 1 and otolith analyses) were collected in pound nets and long haul nets from Pamlico Sound in August and September (Fig. 3). Age 1 fish occurred in the sounds from late May through mid-September, and most specimens were taken by hook-and-line. Food Habits During 1989-1990, 140 cobia stomachs were examined, of which 72.1% ( n - 101) contained representative food items. IrV s were computed from these samples and nine additional stomachs with food items from 1987 to 1988. Twenty-four species groups of crustaceans, 16 species groups of fishes, and one cephalopod were identified from 110 stomachs (Table 5). After pooling data from all three sampling areas, the blue crab, Callinectes sapidus, had the highest iri, followed by the blackcheek tonguefish, Symphurus plagiusa, and unidentified fish remains. Other identifiable fishes in the diet with high iri’s were pipefishes, Syngnathus sp., and the smooth dogfish, Mustelus canis. Items apparently incidentally ingested included eelgrass ( Zostera marina) blades, small fragments of oyster shell ( Crassostrea virginica), and small gastropods. In the Beaufort area, the blue crab (Table 6) had the highest iri, followed by the smooth dogfish, pipefishes, and dasyatid sting rays. Abundant crustaceans included the iridescent swimming crab, Portunus gibbesii, the brown shrimp, Penaeus aztecus, and the mantis shrimp, Squilla empusa. High-ranking food items from the Hatteras- Ocracoke area (Table 6) were the blackcheek tonguefish and the blue crab. Important food items from offshore waters (Table 6) included the coarsehand lady crab, Ovalipes stephensoni, unidentifiable fishes, the blotched swimming crab, Portunus spinimanus, and rock shrimps, Sicyonia sp. Among individual prey taxa, elasmobranchs were the largest prey 12 Joseph W. Smith Table 5. Percent frequency of occurrence (/), percent number ( n ), percent mass (w), and index of relative importance (iri) of food items in cobia stomachs from North Carolina and adjacent waters, 1987-90. Prey Taxa / n w iri Mollusca Cephalopoda Loligo plei 0.9 0.2 <0.1 <1 Arthropoda Crustacea Stomatopoda Squilla sp. 2.7 0.7 0.4 3 S. empusa 3.6 1.3 2.5 14 S. neglecta 1.8 4.2 2.5 12 Decapoda Penaeidae Penaeus sp. 3.6 0.9 0.2 4 P. aztecus 6.4 2.9 3.7 42 P. setiferus 0.9 0.2 0.2 <1 Trachypenaeus constrictus 0.9 1.5 <0.1 <1 Sicyoniidae Sicyonia sp. 3.6 7.9 2.7 38 Palaemonidae Palaemonetes vulgaris 0.9 0.4 <0.1 <1 Crangonidae Crangon septemspinosa 4.5 2.6 0.2 13 Upogebiidae Upogebia sp. 0.9 0.2 <0.1 <1 Albuneidae Albunea gibbesii 0.9 0.2 0.1 <1 Portunidae Ovalipes sp. 6.4 3.7 2.0 36 O. ocellatus 2.7 0.9 1.1 5 0. stephensoni 6.4 4.6 4.0 55 Callinectes sp. 7.3 3.3 1.8 37 C. sapidus 30.0 15.4 19.2 1,038 C. similis 3.6 0.9 0.8 6 Portunus sp. 3.6 1.1 0.5 6 P. gibbesii 5.5 2.2 1.8 22 P. spinimanus 2.7 0.7 1.4 6 Unidentified portunid remains 4.5 2.2 0.3 11 Xanthidae Menippe mercenaria 0.9 0.4 0.1 <1 Unidentified decapod remains 0.9 0.2 <0.1 <1 Chondrichthyes Carcharhinidae Mustelus canis 6.4 6.1 21.2 175 Life History of Cobia 13 Table 5. Continued. Prey Taxa / n w iri Dasyatidae Dasyatis sp. 3.6 0.9 12.7 49 Osteichthyes Clupeidae Opisthonema oglinum 0.9 0.4 0.4 1 Engraulidae Anchoa sp. 0.9 1.3 0.1 1 Synodontidae Synodus foetens 0.9 0.2 0.3 <1 Batrachoididae Opsanus sp. 3.6 1.1 2.7 14 Syngnathidae Hippocampus sp. 0.9 0.2 <0.1 <1 Syngnathus sp. 19.1 7.0 2.2 176 Sparidae Lagodon rhomboides 0.9 0.2 0.2 <1 Uranoscopidae unidentified remains 0.9 0.2 <0.1 <1 Soleidae Trinectes maculatus 0.9 0.7 0.4 1 Cynoglossidae Symphurus plagiusa 17.3 13.2 6.8 346 Balistidae unidentified remains 2.7 1.3 0.1 4 Tetradontidae Sphoeroides maculatus 0.9 0.7 1.9 2 Diodontidae Chilomycterus schoepfi 0.9 0.2 1.2 1 unidentified fish remains 21.8 7.5 4.5 262 ingested. Smooth dogfish pups ( n = 28) averaged 42 g; dasyatid sting rays ( n = 4) averaged 173 g. The largest teleosts consumed were the striped burrfish, Chilomycterus schoepfi (n = 1, 65 g), the northern puffer, Sphoeroides maculatus (n = 3, * = 34 g), and toadfishes, Opsanus sp. (n = 5, x = 29 g). Most portunid crabs were less than 7 cm in carapace width (CW) and were ingested whole; commercial-sized blue crabs (ca. 12.5-cm CW) were rarely consumed. Ovalipid crabs were often macerated. Small balistid fishes occurred in the stomachs of juvenile cobia from offshore trawl catches and were among the smallest teleosts consumed (n = 6, x = 1 g). As cobia increased in size, penaeid shrimps and teleost fishes became relatively less important in the diet, while decapod crabs increased 14 Joseph W. Smith o ON I C" oo ON 03 G 03 u l— o o3 D t-H c3 GA G • i-H ’Oh B 03 c/3 X) C/3 X o 03 s o ■4—* C/3 03 X o o c/3 s (D i-H C/3 D X GO G D '4—4 D X H NO D X 03 H C/3 03 D t-H < CD H o ~4 D t-H Ph O Gt- rH X r- (N X O t-H 10 O X X 4T-H r- 00 On ON CO 00 X X X CM rH 2 2 03 Si 2 ~2 X C/3 03 2 s; 2 2 2 ~SS *4«-4 -*H4 03 Cu 03 2 CJ * 4h4 !s- -*H4 03 2 2 CJ 03 2 2 03 2 • •<» 03 Go « 2 2 ~~~i 2 2 r2 03 2 03 2 is. 2 CO 2 ^ 2 2 G, >3 UOG-OONXCNCNrH lOXCOXlOGfCOCO CO H v— I « 03 5 2 • 2 "2S Go ' X C/3 CJ, Sf <+h •H, T3 Oh 03 2 03 2 Sx* D X C/3 is 2 -2 2 2 2 G D 03 2 •5< Prey 1 to * Hi Hi 2 O T3 G D •H* 2 G O Oh C/3 03 2 ~2 •fca 2 2 Go 2 G Cu C/3 2 2 Oh C/3 03 2 2 ^ 2 G 03 2 B 03 2 2 2 ■4-h N 2 03 2 2 2 2 2 2 03 2 2 * 03 2 -4-h CJ, 2* 03 O, C/3 Co 2 -4-h 2 2 2 2 2 Go 2 ^ C3 "~— in. cs ON «i 2 4h4 03 2 too 2 G D GD 2 -*h4 2 2 * *»*i 2 * 'Ni H** 3 03 • f-H 2 2 3 2 O to 2 Q G 2 cq 2 ft. 2< to 2 O 2 03 2 * *«i Go 2 03 2 Sh. 2 G CJ o >4 X X D G • i-H E G X CL) 1/3 X CJ G E o 9.0 Cobia Size Class (kg) Eiasmobranchs LjTeleosts Crabs Shrimps Fig. 4. Percent iri’s for various prey groups by cobia mass interval (intervals arbitrarily chosen). in importance (Fig. 4). Eiasmobranchs, that is, the smooth dogfish and dasyatid sting rays, were consumed almost exclusively by cobia greater than 9 kg. Seventy-five percent (50 of 67) of female cobia from North Carolina sounds and inlet areas had food in their stomach at capture, suggesting that these areas may be foraging grounds before and after spawning. Reproduction One hundred and twenty-seven male and 113 female cobia were sexed and staged for maturity in the field. Most male cobia were developing or ripe (Table 7). The latter state was characterized by active spermatogenesis and copious amounts of sperm within testicular ducts (Fig. 5a). Mean gsi’s for males increased from 3.0 (SD = 1.2, n = 14) in May, to 4.7 (SD = 1.5, n = 44) in June, then declined slightly to 4.4 (SD = 1.6, n = 7) in July. Most male cobia were sexually mature by 60-65 cm FL (Table 7), or age 2. Most female cobia examined were staged as developing (Table 7), and most were sexually mature by 80 cm FL, or age 2. Histological sections revealed that the ovaries of early developing females had 16 Joseph W. Smith Table 7. Cobia from North Carolina and adjacent waters, 1983-94, in various stages of sexual development by 5-cm-FL intervals. Midpoint 5-cm-FL Males Females Immature Developing and Ripe Immature Developing Ripe < 50 5 0 4 0 0 55 1 0 1 0 0 60 1 3 1 0 0 65 0 6 1 0 0 70 0 5 1 2 0 75 0 22 5 0 0 80 0 23 0 6 0 85 0 25 0 11 0 90 0 11 0 19 1 95 0 17 0 8 0 100 0 7 0 9 0 105 0 0 0 11 0 110 0 1 0 9 0 115 0 0 0 5 0 120 0 6 0 125 0 8 0 130 0 2 0 135 0 2 0 140 0 1 0 Totals 7 120 13 99 1 many small basophilic oocytes with a few early vitellogenic oocytes (Fig. 5b), whereas the ovaries of late developing females had large (ca. 750 ;i m), yolk-filled oocyctes (Fig. 5c). Only one female had hydrated oocytes; it was uncertain if this fish was caught in estuarine or oceanic waters. A few females (collected in early June 1990) showed follicular atresia indicative of a recent spawn, yet also possessed numerous large oocytes, suggestive of an incipient spawn (Fig. 5d). Mean gsi’s for female cobia were high in May at 5.5 (SD = 2.2, n - 8), peaked in June at 5.7 (SD = 2.1, n - 49), and declined slightly in July at 5.3 (SD = 2.2, n = 8). The largest ovaries excised weighed 2.49 kg (7 June) and were in a female weighing 25.4 kg. Peak spawning in June 1989 was confirmed by neuston net collections of cobia eggs from a channel in the lower Newport River estuary about 3 km from Beaufort Inlet (Fig. 1) (L. Settle, National Marine Fisheries Service, Beaufort, North Carolina, unpublished data). During 10 sampling dates between 14 June and 18 August, peak cobia egg Life History of Cobia 17 Fig. 5. Histologic preparations of cobia gonad sections: a) ripe male (70 cm FL), b) early developing female (80 cm FL), c) late developing female (104 cm FL), d) partially spent female (88 cm FL), but with numerous large oocytes. 18 Joseph W. Smith concentrations occurred on 23 June (67 eggs/100 m3), with minor peaks occurring on 11 July (44 eggs/100 m3), and 4 August (28 eggs/100 m3). Moreover, results of a concurrent ichthyoplankton survey (1989) near Ocracoke Inlet indicated that cobia eggs were one of the most common taxa encountered during May and June (W. Hettler, National Marine Fisheries Service, Beaufort, North Carolina, personal communications). DISCUSSION Cobia occurred in the sounds and ocean inlets of North Carolina from May to July, and as Richards (1967) observed in Chesapeake Bay, initial spring catches by sport fishermen were coincident with nearshore and estuarine water temperatures rising above 20 C. Cold fronts during May and June accompanied by strong northeast winds chilled inshore water temperatures and adversely affected spring catches of cobia in North Carolina. During August and into fall, cobia were found primarily in coastal oceanic waters. Cobia reside in other major estuaries along the United States Atlantic coast during spring and summer, e.g., Port Royal and St. Helena sounds in South Carolina (Moore et al. 1980) and Chesapeake Bay (Richards 1967). This contrasts with the northern Gulf of Mexico where most cobia occur along shallow coastal waters of the Gulf and offshore in association with oil and gas platforms and rafts of Sargassum (Ditty and Shaw 1992). It is unclear where cobia from the south Atlantic coast of the United States overwinter. Winter trawl surveys by South Carolina’s Marine Resources Monitoring, Assessment and Prediction Program (South Carolina Marine Resources Research Institute, Charleston, South Carolina, unpublished data) captured cobia ( n = 22, range = 40-127 cm, x = 84 cm) during January and February between Cape Fear, North Carolina, and Cape Canaveral, Florida, in 31-75 m depths where water temperatures ranged from 15.9 to 20.8 C (also see Wenner et al. 1979). Cobia taken by various commercial gears (hand, troll, and long lines) have been processed by port agents in North Carolina during all quarters of the year, 1983-91 (L. Mercer, North Carolina Division of Marine Fisheries, Morehead City, North Carolina, personal communications). These findings suggested that off the south Atlantic coast of the United States cobia may overwinter on the outer half of the continental shelf. Although Richards (1967) used scales to age cobia from Chesapeake Bay, I found that acetate impressions of cobia scales were difficult to interpret for annuli. Alternating translucent and opaque zones of cross-sectioned sagittae were distinct, although I was unable to validate their annual nature. Nevertheless, indirect evidence supported the validity Life History of Cobia 19 of opaque zones as annuli. First, mean size of cobia increased with opaque zone count. Second, young-of-the-year cobia (based on length frequency distributions) had no opaque zone distal to the sagittal core or focus, whereas age 1 fish had one opaque zone distal to the sagittal core. Moreover, recent research in the northern Gulf of Mexico (Franks et al. 1991, Thompson et al. 1991) confirmed the validity of the formation of one translucent and one opaque zone on cobia sagittae each year. Assuming that opaque zones on cobia sagittae were valid annuli, my results indicated that cobia grew rapidly during the first few years of life, and by age 3 mean mass ranged from 6 to 8 kg. Results from public tagging programs report equally dramatic growth for recaptured specimens (Anonymous 1986, Richard 1989, Franks 1995). My study agreed closely with Richards (1967) on mean length for both sexes at age 1 and 2 (Table 2). For age 3 and older, Richards (1967) reported that mean sizes were larger. Eleven specimens were estimated as age 11 to 14, while Richards’ (1967) maximum age for cobia was age 10. Perhaps, erosion on scale edges caused him to underestimate cobia ages, as has been shown in other fishes (Chilton and Stocker 1987). Male cobia have a higher growth coefficient, k, than females, and the difference between sexes was greater for my study (0.37 to 0.24) than previous work (0.28 to 0.23; Richards 1977). Mean asymptotic FLs (Table 3) for both sexes were lower than Richards (1977) reported, possibly reflecting a greater availability of larger cobia in Chesapeake Bay during the 1960s. Age 3 females ( n = 50) predominated in the present study, whereas Richards (1967) found age 5 females ( n = 34) were most numerous. No doubt, estimates of mean asymptotic size in the present study were underestimates as the current North Carolina state record cobia (1988) weighed 46.7 kg. Cobia were primarily demersal feeders along the North Carolina coast, and they preyed on portunid crabs, penaeid shrimps, stomatopods, numerous teleosts, and small elasmobranchs. Overall, the blue crab was the most important food item in the cobia diet, which reinforces the colloquial name of “crab-eater” used along the southeastern coast of the United States (Knapp 1951, Manooch 1984). Most portunids were ingested whole, except for Ovalipes which was usually macerated. Similar to the results of the present study, Knapp (1951) found demersal prey, such as portunids, stomatopods, penaeids, and eels in cobia stomachs from the northern Gulf of Mexico. Cobia from the western Indian Ocean consumed mostly portunids, cephalopods, and eels (Darracott 1977). In the sounds of North Carolina, cobia greater than 9 kg showed a predilection for smooth dogfish pups and small dasyatid sting rays, and these were among the largest prey items ingested. Cobia may 20 Joseph W. Smith be one of the few teleosts that regularly consumed small elasmobranchs. Field inspections and histological sections of cobia gonads indicated that most adult cobia were developing and\or ripe as they entered North Carolina waters in spring. Males became sexually mature by 60-65 cm (age 2), and females by 80 cm (age 2). Richards (1967) stated that the smallest mature male in his collections measured 51.8 cm (“second. ...year of life”) and that the smallest mature female measured 69.6 cm (“third year of life”), but he did not include maturity schedules. Cobia spawned in North Carolina coastal waters from May through July, with peak spawning in June. In Virginia waters, cobia spawned mid-June through mid-August, as determined by ichthyoplankton surveys (Joseph et al. 1964). In the northern Gulf of Mexico, cobia arrived in coastal waters during April and May in prespawning condition and exhibiting peak gsi values (Biesiot et al. 1994). Some female cobia collected during June in North Carolina showed follicular atresia in the ovaries indicative of a recent spawn, yet also had numerous and adjacent, large oocytes, suggesting another potential spawning event. Data on ova diameters presented by Richards (1967) and work by Thompson et al. (1991) and Biesiot et al. (1994) in the northern Gulf of Mexico support the concept of batch spawning in cobia. Precise location of cobia spawning areas along the North Carolina coast was uncertain, although my results suggested that cobia spawned adjacent the state’s major ocean inlets. Likewise, Joseph et al. (1964) found that cobia spawned off the mouth of Chesapeake Bay in Virginia. Collections of cobia eggs in the Gulf Stream off Cape Hatteras, North Carolina, by Hassler and Rainville (1975) (almost 2,000 eggs in 10 collecting trips, May-June 1974) contrast an inlet spawning area hypothesis. In summary, cobia inhabited coastal sounds and inlet areas of North Carolina from May through July. Specimens greater than 15 kg were common, hence the species’ popularity with inshore recreational anglers. Cobia consumed a variety of demersal crustaceans and fishes; of the former, the blue crab was the most important. Spawning probably peaked during June in ocean waters adjacent major inlets. Management regulations adopted by North Carolina in 1991 prohibiting possession of cobia less than 84 cm were effective, and few fish below the minimum possession size were encountered between 1991 and 1994. Migratory routes and overwintering grounds of cobia along the south Atlantic coast of the United States are unclear. Comprehensive tagging of cobia along the south Atlantic coast of the United States and in Chesapeake Bay would help clarify (1) coast-wide migration patterns, (2) ingress and egress from estuaries to ocean, (3) fidelity to specific estuaries, and (4) movements into the northern Gulf of Mexico. Life History of Cobia 21 ACKNOWLEDGMENTS — Cobia are infrequently encountered in creel surveys, thus, I am indebted to the following anglers for allowing me to process their catches, J. DeVane, J. Govoni, S. Hyman, N. Johnson, J. Kenworthy, A. Powell, J. Smith, and K. Smith. Acquisition of cobia samples was greatly enhanced through the cooperation of Arthur, Charlie, Ronnie, and Mac of O’Neal’s Dockside and Sharon and Norman Miller on Ocracoke Island, Steve Hissey at Hatteras, and Donald and Linda Flood at Harkers Island. Others unselfishly provided samples and data from current research projects, R. Beatty and D. Machowski (South Carolina Department of Natural Resources), D. Estes (Virginia Institute of Marine Science), and J. Ross (North Carolina Division of Marine Fisheries). At the Beaufort Laboratory of the National Marine Fisheries Service, J. Merriner and D. Ahrenholz provided encouragement, advice and research facilities; D. Vaughan supplied statistical counsel; L. Settle and W. Hettler shared early life history information; M. Burton and D. Thiesen supplied headboat data and samples; B. Harvey courageously tackled various drafts of the manuscript; C. Lewis developed photographs; N. McNeil helped process samples. L. Mercer, C. Manooch, and several anonymous reviewers provided numerous beneficial comments. Special thanks go to Bill Roumillat (South Carolina Department of Natural Resources) for histological preparations. LITERATURE CITED Anonymous. 1986. Charleston cobia tours Gulf of Mexico. Saltwater Conversation 2:7-8. Biesiot, P. M., R. E. Caylor, and J. S. Franks. 1994. Biochemical and histological changes during ovarian development of cobia, Rachycentron canadum, from the northern Gulf of Mexico. Fishery Bulletin 92:686- 696. Briggs, J. C. 1960. Fishes of world wide (circumtropical) distribution. Copeia 1960:171-180. Chilton, D. E., and M. Stocker. 1987. A comparison of otolith and scale methods for ageing Pacific herring. North American Journal of Fish- eries Management 7:202-206. Darracott, A. 1977. Availability, morphometries, feeding and breeding activity in a multi-species, demersal fish stock of the Western Indian Ocean. Journal of Fish Biology 10:1-16. Ditty, J. G., and R. F. Shaw. 1992. Larval development, distribution, and ecology of cobia Rachycentron canadum (Family: Rachycentridae) in the northern Gulf of Mexico. Fishery Bulletin 90:668-677. 22 Joseph W. Smith Franks, J. S. 1995. Investigations of cobia, Rachycentron canadum, in Mississippi marine waters and adjacent Gulf waters. Study II. Stud- ies on the seasonal movements and migratory patterns of cobia, Rachycentron canadum, in Mississippi marine waters and adjacent Gulf of Mexico. Annual Report, Project No. F-91, Segment No. 6, Sport Fish Resto- ration Program, Gulf Coast Research Laboratory, Ocean Springs, Mississippi. Franks, J. S., J. T. McBee, and M. T. Allen. 1991. Estimations of age in cobia, Rachycentron canadum, from the northern Gulf of Mexico. (Abstract) 55th Meeting of the Mississippi Academy of Sciences, page 55, 21-22 February, Jackson, Mississippi. Gulf of Mexico and South Atlantic Fishery Management Councils. 1985. Final amendment 1, fishery management plan, environmental impact statement for the coastal migratory pelagic resources (mackerels). South Atlantic Fishery Management Council, Charleston, South Carolina. Hassler, W. W., and R. P. Rainville. 1975. Techniques for hatching and rearing cobia, Rachycentron canadum, through larval and juvenile stages. University of North Carolina Sea Grant Program, UNC-SG-75-30. Hrincevich, A. W., and P. M. Biesiot. 1994. Mitochondrial DNA analy- ses of cobia, Rachycentron canadum, from the northern Gulf of Mexico and the Chesapeake Bay. (Abstract) 58th Meeting of the Mississippi Academy of Sciences, page 61, 17-18 February, Biloxi, Mississippi. Humason, G. L. 1972. Animal tissue techniques. Second Edition. Free- man, San Francisco, California. Huntsman, G. R. 1976. Offshore headboat fishing in North Carolina and South Carolina. Marine Fisheries Review 38(3) : 1 3 — 23 . Knapp, F. T. 1951. Food habits of the sergeantfish, Rachycentron canadus. Copeia 1951:101-102. Joseph, E. B., J. J. Norcross, and W. H. Massman. 1964. Spawning of the cobia, Rachycentron canadum, in the Chesapeake Bay area, with observations of juvenile specimens. Chesapeake Science 5:67-71. Manooch, C. S., III. 1984. Fisherman’s guide, fishes of the southeastern United States. North Carolina State Museum of Natural History, Ra- leigh, North Carolina. Manooch, C. S., Ill, L. E. Abbas, and J. L. Ross. 1981. A biological and economic analysis of the North Carolina charter boat fishery. Marine Fisheries Review 43(8): 1—1 1 . McClane, A. J. 1965. McClane’s standard fishing encyclopedia and in- ternational angling guide. Holt, Reinhart and Winston, New York, New York. Moore, C. J., D. L. Hammond, and D. O. Myatt, III. 1980. A guide to saltwater recreational fisheries in South Carolina. South Carolina Wildlife and Marine Resources Department, Charleston, South Carolina. Musick, J. A. 1972. Fishes of Chesapeake Bay and the adjacent Coastal Plain. Virginia Institute of Marine Science Special Scientific Report 65:175-212. Pinkas, L., M. S. Oliphant, and I. L. K. Iverson. 1971. Food habits of Life History of Cobia 23 albacore, bluefin tuna, and bonito in California waters. California Department of Fish and Game, Fishery Bulletin 152. Richard, J. 1989. Tag recovery report. Tide Magazine (March/April 1989):30- 31. Richards, C. E. 1967. Age, growth and fecundity of the cobia, Rachycentron canadum, from Chesapeake Bay and adjacent Mid-Atlantic waters. Transactions of the American Fisheries Society 96:343-350. Richards, C. E. 1977. Cobia ( Rachycentron canadum ) tagging within Chesapeake Bay and updating of growth equations. Chesapeake Sci- ence 18:310-311. SAS Institute, Inc. 1987. SAS/STAT guide for personal computers, Ver- sion 6 edition. SAS Institute, Inc., Cary, North Carolina. Schwartz, F. J., W. T. Hogarth, and M. P. Weinstein. 1981. Marine and freshwater fishes of the Cape Fear estuary, North Carolina, and their distribution in relation to environmental factors. Brimleyana 7:17-37. Shaffer, R. V., and E. L. Nakamura. 1989. Synopsis of biological data on cobia, Rachycentron canadum ( Pisces : Rachycentridae). National Oceanic and Atmospheric Administration, Technical Report, National Marine Fisheries Service 82. Thompson, B. A., C. A. Wilson, J. H. Render, and M. Beasley. 1991. Age, growth, and reproductive biology of greater amberjack and cobia from Louisiana waters. Year one, a report to United States Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service. Cooperative Agreement NA90AA-H-MF089 Marine Fisheries Initiative (MARFIN) Program, Coastal Fisheries In- stitute, Louisiana State University, Baton Rouge. Waltz, W., W. A. Roumillat, and P. K. Ashe. 1979. Distribution, age structure, and sex composition of the black sea bass, Centropristis striata, sampled along the southeastern coast of the United States. South Carolina Marine Resources Center Technical Report 43. Wenner, C. A., C. A. Barans, B. W. Stender, and F. H. Berry. 1979. Results of MARMAP otter trawl investigations in the South Atlantic Bight. IV. Winter-early spring, 1975. South Carolina Marine Resources Center Technical Report 44. Received 27 September 1994 Accepted 21 March 1995 A Review of Stonefly Records (Plecoptera: Hexapoda) of North Carolina and South Carolina Boris C. Kondratieff Colorado State University Department of Entomology Fort Collins, Colorado 80523 Ralph F. Kirchner 5960 East Pea Ridge Ridgeview Apartment 1 Huntington, West Virginia 25705 AND David R. Lenat North Carolina Environmental Management Water Quality Section 4401 Reedy Creek Road Raleigh, North Carolina 27607 Abstract — The stoneflies (Plecoptera) of North Carolina and South Carolina are comprehensively reviewed for the first time. One hundred and thirteen and 83 stonefly species are recorded from North Carolina and South Carolina, respectively. Thirteen new state records are given for North Carolina and two for South Carolina. An additional 22 species are listed that may be eventually collected in either state. Unzicker and McCaskill (1982) presented the first comprehensive checklist of 131 stoneflies known or likely to occur in North Carolina and South Carolina. However, as Lenat and Penrose (1987) pointed out, this list did not distinguish between North Carolina and South Carolina, and validation of individual state records requires examination of the literature. Stark et al. (1986) and Stewart and Stark (1988) have provided recent compilations of stonefly species records for North America, listing 75 and 77 species for North Carolina and 77 and 79 for South Carolina, respectively. However, all three lists contain omissions or list species identified in error (Table 1). For example, Stark et al. (1986) did not list Taeniopteryx burksi Ricker and Ross, T. lonicera Ricker and Ross, and T. metequi Ricker and Ross from North Carolina despite the records published by Ricker and Ross (1968) or by Fullington and Stewart (1980). Notations in Table 1 are included to help clarify taxonomic changes and to distinguish between the lists. Brimleyana 23:25-40, December 1995 25 26 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat In addition to the notations in Table 1, the following species should be deleted from the list of North Carolina and South Carolina stoneflies: (1) Paracapnia opis (Newman) — this species is northeastern in distribution (Stark et al. 1986), previous determinations were in error, and all records are referable to P. angulata Hanson; (2) Leuctra tenella Provancher — a species very similar to L. carolinensis, and L. maria Hanson are also considered northeastern in distribution (Stark et al. 1986); (3) the nearest records for Alloperla imbecilla (Say) are from northwestern Virginia and West Virginia (Baumann 1974, Surdick 1985); (4) Isoperla nana (Walsh) — a small black Isoperla is considered a northeastern and central species (Stark et al. 1986); and (5) Isogenoides doratus Frison — a species that is restricted to the upper Midwest and Northeast (Stark et al. 1986). In addition, Allocapnia granulata (Claassen), A. mystica Frison, and A. pygmaea (Burmeister) were listed by Unzicker and McCaskill (1982), but no verifiable records for North Carolina and South Carolina are available. The following 13 new state records for North Carolina are based on specimens deposited in the C. P. Gillette Museum of Arthropod Diversity, Colorado State University (CSU) or the North Carolina Division of Environmental Management, Water Quality Section (NCDEM) collections. Nine species are also noted that have been reported since Stewart and Stark (1988) for North Carolina: Leuctra ferruginea (Walker) (Huryn and Wallace, 1987). Megaleuctra williamsae Hanson - Macon Co., trib. Cullasaja R., 24 May 1994, CSU; Haywood Co., R. F. Cove Cr., 23 May 1994, CSU; Jackson Co., Mull Cr., 23 May 1993, CSU. Prostoia hallasi Kondratieff and Kirchner - Gates Co., Great Dismal Swamp, 26 March 1992, CSU. Amphinemura nigritta (Provancher) - Avery Co., Linville R., 18 May 1994, CSU; Haywood Co., East Fork Pigeon R., 23 May 1990, CSU; Yancey Co., trib. to Cane R., 18 May 1994, CSU. Zapada chila (Ricker) (Ashe Co., Lenat and Penrose, 1987). Oemopteryx contorta (Needham and Claassen) - Moore Co., Suck Cr., Feb., 1984, NCDEM. Strophopteryx limata (Frison) - Haywood Co., Cataloochee Cr., Great Smoky Mt. Nat. Pk., 23 May 1993, CSU. Agnetina flavescens (Walsh) - Clay Co., Fires Cr., April 1987, NCDEM; Ashe Co., South Fork New R., March 1990, NCDEM. Acroneuria frisoni Stark and Brown - Jackson Co., Dillsboro, 5 Aug. 1982, CSU. Stonefly Records 27 A. lycorias (Newman) - Harnett Co., Barbecue Swamp, Nov., 1988, NCDEM. Paragnetina kansensis Banks (Duplin Co., Robeson Co., Lenat and Penrose 1987). Neoperla clymene (Newman) - Ashe Co., South Fork New River, CSU. Diploperla duplicata (Banks) (Guilford Co., Forsyth Co., Burke Co., Transylvania Co., Lenat and Penrose 1987). D. morgani Kondratieff and Voshell (Surry Co., Lenat and Penrose 1987). Helopicus bogaloosa Stark and Ray (Richmond Co., Robeson Co., Lenat and Penrose 1987). Isoperla burksi Frison (Chatham Co., Davie Co., Duplin Co., Randolph Co., Lenat and Penrose 1987). I. dicala Frison - Ashe Co., South Fork New River, CSU; Jackson Co., CSU. I. frisoni lilies (Cherokee Co., Stokes Co., Lenat and Penrose 1987). I. lata Frison - Clay Co., Fires Cr., 18 April 1988, NCDEM; Big Cr., Haywood Co., Great Smoky Mt. Nat. Pk., CSU. I. namata Frison (Lenat 1983). /. slossonae (Banks) (Ashe Co., Lenat and Penrose 1987); Transylvania Co. NCDEM. I. transmarina (Newman) - Moore Co., Drowning Cr., NCDEM. Pteronarcys dorsata (Say) - Ashe Co., Catawba Co., Scotland Co. NCDEM. Two new South Carolina state records are based on specimens in the C. P. Gillette Museum of Arthropod Diversity, Colorado State University (CSU) from South Carolina, and there is one additional new literature record: Isoperla burksi - Edgefield Co., Stevens Cr., 24 May 1984, CSU. I. davisi James - Edgefield Co., Stevens Cr., 24 May 1984, CSU. Taenionema atlanticum Ricker and Ross (“South Carolina,” Stanger and Baumann 1993). Several undescribed species of Isoperla are known from both states, and S. W. Szczytko (University of Wisconsin, Stevens Point) is presently describing these species. Table 2 lists the species of stoneflies known from North Carolina (113 species) and South Carolina (83 species). This table also includes 22 species marked with a that occur in surrounding states and could be collected in either state. Morse et al. (1993) noted that at least 12 stonefly species, Allocapnia fumosa Ross, Megaleuctra williamsae, Strophopteryx inaya Ricker and Ross, Sweltsa urticae (Ricker), Tallaperla elisa Stark, Acroneuria arida 28 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat (Hagen), Beloneuria georgiana (Banks), B. stewarti Stark and Szczytko, Diploperla morgani, Isoperla bellona Banks, I. distincta Nelson, and Oconoperla innubila (Needham and Claassen) occur in one or both states and are rare and vulnerable species, sensitive to human induced impacts. As Baumann (1979) clearly indicated, stoneflies are good indicators of ecosystem quality at all scales. All the species listed above by Morse et al. (1993) are considered southern Appalachian in distribution. This geographical region is being directly impacted by regional influences (e.g. acid deposition) and local landscape changes (e.g. agriculture, rural developments, and timber harvest). The very diverse stonefly fauna of both states is indicative of a wide range of high quality lotic aquatic habitats, which need active protection. Table 1. A comparison of three stonefly (Plecoptera) species lists for North Carolina (NC) and South Carolina (SC). Unzicker Stark Stewart and McCaskill et al. and Stark (1982) (1986) (1988) Euholognatha Capniidae Allocapnia aurora Ricker A. brooksi Ross A. fumosa Ross A. granulata (Claassen) A. loshada Ricker A. mystica Frison A. nivicola (Fitch) A. pygmaea (Burmeister) A. recta (Claassen) A. rickeri Frison A. stannardi Ross A. virginiana Frison A. wrayi Ross Nemocapnia Carolina Banks Paracapnia angulata Hanson P. opis (Newman) Feuctridae Leuctra alexanderi Hanson L. biloba Claassen L. carolinensis Claassen L. ferruginea (Walker) L. grandis Banks L. maria Hanson X NC,SC NC,SC X1 X NC NC X X1 X X NC NC X X NC,SC NC,SC X NC NC X NC NC X NC,SC NC,SC X NC,SC NC,SC X NC,SC NC,SC X NC NC X X NC,SC NC,SC X NC NC X NC,SC NC,SC X SC SC X NC NC X Stonefly Records 29 Table 1. Continued. Unzicker Stark Stewart and McCaskill et al. and Stark (1982) (1986) (1988) L. mitchellensis Hanson X NC NC L. moha Ricker X SC SC L. monticola Hanson X NC,SC NC,SC L. nephophila Hanson X NC NC L. sibleyi Claassen X NC,SC NC,SC L. tenella Provancher X L. tenuis (Pictet) X SC SC L. triloba Claassen X NC,SC NC,SC L. variabilis Hanson X Megaleuctra williamsae Hanson X SC SC Paraleuctra sara (Claassen) X NC,SC NC,SC Nemouridae Amphinemura delosa (Ricker) X A. nigritta (Provancher) X SC SC A. wui (Claassen) X NC,SC NC,SC Paranemoura perfecta (Walker) X NC NC Prostoia completa (Walker) X NC,SC NC,SC P. similis (Hagen) X SC SC Shipsa rotunda (Claassen) X SC NC2,SC Soyedina carolinensis (Claassen) X NC NC Zapada chila (Ricker) X1 Taeniopterygidae Boltoperla rossi (Frison) X NC NC Oemopteryx contorta (Needham and Claassen) X Strophopteryx appalachia Ricker and Ross X1 NC NC S. fasciata (Burmeister) X NC,SC NC,SC S. inaya Ricker and Ross X NC,SC NC,SC S. limata (Frison) X1 Taenionema atlanticum Ricker and Ross X NC NC Taeniopteryx burksi Ricker and Ross X (3) NC3 T. lit a Frison X NC,SC NC,SC T. lonicera Ricker and Ross X <3> SC (3) SC T. maura (Pictet) X NC,SC NC,SC T. metequi Ricker and Ross X 30 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Table 1. Continued. Unzicker Stark Stewart and McCaskill et al. and Stark (1982) (1986) (1988) X (3> SC <3) sc X1 sc sc T. parvula Banks T. robinae Kondratieff and Kirchner T. ugola Ricker and Ross Systellognatha Chloroperlidae Alloperla atlantica Baumann A. caudata Frison A. chloris Frison A. imbecilla (Say) A. nanina (Banks) A. neglecta Frison A. usa Ricker Haploperla brevis (Banks) Rasvena terna (Frison) Suwallia marginata (Banks) Sweltsa lateralis (Banks) S. mediana (Banks 1911) S. onkos (Ricker) S. urticae (Ricker 1952) Utaperla sp. Peltoperlidae Peltoperla ada Needham and Smith Peltoperla arcuata Needham Tallaperla anna (Needham and Smith) T. Cornelia (Needham and Smith) T. elisa Stark T. laurie (Ricker) T. maria (Needham and Smith) Viehoperla zipha (Frison) Viehoperla ada (Needham and Smith) Perlidae Acroneuria abnormis (Newman) A. arenosa (Pictet) A. arida (Hagen) A. carolinensis (Banks) X NC,SC NC,SC X (4) (5) X1 X6 NC,SC NC,SC X NC NC X1 <7) SC SC X8 NC,SC NC,SC X1 0) (10) X NC,SC NC,SC X NC NC NC11 NC11 X NC NC X12 X1-13 X1 XU4 NC,SC NC,SC NC,SC NC,SC NC,SC NC,SC X14 NC,SC NC,SC X14 NC,SC NC,SC X13 NC,SC NC,SC X NC,SC NC,SC X SC SC X NC NC X NC,SC NC,SC Stonefly Records 31 Table 1. Continued. Unzicker and McCaskill (1982) Stark et al. (1986) Stewart and Stark (1988) A. evoluta Klapalek X1-15 A. filicis Frison X SC SC A. internata (Walker) X1 A. lycorias (Newman) X1 A. mela Frison X15 A. perplexa Frison X1 A. petersi Stark and Gaufin X1 Agnetina annulipes (Flagen) SC A. capitata (Pictet) NC,SC16 A. flavescens (Walsh) sc Attaneuria ruralis (Hagen 1861) X NC,SC NC,SC Beloneuria georgiana (Banks) X NC NC B. steward Stark and Szczytko X NC,SC NC,SC Eccoptura xanthenes (Newman) X NC,SC NC,SC Hansonoperla appalachia Nelson X1 SC Neoperla carlsoni Stark and Baumann X SC SC N. clymene (Newman) X N. freytagi Stark and Baumann X SC N. occipitalis (Pictet)17 SC Paragnetina fumosa (Banks) X NC,SC NC,SC P. ichusa Stark and Szczytko NC,SC NC,SC P. immarginata (Say) X NC,SC NC,SC P. kansensis (Banks) X SC SC P. media (Walker) X Perlesta frisoni Banks X NC,SC NC,SC P. placida (Hagen) X SC P. placida (Hagen) complex18 NC,SC Perlinella drymo (Newman) X SC NC,SC P. ephyre (Newman) X SC SC P. fumipennis (Walsh)19 X SC P. zwicki Kondratieff, Kirchner and Stewart SC Phasganophora capitata (Pictet) X20 Perlodidae Clioperla clio (Newman) (21) NC,SC NC,SC Cultus decisus (Walker X22 NC NC Diploperla duplicata (Banks) X SC SC 32 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Table 1. Continued. Unzicker and McCaskill (1982) Stark et al. (1986) Stewart and Stark (1988) Helopicus bogaloosa Stark and Ray SC SC H. subvarians (Banks) X NC,SC NC,SC Hydroperla fugitans (Needham and Claassen) X23 Isogenoides doratus (Frison) X1 2 I. hansoni (Ricker) X NC NC I. varians (Walsh) X SC SC Isoperla bellona Banks X NC NC I. bilineata (Say) X NC NC I. clio (Newman) X21 I. cotta Ricker X SC SC I. dicala Frison X SC sc I. distincta Nelson X1 NC,SC NC,SC I. holochlora (Klapalek) X NC,SC NC,SC I. lata Frison X1 I. marlynia Needham and Claassen X SC SC I. nana (Walsh) X I. orata Frison X NC,SC NC,SC I. similis (Hagen) X NC,SC NC,SC Malirekus hastatus (Banks) X NC,SC NC,SC Oconoperla innubila (Needham and Claassen)24 NC,SC NC,SC Remenus bilobatus (Needham and Claassen) X NC,SC NC,SC Yugus arinus (Frison) X NC,SC NC,SC Y. bulbosus (Frison) X NC,SC NC,SC Y. innubilus (Needham and Claassen) X24 Pteronarcyidae Pteronarcys biloba (Newman) X25 NC,SC NC,SC P. dorsata (Say) X SC SC P. proteus (Newman) X NC,SC NC,SC P. scotti (Ricker) X NC,SC NC,SC 1 Listed by Unzicker and McCaskill (1982) “as likely to occur in North or South Carolina, but presence has not yet been confirmed.” 2 Listed only by Stewart and Stark (1988) in the “Species of Nymphs Examined” and a specimen is illustrated from Davie County, North Carolina, Yadkin River. Stonefly Records 33 Table 1. Continued (Footnotes). Ricker and Ross (1968) listed records for these species. Additionally, Stewart and Stark (1988) listed Taeniopteryx burksi in the “Species of nymphs examined,” but not in the “North American Species List and Distribution.” 4 Surdick (1985) listed Buncombe County and Great Smoky Mountain National Park, North Carolina. 5 Surdick (1985) listed Buncombe County, North Carolina. 6 Listed by Unzicker and McCaskill (1982) as Sweltsa nanina. Surdick (1985) listed Buncombe County, McDowell County, and Yancey County, North Carolina. 8 Listed by Unzicker and McCaskill (1982) as Hastaperla brevis, Zwick (1977) dis- cusses the generic synonymy. 9 Surdick (1985) listed this species from Great Smoky Mountain National Park, North Carolina. 10Surdick (1985) listed Yancey County and Great Smoky Mountain Park, North Caro- lina. "Surdick (1985) did not list Sweltsa onkos from North Carolina, presumably older records are misidentifications of S. mediana. 12This generic record is based on nymphs, presumably misidentifications of nymphs of Alloperla usa ? Utaperla gaspesiana Harper and Roy is known from West Virginia. "This species is now included in the genus Viehoperla , and V. zipha is considered a synonym of V. ada (Stark and Stewart 1981). "This species is now included in the genus Tallaperla (Stark and Stewart 1981). "Stark and Brown (1991) studied the holotype of Acroneuria evoluta and considered A. mela a synonym. 16 The records for this species in Stark et al. (1986) are composite, including both Agnetina annulipes and A. flavescens, which were distinguished by Stark (1986). The latter reference did not list any records for A. capitata from North Carolina. "Stark (1990) synonymized Neoperla freytagi with N. occipitalis (Pictet) "Stark (1989) divided the Perlesta placida complex into 12 species. Stark (1989) provided records for three species, P. placida, North Carolina; P. frisoni, Haywood County, North Carolina; Oconee County, Pickens County, South Carolina; P. nelsoni, Haywood County, Swain County, North Carolina; Oconee County, South Carolina. "Kondratieff et al. (1988) synonymized Perlinella fumipennis with P. ephyre and de- scribed P. zwicki for the species formerly identified as P. fumipennis from the Southeast. 20Zwick (1984) established the synonymy between Agnetina and Pltasganophora. See note 16. 21Szczytko and Stewart (1981) included Isoperla clio in Clioperla. 22 Stark et al. (1988) reviewed this species complex and recognized two species, Cultus decisus with two subspecies and C. verticalis (Banks). C. d. isolatus (Banks) is known from Madison County, North Carolina and C. verticalis is known from Haywood County and Swain County, North Carolina. 23Kondratieff and Painter (1986) indicated that records of Hydroperla fugitans from North Carolina or South Carolina were in error, and that the only confirmed record of this genus was from South Carolina referable to H. phormidia Ray and Stark. 24 Stark (1985) synonymized Yugus innubilus with Oconoperla weaveri Stark and Stewart. 25 Stark and Szczytko (1982) recognized Allonarcys as a synonym of Pteronarcys. 34 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Table 2. List of stoneflies (Plecoptera) recorded from North Carolina (NC) and South Carolina (SC). Species marked with an “*” occur in surrounding states and could eventually be collected in either state. New state records for both states as indicated in text are marked by a #. NC SC Euholognatha Capniidae Allocapnia aurora Ricker X X * A. brooksi Ross (TN) A. fumosa Ross X * A. granulata (Claassen) (TN, VA) * A. loshada Ricker (TN, VA) * A. mystica Frison (TN, VA) A. nivicola (Fitch) X * A. pygmaea (Burmeister) (TN, VA) A. recta (Claassen) X X A. rickeri Frison X * A. simmonsi Kondratieff and Voshell (VA) A. stannardi Ross X A. virginiana Frison X X A. wrayi Ross X X Nemocapnia Carolina Banks X X Paracapnia angulata Hanson X Leuctridae Leuctra alexanderi Hanson X X L. biloba Claassen X L. carolinensis Claassen X X L. ferruginea (Walker) X X L. grandis Banks X L. mitchellensis Hanson X L. moha Ricker X L. monticola Hanson X X L. nephophila Hanson X L. sibleyi Claassen X L. tenuis (Pictet) X L. triloba Claassen X X * L. variabilis Hanson (VA) Megaleuctra williamsae Hanson x# X Paraleuctra sara (Claassen) X X Nemouridae * Amphinemura delosa (Ricker) (TN, VA) A. nigritta (Provancher) x# X A. wui (Claassen) X X Stonefly Records 35 Table 2. Continued. NC SC * Ostrocerca albidipennis (Walker) (VA) * O. complexa (Claassen) (VA) * O. prolongata (Claassen) (VA) * O. truncata (Claassen) (VA) Paranemoura perfecta (Walker) Prostoia completa (Walker) P. similis (Hagen) P. hallasi Kondratieff and Kirchner Shipsa rotunda (Claassen) Soyedina carolinensis (Claassen) Zapada chila (Ricker) Taeniopterygidae Boltoperla rossi (Frison) Oemopteryx contorta (Needham and Claassen) Strophopteryx appalachia Ricker and Ross S. fasciata (Burmeister) S. inaya Ricker and Ross S. limata (Frison) Taenionema atlanticum Ricker and Ross Taeniopteryx burksi Ricker and Ross T. lita Frison T. lonicera Ricker and Ross T. maura (Pictet) T. metequi Ricker and Ross * T. nelsoni Kondratieff and Kirchner (VA) T. parvula Banks T. robinae Kondratieff and Kirchner * T. ugola Ricker and Ross (GA, TN, VA) Systellognatha Chloroperlidae Alloperla atlantica Baumann A. caudata Frison A. chloris Frison A. furcula Surdick A. nanina (Banks) A. neglecta Frison A. usa Ricker X X X X x# X X X X X x# X X X X X x# X X X X X X X X X X X X X X X X X X X X X X X X X Haploperla brevis (Banks) 36 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Table 2. Continued. Rasvena terna (Frison) Suwallia marginata (Banks) Sweltsa lateralis (Banks) S. mediana (Banks) S. urticae (Ricker) NC SC X X X X X X Peltoperlidae * Peltoperla arcuata Needham (TN, VA) * P. tarteri Stark and Kondratieff (VA) Tallaperla anna (Needham and Smith) X X T. Cornelia (Needham and Smith) X X T. elisa Stark X T. laurie Ricker X X T. maria Needham and Smith X X Viehoperla ada (Needham and Smith) X X Perlidae Acroneuria abnormis (Newman) X X A. arenosa (Pictet) X X A. arida (Hagen) X A. carolinensis (Banks) X X A. evoluta Klapalek X A. filicis Frison X A. frisoni Stark and Brown x# * A. internata (Walker) (GA, VA) A. lycorias (Newman) x# * A. petersi Stark and Gaufin (GA, TN) Agnetina annulipes (Hagen) X * A. capitata (Pictet) (VA) A. flavescens (Walsh) x# X Attaneuria ruralis (Hagen) X X Beloneuria georgiana (Banks) X B. stewarti Stark and Szczytko X X Eccoptura xanthenes (Newman) X X Hansonoperla appalachia Nelson X Neoperla carlsoni Stark and Baumann X * N. catharae Stark and Baumann (TN, VA) N. clymene (Newman) x# N. occipitalis (Pictet) X *N. stewarti Stark and Baumann (TN, VA) Stonefly Records 37 Table 2. Continued. NC sc Paragnetina fumosa (Banks) X X P. ichusa Stark and Szczytko X X P. immarginata (Say) X X P. kansensis (Banks) X X * P. media (Walker) (VA) Perlesta frisoni Banks X X P. nelsoni Stark X X P. placida (Hagen) X Perlinella drymo (Newman) X X P. ephyre (Newman) X X P. zwicki Kondratieff, Kirchner and Stewart X Perlodidae Clioperla clio (Newman) X X Cultus decisus isolatus (Banks) X C. verticalis (Banks) X Diploperla duplicata (Banks) X X D. morgani Kondratieff and Voshell X Helopicus bogaloosa Stark and Ray X X H. subvarians (Banks) X X Hydroperla phormidia Ray and Stark X Isogenoides hansoni (Ricker) X I. varians (Walsh) X Isoperla bellona Banks X I. bilineata (Say) X I. burksi Frison X X* I. cotta Ricker X I. davisi James x# I. dicala Frison x# X I. distincta Nelson X X I. frisoni lilies X I. holochlora (Klapalek) X X I. lata Frison x# I. marlynia Needham and Claassen X I. namata Frison X /. orata Frison X X I. similis (Hagen) X X I. slossonae (Banks) X I. transmarina (Newman) x# Malirekus hastatus (Banks) X X Oconoperla innubila (Needham and Claassen) X X 38 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Table 2. Continued. NC SC Remenus bilobatus (Needham and Claassen) X X Yugus arinus (Frison) X X Y. bulbosus (Frison) X X Pteronarcyidae Pteronarcys biloba (Newman) X X P. dorsata (Say) x# X P. proteus (Newman X X P. scotti (Ricker) X X ACKNOWLEDGMENTS— Vie thank Bill P. Stark of Mississippi College and Richard W. Baumann of Brigham Young University for reviewing the manuscript. Pamela Harrell at Colorado State University, provided editorial assistance. LITERATURE CITED Baumann, R. W. 1974. What is Alloperla imbecilla (Say)? Designation of a neotype, and a new Alloperla from eastern North America (Plecoptera: Chloroperlidae). Proceedings of the Biological Society of Washing- ton 87:257-264. Baumann, R. W. 1979. Nearctic stonefly genera as indicators of ecologi- cal parameters (Plecoptera: Insecta). Great Basin Naturalist 39:241- 244. Fullington, K. E., and K. W. Stewart. 1980. Nymphs of the stonefly ge- nus Taeniopteryx (Plecoptera: Taeniopterygidae) of North America. Journal of the Kansas Entomological Society 53:237-259. Huryn, A. D., and J. B. Wallace. 1987. The exopterygote insect commu- nity of a mountain stream in North Carolina, USA: Life histories, production, and functional structure. Aquatic Insects 9:229-251. Kondratieff, B. C., R. F. Kirchner, and K. W. Stewart. 1988. A review of Perlinella Banks (Plecoptera: Perlidae). Annals of the Entomological Society of America 81:19-27. Kondratieff, B. C., and W. B. Painter. 1986. Two new records of stoneflies (Plecoptera: Perlodidae) from South Carolina. Entomological News 97:17- 20. Lenat, D. R. 1983. Benthic macroinvertebrates of Cane Creek, North Carolina, and comparisons with other southeastern streams. Brimleyana 9:53- 68. Lenat, D. R., and D. L. Penrose. 1987. New distributional records for North Carolina macroinvertebrates. Entomological News 98:67-73. Stonefly Records 39 Morse, J. C., B. P. Stark, and W. P. McCafferty. 1993. Southern Appa- lachian streams at risk: Implications for mayflies, stoneflies, caddisflies and other aquatic biota. Aquatic Conservation: Marine and Freshwa- ter Ecosystems 3:293-303. Ricker, W. E., and H. H. Ross. 1968. North American species of T aeniopteryx (Plecoptera: Insecta). Journal of the Fisheries Research Board of Canada 25:1423-1439. Stanger, J. A., and R.W. Baumann. 1993. A revision of the stonefly ge- nus Taenionema (Plecoptera: Taeniopterygidae). Transactions of the American Entomological Society 119:171-229. Stark, B. P. 1985. Notes on Oconoperla (Plecoptera: Perlodidae). Ento- mological News 96:151-155. Stark, B. P. 1986. The Nearctic species of Agnetina (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 59:437-445. Stark, B. P. 1989. Perlesta placida (Hagen), an eastern nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica 20:263-286. Stark, B. P. 1990. Neoperla clymene revisited: Systematics of the Ne- arctic species complexes (Plecoptera: Perlidae). Pages 299-310 in Mayflies and stoneflies: Life history and biology (I. C. Campbell, editor). Kluwer Academic Publishers. Dordrecht, Holland. Stark, B. P., and L. D. Brown. 1991. What is Acroneuria evoluta Klapalek (Plecoptera: Perlidae)? Aquatic Insects 13: 29-32. Stark, B. P., and K. W. Stewart. 1981. The Nearctic genera of Peltoperlidae (Plecoptera). Journal of the Kansas Entomological Society 54:285-311. Stark, B. P., and S. W. Szczytko. 1982. Egg morphology and phylogeny in Pteronarcyidae (Plecoptera). Annals of the Entomological Society of America 75:519-529. Stark, B. P., S. W. Szczytko, and R. W. Baumann. 1986. North Ameri- can stoneflies (Plecoptera): Systematics, distribution, and taxonomic references. Great Basin Naturalist 46:383-397. Stark, B. P., S. W. Szczytko, and B. C. Kondratieff. 1988. The Cultus decisus complex of eastern North America (Plecoptera: Perlodidae). Proceed- ings of the Entomological Society of Washington 90:91-96. Stewart, K. W., and B. P. Stark. 1988. Nymphs of North America stonefly genera (Plecoptera). Thomas Say Foundation 12:1-460. Surdick, R. F. 1985. Nearctic genera of Chloroperlidae (Plecoptera: Chloroperlidae). Illinois Biological Monograph 54:1-146. Szczytko, S. W., and K. W. Stewart. 1981. Reevaluation of the genus Clioperla. Annals of the Entomological Society of America 74:536- 569. Unzicker, J. D., and V. H. McCaskill. 1982. Chapter 5. Plecoptera. Pages 5.1-5.50 in Aquatic insects and oligochaetes of North and South Carolina. (A. R. Brigham, W. U. Brigham, and A. Gnilka, editors). Midwest Enterprises, Mahomet, Illinois. Zwick, P. 1977. Ergebnisse der Bhutan-Expedition 1972 des naturhistorischen museums in Basel. Entomologica Basiliensia 2:85-134. 40 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat Zwick, P. 1984. Notes on the genus Agnetina ( -Phasganophora ) (Plecoptera Perlidae). Aquatic Insects 6:71-79. Received 9 February 1995 Accepted 20 April 1995 Seasonality in Cetacean Standings Along the Coast of North Carolina Wm. David Webster, P. Dawn Goley, Jessie Pustis, and Joseph F. Gouveia Department of Biological Sciences and Center for Marine Science Research University of North Carolina at Wilmington Wilmington, North Carolina 28403 ABSTRACT — Records of stranding provide an index by which the resident status and local migratory patterns of cetaceans can be ascer- tained, especially along North Carolina’s lengthy coastline, which extends well into the Atlantic Ocean. Stranding records from North Carolina were compiled by month for all cetaceans to test for seasonal trends. Twenty-six cetacean species have stranded, or come ashore intentionally or unintentionally, along the North Carolina coast, 17 of which are year-round residents. The northern right whale ( Eubalaena glacialis), fin whale ( Balaenoptera physalus), humpback whale ( Megaptera novaeangliae ), and harbor porpoise (Phocoena phocoena ) typically strand during the winter and spring months as they migrate along the North Carolina coast. Although stranding records are available for every month, the bottlenose dolphin ( Tursiops turncatus) also strands significantly more frequently in winter and spring, which may be explained, in part, by biases inherent in the use of stranding data. Mammalian diversity in North Carolina exceeds that found in other states and provinces in eastern North America because of the state’s extreme physiographic variability (Webster et al. 1985), and marine mammals clearly exemplify this trend. Many species of marine mammals are year-round residents, but others with subtropical and subarctic affinities, such as the West Indian manatee ( Trichechus manatus Linnaeus) and the harbor porpoise (Phocoena phocoena (Linnaeus)), migrate into inshore and nearshore waters during the summer/fall and winter/spring months, respectively. Some closely related taxa that ostensibly occupy the same niche, such as the long-finned pilot whale ( Globicephala melas (Traill)) and short-finned pilot whale ( G . macrorhynchus Gray), are thought to be latitudinally parapatric along the state’s lengthy (>600 km) coastline, with a dynamic zone of parapatry that shifts relative to the positions of cold-water (Labrador) and warm-water (Gulf Stream) currents. Brimleyana 23:41-51, December 1995 41 42 Wm. David Webster et al. Stranding data can provide a wealth of biological information about marine mammals (Geraci and St. Aubin 1979). Although the cetacean fauna (whales, dolphins, and porpoises) of North Carolina is relatively well known (Caldwell and Golley 1965, Caldwell and Caldwell 1974, Winn et al. 1979, Schmidly 1981, Lee et al. 1983), there has been no attempt to use the state’s stranding records to address the seasonal or distributional ecology of this important component of the marine environment. The purposes of this investigation, therefore, were to describe seasonal periodicity in cetacean strandings in North Carolina and to relate these trends to the zoogeographic significance of North Carolina with regard to the cetacean fauna of the western North Atlantic Ocean. METHODS Cetacean stranding data from North Carolina (Schmidly 1981, and references cited therein; Scientific Event Alert Network Bulletins 1975-1982; J. G. Mead, United States National Museum, personal communication) were compiled by month for each species. These references provided a continuous account of strandings reported from the late- 1800s through 1990; however, most of the records have been accumulated during the last 20 years after the Marine Mammal Stranding Network was established. Stranding records did not always distinguish between live and dead animals, so both were included in our analysis. It was not possible to verify identifications of all specimens associated with these records, especially those of Globicephala and Stenella reported in the Scientific Event Alert Network Bulletins and species of small cetaceans reported in newspapers, because voucher material was some- times not collected (Mead 1977, 1979; Schmidly 1981). Therefore, records were omitted if doubts existed about their veracity. Temporal data were examined statistically (Chi-square) to test the hypothesis that each species exhibited no significant ( P < 0.05) monthly variation in stranding, although sample sizes were small for some species. RESULTS AND DISCUSSION Eight hundred and seventy-two stranding records were available for 26 speices of whales, dolphins, and porpoises (Table 1), nine of which exhibit significant monthly variation in their stranding records. Although the bottlenose dolphin ( Tursiops truncatus (Montagu)) strands in all months of the year, it strands significantly more often in winter and spring. Stranding records for the fin whale ( Balaenoptera physalus (Linnaeus)) and harbor porpoise ( Phocoena phocoena (Linnaeus)) display Cetacean Strandings 43 distinct seasonality, with strandings typically occurring during the winter and spring months. Despite small sample sizes, the northern right whale (Eubalaena glacialis (Miiller)) and humpback whale (Megaptera novaeangliae (Borowski)) also fit into this category. The short-finned pilot whale, long-finned pilot whale, Risso's dolphin ( Grampus griseus G. Cuvier), Atlantic spotted dolphin ( Stenella frontalis G. Cuvier), rough-toothed dolphin ( Steno bredanensis (Lesson)), and dwarf sperm whale ( Kogia simus (Owen)) display significiant monthly variation in stranding with- out exhibiting well-defined seasonal patterns. Not included in Table 1 are Bryde’s whale (Balaenoptera edeni Anderson), blue whale (. B . musculus (Linnaeus)), and short-snouted spinner dolphin ( Stenella clymene (Gray)), species that have stranded to the north and south of North Carolina but not within state boundaries. Also, the pantropical spinner dolphin (, Stenella attenuata (Gray)) was not included because we were unable to verify stranding records in North Carolina. These four species probably inhabit state waters seasonally or as occasional strays (Lee et al. 1983, Webster et al. 1985). Overall, cetaceans strand significantly more frequently during the winter and spring months in North Carolina (Table 1). Several abiotic and biotic factors that are not necessarily related could cause this trend, and examples of each are apparent in these data. Winter storms (known as nor’easters because of the direction from which they blow), coupled with relatively colder water temperatures that slow the process of decomposition, increase the likelihood that a carcass will wash ashore during the winter and spring months. Also, circum- stantial evidence suggests that mortality may be greater for some species during the winter and spring months. Finally, certain species of ceta- ceans are clearly more abundant during the colder months of the year, thus increasing the likelihood of finding stranded animals. Stranding records for the bottlenose dolphin comprise almost 61% of the total number of cetacean strands reported from North Carolina. Bottlenose dolphin strandings increase during the winter and spring months as local neritic populations are augmented by more northerly inshore and pelagic populations (True 1891, Schmidly 1981, Kenney 1990). Increased winter and spring strandings might simply be an artifact of a larger population during those seasons of the year or mortality rates might be greater during the winter months. Significant stranding increases associated with the dolphin die-off of August- October 1987, when the brevetoxin from the dinoflagellate ( Ptychodiscus brevis) weakened dolphins such that they contracted lethal secondary bacterial and fungal infection (Geraci 1989), were clearly evident (Fig. i). Table 1. Monthly frequencies of cetacean strandings along the coast of North Carolina. Significant (P < 0.05) monthly variation is shown with an asterisk for species with 10 or more strands. 44 Wm. David Webster et al a o a o H Q Z o on < < Ph £ 03 tin C/3 o 3 55 -Q 53 2 « 3 * **k» k -4^ >3 C bo 3 3 3 3 r»» K. 3 -4h» 3 3 3 O 00 o N- vo 'Cf 03 3 3 3 3 03 3 k -C O, co od 3 C 3 3 3 3 i-h -4h» Co Lo to 3 3 3 3 3 k §■ * »-k Co Sk 3 O- OO CN VO r- t— i — 4 N- CN oo 00 CO 00 ao 3 >3 3 3 ~3 3 3 C k 3 3 3 C 3 C -3 T3 "C • i—H « -2 3 C/3 bo .. c >4 ^ h0 Oh * CO 3 * *«ki k 3 ■4-, 3 to OV OV — — • — . to cd Cl, Cl, Cl, 3 TO 3 o o —< CO to to ’■k rl 3 3 3 Ci, a,^ ^ ^ N N Total 77 83 154 130 117 35 21 49 53 62 46 45 872 46 Wm. David Webster et al. Tursiops truncatus c n _i < 3 q > Q LL o CC UJ CO MONTH Fig. 1. Monthly stranding records for the bottlenose dolphin ( Tursiops truncatus ) in North Carolina. Significant increases in stranding during the dolphin die- off of 1987 are evident in August, September, and October (*). FALL/WINTER/SPRING MIGRANTS co < 3 a > o u. o oc LU ffl 2 Eubalaena glacialis Balaenoptera acutorostrata Balaenoptera borealis Balaenoptera physalus Megaptera novaeangliae Phocoena phocoena Lagenorhynchus acutus -i — - — i — - — i — ■ — i — i J A S 0 N D MONTH Fig. 2. Monthly stranding records for cetaceans with boreal distributions (see text) that migrate or wander southward along the North Carolina coast during the colder months of the year. Cetacean Strandings 47 Stranding records for several migratory species such as the northern right whale, fin whale, humpback whale, and harbor porpoise indicate when these species are present along the coast of North Carolina (Fig. 2). These North Atlantic taxa migrate southward during the late fall and winter months, and then return northward in the spring, a pattern best demonstrated by the stranding records for the fin whale and hump- back whale. Right whale strandings are confined to the spring months, the season when mothers and their newborn calves migrate northward in shallow nearshore water (Kraus et al. 1986, 1993); the southward winter migration (Reeves et al. 1978) of juveniles, subadults, and adults is farther offshore along the eastern edge of the Labrador Current (Kraus et al. 1993) and the continental shelf (Winn et al. 1986). Stranding records for the harbor porpoise demonstrate a local north-south migra- tion, with the southernmost distributional limits reaching North Carolina during the winter and spring months, rather than the inshore-offshore migratory pattern seen farther north (Neave and Wright 1968, Gaskin et al. 1974, Gaskin and Watson 1985). Based on few stranding records, the sei whale ( Balaenoptera borealis Lesson) and minke whale (B. acutorostrata Lacepede) probably fall into this category as well. These six species account for about 11% of the total number of strandings reported from North Carolina. The common dolphin ( Delphinus delphis Linnaeus), striped dolphin (Stenella coeruleoabla (Meyen)), sperm whale ( Physeter macrocephalus Linnaeus), Blainville’s beaked whale ( Mesoplodon densirostris (Blainville)), and Gervais’ beaked whale ( M . europaeus (Gervais)) have stranded during most months of the year, and we suspect that they are year- round residents in North Carolina waters. The common dolphin inhabits temperate waters adjacent to the 100-fathom isobath where the ocean floor has substantial topographic relief (Leatherwood and Reeves 1983). It does not exhibit a pronounced north-south migration, so increased strandings during the colder months may reflect seasonal inshore-offshore move-ments (Selzer and Payne 1988) or increased mortality during the colder months of the year. Although the sperm whale has a well- documented migration in the North Atlantic Ocean (Townsend 1935), it has stranded in North Carolina in all months except June, October, and December. Mature males migrate northward out of North Carolina waters in the spring, but some immature males and females and their calves remain in North Carolina waters throughout the summer months (Leatherwood et al. 1976). These five species constitute approximately 8% of the total number of strandings reported from North Carolina. Several cetaceans exhibited significant monthly variation in stranding but demonstrated no seasonal periodicity, and each appears to inhabit 48 Wm. David Webster et al. North Carolina waters throughout the year. Monthly variation can be explained by the tendency to mass strand by pilot whales, the rough- toothed dolphin, and possibly Risso’s dolphin. It is difficult to explain significant monthly variation exhibited by the Atlantic spotted dophlin ( Stenella frontalis (G. Cuvier)), which includes stranding records previously attributable to S. plagiodon (Cope), a species once thought to inhabit the Atlantic Ocean. Taxonomic uncertainty in the genus and the difficulty in identifying individuals have been presistent sources of error; however, the recent revision of Stenella in the western North Atlantic Ocean (Perrin et al. 1987) should help alleviate future misidentifications. Significant monthly variation in stranding by Kogia simus might best be explained by a behavior displayed by its close relative, the pygmy sperm whale ( Kogia breviceps (Blainville)). In southeastern North Carolina, we have noticed that strandings of K. breviceps frequently involve females in the process of giving birth or mother-offspring pairs, a behavior also reported by Winn et al. (1979). These seven species account for approximately 17% of the total number of strandings reported from North Carolina. The remaining seven species of cetaceans are relatively rare in North Carolina waters, and scanty stranding records provide little informa- tion about their resident status in the state. The killer whale ( Orcinus orca (Linnaeus)), false killer whale ( Pseudorca crassidens (Owens)), True’s beaked whale ( Mesoplodon mints True), and Cuvier’s beaked whale (Ziphius cavirostris G. Cuvier) are thought to be year-round residents (Leatherwood and Reeves 1983), but the Atlantic white-sided dolphin (Lagenorhynchus acutus (Gray)) inhabits the northern North Atlantic Ocean (Leatherwood and Reeves 1983) and seldom ventures into North Carolina waters. The pygmy killer whale ( Feresa attenuata Gray) and long-snouted spinner dolphin (Stenella longirostris (Gray)) probably enter North Carolina waters during the warmer months of the year (Leatherwood and Reeves 1983). Collectively, this group of species constitutes only about 2% of the total number of strandings reported from North Carolina. Although marine mammal strandings provide a fortuitous source of information on animals that are not typically accessible, there are inherent biases in conclusions derived from stranding data. Neritic species strand more frequently than pelagic species, so stranding fre- quencies are less likely to reflect accurately the abundances of pelagic species. Larger-bodied species and mass strandings are more likely to be reported than small-bodied species or single strandings. Also, the Gulf Stream and Labrador Current could transport dead or dying animals beyond their normal ranges and into North Carolina waters. Cetacean Strandings 49 Conclusions from the North Carolina stranding data, however, agree with information provided by other methods of study for species that are relatively well known. CONCLUSIONS North Carolina has the greatest diversity of cetaceans along the east coast of the United States. Twenty-six species have stranded along the North Carolina coast; four other species might inhabit state waters at least seasonally. Based on stranding records, 17 species appear to be year-round residents, although bull sperm whales leave the area during the warmer months. Seven species with boreal affinities migrate or wander southward into the area during the winter and spring months, and two species with austral affinities migrate northward into the area during the summer and fall months. The status of several species needs additional clarification, and as a matter of protocol, voucher material from deceased marine mammals should always be deposited in museum collections to attain that goal. ACKNOWLEDGMENTS — We thank Dr. James G. Mead for providing us with stranding data from North Carolina. Dave Lee and an anony- mous reviewer provided helpful comments on an earlier draft of the manuscript. Additional support was provided by the Department of Biological Sciences and Center for Marine Science Research (Contri- bution Number 51) of the University of North Carolina at Wilmington. LITERATURE CITED Caldwell, D. K., and M. C. Caldwell. 1966. Observations on the distri- bution, coloration, behavior, and audible sound production of the spotted dolphin, Stenella plagiodon. Los Angeles County Museum, Contribu- tions in Science 104:1-27. Caldwell, D. K., and M. C. Caldwell. 1974. Marine mammals from the southeastern United States coast: Cape Hatteras to Cape Canaveral. Pages 704-772 in A socio-economic environmental baseline summary for the south Atlantic region between Cape Hatteras, North Carolina and Cape Canaveral, Florida. Volume 3. Bureau of Land Management, Contract Number EQ4AC007, Washington, D.C. Caldwell, D. K., and F. G. Golley. 1965. Marine mammals from the coast of Georgia to Cape Hatteras. Journal of the Elisha Mitchell Scien- tific Society 81:24-32. Gaskin, D. E., P. W. Arnold, and B. A. Blair. 1974. Phocoena phocoena. Mammalian Species 42:1-8. Gaskin, D. E., and A. P. Watson. 1985. The harbor propoise, Phocoena 50 Wm. David Webster et al. phocoena, in Fish Harbour, New Brunswick, Canada: Occupancy, distri- bution, and movements. Fisheries Bulletin 83:427-442. Geraci, J. R. 1989. Investigation of the 1987-1988 mass mortality of the bottlenose dolphin. Naval Research News 61(2):2-10. Geraci, J. R., and D. J. St. Aubin. 1979. Biology of marine mammals: insights through strandings. United States Marine Mammal Commis- sion, MMC-77/13, Washington, D.C. Kenney, R. D. 1990. Bottlenose dolphins off the northeastern United States. Pages 369-386 in The bottlenose dolphin (S. Leatherwood and R. R. Reeves, editors). Academic Press, Inc., New York, New York. Kraus, S. D., R. D. Kenney, A. R. Knowlton, and J. N. Ciano. 1993. Endangered right whales of the southwestern North Atlantic. Minerals Manage- ment Service, Contract Number 14-35 0001-30486, Herndon, Virginia. Kraus, S. D., J. H. Prescott, A. R. Knowlton, and S. S. Stone. 1986. Migration and calving in right whales (Eubalaena glacialis) in the western North Atlantic. Pages 139-144 in Right whales: Past and present status (R. L. Brownell, P. B. Best, and J. H. Prescott, editors). Report of the International Whaling Commission, Special Issue 10:1-286. Leatherwood, S., D. K. Caldwell, and H. E. Winn. 1976. Whales, dol- phins, and porpoises of the western North Atlantic. A guide to their identification. National Oceanic and Atmospheric Administration Technical Report, National Marine Fisheries Service CIRC-396, Seattle, Wash- ington. Leatherwood, S., and R. R. Reeves. 1983. The Sierra Club handbook of whales and dolphins. Sierra Club Books, San Francisco, California. Lee, D. S., J. B. Funderburg, Jr., and M. K. Clark. 1983. A distribu- tional survey of North Carolina mammals. Occasional Papers, North Carolina Biological Survey 1982(10):l-70. Mead, J. G. 1977. Records of sei and Bryde’s whales from the Atlantic coast of the United States, the Gulf of Mexico, and the Caribbean. Report of the International Whaling Commission, Special Issue 1:11 3— 116. Mead, J. G. 1979. An analysis of cetacean strandings along the eastern coast of the United States. Pages 54-68 in Biology of marine mam- mals: insights through strandings (J. R. Geraci and D. J. St. Aubin, editors). United States Marine Mammal Commission, MMC-77/13, Washington, D.C. Neave, D. J., and B. S. Wright. 1968. Seasonal migrations of the harbor porpoise (Phocoena phocoena) and other Cetacea in the Bay of Fundy. Journal of Mammalogy 49:259-264. Perrin, W. F., E. D. Mitchell, J. G. Mead, D. K. Caldwell, M. C. Caldwell, P. J. H. Van Bree, and W. H. Dawbin. 1987. Revision of the spot- ted dolphins, Stenella spp. Marine Mammal Science 3:99-170. Reeves, R. R., J. G. Mead, and S. Katona. 1978. The right whale, Eubalaena glacialis, in the western North Atlantic. Report of the International Cetacean Strandings 51 Whaling Commission, 28(SC/29/DOC44):303-312. Schmidly, D. J. 1981. Marine mammals of the southeastern United States coast and the Gulf of Mexico. United States Fish and Wildlife Ser- vice, FWS/OBS-80/41, Washington, D.C. Scientific Event Alert Network Bulletin. 1975-1982. Marine mammals strandings and sightings. Smithsonian Institution, National Museum of Natural History, Volumes 1-7, Washington, D.C. Selzer, L. A., and P. M. Payne. 1988. The distribution of white-sided (Lagenorhynchus acutus ) and common dolphins ( Delphinus delphis ) vs. environmental features of the continental shelf of the northeastern United States. Marine Mammal Science 4:141-153. Townsend, C. H. 1935. The distribution of certain whales as shown by logbook records of American whaleships. Zoologica 19:1-50. True, F. W. 1891. Observations on the life history of the bottlenose porpoise. Proceedings of the United States National Museum 13(1890):197-203. Webster, W. D., J. F. Parnell, and W. C. Biggs, Jr. 1985. Mammals of the Carolinas, Virginia, and Maryland. University of North Carolina Press, Chapel Hill. Winn, H. E., C. A. Price, and P. W. Sorenson. 1986. The distributional biology of the right whale ( Eubalaena glacialis ). Pages 129-138 in Right whales: Past and present status (R. L. Brownell, P. B. Best, and J. H. Prescott, editors). Report of the International Whaling Commission, Special Issue 10:1-286. Winn, L. K., H. E. Winn, D. K. Caldwell, M. C. Caldwell, and J. L. Dunn. 1979. Marine mammals. Chapter 6 in A summary and analysis of environmental information on the Continental Shelf and Blake Pla- teau from Cape Hatteras to Cape Canaveral. Bureau of Land Man- agement, Contract Number AA550-CT7-39, Washington, D.C. Received 9 February 1995 Accepted 28 August 1995 Fishes New or Rare on the Atlantic Seaboard of the United States Fred C. Rohde North Carolina Division of Marine Fisheries 127 Cardinal Drive Extension Wilmington, North Carolina 28405 Steve W. Ross North Carolina National Estuarine Research Reserve 7205 Wrightsville Avenue Wilmington, North Carolina 28403 Sheryan P. Epperly National Marine Fisheries Service Southeast Fisheries Science Center 101 Pivers Island Road Beaufort, North Carolina 28516-9722 AND George H. Burgess Florida Museum of Natural History University of Florida Gainesville, Florida 32611 ABSTRACT — Sampling over the continental shelf of the South At- lantic Bight, especially off North Carolina, continues to produce records of fishes new to or rare in the area. We document the first records for United States continental shelf water (<200 m depth) of seven fish species: Cirrhigaleus asper, Symphysanodon berryi, Pseudocaranx dentex, Lutjanus purpureus, Pristipomoides freemani, Poecilopsetta beani, and Lagocephalus lagocephalus. In addition, we also report on noteworthy collections of our other fishes cap- tured off North Carolina: Synagrops spinosus, Centropristis fuscula, Gonioplectrus hispanus, and Etelis oculatus. Collections along the Atlantic coast of the Carolinas continue to yield fish species never recorded from the area. Most of these new records represent tropical and subtropical species that extend their published ranges from the Caribbean or the eastern Atlantic (Anderson and Gutherz 1964, Burgess et al. 1979, Bohlke and Ross 1981, Ross Brimleyana 23:53-64, December 1995 53 54 Fred C. Rohde et al. et al. 1981). These additions to the ichthyofauna of the Carolinas are a product of increased scientific collecting efforts in a faunistically- rich region that contains diverse habitats and favorable ocean currents. During development of the fifth edition of the American Fisheries Society list of common and scientific names of North American fishes (Robins et al. 1991), we supplied records of several species new to North American waters <200 m deep. These data were cited in Robins et al. (1991) as “Rohde (pers. comm.)” or “Rohde et al. (in press).” Since that manuscript was never published, we herein provide validation for the records cited in the American Fisheries Society list, in addition to documentation of several other noteworthy records. Seven species new to the continental shelf of the United States and four species rare on the Atlantic seaboard are reported. Specimens were collected, often over reef areas, during research cruises using trawls and hook and line, and by intense sampling of the offshore commercial reef fishery where hook and line, and in one instance, a dip net were employed. Museum abbreviations follow Leviton et al. (1985). Squalidae Cirrhigaleus asper (Merrett 1973) The roughskin dogfish is a widespread continental slope species known from the western North Atlantic, southwest Indian, and central Pacific oceans (Compagno 1984). In the Atlantic it has been recorded from the northern Gulf of Mexico (Compagno 1984) and South Carolina (Castro 1983). We note eight additional western North Atlantic records, including one from North Carolina (the northernmost record): UF 37937 (1, 970 mm TL), Atlantic Ocean off North Carolina (32°38'N, 78°14AV) in 201 m, 8 July 1982; UF 44303 (1, 1000 mm TL), Atlantic Ocean off Georgia (30°52'46"N, 79046'11"W) in 374 m, 6 November 1985; UF 47509 (1, 920 mm TL), Straits of Florida south of Big Pine Key in 259 m, 3 June 1987; UF 99624 (1, 968 mm TL), Straits of Florida off Big Pine Key, June 1994; UF 38546 (1, 980 mm TL), Gulf of Mexico off Louisiana (27°42'36"N, 93°14'18"W) in 258 m, 10 August 1983; USNM 217364 (1, 1170 mm TL), Gulf of Mexico off Texas (27°42'N, 94°16'W) in 324 m, 12 May 1973; UF 28535 (1, 1144 mm TL), Campeche Bank, Mexico (21°19'30"N, 92°29'W), in 198- 225 m, 25 August 1980; and UF 47482 (1, 1000 mm TL), Long Bank off Virgin Islands in 183 m, 4 October 1983. The advent of deepwater longline and trap fishing has demonstrated that this fish is widely distributed in waters of 183-457 m in the tropical and subtropical western Atlantic. Generic placement of this New or Rare Fishes 55 species in Cirrhigaleus, rather than Squalus, follows Shirai (1992) and G. H. Burgess (Florida Museum of Natural History, unpublished data). Acropomatidae Symphysanodon berryi Anderson 1970 The first United States record of the slope bass (UF 38899, 61.3 mm SL) was collected by trawl at 35°07'N, 75°07'W (R/V Albatross IV 82-11, station 23) at a depth of 101-256 m on 16 September 1982. Anderson (1970) reported the species from throughout the Caribbean and the Bahamas in 220-476 m. Although Symphysanodon is usually listed as a lutjanid, there is evidence against this placement. It was considered incertae sedis but possibly related to the Acropomatidedae (especially Synagrops) by Johnson (1984), and its taxonomic status is still uncertain (Johnson 1993). Although Eschmeyer (1990) considered it to be in its own family, Symphysanodontidae, Nelson (1994) provisionally placed it in the Acropomatidae. Synagrops spinosus Schultz 1940 The keelcheek bass has been collected extensively in the Gulf of Mexico in depths >60 m (Hoese and Moore 1977, Potts and Ramsey 1987, Mochizuki and Gultneth 1989, Boschung 1992), and is also known from off Suriname and other Caribbean locations (Fujii 1983, Mochizuki and Gultneth 1989), and from scattered locations in the West Atlantic and West Pacific (Mochizuki and Gultneth 1989). Records of this fish, however, are rare along the United States Atlantic east coast. The following trawl collections document the occurrence of S. spinosus on the other continental shelf (<200 m) of the South Atlantic Bight: UF 41747 (1, 68 mm SL), 35°47'N, 74°53'W in 78 m, 14 July 1980; GMBL-74-92 (1, 61 mm SL), 35°10'N, 75°03AV in 221- 229 m, 8 May 1974; GMBL-74-58 (3), 35°02'N, 75°1UW in 238- 256 m, 8 May 1974; UF 40027 (1, 114 mm SL), 34°52'N, 75°27'W (Silver Bay station 1283) in 179 m, 17 September 1959 (Bullis and Thompson 1965); UF 39781 (1, 26 mm SL), 34°4UN, 75°37AV (Delaware II 83-5, station 313) in 154 m, 14 May 1983; UF 39898 (1, 73 mm SL), 34°36'N, 75°39'W (Delaware II 83-5, station 315) in 227 m, 15 May 1983; UF 39853 (3, 98, 99, 106 mm SL), 34°18'N, 75°50'W (Delaware II 83-5, station 317) in 379-402 m, 15 May 1983; UF 39816 (4, 55, 49, 49, 48 mm SL), 34°07'N, 76°09'W (Delaware II 83-5, station 323) in 155 m, 16 May 1983; UF 41084 (2, 100, 101 mm SL), 29°49.6'N, 80°10.8'W in 318 m, 29 May 1984; UF 41229 (4, 96, 102, 104, 108 mm SL), 28°40.6'N, 79°53.8AV in 320 m, 31 56 Fred C. Rohde et al. May 1984; UF 41246 (1, 113 mm SL), 28°29.8'N, 79°50.1'W in 366 m, 31 May 1984. In addition to the above S. spinosus from Silver Bay station 1283, Bullis and Thompson (1965) listed eight other collections (not cited previously) of this species between South Carolina and Cape Canaveral, Florida (82-366 m). Wenner et al. (1979fr, c; 1980) also listed several collections of S. spinosus from northern Florida to South Carolina (128-338 m). The keelcheek bass seems to be common on the outer continental shelf and upper slope from North Carolina through the Gulf of Mexico and western Caribbean. Many previous records were probably confused with the co-occurring congener S. bellus (Goode and Bean). Both species are often collected together, and the most obvious differences between them are the serrations on the anterior edges of the pelvic spines and the second spines of the first dorsal and anal fins of S. spinosus. The genus Synagrops has been variously placed in the families Apogonidae (Cheilodipteridae) and Percichthyidae (Fraser 1972), but is currently placed in the “oceanic percichthyids,” family Acropomatidae (Johnson 1984). Serranidae Centropristis fuscula (Poey 1861) Four individuals (UF 44997, 50 mm SL, 68 mm SL, 107 mm SL gravid female, 117 mm SL gravid female) of the rare twospot sea bass were taken in a single trawl catch at 33°16'N, 77°13AV (Delaware II 82-04, station 153) at a depth of 97-126 m on 9 July 1982. This trawl appeared to have been pulled mostly over very rough bottom as evidenced by severe net damage and captures of reef organisms (e.g., soft corals). An additional specimen, the largest yet reported, (UF 100391, 168 mm SL gravid female) was captured by hook and line at 32°47"N, 78°11'W at a depth of 165 m on 15 July 1995. These five specimens significantly increase the total known specimens and extend the range northward. Previous records of C. fuscula were from Cuba (holotype MCZ 10015, 138 mm SL (Poey 1861); ANSP 94422, 135 mm (Robins and Starck 1961)), Puerto Rico in 183 m (ANSP 144592, 155 mm SL), Gulf of Mexico (1 specimen, G. D. Johnson, United States National Museum, personal communication), and South Carolina (2 collections, listed as Centropristis sp. by Wenner et al. (1979a)). The general rarity of specimens and the bottom type of our collection suggest that this species is a cryptic reef fish. Gonioplectrus hispanus (Cuvier 1828) The Spanish flag, usually considered a Caribbean insular species, New or Rare Fishes 57 has been recorded infrequently from scattered locations in the Gulf of Mexico (Bullock and Smith 1991, Boschung 1992) and is also known from the Bahamas through the Caribbean to Brazil (Bullock and Smith 1991, Heemstra and Randall 1993). Until recently, the only record of Spanish flag outside the above distribution was of a single, pelagic larva collected off Cape Fear, North Carolina (Kendall and Fahay 1979). Intensive sampling of the Carolinian snapper/grouper commercial fishery has yielded the following adult specimens, all collected by hook and line over hard bottoms: UF 45042 (208 mm SL), 33°53'N, 76°35'W in 101 m, 1 July 1987; specimen lost (247 mm TL), 33°31.3'N, 76°56.5AV in 40 m, 15 June 1988; specimen sold (250 mm TL), southern Onslow Bay, North Carolina in 46 m, September 1990; UF98891 (gravid female, 182 mm SL), northern Long Bay, North Carolina, 13 November 1991; specimen released, northern Long Bay in 40 m, January 1993; specimen sold (230 mm TL), northern Long Bay in 30 m, July 1993; UF 98892 (gravid female, 180 mm SL), southern Onslow Bay, 15 November 1993; specimen sold (220 mm TL), northern Long Bay in 36 m, January 1994. This species occurs regularly on hard bottoms of the Carolinian outer continental shelf, and the occurrence of both adults (two in spawning condition) and larvae indicates that a reproducing population exists in the South Atlantic Bight. Carangidae Pseudocaranx dentex (Bloch and Schneider 1801) The circumglobal, antitropical range of the white trevally includes the western Indian Ocean, the Indo-West Pacific, the Mediterranean Sea, the eastern Atlantic, mid-Atlantic islands, southern Brazil, and Bermuda (Smith-Vaniz 1984). The first record (UF 42779, 565 mm FL, 526 mm SL) from the United States continental shelf was taken with hook and line off the Carolinas (33°14'N, 77°16'W) on 19 February 1985 in 91 m. Two other large specimens (both marketed) were taken by hook and line: one on 16 November 1985 (835 mm FL) near 33°15 N, 77°24'W in 46-55 m and one on 3 February 1986 (802 mm FL) near 33°06'N, 77°55'W in 49 m. A fourth specimen (ANSP 159577, 785 mm FL) was collected by hook and line in 88 m between 30 July and 5 August 1987 at 33°16.5'N, 77°15'W. One large P. dentex was captured (hook and line) and released off the Cape Fear, North Carolina area in 42 m on 3 August 1989. Several of the commercial fishermen recognized this species as different (referring to it as “guelly jack”) and, in addition to the above specimens, they had records of other catches of it from similar areas off Cape Fear. The fisherman who produced the first specimen (above) reported that he had seen P. dentex 58 Fred C. Rohde et al. before, near Matanilla Shoal (northwest end of the Little Bahama Bank). Lutjanidae Etelis oculatus (Valenciennes 1828) A single adult specimen of the queen snapper (UF 42778, 673 mm SL) was collected with hook and line 135 km south of Southport, North Carolina at a depth of 201 m on 11 March 1985. Two other adults were landed (not saved) from northern Long Bay, North Carolina: one (915 mm TL) from 219 m in April 1989 and one (685 mm TL) from 164 m in March 1993. These are the first adults of E. oculatus recorded north of Florida. Two other small juvenile specimens are known from off the Carolinas (44 mm SL (Anderson and Fourmanoir 1975) and 30 mm FL, South Carolina Marine Resources Monitoring, Assessment, and Prediction Program collections, 33°02.7'N, 77°55.5'W, 59 m, 3 September 1976). Etelis oculatus ranges in the western Atlantic from North Carolina and Bermuda south through the Gulf of Mexico (Burgess and Branstetter 1985), the Bahamas, the West Indies, and the Caribbean to Brazil (Anderson 1981, Allen 1985). Lutjanus purpureus Poey 1867 The Caribbean red snapper, mainly a continental shelf species, was previously known only from the Caribbean (Yucatan and Cuba) south through the Antilles to northeastern Brazil (Allen 1985). There has been some question whether it may be synonymous with L. campechanus (Poey) (Vergara R. 1978). Our records demonstrate that these two species of red snappers are sympatric at least through the South Atlantic Bight. The South Carolina Marine Resources Monitoring, Assessment, and Prediction Program has collected three specimens by trawl: 320 mm FL, 34°36.4'N, 76°12.8'W in 35 m, 4 May 1974; 30 mm FL, 30°49.7'N, 81°10.7'W in 13 m, 18 August 1974; 40 mm FL, 30°22'N, 81°18.7'W in 12 m, 18 August 1974. The following records of large adults were obtained from the commercial snapper/grouper fishery: 615 mm TL (specimen photographed but lost), 33°31.3'N, 76°56.5'W in 64 m, 15 June 1988; 620 mm TL (specimen sold), northern Long Bay, NC in 54 m, January 1989; 540 mm TL (specimen sold), southern Onslow Bay, NC in 38 m, June 1989; 630 mm TL (specimen sold), southern Onslow Bay in 42 m, August 1989; 610 mm TL (specimen sold), southern Onslow Bay in 42 m, May 1990. The commercial fishermen generally did not recognize that these fish were different from L. campechanus ; however, we verified the identifications of the specimens marketed at the fish houses. We distinguished these two species of Lutjanus by lateral line scale counts and relative body depths New or Rare Fishes 59 (Vergara R. 1978). Pristipomoides freemani Anderson 1966 Only two specimens of the yelloweye wenchman have been collected along the continental shelf of North America. The first specimen (GMBL 78-145, 85.7 mm SL) was collected by trawl off the east coast of Florida at 28°58.4'N, 80°04.4AV in 121-113 m on 18 September 1978 (R/V Dolphin DP 78-07) (W. D. Anderson, Jr., Grice Marine Biology Laboratory, personal communication). The second P. freemani (GMBL 82-197, 82 mm SL) was collected by trawl at the same North Carolina station as the previously discussed C. fuscula (33°16'N, 77°13'W, R/V Delaware II 82-04, station 153) at a depth of 99 m on 9 July 1982. Additionally, Leis and Lee (1994) reported a single larva from off the Florida Keys questionably attributed to this species. The yellow- eye wenchman was previously known from Uruguay to Panama and Barbados (Anderson 1966, 1972; Matsuura 1983) and off Bermuda (60.3 mm SL; 32°09'N, 64°1UW; 24 August 1971; W. D. Anderson, Jr., personal communication). Pleuronectidae Poecilopsetta beani (Goode 1881) This small flounder, called deep-water dab or offshore flounder (Bigelow and Schroeder 1953, Potts and Ramsey 1987), has been reported from water >200 m deep along the United States continental slope from off New York through the northern Gulf of Mexico to Campeche (Goode and Bean 1896, Tyler 1960). Its distribution south of Mexico has been inconsistently reported: from off northern Colombia and St. Kitts, Lesser Antilles (Goode and Bean 1896), possibly to the greater Antilles (Tyler 1960), and from off northern Brazil (Topp and Hoff 1972). In addition to the southernmost Brazilian record, Bullis and Thompson (1965) list several collections from deep water along the Central American coast. We have two records of P. beani from off North Carolina collected by trawl during the same cruise (Delaware II 83-05). The first specimen (UF 39809, 18 mm SL) represents the first report of this species shallower than 200 m and was collected at 34°07'N, 76°09'W in 155 m (station 323) on 16 May 1983. The second specimen (UF 39891, 56 mm SL) was collected at 34°36'N, 75°39'W in 227 m (station 315) on 16 May 1983. Two larval specimens (MCZ 78481 and 78491, Poecilopsetta sp.), probably P. beani , have also been collected off North Carolina. This species was not included in the most recent American Fisheries Society list of common and scientific names of North American fishes (Robins et al. 1991), but 60 Fred C. Rohde et al. based on the above record (UF 39809), it should be added to the continental shelf fauna. Tetraodontidae Lagocephalus lagocephalus (Linnaeus 1758) The first United States oceanic puffers (UF 44194, 169 mm SL; 190 mm SL, specimen mounted) were collected at 34°21.5'N, 75°55'W in 64 m on 6 August 1985. They were dipnetted at night from a school of 6 to 8 individuals swimming at the surface. Lagocephalus lagocephalus is widespread, ranging through the eastern Atlantic, the Mediterranean Sea, and the Pacific and Indian oceans (Shipp 1974). Templeman (1962) reported the first North American occurrence of the species from a single individual collected in Newfoundland. Other isolated western Atlantic records of the species include Bermuda, the Gulf Stream off Florida, and Curacao (Shipp 1974). DISCUSSION The offshore (>20 m) ichthyofauna of North Carolina, particularly on hard bottoms, is dominated (species numbers) by tropical and subtropical species. Expanded sampling of the offshore hard bottom and outer shelf habitats continues to increase the number of these forms known off the Carolinas. Although the Gulf Stream undoubtedly helps dis- perse tropical organisms into the area, many of these southern species apparently maintain self-sustaining populations on North Carolina’s middle to outer continental shelf (Grimes et al. 1977, Grimes and Huntsman 1980, S. W. Ross, North Carolina National Estuarine Research Reserve, unpublished data). Burgess et al. (1994), in fact, proposed that a redefined tropical West Indian zoogeographic province should include the reefs of the outer continental shelf of the South Atlantic Bight to Cape Hatteras. The 11 species documented herein are most common in warm-temperate to tropical waters along the outer shelf or upper slope south of North Carolina. Three of these (C. fuscula, G. hispanus, L. purpureus ) tend to be benthic and tied to reef-like habitats. The remainder are either pelagic or benthopelagic, and are capable of extensive movements. Briggs (1974) noted that, within the warm-temperate Carolinian Region, the northern Gulf of Mexico contained a richer fish fauna (375-400 species) than the Atlantic coast. Since his publication, many new records have been added to both areas. Hoese and Moore (1977) reported over 400 fishes from the northern Gulf of Mexico, and Boschung (1992) listed around 663 marine fishes from the eastcentral Gulf of Mexico. Dahlberg (1975) reported nearly 400 species in and near Georgia New or Rare Fishes 61 coastal waters. With the additions reported herein, the North Carolina marine ichthyofauna in less than 200 m contains over 680 species (S. W. Ross and G. H. Burgess, unpublished manuscript). However, many of these are cold-temperate species that rarely range south of Cape Hatteras, and thus, are not permanent members of the Carolinian Region. The North Carolina ichthyofauna is much richer than previously reported, and future zoogeographic and systematic data will likely prove that the northern Gulf of Mexico and the South Atlantic Bight are not significantly different in fish species richness. ACKNOWLEDGMENTS — We thank the American Fish Company, Davis Fish Company, Ottis Fish House, Sea Coast Seafood, Seafood Source, Southport Fish Company and Scott Blessing, Kenny Brennan, V. P. Brinson, Scott Every, Milton Mathis, and Milton Mullerweiss for their cooperation and donated specimens. We appreciate the assistance of William D. Anderson, Jr., G. David Johnson, William J. Richards, C. Richard Robins, and William Smith-Vaniz in confirming the identity of specimens or providing information on distributions. We thank the South Carolina Marine Resources Monitoring, Assessment, and Prediction Program for allowing us to use their data on the lutjanids and Karsten E. Hartel for providing data from the Museum of Comparative Zoology. LITERATURE CITED Allen, G. R. 1985. Food and Agriculture Organization species catalogue. Volume 6. Snappers of the world. An annotated and illustrated cata- logue of lutjanid species known to date. Food and Agriculture Orga- nization Fisheries Synopsis (125). Anderson, W. D., Jr. 1966. A new species of Pristipomoides (Pisces: Lutjanidae) from the tropical western Atlantic. Bulletin of Marine Science 16:81 4 — 826. Anderson, W. D., Jr. 1970. Revision of the genus Symphysanodon (Pi- sces: Lutjanidae) with descriptions of four new species. Fisheries Bulletin (United States) 68:325-346. Anderson, W. D., Jr. 1972. Notes on western Atlantic lutjanid fishes of the genera Pristipomoides and Etelis. Copeia 1972:359-362. Anderson, W. D., Jr. 1981. A new species of Indo-West Pacific Etelis (Pisces: Lutjanidae), with comments on other species of the genus. Copeia 1981:820-825. Anderson, W. D., Jr., and P. Fourmanoir. 1975. The status of Erythrobussothen gracilis, a percoid fish. Copeia 1975:181-182. Anderson, W. D., Jr., and E. J. Gutherz. 1964. New Atlantic coast ranges for fishes. Quarterly Journal of the Florida Academy of Science 27:299- 306. 62 Fred C. Rohde et al. Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the Gulf of Maine. Fisheries Bulletin (United States) 53:1-577. Bohlke, E. B., and S. W. Ross. 1981. The occurrence of Muraena ro- busta Osorio (Anguilliformes, Muraenidae) in the west Atlantic. Northeast Gulf Science 42:123-125. Boschung, H. T. 1992. Catalogue of freshwater and marine fishes of Alabama. Bulletin of the Alabama Museum of Natural History Number 14. Briggs, J. C. 1974. Marine zoogeography. McGraw-Hill, New York, New York. Bullis, H. R., Jr., and J. R. Thompson. 1965. Collections by the explor- atory fishing vessels Oregon , Silver Bay, Combat, and Pelican made during 1956-1960 in the southwestern North Atlantic. United States Fish Wildlife Service, Special Scientific Report-Fisheries Number 510. Bullock, L. H., and G. B. Smith. 1991. Seabasses (Pisces: Serranidae). Memoirs of the Hourglass Cruises 8 (part 2): 1-243. Burgess, G. H., and S. Branstetter. 1985. Status of Neoscopelus (Neoscopelidae) in the Gulf of Mexico with distributional notes on Caulolatilus chrysops (Branchiostegidae) and Etelis oculatus (Lutjanidae). Northeast Gulf Science 7(2): 157-162. Burgess, G. H., G. W. Link, Jr., and S. W. Ross. 1979. Additional ma- rine fishes new or rare to Carolina waters. Northeast Gulf Science 3:74-87. Burgess, G. H., S. H. Smith, and E. D. Lane. 1994. Fishes of the Cay- man Islands. Pages 199-228 in The Cayman Islands: Natural history and biogeography (M. A. Brunt and J. E. Davies, editors). Kluwer Academic Publishers, Netherlands. Castro, J. I. 1983. The sharks of North American waters. Texas A&M University Press, College Station. Compagno, L. J. V. 1984. Food and Agriculture Organization species catalogue. Volume 4. Sharks of the world. Part 1. Hexanchiformes to Laminformes. Food and Agriculture Organization Fisheries Synopsis 125(4): 1-249. Dahlberg, M. D. 1975. Guide to coastal fishes of Georgia and nearby states. University of Georgia Press, Athens. Eschmeyer, W. N. 1990. Catalog of the genera of recent fishes. Califor- nia Academy of Sciences, San Francisco. Fraser, T. H. 1972. Comparative osteology of the shallow water cardinal fishes [Perciformes: Apogonidae] with reference to the systematics and evolution of the family. Ichthyological Bulletin J.L.B. Smith Institute Ichthyology Number 34:1-105. Fujii, E. 1983. Synagrops spinosus Schultz, 1940. Page 297 in Fishes trawled off Suriname and French Guiana (T. Uyeno, K. Matsuura, and E. Fujii, editors). Japan Marine Fisheries Resource Research Center, Tokyo. Goode, G. B., and T. H. Bean. 1896. Oceanic ichthyology. United States National Museum Special Bulletin. Grimes, C. B., and G. R. Huntsman. 1980. Reproductive biology of the vermilion snapper, Rhomboplites aurorubens, from North Carolina and New or Rare Fishes 63 South Carolina. Fisheries Bulletin (United States) 78:137-146. Grimes, C. B., C. S. Manooch, III, G. R. Huntsman, and R. L. Dixon. 1977. Red snappers of the Carolina coast. Marine Fishery Review 39:12- 15. Heemstra, P. C., and J. E. Randall. 1993. Food and Agriculture Organi- zation species catalogue. Volume 16. Groupers of the world (Family Serranidae, Subfamily Epinephelinae). An annotated and illustrated catalogue of the grouper, rockcod, hind, coral grouper, and lyretail species known to date. Food and Agriculture Organization Fisheries Synopsis 125(16). Hoese, H. D., and R. H. Moore. 1977. Fishes of the Gulf of Mexico. Texas A&M University Press, College Station. Johnson, G. D. 1984. Percoidei: Development and relationships. Pages 464-498 in Ontogeny and systematics of fishes (H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., and S. L. Richardson, editors). American Society of Ichthyologists and Herpe- tologists, Special Publication 1. Johnson, G. D. 1993. Percomorph phylogeny: Progress and problems. Bulletin of Marine Science 52:3-28. Kendall, A. W., Jr., and M. P. Fahay. 1979. Larva of the serranid fish Gonioplectrus hispanus with comments on its relationships. Bulletin of Marine Science 29:117-121. Leis, J. M., and K. Lee. 1994. Larval development in the lutjanid sub- family Etelinae (Pisces): The genera Aphareus, Aprion, Etelis and Pristi- pomoides. Bulletin of Marine Science 55:46-125. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Matsuura, K. 1983. Pristipomoides freemani Anderson, 1966. Page 343 in Fishes trawled off Suriname and French Guiana (T. Uyeno, K. Matsuura, and E. Fujii, editors). Japan Marine Fishery Resource Research Cen- ter, Tokyo. Mochizuki, K., and S. Gultneth. 1989. Redescription of Synagrops spinosus (Percichthyidae) with its first record from the West Pacific. Japanese Journal of Ichthyology 35(4):421-427. Nelson, J. S. 1994. Fishes of the world. Third Edition. John Wiley & Sons, Inc., New York, New York. Poey, F. 1861. Memorias sobre la historia natural de la Isla de Cuba, acompanadas de sumarios lations y extractos en Frances. Habana 1:40- 200. Potts, D. T., and J. S. Ramsey. 1987. A preliminary guide to demersal fishes of the Gulf of Mexico continental slope (100 to 600 fathoms). Alabama Sea Grant Extension Service Publication MASGP-86-009. Robins., C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. Fifth Edition. Ameri- 64 Fred C. Rohde et al. can Fisheries Society Special Publication 20. Robins, C. R., and W. A. Starck, II. 1961. Materials for a revision of Serranus and related fish genera. Proceedings of the Academy of Natural Sciences of Philadelphia 113:259-314. Ross, S. W., G. W. Link, Jr., and K. A. MacPherson. 1981. New records of marine fishes from the Carolinas, with notes on additional spe- cies. Brimleyana 6:61-72. Shipp, R. L. 1974. The pufferfishes (Tetraodontidae) of the Atlantic Ocean. Publication of the Gulf Coast Research Laboratory 4. Shirai, S. 1992. Squalean phylogeny. Hokkaido University Press, Hokkaido, Japan. Smith-Vaniz, W. 1984. Carangidae. in Volume I. Food and Agriculture Organization species identification sheets for fishery purposes, west- ern Indian Ocean (Fishing Area 51) (W. Fischer and G. Bianchi, editors). Food and Agriculture Organization of the United Nations, Rome. Templeman, W. 1962. Record of the oceanic puffer, Lagocephalus lagocephalus (L.), family Tetraodontidae, from Newfoundland. Journal of the Fishery Research Board of Canada 19:811-814. Topp, R. W., and F. H. Hoff, Jr. 1972. Flatfishes (Pleuronectiformes). Memoirs of the Hourglass Cruises 4 (Part 2): 1-135. Tyler, J. C. 1960. Note on the flatfishes of the genus Poecilopsetta oc- curring in Atlantic waters. Stanford Ichthyological Bulletin 7:126-131. Vergara R., R. 1978. Lutjanidae. in Volume III. Food and Agriculture Organization species identification sheets for fishery purposes, west- ern central Atlantic (Fishing Area 31) (W. Fischer, editor). Food and Agriculture Organization of the United Nations, Rome. Wenner, C. A., C. A. Barans, B. W. Stender, and F. H. Berry. 1979 a. Results of the MARMAP otter trawl investigations in the South At- lantic Bight. I. Fall, 1973. South Carolina Marine Resources Center Technical Report Number 33. Wenner, C. A., C. A. Barans, B. W. Stender, and F. H. Berry. 1979b. Results of the MARMAP otter trawl investigations in the South At- lantic Bight. III. Summer, 1974. South Carolina Marine Resources Center Technical Report Number 41. Wenner, C. A., C. A. Barans, B. W. Stender, and F. H. Berry. 1979c. Results of the MARMAP otter trawl investigations in the South At- lantic Bight. IV. Winter-early spring, 1975. South Carolina Marine Resources Center Technical Report Number 44. Wenner, C. A., C. A. Barans, B. W. Stender, and F. H. Berry. 1980. Results of the MARMAP otter trawl investigations in the South Atlantic Bight. V. Summer, 1975. South Carolina Marine Resources Center Techni- cal Report Number 45. Received 8 March 1995 Accepted 29 August 1995 A New Species of Crayfish of the Genus Orconectes, Subgenus Procericambarus (Decapoda: Cambaridae), Endemic to the Neuse and Tar-Pamlico River Basins, North Carolina John E. Cooper1 North Carolina State Museum of Natural Sciences P.O. Box 29555 Raleigh, North Carolina 27626 AND Martha Riser Cooper 209 Lynwood Lane Raleigh, North Carolina 27609 ABSTRACT — Orconectes {Procericambarus) carolinensis is a new species in the Spinosus Group, and the only member of its subge- nus known from east of the Blue Ridge physiographic province. It is endemic to the Neuse and Tar-Pamlico river basins of North Carolina, where it occurs in the Coastal Plain and the eastern edge of the Piedmont Plateau. It is most closely related to O. ( P .) spinosus and O. (P.) putnami, but may be distinguished from these and other members of the Spinosus Group by: the greater length of the ter- minal elements of the form I male gonopod (first pleopod); the shorter and broader areola; the somewhat longer acumen and ros- trum; the occasional presence of multiple cervical spines and small spiniform hepatic tubercles; the smaller size; and various aspects of tuberculation and spination of the palm, dactyl, merus, and is- chium of the cheliped. The species is probably derived from an ancestral Procericambarus stock that inhabited the Tennessee and Teays river basins, gained access to the Atlantic versant follow- ing a breach of the Blue Ridge by headwaters of the young Roanoke River, and later entered the Greater Pamlico River from the Greater Roanoke by stream captures. The Neuse and Tar-Pamlico rivers rise in the eastern Piedmont Plateau of North Carolina, flow southeast across the Coastal Plain of the state, and debouch at saline estuaries of Pamlico Sound on the Atlantic coast. Throughout their lengths they are parallel ripar- ian systems that support nearly identical faunas and, although now separated, during much of the Pleistocene they undoubtedly were a 1 Permanent address: 418 Wayne Drive, Raleigh, North Carolina 27608. Brimleyana 23:65-87, December 1995 65 66 John E. Cooper and Martha Riser Cooper single hydologic unit, the Greater Pamlico River (Lachner and Jenkins 1971:62). Particularly notable in the Neuse and Tar-Pamlico faunas are a number of disjunct endemic species that exhibit closest affinities with congeners whose ranges lie well beyond the two North Carolina rivers. The endemic fish, Noturus ( Rabida ) furiosus Jordan and Meek, is widespread in both rivers, as are its more broadly distributed con- geners, Noturus { Schilbeodes ) gyrinus (Mitchill) and Noturus (, Schilbeodes ) insignis (Richardson). As interpreted by Taylor (1969), however, the closest relatives of N. furiosus are Noturus {Rabida) munitus Suttkus and Taylor, Noturus {Rabida) placidus Taylor, and Noturus {Rabida) stigmosus Taylor, which, along with N. furiosus, comprise the “ furiosus group.” Geographically, the nearest of the three western relatives of N. furiosus is N. munitus, which occurs in the Conasauga River of southeastern Tennessee and in the Pearl, Tombig- bee, and Cahaba rivers of Alabama, Louisiana, and Mississippi (see Rohde 1980 for distribution maps). The large, branchiate salamander, Necturus lewisi (Brimley), is also endemic and widespread in the Neuse and Tar-Pamlico rivers (Braswell and Ashton 1985). Its closest relative is considered to be the sympatric but wider ranging Necturus punctatus (Gibbes) (Ashton et al. 1980; Sessions and Wiley 1985), which inhabits the Atlantic Coastal Plain from southeastern Virginia to central Georgia. The ranges of these two species, although overlapping in the Neuse and Tar- Pamlico basins, are broadly disjunct from those of the other Necturus species. Necturus maculosus (Rafinesque) is the only other member of the genus that occurs in North Carolina, where it appears to be limited to the French Broad River basin in the Blue Ridge (Martof et al. 1980:50). The ranges of several invertebrates exhibit the same phenomenon. The unionid mollusk, Elliptio {Canthyria) steinstansana Johnson and Clarke, is endemic to the Tar-Pamlico River, but its nearest relatives occur in the James River basin to the north and the Altamaha River basin to the south. Among the eight species of crayfishes known to occur in the Neuse and Tar-Pamlico rivers (Cooper and Ashton 1985:9- 10) are two endemic species, Procambarus {Ortmannicus) medialis Hobbs, and the species of Orconectes described herein. Procambarus medialis is most closely related to two other Coastal Plain members of the Planirostris Group in North Carolina: Procambarus {Ortmannicus) plumimanus Hobbs and Walton, of the Northeast Cape Fear and New (White Oak) basins (Cooper and Braswell 1995), and Procambarus {Ortmannicus) pearsci (Creaser), of the Cape Fear, Waccamaw, and New Crayfish Species 67 Lumber-Little Pee Dee basins. These three species of very limited distribution constitute a ‘‘disjunct enclave” whose nearest relatives — Procambcirus ( Ortmannicus ) hybus Hobbs and Walton, Procambarus ( Ortmannicus ) mancus Hobbs and Walton, and Procambarus ( Ortman- nicus) planirostris Penn — inhabit elements of the Gulf drainage in Alabama, Mississippi, and the Florida Parishes of Louisiana (Hobbs and Walton 1958:11; Hobbs 1975:15). The new species of Orconectes described below is yet another Neuse and Tar-Pamlico endemic whose range is widely disjunct from the ranges of its closest relatives. It “is only the third, and southern- most, species of Orconectes recorded from anywhere on the Atlantic seaboard” (Cooper and Cooper 1977:199), and has its closest affinities with species found in and west of the Blue Ridge rather than with its geographically close congener, Orconectes ( Crockerinus ) virginiensis Hobbs, of southeastern Virginia and northeastern North Carolina. First reports of an Orconectes in the Tar-Pamlico River appeared in the late 19th and early 20th centuries, but the animal was assigned to Bundy’s Cambarus (= Orconectes) spinosus (Bundy 1877; Faxon 1884, 1890; Harris 1903; Ortmann 1905, 1931), a creature of the Ten- nessee and Coosa river basins. Bundy’s specimens, which came from the Tar River at Rocky Mount, Edgecombe County, apparently were the only ones on which the several subsequent reports were based. Later papers (e.g., Hobbs 1972, 1974, 1989; Hobbs and Peters 1977) did not comment on an Orconectes in this Atlantic drainage river, although Hobbs (1981:294) repeated Ortmann’s (1931) statements (based on Faxon) that “C. spinosus ” occurred in the Tar River. Over 200 specimens of this crayfish, from 43 localities, have now been collected. Examination of this material verifies our opinion that the animal is quite distinct from other members of the genus Orconectes. Orconectes ( Procericambarus ) carolinensis, new species Figure 1 Cambarus spinosus Bundy. — Faxon, 1890:632 [p.p.: “Tar River Rocky Mount, North Carolina.”]. — Harris, 1903:180 [p.p.: “North Carolina. 1. Tar River, Rocky Mount (Nash County). F., ’90”]. — Ortmann, 1905:115 [p.p.: “Atlantic drainage ... North Carolina”]. — Hobbs, 1981:294 [p.p.: quoting Ortmann 1905]. C. spinosus Bundy. — Ortmann, 1931:87 [p.p.: “Tar River, Rocky Mount, Edgecombe Co., North Carolina, according to Faxon”], 88 [p.p.: “(from ... North ... Carolina)” ...]. — Hobbs, 1981:294 [p.p.: quoting Ortmann, 1931:87]. 68 John E. Cooper and Martha Riser Cooper “recently discovered species of Orconectes — Cooper and Cooper, 1977:199. Orconectes sp. A. — Cooper and Ashton, 1985:9. — Cooper and Braswell, 1995:88, 89, 123, 126. Diagnosis — Body and eyes pigmented, eyes large. Rostrum acarinate, deeply excavate, with conspicuous acumen, delimited basally on each side by strong, acute marginal spine; margins of rostrum thickened, elevated, parallel from base to marginal spines; acumen 42.6 to 57.3 (x = 50.1, n = 113) percent length of rostrum. Areola 3.0 to 5.8 (* = 4.2, n = 112) times longer than wide, constituting 25.5 to 31.1 (x = 28.5, n = 113) percent of total carapace length (TCL) and 39.0 to 43.3 (x = 41.2, n = 113) percent of postorbital carapace length (PCL), with 4 to 8 (mode 6) punctations across narrowest part. Cervical spines long, acute, usually single spine each side of carapace (70.7% of 123 specimens). Branchiostegal spine prominent, acute; hepatic spines or spiniform tubercles only occasionally present. Suborbital angle vestigial to nearly obsolete; postorbital ridge well developed, with prominent cephalic spine. Antennal scale 2.7 to 3.6 (x = 3.1, n = 113) times as long as broad, widest just distal to midlength, lateral margin thickened and terminating cephalically in long spine. Palm of chela inflated (ratio width to depth 1.4-1. 8, j = 1.6, n = 108), lateral margin costate for most of length; dorsal surfaces of palm and fingers usually with dense setae; mesial margin of palm with 2 rows of tubercles, mesialmost row subserrate, with 6 to 11 (mode 7-8) prominent, acute tubercles; irregular row of 4 to 10 (mode 7-8) smaller, truncate or subsquamous (but often acute) tubercles subtending mesial row dorsally; conspicuous tuft of setae at opposable bases of both fingers; fixed finger with well defined ridges dorsally, mildly costate dorsolaterally; opposable surface of finger with row of 4 to 8 (mode 5-6) knoblike tubercles between base and about midlength of finger, and isolated subconical tubercle ventral to continuous row of denticles at base of distal one- third or one-fourth; opposable surface of dactyl moderately excised in basal one-third in adult males; dactyl with well defined dorsal ridges flanked by rows of setiferous punctations; proximal one-half to two- thirds of mesial margin of dactyl with 2 rows of prominent tubercles, ventral row often subserrate in dorsal outline, with 4 to 10 (mode 6) somewhat depressed, often spiniform tubercles; dorsal row with 3 to 8 (mode 4) tubercles, and sometimes subtended laterally by third short, irregular row; opposable surface of dactyl with row of 4 to 9 (mode 6) knoblike tubercles between base and about midlength. Merus of cheliped with at least 2 long subdistal spines dorsally, and 1 to 4 short, acute spines proximal to them; ventromesial ridge New Crayfish Species 69 of merus with longitudinal row of 2 to 8 (mode 4-5) prominent, acute spines in addition to long distal spine, ventrolateral ridge with longitudinal row of 2 to 7 (mode 2-3) prominent, acute spines in addition to long distal spine. Ventral surface of carpus of cheliped almost always with strong distomedian spine, and larger acute spine on lateral articular condyle; mesial surface of carpus with long, procurved spine at about midlength, usually smaller spine distal to it, and tubercle, often spiniform, near proximal margin. Hook on ischium of third pereiopod of form I male usually extending beyond basioischial articulation, sometimes opposed by low, vestigial tubercle on basis. Length of gonopod (first pleopod) of form I male divisible into TCL 2.1 to 2.5 ( * = 2.2, n - 54) times; cephalic surface of gonopod with prominent angular shoulder, distal to which both rami inclined cephalically; terminal elements long, slender, subparallel to about midlength of mesial process; central projection tapering smoothly from base to acute tip, distal portion gently recurved, tip directed caudodistally and reaching to or beyond midlength of coxa of first pereiopod when abdomen flexed; length of central projection 46.5 to 57.9 ( x = 51.6, n = 54) percent of total length of gonopod; tip of mesial process somewhat spatulate and mildly excavate cephalically; length of mesial process 34.7 to 44.5 (* = 41.5, n = 54) percent of total length of gonopod, and 73.3 to 89.2 (x = 80.4, n = 54) percent of length of central projection. In caudal (ventral) aspect, bases of gonopods and tips of mesial processes contiguous; tip of central projection directed slightly distolaterally, that of mesial process directed distomesially. Annulus ventralis inflexibly fused to sternum; annulus 1.3 to 1.4 times wider than long, with deep, broad transverse fossa; cephalic border of annulus convex, caudal border subangular; cephalomedian wall with short longitudinal trough, flanked at each side by ridge, ridges diverging caudally and each terminating in large, rounded cephalo- lateral prominence, prominences in ventral aspect obscuring cephalic half or more of fossa; caudomedian wall dissected by sinus; obvious sulcus cephalic to caudal margins. Measurements of type specimens provided in Table 1. Holotypic male, form I — Body and eyes pigmented, eye 2.5 mm diam. Cephalothorax (Fig. 1A, D) subcylindrical, somewhat depressed dorsally; carapace widest at midlength, width greater than depth at caudodorsal margin of cervical groove; abdomen narrower than carapace. Areola 4.1 times longer than wide, constituting 29.5 percent of TCL (42.0% of PCL), with 5 punctations across narrowest part and low, rounded eminence at caudal base. Rostrum acarinate, length 31.2 percent of TCL (44.4% of PCL), deeply excavate, with thickened, elevated 70 John E. Cooper and Martha Riser Cooper Table 1. Measurements (mm) of types, Orconectes ( P .) carolinensis, new species. Holotype Allotype Morphotype Carapace Total length 29.2 29.7 23.9 Postorbital length 20.5 20.9 17.1 Width 14.1 14.0 10.9 Depth 11.4 12.7 9.8 Rostrum Length 9.1 9.3 7.3 Width at base 4.2 4.4 3.7 Length acumen 4.4 4.5 3.5 Areola Length 8.6 9.1 7.1 Width 2.1 2.3 1.7 Antennal scale Length 7.8 8.2 6.6 Width 2.5 2.8 2.3 Abdomen Length 33.5 34.7 27.1 Width 10.9 11.5 9.0 Cheliped (right) Length lateral margin chela 24.5 17.6 16.7* Length mesial margin palm 7.2 5.7 5.1* Width palm 9.2 7.0 6.3* Depth palm 5.6 4.4 3.7* Length dactyl 13.7 9.5 9.1* Length carpus 7.9 7.3 5.9 Width carpus 5.4 4.7 4.3 Depth carpus 4.7 4.0 3.3 Length dorsal margin merus 10.7 9.3 7.8 Depth merus 5.0 5.1 4.1 Gonopod length 13.1 NA 10.5 * Left chela. margins bearing long marginal spine at base of acumen each side; rostrum subrectangular, margins converging very slightly to base of marginal spines; walls and floor of rostrum with setose punctations, proximal portion of floor cephalodorsally acclivous, floor of acumen plane. Acumen length 48.4 percent of rostrum length, strongly tapering from base to acute, corneous tip, which extending to distal margin of second article of antennular flagellum. Subrostral ridge strong, evident in dorsal aspect from base of rostrum to marginal spines. New Crayfish Species 71 Fig. 1. Orconectes (Procericambarus) carolinensis, new species (all from holotypic male, form I, except C, E, from morphotypic male, form II, and G, from allotypic female; setae not illustrated): A, lateral aspect of cara- pace; B, C, mesial aspect of gonopod (first pleopod); D, dorsal aspect of •j carapace; E, F, lateral aspect of gonopod; G, annulus ventralis and postannular sclerite; H, epistome; I, basal podomeres of third pereiopod; J, dorsal as- pect of distal podomeres of cheliped; K, antennal scale; L, caudal aspect of in situ gonopods. 72 John E. Cooper and Martha Riser Cooper Postorbital ridge strong throughout most of length, with shallow, setiferous dorsolateral groove; cephalic margin terminating in strong spine. Suborbital angle nearly obsolete, delimited ventrally by shallow rounded notch at proximal base of antennal peduncle. Cervical spines strong, acute, directed cephalolaterally, 1 each side, plus single, small spiniform tubercle just dorsal to spine. Branchiostegal spine prominent, acute. Hepatic region punctate, with some small subspiniform tubercles. Carapace densely punctate dorsally (including gastric region) and laterally, somewhat granulate ventrolaterally; cluster of tubercles just caudal to branchiostegal spine and ventral to cephalolateral portion of cervical groove, which with row of tubercles extending along ventral margin; cervical groove deep, uninterrupted, with shallow, curved sulcus (tributary to groove) ventral to cervical spine, creating subglabrous mound below spine. Abdomen slightly longer than carapace; pleura well developed, subtruncate, with smoothly rounded cephaloventral margins and sub- angular caudoventral margins; minute atypical notch in apex of second through fourth pleura. Cephalic section of telson with large, acute, immovable spine at caudolateral corner, and smaller, movable spine just mesial to it (that on right side small, regenerate); cephalic and caudal sections of telson partly separated by oblique lateral incisions and shallow transverse sulcus. Promixal podomere of uropod with long, corneous spine on mesial lobe and slightly smaller spine on lateral lobe; mesial ramus of uropod with distolateral marginal spine and relatively broad median keel terminating distally in acute premarginal spine; cephalic section of lateral ramus with median keel terminating in acute spine at transverse flexure, which with total of 13 fixed spines and 1 large movable spine at lateral margin. Cephalic lobe of epistome (Fig. 1H) spatulate, tilted cephaloventrally, with moderate constriction and deep transverse groove at caudal base; cephalomedian margin with slight notch ventral to short projection; cephalolateral margins elevated (ventrally), broad, sloping onto ventral surface and with thin, emarginate rim bearing short setae; lateralmost extremities mildly flanged; ventral surface of lobe concave, punctate, with short, sparse setae, concavity continuing into cephalomedian notch; caudal one-third of ventral surface with low, subtriangular eminence; main body of epistome relatively glabrous, cephalomedian and cephalo- lateral margins forming a somewhat hemitubular, curved ridge; central depression of body broad, deep, with deep fovea situated in midline at cephalic margin of depression; zygoma moderately arched, wider than space between renal apertures; pits at cephalolateral borders of zygoma elongate and relatively shallow. New Crayfish Species 73 Proximal podomere of antennular peduncle with strong spine on ventral surface slightly proximal to midlength; antennal peduncle with acute ventromedial spine on ischium and larger distolateral spine on basis. Antennal flagellum about 49 mm long, tip reaching midlength of fourth abdominal tergite when flagellum adpressed. Antennal scale (Fig. IK) 3.1 times as long as wide, greatest width just distal to midlength; distal margin of lamella steeply declivous to widest part, row of setae not encroaching on base of apical spine, mesial margin narrowly curved; lateral margin thickened, gently bowed and with strong apical spine, tip of which directed distolaterally and reaching distal margin of ultimate podomere of antennular peduncle and base of distal one-third of same podomere of antennal peduncle. Third maxilliped with tip reaching about midlength of basal podomere of antennal peduncle, tip of exopodite reaching base of distal one- fourth of merus of endopodite; cephalolateral margin of ischium produced as acute spine; ventrolateral ridge flanked mesially by row of punctations bearing short setae; most of ventrolateral half with sparse punctations and short setae, but distomedial margin with longer setae; ventromesial half of ischium with long, stiff setae, longer and more dense proximally than distally and obscuring proximal portion; surface between bases of setae and dentate mesial margin glabrous, margin with 23 denticles. Right mandible with incisor ridge bearing 7 denticles (6 on left). Right chela (Fig. 1J) with moderately inflated palm, ratio width to depth 1.6, ratio length to depth 1.3; lateral margin of palm and proximal fixed finger visibly but narrowly costate dorsally and ventrally; chela 2.7 times longer than wide, shorter than carapace (ratio TCL to chela length 1.2). Dorsal surface of palm densely punctate, most punctations with short setae, but longer and more plumose ones laterally and mesially. Mesial margin of palm with 2 obvious rows of tubercles; mesialmost row subserrate, with 8 acute, subconical tubercles (9 on left chela), and distomesial margin of articular ridge produced as additional rounded tubercle; single small, rounded tubercle at proximolateral base of proximalmost tubercle; mesial row of tubercles flanked dorsally by irregular row of 7 (8 on left) anteriorly rounded to subsquamous tubercles; 2 rounded tubercles dorsal to dorsal row at about midlength of palm, and 4 subsquamous to squamous tubercles lateral to these; all tubercles with group of setae originating at distal base. Ventral surface of palm subglabrous, with sparse, shallow punctations and minute setae. Fingers moderately gaping in proximal half, opposable surfaces contiguous along distal half. Dactyl subovate in cross-section; mesial surface gently bowed, with 2 rows of anteriorly elevated and rounded 74 John E. Cooper and Martha Riser Cooper tubercles, somewhat subserrate in dorsal outline; 10 tubercles (13 on left dactyl) in ventromesial row, extending to base of distal two-fifths of finger; single tubercle proximoventral to first in ventromesial row (on left, similar tubercle ventral to first); dorsomesial row with 6 tubercles (7 on left); single small, rounded tubercle dorsal to base of first one in dorsomesial row. Ventral surface of dactyl rounded, with sparse punctations bearing setae. Opposable surface of dactyl with mat of plumose setae at base, and moderate excision extending from first to fourth tubercles; margin with row of 9 knoblike tubercles; denticles in 3 to 4 rows, extending along distal two-thirds of finger. Fixed finger of propodus subtriangular in cross-section, with well developed, fairly broad middorsal ridge, which essentially glabrous, but with sparse punctations on proximal one-fifth; ridge mesially and laterally subtended by groove containing row of setiferous punctations extending nearly to base of tip, and flanked each side by somewhat abbreviated ridge, ridges merging at about base of distal one-fourth of finger; rudimentary dorsolateral ridge in proximal half of finger, extending to about midlength; lateral margin proximally costate. Opposable surface of fixed finger with setae at base, margin with row of 8 knoblike tubercles extending to about midlength; subconical tubercle at base of distal one-third of finger, ventral to denticles, which in 3 to 4 continuous rows. Right carpus (Fig. 1J) 1.5 times as long as wide, 1.7 times as long as deep, ratio width to depth 1.1; carpus punctate dorsally, with oblique median sulcus, several squamous tubercles on dorsomesial surface; mesial surface sparsely punctate, with long procurved spine at midlength, flanked proximally by smaller, somewhat acute tubercle near mesial base of podomere; second long, acute spine situated proximomesial to articular eminence; ventromesial surface with sparse, setiferous puncta- tions and several very small tubercles; ventral surface with large disto- median spine and larger spine on distal end of lateral articular condyle; lateral surface of carpus with scattered setiferous punctations. Right merus 2.1 times as long as deep; dorsal surface with 2 long subdistal spines; ventromesial ridge with acute distal spine and 5 additional ones, ventrolateral ridge with acute distal spine and 4 additional ones, none on distolateral articular condyle; lateral and mesial surfaces of merus essentially glabrous. Right ischium with row of 3 low, subacute tubercles on ventral ridge, distal to suture line; small hooklike sufflamen adjacent to large articular condyle of coxa. Palm of chela and carpus of second pereiopod with row of long setae on dorsomesial and ventromesial surfaces, no stiff setae on mesial margin; other podomeres with rows of long setae on ventral or ventromesial New Crayfish Species 75 surfaces and sparse setae on dorsomesial surface, none stiff; distal margin of merus of second through fourth pereiopods with small, acute ventrolateral spine. Coxa of fourth pereiopod without boss, ventral surface punctate and with long setae; ventral membrane on coxa of fifth pereiopod studded with short setae. Ischium of third pereiopod (Fig. II) with simple hook extending past basioischial articulation by nearly half its length, hook opposed by vestigial tubercle on basis. Gonopods (Fig. IB, F, L) symmetrical (see “Diagnosis” for description). In addition, left gonopod length 44.9 percent of TCL (63.9% of PCL). Allotypic female — Differing from holotype in following respects: areola 4.0 times as long as wide, length 30.6 percent of TCL (43.5% of PCL). Tip of acumen extending to midlength of second article of antennular flagellum. Small acute tubercle ventrocephalic to cervical spine. Total carapace length 85.6 percent of abdomen length. Antennal scale 2.9 times longer than wide, tip of apical spine extending to about midlength of second article of antennular flagellum. Palm of right chela 1.6 times wider than deep; chela much shorter than carapace (ratio chela length to TCL 1.7), and 2.5 times longer than wide; mesial row of tubercles on palm subtended dorsally by irregular row of 6 tubercles. Mesial surface of dactyl not bowed, opposable base not excised, closed fingers without proximal gape. Mesial surface of dactyl with ventromesial row of 4 acute but depressed tubercles. Opposable surface of fixed finger with row of 6 (5 on left) knoblike tubercles extending just distal to midlength of finger; subconical ventral tubercle situated just proximal to base of distal one-third of finger. Right carpus 1.8 times as long as deep, ratio width to depth 1.2. Right merus 1.8 times longer than deep. Right ischium with row of 4 acute tubercles on ventral ridge. Annulus ventralis (Fig. 1G) as described in “Diagnosis.” In addition, first pleopod well developed, extending beyond cephalic margin of annulus when abdomen flexed. Postannular sclerite (Fig. 1G) 3.2 times wider (2.9 mm) than long (0.9 mm). Morphotypic male, form II — Differing from holotype in following respects: rostrum 30.5 percent of TCL (42.7% of PCL); acumen length 47.9 percent of rostrum length. Total carapace length 88.2 percent of abdomen length. Left chela (right regenerate) 2.7 times longer than wide; mesial margin of palm with subserrate mesial row of 7 spiniform tubercles, distal margin developed as eighth. Mesial surface of dactyl with subserrate row of 6 acute tubercles extending to about midlength of finger, dorsally subtended by irregular row of 5 squamous tubercles and ventrally by several squamous tubercles; opposable surface of dactyl with 4 knoblike tubercles extending to about base of distal 76 John E. Cooper and Martha Riser Cooper one-third of finger, fourth slightly offset ventrally, small vestigial fifth tubercle dorsal to row of denticles just distal to fourth tubercle, another small tubercle interrupting denticles just proximal to midlength of finger, and a minute, conical tubercle (smaller than denticles and among them) distal to midlength. Opposable surface of fixed finger with 4 knoblike tubercles, smaller fifth tubercle at about midlength of finger. Antennal scale 2.9 times longer than wide. Tip of antennal flagellum reaching nearly to caudal margin of third abdominal tergite. Merus with 3 spines each on ventrolateral and ventromesial ridges. Hook on ischium of third pereiopod greatly reduced. Left gonopod (Fig. 1C, E) somewhat aberrent, mesial process in mesial aspect slightly longer than on right, tip curving cephalically, recurving, directed distocephalically; terminal elements of gonopod shorter and more robust, both less acute, central projection not corneous, cleft between terminal elements much shorter; cephalic border of gonopod without prominent angular shoulder. Right gonopod 43.9 percent of TCL (61.4% of PCL). Color notes — Base color varies from tan to forest green. Dark brown to black saddle, often mottled, on posterior carapace, narrowest between caudal bases of branchiocardiac grooves and caudal ridge; horns of saddle produced along ventrolateral margin of carapace as far cephalically as anteroventral branchiostegal region below spine; anterolateral branchiostegite with somewhat reticulated blotches of brown, olivaceous, or black pigment; antennal region of carapace cream; mandi- bular adductor region with irregular dark brown to black splotches or mottlings; tips of acumen and marginal, cephalic, and cervical spines crimson or orange, subtended proximally by black band; marginal ridges of rostrum black; most of carapace and abdomen with fine dark flecking. Cephalic portion of first abdominal segment with paired, subtriangular, dorsolateral blotches, dark brown, olivaceous, or black in color; second through fourth abdominal segments with short, paired dorsolateral bars inclined cephalolaterally from cephalic margin, imparting in dorsal aspect “interrupted chevron” pattern; pleura of second through fifth segments with dark oblique blotch extending caudoventrally from base; caudal rim of each abdominal segment with narrow, light brown, orangish, or red band. Caudal portion of proximal segment of lateral ramus of uropod with transverse light brown band; ventral surfaces of uropods and telson with fine dark speckling. Dorsal surface of palm of chela orangish tan, dorsal surfaces of fingers darker, all with small flecks and some irregular spots or mottling; tips of fingers with crimson or orange band, subtended proximally by somewhat broader black band; large tubercle on dorsal palm proximal to articulation of dactyl crimson; New Crayfish Species 77 ventral surface of cheliped oyster with black flecking, tubercles at base of dactyl crimson; lateral margin of propodus with thin iodine or black line; articular eminences of chela and carpus of cheliped pale orange; distolateral spine of merus crimson. Pereiopods base color, with somewhat darker, mottled bands; margins and articulations crimson. Annulus ventralis and postannular sclerite gunmetal blue, except outer surfaces of cephalolateral prominences and region of sinus in caudal wall white. Antennular and antennal flagellae greenish brown proximally, changing to reddish tan distally. Type locality — North Carolina, Jones County, Trent River (Neuse River basin) at State Road (SR) 1129 near junction SR 1131, ca. 4.5 air mi (7.2 air km) NNE of Comfort (Phillips Crossroads USGS Quadrangle, UTM coordinates 3882150/275010). On 5 October 1978, when the holotype and several paratypes were collected, the river was about 10 m wide between banks, the abnormally low water was clear and shallow, and there was little or no visible flow. Pitted limestone outcrops were abundant at the site, and the limestone substrate was covered with fine silt, organic debris, and pale yellowish flocculence. Most of the 16 specimens of O. carolinensis collected were found under rocks. They were darkly encrusted and a great deal of flocculence was clinging to their setae. No other crayfishes were found at this site, but other aquatic invertebrates observed included Hemiptera of the families Nepidae, Notonectidae, and Belostomatidae; two kinds of Odonata nymphs; at least two kinds of mussels, one of them large and very abundant; several kinds of gastropods; aquatic Coleoptera, including a species of dytiscid; several kinds of unidentified insect larvae; and abundant Palaemonetes paludosus (Gibbes). A number of N. lewisi also were collected or observed. Disposition of types — The holotype, allotype, and morphotype are in the crustacean collections of the North Carolina State Museum of Natural Sciences (NCSM), Raleigh (catalogue numbers NCSM C- 2462, C-2486, and C-2463, respectively), as are the following paratypes: 1 6 II, 3 j 6, and 2 j 9 (NCSM C-78); paratypes consisting of 8 8 I and 1 ovig 9 are in the United States National Museum of Natural History (USNM), Smithsonian Institution, Washington (USNM 332038). Range and specimens examined — Endemic to the Neuse and Tar- Pamlico river basins of North Carolina. Within the Neuse basin, O. carolinensis occurs from near Willow Springs in southern Wake County, southeast to the upper reaches of the Trent River in central Jones County to Swift Creek on the line between Pitt and Craven counties. It appears to be absent from some of the Coastal Plain and most of the Piedmont Plateau of the Neuse River basin. Within the Tar-Pamlico 78 John E. Cooper and Martha Riser Cooper basin, the species occurs from headwater streams in the Piedmont of Granville County, east to western Halifax County and southeast to Pitt County. Specimens have been collected at the following localities (nearly all specimens are catalogued in the collections at NCSM, some are at USNM): TAR-PAMLICO RIVER BASIN. Edgecombe Co.—( 1) Tar R at US 64 bridge in Tarboro; 2 9, 19 Jan 1980, R. W. Mays; (2) Tar R at NC 42 E of Old Sparta, 5 air mi (8 air km) E center Pinetops; 4 8 I, 13 9, 30 Oct 1984, A. L. Braswell, JEC; (3) Tar R at Tarboro; 1 j c3, 2 j 9, ? Aug 1983, D. R. Lenat; (4) Tar R at NC 44 bridge, I. 6 air mi (2.6 air km) NNW Tarboro; 1 8 II, 5 9, 18 May 1986, ALB, D. Smith. Franklin Co. — (5) Shocco Crk at NC 58, 1.6 air mi (2.6 air km) N Centerville; 1 <5“ I, 17 Jan 1980, E. Rawls. Franklin- Vance Co. line— (6) Tar R at US 1; 1 8 II, 24 Jul 1993, D. G. Cooper, D. Jackan. Granville Co. — (7) Tar R at SR 1141, 0.9 air mi (1.4 air km) SSW Berea; 1 8 I, 1 8 II (molted to 8 I), 6 ovig 9,1 9 & lst-instar young, 6 May 1981, ALB; 1 8 II, 1 9, 17 Jun 1980, ALB, J. Cannon; 1 8 II, 1 9,1 9 with exuvium, 4 Jun 1994, ALB, JEC; (8) Tar R ca. 0.5 mi (0.8 km) above SR 1133 bridge, ca. 1.8 air mi ( 2.9 air km) SE Providence; 2 9, 18 Aug 1980, ALB, J. H. Reynolds, C. Carnes; (9) Tar R at SR 1622, ca. 5.1 air mi (8.2 air km) SSW Dickerson; 1 <3 I, 25 Feb 1980, ER; 1 8 I, 28 Jan 1980, ER; (10) Tar R at SR 1138, ca. 2.3 air mi (3.7 air km) N Culbreth; 1 8 I, 14 Nov 1981, R. E. Ashton, Jr., DS, P. Kumyhr; (11) Tar R at NC 96, 3.7 air mi (5.9 air km) NNW Wilton; 8 8 I, 1 ovig 9, 16 Apr 1977, R. Thoma; 2 8 II, 1 9 with young, 18 May 1986, ALB, DS, D. Etnier. Halifax Co. — (12) Little Fishing Crk, ca 0.3 mi (0.5 km) below SR 1322 bridge, ca 2.7 air mi (4.3 air km) E Hollister; 4 <3 I, 8 c3 II, 12 9,6 Aug 1980, ALB; (13) Bear Swamp at SR 1300, 6.7 air mi (10.7 air km) NNE Hollister; 1 8 I, 28 Feb 1980, RWM. Nash Co. — (14) Swift Crk at SR 1003, 3.3 mi (5.3 km) NE center Red Oak; 1 8 II, 8 Mar 1980, RWM; (15) Stony Crk at SR 1603, S jet SR 1609, 3.5 mi (5.6 km) S Red Oak; 1 8 I, 8 Mar 1980, RWM; (16) Tar R at SR 1746, 5 mi (8 km) SW Rocky Mount; 1 8 I, 25 Jan 1980, RWM. Pitt Co. — (17) Tar R at SR 1560, 1.3 air mi (2.1 air km) SE Pactolus; 1 9, 24 Mar 1980, JHR; (18) Tar R ca. 4 air mi (6.4 air km) E jet US 264 Bypass, 2 air mi (3.2 air km) NNE Simpson; 1 8 I, 29 Feb 1980, JHR; (19) Tar R ca. 2 air mi (3.2 air km) E jet SR 1565, ca. 2.8 air mi (4.5 air km) ENE center Grimesland; 1 9 , 29 Feb 1980, JHR. Warren Co. — (20) Little Fishing Crk at SR 1532, ca. 3 air mi (4.8 air km) NNE Grove Hill; 2 8 I, 24 Mar 1980, ER; 2 8 I, 21 Mar 1980, New Crayfish Species 79 ER; (21) Reedy Pond Crk at SR 1510, ca. 1.6 air mi (2.6 air km) NNE Grove Hill; 2 8 I, 1 9, 19 Mar 1980, ER; 2 8 I, 17 Mar 1980, ER; (22) Shocco Crk at SR 1613, ca. 2.7 air mi (4.3 air km) NW Lickskillet; 3 8 I, 1 9, 10 Mar 1980, ER; (23) Fishing Crk at SR 1640, ca. 4.3 air mi (6.9 air km) SE Inez; 1 8 I, 14 Mar 1980, ER; (24) Fishing Crk at SR 1600, 2.9 mi (4.6 km) SSE Warrenton; 1 8 I, 27 Mar 1980, ER; (25) Shocco Crk at SR 1133, ca. 2.3 air mi (3.7 air km) SSE Vicksboro; 1 8 I, 14 Mar 1980, ER; (26) Fishing Crk at SR 1609, 4.4 mi (7.0 km) SSE Warrenton; 1 9, 26 Mar 1980, ER; (27) Possumquarter Crk at SR 1606, 3.5 mi (5.6 km) SSE Warrenton; 1 9, 17 Mar 1980, ER. NEUSE RIVER BASIN. Craven-Pitt Co. line. — (28) Swift Crk at SR 1465 (Craven Co.), ca. 7.3 air mi (11.7 air km) W Vanceboro; 2 8 I, 19 Mar 1979, JHR. Greene-Lenoir Co. line. — (29) Contentnea Crk at SR 1004 (Greene Co.), 5.1 mi (8.2 km) SSE Hookerton; 1 8 I, 20 Mar 1979, P. S. Freed, E. Flowers. Greene-Pitt Co. line. — (30) Little Contentnea Crk at SR 1311, 2.5 mi (4.0 km) NNE Walstonburg; 2 (5 I, 1 9, 9 Mar 1979, PSF. Johnston Co.— (31) Middle Crk at SR 1507, ca. 3.2 air mi (4.1 air km) ENE Willow Springs; 1 8 I, 1 8 II, 4 Apr 1979, A. P. Capparella; 1 8 I, 1 9 with young, 8 Apr 1979, APC; (32) Middle Crk at NC 210, ca. 3.2 air mi (5.1 air km) W Smithfield; 1 8 II, 3 j 8,2] 9, 28 Jul 1976, D. S. Lee, R. Franz; 16 j 8, 4 9, 16 j 9,2 Aug 1976, DSL, M. M. Browne, Z. Sykes, MRC; (33) Middle Crk at SR 1504, 6.8 air mi (10.9 air km) W. Smithfield; 2 j 8,1 9,2 Aug 1976, DSL, MMB, ZS, MRC; (34) Neuse R at SR 1201, 6.7 mi (10.7 km) SSW Princeton; 1 8 I, 18 Mar 1979, PSF. Jones Co. — (35) Beaver Crk at SR 1316, 6.2 air mi (9.9 air km) NW Trenton; 1 8 I, 6 Feb 1979, JHR; (36) Trent R at SR 1129, 4.5 air mi (7.2 air km) NNE Comfort (TYPE LOCALITY); 9 8 I, 7 9,5 Oct 1978, ALB, REA, Jr., JEC; (37) Trent R at SR 1300, 4.8 air mi (7.7 air km) NW center Trenton; 9 8 I, 1 j 8,6 9, 1 j 9,1 Oct 1983, B. M. Burr, P. A. Burr; (38) Big Chinquapin Br at SR 1129, 0.8 air mi (1.3 air km) NE Phillips Crossroads; 1 8 I, 6 Feb 1979, JHR; (39) Beaver Crk at SR 1303, 5 air mi (8 air km) S Wyse Fork; 1 8 I, 14 Feb 1979, JHR; (40) Trent R at NC 58, 1.8 air mi (2.9 air km) ESE Phillips Crossroads; 1 8 I, 3 Sep 1985; DRL; (41) Trent R, 2.5 air mi (4.0 air km) WNW Pollocksville at Marine Corps Facility Oak Grove; 1 8 l, 22 Feb 1993; ALB, J. C. Beane. Wake Co. — (42) Middle Crk at SR 2739, 3.6 air mi (5.8 air km) E Willow Springs; 1 1,13 Mar 1979, APC. Wilson Co. — (43) Turkey Crk just S Nash Co line, 1.0 air mi (1.6 air km) W Conner; number & sexes not available, 10 Jul 1985, V. Schneider. 80 John E. Cooper and Martha Riser Cooper Variations and anomalies — Individual variation in a number of characters is common, but no consistent hydrologic or geographic pat- terns are evident. Significant meristic and proportional variations are addressed in the “Diagnosis,” but others also require notation. The distomedian eminence on the ventral surface of the carpus of the cheliped varies in development from a broad, rounded tubercle (rarely) to the usual prominent, acute spine that often is as long as the lateral and mesial spines of the podomere. Most individuals have a single cervical spine on each side of the carapace, but in 8.9 percent of 123 specimens there are 2 spines per side, and some animals have a single spine on one side and 2 on the other. One form I male has a tubercle and 2 spines on the left, 3 spines on the right. In some individuals, one or another of the cervical spines is bifurcate, and one female has two bifurcate spines on each side. In the same female, the right eye was yellow in life, and there are 3 spines instead of the normal 2 in each caudolateral corner of the cephalic section of the telson. There is also variation in the latter character in other specimens, but only one other individual, a form I male, has 3 spines in each corner. Small hepatic spines or spiniform tubercles are present in 6 of 123 specimens. One form I male shows a congenital lack of marginal spines on the rostrum, and in the same animal the mesial process of the left gonopod has a very acute tip with a minute notch in its caudal surface. Another form I male has a full-sized but deformed antennal scale projecting at nearly 90 degrees dorsally from the base of the normal left antennal scale. One female has a copulatory hook on the ischium of one of the third pereiopods. The fingers of the chela of females and form II males are essentially contiguous throughout their lengths when closed, but in form I males the fingers gape in the proximal half, and the proximal one-third of the opposable margin of the dactyl is moderately excised. In dorsal aspect, the central one-third of the dactyl in form I males is gently concave, then strongly recurved. In females, a single row of denticles extends distally along the opposable margin of the dactyl to the base of the cornified tip. In form II males there are one or two such rows of denticles, and in form I males there are three or four rows. The antennae of form I males are slightly longer than those of form II males and females. Size — The largest specimen in our samples is a form I male of 34.1 mm TCL (23.6 mm PCL). Only eight other adult males have TCLs in excess of 30 mm. The smallest form I male measures 15.8 mm TCL (10.8 mm PCL). Lourteen other form I males have TCLs of less than 20 mm, and the mean of 55 form I males is 24.6 mm. New Crayfish Species 81 Only two large form II males have been collected; one (the morphotype) is 23.9 mm TCL (17.1 mm PCL), and the other is 24.3 mm TCL (16.7 mm PCL). The largest female measures 32.8 mm TCL (24.1 mm PCL). Five other females have TCLs greater than 28 mm. The smallest female with attached ova or young measures 17.1 mm TCL (11.3 mm PCL), and the largest female in this condition measures 25.3 mm TCL (18.1 mm PCL). Assuming 17.1 mm as the lower limit of TCL for sexually mature females (range = 17.1-32.8 mm; n = 46), the mean TCL for this group is 23.4 mm. The TCLs of 101 form I males and mature females range from 15.8 to 34.1 mm (both extremes are males) and the mean is 24.1 mm. Life history notes — Form I males have been collected in every month except June, July, and December, but were preponderant in the spring and fall. Of 71 such males, 22 were collected in March, 10 in April, and 22 in October. A form II male collected on 6 May 1981 molted to form I on 3 July 1981 in the laboratory. Females bearing ova or young have been collected only in April and May. An unmeasured female taken on 8 April 1979 had two third- instar young attached, and a female measuring 18.7 mm TCL (13.4 mm PCL), collected on 16 April 1977, was carrying 39 ova of about 1.8 mm diameter. The latest date on which a female carrying young has been found was 18 May 1986. Table 2 provides data for six laden females collected at the same site on 6 May 1981. Crayfish associates — Cooper and Ashton (1985) and Cooper and Braswell (1995) briefly discussed the decapod fauna of the Neuse and Tar-Pamlico basins. The crayfish associates encountered with greatest frequency in O. carolinensis samples were Cambarus (Depressicambarus) latimanus (LeConte) and Cambarus ( Puncticambarus ) acuminatus Faxon (s.l.). Procambarus ( Ortmannicus ) acutus (Girard) was the third most often collected species in these samples, and all three of these associates appeared together in some of them. Cambarus ( Lacunicambarus ) diogenes Girard also was included in several of the collections. Generally, C. acuminatus (s.l.) and C. latimanus far outnumbered O. carolinensis in samples where they were taken together. At a few localities, however, O. carolinensis either outnumbered any other species found, or was the only species collected. Although Fallicambarus ( Creaserinus ) fodiens (Cottle), Cambarus ( Depressicambarus ) reduncus Hobbs, and P. medialis have been found in open water in both river basins, they have not been taken at any O. carolinensis site. Relationships — The general features of the form I male gonopod (terminal elements long and of subequal length, subparallel, subsetiform) and the female annulus (well defined sulcus, deep fossa, large and 82 John E. Cooper and Martha Riser Cooper Table 2. Data for six laden female Orconectes ( P .) carolinensis, new spe- cies, collected at the same site on 6 May 1981. TCL (PCL) Condition 17.1 (11.5) 28 ova (1.8-1. 9 mm diam) 19.5 (13.7) 59 ova (1.7-1. 9 mm diam), 3 of them beginning eclosion, and several empty egg cases 20.4 (14.3) 76 ova (1.7-1. 8 mm diam) 21.2 (14.9) 29 first-instar young and 1 unhatched ovum 23.8 (16.7) 16 first-instar young and 60 ova (1. 9-2.0 mm diam) undergoing eclosion 25.3 (18.0) 109 first-instar young and 2 unhatched ova lobiform cephalolateral prominences overhanging fossa and separated by deep trough) are typical of subgenus Procericambarus (Fitzpatrick 1987:57-58). However, the upper limit of the range for length of the terminal elements of the mature gonopod, expressed as percent of total length of the appendage, is slightly higher for O. carolinensis (range = 46.5-57.9% * = 51.6%, n = 54) than for the subgenus (range = 34-55%). In addition, the range for areola length, expressed as percent of TCL, is considerably lower for O. carolinensis (range = 25.5-31.1%, x = 28.5%, n = 113) than for the subgenus (range = 29-37%). Within Procericambarus , O. carolinensis clearly has its greatest affinities with several members of the Spinosus Group, as defined by Fitzpatrick (1987:58). Its closest relatives almost certainly are Orconectes (. Procericambarus ) spinosus (Bundy), of the Tennessee and Coosa river basins, and Orconectes (Procericambarus) putnami (Faxon), which occurs in parts of the Ohio River drainage. The new species may be distinguished from its relatives in the Spinosus Group by the following: (1) the greater length of the terminal elements of the mature gonopod (range for the Spinosus Group is 40-48% of total gonopod length); (2) the shorter, broader areola, and the greater number of punctations across the narrowest part; (3) the somewhat longer acumen and rostrum, the latter being deeply excavate, much more punctate and setiferous, and with thicker margins; (4) the smaller size (see section on “Size” for data); (5) the lack of a clearly defined ventral row of tubercles subtending the row on the mesial margin of the palm; (6) fewer tubercles in the ventralmost and dorsalmost rows of the mesial margin of the dactyl; (7) the far greater number of spiniform tubercles on the ventrolateral ridge of the merus, these being generally strong, acute spines, especially in the distal half to New Crayfish Species 83 two-thirds of the podomere, as opposed to small (often minute) tubercles; (8) fewer but much more highly developed tubercles on the ventromesial ridge of the merus; (9) the presence of 3 or more, usually spiniform and sometimes bifurcate, tubercles on the ventromesial ridge of the ischium; and (10) the occasional presence of multiple cervical spines (and small, spiniform hepatic tubercles). A few of the characters exhibited by O. carolinensis could be interpreted as plesiomorphies within the subgenus. These include multiple cervical spines and spiniform hepatic tubercles in some specimens, a generally more spinose mien, a short, broad areola, and a color pattern that includes a “saddle” on the carapace. It is tempting to conclude that it is one of the phylogenetically older species within the Spinosus Group and perhaps within the subgenus. On the other hand, the various spines and spiniform tubercles could also represent periodic recurrences of atavisms. Until such time as the other members of the Spinosus Group, especially O. spinosus and O. putnami, have been described and defined more thoroughly, any conclusions as to the relative “age” of O. carolinensis would be premature. Remarks — Fitzpatrick (1987:69) hypothesized that the progenitors of Procericambarus occupied the southern extremities of the eastern part of the Tennessee River, and that their establishment there “may have taken place in early Quaternary times.” From this center they spread over the Cumberland Plateau, entered the Ohio system, and expanded west. Today the subgenus occupies a broad range that extends from the Blue Ridge into eastern Oklahoma and Kansas, and the disjunct O. carolinensis is the only representative inhabiting Atlantic Coast drainages. In North Carolina, the easternmost montane populations of the Spinosus Group occur in the New and upper Little Tennessee river basins (Cooper and Braswell 1995), from approximately 270 to 480 air km (170 to 300 air mi), respectively, west of the westernmost populations of O. carolinensis. It seems reasonable, considering the incontrovertible affinities of the species, that it derived from an ancestor that was part of an early and aggressive Procericambarus stock that was widespread in the pre-glacial Teays River and the upper reaches of the Tennessee River. If true, this upland stock could only have gained access to the Atlantic versant by means of a breach of the Blue Ridge and subsequent capture of some upper Teays headwaters by an east-flowing Piedmont stream. Ross (1969:283-290) persuasively argued that, in the area under consideration, this probably was accom- plished “in early Pleistocene time” by headwaters of the young Roanoke River. Subsequent southeastward dispersal of this putative ancestor in the Greater Roanoke system would have brought it into waters 84 John E. Cooper and Martha Riser Cooper contiguous or interdigitating with those of the adjacent Greater Pamlico River, later (probably late in the Pleistocene) to become the separate but twin systems now known as the Neuse and Tar-Pamlico rivers. Jenkins et al. (1971:45) indicated the presence of a theater of stream capture between the Roanoke and what are the headwaters of today’s Tar and Neuse rivers, which we interpret to have been in the area of present Person and Granville counties. It is reasonably parsimonious to conclude that it was through such piracies, from Teays to Greater Roanoke to Greater Pamlico, that the ancestor of O. carolinensis gained access to and became isolated in the latter system, there to evolve allopatrically into the present species. This dispersal, particularly between the contiguous Roanoke and Neuse-Tar systems where downslope differ- ences in elevation occurred and drainage divides were not of excessive magnitude, could have been abetted by the flooding that likely would have been prodigious during interglacial periods, and perhaps even by relatively minor tectonic events. Jenkins et al. (1971:82) postulated a generally similar dispersal history for the ancestor of N. furiosus. On the other hand, Sessions and Wiley (1985), on the basis of their karyological studies and electro- phoretic analyses provided by Ashton et al. (1980), considered Necturus lewisi to be the most primitive of the extant species of the genus Necturus , and the widespread western N. maculosus to be the most derived. They suggested that the Necturus stock initially spread south in the Atlantic Coastal Plain, then west through the Gulf regions and around the southern Appalachians, and finally north in the Ohio and Mississippi drainages. As earlier mentioned, O. carolinensis occupies the entire Tar- Pamlico watershed, but appears to be absent from some parts of the Neuse watershed, including nearly all of its Piedmont streams. If not a sampling deficiency, this could indicate that the initial entry of its ancestor into the Greater Pamlico River occurred in an extensive northernmost (Tar River) portion, and expansion of the species in the current Neuse basin is an ongoing process. If so, this stream dweller likely will never extend it range upriver into the Eno, Little, and Flat rivers, since the Neuse River has been impounded to create Falls Lake, which stretches for about 35 km (22 mi) from northwestern Wake County into Granville and Durham counties, and has converted most of the Neuse and its tributaries in those areas to lacustrine habitats. Etymology — Carolinensis , after North Carolina, to which the new species is endemic. Suggested vernacular name: North Carolina Spiny Crayfish. New Crayfish Species 85 ACKNOWLEDGMENTS — We are indebted to all those persons who collected or assisted in collecting crayfishes in the Neuse and Tar-Pamlico basins; their names are provided elsewhere in this paper. We are particularly indebted to Alvin L. Braswell and Ray E. Ashton, Jr., who supervised the field work for the N. lewisi project and made certain that crayfishes received considerable attention. The North Carolina Wildlife Resources Commission and the Office of Endangered Species, United States Fish and Wildlife Service, provided major funding for the N. lewisi project, and other funds were provided by the North Carolina State Museum of Natural Sciences. We thank Joseph F. Fitz- patrick for his invaluable review of the manuscript. JEC expresses his sincerest personal gratitude to J. E. Cooper, Jr., R. E. Ashton, Jr., L. Ferguson, J. R. Holsinger, G. Leake, J. Perry, W. J. Rishel, and his co-author, and especially to D. Howard and A. L. Braswell, without whose assistance this paper could never have been completed. Finally, we cannot adequately express our gratitude to the late Horton H. Hobbs, Jr., who provided invaluable guidance and construc- tive criticism in our studies of crayfishes. He reviewed an early draft of this paper. His generosity, although legendary, was ever a source of amazement, and he is sorely missed. LITERATURE CITED Ashton, R. E., Jr., A. L. Braswell, and S. I. Guttman. 1980. Electrophoretic analysis of three species of Necturus (Amphibia: Proteidae), and the taxonomic status of Necturus lewisi (Brimley). Brimleyana 4:43-46. Braswell, A. L., and R. E. Ashton, Jr. 1985. Distribution, ecology, and feeding habits of Necturus lewisi (Brimley). Brimleyana 10:13-35. Bundy, W. F. 1877. On the Cambari of northern Indiana. Proceedings of the Academy of Natural Sciences of Philadelphia 29:171-174. Cooper, J. E., and R. E. Ashton, Jr. 1985. The Necturus lewisi study: Introduction, selected literature review, and comments on the hydro- logic units and their faunas. Brimleyana 10:1-12. Cooper, J. E., and A. L. Braswell. 1995. Observations on North Caro- lina crayfishes (Decapoda: Cambaridae). Brimleyana 22:87-132. Cooper, M. R., and J. E. Cooper. 1977. A comment on crayfishes. Pages 198-199 in Endangered and threatened plants and animals of North Carolina (J. E. Cooper, S. S. Robinson, and J. B. Funderburg, edi- tors). North Carolina State Museum of Natural History, Raleigh. Faxon, W. 1884. Descriptions of new species of Cambarus, to which is added a synonymical list of the known species of Cambarus and Astacus. Proceedings of the American Academy of Arts and Sciences 20:107- 158. 86 John E. Cooper and Martha Riser Cooper Faxon, W. 1890. Notes on North American crayfishes, family Astacidae. Proceedings of the United States National Museum 12(785):619-634. Fitzpatrick, J. F., Jr. 1987. The subgenera of the crawfish genus Orconectes (Decapoda: Cambaridae). Proceedings of the Biological Society of Washington 100(1):44- 74. Harris, J. A. 1903. An ecological catalogue of the crayfishes belonging to the genus Cambarus. Kansas University Science Bulletin II (3) :5 1 — 187. Hobbs, H. H., Jr. 1972. Crayfishes (Astacidae) of North and Middle America. Biota of freshwater ecosystems identification manual 9. United States Environmental Protection Agency, Washington, D.C. Hobbs, H. H., Jr. 1974. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambaridae). Smithsonian Con- tributions to Zoology 166:1-161. Hobbs, H. H., Jr. 1975. New crayfishes (Decapoda: Cambaridae) from the southern United States and Mexico. Smithsonian Contributions to Zoology 201:1-34. Hobbs, H. H., Jr. 1981. The crayfishes of Georgia. Smithsonian Contri- butions to Zoology 318:1-549. Hobbs, H. H., Jr. 1989. An illustrated checklist of the American cray- fishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology 480:1-236. Hobbs, H. H., Jr., and D. J. Peters. 1977. The entocytherid ostracods of North Carolina. Smithsonian Contributions to Zoology 247:1-73. Hobbs, H. H., Jr., and M. Walton. 1958. Procambarus pearsei plumimanus , a new crayfish from North Carolina (Decapoda, Astacidae). Journal of the Elisha Mitchell Scientific Society 74:7-12. Jenkins, R. E., E. A. Lachner, and F. J. Schwartz. 1971. Fishes of the central Appalachian drainages: their distribution and dispersal. Pages 43-117 in The distributional history of the biota of the southern Ap- palachians, part III: vertebrates (P. C. Holt, editor). Research Divi- sion Monograph 4, Virginia Polytechnic Institute and State University, Blacksburg. Martof, B. S., W. M. Palmer, J. R. Bailey, and J. R. Harrison, III. 1980. Amphibians and reptiles of the Carolinas and Virginia. University of North Carolina Press, Chapel Hill. Ortmann, A. E. 1905. The mutual affinities of the species of the genus Cambarus, and their dispersal over the United States. Proceedings of the American Philosophical Society 44:91-136. Ortmann, A. E. 1931. Crawfishes of the southern Appalachians and the Cumberland Plateau. Annals of the Carnegie Museum 20:61-160. Rohde, F. C. 1980. Noturus furiosus, Noturus munitus, Noturus placidus, Noturus stigmosus. Pages 457, 465, 468, 469 in Atlas of North American freshwater fishes (D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr., editors). North Carolina State Museum of Natural History, Raleigh. New Crayfish Species 87 Ross, R. D. 1969. Drainage evolution and fish distribution problems in the southern Appalachians of Virginia. Pages 277-292 in The distri- butional history of the biota of the southern Appalachians, part III: vertebrates (P. C. Holt, editor). Research Division Monograph 4, Virginia Polytechnic Institute and State University, Blacksburg. Sessions, S. K., and J. E. Wiley. 1985. Chromosome evolution in sala- manders of the genus Necturus. Brimleyana 10:37-52. Taylor, W. R. 1969. A revision of the catfish genus Noturus Rafinesque with an analysis of higher groups in the Ictaluridae. United States National Museum Bulletin 282. Received 2 August 1995 Accepted 24 August 1995 Nine-banded Armadillo, Dasypus novemcinctus (Mammalia: Edentata), in South Carolina: Additional Records and Reevaluation of Status Steven G. Platt Department of Biological Sciences 132 Long Hall, Clemson University Clemson, South Carolina 29634-1903 AND William E. Snyder Center for Ecology, Evolution, and Behavior, and Department of Entomology S-255 Agricultural Science Building University of Kentucky, Lexington, Kentucky 40546-0091 ABSTRACT — The nine-banded armadillo (Dasypus novemcinctus Linnaeus) has been undergoing range expansion and is now estab- lished throughout much of the southeastern United States. Previ- ous records for South Carolina are widely scattered, and no evi- dence of an established population has been reported. We present an additional museum record, not previously reported, and field obser- vations of living and road-killed animals that strongly suggest a population of armadillos is established in southwestern South Carolina. This range extension into South Carolina probably occurred within the past ten years. Henceforth, the nine-banded armadillo should be considered an established member of South Carolina’s mammalian fauna. The nine-banded armadillo has been undergoing a natural expansion into the southeastern United States over the past 100 years. This expansion began when animals moved into Texas from northern Mexico in the mid-1800’s, perhaps in response to changing land use practices. By 1954 armadillos had reached the Mississippi River, and by 1972 were in the western Florida panhandle. Armadillos were also introduced into peninsular Florida between 1920 and 1936, and the two subpopulations merged in the mid-1970’s (Talmage and Buchanan 1954, Humphrey 1974). This species is now established in eight states (Texas, Oklahoma, Arkansas, Louisiana, Mississippi, Alabama, Florida, and Georgia), and is expected to continue to move northward and eastward until limited by low winter temperatures (Humphrey 1974). Brimleyana 23:89-93, December 1995 89 90 Steven G. Platt and William E. Snyder The distribution and status of the nine-banded armadillo in South Carolina is problematical. Webster et. al. (1985) stated that South Carolina might represent the northernmost limit of the armadillo’s expanding range, but concluded its status was uncertain. Hall’s (1981) range map included most of South Carolina based on three records from Golley (1966), which the latter believed to be translocated animals. Humphrey (1974) listed a single occurrence based on a widely circulated questionnaire, and Sanders (1978) reported ten records from scattered Coastal Plain and Piedmont locations. More recently Mayer (1989) summarized all previous state records, reported two additional animals, and 15 recent sightings listed by respondents to a questionnaire (Mayer 1989). Given the proximity of many previous records to major highways, some believe these reports represent escaped or released animals rather than pioneering individuals at the forefront of an expanding range (Golley 1966, Sanders 1978, Mayer 1989). Mayer (1989) concluded that because direct evidence of an established population is lacking, the status of the species in South Carolina remains uncertain. METHODS We present an additional museum record, not previously reported, and field observations from southwestern South Carolina. The museum specimen (Clemson University Vertebrate Collection #126) was collected 9 December 1978 on Port Lamar Road, Cessionville, Charleston County. Field observations were made in Jasper, Allendale, and Barnwell counties during April, May, August, and October 1995 (Table 1). Specific locality data for all records were deposited in the files of the Clemson University Vertebrate Collection (CUSC). RESULTS AND DISCUSSION Four road-killed and two living animals were found in five nights (ca. 9.5 hours) of collecting along a 17.6-km segment of Sandhills Road (County Road 119), due west of Tillman. Furthermore, numerous tracks and probe-holes made by foraging armadillos (Murie 1954) were noted at the Tillman Sand Ridge Natural Heritage Preserve on Sandhills Road. This road is a popular collecting location for reptile enthusiasts, and others also report frequent sightings of road-killed and living armadillos here (Todd Kuntz, United States Forest Service, personal communication). Another road-killed armadillo was found in Jasper County on Cohen Road (County Road 22), ca. 10 km northeast of Tillman. Two additional road-killed animals were also found in Barnwell and Allendale counties. We are unaware of any other reports of such a temporal and spatial Table 1. Summary of recent nine-banded armadillo (Dasvpus novemcinctus) records from southwestern South Carolina. Nine-banded Armadillo Records 91 H •> k C/5 M (-H G a D Pi x D C/3 G o ■rd o u o "o u o c r\ *o d bJO G a G T3 T3 03 CQ T3 D 00 CO 00 O =tfc D o o o c c-. on G u 00 u Td o o o C/2 g - E ° o dp O 3 « X Q 00 G O a) o cd Dh cd Vh cd O 5/5 p 03 0) pi T3 o h— » o o ”o a h— > o G C/2 G 03 E o 1— Td o • -"H i-H Q id D o o o o o G C/3 o G > -D G D 03 HO G G _T Gh td D W) bD G G E G Td id G PQ o 6 6 o o o o U U U U U U U Vh *-< i-H i-H (D o U —5 ^5 cd Dh rs CQ C/5 cd 'd ■d ■d ■d ~6 "d rs —5 Pi Pi Pi Pi Pi' Pi On CO *d G C/5 C/5 C/5 C/5 C/5 C/5 Pi o • f-H '4— * • i-H -G • *-H -C JG • hH -G • hH -G • F-H -G £ X G cd Td -a ■a T3 Td Td n Q n rH rH t-H — G 0 D _> < 1 92 Steven G. Platt and William E. Snyder concentration of records within this state. We believe these records strongly suggest that the nine-banded armadillo is now established in extreme southwestern South Carolina. The timing of this range extension into South Carolina remains speculative, but probably occurred within the past ten years. Humphrey (1974) places the northernmost range limit in the Coastal Plain near McIntosh County, Georgia, approximately 80 km south of the Savannah River. In the absence of physical or climatic barriers, an average invasion rate of four to 10 km per year has been estimated (Humphrey 1974). This model predicts an extension into South Carolina by the early to mid-1980’s. The Savannah River could have slowed immigration, but armadillos are known to cross major rivers by swimming or bottom walking (Talmage and Buchanan 1954). Wright (1982) did not find armadillos among mammals using gopher tortoise ( Gopherus polyphemus) burrows on Tillman Sand Ridge Natural Heritage Preserve, although they are reported to inhabit tortoise burrows elsewhere (Jackson and Milstrey 1989). Thus, we speculate armadillos did not become established in Jasper County until 1985 or later. This coincides with the 1985 and 1986 records reported by Mayer (1989) further north in Aiken and Barnwell counties. Invasion is especially rapid in river valleys, and the Savannah River may be functioning as a dispersal corridor. Range expansion is expected throughout South Carolina, with the ex- ception of the mountainous northwestern corner of the state where low temperatures likely preclude overwinter survival (Humphrey 1974, Mayer 1989). CONCLUSIONS It appears that numbers of armadillos are present in at least one area of southwestern South Carolina. Further range expansion can be expected in the state, particularly throughout the Savannah River drainage and Coastal Plain. Henceforth the nine-banded armadillo should be considered an established member of South Carolina’s mammalian fauna. ACKNOWLEDGMENTS — Stanlee Miller provided access to the Clemson University Vertebrate Collection. John Scavo, Todd Kuntz, Hong Liu, and B. Anne Ditte assisted in field work. Comments by Richard R. Montanucci and David Lee greatly improved this manuscript. Nine-banded Armadillo Records 93 LITERATURE CITED Galbreath, G. J. 1982. Armadillo, Dasypus novemcinctus. Pages 71-79 in Wild mammals of North America (J. A. Chapman and G. A. Feldhamer, editors). John Hopkins University Press, Baltimore, Maryland. Golley, F. B. 1966. South Carolina mammals. Charleston Museum Press, Charleston, South Carolina. Hall, E. R. 1981. The mammals of North America. Volume 1. John Wiley and Sons, New York, New York. Humphrey, S. R. 1974. Zoogeography of the nine-banded armadillo ( Dasypus novemcinctus ) in the United States. Bioscience 24:457-462. Jackson, D. R., and E. C. Milstrey. 1989. The fauna of gopher tortoise burrows. Pages 86-98 in Proceedings of the gopher tortoise reloca- tion symposium. (J. E. Diemer, D. R. Jackson, J. L. Landers, J. N. Layne, and D. A. Wood, editors). Florida Nongame Wildlife Program Technical Report 5, Florida Game and Freshwater Fish Commission, Tallahassee, Florida. Mayer, J. J. 1989. Occurrence of the nine-banded armadillo, Dasypus novemcinctus (Mammalia: Edentata), in South Carolina. Brimleyana 15:1-5. Murie, O. J. 1954. A field guide to animal tracks. Houghton Mifflin Company, Boston, Massachusetts. Sanders, A. E. 1978. Mammals. Pages 296-308 in An annotated check- list of the biota of the coastal zone of South Carolina. (R. G. Zingmark, editor). University of South Carolina Press, Columbia. Talmage, R. V., and G. D. Buchanan. 1954. The armadillo ( Dasypus novemcinctus ): A review of its natural history, ecology, anatomy, and reproductive physiology. Rice Institute Pamphlet 41:1-135. Webster, W. D., J. F. Parnell, and W. C. Biggs, Jr. 1985. Mammals of the Carolinas, Virginia, and Maryland. University of North Carolina Press, Chapel Hill. Wright, J. S. 1982. Distribution and population biology of the gopher tortoise, Gopherus polyphemus, in South Carolina. Masters Thesis. Clemson University, Clemson, South Carolina. Received 24 October 1995 Accepted 31 October 1995 Post-hibernation Movement and Foraging Habitat of a Male Indiana Bat, Myotis sodalis (Chiroptera: Vespertilionidae), in Western Virginia Christopher S. Hobson1 Department of Biology, Tennessee Technological University Cookeville, Tennessee 38505 AND J. Nathaniel Holland2 Division of Life Sciences, Ferrum College F err urn, Virginia 24088 ABSTRACT — We investigated departure patterns of bats from a hibernaculum and use of tree roosts and foraging habitat by a male Indiana bat {Myotis sodalis ) in western Virginia with radio-telem- etric techniques, cave population surveys, and habitat assessment. Although hibernating Indiana bats are well documented in Virginia, our study is the first to report foraging and roosting habitat in Virginia, which is along the eastern periphery of the range of the species. After departure from the hibernaculum in late April, a radio-tagged male M. sodalis moved 16 km southwest where it remained for two weeks until the radio transmitter failed. This bat used a ma- ture, live, shagbark hickory {Carya ovata) tree as a diurnal roost; up to 10 other bats roosted in the same tree. The bat primarily foraged among tree canopies within 625-ha area of an 80-year-old, oak-hickory forest. Our study suggests that male M. sodalis use foraging areas and tree roosts found in the area of hibernacula. Thus, we recommend that conservation efforts protect and manage foraging and tree roosting habitat in the vicinity of M. sodalis hibernacula. Approximately 85% of all Indiana bats {Myotis sodalis Miller and Allen) hibernate in seven caves located in Missouri, Indiana, and Kentucky (Harvey 1992). Indiana bats are most common in the Midwest, with peripheral populations in northeastern (e.g., Pennsylvania), Atlantic (e.g., Virginia), southeastern (e.g., Georgia), and northern midwestern (e.g., Michigan) states (Humphrey 1978). Peripheral populations may become increasingly important in the management of this federally 1 Present Address: Department of Conservation and Recreation, Division of Natural Heritage, 1500 East Main Street, Suite 312, Richmond, Virginia 23219. 2 Present Address: Department of Biology, University of Miami, P.O. Box 249118, Coral Gables, Florida 33124-0421. Brimleyana 23:95-101, December 1995 95 96 Christopher S. Hobson and J. Nathaniel Holland endangered species if larger populations continue to decline. Populations of M. sodalis hibernating in Virginia caves typify a peripheral population, as they occur on the eastern edge of the range and represent <1% of the total estimated population (Dalton 1987). Ten known M. sodalis hibernacula occur in Virginia accounting for 2,500 individuals. Knowledge of summer foraging areas and roost sites of M. sodalis is as important to conservation efforts as identification and protection of hibernacula, but the former remain poorly documented (Humphrey et al. 1977, Thomson 1982). Most information on summer foraging and roosting habitat of M. sodalis comes from the central area of the range of the species (i.e., midwestern United States), with an emphasis on maternity sites. In Illinois and Indiana, female M. sodalis roost in several species of trees including shagbark hickory ( Carya ovata), bitternut hickory (C. cordiformis), green ash ( Fraxinus pennsylvanica), eastern cottonwood ( Populus deltoides), northern red oak ( Quercus rubra), post oak ( Q . stellata), shingle oak (Q. imbricaria), and sycamore {Platanus occidentalis) (Humphrey et al. 1977, Gardner et al. 1990, Kurta et al. 1993). In addition, Humphrey et al. (1977) identified a maternity roost under loose bark of a dead bitternut hickory tree. Females and newly volant young M. sodalis forage in riparian habitat, along the edge of floodplain forest and within forest canopies (Humphrey et al. 1977, Laval et al. 1977); however, roosting and foraging habitats of male M. sodalis are less well known. Observations in Missouri indicate that males forage along ridges and hillsides around forest canopies (Laval et al. 1977). Migratory behavior may differ among male and female M. sodalis (Hall 1962). Females migrate from hibernacula to maternity sites, whereas males either move away from or remain near the hibernacula during spring and summer. This suggests that suitable foraging and roosting habitat for M. sodalis occurs in the vicinity of some hibernacula. Although the distribution and abundance of M. sodalis hibernating in Virginia caves has been well documented, no summer roost sites, maternity colonies, or summer foraging habitats have been identified in the state prior to our study. Our objectives were to determine if male M. sodalis wintering in a Virginia cave remained in the vicinity of the hibernaculum during spring and summer months, and to charac- terize foraging and roosting habitats of male M. sodalis. METHODS We studied a M. sodalis colony that hibernates in a cave in Bath County, Virginia. Bath County is located in a rural area of the Appalachian Mountains in western Virginia within the George Washing- Male Indiana Bat 97 ton National Forest. An active timber management program is practiced in this forest. We monitored emergence and departure patterns at the cave entrance between March and early May 1993. Using night-vision goggles, emergence patterns at the cave entrance were monitored by observing the numbers of individuals entering and leaving the cave. Mist nets (one net covering cave entrance) and harp traps were used to census species emerging from the cave (7 net nights, from sunset to approximately 0100 hours; one net night = 1 open net per night). We used head lamps with red filters and infrared goggles to aid in estimating numbers of individuals for all species in the cave. Myotis lucifugus, M. septentrionalis, M. leibii, Eptesicus fuscus, Pipistrellus subflavus , and M. sodalis were known to hibernate in the cave. The abundance of M. sodalis in the cave was determined five times (3, 17, 24, 28 April, and 1 May) to establish spring departure patterns for this species. On 28 April 1993, two male M. sodalis were captured in the cave and fitted with 0.65-g radio transmitters (Holohill Systems, Ltd., 3387 Stonecrest Road, Woodlawn, Ontario, Canada). Transmitters were equivalent to 6% of the body mass of each bat. Transmitters were attached between the shoulder blades with eyelash glue (no hair was removed). Each bat was placed in a cloth sack for approximately 30 minutes to allow the glue to dry before they were released. The bats were released 15 minutes apart approximately 200 m south of the cave entrance, and tracked with radio receivers (Wildlife Materials, Model TRX2000S, Route 1 Box 427A, Carbondale, Illinois 62901) for the life of the transmitter batteries. Triangulation techniques using two or three observers, direct observation, and the receiver’s attenuator were used to identify the roost site, to delineate foraging areas, and to identify movement patterns. Vegetative characteristics of foraging and roosting habitats were assessed with the point-quarter sampling technique (Brower et al. 1989) and visual observation. Points of vegetative sampling occurred along seven 100-m transects. For each transect, five points were determined, and four trees were sampled at each point for a total of 140 trees sampled for all seven transects. Additional data were collected at the tree roost using mist nets (2 net nights; see Gardner et al. 1989 for description of mist netting system) and infrared goggles, the latter to determine activity patterns (time of emergence from roost) and numbers of bats associated with the tree roost. The tree roost was not mist netted due to its height above ground and the steep slope of the terrain. Forest habitat, streams, and roads surrounding foraging and roosting habitats were sampled with mist nets (11 net nights). 98 Christopher S. Hobson and J. Nathaniel Holland RESULTS AND DISCUSSION In winter 1992, 38 M. sodalis and 1,686 bats were hibernating in the cave (Leffler et al. 1993). Few individuals left the cave in early April, with most bats departing by early May. Trapping at the cave entrance resulted in 56 captures (all adults), including 19 Myotis lucifugus, 17 M. septentrionalis, 5 M. leibii, 4 Eptesicus fnscus , and 11 Pipistrellus subflavus. M. sodalis were not captured during this period. The 56 bats captured account for only 4% of the hibernating population of bats in the cave. Assuming that no M. sodalis had left the cave before our first census, the M. sodalis population in the cave declined from 31 to 18, 8, 6, and 0 individuals on 3, 17, 24, 28 April and 1 May, respectively. By mid-April the single cluster of M. sodalis had broken into several small clusters within a 2-m2 area. Cope and Humphrey (1977) reported similar trends in departure patterns of M. sodalis, where females left the hibernaculum before males, and most bats departed by late April and early May. Two radio-tagged male M. sodalis were tracked for approximately one hour after release near the cave, at which time signals of both individuals were lost. Bat #440 was never located from the ground after release, but its signal was detected by an aircraft in the cave area on 8 and 10 May. Bat #458 was relocated by ground in the George Washington National Forest on 1 May, approximately 16 km SW of the hibernaculum. Bat #458 roosted and foraged in George Washington National Forest until 20 May when the transmitter battery failed. For 19 nights, bat #458 roosted on a north facing slope (0° to 5° east of north) at 700 m elevation, beneath the bark of a mature shagbark hickory (ca. 30 m in height, 61 cm DBH). The bat roosted at a height >8 m in the shagbark hickory. Other tree species within a 10-m radius of the roost tree included basswood (Tilia spp.), red maple (Acer rubrum), eastern hophornbeam (Ostrya virginiana), tulip poplar (Liriodendron tulipifera), and pignut hickory (Carya glabra ). Bat #458 was not the only bat using the shagbark hickory for a roost; on separate occasions at dusk 5, 10, and 3 bats emerged from the roost tree. Leaving the roost tree around 2030 hours nightly, bat #458 was one of the first bats to emerge, with the other bats emerging by 2100 hours. Bat #458 immediately left the area of the roost tree upon emergence and flew to foraging habitat, located within 1 km of the roost tree. Mist netting the roost site and foraging habitat resulted in capture of 2 Lasionycteris noctivagans, 3 Lasiurus borealis, 4 Eptesicus fuscus, 6 Pipistrellus subflavus, 6 Myotis lucifugus, and 3 M. septentrionalis', but, no M. sodalis were captured. After emerging from the roost tree, bat #458 foraged persistently Male Indiana Bat 99 throughout the night, often until after 0200 hours. On 2 of 19 nights, the bat ceased foraging for approximately one hour around 2300 hours. Foraging habitat for bat #458 encompassed approximately 625 ha. A small, two-lane road and an unimproved forest service road transected the habitat. A small first-order stream ran parallel to the two-lane road. Foraging habitat was an 80-year-old, mature oak-hickory mixed deciduous forest with a conifer component (Table 1). When foraging in the 625 ha habitat, this bat spent the majority of its time flying in an elliptical pattern at canopy height. The ellipse was transected by a small two-lane road, which was occasionally used as a flyway. Bat #458 was also observed flying in an elliptical pattern along a ridge containing a patch of mature hemlocks. On two occasions at dusk, the bat was observed foraging along a water course within 0.5 km of the roost site. In Missouri, Laval et al. (1977) found that male M. sodalis forage in elliptical patterns among treetops of dense forest along ridges and hillsides instead of over water. Humphrey et al. (1977) also reported M. sodalis foraging around tree canopies. Table 1. Diameter at breast height (DBH), density, and relative frequency of tree species within the foraging area of a male M. sodalis in Bath County, Virginia. A total of 140 trees were sampled within the foraging habitat. Tree Species Mean (: DBH t SE) (cm) Range Density Relative (Trees/ha) Frequency Acer rubrum 13.3 + 2.0 33 - 8 66 0.091 Carya glabra 22.5 + 1.3 24 - 20 17 0.027 C. ovata 20.0 + 3.0 30 - 14 28 0.045 C. tomentosa 30.0 + 3.5 38 - 21 50 0.082 Cornus florida 11.8 + 1.0 17 - 9 50 0.064 Fagus grandifolia 17.3 + 4.3 38 - 10 33 0.045 Juglans nigra 19.5 + 3.3 27 - 10 28 0.027 Liriodendron tulipifera 37.8 + 8.8 84 - 9 50 0.082 Pinus strobus 23.5 + 2.5 46 - 9 89 0.091 Quercus rubra 40.0 + 3.8 72 - 9 94 0.127 Q. alba 36.8 + 4.0 52 - 24 39 0.055 Tilia spp. (unidentified) 23.0 + 2.5 32 - 14 39 0.064 Tsuga canadensis 28.3 + 3.8 64 - 8 111 0.136 other1 12.5 + 1.8 20 - 8 44 0.064 1 Includes infrequent occurrence of Acer pennsylvanicwn, Betula lenta, Carpinus caroliniana, and Ulmus rubra. 100 Christopher S. Hobson and J. Nathaniel Holland CONCLUSIONS Although foraging habitat and roost site use by M. sodalis in Virginia are based on data from one bat, our study is consistent with studies conducted in other geographic regions. Because the male M. sodalis remained in the vicinity of the hibernaculum during spring, our study suggests that foraging areas and tree roosting habitats for male M. sodalis may be found near hibernacula. Conservation practices dictate that identification and protection of roosting and foraging habitat is necessary for bat conservation efforts. ACKNOWLEDGMENTS— We thank J. W. Leffler, R. Reynolds, L. West, and L. Miller for assistance with field work, and the U.S. Forest Service for providing needed resources. This research was funded by the Non-game Division, Virginia Department of Game and Inland Fisheries, and the Center for Protection, Utilization, and Management of Water Resources, Tennessee Technological University. We are grateful for comments provided by J. R. Belthoff, J. W. Leffler, R. Reynolds, and S. M. Roble on a previous version of this manuscript. LITERATURE CITED Brower, J. E., J. H. Zar, and C. N. von Ende. 1989. Field and labora- tory methods for general ecology. Third Edition. Wm. C. Brown Publishers, Iowa. Cope, J. B., and S. R. Humphrey. 1977. Spring and autumn swarming behavior in the Indiana bat, Myotis sodalis. Journal of Mammalogy 58(1):93— 95. Dalton, V. M. 1987. Distribution, abundance, and status of bats hiber- nating in caves in Virginia. Virginia Journal of Science 38:369-379. Gardner, J. E., J. D. Garner, and J. E. Hofman. 1989. A portable mist netting system for capturing bats with emphasis on Myotis sodalis (Indiana bat). Bat Research News 30:1-7. Gardner, J. E., J. D. Garner, and J. E. Hofman. 1990. Progress report: 1989 and 1990 investigations of Myotis sodalis (Indiana bat) distri- bution, habitat use, and status in Illinois. Unpublished report. United States Fish and Wildlife Service, Twin Cities, Minnesota. Hall, J. S. 1962. A life history and taxonomic study of the Indiana bat, Myotis sodalis. Reading Public Museum and Art Gallery, Scientific Publication 12:1-68. Harvey, M. J. 1992. Bats of the eastern United States. Arkansas Game and Fish Commission, United States Fish and Wildlife Service, and Tennessee Technological University. Humphrey, S. R. 1978. Status, winter habitat, and management of the endangered Indiana bat, Myotis sodalis. Florida Scientist 41(2):65-76. Male Indiana Bat 101 Humphrey, S. R., A. R. Richter, and J. B. Cope. 1977. Summer habitat and ecology of the endangered Indiana bat, Myotis sodalis. Journal of Mammalogy 58(3):334- 346. Kurta, A., J. Kath, E. L. Smith, R. Foster, M. W. Orick, and R. Ross. 1993. A maternity roost of the endangered Indiana bat ( Myotis sodalis ) in an unshaded, hollow, sycamore tree ( Platanus occidentalis ). Ameri- can Midland Naturalist 130(2):405-407. Laval, R. K., R. L. Clawson, M. L. Laval, and W. Caire. 1977. Forag- ing behavior and nocturnal activity patterns of Missouri bats, with em- phasis on the endangered species Myotis grisescens and Myotis sodalis. Journal of Mammalogy 58(4):592-599. Leffler, J. W., R. Powers, C. Hobson, R. Reynolds, G. Nussbaum, J. Hol- land, and K. Terwilliger. Bat investigations annual report. Pages 55- 71. In Virginia nongame and endangered wildlife investigations. (K. Terwilliger, editor). Virginia Department of Game and Inland Fish- eries, Richmond. Thomson, C. E. 1982. Myotis sodalis. Mammalian Species 163:1-5. Received 13 September 1995 Accepted 12 November 1995 5 % *■ ■ \ . • i . ■ • - - *? 1 - • » ’ ' - ' ; - ' : ■ : •- ■ ■ • , ■ - ■ ■ '■ . / ' I ■ • ! . • • The Milliped Family Hirudisomatidae in the New World (Polyzoniida) Rowland M. Shelley North Carolina State Museum of Natural Sciences, P. O. Box 29555, Raleigh, North Carolina 27626-0555 Abstract — In the New World, the milliped family Hirudisomatidae is represented by Octoglena Wood, with five species, and the monotypic new genus, Mexiconium. Octoglena bivirgata Wood, O. anura (Cook), n. comb., and O. prolata, n. sp., are contiguous along the Pacific Coast from British Columbia to Santa Cruz County, California; O. sierra, n. sp., is a localized, allopatric species in the Sierra Ne- vada foothills; and O. gracilipes (Loomis), n. comb., occurs in the eastern United States from South Carolina to Tennessee and Ala- bama. Mexiconium absidatum, n. sp., the first record of the fam- ily from Mexico, occurs at a high elevation in the Sierra Madre Oriental, Vera Cruz. Octoglena bivirgata displays three dark dor- sal stripes, and M. absidatum exhibits a dark, middorsal stripe; the other species are pale yellowish to white. The following new syn- onymies are proposed: Hypozonium Cook and Euzonium Chamberlin under Octoglena, and H. arnaudi and E. crucis, both by Chamberlin, under O. bivirgata. The Hirudisomatidae represents an Ancient Holarctic faunal assemblage that spread across North America from east to west, and southward into Mexico, and has experienced consider- able extinction. Octoglena is one of five Nearctic genera exhibit- ing east/west disjunctions, and a secondary center of evolution exists along the Pacific Coast. Relationships within Octoglena are gracilipes + ( sierra + (anura + (prolata + bivirgata))). In the Western Hemisphere, the milliped order Polyzoniida is represented by the pantropical family Siphonotidae, with two genera in South America and the common synanthrope, Rhinotus purpureus (Pocock), in the West Indies, Florida, Louisiana, and Central America (Hoffman 1977, 1980), and the Holarctic families Polyzoniidae and Hirudisomatidae, in the eastern and western United States. The latter spreads northward into coastal British Columbia, and an allopatric species occurs in the Sierra Madre Oriental, Vera Cruz, Mexico (Fig. 1). The east-Nearctic polyzonioid fauna comprises six species of Poly- zonium Brandt (Polyzoniidae) (Loomis 1971; Shelley 1976, 1988) and one hirudisomatid that was erroneously placed in this genus and family. The western fauna of this order has never been examined and presently consists of seven genera and ten nominal species (Chamberlin 1954, Brimleyana 23:103-143, December 1995 103 104 Rowland M. Shelley Fig. 1. Distribution of the Hirudisomatidae in the New World, showing the areas occupied by Octoglena in the eastern and western United States and British Columbia, and the single site of Mexiconium, denoted by the dot and arrow, in Mexico. The dot in California represents the two localities of O. sierra. Chamberlin and Hoffman 1958). Hoffman (1980) and Shelley (1988) referred Octoglena and O. bivirgata , both authored by Wood, to the western fauna and the Hirudisomatidae, because its striped color pattern (Wood 1864, 1865) is displayed by two California hirudisomatids, Euzonium crucis and Hypozonium arnaudi, both authored by Chamberlin, whereas no eastern polyzonioid is so marked. Studies are progressing on the west-Nearctic Polyzoniidae, so this contribution addresses the Hirudisomatidae and transfers the east-Nearctic representative into Octo- glena; I also erect a new genus, Mexiconium , to accommodate the Mexican species. Octoglena is therefore a continental taxon and similar to Brachycybe Wood (Platydesmida: Andrognathidae), Orinisobates Lohmander (Julida: Nemasomatidae), Scytonotus Koch (Polydesmida: Polydesmidae), and Ergodesmus Chamberlin (Polydesmida: Nearctodes- midae) in exhibiting east/west disjunctions (Fig. 1) (Gardner 1975, Enghoff 1985, Shelley, 1993, 1994a). I do not address here the larger Hirudisomatid Millipeds 105 question of the distinction(s) between the New World hirudisomatids and the European genus, Hirudisoma Fanzago, which requires comparative material of its eight species (keyed by Mauries 1964), nor do I assess differences with Orsiboe Attems and Kiusiozonium Verhoeff in Japan, and the former may belong to another family (Hoffman 1980, personal communication). Polyzonioid gonopods tend to be structurally conservative and lack the dramatic elaborations of polydesmoids that typically form the bases of generic diagnoses; consequently, genera are often distin- guished by subjective somatic features. Octoglena gracilipes (Loomis), in the eastern United States, differs somatically from the western species in its narrower telson and the absence of a slight caudolateral extension to the midbody metatergites (Figs. 8, 12, 23-24). Its anterior gonopod differs from those of the Pacific Coastal species, but this anatomical gap is bridged by O. sierra , n. sp., in the Sierra Nevada, which occupies an intermediate geographical position. Separate generic status for gracilipes could be based on the somatic features, but I think these differences are insignificant when compared to the gonopodal linkage that unites gracilipes with the Pacific Coastal components; I therefore opt for a single genus, for which Octoglena is the oldest available name. Similarly, the anterior gonopod of the Mexican hirudisomatid resembles that of O. sierra , but its somatic differences are much greater and, in my view, require generic recognition. The metatergites do not extend laterad, and the body form is fundamentally different, the segments being narrower and more vaulted than the flattened, “bell shaped” segments of Octoglena (Figs. 7, 35). There is also one broad, middorsal stripe rather than three, as in O. bivirgata (Figs. 2, 31-33), and there are two pairs of ocelli rather than three (Figs. 6, 22, 34). Hirudisomatids are not readily distinguished from polyzoniids, as most characters have exceptions. For example, the caudal metatergal margins of O. bivirgata are strongly upturned and clearly differentiate it from the flush condition in sympatric polyzoniids, but this feature is less distinctive for O. anururn and O. gracilipes , which can be confused with polyzoniids. The margins of M. absidatum are slightly elevated but not upturned, and hence resemble the closely appressed tergites of polyzoniids. West-Nearctic hirudisomatids are diagnosed by the broad telson, but this structure is narrower in O. gracilipes and subequal in breadth to that of species of Polyzonium (Figs. 12, 24). The collum overhangs the epicranium and the uppermost ocelli in hirudisomatids, but it likewise overlaps part of the head in some western polyzoniids and thus does not discriminate the families. The 106 Rowland M Shelley ywwwB ■“r/j&LSrr •u!^ig»wra^.iT>T- .I^JIUWW >v?r«p» Fig. 2. Octoglena bivirgata. Dorsal view of female from Marin County, California Scale line = 1.00 mm. Hirudisomatid Millipeds 107 only inviolable characters involve the size and position of the penes on the second male coxae, which must be dissected and examined under a compound microscope. In his key to North American diplopod families, Hoffman (1990) placed the penes on the ventral coxal surface in the Hirudisomatidae and caudal to this podomere in the Polyzoniidae. To determine the correct family for Mexiconium, I had to examine the penes and compare their shape and location with these attributes in definite representatives of both families. As shown in figs. 3-4, the short, subconical penes of M. absidatum resemble those of O. bivirgata, and both are positioned caudoventrad on the coxae; in Poly- zonium rosalbum (Cope), however, the structures are longer, “bottle shaped,” and arise more dorsad (Fig. 5). Enghoff and Golovatch (1995) provide SEM photos of a European hirudisomatid and polyzoniid that also show the penes in these positions, so couplet 16a of Hoffman’s key (1990) should be amended to read “caudoventrad” as to the location of the penes in the Hirudisomatidae. Acronyms of sources of preserved study material are as follows: AMNH — American Museum of Natural History, New York, New York. CMN — Canadian Museum of Nature, Ottawa, Ontario, FSCA — Florida State Collection of Arthropods, Gainesville. NCSM — North Carolina State Museum of Natural Sciences, Raleigh. NMNH — National Museum of Natural History, Smithsonian Institution, Washington, DC. RBCM — Royal British Columbia Museum, Victoria. VMNH — Virginia Museum of Natural History, Martinsville. WAS — Private collection of William A. Shear, Hampden-Sydney, Virginia. Literature Review The history of the Hirudisomatidae in the New World begins with the proposal of Octoglena by Wood (1864) for O. bivirgata, a new species with fuscous stripes believed to inhabit the mountains of Georgia. He (Wood 1865) repeated these accounts and provided illustrations of the ventral and dorsal surfaces of the head. In the only other reference of the 19th century, Bollman (1893) included Octoglena in a key to North American myriapod genera. In the twentieth century, Cook (1904) proposed Hypozonium for H. anurum, a new species from Seattle, King County, Washington, and Chamberlin (1911) recorded it from Bremerton, Kitsap County. Cook and Loomis (1928) reiterated these records and transferred bivirgata into Polyzonium, thereby relegating Octoglena to the generic synonymy, 108 Rowland M. Shelley where it remained for 52 years. Chamberlin (1950) erected Euzonium for E. crucis, a striped species from Felton, Santa Cruz County, California, and (1954) proposed Hypozonium arnaudi, for a striped female from this locality that was taken on the same date and by the same collector as the type of E. crucis. Chamberlin and Hoffman (1958) included Octoglena in the synonymy of Polyzonium and reported Euzonium, Hypozonium, E. crucis, and H. anurum, inadvertently omitting H. arnaudi. However, Buckett (1964) included this species in his listing of California diplopods, and Jeekel (1971) cited all the genera along with their type species. Hoffman (1980) assigned Hypozonium and Euzonium to the Hirudisomatidae and revived Octoglena for a California hirudisomatid, because the collector, John Lawrence LeConte, sampled in California as well as Georgia, and because the striped pigment pattern fits Californian, rather than Georgian, polyzonioids. Hoffman (1980) and Shelley (1988) suggested that O. bivirgata Wood may be a senior name for E. crucis and H. arnaudi, which the present study confirms. Kevan (1983) reported H. anurum from unspecified sites in British Columbia, the first ordinal records from western Canada, and Kevan and Scudder (1989) included the milliped in their key to Canadian myriapods. Shelley (1990) reported five localities for H. anurum in the southwestern corner of the British Columbia mainland, which were reiterated by Scudder (1994). Key to North American Families of the Polyzoniida (adapted from that by Hoffman (1990)) I. Tarsal claws with prominent, overhanging paronychium; animals relatively quick and active, color pink; south Florida and Louisiana. Siphonotidae Tarsal claws simple, without paronychium; animals relatively sluggish, color pale white, yellowish, or with one or three dark longitudinal stripes 2 2. Caudal edges of metaterga detached from succeeding tergite, elevated or variably upturned; telson broad or narrow; penes short and subconical, located caudoventrad on 2nd male coxae (Figs. 3-4, 8, 12) Hirudisomatidae Caudal edges of metaterga not upturned, smoothly overlying and closely appressed to succeeding tergite; telson narrow; penes relatively long, located well dorsad on caudal surface of 2nd male coxae (Fig. 5) Polyzoniidae Family Hirudisomatidae Genus Octoglena Wood Octoglena Wood, 1864:186; 1865:229. Bollman, 1893:117, 137, 187. Hirudisomatid Millipeds 109 Figs. 3-5. Comparison of penes, caudal views of 2nd male coxae. 3, O. bivirgata , Sonoma County, California. 4, M. absidatum holotype. 5, Polyzonium rosalbum, Dade County, Georgia. Scale line = 0.25 mm for all figs. 110 Rowland M. Shelley Jeekel, 1971:41. Hoffman, 1980:73. Hypozonium Cook, 1904:62. Cook and Loomis, 1928:17. Chamberlin and Hoffman, 1958:187. Buckett, 1964:29. Jeekel, 1971:39. Hoffman, 1980:73. Kevan, 1983:2962. NEW SYNONYMY. Euzonium Chamberlin, 1950:1. Chamberlin and Hoffman, 1958:187. Buckett, 1964:29. Jeekel, 1971:38. Hoffman, 1980:73. NEW SYNONYMY. Type species — Of Octoglena, O. bivirgata Wood, 1864, by monotypy; of Hypozonium , H. anurum Cook, by monotypy; of Euzonium , E. crucis Chamberlin, 1950, by original designation. Diagnosis — Dorsum granular, with or without three dark, longitudinal stripes; caudal metatergal margins detached from succeeding tergite, upturned to varying degrees, body broad (W/L ratio generally 28- 38%), flattened “bell shaped” in profile, sides extending strongly laterad; collum broad, overhanging epicranium and at least one pair of ocelli; telson variable, either broad and comprising nearly entire caudal width, or relatively narrow and comprising around half of caudal width; head subtriangular, with three ocelli on each side arranged linearly in angular rows beginning at levels of antennal sockets; sternum of anterior gonopods with strong, apically hirsute lobes, segregated to varying degrees; anterior gonopods curving variably anteromediad distad, ultimate podomere divided, with broad, hirsute, ventral lobe of variable size usually over- hanging dorsal glabrous branch, latter either short, broad, and apically blunt, curved slightly ventrad, and directed anteromediad, or long, narrow, and acuminate, slightly sinuate and directed sublaterad; corners of 4th and 5th podomeres extended on caudal side; coxa with or without hirsute anterior lobe, length variable; posterior gonopod with ultimate podomere simple and acicular, apically acuminate, fimbriate, or lightly hirsute, projecting anteriad between solenomere and ventral lobe. Species — Five. Distribution — Along the Pacific Coast from the southwestern corner of the British Columbia mainland to central Santa Cruz County, California, extending inland to the western slope of the Cascade Mountains from British Columbia to central Oregon and the eastern slope of the Coast Range in southern Oregon and California, with a localized allopatric species some 75 mi (120 km) to the east in the Sierra Nevada foothills, Placer County, and one 1,897 mi (3,035 km) farther east in the eastern United States, extending from westcentral South Carolina to southcentral Tennessee and northwestern Alabama (Figs. 1, 28-29). The coastal species are contiguous and demonstrate parapatric spatial relationships and a sublinear, north to south, arrangement, the area being wider from central Oregon northward. Dimensions are approximately 850 Hirudisomatid Millipeds 111 mi (1,360 km), north/south, and 110 mi (176 km), east/west, for the coastal area, and 220 mi (352 km), north/south, and 380 mi (608 km), east/west, for the eastern. Relationships — With the somatic differences and the apomorphic absence of the coxal lobe, O. gracilipes is sister to the western species (Fig. 30). Octoglena sierra is sister to those along the Pacific Coast, which share the configuration of the ultimate anterior gonopod podomere, and O. anura, with the short, apically linear coxal lobe, is sister to 0. prolata plus O. bivirgata, in which the lobe is extended and apically rounded. Remarks — The anterior gonopods of each species are quite uniform and show little intraspecific variation, so I analyze segment numbers and lengths of measurable specimens in the variation sections. As per the recommendation of Enghoff et al. (1993), I exclude the telson from the segment counts. Key to Species of Octoglena 1. Caudolateral corners of midbody metatergites slightly but distinctly extended and rounded; telson broad, extending for nearly entire breadth of caudal extremity; anterior gonopod coxae with hirsute lobes of varying lengths (Figs. 9, 13, 16, 19); southwestern British Columbia to central California 2 Caudolateral corners of midbody metatergites not extended, continuous with middorsal margins, apically blunt; telson narrow, comprising no more than half of breadth of caudal extremity; anterior gonopod coxae without lobes (Fig. 25); North and South Carolina to Tennessee and Alabama gracilipes (Loomis) 2. Ultimate podomere of anterior gonopod with broad, distinct lobe ventral to and overhanging dorsal glabrous branch; latter rela- tively short and broad, curved slightly ventrad, directed antero- mediad (Figs. 9, 13, 16); along Pacific Coast from British Columbia to Santa Cruz County, California 3 Ultimate podomere of anterior gonopod with at most short, indistinct, ventral lobe, not overhanging dorsal branch; latter relatively long and narrow, slightly sinuate, directed sublaterad (Fig. 19); western Placer County, California sierra , new species 3. Coxal lobe of anterior gonopod short and apically sublinear, not overlying distal podomeres; ultimate podomere of posterior gonopod apically narrow and with numerous short hairs (Figs. 13, 15); southwestern British Columbia to northern Douglas County, Oregon anura (Cook) 112 Rowland M. Shelley Coxal lobe of anterior gonopod long and apically rounded, overlying distal podomeres; ultimate podomere of posterior gonopod apically fimbriate (Figs. 9, 11, 16, 18) 4 4. Coxal lobe of anterior gonopod leaning slightly mediad, barely overlapping distal podomeres; dorsum pigmented, with three dark, longitudinal stripes (Figs. 2, 9); Curry County, Oregon, to Santa Cruz County, California bivirgata Wood Coxal lobe of anterior gonopod leaning strongly laterad, clearly overlapping distal podomeres; dorsum pale yellow to white, without stripes (Fig. 19); Douglas, Jackson, and Josephine counties, Oregon prolata, new species Octoglena bivirgata Wood Figs. 2, 3, 6-11 Octoglena bivirgata Wood, 1864:186; 1865:229-230, figs. 58-59. Bollman, 1893:117, 187. Euzonium crucis Chamberlin, 1950:1-2. Chamberlin and Hoffman, 1958:187. Buckett, 1964:29. NEW SYNONYMY. Hypozonium arnaudi Chamberlin, 1954:233. Buckett, 1964:29. NEW SYNONYMY. Type specimens — Male neotype (AMNH) collected by V. Roth, 19 July 1962, 2 mi (3.2 km) N Ft. Ross, along California highway 1, ca. 15.3 mi (24.5 km) N Bodega Bay, Sonoma County, California. Citing a letter from R. L. Hoffman, Loomis (1971:155) reported that the type, which lacked the anterior end, was at the Academy of Natural Sciences, Philadelphia (ANSP), but extensive searches in June 1995 by the entomology collection manager failed to locate it. The specimen therefore appears to be lost, and neotype designation is needed to stabilize the name, necessitating library research on the collector, the entomologist and medical doctor, John Lawrence LeConte, and the origin of the confusion between Georgia and California. In the original accounts, Wood (1864, 1865) indicated uncertainty about the type locality by stating that he believed it to be the mountains of Georgia. That the site was probably in California is shown by unquestionable California material, collected by LeConte, that incorrectly carries a Georgia label. For example, Chamberlin (1947) reported several ANSP males of the California genus Xystocheir Cook (Polydesmida: Xystodesmidae) that were taken by Leconte and erroneously labeled “Georgia.” Wood (1867) explained that LeConte collected in both Georgia and California, that he presented all of his material to the ANSP, but that there was only one bottle, labeled Georgia. Thus, Wood originally concluded that the California specimens were missing, Hirudisomatid Millipeds 113 Figs. 6-11. O. bivirgata, neotype. 6, head and collum, anterior view, setation and pigmentation omitted. 7, profile of midbody segment, caudal view. 8, midbody segments, lateral view of left side, pigmentation omitted. 9, left anterior gonopod and sternum, anterior view. 10, right anterior gonopod, caudal view. 11, left posterior gonopod, caudal view. Upper scale line = 0.50 mm for figs 6-8; lower line = 0.25 mm for figs 9-11. 114 Rowland M. Shelley but he later realized that they were probably combined with the Georgia material. According to Horn (1884a, b), LeConte visited California in 1850, staying in San Francisco, San Jose, and San Diego, after which he traveled to central Arizona. He then returned to California and New York, moving to Philadelphia in 1852. Consequently, the only places where LeConte is likely to have collected in California were around these cities, and San Diego is eliminated as a potential site because of its aridity. Polyzonioids do not occur nearly this far south, the southernmost hirudisomatid record being from central Santa Cruz County, some 330 mi (528 km) to the north. The original material therefore probably came from the vicinities of San Francisco and/or San Jose, which are within the ranges of the family and the striped species. Diagnosis — Dorsum with dark medial and two lighter lateral stripes; caudal metatergal margins strongly upturned, caudolateral corners of midbody metatergites slightly but distinctly extended and rounded; telson broad, comprising entire breadth of caudal extremity; sternal lobes of anterior gonopods relatively short, moderately segre- gated; coxal lobe of latter moderately long and broad, apically rounded, leaning mediad and overlapping corners of 4th and 5th podomeres; dorsal branch of ultimate podomere short and broad, apically blunt, curved slightly ventrad and directed submediad; ventral lobe of ulti- mate podomere distinct, clearly overhanging dorsal branch; ultimate podomere of posterior gonopod apically fimbriate (Figs. 2, 6-11). Variation — Males with seemingly mature gonopods have from 21 to 55 segments and vary in length from 2.8 to 19.1 mm; female segment numbers vary from 12 to 58, while lengths range from 1.6 to 19.7 mm. These data are presented in table 1, with localities arranged in a general north to south sequence. Individuals are slightly longer toward the south of the range. Ecology — Habitat notations on vial labels include “under wet, rotting branches,” “redwood litter,” “under a piece of redwood log,” “on a redwood stump,” and “under rock on damp, muddy floor” in a cave. Distribution — The southernmost western species, O. bivirgata traverses San Francisco Bay and extends from coastal Curry County, Oregon, to central Santa Cruz County, California; although primarily hugging the coastline, the distribution extends eastward to the eastern slope of the Coast Range in western Colusa County (Fig. 28). Maximum dimensions are around 385 mi (616 km), north/south, and 67 mi (107 km), east/west. Specimens were examined as follows; the initials AKJ in this and the succeeding account denote samples collected by Hirudisomatid Millipeds 115 Table 1. Segment Numbers and Average Lengths (mm) of O. bivirgata (samples at one locality are combined; lengths averaged for individuals with same seg- ment number, n in parentheses). Males Females County Locality Segs. Lengths Segs. Lengths Humboldt, CA ■’R 2 mi E 55 19.1 58 19.7 Trinidad 46 12.2 51 15.3 Humboldt, CA 2.5 mi S Blue Lake 49 15.6 Humboldt, CA Areata 44 13.2 (2) 50 14.8 43 12.3 49 15.0 42 11.9 41 9.2 (2) 38 8.7 37 8.2 Humboldt, CA 2.75 mi NE 50 12.4 52 16.9 Orleans (2) 45 11.0 48 12.8 41 9.1 45 15.3 34 5.4 39 9.1 28 4.7 38 8.3 36 7.1 Humboldt, CA 11 mi W Willow Creek 41 9.1 Humboldt, CA 3.5 mi “up” Fickle Hill Rd. 41 15.1 Humboldt, CA Redcrest (2) 43 11.9 34 8.7 (2) 39 11.6 33 8.2 33 7.3 (2) 24 4.4 30 5.3 (2) 23 4.3 28 5.0 (3) 18 2.6 17 2.5 12 1.7 Humboldt, CA 4.5 mi N 41 10.3 40 11.6 Pepperwood 39 9.8 38 11.2 24 4.0 37 9.1 36 8.1 34 7.0 Humboldt, CA Weitchpec (3) 40 9.7 40 9.0 39 9.0 39 9.5 116 Rowland M. Shelley Table 1. Continued. Males Females County Locality Segs. Lengths Segs. Lengths 27 4.6 (2) 27 4.7 (2) 25 4.3 (3) 26 4.5 (2) 25 4.0 24 3.7 (2) 23 3.7 17 2.3 12 1.6 Mendocino, CA Elk 37 10.2 Mendocino, CA 12.7 mi SSW (2) 35 9.9 42 14.0 Leggett (2) 34 9.3 39 12.0 33 9.5 (2) 37 11.9 32 8.1 36 12.9 30 7.7 (4) 35 11.0 29 6.5 (2) 34 9.9 27 6.5 (4) 33 9.2 (4) 32 8.4 (6) 31 8.2 29 6.9 28 6.0 27 6.8 Lake, CA Kelsey Cr. 32 6.5 33 7.9 Colusa, CA 2 mi W jet. Hys. 20 & 16 34 6.0 Sonoma, CA El Verano 36 10.2 38 11.1 Sonoma, CA 2 mi N Ft. Ross 32 6.3 Napa, CA Clay Cave 44 13.7 Marin, CA Bolinas Jet. 36 7.8 34 8.9 30 6.0 33 6.8 32 6.3 30 5.6 Marin, CA Inverness Ridge 27 6.8 30 8.0 29 5.7 Hirudisomatid Millipeds 117 Table 1. Continued. County Locality Males Segs. Lengths Females Segs. Lengths Marin, CA 1 mi S 35 9.6 Inverness Marin, CA 2 mi SSW 29 5.8 Inverness Marin, CA Lagunitas Cyn. (2) 37 11.4 Marin, CA Sam P. Taylor 32 7.7 (3) 34 9.5 St. Pk. (3) 31 8.0 33 9.5 29 7.1 32 9.3 Marin, CA Mt. Tamalpais 40 9.7 San Mateo, CA 6.5 km ESE 30 6.1 Half Moon Bay Santa Clara, CA Stevens Cr. 29 6.0 35 7.7 (2) 28 6.3 34 7.1 (2) 33 8.4 32 7.3 31 6.5 29 5.8 28 6.1 26 6.4 (2) 25 5.0 24 4.7 Santa Cruz, CA 9.5 mi NE (2) 31 6.6 32 7.7 Soquel 29 6.3 30 5.7 26 4.5 29 5.8 25 4.6 19 3.2 21 2.8 A. K. Johnson. OREGON: Curry Co., Gold Beach, M, 19 August 1961, W. Suter (FSCA). CALIFORNIA: Colusa Co., 2 mi (3.2 km) W jet. hwys. 20 & 66, F, juv., 30 November 1960, T. Fenner (FSCA). Del Norte Co., 2 mi (3.2 km) N, 7 mi (11.2 km) E Gasquet, Cedar Rustic Cpgd., 118 Rowland M. Shelley 3M, 22 December 1977, AKJ (VMNH); 7 mi (11.2 km) ENE Gasquet, Patrick Cr. Cpgd., 3M, 5F, 22 December 1977, AKJ (VMNH); 5 mi (8 km) E Gasquet, Grassy Flat Cpgd. along CA hwy. 199, M, 25 March 1976, AKJ (VMNH); and Gasquet, M, 1 November 1977, and M, 2F, 22 December 1977, AKJ (VMNH). Humboldt Co., Areata, Hum- boldt St. Univ., Fern L. vie., 2F, 25 January 1976, AKJ (VMNH); Redcrest,7M, 3F, 5 juvs., 25 November 1977, AKJ (VMNH); Tish Tang Rec. Area, along CA hwy. 299 N Willow Creek (town), 4M, F, 21 February 1976, and 2F, 20 December 1979, AKJ (VMNH); 11 mi (17.6 km) W Willow Creek, M, 2F, 28 March 1976, AKJ (VMNH); Cheatham Redwood Grove, along CA hwy. 36 at Van Duzen R., F, 19 December 1977, AKJ (VMNH); 2.75 mi (4.4 km) NNE Orleans, MM, FF, juvs., 21 December 1976, AKJ (VMNH); 4.5 mi (7.2 km) N Pepperwood, along CA hwy. 36, 5M, 5F, 2 juvs., 12 November 1977, AKJ (VMNH); 3.5 mi (5.6 km) N, 1 mi (1.6 km) E Pepperwood, along CA hwy. 36, F, 12 November 1977, AKJ (VMNH); Big Lagoon, M, 30 November 1974, AKJ (VMNH); 1.5 mi (2.4 km) S Scotia, along US hwy. 101, M, juv., 3 January 1977, AKJ (VMNH); 3.5 mi (5.6 km) “up” Fickle Hill Rd., F, 29 November 1975, W. B. Dean (VMNH); 1.5 mi (2.4 km) NE, 1.75 mi (2.8 km) SE Orick, along Redwood Cr., 2M, F, 27 November 1976, and F, juv., 26 March 1977, AKJ (VMNH); 5 mi (8 km) E, 2.5 mi (4 km) S Blue Lake, along Canon Cr., M, F, 5 February 1976, AKJ (VMNH); 2 mi (3.2 km) E Trinidad, 2M, 9 March 1976, G. Panting (VMNH); Patrick’s Pt. St. Pk., M, 2F, 7 December 1974, AKJ (VMNH); Redwood Nat. Pk., Bald Hills Rd., M, 27 March 1977, AKJ (VMNH); and Weitchpec, along Klamath R., MM, FF, 30 November 1974, AKJ (VMNH). Lake Co., Cobb Mtn., along Kelsey Cr., 2F, 13 March 1962, J. S. Buckett (FSCA). Marin Co., Bolinas Jet., 2M, 4F, 29 September 1965, W. Ivie (AMNH); Inverness Ridge, 2M, 2F, 9 October 1963, J. S. Buckett (FSCA); 1 mi (1.6 km) S Inverness, F, 21 March 1959, C. W. O’Brien (FSCA); 2 mi (3.2 km) SSW Inverness, F, 30 December 1961, J. S. Buckett (FSCA); Lagunitas Cyn., 2F, 7April 1946, H. P. Chandler (FSCA); Sam P. Taylor St. Pk., M, 28 September 1959, J. Wagner (FSCA) and 2M, 5F, 19 September 1962, N. B. Causey (FSCA); and Mt. Tamalpais St. Pk., F, 1 April 1958, Lange (VMNH). Mendocino Co., 12.7 mi (20.3 km) SW Leggett, along CA hwy. 208, MM, FF, 21 March 1976, G. & J. Parkinson, AKJ (VMNH); 4 mi (6.4 km) S Rockport, F, 19 August 1959, V. Roth (NMNH); 1 mi (1.6 km) SE Caspar, M, 13 September 1961, W. J. Gertsch, W. Ivie (NMNH); along Little R., 3 August 1957, F, juv., 3 August 1957, J. R. Heifer, G. A. Marsh (NMNH); and Elk, F, 16 February 1967, V. Roth (AMNH). Hirudisomatid Millipeds 119 Napa Co., Clay (Kiel) Cave, 3 mi (4.8 km) N St. Helena, F, 26 November 1959, R. Graham (FSCA). San Mateo Co., 6.5 mi (10.4 km) ESE Half Moon Bay, along Purisima Cr., juv., 25 December 1974, AKJ (VMNH); 6 mi (9.6 km) SE Half Moon Bay, 2M, 3F, 1 June 1957, juv., 21 July 1957, R. O. Schuster (NMNH); and Woodside, M, 2F, 18 January 1947, P. H. Arnaud (NMNH). Santa Clara Co., along Stevens Cr., M, 14F, 2 June 1957, R. O. Schuster (NMNH, VMNH). Santa Cruz Co., Felton, F, 6 February 1949, P. H. Arnaud (NMNH); and 9.5 mi (15.2 km) NE Soquel, 5M, 4F, 31 December 1956, S. M. Fidel (VMNH). Sonoma Co., 2 mi (3.2 km) N Ft. Ross, M, 19 July 1962, V. Roth (AMNH) NEOTYPE LOCALITY; and El Verayo, along Fowler Cr., exact location unknown, 2F, 29 November 1975, J. DeMartini (VMNH). Remarks — The holotype of E. crucis is lost, and its sex is unknown, as Chamberlin (1950) merely states that there was “one specimen.” The female holotype of H. arnaudi, collected at the same time and place, and by the same collector, is available and confirms Hoffman’s (1980) and Shelley’s (1988) beliefs that both names are synonyms of O. bivirgata. It displays the diagnostic striped color pattern (Fig. 2), and the gonopods of proximate males agree with those of males from throughout the range of the striped species. Unlike most diplopod pigmentations, the stripes of O. bivirgata persist and are usually visible after 30-40 years in alcohol. Occasional specimens are pallid or nearly so, displaying only a trace of the stripes, but most individuals of O. bivirgata can be distinguished from sympatric polyzonioids by this distinctive pattern. Octoglena anura (Cook), new combination Figs. 12-15 Hypozonium anurum Cook, 1904:63, pi. V, figs. la-d. Chamberlin, 1911:262. Cook and Loomis, 1928:17. Chamberlin and Hoffman, 1958:187. Kevan, 1983:2962. Scudder, 1994:22. Type specimens — The holotype, from Seattle, Washington, was type no. 791 at the NMNH (Cook 1904, Chamberlin and Hoffman 1958), but it is now lost. Male neotype and female paraneotype (FSCA) collected by W. Suter, 15 August 1961, in Saltwater State Park, ca. 18 mi (28.8 km) S Seattle, King County, Washington. Diagnosis — Dorsum without stripes, color pale yellow to white; caudal metatergal margins indistinctly upturned, caudolateral corners of midbody metatergites slightly but distinctly extended and rounded; telson broad, comprising entire breadth of caudal extremity; sternal lobes of anterior gonopods relatively long, widely segregated; coxal 120 Rowland M. Shelley / Figs. 12-15. O. anura, neotype. 12, telson and caudal tergites, dorsal view. 13, right anterior gonopod and sternum, anterior view. 14, left anterior gonopod, caudal view. 15, ultimate podomeres of posterior gonopods, caudal view. Scaleline for fig. 12 = 1.00 mm; line for other figs. = 0.25 mm for each. lobe of latter short, broad, and upright, apically linear, not overlapping distal podomeres; dorsal branch of ultimate podomere short and broad, apically blunt, curved slightly ventrad and directed submediad; ventral lobe of ultimate podomere distinct, clearly overhanging dorsal branch; ultimate podomere of posterior gonopod narrowing apically, lightly hirsute (Figs. 12-15). Variation — Males with seemingly mature gonopods have from 16 to 40 segments and vary in length from 3.4 to 13.3 mm, the latter, a specimen with 38 segments, being slightly longer than that with Hirudisomatid Millipeds 121 Table 2. Segment Numbers and Average Lengths (mm) of O. anura (samples at one locality are combined; lengths averaged for individuals with same segment number, n in parentheses). Males Females County Locality Segs. Lengths Segs. Lengths BC 1.9 mi SE 28 8.3 30 9.4 Hope (2) 25 6.1 26 6.9 24 5.7 25 6.1 23 4.9 (2) 24 5.8 22 5.6 (3) 23 5.9 19 4.1 22 5.0 19 3.7 (2) 18 3.9 (2) 14 2.6 BC Manning 26 6.4 26 7.1 Prov. Pk. 22 4.4 25 5.5 Clallam, WA Olympic 27 6.5 27 5.0 Hot Spgs. 25 5.1 24 4.8 23 4.0 Jefferson, WA 5.5 mi S 34 9.1 27 7.1 Brinnon 31 8.0 (2) 26 7.0 29 8.8 (5) 27 6.8 26 6.0 25 6.0 23 5.5 Mason, WA Kamilche Pt. 40 12.9 21 4.3 25 6.0 22 5.0 King, WA Federation For. 32 9.3 St. Pk. King, WA Saltwater St. Pk. 25 5.7 31 8.6 Thurston, WA Puget 28 7.3 30 7.0 25 6.1 29 7.1 25 6.4 Pierce, WA Mt. Rainier 22 5.0 29 8.1 Nat. Pk. 122 Rowland M. Shelley Table 2. Continued. County Locality Males Segs. Lengths Females Segs. Lengths Cowlitz, WA 1 mi E Touttle 31 9.4 Multnomah, OR Portland (2) 28 7.1 31 6.7 27 6.7 25 5.6 24 5.7 Washington, OR 5 mi SW 29 8.1 28 6.9 Tualatin 28 6.5 Clackamas, OR 10 mi E 30 8.0 29 9.0 Zigzag 28 7.5 (2) 28 8.1 (2) 26 6.7 (2) 26 6.7 25 6.2 (2) 25 7.0 (4) 24 5.9 24 5.6 (2) 23 5.2 (6) 23 5.4 22 5.9 21 4.9 (2) 20 4.3 20 3.8 19 4.0 19 4.4 (2) 18 3.5 (2) 16 2.9 (2) 16 2.9 14 2.4 Yamhill, OR 5 mi E 34 9.3 Yamhill (2) 28 7.1 Lincoln, OR 0.6 mi NW 29 9.3 28 8.2 Elk City 28 7.6 27 8.6 22 5.3 Benton, OR 6 mi N 28 6.8 (2) 28 6.9 Corvallis 27 6.9 27 5.8 (2) 24 5.8 26 6.0 (4) 23 5.4 25 5.5 (2) 22 4.1 24 4.8 21 4.4 (3) 23 5.1 (6) 19 3.2 21 4.3 (4) 18 3.3 20 3.5 (2) 17 3.1 19 3.3 16 3.4 (4) 18 3.3 (5) 17 3.0 16 2.8 (4) 13 2.1 (3) 12 2.0 9 1.5 (5) 5 1.0 Hirudisomatid Millipeds 123 Table 2. Continued. County Locality Males Segs. Lengths Females Segs. Lengths Lane, OR 11 mi NE 38 13.3 Blue R. Lane, OR 18.5 mi ESE 31 8.4 Springfield 30 9.8 (4) 28 7.5 25 5.9 Douglas, OR Comstock 34 10.0 32 9.6 40 segments, which is 12.9 mm. Female segment numbers vary from 5 to 22, while lengths range from 1.0 to 9.6 mm. These data are presented in table 2, with localities arranged in a general north to south sequence. Lengths appear roughly comparable throughout the range, and no geographic trends are evident. Ecology — The neotype was recovered from maple litter; habitat notations on labels in other vials include “under moss on forest floor,” “berlese conifer duff,” “willow, maple, fir duff, moss,” “in littler under firs, cedars,” “deciduous litter, grass,” “under log,” “bark, moss, debris,” “ash, oak, conifer duff,” “oak litter, rotted wood,” and “berlese beach grass debris, dried seaweed, spruce duff.” Distribution — The northernmost species, O. anura extends along the Pacific Coast from the southwestern corner of the British Columbia mainland to northern Douglas County, Oregon, and ranges inland to the western slope of the Cascade Mountains (Fig. 28); dimensions are approximately 385 mi (616 km), north/south, and 110 mi (176 km), east/west, the former being about the same distance as the north/ south dimension of O. bivirgata. Specimens were examined as follows; to consolidate records, I repeat the five from British Columbia cited by Shelley (1990). The initials EMB in this and the following account denote samples collected by E. M. Benedict, primarily from berlese extracts. CANADA: BRITISH COLUMBIA: Burnaby Mtn., Simon Fraser Univ., 2M, 2F, 1972, R. G. Holmberg (CMN); Burquitlam, F, 10 March 1940, W. Dale (NMNH); Steelhead, F, 2 June 1933, H. B. Leech (NMNH); Agassiz, F, 7 March 1931, H. B. Leech (NMNH); 1.9 mi (3 km) SE Hope, Silver Skagit Rd., 8M, 13F, 4 juvs., 30 June 1988, S. & J. Peck (NCSM); and Manning Prov. Pk., West Gate, 2M, 2F, 124 Rowland M. Shelley 1 July 1988, S. & J. Peck (NCSM). USA: WASHINGTON: Clallam Co., Olympic Hot Spgs., 3M, 2F, 15-16 August 1961, W. Suter (FSCA). Cowlitz Co., 1 mi (1.6 km) E Touttle, M, F, 4 juvs., 16 April 1960, B. D. Ainscough (RBCM). Jefferson Co., 5.5 mi (8.8 km) S Brinnon, along US hwy. 101, 9M, 4F, 23 September 1978, AKJ (VMNH). King Co., Federation Forest St. Pk., F, 28 August 1990, R. M. Shelley (NCSM); and 18 mi (28.8 km) S Seattle, Saltwater St. Pk., M, F, 15 August 1961, W. Suter (FSCA) NEOTYPE LOCALITY. Mason Co., Kamilche Pt., 3M, F, 25 November 1967, EMB (WAS). Pierce Co., Mount Rainier Nat. Pk., Longmire Cpgd., M, F, 18 August 1961, W. Suter (FSCA). Thurston Co., Puget, 2M, 3F, 28 October 1967, EMB (WAS); and Millersylvania St. Pk., M, 28 October 1967, EMB (WAS). OREGON: Benton Co., 6 mi (9.6 km) N Corvallis, McDonald For., F, 23 October 1969, 17M, 14F, 12 juvs., 21 February 1971, and 2M, 4F, 20 juvs., 21 February 1973, L. Russell (VMNH); and Mary’s Peak, M, 30 October 1972, L. Russell (VMNH). Clackamas Co., 10 mi (16 km) E, 3.5 mi (5.6 km) S Zigzag, Still Cr. Cpgd., 19M, 23F, 14 September 1977, AKJ (VMNH). Clatsop Co., Ft. Stevens St. Pk., M, 22 November 1971, EMB (WAS). Columbia Co., Locoda Sta. nr. Clatskanie, M, 2F, juv., 31 March 1937, J. Schuh (NMNH); and Scappoose, M, 7 May 1937, J. C. Chamberlin (NMNH). Douglas Co., 6 mi (9.6 km) S Cottage Grove, 3M, 4F, 28 April 1937, J. C. Chamberlin (NMNH); Comstock, M, F, 7 January 1950, V. Roth (VMNH); 1 mi (1.6 km) SE Tiller, along Elk Cr. on OR hwy. 227, 6M, 3F, 6 November 1971, EMB (WAS); and 3 mi (4.8 km) SE Tiller, Umpqua R. Val., along OR hwy. 227, F, 6 November 1971, EMB (WAS). Lane Co., 18.5 mi (29.6 km) ESE Springfield, Dolly Varden Cpgd. in Willamette Nat. For., along USFS 18, ca. 10 mi (16 km) E Fall Creek (town), 7M, 4 March 1972, EMB (WAS); and 11 mi (17.6 km) NE Blue River, Andrews Exp. For., M, 18 October 1983, C. L. Parsons (VMNH). Lincoln Co., Saddlebag Mtn., ca. 14.5 mi (23.2 km) ENE Lincoln, 2F, 3 March 1960 & 6 January 1961, J. C. Dirks-Edmunds (FSCA); and 0.6 mi (1.0 km) NW Elk City, along Yaquina R., 2M, 3F, 20 December 1971, EMB (WAS). Multnomah Co., Portland, Lewis & Clark Col., 2M, 11 May 1957, R. Ennis, M, F, March 1961, R. Anderson, and 3M, 1 May 1968, A. Ashwanden (FSCA). Washington Co., 3 mi (4.8 km) SW Tualatin, 2M, F, 1 January 1972, EMB (WAS). Yamhill Co., 5 mi (8 km) E Yamhill, along OR hwy. 240, M, 2 October 1971, EMB (WAS). Literature Record: WASHINGTON: Kitsap Co., Bremerton (Chamberlin 1911). Hirudisomatid Millipeds 125 Remarks — Cook (1904) provided few clues as to this species’ identity, but his illustration of the broad telson indicates a hirudisomatid, and only one ordinal representative, a hirudisomatid, occurs around Seattle and Puget Sound. It must therefore carry Cook’s name. Octoglena prolata, new species Figs. 16-18 Type specimens — Male holotype and 3 male and 2 female paratypes (NMNH) collected by E. M. Benedict, 6 November 1971, along Oregon highway 227 in Canyonville, Douglas County, Oregon; 4 male and Figs. 16-18. O. prolata, holotype. 16, left anterior gonopod and sternum, anterior view. 17, the same, caudal view. 18, distal podomeres of right posterior gonopod, caudal view. Scale line = 0.50 mm for all figs. 126 Rowland M. Shelley one female paratypes (VMNH) taken by same collector on same date in Canyonville County Park, 2 mi (3.2 km) E Canyonville. Diagnosis — Dorsum without stripes, color pale yellow to white; caudal metatergal margins indistinctly upturned, caudolateral corners of midbody metatergites slightly but distinctly extended and rounded; telson broad, comprising entire breadth of caudal extremity; sternal lobes of anterior gonopods relatively long, widely segregated; coxal lobe of latter long and narrow, apically rounded, leaning laterad and overlapping 4th and 5th podomeres; dorsal branch of ultimate podomere short and broad, apically blunt, curved slightly ventrad and directed submediad; ventral lobe of ultimate podomere distinct, clearly overhanging dorsal branch; ultimate podomere of posterior gonopod apically fimbriate (Figs. 16-18). Table 3. Segment Numbers and Average Lengths (mm) of O. prolata (no. indi- viduals averaged in parentheses). County Locality Males Segs. Lengths Females Segs. Lengths Douglas, OR Canyonville 34 6.8 (2) 34 7.3 30 5.6 31 6.8 (2) 24 3.8 Douglas, OR 2 mi E 38 9.2 32 7.2 Canyonville 36 8.4 31 6.0 30 5.4 Josephine, OR SW Wolf 45 9.2 Creek Josephine, OR 1 mi S 30 5.6 O’Brien Josephine, OR 2.5 mi S 35 7.9 O’Brien Jackson, OR 6 mi S Ruch 32 6.6 37 8.5 29 6.9 31 6.2 27 5.7 26 6.3 25 ' 4.8 21 5.0 (2) 23 4.2 22 4.2 Hirudisomatid Millipeds 127 Variation — Males with seemingly mature gonopods have from 22 to 38 segments and vary in length from 4.2 to 9.2 mm. Female segment numbers vary from 21 to 37 and lengths, from 5.0 to 8.5 mm. These data are presented in table 3, with localities arranged in a general, north to south, sequence; no geographic trends are evident. Ecology — The types were retrieved from alder litter and moss, wood, and soil; another sample was discovered under “rotten madrone wood.” Distribution — A small subtriangular area in southwestern Oregon, slightly to the east of the northern range periphery of O. bivirgata (Fig. 28); dimensions are about 63 mi (100.8 km), north/south, and 35 mi (56 km), east/west. In addition to the types, specimens were examined as follows: OREGON: Jackson Co., 6 mi (9.6 km) S Ruch, 7M, 4F, 13 November 1971, EMB (WAS). Josephine Co., along Grave Cr., SW Wolf Creek (town), M, 30 May 1952, V. Roth (FSCA); 1 mi (1.6 km) S, 0.5 mi (0.8 km) W O’Brien, F, 18 December 1971, EMB (WAS); and 2.5 mi (4 km) S, 1 mi (1.6 km) W O’Brien, M, 18 December 1971, EMB (WAS). Octoglena sierra, new species Figs. 19-21 Type specimens — Male holotype and one male and one female paratypes (VMNH) collected by Smith and R. O. Schuster, 15 April 1958, 4 mi (6.4 km) W Newcastle, Placer County, California; other paratypes from this locality include 5 males and 3 females (VMNH) by same collectors, 12 March 1958; one male (NCSM) by same collectors, 10 March 1959; and 2 females (VMNH) by Lange, Smith, and R. O. Schuster, 21 March 1958; one female paratype (VMNH) by Smith and R. O. Schuster, 19 March 1959, 4 mi (6.4 km) N Newcastle. Diagnosis — Dorsum without stripes, color pale yellow to white; caudal metatergal margins strongly upturned, caudolateral corners of midbody metatergites slightly but distinctly extended and rounded; telson broad, comprising entire breadth of caudal extremity; sternal lobes of anterior gonopods relatively short, widely segregated; coxal lobe of latter long and relatively broad, leaning mediad, overlapping 4th-6th podomeres; dorsal branch of ultimate podomere long, narrow, and sinuate, apically acuminate, directed sublaterad; ventral lobe of ultimate podomere short and indistinct, only slightly overhanging dorsal branch; ultimate podomere of posterior gonopod apically fimbriate (Figs. 19-21). Variation — Males with seemingly mature gonopods have from 128 Rowland M. Shelley Figs. 19-21. O. sierra, holotype. 19, right anterior gonopod and sternum, anterior view. 20, left anterior gonopod, caudal view. 21, distal podomeres of right posterior gonopod, caudal view. Scale line = 0.25 mm for all figs. Table 4. Segment Numbers and Average Lengths (mm) of O. sierra (samples at one locality are combined; no. individuals averaged in parentheses) County Locality Males Females Segs. Lengths Segs. Lengths Placer, CA Placer, CA 4 mi W (2) 49 11.7 41 11.9 Newcastle 42 9.3 39 11.0 38 9.0 38 9.8 36 7.1 (3) 35 7.2 34 7.8 4 mi N 43 11.5 Newcastle Hirudisomatid Millipeds 129 34 to 49 segments and vary in length from 7.1 to 11.7 mm, the shortest individual having 36 segments and being 0.7 mm shorter than that with the least segments. Female segment numbers vary from 38 to 43 and lengths, from 9.8 to 11.9, the latter, of an individual with 41 segments, being 0.4 mm longer than the female with the most segments (table 4). Ecology — One paratype sample was encountered in “litter under oak.” Distribution — A localized species known only from the type and paratype localities in the foothills of the Sierra Nevada (Fig. 28), O. sierra is detached from the coastal representatives and occurs some 75 mi (120 km) east of the nearest site of O. bivirgata, in Colusa County. It is the easternmost western species and occupies an inter- mediate geographical position, albeit far to the western side of the generic range. By combining gonopodal attributes of O. gracilipes, the long, sinuate dorsal branch and the indistinct ventral lobe, with a trait of the coastal species, the coxal lobe, O. sierra links the ana- tomical extremes, which justifies congeneric status. Octoglena gracilipes (Loomis), new combination Figs. 22-27 Polyzonium gracilipes Loomis, 1971:157-159, figs. 18-23. Type specimens — Male holotype and one male and one female paratypes (NMNH) and one male and one female paratypes (FSCA) collected by H. R. Steeves, 17 June 1962, at Cloudland Canyon State Park, Dade County, Georgia. The NMNH sample also includes one male and three female polyzoniids. Diagnosis — Dorsum without stripes, color pale yellow to white; caudal metatergal margins moderately upturned, caudolateral corners of midbody metatergites not extended, contiguous with middorsal margins, blunt; telson narrow, comprising about half of breadth of caudal extrem- ity; sternal lobes of anterior gonopods relatively long, narrowly segregated; coxae of latter without lobes; dorsal branch of ultimate podomere long, narrow, and sinuate, apically acuminate, directed sublaterad; ventral lobe of ultimate podomere indistinct, not overhanging dorsal branch; ultimate podomere of posterior gonopod apically narrow and attenuated (Figs. 22-27). Variation — Measurable males with seemingly mature gonopods have from 19 to 33 segments and vary in length from 2.8 to 7.9 mm; female segment numbers vary from 13 to 35, while lengths vary from 1.7 to 8.5 mm. These data are presented in table 5 with localities arranged in a general east to west sequence. Lengths appear roughly 130 Rowland M. Shelley Figs. 22-27. O. gracilipes, holotype. 22, head and collum, anterior view, setation omitted. 23, midbody segments, lateral view of left side. 24, tel- son and caudal tergites, dorsal view. 25, right anterior gonopod and ster- num, anterior view. 26, the same, caudal view. 27, distal podomeres of left posterior gonopod, caudal view. Upper scale line = 0.50 mm for figs. 22- 24; lower line = 0.25 mm for figs. 25-27. comparable throughout the range, and no geographic trends are evident. Ecology — Habitat notations on vial labels include “virgin cove forest,” “under logs,” and “wooded hillside.” Specimens from Cleburne County, Alabama, and Greenwood County, South Carolina, were encountered in ravines. Distribution — The Piedmont Plateau, Blue Ridge, Ridge and Valley, and eastern Cumberland Plateau Physiographic Provinces of Tennessee, Hirudisomatid Millipeds 131 Table 5. Segment Numbers and Average Lengths (mm) of O. gracilipes (samples at one locality combined; lengths averaged for individuals with same segment number, n in parentheses). County Locality Males Segs. Lengths Females Segs. Lengths Chester, SC ■=) 13.6 mi 30 8.2 W Chester Greenwood, SC Ware Shoals 28 7.9 Polk, NC Saluda 25 4.8 Transylvania, NC SE L. Toxaway 33 7.9 Macon, NC Ellicott Rock 19 2.8 22 4.1 19 2.8 Macon, NC 3.6 mi E 20 3.5 19 2.6 Highlands (2) 18 2.5 16 2.0 15 2.2 13 1.9 Rabun, GA E of Satolah 28 7.7 Graham, NC 6 mi SE 31 7.6 32 8.6 Beech Gap 30 6.3 Graham, NC Kilmer Mem. For. 27 5.8 Sevier, TN 1.4 mi N 28 7.7 Gatlinburg Cumberland, TN Ozone 30 6.7 29 7.3 21 3.5 Dade, GA Cloudland Cyn. 31 7.3 St. Pk. 30 6.7 27 6.0 Jackson, AL Princeton 23 4.6 Cleburne, AL N of Heflin 26 4.4 25 5.1 132 Rowland M. Shelley Table 5. Continued. County Locality Males Segs. Lengths Females Segs. Lengths Clay, AL Talladega 28 6.4 Nat For. Franklin, AL The Dismals 35 8.5 20 2.9 15 1.7 North and South Carolina, Georgia, and Alabama (Fig. 29), some 1,897 mi (3,035 km) east of the most proximate western species, O. sierra. The area is oriented generally east to west, extends from westcentral South Carolina to southcentral Tennessee and northwestern Alabama, and covers approximately 220 mi (352 km), north/south, and 380 mi (608 km), east/west. Specimens were examined from the following localities: SOUTH CAROLINA: Chester Co., 13.6 mi (21.8 km) W Chester, Woods Ferry Rec. Area, Sumter Nat. For., along USFS rd. 574, 3 mi (4.8 km) W jet. SC hwy. 49, F, 4 August 1976, R. M. Shelley (NCSM). Greenwood Co., Ware Shoals, F, 29 May 1960, L. Hubricht (VMNH). Oconee Co., along SC hwy. 28, exact site unknown, F, 29 July 1960, collector unknown (AMNH). NORTH CAROLINA: Graham Co., Joyce Kilmer Mem. For., M, 20 May 1970, W. A. Shear (WAS): 6 mi (9.6 km) SE Beech Gap, 2M, F, May 1958, L. Hubricht (VMNH); and Stratton Gap, 2F, 27 May 1959, L. Hubricht (VMNH). Jackson Co., Whiteside Cove, F, 26 July 1958, R. L. Hoffman (VMNH). Macon Co., Highlands, Highlands Biol. Sta., M, 10 August 1955, P. J. Darlington (VMNH); 3.6 mi (5.8 km) E Highlands, Horse Cr. Clearcut, M, 6F, 16 June 1976, F. A. Coyle (NCSM); and 7.5 mi (12 km) SSE Highlands, along Bull Pen Rd. nr. Ellicott Rock, M, 2F, 6 July 1976, F. A. Coyle (NCSM). Polk Co., Saluda, 2F, 5 August 1910, collector unknown (FSCA). Transylvania Co., Thompson R. Gorge SE L. Toxaway, M, 5 September 1961, R. L. Hoffman (VMNH). GEORGIA: Dade Co., Cloudland Cyn. St. Pk., 3M, 2F, 17 June 1962, H. R. Steeves (FSCA, NMNH) and 3F, 16 May 1972, S. Peck (WAS) TYPE LOCALITY. Rabun Co., along Glade Mtn. Rd. E of Satolah, F, 6-8 September 1961, R. L. Hoffman (VMNH). Hirudisomatid Millipeds 133 TENNESSEE: Cumberland Co., Ozone, below Ozone Falls, 5F, 21 May 1961, L. Hubricht (VMNH). Sevier Co., 1.4 mi (2.2 km) N Gatlinburg, F, 17 May 1961, L. Hubricht (VMNH). ALABAMA: Clay Co., Talladega Nat. For., exact site unknown, F, 16 April 1960, H. R. Steeves (FSCA). Cleburne Co., ravine nr. Skyway N of Heflin, 2.4 mi (3.8 km) N Bankhead Fire Tower, 2F, 23 October 1960, L. Hubricht (VMNH). Franklin Co., “The Dismals,” ca. 13 mi (20.8 km) SSW Russellville, M, F, 18 July 1959, 2F, 28 May 1960, and 2M, F, 17 June 1961, H. R. Steeves (FSCA). Jackson Co., Princeton, F, 29 October 1966, H. R. Steeves (FSCA). Winston Co., nr. Natural Bridge Cave, ca. 9 mi (14.4 km) S Haleyville, 2M, F, collector unknown (FSCA). Fig. 28. Distributions of the west-Nearctic Hirudisomatidae. Dots, O. bivirgata\ triangles, O. anura; diamonds, O. prolata\ squares, O. sierra. The open triangle denotes the literature record of O. anura from Bremerton, Washington (Chamberlin 1911). Fig. 29. Distributions of O. gracilipes and the Hirudisomatidae in eastern North America. 134 Rowland M. Shelley Although O. gracilipes was officially named and described by Loomis (1971), its existence was first mentioned by Hoffman (1969), who called it an undescribed and possibly relictual representative of the Hirudisomatidae in the high mountains of North Carolina, Georgia, and Tennessee. Mexiconium, new genus Type species — Mexiconium absidatum, new species. Diagnosis. Dorsum smooth and glossy, polished, with broad, dark middorsal stripe arising on collum and terminating on penultimate segment; body narrow (W/L ratio 16.7%) and vaulted in profile, sides not extending laterad; metaterga with caudal margins slightly elevated but not upturned; collum moderately broad, over-hanging epicranium and part of one pair of ocelli; telson broad, comprising entire breadth of caudal extremity; sternum of anterior gonopods with strong, sparsely hirsute lobes, widely separated from each other; anterior gonopod curving strongly anteriad distad, ultimate podomere with lightly hirsute lobe ventral to and overhanging glabrous dorsal branch, latter long, narrow, and acuminate, extending nearly directly laterad; corners of 4th and 5th podomeres slightly extended on caudal side, latter expanding distad and subuncinate; coxa with broad, glabrous lobe on anterior side, apically rounded; posterior gonopod with ultimate podomere simple and acicular, apically trifurcate. Species — One is known; others may exist in remote pockets in the Sierra Madre Oriental. Distribution — Known only from the type locality of the one component species in Vera Cruz, Mexico. Fig. 30. Relationships in Octoglena. Hirudisomatid Millipeds 135 Mexiconium absidatum, new species Figs. 4, 31-38 Type specimens — Male holotype and two male paratypes (VMNH) collected by R. E. Leech, 25 August 1967, 13.3 mi (21.8 km) S La Vigas [ca. 18 mi (28.8 km) WSW Jalapa], on the north side of Cofre de Perote, Tembladera, Vera Cruz, Mexico. Diagnosis — With the characters of the genus (Figs. 31-38). Variation — The holotype has 27 segments and is 6.7 mm long; the paratypes have 29 and 35 segments and measure 7.7 and 10.8 mm, respectively. Ecology — According to the vial label, the types were collected between 11,500 and 13,500 ft., an extremely high elevation for a North American milliped. The habitat is not indicated. Distribution — Known only from the type locality, in the Sierra Madre Oriental in the interior of Vera Cruz (Fig. 1), some 440 mi (704 km) south of the Rio Grande and the United States border and around 1,200 and 2,117 mi (1,920 and 3,387 km) from the most proximate sites of O. gracilipes and O. bivirgata, respectively. Remarks — This species, the first record of the family from Mexico, is isolated from an ancient dispersal of the Hirudisomatidae across North America that extended southward for an unknown distance into Mexico. The dorsal branch of its ultimate anterior gonopod podomere is similar to those of O. gracilipes/ sierra, instead of the Pacific Coastal species, and this is further evidence that the Hirudisomatidae dispersed from east to west across North America, as is its occurrence on the eastern side of Mexico rather than the west (see below). The specific name refers to the vaulted body form. Discussion The two great allopatries in the Hirudisomatidae merit elaboration because they exemplify broader patterns among Nearctic Diplopoda. The family’s occurrence in Mexico is evidence of southward expansion of ancestral stock, a dispersal also demonstrated by the Glomeridae, Spirobolidae, Parajulidae, Nearctodesmidae, and Xystodesmidae. The Mexican xystodesmid fauna exhibits Appalachian affinities and also immigrated from the east-Nearctic (Hoffman 1969), but the austral representatives of the other families have west-Nearctic relationships and occur more in western Mexico, aside from cave inhabiting species of Glomeroides Chamberlin (Glomeridae) in Nuevo Leon, Tamaulipas, and Vera Cruz. Like the Hirudisomatidae, the species of the Spirobolidae, Parajulidae, and Nearctodesmidae show further evidence of Nearctic ancestry in their occurrences at high elevations, in “Nearctic” environments, 136 Rowland M. Shelley C/3 03 3 Cn to 03 • F— < c 3 c CO rn ON MO’ co co t-H mo ON r\ CM CM v n 00 • ^ • »—h • ^ • ^ £ £ £ £ o o £ CM o v-H o r— s 00 M0 t-H o U OX 3 3 3-1 Po < U 3 Q. 3 3 o r U N 3 3-H u 3 3 3 00 a 3 03 X 3 H , r 6 6° 3 >X» O _C 3 0 : — i 3 ^ O x 03 o 03 C/3 ”c3 cd £ u u 0) _r ^ 8 3 3 3 « -c o 03 > S >3 2 X 03 Q o O • X 03 3 'O J < ^ U o > r 3 O 03 (J z Z CQ O M s u 03 c3 3 is 3 3 3 O U U <30 2 0) o 2 03 3 TO • fH £ C/3 03 2 3 2 5 2 X 2 X 2 ~2 • ^ c>, c5 O 2 C/3 E Ergodesmus 1 sp. 1 sp. Lewis & Clark, Pike, IL MT 1,150 mi 1,840 km W— >E Chonaphini 5 gen. 11 spp. 1 gen. 1 sp. Missoula, MT Rice, MN 1,060 mi 1,696 km W— >E Scytonotus 6 spp. 3 spp.* Sublette, WY Shawnee, KS 984 mi 1,574 km W-^E *One eastern species is widespread; the others are Appalachian endemics and reflect a secondary center of evolution in the southern Blue Ridge Province (Shelley 1993). The east/west allopatry in the United States is a consistent pattern among Nearctic Diplopoda, not only in the disjunction, but also in the greater western diversity. Scytonotus, Orinisobates, Brachycybe, and Octoglena have more western than eastern species, as does the transcontinental xystodesmid tribe Chonaphini, with five western genera and eleven species, versus one of each in the east (Table 7) (Gardner 1975, Enghoff 1985, Shelley 1993, 1994b). Ergodesmus is an exception, but the western species occupies a much larger area and demonstrates more variation (intraspecific diversity) than the eastern species, which comprises small populations and is restricted to caves in Illinois (Shelley 1994a). Western origins have been postulated for Scytonotus and the Chonaphini (Shelley 1993, 1994b), and Enghoff (1985) concluded that Orinisobates probably arose in the eastern Palearctic and invaded the Nearctic via the Bering Bridge, so it too probably spread from west to east across North America. Consequently, the east-Nearctic components of these taxa appear to represent the results of range expansions, rather 140 Rowland M. Shelley than remnants that remained in the original source area. Octoglena conforms to this diversity pattern, but the eastern component is sister to the western species, implying dispersal from east to west. The overall distribution of the family, with the greatest diversity in Europe, supports this scenario and suggests a Laurasian origin for the New World fauna. Futhermore, the continuous, parapatric ranges and the absence of extinc- tions along the Pacific Coast contrast so strongly with the allopatries and lacunae elsewhere in the New World that they probably reflect recent evolution and the end products of range expansion. Octoglena therefore represents the northern part of an Ancient Holarctic faunal assemblage that probably spread from east to west across the United States, and southward into Mexico, in one or possibly two dispersals. There has been considerable extinction, as evidenced by the extensive lacunae; that between O. sierra and O. gracilipes is greater than those in other disjunct taxa in the United States (Table 7), and coupled with the unique somatic features of O. gracilipes , implies lengthy isolation of the eastern and western hirudisomatid faunas. Hoffman (1969) cited three distribution patterns that have impacted Appalachian Diplopoda — Ancient Holarctic, Tertiary Nearctic Endemi- city, and Late Coenozoic Austral Immigration. These also affected lowland areas, and “West-Nearctic Immigration”, from beyond the Continental Divide, seems equally important in understanding the origins of the fauna east of the Plains. Causey (1974) postulated a west- Nearctic origin for the Parajulidae, with dispersal centers in central California and around Puget Sound that gave rise to the east-Nearctic and Mexican/Guatemalan faunas. This hypothesis is consistent with the conclusion of Shelley (1994c) that the exclusively western julidan superfamily Paeromopodoidea probably arose from a parajuloid-like ancestor along what is now the border between Oregon and California. This is a key area in the evolution of west-Nearctic diplopods (Shelley 1994c), being the apparent primary centers of evolution of the Paeromopodoidea and Scytonotus, and harboring a peripheral relict species of Chonaphe Cook (Shelley 1993, 19946, c). The area also seems to be a secondary center of evolution within Octoglena , thereby accounting for the greater diversity, parapatry, and abundance of the Pacific Coastal components. ACKNOWLEDGEMENTS — I thank Jonathan Coddington (NMNH) for loan of the holotype and paratypes, and G. B. Edwards (FSCA) for additional paratypes, of Polyzonium gracilipes; they also loaned non-typical specimens from these holdings. The following curators loaned material from the indicated collections: N. I. Platnick (AMNH), Hirudisomatid Millipeds 141 P. F. Frank (CMN), R. A. Cannings (RBCM), and R. L. Hoffman (VMNH); my colleague W. A. Shear provided specimens in his private holdings. Figures 2, 31-33 were prepared by R. G. Kuhler, NCSM scientific illustrator, who assisted with the other illustrations. I thank V. F. Lee & B. W. Rogers, for localiting obscure California localities; S. Bauer, for segment counts and measurements; C. Wood, for word processing; B. Randall-Schadel, for use of a compound microscope; and A. R. Hardy, for information about John Lawrence LeConte. LITERATURE CITED Bollman, C. H. 1893. The Myriapoda of North America. United States National Museum Bulletin Number 46:1-210. Buckett, J. S. 1964. Annotated list of the Diplopoda of California. Sim- mons Publishing Company, Davis, California. Causey, N. B. 1974. The phylogeny of the family Paraiulidae (Paraiuloidea: Blaniulidea: Iulida: Diplopoda). Symposium of the Zoological Soci- ety of London No. 32:23-39. Chamberlin, R. V. 1911. Notes on myriopods from Alaska and Washing- ton. Canadian Entomologist, 43:260-264. Chamberlin, R. V. 1947. Some records and descriptions of diplopods chiefly in the collection of the Academy. Proceedings of the Academy of Natural Sciences, Philadelphia, 99:21-58. Chamberlin, R. V. 1950. Three new genera and eight new species of western millipeds. Chicago Academy of Sciences Natural History Miscellanea Number 68:1-6. Chamberlin, R. V. 1954. Six new California millipeds. Proceedings of the Biological Society of Washington, 67:231-234. Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. United States National Museum Bulletin Number 212:1-236. Cook, O. F. 1904. Myriapoda of northwestern North America. In: Harriman Alaska Expedition, 8(Insects, pt. l):47-82. Cook O. F., and H. F. Loomis. 1928. Millipeds of the order Colobognatha, with descriptions of six new genera and type species, from Arizona and California. Proceedings of the United States National Museum, 72:1-26. Enghoff, H. 1985. The millipede family Nemasomatidae. With the de- scription of a new genus, and a revision of Orinisobates (Diplopoda: Julida). Entomologica Scandinavica, 16:27-67. Enghoff, H., and S. I. Golovatch. 1995. A revision of the Siphonocryptidae (Diplopoda, Polyzoniida). Zoologica Scripta, 24:29-41. Enghoff, H., W. Dohle, and J. G. Blower. . 1993. Anamorphosis in milli- pedes (Diplopoda) - the present state of knowledge with some devel- opmental and phylogenetic considerations. Zoological Journal of the Linnean Society, 109:103-234. 142 Rowland M. Shelley Gardner, M. R. 1975. Revision of the millipede family Andrognathidae in the Nearctic region. Memoirs of the Pacific Coast Entomological Society, 5:1-61. Hoffman, R. L. 1969. The origin and affinities of the southern Appala- chian diplopod fauna. Pages 221-246. In The Distributional History of the Biota of the Southern Appalachians, Part I: Invertebrates (P. C. Holt, editor). Research Division Monograph 1, Virginia Polytech- nic Institute, Blacksburg. Hoffman, R. L. 1970. Random studies on Rhysodesmus. I. Notes and re- descriptions of miscellaneous species. The Radford Review, 24:143- 162. Hoffman, R. L. 1977. On the status of Siphonotus brasiliensis Brandt, and of the diplopod family Siphonotidae (Polyzoniida: Diplopoda). Deutsche Entomologische Zeitschrift, NF, 24:425-431. Hoffman, R. L. 1980 (“1979”). Classification of the Diplopoda. Museum d’Histoire Naturelle, Geneva, Switzerland, 237 pp. Hoffman, R. L. 1990. Diplopoda. pp. 835-860, In: Dindal, D. L., ed., Soil Biology Guide. Wiley Interscience, New York. Hoffman, R. L., and W. A. Shear. 1969. Nomenclatorial alterations in the milliped genus Boraria (Polydesmida: Xystodesmidae). The Radford Review, 23:93-101. Horn, G. H. 1884a. John Lawrence LeConte. Daedalus, 19:511-516. Horn, G. H. 18846. Memoir of John L. LeConte, M. D. Proceedings of the American Philosophical Society, 21:294-299. Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monografieen van de Nederlandse Entomologische Vereniging Number 4:1-412. Keeton, W. T. 1960. A taxonomic study of the milliped family Spirobolidae (Diplopoda: Spirobolida). Memoirs of the American Entomological Society No. 17:1-146. Kevan, D. K. McE. 1983. A preliminary survey of known and potentially Canadian millipedes (Diplopoda). Canadian Journal of Zoology, 61:2956- 2975. Kevan, D. K. McE., and G. G. E. Scudder. 1989. Illustrated keys to the families of terrestrial arthropods of Canada 1. Myriapods (Millipedes, Centipedes, etc.). Biological Survey of Canada Taxonomic Series No. 1:1-88. Loomis, H. F. 1971. The diplopod genus Polyzonium in North America. Florida Entomologist, 54:151-163. Mauries J.-P. 1964. Sur quelques diplopodes de la peninsule Iberique. Bulletin de la Societe d’Histoire Naturelle de Toulouse, 99:157-170. Scudder, G. G. E. 1994. An annotated systematic list of the potentially rare and endangered freshwater and terrestrial invertebrates in British Columbia. Entomological Society of British Columbia Occasional Pa- per Number 2:1-92. Hirudisomatid Millipeds 143 Shear, W. T. 1973. Millipeds (Diplopoda) from Mexican and Guatema- lan caves. Accademia Nazionale dei Lincei, Problemi Attuali di Scienza e di Cultura, 171:239-305. Shear, W. T. 1986. Millipeds from caves in Mexico and Central America. V. New species and records of Glomeridae, Trichopetalidae, Cleidogonidae, Furmannodesmidae, Cryptodesmidae, Cambalidae, Typhlobolellidae, Rhacodesmidae, and Sphaeriodesmidae. Texas Memorial Museum, Speologi- cal Monograph, 1:63-86. Shelley, R. M. 1976. Two new diplopods of the genus Polyzonium from North Carolina, with records of established species (Polyzoniida: Polyzoniidae). Proceedings of the Biological Society of Washington, 88:373-382. Shelley, R. M. 1987. The milliped Stenodesmus tuobitus (Chamberlin) (Poly- desmida: Xystodesmidae) in Texas and New Mexico. National Geo- graphic Research, 3:336-342. Shelley, R. M. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Canadian Journal of Zoology, 66:1638-1663. Shelley, R. M. 1990. A new milliped of the genus Metaxycheir from the Pacific Coast of Canada (Polydesmida: Xystodesmidae), with remarks on the tribe Chonaphini and the western Canadian and Alaskan diplopod fauna. Canadian Journal of Zoology, 68:2310-2322. Shelley, R. M. 1993. Revision of the milliped genus Scytonotus Koch (Polydesmida: Polydesmidae). Brimleyana, 19:1-60. Shelley, R. M. 1994a. The milliped family Nearctodesmidae in northwestern North America, with accounts of Sakophallus and S. simplex Chamberlin (Polydesmida). Canadian Journal of Zoology, 72:470-495. Shelley, R. M. 19946. The Chonaphini, a biogeographically significant milliped tribe in eastern and western North America (Polydesmida: Nearctodesmi- dae). Brimleyana, 20:111-200. Shelley, R. M. 1994c. Revision of the milliped family Paeromopodidae, and elevation of the Aprosphylosomatinae to family status (Julida: Paeromopo- doidea). Entomologica Scandinavica, 25:169-214. Wood, H. C. 1864. Description of new genera and species of North American MYRIAPODA. Proceedings of the Academy of Natural Sciences, Phila- delphia, 16:186-187. Wood, H. C. 1865. The Myriapoda of North America. Transactions of the American Philosophical Society, 13:137-248. Wood, H. C. 1867. Notes on a collection of California Myriapoda, with the descriptions of new eastern species. Proceedings of the Academy of Natural Sciences, Philadelphia, 19:127-130. Received 5 June 1995 Accepted 3 October 1995 DATE OF MAILING Brimleyana 22 was mailed on 19 October 1995. Publications of the North Carolina Biological Survey Endangered, Threatened, and Rare Fauna of North Carolina Part I. A Re-evaluation of the Mammals Mary K. Clark, editor, 52 pages, 1987, $5 postpaid Part II. A Re-evaluation of the Marine and Estuarine Fishes Steve W. Ross, Fred C. Rohde, and David G. Lindquist, 24 pages, 1988, $3 postpaid Part III. A Re-evaluation of the Birds David S. Lee and James F. Parnell, editors, 52 pages, 1990, $8 postpaid Atlas of North American Freshwater Fishes David S. Lee et al., 867 pages, 1980, $25 postpaid Atlas of North American Freshwater Fishes, 1983 Supplement D. S. Lee, S. P. Platania, and G. H. Burgess, 68 pages plus looseleaf additions and corrections, 1983, $10 postpaid A Distributional Survey of North Carolina Mammals David S. Lee, John B. Funderburg, and Mary K. Clark, 70 pages, 1982, $5 postpaid s> The Seaside Sparrow, Its Biology and Management Thomas L. Quay et al., editors, 174 pages, 1983, $15 postpaid Autumn Land-bird Migration on the Barrier Islands of Northeastern North Carolina Paul W. Sykes Jr., 50 pages, 1986, $5 postpaid Potential Effects of Oil Spills on Seabirds and Selected Other Oceanic Vertebrates Off the North Carolina Coast David S. Lee and Mary C. Socci, 64 pages, 1989, $8 postpaid Bird Life of North Carolina’s Shining Rock Wilderness Marcus B. Simpson, Jr., 32 pages, 1994, $5 postpaid Send orders to: N.C. State Museum of Natural Sciences Attention: Beverly Craven P.O. Box 29555, Raleigh, NC 27626-0555 Please make checks payable in U.S. currency to Museum Extension Fund. 96-5869 BRIMLEYANA NO. 23, DECEMBER 1995 CONTENTS Life History of Cobia, Rachycentron canadum (Osteichthyes: Rachycentridae), in North Carolina Waters. Joseph W. Smith 1 A Review of Stonefly Records (Plecoptera: Hexapoda) of North Carolina and South Carolina. Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat 25 Seasonality in Cetacean Strandings Along the Coast of North Carolina. Wm. David Webster, P. Dawn Goley, Jessie Pustis, and Joseph A. Gouveia 41 Fishes New or Rare on the Atlantic Seaboard of the United States. Fred C. Rohde, Steve W. Ross, Sheryan P. Epperly, and George H. Burgess 53 A New Species of Crayfish of the Genus Orconectes, Subgenus Procericambarus (Decapoda: Cambaridae), Endemic to the Neuse and Tar-Pamlico River Basins, North Carolina. John E. Cooper and Martha Riser Cooper 65 Nine-banded Armadillo, Dasypus novemcinctus (Mammalia: Edentata), in South Carolina: Additional Records and Reevaluation of Status. Steven G. Platt and William E. Snyder 89 Post-hibernation Movement and Foraging Habitat of a Male Indiana Bat, Myotis sodalis (Chiroptera: Vespertilionidae), in Western Virginia. Christopher S. Hobson and J. Nathaniel Holland 95 The Milliped Family Hirudisomatidae in the New World (Polyzoniida). Rowland M. Shelley 103 Miscellany 144