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number 23

december 1995

EDITORIAL STAFF

Richard A. Lancia, Editor

Suzanne A. Fischer, Assistant Editor

Eloise F. Potter, Production Manager

EDITORIAL BOARD

James W. Hardin

Professor of Botany

North Carolina State University

Rowland M. Shelley

Curator of Invertebrates

North Carolina State Museum

of Natural Sciences

William M. Palmer, Emeritus

Director of Research and Collections

North Carolina State Museum

of Natural Sciences

Robert G. Wolk

Director of Programs

North Carolina State Museum

of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State

Museum of Natural Sciences, appears twice yearly in consecutively

numbered issues. Subject matter focuses on systematics, evolution,

zoogeography, ecology, behavior, and paleozoology in the southeastern

United States. Papers stress the results of original empirical field studies,

but synthesizing reviews and papers of significant historical interest

to southeastern zoology are also included. Brief communications are

accepted.

All manuscripts are peer reviewed by specialists in the Southeast

and elsewhere; final acceptability is determined by the Editor. Address

manuscripts and related correspondence to Editor, Brimleyana, North

Carolina State Museum of Natural Sciences, P.O. Box 27555, Raleigh,

NC 27626. Information for contributors will be sent upon request.

Address correspondence pertaining to subscriptions, back issues,

and exchanges to Brimleyana Secretary, North Carolina State Museum

of Natural Sciences, P.O. Box 27555, Raleigh, NC 27626-0555.

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Environment,

Health, and Natural Resources

James B. Hunt Jr., Governor

Jonathan B. Howes, Secretary

CODN BRIMD 7

ISSN 0193-4406

Life History of Cobia, Rachycentron canadum

(Osteichthyes: Rachycentridae), in North Carolina Waters

Joseph W. Smith

National Marine Fisheries Service

Southeast Fisheries Science Center

Beaufort Laboratory

101 Fivers Island Road

Beaufort, North Carolina 28516-9722

ABSTRACT. — Cobia ( n = 416) were collected primarily along the

central North Carolina Atlantic coast from recreational anglers between

1983 and 1994. Males ( n = 174) ranged up to 136-cm fork length

(FL) and 32.0 kg, and females ( n = 182) up to 142-cm FL and

32.2 kg. Most cobia greater than 100-cm FL were females. Ages

of cobia (to age 14) were estimated by counting opaque zones on

cross-sectioned sagittal otoliths. Von Bertalanffy growth parameter

(k) estimates were 0.37 for males and 0.24 for females. Adult co-

bia occurred in major sounds and coastal Atlantic waters of North

Carolina from May through July, and in nearshore oceanic waters

through fall. Cobia may overwinter between Cape Fear and Cape

Canaveral at depths of 30-75 m. Cobia fed chiefly on demersal

crustaceans and fishes in the study area. Cobia may be one of the

few teleosts that regularly consumed small elasmobranchs. Male cobia

were sexually mature at 60-65-cm FL (age 2), and females at 80-

cm FL (age 2). Cobia spawned May through July along the North

Carolina coast, and ocean waters adjacent major coastal inlets were

probable sites for cobia spawning activity.

Cobia, Rachycentron canadum, a large, coastal fish of the monotypic

family Rachycentridae, has a cosmopolitan distribution in tropical to

warm temperate seas, except for the eastern Pacific Ocean (Briggs

1960, Shaffer and Nakamura 1989). Cobia occur during summer in

the United States coastal waters of the northern Gulf of Mexico and

along the Eastern Seaboard from the Florida Keys north to Cape Cod

(McClane 1965), although they are uncommon north of Chesapeake

Bay (personal observations). Cobia migrate north along the Atlantic

coast from northern Florida to the Carolinas, and then into Chesapeake

Bay (McClane 1965, Shaffer and Nakamura 1989) during spring and

summer. By late spring and early summer cobia enter polyhaline to

mesohaline areas of major coastal bays, sounds and river systems in

the Carolinas and Virginia (Musick 1972, Moore et al. 1980, Schwartz

et al. 1981). Lone fish or “pods” of several cobia often hover in

the shadow of near-surface objects, such as buoys, boats, sharks, and

Brimleyana 23:1-23, December 1995

1

2

Joseph W. Smith

rays (Joseph et al. 1964, McClane 1965, Shaffer and Nakamura 1989).

Their size, commonly exceeding 23 kg (McClane 1965), and nearshore

residence during spring through summer, make them a favorite of

coastal recreational fishermen. Recent estimates (1991) place recreational

cobia landings along the United States south Atlantic coast (292,600

kg) at five times that of commercial landings (58,000 kg)(Isley 1992).

To date, Richards (1967) conducted the most comprehensive life

history study of cobia on the Atlantic coast of the United States,

collecting specimens during the mid-1960s in lower Chesapeake Bay.

Various facets of cobia biology have been examined, including feeding

habits (Knapp 1951, Darracott 1977), reproduction (Biesiot et al. 1994),

spawning areas and season (Joseph et al. 1964), movements and growth

(Richards 1977, Franks 1995), rearing eggs and larvae (Hassler and

Rainville 1975), and egg and larval distributions (Ditty and Shaw

1992). Recent mitochondrial DNA analyses (Hrincevich and Biesiot

1994) suggested that cobia from the northern Gulf of Mexico and

the south Atlantic coast of the United States should be considered

a unit stock. Shaffer and Nakamura (1989) compiled a biological synopsis

of the species.

My interest in cobia stems from (1) a perceived increase in fishing

effort for the species along the North Carolina coast during the 1980s,

including a directed charter boat fishery for cobia at Ocracoke Inlet

and the establishment of a cobia fishing tournament in Carteret County,

and (2) the lack of contemporary fishery statistics on which to base

cobia stock assessments (Gulf of Mexico and South Atlantic Fishery

Management Councils 1985, Isley 1989). Objectives were to elucidate

various aspects of cobia life history in North Carolina waters, in particular,

age and size composition of the recreational catch, distribution, feeding

habits, and reproduction.

MATERIALS AND METHODS

Recreational fishermen in the Morehead City-Beaufort area (Carteret

County) of the central North Carolina coast (Fig. 1) were the major

sources of specimens from 1983 to 1994. Beginning in June 1987

and each spring thereafter, fish were processed at a local cobia tournament.

Additionally, during 1989-92 charter boat captains and tackle shop

proprietors at Ocracoke Island and Hatteras, North Carolina, provided

frozen cobia carcasses, individually labeled with date, location of capture,

and whole (round) mass; for most of these specimens the head, axial

skeleton and viscera were intact. Carcasses were returned to the laboratory

biweekly for processing. Additional specimens came from pound nets

and haul seines in Pamlico Sound near Cape Hatteras, ocean research

Life History of Cobia

3

Fig. 1. Major sampling sites (arrows) for cobia along the North Carolina

coast, 1983-94.

4

Joseph W. Smith

cruises between Cape Lookout, North Carolina, and northern Florida,

research trawls in lower Chesapeake Bay, and port agents in South

Carolina and northeast Florida.

Whole cobia were weighed to the nearest 0.1 kg. Carcasses and

whole cobia were measured for total (TL) and fork length (FL) in

centimeters and sexed. Gonads were staged for maturity based on criteria

in Waltz et al. (1979), then excised and weighed to the nearest gram.

Subsamples of fresh gonadal tissue from 99 cobia were preserved in

10% buffered formalin, and later sectioned by standard histological

techniques (Humason 1972) to verify maturity staging in the field.

A gonadosomatic index ( gsi ) was computed for sexually mature specimens,

whereby gsi = (gonad mass/body mass) x 100. Axial skeletons were

missing from some frozen specimens, as catches were “steaked” versus

filleted. Fork lengths for fish lacking an axial skeleton were estimated

by calculating a regression of FL on intraorbital distance (measured

with a caliper in mm) from whole fish (Table 1). Fork length was

then assigned to carcasses based on this regression.

Table 1. Mass-length (In) and length-length regression equations for cobia from North

Carolina and adjacent waters, 1983-94.

a W = fish mass in kg, 10 = intraorbital distance in mm, FL and TL in cm.

b I = undifferentiated specimens.

Stomachs were examined and the contents were preserved in 10%

formalin and later transferred to 50% isopropanol. Bait or chum (fish

that had obviously been sliced or cut by anglers, mostly Atlantic menhaden,

Brevoortia tyrannus, pinfish, Lagodon rhomboides, and various sciaenids)

occurred in 37 stomachs; these items were eliminated from any analyses,

as were 15 stomachs where bait or chum was the only food item

Life History of Cobia

5

present. Represented food items were drained, identified, counted, and

weighed to the nearest gram.

Importance of each prey item to the cobia diet was based on

an index of relative importance (iri; Pinkas et al. 1971). Percent frequency

of occurrence for each item in non-empty stomachs (/), percent total

number of prey items («), and percent total mass of prey items (w)

were calculated. The original iri formula was modified to use the

mass of a prey item instead of volume, ( iri = f(n+w)). The results

were examined for areal differences in diet (Beaufort Inlet and vicinity,

Ocracoke and Hatteras inlets and vicinity, and offshore oceanic waters).

To determine changes in cobia food habits with growth, specimens

with food items were partitioned into arbitrary size classes (<4.5, 4.5-

9.0, and >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

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Number Number

Life History of Cobia

9

Fork Length (mldpts. of 5 cm Intervals)

Fig. 3. Fork length frequency distributions (all gears and hook-and-line) by

5-cm increments for male and female cobia from North Carolina and adjacent

waters, 1983-94.

10

Joseph W. Smith

Table 3. Von Bertalanffy parameter estimates by sex describing the growth of cobia

from North Carolina and adjacent waters, 1983-94; CL = 95% confidence limits. Richards’

(1977) estimates shown for comparison.

coral patches of low relief (Huntsman 1976). By early May, cobia

were found on nearshore artificial reefs and under navigation buoys

in the vicinity of Beaufort, Ocracoke, and Hatteras inlets (Manooch

et al. 1981). The earliest record for cobia caught by hook-and-line

in North Carolina estuarine waters during the study was 8 May 1990.

Initial spring catches in the sounds coincided with inshore water temperatures

reaching 20 C and higher. Most “inshore” angling activity for cobia

was concentrated in Bogue and Back sounds adjacent to Beaufort and

Bardens inlets near Cape Lookout, and Pamlico Sound adjacent to

Ocracoke and Hatteras inlets near Cape Hatteras (Fig. 1). Traditional

fishing locations for cobia in North Carolina’s inlets, sounds, and

coastal rivers were poly- to mesohaline waters >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).

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

Life History of Cobia

13

Table 5. Continued.

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

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Life History of Cobia

15

All Areas (N = 94 stomachs)

% iri

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< 4.5 4.5 - 9.0 > 9.0

Cobia Size Class (kg)

Eiasmobranchs

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Crabs

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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.

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

Stonefly Records

29

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

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)

X12

X1-13

X1

Stonefly Records

31

Table 1. Continued.

32 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat

Table 1. Continued.

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

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

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

Peltoperlidae

* Peltoperla arcuata Needham (TN, VA)

* P. tarteri Stark and Kondratieff (VA)

*N. stewarti Stark and Baumann (TN, VA)

Stonefly Records

37

Table 2. Continued.

38 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat

Table 2. Continued.

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

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Table 1. Continued.

Cetacean Strandings

45

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46

Wm. David Webster et al.

Tursiops truncatus

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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

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oc

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Balaenoptera acutorostrata

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-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.

* 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.

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

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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.

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

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

116

Rowland M. Shelley

Table 1. Continued.

Males Females

Hirudisomatid Millipeds

117

Table 1. Continued.

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.,

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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

Nat. Pk.

122

Rowland M. Shelley

Table 2. Continued.

Hirudisomatid Millipeds

123

Table 2. Continued.

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,

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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).

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

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).

132

Rowland M. Shelley

Table 5. Continued.

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

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Hirudisomatid Millipeds

137

Figs. 31-38. M. absidatum, holotype. 31-33, pigmentation patterns of ante-

rior, midbody, and caudal segments, respectively. 34, head and collum, anterior

view, setation and pigmentation omitted. 35, profile of midbody segment,

caudal view. 36, left anterior gonopod and sternum, anterior view. 37, the

same, caudal view. 38, left posterior gonopod, caudal view. Scale line for

figs. 31-33 = 1.00 mm for each; line for figs. 34-35 = 0.50 mm for each;

line for figs. 36-38 = 0.25 mm for each.

138

Rowland M. Shelley

at sites that are latitudinally in the Neotropics or the transition zone

between the two biogeographic regions. In the Parajulidae, Causey

(1974) reported that Mexican forms occur in the temperate zone near

the coast but are absent from the “tierra caliente” lowlands; they are

most abundant on the plateau and follow the mountains southward

into western Guatemala, where they are restricted to the mountains.

Likewise in the Spirobolidae, Hiltonius Chamberlin extends southward

from southern Arizona to southwestern Guatemala, where forms occur

only “in high mountainous regions” (Keeton 1960). Finally, Sakophallus

simplex Chamberlin, a species with west-Nearctic affinities provisionally

assigned to the Nearctodesmidae, is known only from high elevations

in western Jalisco and Michoacan (Shelley 1994 a).

The Xystodesmidae, Nearctodesmidae, and Glomeridae also exhibit

substantial Mexico/United States range disjunctions (Table 6). The

degree of continuity is unknown in the Parajulidae, which has never

been comprehensively studied at the generic and specific levels, and

although Keeton (1960) revised Hiltonius, enough new material exists

in American repositories to reassess the Mexican forms, which range

northward into Santa Cruz County and adjacent parts of southern Arizona.

A hiatus appears to exist between southern Arizona and coastal California,

and there may or may not be one in Mexico. The Mexican representatives

of the Xystodesmidae, Rhysodesmus Cook and Stenodesmus Saussure,

also occur in the adjacent fringe of the United States, and there is

a sizeable lacuna between them and the east-Nearctic species, the

most proximate localities being in the lower Rio Grande Valley of

Texas and northern Louisiana (Hoffman and Shear 1964, Hoffman

1970, Shelley 1987). 1 An even larger gap exists between the most

proximate sites of Sakophallus Chamberlin, in Jalisco, and Nearctodes-

mus Silvestri, in Marin County, California (Shelleyl994fl). Regarding

Glomeroides, the only United States species, G. prima (Silvestri), occurs

in the San Francisco Bay area. The type locality is in Marin County;

Shear (1986) reported Pfeiffer Big Sur State Park, Monterey County;

and I now add Redwood Regional Park, Contra Costa/Alameda counties,

based on juveniles collected on 18 May 1953 by R. O. Schuster and

E. E. Gilbert (NMNH).

1 The Louisiana species is Boraria profuga (Causey), known previously only from the

type locality in Montgomery County, Arkansas, in the Ozark-Ouachita Physiographic

Province. The Louisiana locality, in the Gulf Coastal Plain, is Monroe, Ouachita County,

based on unreported males and females collected in December 1974 and 1978 by

M. R. Cooper (NCSM).

Hirudisomatid Millipeds

139

Table 7. Generic and Tribal, East/West Nearctic Disjunctions.

Proximate Probable

No. Components Localities Breadths Dispersal

Taxon West East (County, State) of Lacunae Directions

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.

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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