fhe Journal of the North Carolina
State Museum of Natural Sciences
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RICHARD A. LANCIA, Editor
SuzANNE. A. FISCHER, Assistant Editor
ELOISE F. POTIER, Production Manager
EDITORIAL BOARD
JAMES W. HARDI
Professor of Botany
ROWLAND M. SHELLEY
Curator of Invertebrates
North Carolina State Museum North Carolina State University
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Director of Research and Collections
North Carolina State Museum
of Natural Sciences
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 re sults of original empirical field studies,
but synthesizing reviews and papers of significant historical interest
to southeastern zoology are also included. Brief communications are
accepted.
A ll 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 i ssues,
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
CODN BRIMD 7
ISSN 0193-4406
NORTH CAROLINA 0FPARTMENT OF ENVIRONMENT,
H EALTH, AND NATURAL RESOURCES
lAMEs B. HuNT JR., GovERNoR
JoNATHAN B. Howes, SecRETARY
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 Pivers 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 co­bia
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
7 <r
~
w
u . ,-' 0
0 u \ ~ f-
~) z
<[
'6 _j
~~ 1-
;...-'..>-
<[
%.
o-
<[
<[ z z: :I:::::i \
(3 1-0
"'- "' "' ' '
0:: 0<[ > z u
<[ 4A z
"'- :I:::::i ~ ( 1-0
:>a::
<-., O «
U'IU
-:z: !_,--'
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.
Variables• Sexb n Equation Rz Range
W-FL 0 86 log W=3.4 log FL -13.3 0.972 0.5- 32.0 kg
~ 94 log W=3.2 log FL -12.3 0.949 0.7- 32.2 kg
o+'?+l 194 log W=3.4 log FL -13 .0 0.987 0.5- 32.2 kg
TL-FL 0 105 TL" l.l FL -1.1 0.989 39- 136 em FL
'? 97 TL,.l.1 FL +0.7 0.993 44- 142 em FL
o+'?+I 217 TL"l.l FL -0.9 0 .995 39- 142 em FL
FL-10 0 75 FL" 0.8 10 + 17.3 0.929 39-136 em FL
9 65 FL" 0.8 10 + 18.5 0.956 44- 142 ern FL
• W = fish mass in kg, IO = intraorbital distance in mm, FL and TL in em.
b I = undifferentiated specimens.
Stomachs were examined and the contents were preserved i:J 10%
formalin and later transferred to 50% isopropanol. Bait or cht;m (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 (n), 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 NUN 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
a)
Fig. 2. Cross-sections (0.5 mm thick) of cobia sagittae: a) sagitta from age
3 fish (89 em FL male, 18x magnification), b) sagitta from age 8 fish (125
em 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 recr eational hook -and-line fi shermen , 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 Caro lina (n = 11), and northeast Florida (n = 15). Research
trawls (75 -ft high-rise mongoose net) from Daytona Beach, Florida,
to Cape Lookout, 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 em and 0.47 to 32.0 kg, and 182 females ranged
from 44 to 142 em 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
em; conversely 91 of 174 (52.3 %) of the fema l es caught by the same
gear were greater than 100 em (Fig. 3).
North Carolina enacted bag (2 fish / angler/day) and minimum size
limits (33 inches [84 em] 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 exam ined, and 65 (24. 9 %) were le ss than 84 em. Between 1991
and 1994, only five (5.3 %) of 93 fish caught by hook-and-line were
less than 84 em, and four of these were 82-83 em.
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 f rom young-of-th e-year
cobia that averaged 31-cm (n = 1 7, range = 21-46-cm). Age 1 cobia,
or those with one opaque zone distal to the cor e, averaged 51 em
(n = 9, range = 39-64-cm) . Mean length of females was larger than
mean length for males at a given est imated 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 female s, although
mean asymptotic size was large r for females (Table 3).
SEASONALITY AND DISTRfBUTlON
Initial catches of cobia by North Carolina anglers usually occurred
in March or April 50- 65 km offshore over rocky outcroppings and
Table 2. Samp le s ize (n), fo rk length range, mean observed fork lengths (:t1 SE), mean observed mass, and von Bertalanffy estimates of
fork lengths (VB FL) for each sex of cobia from North Carolina and adjace nt waters, 1983 -94, by estimated age . Results of Richards'
(1967 and 1977) studies are presented for compari son. All lengths are in ern. 00
Prese nt Study Richards (1967)1 Richards ( 1977)
X X X X
Estimated FL observed mass VB 2 observed mass
Sex Age n ran ge FL :t1SE kg n FL FL kg VB FL
Males 1 6 39- 64 so 4 1.3 6 56 54 1.5 31
2 22 63- 93 74 2 3.9 13 71 71 3 .8 5 3
3 41 68- 102 82 1 6.7 23 82 84 7.6 69
4 32 82- 97 88 1 8.3 14 89 94 9.6 82
5 20 78- 99 92 1 9.4 12 94 101 11. 9 91
6 7 90- 103 95 2 10.6 7 97 104 13.7 99
7 6 94- 108 100 2 11 . 5 5 100 105 14.5 104 ......
8 8 89- 107 99 2 13.6 5 101 108 0
CIJ
9 6 99- 136 107 6 18.3 4 103 109 17.7 (1)
'0
10 5 101- 109 105 1 12.2 1 103 119 19.0 ::::r
11 3 102- 109 105 2 104 ~
12 0 104
13 1 113 19.3 1 104 C/)
14 1 106 20.0 1 105 2.
Fema les 1 3 49- 63 55 4 1.0 1 61 58 1.8 3 6 :;.
2 18 57-106 81 3 6.1 16 77 78 4.6 61
3 5 0 79- 99 89 1 8.4 36 89 95 11.1 82
4 23 88- 132 102 2 12.5 1 2 99 104 13.2 99
5 13 98- 113 106 1 14.9 10 1 07 116 19.7 112
6 20 99-126 111 2 17 .8 14 113 120 21.8 122
7 11 110- 12 6 117 2 21.0 10 117 126 25.1 131
8 8 114- 128 123 2 23 .9 6 121 130 2 8.4 137
9 7 114- 134 125 2 24.6 5 124 133 30 .5
10 3 117- 133 127 5 29.7 2 126
11 1 121 18.1 1 128
12 3 125- 130 127 2 28.6 1 130
13 2 134- 14 2 138 4 32.0 2 131
1 Values converted to metric units from published English units.
2 Values are von Bertalanffy estimates of FL based on individua l FL s at estimated age .
35
30
25 -
:u 20 -
.0
E
~ 1 5 -
1 0 -
5 -
0
30
25
20 --'
:u
.0
E 15 -
:I z
10 -
Life History of Cobia
.Hook & Un• (n•112)
O.t.u O••n (nc 114) Males
nnn~ I I., .,
I I I
35 45 55 65 75 85 95 1 05 115 125 135 145
I . Hook & U n• (n•174)
0AII a •• ,. (n•182) Females
35 45 55 65 75 85 95 105 115 125 135 145
Fork Length (mldpts. of 5 em Intervals}
9
Fig. 3 . Fork length frequ e ncy distribut ions (all gears and hook-and -line) by
5-cm increment s fo r male and f emale cobia from North Carolina a nd adjacent
wa ters, 1983-94.
10 Joseph W. Smith
Table 3. Von Bertalanffy parameter estimates by sex describing the growth of cobia
from North Ca rolina and adjacent waters, 1983-94; CL = 95 ~o confidence limits. Richard s'
(1977) es timates shown for comparison.
Asymptotic Asymptotic CL Ri c hards ' (1977)
Sex Parame ter Estimate SE lower upper estimates
Males I~ 105 1.85 101 108 121
k 0.37 0 .04 0.29 0.45 0.28
t, - 1.08 0 .29 -1.65 -0.51 -0.06
Fema les I ~ 135 3.82 127 142 164
k 0.24 0.03 0.18 0.31 0.23
t, -1.53 0 .39 -2.30 - 0.77 -0.08
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
si tes 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 ne arshore ocean adjacent to buoys and fishing piers, and over
artificial r eefs 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
s ummer. 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 a rbitrarily chosen).
Dates May June July August September
1-7 1 45 8 0 0
8-15 5 100 5 0
16 - 22 27 6 3 0
23 - 31 20 33 5 0 0
Totals 53 184 21
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.
Fooo HABITS
During 1989-1990, 140 cobia stomachs were examined, of which
72.1 % (n = 101) contained representative food items . lri's were computed
from these samples and nine addi tional 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, t he blue c r ab,
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, followe d by the smooth dogfish, pipefishes , and dasyatid sting
rays. Abundant crustaceans included the iridescent swimming crab,
Portunus gibbesii, the brown shrimp, Penaeus aztecu s, and the mantis
shrimp, Squilla empusa. High-ranking food items from the Hatteras­Ocracoke
area (Table 6) were the blackcheek tonguefisb and the blue
crab. Important food items from offshore waters (Table 6) included
the coarsehand lady crab, Ovalipes stephensoni, unidentifiable fishes ,
the blotc hed 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 frequen cy of occurrence (f), percent number (n), percent mass (w),
and index of re lative importan ce (iri) of food items in cobia stomachs from North Carolina
and adjacent waters, 1987-90.
Prey Taxa f II IV iri
Mollusca
Ceph a lopoda
Loligo plei 0.9 0.2 <0.1 <1
Art hrop od a
Cr ustacea
Stomatopoda
Squi/la sp. 2.7 0 .7 0.4 3
S. empusa 3.6 1.3 2.5 14
S. neglecta 1.8 4 .2 2.5 12
Decapoda
Penaeid ae
P ena eus sp. 3.6 0. 9 0 .2 4
P. aztecus 6.4 2.9 3.7 4 2
P. setiferus 0.9 0.2 0 .2 < 1
Trachy penaeus co nstrictus 0.9 1.5 <0. 1 <1
Sicyoniidae
Sicyonia sp. 3.6 7.9 2. 7 38
Palaemonidae
Palaemonetes vulgaris 0.9 0.4 <0.1 <I
Cra ngoni dae
Crangon sept emspinosa 4.5 2.6 0. 2 13
Upogcbiidae
Upog ebia sp. 0.9 0.2 <0 .1 <1
Albunci dae
Albunea g ibbesii 0.9 0 .2 0.1 <1
Portunidae
Ova lipes sp. 6.4 3.7 2. 0 36
0. ocellaws 2.7 0 .9 1.1 5
0 . stephe nso ni 6.4 4 .6 4.0 55
Callinectes sp. 7.3 3.3 1.8 37
C. sapidus 30.0 15.4 19.2 1,038
C. simi/is 3 .6 0 .9 0 .8 6
Portwws sp. 3.6 1.1 0 .5 6
P . g ibb esii 5.5 2.2 1.8 22
P . spinimanus 2.7 0.7 1.4 6
Unidentified portunid remains 4.5 2.2 0 .3 1 1
Xanthidae
Menipp e mercenaria 0.9 0.4 0.1 <1
U nide ntified de capod r emains 0.9 0.2 <0.1 <1
Chon dri c ht hyes
Carcharhinida e
Mustelu s ca nis 6.4 6.1 21.2 175
Life History of Cobia 13
Tab le 5. Contin ued .
Prey Taxa f II w iri
Dasyatidae
Da syaris sp. 3.6 0.9 12.7 49
Osteich thyes
Clupeidae
Opistlronema oglinum 0.9 0 .4 0.4
Engraulidac
A11 clroa sp. 0 .9 L.3 0.1
Synodon tidac
Synodus foerens 0.9 0. 2 0.3 <1
Bat rachoididae
Opsa11us sp. 3.6 1.1 2.7 14
Syngnathidae
Hippocampus sp. 0.9 0.2 <0.1 <1
Syn g narlrus sp. 19.1 7.0 2.2 176
Sparidae
Lagodon rhomboides 0.9 0.2 0.2 <1
Uranoscopidae
unide nt ified re mains 0.9 0 .2 <0.1 <I
Soleidae
Trinecres macu/atus 0 .9 0.7 0.4
Cynoglossid ae
Symplrurus plag iusa 17.3 13.2 6.8 346
Ba listidae
unidentified remains 2.7 1.3 0. 1 4
Tet radontidae
Sphoeroides macularus 0 .9 0.7 1.9 2
Diodontida e
Chilomycrerus schoepfi 0.9 0 .2 1. 2
unidentified fish rema ins 2 1.8 7.5 4.5 262
inges ted. Smooth dogfi s h pups (n = 28) averaged 4 2 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, x = 34 g), and toadfishes , Opsanus
sp . (n = 5, x = 2 9 g). Most portunid crabs were less than 7 em in
carapace width (CW) and were ingested whole; commercial-sized blue
crabs (ca. 12.5-cm CW) were rarely consumed. Ova l ipid c r abs wer e
often macerat ed. Small balistid fis h es occurred in the stomachs of
juvenile cobia fro m offshore trawl catches and w ere among the smalles t
teleosts consumed (n = 6, :x = 1 g) .
As cobia increased in size, penaeid shrimps and teleost fishes
became relativ ely less importan t in the diet, while decapod crabs inc reased
Table 6. The ten highest ranked prey items in cobia stomachs by sampling area m North Carolina' , 1987- 90.
Beaufort Inlet (n 40) Ocracoke-Hatteras Inlets (n 56) Offshore Areas (n 14)
Prey iri Prey iri Prey
Ca llinectes sapidus 1,609 Symphurus plagiusa 1,311 Ovalipes stephensoni
Mustelus canis 802 Callinectes sapidus 1,111 Unidentified fish
Syngnathus sp. 236 Unidentified fis h 355 Portunus spinimanus
Dasyatis sp. 208 Ovalipes sp. 164 Sicyonia sp.
Unidentified fish 133 Syngnathus sp. 130 Unident. Balistidae
Portunus gibbesii 89 Sicyonia sp. 69 Syngnathus sp.
Penaeus aztecus 73 Opsanus sp. 56 Trachypenaeus constrictus
Squilla empusa 69 Penaeus aztecus 42 Anchoa sp.
Callinectes sp. 55 Crangon septemspinosa 32 Synodus foetens
Symphurus plagiusa 38 Callinectes sp. 31 Symphurus plagiusa
1 n = number of stomachs examined by locality.
iri
870
654
561
516
277
182
95
90
41
35
'-<
0
(/)
0>
'0 ::r
(/)
3 -· -::r
70
60
50
40
30
20
10
0
Life History of Cobia
All Areas (N = 94 stomachs)
% iri
< 4. 5 4.5- 9.0 > 9.0
Cobia Size Class (kg)
15
• E:usmobrqnchs
TI-clcoct::;
Dcrabs
~Shrirrps
Fig. 4. Percent iri's for various prey groups by cobia mass interva l (intervals
arbi trarily chosen).
in importance (Fig. 4). Elasmobranchs, 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.
REPR ODUCTION
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. Sa). 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 em FL (Table 7), o r age 2 .
Most female cobia examined were staged as developing (Table
7), and most were sexually ma t ure by 80 em FL, or age 2. Histological
sections revealed that the ovaries of early developing females had
16 Joseph W. Smith
Tab le 7. Cobia from Non h Carolina and adjacent waters, 1983-94, in various stages
of sexual development by 5-cm-FL intervals.
Midpoint Males Femal es
5-cm-FL Immature Develop ing and Ripe Immature Developing Ripe
:S 50 5 0 4 0 0
55 0 1 0 0
60 1 3 0 0
65 0 6 0 0
70 0 5 1 2 0
75 0 22 5 0 0
80 0 23 0 6 0
85 0 25 0 11 0
90 0 11 0 19 1
95 0 17 0 8 0
10 0 0 7 0 9 0
105 0 0 0 11 0
110 0 l 0 9 0
1 15 0 0 0 5 0
120 0 6 0
125 0 8 0
130 0 2 0
135 0 2 0
140 0 0
Totals 7 120 13 99
many small basophilic oocyte s with a few early vitellogenic oocytes
(Fig. 5 b) , whereas th e ovaries of la te de veloping females had large
(ca. 750 pm), yolk-filled oocycte s (Fig. 5c). Only one female had
h yd r ated oocytes; it was uncertain if th is fi sh was caught in estuarine
or oceanic wa ters. A few females (collected in early June 1990) showed
follicular atresia indicative of a recent spawn, yet also possessed numerous
large oocytes, suggestive of a n incipient spawn (Fig. 5d).
Mean gsi's for female cobia were hig h 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 a t 5 .3 (SD = 2 .2, n = 8). The largest ovaries excised
weighed 2 .49 kg (7 June) and were in a female weig hing 25 .4 k g.
Peak spawning in June 1989 wa s confirmed by neuston net collections
of cobia eggs fr om a channel in the lo wer Newport River estuar y
about 3 km from Beaufort Inlet (Fig . 1) (L. Settle, National Marine
Fisheries Service , Beaufort, North Car o lin a , unpublis hed data). During
10 sampling da tes be t ween 14 June and 18 August, peak cob ia egg
Life History of Cobia 17
d)
Fig. 5. Histologic preparations of cobia gonad sections: a) ripe male (70
em FL), b) early developing female (80 em FL), c) late developing female
(104 em FL), d) partially spent female (88 em 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 m 3), and 4 August (28 eggs/100
m 3). 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 a!. 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 fr om the south Atlantic coast of the
United States overwin t er. Winter trawl s urveys 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 em, x =
84 em) 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 a!. 197 9). 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 y ear, 1983-91 (L. Mercer, North Carolina Division of Marine
Fis heries, 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 Richa rds (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 pres ent 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 teleos ts, 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
pre y, such as portunids, stomatopods, penaeids, and eels in cobia stomachs
from the northern Gulf of Mexico. Cobia from the western Indian
Ocean consumed mos tly 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 thes e were among the largest prey items ingested. Cobia may
20 Joseph W. Smith
be one of the few teleosts that regularly consumed small e lasmobranchs.
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 em (age 2), and females by 80 em (age 2). Richard s (1967)
s tated that the smallest mature male in hi s collections measured 51.8
em ("second .... year of life") and that the smallest mature fema le measured
69.6 em ("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 No rth 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
o f Mexico s uppo rt the concept of batch s pawning in cobia.
Precise location of cobia spawning areas along the North Carolina
coast was uncertain, although my results suggested that cobia spawned
adjacen t the state's major ocean inlets. Likewise, Joseph et a l. (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 inle ts. Management
regulations adopted by North Carolina in 1991 prohibiting possession
of cobia Jess than 84 em were effective, and few fish below the minimum
possession size were encountered between 1991 and 1994. Migrato ry
routes and overwintering grounds of cobia along the south Atlantic
coast of the United States are uncl ear. 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 su rveys, 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 samp l es 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 sampl es 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 Labo ratory
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 s hared early life
hi story 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.
Bies iot, P. M., R. E . Caylor, and J. S. Franks. 1994. Biochemical and
histological changes during ova rian development of cobia , Rachycentron
canadum , from the northern Gulf of Mexico. Fishery Bu llet in 92:686-
696.
Briggs , J . C . 1960. Fishes of world wide (circumtropical) distribution.
Copeia 1960:171-180.
C hilton, D . E. , and M. Stocker. 1987. A comparison of otolith and scale
methods for ageing Pacific herring. North American J ournal of Fish­eries
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Darracott, A. 1977. Availability, morphometries, feeding and breeding activity
in a multi-species, demersal fish s t ock of the Western Indian Ocean.
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Di tty, J. G., and R. F. Shaw. 1992. Larva l development, distri bution, and
eco logy of cobia Rachycentron canadum (Family: Rachycentridae) in
the northern Gulf of Mexico. Fishery Bulletin 90:668-677.
22 Joseph W. Sm i th
Frank s, J. S. 1995. Investigat ions o f cobia, R achycentro n canadum, in
Mississippi marine waters and adjacent G ulf waters. Study II. Stud­ies
on the seasonal movements and migratory patterns of cobia, Rachycentron
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Ann ual Report, Project No . F-91, Segment No. 6, Sport Fish Resto­rat
ion P rogram, Gu lf Coast Research Laboratory, Ocean Springs, Mississippi.
Franks, J. S., J. T. McBee, and M. T. Allen. 1991. Estimations of age
in cobia, Ra chycentron canadum , from the northern Gulf of Mexico .
(Abstra ct) 55th Meeting of the Mississippi Academy of Scien ces, page
55, 21-22 February , Jackson, Mississippi.
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F inal amendment 1, fishery management plan, environmental impact
statement for the coasta l migratory pelagic resources (mackerels) . South
Atlant ic Fi she r y Managemen t Council, C harleston, South Carolina.
Hassler, W. W. , and R. P. Rainville. 1975. Techniques for hatc hing and
rearing cobia, Rachycentron ca nadum, t hrough larval and juvenile stages.
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Hrincev ich, A. W., and P. M. Biesiot. 1994 . Mitochondrial DNA a nal y ­ses
of cobia, Ra chycentron canadum , from the northern Gulf o f Mexico
and the Chesapeake Bay. ( A bstract) 58th Meeting of the Mississippi
Academy of Sciences, page 61, 17-18 February, Biloxi, Mississippi.
Humason, G. L. 1972. Animal tissue techniques. Second Edition. Free­m
an, San Francisco, California.
Hunt s ma n, G. R. 1976. Offshore he adboat fi shing in North Carolina and
South Carolin a. Marine Fisheries Review 38(3): 13-23.
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Copeia 1951:101- 102.
Joseph, E. B., J. J. Norcross, and W. H. Massman. 1964. Spawn ing of
the cobia, Ra chycenlrorz canadum, in the Chesapeake Bay area, wit h
observa tions of juvenile specimens. Chesapeake Science 5 :67-71.
Manooch, C. S., III. 1984. Fis h erman 's guide, fishes of the southeastern
United States. North Carolin a State Muse um of Natural History, Ra­leigh,
No rth Carolina.
M anooch, C. S., III, L. E. Abbas, and J. L. Ross. 1 981. A bi o l ogi cal
and economic analysis of the North Carolina charter boat fish ery. Marine
F isheri es Review 43(8): 1- 11.
McClane, A. J . 1965. McCla ne's standard f ishing encyclopedia and in­ternational
a ngling gui de. Ho lt, Reinhart and Winston, New York, New
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Moore, C. J., D. L. H a mmond , and D. 0. Myatt, III. 1980 . A guide to
saltwater recrea tional fisheries i n South Carolina. South Caroli na Wildlife
and Marine Resources Departm ent, Charles to n, South Carolina.
Musick, J. A. 1972. Fishes of Chesapeake B ay a nd the adjacent Coast a l
Plain. Virginia Institute of Marine Scien ce Special Scientific Report
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Life History of Cobia 23
albacore, bluefin tuna, and bonito in California waters. Ca li forni a Department
of Fish and Game, Fishery Bulletin 152.
Richard, J. 1989. Tag recovery report. Tide Magazine (March/Apri l 1989):30-
31.
Richard s, C. E. 1967. Age, growth and fecundity of the cobia, Rachycentron
canadum, from Chesapeake Bay and adjacent Mid-Atlantic waters. Transactions
of the American F isheries Society 96:343-350.
Richards, C. E. 1977. Cobia (Rachycentron ca nadum) tagging within
Chesapeake Bay and updating of growth equations. Chesa p ea ke Sci­ence
18:310- 311.
SAS Institute, Inc. 1987. SAS/STAT guide for pers onal computers, Ver­sion
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freshwater fishes of the Cape Fear estuary, Nor th Carolina, and their
distribution in relation to environmental factors. Brim leyana 7:17-37.
Shaffer, R . V. , and E. L. Nakamura. 1989. Synopsis of biological data
on cobia, Ra chycentron canadum (Pisces: Rachycentridae). National
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Waltz, W., W. A. Roumillat, and P. K. Ashe. 1979. Distribution, age
struc ture, and sex composition of the black sea bass, Centropristis
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Received 27 September 1994
Accepted 21 March 1995
A Review of Stonefly Records (P l ecopter a: Hexapod a) of
North Carolina and South Carolina
B OR IS c. KON D RATI EFF
Colorado S tate Unive rsity
Department of En tomology
Fort Collins, Co l orado 80523
R ALPH F. KIRCHNER
5960 East Pea Ridge
Ridgeview Apartment 1
Hun tington, West Virginia 25705
AND
DA VID R. LENAT
North Ca ro lina Environmental Management
Water Quality Sec tion
4401 Reedy Cr eek Road
Raleigh, North Carolina 27607
Abstract-The s to neflies (Piecoptera) of North Car o lina and South
Carol ina are comprehensively reviewed fo r the firs t time. One hundred
and thirteen an d 83 stonefly species are recorded from North Carolina
and South Carolina, respectively. Thirteen n ew s ta te records are
g i ven fo r North Carolina and two for Sou th Carolina. An a dditional
22 speci es are listed that may be eve ntu ally coll ected in ei ther sta te.
U nzicker and McCaskill (19 82) presented the fi rs t compr ehensive
checklist of 131 stoneflies known or like ly to occur in North Carolina
and Sou th Carolina. However, as Lenat and Penrose (1987) pointed
out, thi s list di d not distinguish between North Carolina and South
Caro lina, and validation of individual state records requires examin a tio n
of the l iterature. Stark et al. (1986) and S t ewart and Stark (1988)
have provided recent compilations of s tonefly s pecies records for North
America, listing 75 and 77 s p ecies for North Carolina and 77 and
79 for South Caroli n a , respectively. However, all three lists contain
omissions or list sp ecies identified in e r ror (Table 1). For example,
Stark e t al. (1986) did not lis t Taeniopteryx burksi Ricker and Ross,
T. lonicera Ricker and Ross, and T. mete qui Ricker a n d Ross from
No rth Carolina despite the record s published by Ricker and Ross (1968)
or by Fulli ngto n and Stewart (1980). Notations in Table 1 are included
to help clarify taxonomic changes and to distinguish between th e lists.
Brimlcyana 23:25-10. 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: (I) 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 neares t records for Alloperla imbecilla (Say)
are from northwestern Virginia and West Virginia (Baumann 1974,
Surdick 1985); (4) Jsoperla nana (Walsh)- a small black Isoperla is
considered a northeastern and central species (Stark et al. 1986); and
(5) Jsogenoides 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 a lso noted that have been reported since Stewart
and Stark (1988) for North Carolina:
Leuctra f erruginea (Walker) (Huryn and Wallace, 1987).
Megaleuctra williamsae Hanson - Macon Co. , trib. C ullasaja 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 K ondra tieff 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. , Len at and Penrose, 1987 ).
Oemopteryx contorta (Needham and Claassen) - Moore Co., Suck Cr.,
Feb. , 1984, NCDEM.
Strophopt eryx 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).
!soper/a 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 Illies (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).
I. 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:
!soper/a 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 !soper/a 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 stonef!ies 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
Jumosa Ross, Megaleuctra williamsae, Strophopteryx inaya Ricker and
Ross, Sweltsa urticae (Ricker), TaUaperla 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, /soper/a bellona Banks, /. di stincta Nelson, and
Ocon operla innubila (Needham and Claassen) occur in one or both
s tates and are rare and v ulnerable species, sensitive to human induced
impacts. As Baumann (1979) clearl y indicated, stonefl ies are good
indicators of ecosystem quality at all scales. All t he s pecies 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, ru ral developments , and timber harvest). The very
diverse stonefly fauna of both states is indicative of a wide range
of hig h quality lotic aquatic habitats, which need active protection.
Table 1. A comparison o f three stonefly (Piecop te ra) species li sts for North Carol ina
(NC) and So uth Carolina (SC).
Unzicke r St ark Stewart
and M cCaskill e t al. and Stark
( 198 2) ( 1986) ( 1988)
Euholognatha
Capniidae
Allocapnia aurora Ricker X NC,SC NC,SC
A . brooks i Ross X'
A. fumosa Ross X NC NC
A. granulata (Claasse n) X
A. lo slwda Ricker X'
A. mystica Frison X
A . nivicola (Fitc h) X NC NC
A . pygma ea (Burm e is ter) X
A. recta (Claassen) X NC,SC NC,SC
A. rickeri Frison X NC N C
A . stannardi Ross X NC NC
A . virginiana Frison X NC,S C NC,SC
A. wroyi Ross X NC,SC NC,SC
Ne mocapnia carolina Banks X NC,SC NC,SC
Paracapnia ang ulata Hanson X NC NC
P. opis (Newman) X
Leuctridae
Lcuctra alexanderi Hanson X NC,SC NC,SC
L . hiloba C laasse n X NC NC
L. caroline11SiS C laasse n X N C,SC N C,SC
L. ferruginea (Walker) X sc sc
L. g randis Banks X NC NC
L . maria Hanso n X
Stonefly Records
Table I. Continued.
L. mirche/lensis Hanson
L. moha Ricker
L. monricola Hanson
L. nephophila Hanson
L. sibleyi Claassen
L. rene/la Provancher
L. re nuis (Pictet)
L. rriloba Claassen
L. variabilis Hanson
Megaleucrra williamsae Hanson
Paraleucrra sara (Claassen)
Nemouridae
Amphinemura delosa (Ricker)
A. nigrirra (Provancher)
A. wui (Claassen)
Paranemoura perfecra (Walker)
Prosroia complera (Walker)
P. similis (Hagen)
Shipsa ro tunda (Claassen )
Soyedina carolinensis (C laassen)
Zapada chi/a (Ricker)
Taeniopterygidae
Bolroperla rossi (Fri son )
Oemopreryx conrorra
(Needham and C laassen)
Srrophopreryx
appalachia Ricker and Ross
S. fasciara (Burmeister)
S . inaya Ricker and Ross
S. /imara (Friso n)
Taen i onema
arlanricum Ricker and Ross
Un zicker
and McCaskill
( 1982)
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X'
X
X
X'
X
X
X'
X
Taeniopreryx burksi Ricker and Ross X
T. lira Fr ison X
T. l onicera Ricke r and Ross
T. maura (Pictet)
T. merequi Ricker and Ross
X
X
X
Stark
et al.
( 1986)
NC
sc
NC,SC
NC
NC,SC
sc
NC,SC
sc
NC,SC
sc
NC,SC
NC
NC,SC
sc
sc
NC
NC
NC
NC,SC
NC,SC
NC
())
NC,SC
()) sc
NC,SC
Stewart
11nd Stark
( 1988)
NC
sc
NC,SC
NC
NC,SC
sc
NC,SC
sc
NC,SC
sc
NC,SC
NC
NC,SC
sc
NC2 ,SC
NC
NC
NC
NC,SC
NC,SC
NC
NC 1
NC,SC
()) sc
NC ,S C
29
30 Boris C. Kondratieff, Ralph F. Ki rchner, and David R. Lena!
Table 1. Continued.
T. parvula Banks
T. robinae Kondratieff
and Kirchner
T. ugola Ricker and Ros s
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 lerna (Frison)
Suwallia marginata (Banks)
Sweltsa latera/is (Banks)
S. mediana (Banks 191 1)
S. onkos (Ricker)
S. urticae (Ricker 1952)
Unzicker
and McCaskill
( 1982)
X
X'
X
X
(S)
X'
x•
X
X'
X'
(10)
X
X
X
Utaperla sp . X 12
Peltoperlidae
Peltoperla ada
Needham and Smi th X 1•13
Peltoperla arcuata Needham X'
Tallaperla anna
(Needham and Smith)
T. corn elia (Need ha m and S mi th)
T. elisa S tark
T. laurie (Ricker)
T. maria (Needham and Smith)
Vielwperla ziplw (Frison)
Vielzoperla ada
(Needham and Smi th)
Perlidae
Acron euria abnormis (Newman)
A. arenosa (Pictet)
A. arida (Hagen)
A . carolinensis (Banks)
X
X
X
X
Stark
et a!.
(1986)
PI SC
sc
NC,SC
(4 )
NC,SC
NC
171 sc
NC,SC
NC,SC
NC
NC"
NC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
sc
NC
NC,SC
Stewart
and Star k
(1988)
m SC
sc
NC,SC
NC,SC
NC
sc
NC,SC
NC,SC
NC
NC"
NC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
sc
NC
NC,SC
Stonefly Records 3 1
Table 1. Continued.
Unzicker Stark Stewart
and McCaskill et al. and Stark
(1982) ( 1986) (1988)
A. evoluta Klapalek X'·"
A. filicis Frison X sc sc
A. intemara (Walker) X'
A . lycorias (Newman) X'
A . me/a Frison X' 5
A . perplexa Frison X'
A. petersi Stark and Gaufin X'
Agnetina annulipes (Hagen) sc
A . capita/a (Pictet) NC,SC16
A. flavescens (Walsh) sc
Attaneuria ruralis (Hagen 1861) X NC,SC NC,SC
B eloneuria georgiana (Banks) X NC NC
B. stewarti Stark and Szczytko X NC,SC NC,SC
Eccoptura xanthenes (Newman) X NC,SC NC,SC
Hansonoperla appalachia Nelson X' sc
Neoperla carlsoni Star k
and Baumann X sc sc
N. clymene (Newman) X
N. freytagi Stark and Baumann X sc
N. occip italis (Pictet)17 sc
Paragnetina fumosa (Banks) X NC,SC NC,SC
P. ichusa Stark and Szczytko NC,SC NC,S C
P. immarginata (Say) X NC,SC NC,SC
P. kansensis (Banks) X sc sc
P. media (Walker) X
Perlesta frisoni Banks X NC,SC NC,SC
P . placida (Hagen) X sc
P . placida (Hagen) complex'" NC,SC
P erlinella drymo (Newman) X sc NC,SC
P . ephyre (Newman) X sc sc
P . fumipennis (Walsh)'9 X sc
P. zwicki Kondratieff,
Kirchner and Stewart sc
Ph asganophora capita/a (Pictet) x2o
Pcrlodidae
Clioperla clio (Newman) (~I) NC,SC NC,SC
Cultus decisus (Wa lk e r Xl~ NC NC
Diploperla duplicata (Banks) X sc sc
32 Boris C. Kondra tieff, Ralph F. Kirchner, and David R. Lenat
Tabl e I . Contin ued.
Unzicker
a n d McCaskill
( J 982)
Hel opicus bogaloosa S tark and Ray
H. s ubt ·arian s (Bank s) X
Hy dr oper/a fugitan s
(Needham a nd Claassen) x ~·'
l sogenoides doratus (Fri so n) X'
I. han so ni (Ricker) X
1. varians (Walsh) X
! sop er/a bellona Banks X
! . bilineata (Say) X
! . clio (Newman) X"
I . cotta Ri cker X
I. dicala F rison X
I. d is tincta Nelson X '
I. h olochlora (Klapalek) X
I. l ata Frison X'
I . marlynia Needham
and Claassen X
I . nana (Walsh) X
I . orata Friso n X
!. simi/is (Hagen) X
Malirekus hastatus (Banks) X
Ocon operla innubila (Needham
and C laassen)"
Remenus bilobatus (Need ham
and C laassen) X
Yugus arinus (Frison) X
Y. bulbosus (Frison) X
Y. innubilu s (Nee dham
and Claassen) X 24
Ptcronarcyi dac
Pteronarcys biloba (Newman)
P. d orsata (Say)
P. proteus (Newman)
P. SCO tti (R icker)
x~'
X
X
X
Stark
et al.
(1986)
s c
NC ,S C
NC
sc
N C
N C
sc
sc
NC,SC
NC,SC
sc
NC,SC
NC,SC
NC,SC
NC,SC
NC,S C
NC,SC
NC,SC
N C,SC
sc
NC,SC
NC,SC
Stewart
and Stark
( l 988)
sc
NC,SC
NC
sc
NC
NC
sc
sc
NC ,SC
NC,SC
sc
N C,SC
NC,SC
NC,S C
NC,SC
NC,SC
NC,SC
NC,SC
NC,SC
sc
NC,SC
NC,SC
' Li sted by U n zic k er and McCaski ll (1982) " as likely to occur in North or South Caro lina,
but presence bas no t yet be en confirmed."
0 Listed onl y b y S tewart and Stark (1988) in the '"Speci es of Nymphs Examined " and
a s pecimen is i llu st rated from Davie Co unty, Nort h Ca ro lina , Yadkin Rh er.
Stonefl y R eco rds 33
Table I. Continued (Foot notes).
Rick e r and Ross ( 1968) li sted records for th ese s pecies. Additi ona ll y, Stewart and
Star k (1988) listed Taeniopteryx burksi in th e "Species of nymphs exami ned ," b ut
not in the ··Nort h Ame rican Species Li s t and Dist ribution. "
• Su rdi ck ( 1985) lis ted Buncombe Coun ty and Great Smoky Mountain National Park,
Nor t h Carol in a.
~ Surdick (1 985) listed Bunc ombe Cou nty, No rth Carolin a.
1' L is ted by Unz icker and McCask ill (1982) as S weltsa nanina .
7 Surdick (1985) listed Bunco mb e County, Mc Dowell Co unt y, and Ya ncey Count y, North
Carolina.
~ Li sted b y Unzicker and McCask ill (1 982) as Ha st aper/a ilre\'is , Zw ic k { 1977) dis­c
usses the gener ic sy nonym y.
• Su rdick ( 1985) li ste d th is species from Great Smoky Mountain Nation al Pa rk, North
Carol ina .
'" Surdick ( 1985) li s ted Yancey County and Grea t S moky Mountain Park. North Caro­li
na.
11 Surdick (1985) d id not li st Swelrsa onkos from North Carolina, presu mably older records
are mis ident ificat ions of S. mediana.
'~ This generic record is based o n n ym phs, presumab ly misidentificat io ns of nymphs
of Alloperla usa? Uraperla gaspes iana Harper and Roy is known fr om We s t Virgin ia.
'-'Thi s species is now included i n the g~:n us Vielloperla, an d V. zipha is considered a
syno nym of V. ada (Stark and Stewart 1981 ).
"This s pecies is now included in the genus Tallaperla (Stark and Stewart 1981).
"Stark and Brown ( 1991) stud ied the hol oty pc of Acroneuria evoluta and co nsid ered
A. m e/a a sy nony m.
"'The record s for this s pecie s in Stark et al. (1986) are com posit e, including bot h Agnetina
ann u /ipes and A. flnvcscens, which were di st i nguished b y S tark ( 1986) . The !alter
re ference did no t list a ny record s for A . capita ta fro m North Ca ro l ina.
" Stark (1 990) sy non ym ized Neoper/a freytag i \\ ith N. occipitalis ( Pi cte t )
' "Stark (1989) divided th e Perles ta placida complex into 12 sp ec ies. Stark ( 1989) provided
records for t hree spec ies, P. placida. Nort h Carol ina ; P. friso ni, Haywood Co unt y,
North Ca rol i na ; Ocon ee Co u nty, Pickens County, Sout h Carolin a ; P. nelsoni , Haywood
County , Swai n County, Nort h Carolina; Oconee Co un ty , Sout h Caro l ina.
•• Ko nd rat ieff et al. (1988) synonymized Perline/la f umipr:nnis wit h P. ephyre and de ­scribed
P. zwicki for the species formerly identi fi ed as P. fumipennis from the Southeast.
~0 Zw ick ( 1984) established the synonymy be tween Agnetina and Pha sganophora . Sec
note 16.
~ · Szczytko and Stewart ( 198 1) included ! soper/a clio in Cliopula .
" Stark ct at. ( !988) re view ed this speci es complex and recognized two species. Cull/Is
decisu s with two subspecies and C. verti calis (Banks). C. d. isola tu s (Ba uk s) is known
from Madi so n Count y, North Caro lina and C. wmica lis is know n from Haywood Co un ty
and Swain Co unt y, Nor th Ca ro li na.
" Kondratieff and Painter (1986) indi ca ted t ha t records o f Hydroper/a fugiratts fro m
North Caro lin a or So ut h Caro lina were in er r or, and t hat t he on l y confirmed record
of this ge nu s was from Sout h Caro lin a refe rable to H. p hormidia Ray and Stark.
!<Stark ( 198 5) synon ymized Yu g us innu bilus with Oconoperla we averi S tark and Stewarl.
~~ St ark and Szczy t ko ( 1982) recognized Allonarcys as a sy nonym o f Pt er onarcys.
34 Boris C. Kondratieff, Ralph F. Kirchn er, and David R. Lenat
Table 2. List of s tone fli es (Plecoptera) recorded from North Carolina (NC) a nd South
Ca rolina (SC). Species marked with a n " " occur in surrou nding states and could eventually
be collected in ei ther state. New state records for bot h states as indicated in text are
marked b y a #.
Euholognatha
Capniidae
Allocapnia aurora Ri cker
A. brooksi Ross (TN)
A. fumosa Ross
• A. granulata (Claasse n) (TN, VA)
• A. loshada Ri c ker (TN, VA)
A. mystica Fri son (TN, VA)
A. nivicola (Fitch)
• A. p y gmaea ( Burmeister) (TN , VA)
A. recta (Claassen)
A . rickeri Frison
* A. simmonsi Kondrat ieff and Voshell (VA)
A. stannardi Ross
A. virginiana Frison
A. wrayi Ross
Nemocapnia carolina Banks
Paracapnia angulata Hanson
Leuctridae
Leuctra alexanderi Hanson
L. biloba Claassen
L. ca rolinensis Claassen
L. ferruginea (Walker)
L. grandis Banks
L. mitchellensis Hanson
L. moha Ricker
L . monticola Hanson
L. nephophila Hanson
L. sibleyi Claassen
L. tenuis ( Pi ctet)
L . triloba Claassen
• L. variabilis Hanson (VA)
M egale uctra williamsae H anson
Para/euctra sara (Claassen)
Ncmouri dae
Amphinemura delosa (R icker) ( TN, VA)
A. nigritta (Provancher)
A. wui (Claassen)
NC
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X'
X
x•
X
sc
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Stonefly Records
Table 2. Continued.
• Ostrocerca albidipennis (Walker) (VA)
0. complexa (Claassen) (VA)
• 0. prolongata (Claassen) (VA)
• 0. truncara (Claassen) (VA)
Paranemoura perfecto (Walker)
Prostoia completa (Walker)
P. simi/is (Hagen)
P. hallasi Kondratieff and Kirchner
Shipsa rownda (CI~ assen)
Soyedina carolinensis (Claassen)
Zapada clzi/a (Ricker)
Taeniopterygidae
Boltoperla ros si (Frison)
Oemopreryx contort(! (Needham and Claassen)
Strophopreryx appalachia Ricker and Ross
S. fasciata (Burmeister)
S. inaya Ricker and Ross
S. limata (Frison)
Taenionema arlanticum Ricker and Ross
Taeniopteryx burksi Ricker and Ross
T. lira 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 atlamica Baumann
A. caudata Frison
A. chloris Frison
A. furcula Surdick
A. nanina (Banks)
A. neglecta Frison
A. usa Ricker
Haploperla brevis (Banks)
NC
X
X
x•
X
X
X
X
x•
X
X
X
x•
X
X
X
X
X
X
X
X
X
X
X
X
X
X
sc
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
35
36 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat
Table 2. Cont inued.
Rasvena tema (Frison)
Swvallia marginata (Banks)
Sweltsa lat era /is (Banks)
S. m ediana (Banks)
S. urticae (Ricker)
Peltopcrlidae
• Peltoperla arcuata Needham (TN, VA)
P. tarteri Stark and Kondratieff (VA)
Tallaperla anna (Needham and Smith)
T. comelia (Needham and Smith)
T. elisa Stark
T. laurie Ricker
T. maria Needham and Smith
Vie/roper/a ada (Needham and Smith)
Perlidac
Acroneuria abnormis (Newman)
A. are nosa (Pict e t)
A. arida (Hagen)
A. carolinensis (Banks)
A. el·o luta Klapalck
A. filicis Frison
A. frisoni Stark and Brown
*A. internata (Walker) (GA, VA)
A. /ycorias (Newman)
• A. petersi Stark a nd Gaufin (GA, TN)
Agnetina ammlipes (Hagen)
• A. capita/a (Pictct) (VA)
A. flavescens (Walsh)
Allaneuria ruralis ( Hagen)
Beloneuria georgiana (Ba n ks)
B. stewarti Star k und Szczytko
E c coptura xanthenes (Newman)
Hansonoperla appalachia Nelson
Neoperla carlsoni Stark and Baumann
N. c atharae Stark and Baumann (TN, VA)
N. clymene (Newman)
N. occipitalis (Pictct)
N. .1/ewarti Stark and B aumann (TN, VA)
NC SC
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x•
x•
x•
X
X
X
X
x•
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Table 2 . Continued.
Paragnetina fumosa (Banks)
P . ichusa Stark a nd Szczytko
P. immarginata (Say)
P. kan sensis (Banks)
P. media (Walker) (VA)
Perlesta frisoni Banks
P. ne lsoni Stark
P. placida (Hagen)
P erlinel/a drymo (Newman)
P. ephyre (Newman)
Stonefly Records
P. ZIVick i Kondratieff, Kirchner and Stewart
Pcr!od idae
Clioperla clio (Newma n )
Cultus d ecisus isolatus (Banks)
C. 1-erticalis ( Banks)
Diploper/a duplicata (Banks)
D . morgani Kondratieff and Voshell
Helopicus bogaloosa Stark and Ray
H. s rtbvarians (Banks)
Hydroperla phormidia Ray and Stark
Jsogenoid es hansoni (Ricker)
/. va ri ans (Wa lsh)
/ soper/a bellona Banks
I . bilineata (Say)
I . burksi Frison
I . colla Ricker
/. da visi James
I . dic a/a Frison
I . distincta Nelson
I . frisoni lilies
I . holoch/ora (Kiapalek)
I . lata Frison
I . marlynia Needham and Claassen
I . namata Frison
I . orata Frison
I. simi/is (Hagen)
I. slossona e (Banks)
I. transmarina (Newm an)
Malirekus lrastatu s (Banks)
Oconopcr/a innrtbila (Needham and Claassen)
NC
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x•
x·
X
X
X
x•
X
X
X
X
x•
X
X
sc
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x•
X
X
X
X
X
X
X
X
X
37
38 Boris C. Kondratieff, Ralph F. Kirchner, and David R. Lenat
Table 2. Continued.
NC sc
Remenus bilobar us (Needham and Claassen) X X
Yugus arinus (Frison) X X
Y. bu/bosus (Frison) X X
Pteronarcyidae
Pteronarcys biloba (Newman) X X
P. dorsara (Say) x• X
P. proteus (Newman X X
P. SCOtti (Ricker) X X
ACKNOWLEDGMENTS-We 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.
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Baumann, R. W. 1979. Nearctic stonefly genera as indicators of ecologi­cal
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Fullington, K. E., and K. W. Stewart. 1980. Nymphs of the stonefly ge­nus
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Stonefly Records 39
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(Plecoptera: Perlidae)? Aquatic Insects 13: 29- 32.
Stark, B. P., a n d K. W. Stewart. 1981. The Nearctic gen era of Peltoperlidae
(Plecoptera). Journal of the Kan sas E nto mological Society 54:285-311.
Stark, B. P., and S. W. Szczytk o. 1982. Egg morpho logy and phylogeny
in Pteronarcyidae (Plecoptera). Annals of th e Ento mo logical Society
of America 75:51 9-529.
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sto neflies (P lecoptera): Systematics, distribution, a nd taxonomic
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complex of eastern North America (Pl eco ptera: Perlodidae). Proceed­ings
of the Entomological Society of Washington 90:91-96.
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Received 9 February 1995
Accepted 20 April 1995
Seasonality in Cetacean Strandings
Along the Coast of North Carolina
WM. DAVID WEBSTER, P. DAWN GOLEY, JESSIE P USTIS,
AND JOSEPH F. GOUVE IA
Department of Biological Sciences
and Center for Marine Science Research
U11iversity of North Carolina at Wilmington
Wilmington, North Carolina 28403
ABSTRACT - Records of strand ing provide an index by which the
resident s tatus and loca l migratory patterns of cet aceans can be ascer­tained,
especia lly along North Carolina's l engthy 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
in tentionally 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 a long
the North Caro lina coast. Although stranding records are available
for every month, the bottlenose dolphin (Tursiops tumcatus) 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 a nd provinces in eastern North America because of the
state's extreme physiographic variability (Webster et al. 1985), and
marine mammals clearly exemplify this tre nd . Many species of marine
mammals are year-round residents, b ut others with s ubtropical and
subarctic affinities, s uch 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 c losely related
taxa that ostensibl y 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.
Brimle yana 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 fa una (whales, dolphins, and porpoises) of North Carolina
is relatively well known (Caldwell and Golley 1965, Caldwell and
Ca ldwell 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 th ese 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 fro m 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
Jive and dead animals, so both were included in our analysis. It was
not possible to verify identifications of all specimens associated with
th ese 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 some­times
not collected (Mead 1977, 1979; Schmidly 1981). Therefore,
records we re omitted if doubts existed about their veracity. Temporal
data were examined sta tistically (Chi-square) to test the hypothesis
that each species exhibited no significant (P < 0.05) monthly variation
in stranding, a lth ough sample sizes were small f or some s pecies.
RESULTS AND DISCUSSION
E ight 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 stra nding records.
Although the bottl e no se dolphin (Tursiops truncatus (Montagu)) strands
in all months of the year, it s trands significantly more often in winter
and s pring. Stranding records fo r 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 (Stene/la frontalis G. Cuvier), rough-toothed
dolphin (Steno bredanensis (Lesson)), and dwarf sperm whale (Kogia
simus (Owen)) display significiant monthly variation in stranding with­out
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 a!. 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, circum­stantial
evidence suggests that mortality may be greater for some species
during the winter and spring months. Finally, certain species of ceta­ceans
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 August­October
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.
1).
.j:>.
.j:>.
Table 1. Mont hl y frequencies of cetacean strandings a long the coast of North Carolina. Significant ( P < 0.05)
monthly var iation is shown with an asterisk for spec ies with 10 or mo re strands.
Fami ly Month
Species J F M A M J J A s 0 N D Total
~
Balaenidae 3
Eubalaena glacia/is 2 1 4 t1
Ba l aenopteri dae 00
<
Balaenoptera acutorostra ta 2 2 Q..
Ba/aenoptera borealis 1 ~
Balaenoplera physalus 5 3 2 2 14 (1>
0"
Megaptera novaeangliae 2 2 3 9 "' (';
Delphinidae ...,
D elphinus d elphis 3 2 2 8 ~
Feresa attenuata I I 00 :-
Globicephala macrorhynchus 2 1 5 4 2 2 J H
Globicephala me/as 4 4 L 10
Grampus griseus 6 J 4 5 2 21
Lagenorhynchus acutus 2 2
Orcinus orca
Table I . Continued.
Family Month
Species J F M A M J 1 A s 0 N D Total
Pseudorca crassidens 1 I
Stene/la coerufeoafba 4 1 3 1 3 I 3 2 2 20
Stene/la fron talis 3 7 1 4 I 2 4 3 25*
Stene/la longirostris 1 I 2 (")
Steno bredanensis 1 1 12 1 15 (1)
PJ
Tursiops truncatus 46 40 77 80 79 28 8 24 36 52 35 26 531 ('l
(1)
Phocoen idae "' ::l
Phocoena phocoena 2 17 37 14 7 77 (/)
Physeteridae
...,
"' Kogia breviceps 3 7 3 4 7 4 8 5 2 l 4 48 ::l a.
Kogia simus 1 3 2 I 1 4 12 ::l
(JQ
Physeter macrocephalus I 1 4 3 1 I 3 I I 16 Vl
Ziphiidae
M esopfodon densirostris 2 2 2 1 2 9
Mesoplodon europaeus 3 1 2 3 4 2 1 1 2 19
Mesoplodon mirus 1 l 2
Ziphius cavirostris 2 1 I 4
Total 77 83 154 130 117 35 21 49 53 62 46 45 872
.,.
V'1
46 Wm. David Webster et a l.
100 Tursiops truncatus
80 C3 AU. YEARS COMBINED
Cll
0 AU. YEARS EXCEPT 1987
..J
c(
:;)
c 60 >
0
3;
1!.
0 40
a:
UJ
aJ
::;::
:;) z 20
F M A M A s 0 N 0
MONTH
Fig. 1. Monthly s tranding records for the bottlenose d olphin (Tursiops truncatus)
in North Car olina. Significant increases in s tra nding during the dolphin di e­off
of 1987 are evident in August, September, and October ( " ).
50
en
..J
40 c(
:J
0 > 0 30
z
u..
0 20
a:
w
m
::E 10
:J z
0
J
FALL/WINTER/SPRING MIGRANTS
F M A
~ Eubalaena glacialis
13 Balaenoptera acutorostrata
Uil Ba/aenoptera borealis
0 Ba/aenoptera physa/us
l2l Megaptera novaeang/iae
• Phocoena phocoena
~ Lagenorhynchus acutus
M J J A
MONTH
s 0 N D
F i g . 2. M o nthly stranding records fo r ce tacean s with boreal distributions
(see text ) that migrate or wa nder southward a long the North Car o lina coast
during th e co lder months of the year.
Cetacean Strandings 47
Stranding records for several migratory species s uch as the northern
right whale, fin whale, humpback w h ale, and harbor porpoise indicate
when these species a re 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 w hale and hump­back
whale. Right w hale stran dings are confined to the spring months,
the season when mothers and their n ewborn ca l ves migrate northward
in· shallow nearshore water (Kraus et al. 1986, 1993); the sou thward
winter migr ation (Reeves et al. 1978) of juveniles, subadults, and
adults is farther offshore a long the eastern edge of the Labrador Current
(Kraus et al. 1993) and the continental shelf (Winn e t al. 1986). Stranding
records for the harbor porpoise demonstrate a local north-south migra­tion,
with the southernmost distributional limits reaching Nort h Carolina
during the winter and spring months, rather than the insho re-offshore
migratory pattern seen farther no rth (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 (De lphinus de /phis Linnaeus), striped dolphin
(Stenella coeruleoabla (Meyen)), sperm whale (Physet er macrocephalu s
Li nnaeus), Blainville's beaked whale (Mesoplodon densirostris (Blainville)),
and Gervais ' beaked whale (M. europaeus (Gervais)) have stra n ded
during most months of the year, and we suspect t hat they are year­rou
nd residents in North Carolina waters. The common dolphin inhabits
temperate waters adjacent to the 100-fath om isobath where the ocean
floor has substantial topographic relief (Leatherwood and Reeves 1983).
It does n ot exhibit a pronounce d nort h-south migration, so increased
s trandings during the colder months may reflect seaso nal inshore-offshore
move-ments (Selzer and Payne 1988) or increased mortality during
the colder months of the year. Although the sperm whale has a well­documented
migration in t he North Atlantic Ocean (Townsend 1935),
it has stranded in No rth Carolina in all months except June, October ,
a nd December. Mature males migrate northward out of North Carolina
waters in the spring, but so me immature males a nd females and their
calves remain in North Carolina waters throughout the summer months
(Leatherwood et al. 1976). These five species constitut e approximately
8~ of the tota l numbe r of s trandings reported from North Carolina .
Several ce taceans exhibited significant monthly variation in stranding
bu t demonstrated no seasonal pe ri odicity, and each appears to inhabit
48 Wm. David Webster et al.
North Ca r olina wa t ers t hroughout the year. Monthly va n at10n can be
explaine d by th e tendency to mass strand by pilot wha les, the rough­toothed
dolphin , and possibly Risso ' s dolphin. It is difficult to explain
significan t monthl y variation exhibited by the Atla n tic spotted dophlin
(Stenella frontalis (G. Cuvier)) , which includes stranding records previously
a ttri butable to S. plagiodon (Cope), a species o nce th ought to inhabit
the A tlantic Ocean. Taxonomic uncertainty in the genus an d the difficulty
in identifying individuals have b een presistent sour ces of e rro r; however,
the recent revision of Stenella in the western North Atlantic Ocean
(Perrin et a!. 1987) s h ou ld help alleviate future misidentifications.
Significant monthly va ri a tion in str anding by Kogia simus might best
be explained by a behavior displayed by i ts c lose relative, t he pygmy
sperm whale (Kogia breviceps (Blainville)). In so utheast ern North Carolina,
we have notice d that strandings of K. breviceps freque ntly involve
females in th e p rocess of givin g birth or mother -offspring pairs, a
behav i or also reported by Winn e t al. (1979). T h ese seven species
account for approximatel y 17% of t h e to tal number of s tra nd i ngs reported
from North Carolin a.
T he remaining seven species of cetaceans are re latively ra r e in
North Carolin a waters, and scant y stranding records provide little informa­t
ion about th eir resident status in the state. The killer whale (Orcinus
orca (Linnaeus)), false killer whale (Pseudorca cr assidens (Owens)),
True's beaked whale (Mesoplodon mirus T ru e), and Cuvier 's beaked
whal e (Ziphius cavirostris G. Cuvier) are thought t o be year-round
residents (Leatherwood and Reeves 1983), but th e Atlantic w hite-sided
dolphin (Lagen orhynchus acutus (Gray)) inhabits the northe rn North
A tl antic Ocean (Leath erwood and Reeves 1983) and seldom ventures
into No rt h Carolina waters. The pygmy killer w hal e (Feresa attenuata
G ray) and long-snouted spinner dolphin (Stene/la longirostris (Gray))
probably enter North Carolina water s during the war mer m onths of
th e year (Leatherwood and Reeves 1983). Coll ect ively , th i s group of
species constit utes onl y about 2% of the total number of s trandings
repo rte d from North Carolin a.
Although ma rine mammal str andings provide a fortuitous source
of information on animals that are not typically access ib le , t h ere are
inhe r e nt biases in conclusions derived from stranding data. Neritic
species s trand more frequently th an pelagic species, so stranding fre­qu
en cies are less likely t o r eflect accuratel y the abu nd ances of pel agic
s pecies. Larger -bodied speci es and m ass strandings are more likely
to be reported th an sma ll - bodied sp ecies or sing le strandings. A lso,
the Gulf Stream and Labrad or Current could transport dead or dying
animals beyond their normal ranges and into North Carolin a waters.
Cetacean Strandings 49
Conclusions from the Nort h Carolina str anding da t a, h oweve r, agree
with information provided by other methods of study fo r species that
are re latively well known.
CONCLUSIONS
North Carolina has t h e greatest dive rsi ty of ce taceans along the
east coast of the Un ited States. Twenty-six species have stranded along
the North Carolina coast; four other species might inhabit state waters
at least seasonally. Based on s tranding records, 1 7 species appear
to be year- r ound residents, although bull sperm whales l eave 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 affin iti es migrate northward into the area
during the s ummer and fall months. The status of several s pecies
needs additional clarification, and as a matter of protocol, voucher
material from deceased marine mammals should a lways 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 a n anony­mous
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 (Con t ri­bution
Number 51) of the University of North Carolina at Wilmington.
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Received 9 February 1995
Accepted 28 August 1995
Fishes New or Rare on the Atlantic Seaboard
of the United States
FRED c. R OHDE
North Carolina Division of Marin e Fis heri es
127 Cardinal Drive Exten s ion
Wilmington, North Carolina 28405
S TEVE W. Ros s
North Carolina Nationa l Estuarine Research Reserve
7205 Wrightsville A venue
Wilmington, North Carolina 28403
SHERYAN P. EPPERLY
National Marine Fisheries Service
Southeast Fisheries Scienc e Center
101 Pivers Island Road
B eaufort, North Caro lina 285I6-9722
A ND
GEORGE H. BURGESS
Florida Museum of Natural Hist ory
University of Florida
Gainesville, Florida 326I I
ABSTRACT-Sampling over the con tinental s helf of the South At­lantic
Bight, especial ly off North Carolina, c ontinues to produce
records of fis hes n e w to or rare in the area. We docume n t the f irs t
records for United States continental s helf water ( <200 m depth)
of seven fi sh species: Cirrhigaleus asper, Symphysanodon berryi,
Pseudocaranx dentex , Lutjanus purpureus, Pristipomo ides freemani,
Poecilopsetla beani, and Lagocepha lus la gocephalus. In addition ,
we also repo rt on n o teworthy collections of o ur other fi s hes cap­tured
off North Carolina : S ynagrops spinos us , Centropris tis fu sc ula ,
Gonioplectrus hispanus, and Etelis ocu/atus.
Coll ectio ns along the Atlantic co ast of th e Carolinas continue
to y ie ld fish s pec ies never recorded from the area. Most of these
new records represen t tropical and subtropical species that extend their
published ranges from the Caribbean or the eastern Atlantic (Anderson
and Guthe rz 1964, Burgess et al. 1979, Bohlke and Ross 1981, Ross
Brim Ievana 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 faunistica lly­rich
region that contains diverse habitats and favorable ocean currents.
During development of the fifth edition of the American F isheries
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 addi tion
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).
SQU ALIDAE
Cirrhigaleus asper (Merrett 1973)
The roughskin dogfish is a widespread continental s lope species
known from the western North Atlantic, sou thwest 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° 14 ' W)
in 201 m, 8 July 1982; UF 44303 (1 , 1000 mm TL) , Atlantic Ocean
off Georgia (30°52 ' 46"N, 79°46 ' 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 Iongline and tr ap fishing has demonstrated
that this fish is widely distributed in waters of 183- 457 m in the
tropical and subtropical wes tern Atlant ic. Generic placement of this
New or Rare Fish es 55
species in Cirrhigaleus, rather than Squalus, fo ll ows Shirai (1992)
and G. H. Bu rgess (Florid a Museum of Natural History, unpublished
data).
ACROPOMATIDAE
Symphysanodon berryi A nder son 1970
The fi rs t United Sta tes rec ord of the s lope 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 Septemb er 1982.
A nderson (1970) repo rted the species from throug hout the Ca r ibbea n
a nd the Bahamas in 220-476 m. Alt hough Symphysanodon is usu ally
lis ted as a lu tjanid, there is evidence against this placeme n t. It was
considered incertae sedis but possibly related to the Acropomatidedae
(especially Synagrops) by Johnson (1984), and its taxonomic stat us
is still uncerta in (Johnso n 1993). Although Eschmeyer (1990) considered
i t to be in its own family, Symp hysanodon tidae, 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 (Roese an d Moore 1977, Potts and Ra msey
1987, Mochizuki and Gultneth 1989, Boschung 1992), and is also
known from off Suriname and ot her Car ibbea n locations (Fujii 1983,
Mochizuki and Gultneth 1989), an d from scattered locations in th e
West Atlantic and West Pacific (Mochizuki and Gultne th 1989). Records
of this fish, however, a re rare alo ng t he United S tates Atlantic east
coast. The following trawl collections document the occurrence of
S. spinosus on the other continental shelf (s200 m) of the South Atlantic
Bight: UF 41747 (1 , 68 m m SL), 35 °47 ' N, 74°53'W in 78 m, 14
July 1980; GMBL-74-92 (1 , 61 mm SL) , 35° 10 ' N, 75°03'W in 221-
229 m, 8 May 1974; GMBL-74-58 (3), 35°02'N, 75°ll ' W 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°41 ' N, 75°37'W (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, s ta t ion 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 1 984; UF 4 1229
(4, 96, 102, 104, 108 mm SL), 28° 40.6 ' N , 79°53.8'W in 320 m , 31
56 Fred C. Rohde et a!.
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 sta ti on
1283, Bullis a nd Thompson (1965) listed eight other coll ections (not
cited previously) of thi s species between South Carolina and Cape
Canaveral , F lor id a (82-366 m). Wenner et al. (1979b, c; 1980) also
listed several collections of S. spinosus from northern Florida to Sout h
Ca r o lina (128-338 m). The keelcheek b ass see ms to be common on
the outer continental shelf and upp er slope from North Carolina through
the G ul f 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 th e pelvic spines and the secon d spin es of the first dorsal
and a nal fins of S. spinosus. The gen us Synagrops has been variously
placed in the families Apogonidae (Cheilod i pteridae) and Percichthyidae
(Fraser 1972), but is currently placed in the " oceanic percichthyids,"
family Acropomatidae (Johnson 1984).
SERRANIDAE
Cent r opristis fuscula (Poey 1861)
Four individuals (UF 44997, 50 mm SL, 68 mm SL, 107 mm
SL g ravid fem ale, 1 17 mm SL gravid female) of the rare twospot
sea bass were taken in a single trawl catch at 33°16'N, 77°13'W (Delaware
II 82-04 , s tation 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 r eef o rganisms
(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°4TN, 78°11 ' W at a depth of 165 m on 15 July 1995. T hese
five specimens signific antly increase the total k nown specimens and
extend the range northward. Previous records of C. fu scula were from
Cuba (holotype MCZ 10015, 138 mm SL (Poey 18 61 ) ; ANSP 94422,
135 mm (Robin s and Starck 1961)), Puerto Rico in 183 m (ANSP
144592, 155 mm SL), G ul f of Mexico (I specimen, G . D. Johnson,
United States National Museum, personal commun ication) , and Sou th
Carolina (2 collections, listed as Cen tropristis sp. by Wenner et al.
(1979a)) . The genera l rarity of specimens and th e bottom type of
our collection suggest t hat this species i s a c ryptic reef fish.
Goniople ctrus hispanus (Cuvier 1828)
The Spanish flag, usually considered a Caribbean in s ular 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 s in gle, pelagic
larva collected off Cape Fear, North Carolina (Kendall and Fahay
1979). Intensive sampling of the Carolinian snapper/gr ouper 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, I July 1987; specimen lo st (247 mm TL), 33°31.3'N,
76°56.5'W in 40 m, 15 June 1988; s pecimen sold (250 mm TL), southern
Onslow Bay, North Carolina in 46 m, September 1990; UF98891 (gravid
female , 182 mm SL), northern Long Bay, North Ca ro lin a, 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), south ern 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 trevaUy includes
the western Indian Ocean, the Ind o- West Pacific, the Mediterranean
Sea, the eastern Atlantic, mid-Atlantic islands, southern Brazil, and
Bermuda (Sm ith-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, 7rl6 ' W) on 19 February
1985 in 91 m. Two other large specimens (both marketed) were taken
by hook and line: one on 16 Novembe r 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 coll ected by hook and line in 88 m between 30 July
and 5 August 1987 at 33 °16.5'N, 7r15'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 co mmercial fishermen
recogn ized this species as different (referring to it as "gu elly jack")
and, in addition to the above specimens, th ey 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 Ba hama Bank).
LUTJANIDAE
Etelis oculatus (Valenciennes 1828)
A single adult specimen of the queen snapper (UF 42778, 673
mm SL) was collected with hook and li ne 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, 7r55.5'W,
59 m, 3 September 1976). Etelis oculatus ranges in t he 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°l2.8'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 fis h 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 continen tal shelf of North America . The first specimen (GMBL
78-145 , 85.7 mm SL) was collected by trawl off the eas t coast of
Florida at 28°58.4'N, 80°04.4'W 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 Nor t h 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 yellow­eye
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°11 ' W; 24 August 1971 ; W. D. Anderson,
Jr. , personal communication).
PLEURONECT!DAE
Poecilop setta 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 d eep 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 d eep 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 s pecies 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 o n the abo ve re cord (UF 39809), it s h o uld be added t o the
continental shel f fa una.
TETRAODONTIDAE
Lagocephalus la gocephalus (Linnaeus 1758)
T h e first United States oceanic puffers (UF 44194, 169 mm SL;
190 mm SL, specimen mounted) were collected at 34 °2l.5'N, 75°55'W
in 64 m o n 6 Augus t 1985. They were dipnett e d at night from a
school of 6 to 8 individuals swimming a t the surface. Lagocephalus
lagocephalus is widespread, ranging through the eastern Atlantic, the
Mediterranean Sea, and the Pacific and Indian oceans (S hipp 1974).
Templeman ( 1962) reported t he fi rst North American occurrence of
the speci es from a singl e individual collected in Newfoundland. Other
is ol ated wes te rn Atlantic records of the species include Bermuda, the
Gulf Strea m off Florida, and Curacao (Shipp 1974).
DISCUSSION
The offshore (>20 m) ichthyofauna of No rth Carolina, particularly
on ha rd bottoms, is dominated (species numbers) by tropical and subtropical
s pec i es. Expa nded sampling of th e offshore hard bottom and ou ter
she l f habitats co ntinues to increase the numbe r of these forms known
off th e Carolinas. Although t h e Gulf Stream undoubtedly helps dis­perse
tropical organisms into the area, many of th ese so uthern s pecies
apparently maintain self-sust aining populations on North Carolina 's
middle to outer continental shelf (G ri mes et a l. 1977 , Grimes and
Huntsman 1980, S . W. Ro ss, North Caro lina National Estuarin e
Research Reserve, unpubl ish ed data). Burgess et al. (1994), in fac t ,
pro posed that a redefined tropical West Indian zoogeographic province
shou ld include the r eefs of th e outer continental shelf of the South
Atlantic Bight to Cape Hatteras. The 11 s pecies documented h erein
are most common in warm-temperate to t ropical waters along the outer
s helf or upper slop e south of Nort h Caro lina . T hree of these (C. fuscula,
G. hispanus, L. purpureus) tend to be benthic a nd t ied to reef-like
hab i ta ts. The re maind e r are eithe r pelagic or ben th ope lagic, an d ar e
capa ble of extensive movements.
B ri ggs (1974) n oted that , within the warm -temperate Caro linian
Region, the northern G ulf of Mexico contain ed a riche r fis h fauna
(375-400 species) than the Atlantic coast. Since his publication , many
n ew records have been add e d to both areas. Hoese and Moore ( 1977)
re ported ove r 400 fi shes from the northern Gulf of Mexico, and Boschung
(1 992) lis ted around 663 marine fishes from th e eastcentral Gulf o f
Mexico. Dahlberg ( 1975) report ed nearly 400 species in an d near Georgia
New or Rare Fisht:s 61
coastal waters. With the additions reported herein , the North Carolina
m a rine ichthyofauna in less than 200 m contains over 680 species
(S. W. Ross and G. H. Burgess, unpublished manuscript). However,
many of these a re 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 r ichness .
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-Yaniz in confirming the identity
of specimens or providing information on distributions. We thank the
South Carolina Marine Resources Monitoring, Assessment, and Prediction
Program for a llowing us to use their data on the lutjanids and Karsten
E. Hartel for providing data from the Museum of Comparative Zoology.
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