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1975
Contributions to the Biology of Bathynectes superbus (Costa) (Decapoda:Portunidae) from the Chesapeake Bight of the Western North Atlantic.
Elizabeth Gayle Lewis College of William and Mary
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Recommended Citation Lewis, Elizabeth Gayle, "Contributions to the Biology of Bathynectes superbus (Costa) (Decapoda:Portunidae) from the Chesapeake Bight of the Western North Atlantic." (1975). Dissertations, Theses, and Masters Projects. Paper 1593092163. https://dx.doi.org/doi:10.21220/m2-9n8q-sj40
This Thesis is brought to you for free and open access by the Theses, Dissertations, & Master Projects at W&M ScholarWorks. It has been accepted for inclusion in Dissertations, Theses, and Masters Projects by an authorized administrator of W&M ScholarWorks. For more information, please contact [email protected]. CONTRIBUTIONS TO THE BIOLOGY OF Bathynectes superbus (Costa)
(DECAPODA;PORTUNIDAE) FROM THE CHESAPEAKE BIGHT OF
THE WESTERN NORTH ATLANTIC
A Thesis
Presented to
The Faculty of the School of Marine Science
The College of William and Mary in Virginia
In Partial Fulfillment
Of the Requirements for the Degree of
Master of Arts
by
Elizabeth Gayle Lewis
1975 APPROVAL SHEET
This thesis is submitted in partial fulfillment of
the requirements for the degree of
Master of Arts
El yie
Approved, May 1975
A , Musick, (PhiD,
Maurice P. Lynch,
Frank 0, Perkins, Ph,D. TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS...... iv
LIST OF TABLES ...... vi
LIST OF FIGURES ...... ix
ABSTRACT ...... xii
INTRODUCTION ...... 2
SYNONOMY ...... 3
SYSTEMATICS ...... 5
SECTION I. Distribution and abundance of Bathynectes superbus (Costa) ...... 9
SECTION II.Relative growth and morphometries of Bathynectes superbus ...... 39
SECTION III .Reproductive biology of Bathynectes superbus (Costa) ...... 63
SECTION IV.Feeding habits of Bathynectes superbus ( C o s t a ) ...... 112
SECTION V.Epizoites associated with Bathynectes s u p e r b u s ...... 121
SECTION VI.Exoskeletal deformities of Bathynectes s u p e r b u s ...... 129
SECTION Vll.Setal variation in Bathynectes superbus ( C o s t a ) ...... 137
SUMMARY ...... 144
APPENDICES ...... 147
LITERATURE CITED ...... 164
VITA ...... 176
iii 623999 ACKNOWLEDGMENTS
I wish to thank Dr, Paul A, Haefner, Jr, (VIMS) for the collection of specimens from the R/V Columbus Iselin (73-10) cruise (NSF Grant No. GA-37561); Frank Holland (North Carolina
Division of Commercial and Sports Fisheries) from the R/V
Dan Moore (030); and Charles Wenner (VIMS) from the R/V
Albatross IV (74-4) cruise. The R/V Eastward cruise (E-2-74) was supported by an NSF Oceanographic Training Grant to
Duke University. In addition, I thank Drs, Thomas Bowman and Fenner Chace of the U.S. National Museum for providing specimens for morphometric analysis. Dr. Austin Williams of USNM assisted in the loan of specimens for the study of setal variation. Dr. Dorothy Bliss supplied information on
superbus in the collections of the American Museum of
Natural History,
Dr. Dale Calder of the South Carolina Marine Resources
Laboratory identified the hydroids found on B. superbus and
Ms. Mariana Doyle of USNM confirmed identification of
Poecilasma inaeguilaterale.
For technical assistance, I gratefully acknowledge
Sidney Powell of the National Aeronautics and Space Admins istration at Langley Air Force Base for his scanning electron photomicrographs of setae. Dr. Frank Perkins assisted with the interference photomicroscopy. Most of the photographs
iv were taken by Ken Thornberry, Bill Jenkins handled the
development and printing, A1 Zwirko and Jim Amon also helped with the photography, Kay Stubblefield drafted all of the
figures. Histological preparations were largely the work of
Ms. Patricia Berry, who did an admirable and competent job
in spite of the large number of samples. Frank Wojcik and the
data processing staff performed the computer analyses for the morphometries section. I also thank Dr, Gerald Engel for
his help and persistance on the regression analysis program.
Ms, Ann Jamison and Ms. Susan Barrick acquired much of the
literature for the systematics and synonomy section, in spite
of the obscurity of the literature mentioned. Jim Lesofsky
and Bobby Harris helped in maintenance of the live crabs,
I express special appreciation to my major professor,
Dr. Paul A. Haefner, Jr. for introducing me to the topic and
for his helpful suggestions and calmative influence on my
overly zealous nature. I also thank Drs. John Musick, Frank
Perkins, George Grant and Maurice Lynch for reviewing the manuscript. In addition Drs. Austin Williams and John
McDermott (Franklin and Marshall College) made helpful comments
on the study. I would also like to thank Dr. Waldo Schmitt
(USNM), a crustaceologist par excellence, for sharing a few
hours of his time and knowledge with me.
I owe special thanks to my parents for their encourage^ ment in so many ways and to my friends, especially C.W,, J,W, and N^M.W,, who have provided both psychological and academic
support. v LIST OF TABLES
Table Page
1 Catch of Bathynectes superbus and bottom hydrography at CI-73-10 trawl stations in June 1973 ...... 12
2 Intermolt stages of Bathynectes superbus . . 16
3 Sex ratios by size group for Bathynectes superbus captured in the 150-400 m depth stratum during June 1973...... 24
4 Percentage occurrence of brachyuran intermolt stages among four size groups of Bathynectes superbus during June 1973. . . 26
5 Contingency tables used in computing Cole's coefficient of association (C7) for June 1973...... 28
6 Contingency tables used in computing Cole's coefficient of association (C7) for November 1973 ...... 29
7 Catch of Bathynectes superbus and bottom temperature at Alb.IV-74-4 stations in March 1974...... 31
8 Contingency tables used in computing Cole's coefficient of association (C7) for March 1974 32
9 Regression analysis of Bathynectes superbus morphometric d a t a ...... 45
10 Covariance analysis of morphometric data from male and female Bathynectes superbus .... 47
11 Stages of gonadal development of Bathynectes s u p e r b u s ...... 66
12 Number of Bathynectes superbus from November 1973, March 1974, April 1974 in each stage of gonad development for the short carapace widths i n d i c a t e d ...... 85 vi Table Page
13 Relationship of intermolt stage to gonad development in 132 Bathynectes superbus captured in June 1973 ...... 87
14 Relationship of intermolt stage to gonad development in crabs captured during November 1973, March 1974 and April 1974. . . 89
15 Incidence of condition of the spermathecae in relation to size of non-ovigerous female Bathynectes superbus captured in June 1973, March 1974, November 1973 and April 1974 91
16 Incidence of the condition of the sperma thecae relative to gonad development of non-ovigerous female Bathynectes superbus captured in June 1973, March 1974, November 1973 and April 1974 ...... 92
17 Incidence of the condition of the sperma thecae to molt stage of non-ovigerous Bathynectes superbus captured during June 1973, March 1974, November 1973 and April 1974 94
18 Fecundity of Bathynectes superbus ...... 95
19 The incidence of the presence of sperm or spermatophores relative to gross development stage of gonad and size of c r a b ...... 96
20 Frequency of occurrence of stomach contents of hard (C^-C^) , peeler (D^-D^) , soft (A-^) and papershell (A2-B2 ) Bathynectes superbus from June 1973 ...... 114
21 Frequency of occurrence of stomach contents of hard, peeler and paper Bathynectes superbus from November and December 1973. . . 116
22 Frequency of occurrence of stomach contents of hard and papershell Bathynectes superbus from March 1974 ...... 117
23 Frequency of occurrence of stomach contents of hard, peeler and papershell Bathynectes superbus from April 1974 118
24 Incidence and type of fouling organisms observed on Bathynectes superbus ...... 123
yii Table Page
25 Geographic locations at which specimens of Bathynectes superbus with pubescent covering were collected ...... 139
vi ii LIST OF FIGURES
Figure Page
1 Chart of Middle Atlantic Bight showing locations at which Bathynectes superbus were captured in June 1973, November 1973 and March 1974 ...... 10
2 Width frequency distribution of Bathynectes superbus captured in Norfolk Canyon and the adjacent slope during June 1973 20
3 Index of transformed mean abundance of four size groups of male and female Bathynectes superbus captured in Norfolk Canyon and the adjacent slope during June 1973 in 250- 400 m ...... 21
4 Index of transformed mean abundance of four size groups of male and female Bathynectes superbus captured within two degree Celsius temperature strata during June 1973 22
5 Dimensions used in the morphometric analysis of Bathyhebtes superbus...... 43
6 The morphometric relationships of carapace measurements for male and female Bathynectes superbus ...... 49
7 The morphometric relationships of appendage measurements for male and female Bathynectes s u p e r b u s ...... 51
8 Ventral series of male Bathynectes superbus. . . 52
9 Ventral series of female Bathynectes superbus showing abdominal shape change with size and gonad development ...... 53
10 The morphometric relationships of abdominal segment measurements for male and female Bathynectes superbus ...... 54
11 The relationship of penis length to short carapace width for Bathynectes superbus...... 56 ix Dorsal series of female Bathynectes superbus showing fifth anterolateral spine variation with size ...... 57
Male dorsal series of Bathynectes superbus. . . 58
Gross anatomy of the male and female reproductive system of Bathynectes superbus . . 70
Gross reproductive anatomy of ovaries from female Bathynectes superbus ...... 71
Reproductive histology of male Bathynectes superbus ...... 73
Reproductive histology of male Bathynectes superbus ...... 75
Reproductive histology of female Bathynectes s u p e r b u s ...... 77
Ova from female Bathynectes superbus at two stages of gonad development ...... 79
Ventral view of mature and immature male Bathynectes superbus ...... 81
Ventral view of female Bathynectes superbus showing relative change in abdomen shape with size and gonad development ...... 82
Cumulative percentage occurrence of various stages of gonad development in relation to size of male and female Bathynectes superbus captured in June 1973 ...... 83
The indidence of Bathynectes superbus in each stage of gonad development in relation to the mean diameter of spermatophores (from hanging drop preparations) observed at each stage of gonad development ...... 97
The maximum width of the anterior ovarian horns and the incidence of female Bathynectes superbus in each stage of gonad development . . 99
The incidence of Bathynectes superbus in each stage of gonad development in relation to the mean diameter of ova at each stage ...... 100
Size frequency distribution of eggs at various stages of development ...... 102 x Figure Page
27 Malformations of abdominal segments in Bathynectes superbus ...... 131
28 Dorsal view of female Bathynectes superbus . . 132
29 Abnormalities of the frontal region of the carapace of Bathynectes superbus ...... 133
30 Carapace abnormalities of Bathynectes s u p e r b u s ...... 135
31 Scanning electron photomicrographs of selected regions of the carapace of Bathynectes s u p e r b u s ...... 143
xi ABSTRACT
Bathynectes superbus is contagiously distributed on the lower shelf-upper slope areas of the Chesapeake Bight and is consistently found at 122-390 m (5.1-11.3 C) in the Middle Atlantic Bight, Crabs are found deeper (252-390 m) in June, March and April than during November and December (122-223 m). The extension of B, superbus into shoaler or deeper water depends on the hydrographic conditions of the area.
Bathynectes is associated with Cancer borealis, Homarus americanus and Geryon quinquedens. Low association with G. quinquedens and H, americanus in March, November and December may be related to migrations of these crabs or may be an artifact of sampling.
Male crabs ranged from 21-70 mm short carapace width; females from 28-62 mm, Bathynectes superbus molt throughout the year. Male crabs continue to molt at sizes in the 46-57 mm (short carapace width) range while female crabs in this range will not likely molt again. There is some indication that hard and peeler crabs are more abundant in Norfolk Canyon than on the adjacent continental slope.
Morphometric and relative growth analyses reveal simple allometric relationships between short carapace width, the reference measurement, and other carapace dimensions. The relationship between short carapace width and long carapace length is nearly isometric. Male chelipeds attain a larger absolute size than females. No sexual differences in the allometric relationship of chela length and depth were observed. Growth of abdominal segments 3 and 6 demonstrated true sexual dimorphism. A change in growth rate of abdominal segment 6 occursr in females at ^30 mm short carapace width, suggesting a puberty molt associated with the size at which copulation, increased pleopod setation and widening of the vulvae first occur.
The reproductive anatomy and histology of B. superbus is similar to that of other portunids. Synchronous' reproduction is apparently present with some element of the population ovigerous year-round. Maturity within male and female crabs apparently occurs near 28 mm short carapace width based on the presence of sexual products within the reproductive system. Fecundity estimates of number of external eggs ranged from 20 x 103 to 192 x 103 .
xii B. superbus is an opportunistic feederf preying on mollusks, arthropods, echinoderms and polychaetes, Sediment was also ingested.
Epizoites of B. superbus include Poecilasma inaegu ilaterale (Cirripedia: Lepadidae), Anomia aculeata (Pelecypoda) and the hydroids Stegopoma piicatile and Perigonimus sp,
Exoskeletal malformations which were associated with the carapace and abdomen are attributed to injuries sustained by the crab.
Intraspecific setal variation among two allopatric pop ulations of B. superbus is noted. Populations from the lower latitudes of the eastern and western Atlantic are found to be totally pubescent whereas those crabs from the Middle Atlantic Bight are setose along the ventral anterolateral border only.
xiii CONTRIBUTIONS TO THE BIOLOGY OF Bathynectes superbus (Costa)
(DECAPODA:PORTUNIDAE) FROM THE CHESAPEAKE BIGHT OF
THE WESTERN NORTH ATLANTIC INTRODUCTION
Bathynectes superbus(Costa, 1853) is an archibenthic portunid crab that inhabits the shelf edge-upper slope areas of the eastern and western Atlantic. It has a recorded geographic range from Massachusetts to the Florida Straits in the western Atlantic Ocean and from Norway to Angola,
Africa in the eastern Atlantic (Christiansen, 1969). There have been numerous accounts of its occurrence and descriptive characters (Appendix 1), but there have been few contri butions on other aspects of the biology of the species.
Musick and McEachran (1972) established co-occurrence of B. superbus with Nephropsis aculeata, Rochinia crassa, Munida iris, Homarus americanus and Cancer borealis in the area from Cape Hatteras to Cape Henlopen. Roberts (1969) des cribed zoeal stages reared in the laboratory. Garstang
(1897) observed burrowing behavior in Bathynectes longipes, the only other member of the genus and an inhabitant of the
Mediterranean area.
This study considers the following aspects of the biology of Bathynectes superbus: the synonomy, taxonomy and systematics, distribution and abundance, feeding habits, commensal associations, morphometries, reproductive anatomy, gonadal condition, morphological abnormalities, and intra specific setal variation. 2 SYNONOMY
Portunus superbus^-Costa, 1853, p r 1 9 r pi, 7; Carus, 1885,
p, 517, Vol. 1.
Bathynectes longispina (male)-- Stimpson, 1871, pp. 146-147,
Vol. 2; Milne-Edwards, 1873-1881, p. 234, pi, 42, fig.
1; 1881, p. 10; Smith, 1881, p. 418; 1883, p. 17, Vol.
6 ; Bourne, 1890, p. 314; Milne-Edwards and Bouvier,
1894, pp. 29-30.
Bathynectes brevispina (female)'— Stimpson, 1871, p. 147, Vol.
2; Milne-Edwards, 1873-1881, p. 235.
Thranites velox— Bouvier, 1876, p. 60, pis. 14, 15; Bovallius,
1881, p. 9, pi. 2; Sars, 1886, p. 1, Vol. 15,
Bathynectes superba— Norman, 1891, p. 274; Adensamer, 1898,
p. 612; Milne-Edwards and Bouvier, 1923, p. 311, Vol.
47; 1899, pp. 25-27, pi. 2; 1900, pp. 65-67; Bouvier,
1922, pp. 59-60; Bouvier, 1940, pp. 247-248, fig. 159,
pi, 9; Rathbun, 1930, pp. 28-30, pis. 9-10; Monod, 1933,
p. 510; Nobre, 1936, pp. 39-40, pi. 15, fig. 36; Zariquiey,
1946, p. 158; 1948, pp. 279-280, pi. 23, fig. 1; 1952,
p. 35; Capart, 1951. pp. 121-123, fig. 42; Dieuzeide,
1955, pp. 48-50, fig. 6-7; Holthuis and Gottlieb, 1958,
p. 118; Forest and Gantes, 1960, p. 351; Rae and Lamont,
1965, pp. 23-28. 3 4
Bathynectes superbus — Monod, 1956, p, 183; Zariquiey,
1968, pp, 380-382; Williams, McCloskey and Gray, 1968,
pp. 50-51; Christiansen, 1969, pp, 69-71, SYSTEMATICS
The genus Bathynectes contains two species; B. superbus
(Costa) which ranges from south-west Africa to the coast of
Norway in the eastern Atlantic and from Massachusetts to the
Florida Straits in the western Atlantic (Christiansen, 1969)? and B, longipes Risso (1816) which is found in the Atlantic
Ocean near southern Englandf the Mediterranean Sea, and the
Black Sea (Pesta, 1918; Norman, 1891).
The following summation of diagnostic and descriptive characters for B. superbus is largely taken from Christiansen
(1969), Bouvier (1940), Zariquiey (1968), Norman (1891),
Rathbun (1930) and Capart (1951);
The frontal margin is divided into four teeth with the middle pair narrower and more protruding than the outer which are broad and triangular. In young crabs, the middle teeth have a shallow diastema which widens in older crabs and results in two equal teeth. The frontal area is wide relative to the entire carapace width. There are five antero-lateral teeth with the first and second teeth less acute than the third and fourth; the fifth tooth is much longer, more conical, more spinelike than the others? about twice as long as the fourth spine. The fifth spine is generally much stronger than in the large specimens, although variability does occur. Antero-lateral teeth 5 6
2-4 are not strong. The merus of the chelipeds has one tooth on the inner margin and a short tooth on the upper edge; the carpus is distally formed on the inner side into a large, acute tooth which bears three smaller teeth on the anterior edge. The external edge has a short tooth and the superior surface has three other spines and many tuberculate processes on the upper-outer surface. The propodus consists of five carinae on the dorsal surface: The inner with an anterior denticulated spine; the next carina with 5-7 small spines; the middle carina tuberculated and the outer two nearly smooth. Norman (1891) classifies a sixth carina which is faintly marked and passes down the middle of the inferior surface and runs to the end of the fixed finger. The ventral surface of the propodite is smooth with two indistinctly marked ridges; fingers are nearly as long as the propodus and strongly carinate. Pereiopods 2-5 are slender, laterally compressed with 2-4 being the longest and 5 the shortest.
Carinae are present on the last three segments. The dactyl and propodus of the fifth pereiopod are flat and broad with hairy margins; the dactyl is lanceolate. The carapace is hexagonal in shape; the surface is granulated. A well- marked ridge extends laterally across the middle to unite the bases of the fifth lateral spines. The other regions of the carapace are less well-defined with a short ridge on each of the cardiac and gastric regions; the latter is interrupted in the middle by a longitudinal depression projecting to the frontal margin. Rathbun (1930), Bouvier 7
(1940) and Capart (1951) state that the carapace is almost entirely covered with short hair or pubescence. This pubescence was not present on the specimens taken in
Chesapeake Bight which I examined, but it was observed on
USNM specimens from West Africa.
The abdomen in male crabs is triangular? somites 3-5 are fused. The female abdomen becomes semi-circular in shape with age? all somites are freely articulated.
The in vivo color of the dorsal carapace is reddish orange while the ventral surface is white. The spines are generally very red. The pereiopods are marked with alternating white and orange bands. The chelipeds are specked with red? tips of the dactyl and fixed finger of the chelipeds are dark- brown to black with color extending along the finger to a junction with the propodus in larger specimens. The merus and carpus of the legs and the propodus of the last leg are specked with acarlet. The propodus of the pereiopods is mostly scarlet except for light brown at the extreme distal tips.
According to Pesta (1918), Bouvier (1940), Bell (1853),
Milne-Edwards (1834, 1861), Risso (1816), and Roux (1828), the diagnostic characters of Bathynectes longipes are as follows:
The frontal area is only slightly quadrilobed. The first four anterolateral teeth are similar to those of B. superbus but the fifth is much shorter and not more than half as long again as the fourth. The chelipeds are armed 8 only with a simple carpal spine and a spine at the distal end of the most internal of the two longitudinal carinae of the cheliped. This spine bears no lateral teeth; the propodus bears one distal tooth at the extreme inner margin. The propodus has two carinae above, the inner one terminating in a small spine. The dactyl has 3 carinae. SECTION I
DISTRIBUTION AND ABUNDANCE OF Bathynectes superbus(Costa)
Introduction
Bathynectes superbus is an archibenthic crab that ranges
from Massachusetts to the Florida Straits in the western
Atlantic Ocean and from Norway to Angola, Africa in the
eastern Atlantic (Christiansen, 1969). Since the species
description by Costa in 1853, there have been numerous
accounts of its geographic distribution (Appendix 1) but no
information on relative abundance and bathymetric distribution.
This study describes the distribution and abundance
with depth and temperature of B. superbus off the Middle
Atlantic Seaboard of the U.S.A.
Materials and Methods
The R/V Columbus Iselin (University of Miami) trawled in Norfolk Canyon (37900'-37°10'N; 74°10'-75°45'W) and on the adjacent continental shelf (36056'-37°09'N; 74°50'-75051'W) and slope (36032'-36°48'N; 74°25'-74°461W) from June 4-16,
1973 (CI-73-10) (Figure 1). Forty -nine half-hour tows were made from 15-2000 meters. Trawling was conducted according to depth strata of 1 0 -2 0 , 20-40, 40-60, 75-150, 150-400,
9 Figure 1.
Chart of Middle Atlantic Bight showing locations
(darkly shaded) at which Bathynectes superbus were captured in June 1973, November 1973 and March 1974.
The 40, 200 and 2000 m depth contours are shown.
Lightly shaded areas indicate bathymetric and geographic ranges of fishing effort.
10 '■ WASHINGTON CANYON
NORFOLK CANYON
30 60
N. M I L E S 11
400-1000, 1000-2000 m (Table 1).
A 15 meter semi-balloon, four-seam trawl equipped with plastic mud rollers, and steel China V-doors was used during the cruise. Conventional metal trawl floats were effective to about 1200 m. In deeper water, these were replaced with
2 liter plastic Nalgene bottles filled with gasoline. The nets were constructed of nylon netting with stretch mesh:
44 mm in the body, 37 mm in the intermediate part, 36 mm in the cod end and a 12 mm inner liner.
Bottom temperature, salinity and dissolved oxygen samples were taken before each trawl. Water samples were obtained with 1.7 liter Niskin bottles equipped with reversing thermometers. Bathythermograph or expendable bathythermograph casts were made at each station. Salinity was determined on an induction salinometer (Beckman RS-7B) and dissolved oxygen was determined by the azide modification of the
Winkler titration method (Strickland and Parsons, 1968).
Eighteen successful trawls were made by the R/V Dan
Moore (North Carolina Division of Commercial and Sport
Fisheries) from 17 November to 14 December 1973 (DM-030)
(Figure 1). Six deep water tows were made on a transect from approximately 36 nautical miles ESE of Oregon Inlet
(35°29'N, 74°44'W). Towing depths ranged from 387 meters to 1134 meters with an average between 630 and 720 meters.
Twelve tows were also attempted along the 180 meter contour line extending from 48 nautical miles east of Currituck
Beach (36°18'N, 74°49'W) to 9 nautical miles NNE of Wash ington Canyon (37°20’N, 74°15'W). Actual towing depths Table 1. Catch of Bathynectes superbus and bottom hydrography at Cl-73-10 trawl stations in June 1973. — A A A rH O rH o o co Q Q \ U S U S Q A E ■P -H O CD O 0 0 g g h e 10 d rd rd rd S' ft d) u s — ft— 0 rd p d \ (d h o •
0 H £ •rH - i—I 1 co A +J < H 0 rd o P CD rd 0 P £ e On 11
H OH rH 00 M—I o o o o o o A rH o o oo oo oo C"- "vt1 IS n *3* o 00 00 00 oo oo oo A rH *+H nin s o is on is oo co r~- on oo oo CO o o oo oo oo oo oo oo r O o o o o o o CD id -1 i— I 12 X! rH MH * CO oo oo i—l on 00 00 CM IS o o o o o o o o o o CD 3 in i * 00 00 i—I o o q o o H oo co oq oo co i c oo an an oo c i CO in n i oo oo oo oo o s o ~ is oo s i r~* oo Is* oo O O D M O D I O I 00 m n n n oo in 00 in in m m in 0 ft 0 o o o o o o CM CM rH CM OH ^ 00 H i—I rH rH 00 in o 00 h 00 in * n i (Ti 00 rH N n O' in H 1—1 o n H oo rH o an oo ID ' O o o n i oo oo ^ n i rH CO i —I i —I i —I C M i —I n i n i in in in in in OO OO o 0 ft n i i in in m in in ■- o r-'- " f co " r [■-- 10 H O O O O rH o o o o o n m in n i ^ 0 0 0 0 oj ^ H rH s i h 00 0 0 0 i o ( 0 00 00 h ^ .Q U ■P E 0 G td h Mean Depth Temp. Sal. DO Catch (numbers) ■H ■H o o S -p -P ■p < cd cu u e &) o o o r n i H oo H oo o r o r ro o r ro ro ro ro o r i— rH in o') oo m in n i n i n i n i n i O S O CO h VO O OSCO o o o o o o S O S H OS O O CO OS H H H H H H ix) « r " r ix> vo n i o o o co oo oo vo in in o o < ro 13 o r o r ro o r ro ro o r ro ro < s s vo - r m os o os — o I o o o n i oo rH Is o i—i i—f n in m in in in in n i n i n i n i m vo n i ^ in ro ^ ro N s C C C C ro CN CN CN CN h OS l r* cm 0 0 p - ...... o r» o p- o - p ro r*» ro ~ p- o oo oo - p - p p~ 00 00 H 0 0 00 ON rH rH ON as r-- r - as vo r-' vo as - r r-- o o rH o o o o c o o o o s o o - r oo co as oo o ro ro ro ro o r o r o r ro o ^ H m o r o n m m m in n i n i '? in in o o o o o o co » r oo - r o o o o o cd >i 0 G G cm • n d v F o m ro "vF 'S’ 00 vd 3 vo i O- o r vo vo s s s as as as as o os ■ ' r —i t H —I co i —I < —i oo as vo vo as oo os os o o r o ro ro ro ro O o ro ro ro o r o r ro m m m n i i —I i —I i —I i —I o o o o o o o o o o o o o o o o O G cd >i G o in co - r h d i ^ ^ ^ 'H' oo m cn in h 14 ranged from 115 meters to 525 meters with an average depth of 228 meters. Fishing time varied in duration from 1 to 3 hours. Tows were made using a Yankee # 36 fish trawl with an 18.3 m headrope. This gear was lost prior to the last two stations and a Yankee # 41 fish trawl was substituted. The Yankee # 41 two-seam net consisted of 127 mm mesh except for the cod end which was constructed of 114 mm mesh and lined with 13 mm stretch mesh. The head rope was 24 meters in length; the footrope was 30 meters long and roller-rigged. Hydrographic data were not available from this cruise. Fifty-four one-half hour tows were made on the conti nental shelf (35°58.5'-41°04.5'N; 71°18'-75°031W) during the National Marine Fisheries Service groundfish survey (R/V Albatross IV cruise 74-4) from 12-21 March 1974 (Figure 1). Fishing effort was distributed according to depth as follows: 19 tows in 22-60 m; 16 tows in 61-100 m; 9 tows in 101-150 m and 10 tows in 151-338 meters. A Yankee # 41 two seam net was used. Bottom temperatures were taken by bathythermograph casts. Except for a few specimens which were processed at sea during CI-73-10, all B. superbus were preserved in 10% sea- water-formalin for later examination. The following ob servations were made on the preserved individuals: morpho- metric measurements (see Section II), sex, molt stage, reproductive anatomy(see Section III), stomach contents (see Section IV), fouling organisms (see Section V) and morphological abnormalities (see Section VI). Usually only 15 sex, molt stage, carapace width and length data were re corded in specimens processed on board ship. Molt stages were determined from descriptions (Table 2) modified from Drach (1939). Short carapace width, i.e., the distance between the notches at the bases of the fifth antero-lateral spines excluding curvature of the carapace, is used in tables or figures relating to width frequency. In addition, U. S. National Museum (USNM) records of B. superbus collections were examined (Appendix 2). Areas of collection for the Western Atlantic ranged from Martha's Vineyard to the Tortugas. Collection log information on USNM specimens is used only as supportive data for the re sults of the cruises already mentioned and is not analysed in the results. Table 2. Intermolt stages of Bathynectes superbus. Modified from Drach (1939). Condition Stage Description Hard C-,“C. Exoskeleton very firm and strong; 1 4 pressure on sub-branchial area of carapace does not cause indentation or flexing of shell Peeler D-D. Advanced premolt. Shell firm, but 2 4 ecdysial suture indents when pressure is applied. In crabs actively molting, the ecdysial suture is clearly separated. Where the carapace is broken,the newly formed shell is evident Soft A Newly molted, skeleton very soft 1 Paper A2~B2 In early paper condition, the exoskeleton is leathery, pliable. The merus and propodus bend without breaking. In late paper condition, the exoskeleton is brittle, not easily broken. The anterolateral spines are hard. Merus and propodus crack when bent. 16 17 Results In June 1973, slope water ( >35/0) was found at depths >75 m in both the Norfolk Canyon and the adjacent conti nental slope. Bottom salinity was lowest (31. 9-32. 0/o) at the stations nearest shore and increased with distance off shore (Table 1). Warmer water (11-13 C) was found in the 75-150 m stratum. The slope and canyon temperatures in both areas gradually decreased with depth: 8-11 C in the 150-400 m stratum and a range of 3-5 C at depths >700 m. The pattern of oxygen distribution was that of an oxygen maximum at or near the surface with a gradual decrease with depth to the oxygen minimum followed by an increase with depth until near-surface values were reached. The level at which dissolved oxygen values were below 60% sat uration (or oxygen minimum) (Seiwell, 1937) occurred at 250 or 400 m with oxygen values between 4.5-4.7 mg/1 in both the canyon and adjacent slope. In June 1973, 139 Bathynectes superbus were caught in nine trawls within a depth range of 252-390 m (Table 1). They appeared to be contagiously distributed over the geo graphic area sampled, since there was an excess of stations at which no crabs were captured. Contagious distribution was also indicated by a sample variance divided by the sample mean>1 (12/1.5 = 8) (Woolf, 1968). 18 Bathynectes superbus were present only in 250-400 m of the canyon and slope areas, except for one crab recorded at 751 m (400-1000 ra depth stratum) on the slope. Since it is likely that this crab was left in the net from the previous trawl catch made in the 150-400 m stratum, it is not in cluded in further analyses. Crabs were contagiously dis tributed within the 150-400 m stratum since the sample variance (226) divided by the sample mean (11.5) gave a value (19) >1. Crabs were most abundant at depths between 250 and 350 m. The apparent presence of crabs in shallower depths in the canyon (250-300 m) than on the slope (300-350 m) is an artifact resulting from more intensive sampling in the can yon at 250-300 m and on the slope at 300-350 m (Table 1). Bathynectes were captured within a range of 8.0-11.3 C (Table 1) but were most abundant at 8.0-9.9 C. The 10.0- 11.9 C range was associated with one station in the 150-400 m stratum. The short-width frequency distribution of male and female crabs is shown in Figure 2. For specimens on which short width was not measured, this dimension was derived by application of the regression basdd on morphometric data presented in Section II, Table 9: x: short carapace width Male: y = 6.020 + 1.121x y: long carapace width Female: y = 1.677 + 1.124x Male crabs ranged from 21-70 mm in short carapace width; females from 28-62 mm. Four modal groups are apparent in 19 Figure 2. Further analyses of the population structure are based on the arbitrary designation of these four groups, i. e., those crabs <35 mm, 36-45 mm, 46-57 mm and >58 mm. An index of transformed mean abundance: Y = In (x + 1), where x is abundance in number of crabs, was applied to the data to normalize the contagious distribution and eliminate the proportional relationship between the mean and variance (Pereya, Heyamoto and Simpson, 1967). The transformed index of abundance by size group for male and female crabs in the 150-400 m stratum is shown in Figure 3. Among male crabs, those 46-57 mm were most num erous at 250-300 m in the canyon and at 300-350 m on the slope. In the remaining size groups, fewer male crabs were found in the canyon except for smaller individuals (- 35 mm). Larger crabs (46-57 mm) were likewise most abundant among females in the canyon at 250-300 m but females in the 36- 45 mm size group were most abundant on the slope at 300-350 m. One ovigerous female, 42 mm, was found in the 250-3 00 m depth in the canyon. No female crabs >58 mm were found at the slope stations of 300-350 m. One male 45 mm and two female crabs 52 and 45 mm were found in the 350-4 00 m depths of the slope. A transformed index of abundance was also calculated by size group for temperature ranges encountered at the trawl stations (Figure 4). Male crabs 46-57 mm and 36-45 mm females were most abundant in the 8 .0-9.9 C range. Males and females <35 mm were nearly equal in abundance at 8.0-9.9C Figure 2. Width frequency distribution of Bathynectes superbus captured in Norfolk Canyon and the adjacent slope during June 1973* Intermolt (C^-C^) stage is rep resented by the blank areas; peeler (D-^-D^) by the solid area; soft or papershell (A^-B2) by the cross- hatched area. Males are plotted above the line, females below. 20 o _ o SHORT SHORT CARAPACE WIDTH (mm) Q. cx Q. _ o o CM (D s ID If) ^ IO W o w IO ^ 10 ID sivnoiAiQNi jo djgw nN Figure 3. Index of transformed mean abundance of four size groups ( £38 mm, 36-45 mm, 46-57 mm, ^58 mm) of male and fe male Bathynectes superbus captured in Norfolk Canyon and the adjacent slope during June 1973, in 250-400 m. The fraction above each stratum is the ratio of the number of stations at which the species was captured to the number of stations in the stratum. 21 CANYON 1 I SLOPE ro|ro 10 IO 0vNa J X 3 0 JON I 30NvoNnav CO-1 ro|ro 110 CM or Ol i _ 0 C z o DEPTH (m) Figure 4. Index of transformed mean abundance of four size groups ( ^ 35 mm, :36-45 mm, 46-57 mm, ^ 58 mm) of male and female Bathynectes superbus captured within two degree Celsius temperature strata during June 1973. The fraction above each stratum is the ratio of the number of stations at which the species was captured to the total number of stations in the stratum. 22 □ < 3 5 7_ £3 36-45 15 ESI 46-57 m > 58 3 - Cf IzJ i O Z I _0 < 10 a ~T~ 6SI-0-bl-iO|cM z 0) O) 01 0> G) to in cr> to CO 1 I 1 1 °C 1 1 I < o o o o o O cvi 9 3 “ 7 15 4-J and were equal at 10.0-11.9 C, No female crabs >46 mm were present in the 10.0-11.9 C range. Males slightly outnumbered females in the slope and canyon collections (Table 3). The maleifemale (M:F) ratios for the canyon (1.1:1) and for the slope (1.3:1) were not significantly different from 1:1. Female crabs predominated or were equal in number to males in the -35 mm and 36-45 mm size groups for the canyon and slope areas. Male crabs ^ 46 mm were generally more numerous than f emale- B ,, superb'us of the same size range. A Chi-square test using Yate's correction for continuity (Woolf, 1968) indicated that the ratios differed from the expected 1:1 ratio for the slope stations in the 46-57 mm and ^ 58 mm groups. Total catch was nearly equally divided between true hard shell (C^-C^) crabs (47%) and those showing evidence of recent or incipient ecdysis (53%). Relatively few peelers (D^-D2) (11% of total catch) were represented in the latter group. Male and female hard crabs were more abundant on the slope than in the canyon. Male hard crabs were most abundant in the larger (36-45mm, the 46-57 mm and the >58 mm) size groups whereas hard females were most abundant in the smaller ( < 35 mm and the 36-45 mm) size groups. Peeler crabs were more numerous on the slope for both sexes. Male peeler crabs were present throughout the size ranges of ^ 35-> 58 mm. Female peeler crabs were present only in the range of < 35-57 mm. No female peeler crabs were found in the canyon for the < 35 mm and the 46-57 mm size groups. 4 'd tn 0 CN 03 CO CO ■H CO •p X P P p 0 p g o 0 o a o X 0 0 r-l rH 1—1 i—1 H i ■rl •• • • 44 •4 *• o 0 -p CO ID 00 ID CN ID X p 4 •• 4 4 rH -p (Z o o 1—1 P" rH 0 g H X 0 P -p p -P 4-1 0 P4 1—I VO CNVO p rH CN rH ID ■H CO P • 'd 0 P 0 ■H S p -p P £ CD O ID 00 P 0 H i—! ro i—1 vo ■p ■H 0 a > o rd 0 O Q 0 P CN CO CO 03 CO 03 CO rd X P P P p P P • P XI ro p r- co o oi 1 a. i—I *H 0 tH i—1 i—1 rH i—1 p XU ■H • • •• • • • • 44 CO 0 u -P i—1 VD i—1 ID 1- 1 p P • 4 4 CO p 0 O ro 1- 1 0 I“D X! P -p -P o o K >1 0 0 P P Cm VO r-' CNCNO' p CO -H P .H CN •H U P -P -p P P rd O rd £ VO CN CD PQ OH CN O *H p h m 0 1 -H • 4 4-1 ro P d> tn CN CO 03 •H ■H CO a 1 -H X P P CO CO P P H CO 0 u p •d tn 0 P r-l 0 i—1 i—I tH o i—I 0 •H 0 ■H •t •» • t 44 4 4 N p ,Q +j VO id VO 00 ro ■H P rd p • • • 4 CO 'd iH & O o ro 1- 1 >i g 0 0 X P P Q a -P rd 0 03 rd iH El, LT> cn ID o CD 0 P o 03 i—I CN ■H ■P *H -P 01 -P 0 rd P X P -p £ ro o 00 00 o X C4 H ■—l rH ro 0 0 •• 03 fd H • g ro g LO i—1 '— LO ID 00 P 0 CO 1 1 LO -P 1—1 0 VO vo o N X VI ro Al Eh rd •H E-< 03 24 25 The incidence of soft and papershell crabs was higher in the canyon than on the slope area. Papershell crabs predomi nated in the < 35 mm group for males and in the 46-57 mm group for females (Table 4). Cole's (1949) coefficient of association (C7) was cal culated to determine whether there was co-occurrence of B. superbus with other selected crustaceans (Table 5). Since only a determination of the presence of absence of assoc iation was desired, was computed instead of the Cg coeffi cient of Hurlbert (1969). Analysis for revealed that B. superbus was most positively associated with Cancer borealis in June but was also associated with Homarus americanus and Geryon quinquedens. No significant association was found with Cancer irroratus. Five ovigerous females and one male B. superbus were collected in three 2 hour tows in November 1973. Four ovigerous crabs were collected in depths that ranged from 122 to 165 m. The other two specimens were captured at a mean depth of 223 m. The ovigerous females, all interm°l'tr were 47, 56, 57, 57 and 63 mm (short carapace width). The male crab, a papershell (B2), measured 66 mm. Analysis of the C coefficient of association (Cole, 7 1949) revealed that B. superbus was not associated with Homarus americanus (C7 = 0.127; X 2 = 329) or Geryon quinquedens (C? = -1; x2 = 1.37) for the areas trawled on this cruise (Table 6). Information on Cancer borealis and CM O' CO o OO' rH ffl • • 1 rH ro o LO rH rH O' LO LO < W 0 f t p p p P o 00 0 1-3 i lo ro O' LO P g IN • • • o rH • tP CP g Q lo oo LO -5? o 1—1 00 p CD -H O' N P LO -H P 1 ^ in np LO U o o O o o LO ^ 1 • • < • • • • P W i—1 o o o o LO CO o P P a LO LO LO CM CM CM O 42 LP P CD CP ft • P P 0 00 CM o CM O' O' 0 M 13 i—1 i—1 CO LO rH rH PiC! rd in CD CM in -P P2 o o as LO LO 00 o a) o 1 • • ••• • • CP CD rH O LO CM i—1 00 o LO p P < ro O' CO CM CO CO -p > i in xi -p •P id rH CQ Q rH LO I—1 o 0 1 •• • • g MH g og rH 00 CO LO P 0 g Q o o o CM CN CM 1—1 CD -P - v LO P g •St* *P g 1 ^ LO u o o rH 00 i—1 o P 00 to 1 • • ••••• rd lo 1—1 O LO O' O' CM LO o u O' CM LO "vT LO £j Al 43 - 0 g • rd g 0 o as LO o P s rH ^ rH rH O' CM 42 O' LO m i CN 0 LO PQ CO O LO O o o "cr 1 • • ••••• CD rH CO o o o LO o O ' 0 CO LO as LO LO 20 !3 p p 0 0 0 0 rH 0 0 rH i—1 i—i H 0 f t >1 0 0 f t >, 0 0 0 42 0 o p -P «—1 0 p +J g -P 0 p rH 0 O 0 rH 0 0 0 o Eh < cn u Eh g C/3 o EH ft Eh 26 CN DQ CO CN CN O LO 00 as | c ■ ■ •«• • • rH cn rH rH rH in CN ■—i c CO TP TP CO in TP TP TP Q CN i—1 CO 1 i" CO CO • t p •• CN •••• LO 1—1 Q r- O' c- CN n> i—1 rH rH <0 -p o Eh t P U o in in 00 o i—1 00 1 ••• • •• • 1—1 ■—i rH t P o rH LO o m in in •S3* CO TP TP TP 0 cn 00 00 OL LO CN z co LO LOCN CN LO i— 1 CN ffl in ■—i CO O O CO 1 •••••• rH CNCO o o in c i— 1 in CO OLO in CO g ^ g Q in 1 • r" cn 00 CN CN •• in q i—! o LO O o o m Al tp u O as O o o 00 1 • • •• •• rH LOCN o o o 00 u t" tp LO O in in in 0 in r - ■p s 00 rH O CN CN ■—I c o u TP a C CD CD CD 0 rH CD 0 i—1 i—1 H 0} I—I cd Oj >1 cd CD ft >1 cd cd cd 43 CD 0 c -P rH 0 c -P g •P cd cti P rH cd O cd H (ti 0 CD 0 n EH < CO u Eh g CO u Eh ft Eh O 27 to -P 0 +> p p P P XI 0 <31 o X! 0 CN r - ._* P 0 rH CN P 0 rH CN 0 X 0 XI u a < rH a < '— p <31 p 0 • 0 O' p -P r - -P ID 0 0 P 0 P • •H 0 0 II 0 0 -P -P 0 Cl o -p 0 in Cti o 0 CM 0 0 II •H 0 P X 0 P 0 p PH p a CN 0 >< > i X CO xj XI CO ■p -P to -p -p cd p P ■p 00 p p ■P PQ 0 p in PQ 0 P ID tH 0 0 o 0 0 00 0 0 0 • 0 0 i—1 P X! o PX • +> PH << a < o p + l 0 0 p +1 •H 0 00 0 U P ■d -P in 0 cn P i—I p CN HH P • a • 0 0 o p O 0 P •p OP II P II ■H d 1 CO - p u p U 0 0 0 H o > i 0 p p U p 0 u o CT> P 0 0 "H P P ■p XI XI p P -P P -p a 0 P 0 p g a 0 CT> O a 0 ID 00 0 p 0 00 p 0 00 o 0 X2 0 X! < < p 0 ID 0 •H 0 O 0 ■p ■P o i -p -P in 'd u P oo o P • 0 0 0 • 0 0 rH CO p 0 cn o p 0 00 ID 3 0 II 0 II X P X! P CO •P PH CN -P a CN 0 P X P X rH pq PQ XI • -P ■P cd oo P -p P ■P -p r - 0 p ID 0 P Cl 0 0 i—1 0 0 > lH 0 0 ID 0 0 1—1 o P X! • P X! • P PH <1 O a < o 0 0 0 tn P +1 p +1 P P p •H 1-0 0 LO p <31 -P *H 00 u O' P P f ■[ 00 ■rH i—1 0 O (tf • P U MH 0 o 0 O P £ 0 II P II • XI ID 0 r - p U P U 0 0 p i— 1 0 P rQ p £ 0 p 0 EH 1 u td 28 Table 6. Contingency tables used in computing Cole’s coefficient of association (C_) for November 1973. Bathynectes superbus Present Absent Present 3 8 Homarus americanus Absent 0 7 C? = 0.127 ± 0.007 X2 - 329.2 Bathynectes superbus Present Absent Present 0 5 Geryon quinquedens Absent 3 10 C ? = - 1 + 0.850 X 2 = 1.37 29 30 C. irroratus was unavailable. In March 1974, thirteen Bathynectes were captured in three tows from 236-300 m and at bottom temperatures that ranged from 8.7-10.1 C (Table 7). Most crabs (6) were captured in the 280 m, 10.1 C haul. Most (9) of the male and female B. superbus were hard shell crabs; three were in the pre-molt condition and only one was papershell or soft (Table 7). All crabs > 60 mm and 6 of the crabs < 60 mm were hard. Only one papershell (56 mm) was captured. Peeler crabs ranged from 52-56 mm in size. Male crabs outnumbered female crabs but the ratio (1.6:1) was not significantly different from the expected 1:1 ratio. Low positive association was indicated .between Bathynectes superbus and Geryon quinquedens and Cancer borealis. There was no significant association with Homarus americanus or Cancer irroratus. The C^ value (-1) for C. irroratus suggests that this species and B. superbus are distributed randomly with respect to each other (Table 8) . cd tp TP tp Tp TP •p tn U u U P 0 rH fd 1 1 t>1 1 | 1 l 1 O -P rH 1—1 rH 1 l 1 (N I rH < u S w u u u P U rd 42 g rH CD <1 CP CD — N g 00 00 1 I I VO I TP -P TP tp vo 1 I 1 in I in fd CO w CD u p -p CD TP tp TP CN TP TP TP TP rd +J Cn p P U PQ u 0 U u P i—i td 1 i 1 1 1 1 1 1 ■ CD O -P CN CN tH CM 1—1 1—1 1-1 1—1 i g co Q P u < u o u u & CD -P td g g CN Tp TP vo TP r» o rH I o in in LO in vo r* m VO i -p -p o 42 fP — r- c g u (0 CD co Eh CA P 42 P 45 CD -P Pj| &. P CD CO Q o vo o g 00 ro o CO s5 ~ CN CN ro 0 td -p CD o g 0) C S3 J3 >i Cn o CO ro -P fi o ro TP tCJ O o o pq i-5 CN CN TP r - r- MH 0 45 O 5 -P rd r"» ro LO U H •P ro ro fd i-5 42 fd ro td -P tp ro Eh co i-H ro 31 Table 8. Contingency tables used in computing Cole's coefficient of association (C7) for March 1974. Bathynectes superbus Present Absent Present 3 18 Cancer borealis Absent 0 33 C ? = 0.0907 ± 0.0410 X 2 = 4.84 Bathynectes superbus Present Absent Present 3 13 Homarus americanus Absent 0 38 C ? = 0.140 + 0.0509 X 2 = 7.56 Bathynectes superbus Present Absent Present 1 2 Geryon quinquedens Absent 2 49 C? = 0.294 + 0.136 X 2 = 4.67 Bathynectes superbus Present Absent Present 0 18 Cancer irroratus Absent 3 33 C ? = - 1 + 0.793 X 2 = 1.59 32 33 Discussion The data in this paper suggest that Bathynectes superbus is contagiously distributed on the lower shelf- upper slope areas of the Chesapeake Bight. Its bathymetric extension into shoaler or deeper water depends on the hydro- graphic conditions of the area. Although B. superbus has an extensive bathymetric range for the western (159-633 m) and eastern Atlantic (85-1455 m), it was consistently found on the shelf edge-upper slope areas at 122-390 m in the Middle Atlantic Bight. The USNM specimens examined were captured at 35 out of 43 stations in both the eastern and western Atlantic in which the depth was within the 150-400 m range. Capart (1951) and Monod (1956) give extensive infor mation on the occurrence of Bathynectes superbus in the eastern Atlantic, off the African coast. Crabs were dis tributed from 290 to 660 m with one occurrence at 85 m (Monod, 1956). No hydrographic conditions were given, but it is likely that the water of the Benguela current greatly affects the bathymetric distribution of the African coast fauna. The dense cold water of this current is found close to the African coast and upwelling of water from mod erate depths takes place as the surface water layers are carried away from the coast by the southerly and south easterly winds (Sverdrup, Johnson and Fleming, 1942) . Since B. superbus is probably limited in its vertical distribution by temperature, it may extend its range into water as shoal as 85 m in the area of upwelling. My data suggest that B. superbus is most abundant in the 8.0-11.9 C range. Rathbun (1930) gives a temperature range of 5.5-11.1 C for the occurrence of B. superbus off Nantucket Shoals and Martha's Vineyard during the months of August to October and 8.3-14.4 C during September and October in Chesapeake Bight. In the more southerly stations off Florida and Georgia, crabs were captured in the 6.5-12.1 C range during March, April and May. Despite the occurrence of B. superbus at the higher temperatures of 12.1 and 14.4 C, 10 out of 17 total stations and 74% of the crabs from these stations were captured within the 8.0-11.9 C range (Rathbun, 1930). Seasonal changes in bathymetric distribution may be related to water temperature changes above the permanent thermocline. In June 1973, B. superbus were consistently captured below 200 m, the depth of the permanent thermo cline (Wenner, unpublished manuscript), In November and December 1973, crabs were collected at depths of 122-165 m; in March 1974, at 235-331 m ; and in April 1974 at 315 m. This suggests that crabs are found deeper (252-3H0 m) in June, March and April than during the winter months. How ever, the vulnerability of B. superbus to the large mesh gear in November and March is low. Also, trawling effort on the Albatross IV was primarily made on the continental shelf with very little work on the upper slope. Therefore, 35 bathymetric records of B« superbus from these cruises must be considered incomplete. Observations suggest that B. superbus is capable of withstanding temperatures well above the range of maximum abundance. Bathynectes superbus kept in a shipboard aquarium during the R/V Eastward cruise (2-74) in April 1974 (see Section II for gear and station descriptions), indicate that crabs can withstand temperatures approaching 21.9 C. After having been trawled from a depth of 350 m at 9.4 C, crabs were exposed to continuous flowing surface water ranging from 14.8-21.9 C for five days. Prolonged survival in unusually high or low temperatures is questionable and reproduction would be unlikely in these extreme temper atures. The apparent association of B. superbus with the shelf edge-upper slope habitat therefore is most likely a result of relatively stable temperatures associated with slope water rather than the eurythermal conditions that occur with coastal water along the inner shelf. Inferences concerning tropical submergence of B. superbus are difficult to make since there has been no consistent seasonal or latitudinal collection of specimens and hydrographic data. Furthermore, collection methods have not been standardized. Rathbun1s (1930) bathymetric data for B. superbus in the western Atlantic are somewhat confusing since crabs have been reported at similar depths in boreal and tropical provinces. A depth range of 140-405 m has been recorded for areas from North Carolina to Martha's 36 Vineyard. Crabs were collected in deeper water (496-633 m) off Georgia and Florida, although at Sand Key Light and off Key West, B. superbus were captured in 162-228 m depths. Depth distribution latitudinally could result from water temperature fluctuations over the seasons sampled. The values indicate that in June 1973, B. superbus was associated with Cancer borealis, Homarus americanus and Geryon quinquedens. Musick and McEachran (1972) found that the bathymetric distribution of Homarus americanus, Cancer borealis and Bathynectes superbus were similar for Chesapeake Bight in autumn and winter. Some co-occurrence with Geryon quinquedens was observed in June 1973, although this species was most abundant at 400-1000 m. This could explain the low association in November and December. Bathynectes superbus has been captured in the same tow as Geryon quinquedens off the West African coast between 300-500 m at 7.8-9.9 C (Capart, 1951) . The lack of association of B. superbus with Homarus americanus in March, November and December may be related to seasonal migration of the lobster in Chesapeake Bight (Cooper and Uzmann, 1971). Lobsters were generally found within a temperature regime of 10-17.5 C. Offshore lobsters migrate to optimum temperatures according to season, moving into shoaler water in late spring and early summer and re turning to deep water at the shelf edge and slope in late fall and early winter. The non-significant association of Cancer irroratus, 37 the rock crab, with B. superbus in June, indicates that the deeper end of the distribution of C. irroratus does not over lap the range of B. superbus. Rock crabs were most abundant in the 40-60 m stratum although common from 18-348 m in June (Haefner, personal communication). Rock crabs have been found to inhabit offshore ( ^ 40 m) waters during the late spring and summer. The lack of association between these species in March may be due to the increased inshore abun- cance of C. irroratus during the winter and early spring months (Shotton, 1973). Width frequency plots indicate that male B. superbus achieve a larger size than females. No females were cap tured that were > 63 mm short carapace width whereas males as large as 70 mm short carapace width were found. Females > 54 mm were few in number and may represent mature females that will not molt again. The data from this paper suggest that some element of the population is molting throughout the year. In June, molting was most prevalent in crabs < 45 mm for both sexes and papershell crabs were most abundant in the 46-57 mm size group. Data indicate that male crabs may continue to molt at sizes in the 46-57 mm range while female crabs will not likely molt again in this range. From the sex ratios calculated for June 1973, there is apparently no real segregation of males and females. Al though there were more males located on the slope for the 46-57 mm and — 58 mm size groups, this is a result of the 38 larger body size of male crabs. Females do not attain sizes much larger than 57 mm and are predominant within the 36-45 mm group. With regard to molt stage, hard and peeler crabs were more abundant in the canyon. This may be an artifact of sampling but it does suggest that the most vulnerable part of the molt cycle is spent among the mud overhangs and rocks of the canyon. Certainly, the canyon would provide more sheltered areas than the open slope and may provide an avenue for concentration of food sources as Rowe (1971) has suggested. SECTION II RELATIVE GROWTH AND MORPHOMETRICS OF Bathynectes superbus Introduction Among the Brachyura, morphometric studies have been used to study growth and form (Thompson, 1917; Cott, 1929) and attainment of sexual maturity (Haley, 1967, 1969, 1973; Ritchie, 1973; Sandon, 1937; Terretta, 1973; Shotton, .1973; .Olmsted;.and Baumberger, 1923; Hartnoll, 1965 a,,b). Among the Portunidae, relative growth of Callinectes sapidus has been studied by Gray and Newcombe (1938); Newcombe, Sandoz and Rogers-Talbert (1949) and Tyler and Cargo (1963) . Ryan (1967 a) used morphometric and morphological criteria to determine sexually mature instars in Portunus sanguinolentus. In this study, I have used morphometric analysis to de fine growth patterns of male and female Bathynectes superbus and to show changes of form with onset of sexual maturity. Materials and Methods Crabs used in the relative growth and morphometric analyses were obtained from cruises of the R/V Columbus Iselin (73-10) in June 1973, the R/V Dan Moore (73-030) in November and December 1973, the R/V Albatross IV (74-4) in 39 40 March 1974 and the R/V Eastward (2-74) in April 1974 (see Appendix 2 for station location). Methods of sampling have been described in the previous section for June 1973, Novem ber and December 1974 and March 1974. Twenty successful tows were also made from April 15-21, 1974 on the R/V Eastward, a vessel operated by Duke University for the National Science Foundation. Sampling was conducted on the continental shelf, slope and within the Norfolk Canyon from 34°22.91-37°08.51N to 73°39.5'-76°37.6'W at depths of 15- 2550 meters. Tows were one-half hour each except for two one-hour tows at 2125 and 2160 meters respectively. The gear used was a 9.1 meter semi-balloon otter trawl consisting of 38 mm mesh in the body and 13 mm mesh liner in the cod end. In addition to the specimens collected during the cruises mentioned above, morphometric data were also collected from U.S. National Museum specimens (Appendix 3). The following measurements (Figure 5) were made with metric vernier calipers to the nearest 0.1 mm: LCW: Long carapace width. The distance between the fifth anterolateral spines excluding curvature of the carapace. SCW: Short carapace width. The distance between the notches at the base of the fifth antero lateral spines excluding curvature of the carapace. SL; Length of the fifth anterolateral spines. The difference between maximum and minimum carapace length. A derived measurement. LCL: Long carapace length. The distance from the tips of the two frontal teeth to the extreme posterior margin of the carapace, excluding curvature of the carapace. SCL: Short carapace length. The distance from the base of the two frontal teeth and the extreme posterior margin of the carapace, excluding curvature of the carapace. RC and LC: Length of the right and left cheliped. The distance from the articulation of the coxa with the sternum to the most distal part of the propodus. RCP and LCP: Length of the right and left cheli ped propodus. The distance from the tip of the propodus to the articulation with the carpus. RCH and LCH: Right and left cheliped propodus height. The distance between the long dorsal spine and the base of the propodus. FOW: Fronto-orbital width. The distance between the outer angles of the orbits. FW: Frontal width. The distance across that portion of the carapace between but not including the orbital area. BD: Body depth. The distance between the high est part of the dorsal metagastric region 42 and a point along the ventral median line just anterior to the flexed abdomen. FABW-, ^: Maximum width of abdominal segments 3 and 3, o 6 in females. MABW3 6: Maximum width of abdominal segments 3 and 6 in males. Short carapace width was chosen as the reference measure ment. It seemed more accurate than long carapace width since the fifth anterolateral spines were frequently broken. Preliminary plots were made to determine if the data were linear. Linear relationships were expressed by the least-squares regression, y = a + bx. Linearity was also tested by analysis of variance and the comparison of deviations between a simple linear regression and a second- degree quadratic regression. Male and female measurements were compared by analysis of covariance to determine if sig nificant differences were present between regression co efficients, adjusted means and variances. The right and left cheliped lengths were compared for both males and females by Student's t-test. Statistical results and data plots were used to investigate sexual differences in relative growth and the size of crabs at sexual maturity. Figure 5. Dimensions used in the morphometric analysis of Bathynectes superbus. See text for description. A. Dorsal view of carapace. B. Female abdomen. C. Male abdomen. D. Chela. E. Lateral view of crab, appendages removed. F. Cheliped. 43 A sew ■jFOW I I— H FW MABW LCW B 3 FABW MABW RCH and LCH RCP and LCP RC and LC 44 Results Regression equations for all relationships examined are found in Table 9, Results of the covariance analysis for all relationships examined in the study are found in Table 10. The relationships between short carapace width and short and long carapace length were linear for the size range of specimens examined (Figure 6 a,b). The relationship between short carapace width and long carapace length was significantly different for male and female crabs. The regressions of short carapace width and short carapace length for males and fe males differed significantly in slope, adjusted mean and variance. A significant difference in the variances was most likely due to the smaller size range of female crabs sampled. The relationships between short carapace width and long carapace width (Figure 6 c ) , body depth (Figure 6 d) and fronto-orbital width (Figure 6 e) were linear (Table 9) and significantly different for male and female crabs at the 0.05 level. The regressions of frontal width on short carapace width (Figure 6 f) for male and female crabs were not sig nificantly different at the 0.05 level (Table 10), Hence, the data were pooled. The chelipeds of B. superbus were sexually dimorphic. Analysis of covariance revealed significant differences between sexes in the linear relationship of both the left and right cheliped length and short carapace width (Table 10). Table 9. Regression analysis of Bathynectes superbus morphometric data. CN •H ■H •H •H 1—1 P■ -p ■p *P N •H o o C/2 + w -P P3 w 'P i—I -P P3 (0 cn P 0 P 0 S3 td P H S3 ■H o o P Cn •P rC oo - p O 01 N nC c r—1 Ol CM cn i > i td Oi i P 0 rH 0 S3 0 0 O i p P I II II >1 X o to 0 • • .. •• • • ...... II I • or 0 'sF 00 ro ro no o o in ro fh id n u £ F .. .. a . . . . • > . . . . . ■r| 3 0 43 'p 34 43 43 43 cn ■P a -p id 0 CN 00 O HI 00 ID UO rH ' P 00 m oo r ni in in ro o ro - r 00 o o - r n c F 00 rF cn in 43 0 34 43 43 43 0 0 0 43 0 0 >P 0 0 0 0 i O P 0 £ >1 >1 0P S3 P >0 P S i P u 0 0 I II II >i > X t •« t • h . . -P in «? 45 'P fh — N N0 'FC CN CN 'sF 00 CN H CN i—I id f H *H •H 3 P 43 >p fp N NrH CN CN p p• -P •P -p •p *^F 00 a id P -P -P D rH VD n ncn X m CN ID p~ ,'P CN -i in p- or o n in oo ro ro tn + + + HrH CN CD rH 00 1>i i > i > >1 iih y 0 3 0 43 H S3 ■H p pi p fp 3 43 43 — • ■—i H n • in ' +J n r o nr- - r cn oo ro in n left cheliped length M y = -0.753 + 0.543x 99.6 72.702 left chela length F y = 0.063 + 0.526x 99.6 46.516 00 ["■ o LO LO CN LO o o CO i—1 LO rH m cn • • • • • • • • 00 LO O O LO ro CXX X X 0 X O X X LO X ro r—I •H t" ^ o L0 i—1 CN o ro P o ^ LO LO O LO I—1 0 • ro • CN • • d • o • • O • o o d 1 o o O o w + + + + + + + + Cl LO LO LO ro 0 CN 00 CO ^ LO O -sF ro ■H CTi CO 1— 1 0 \ LO rH CN CN cn rH • o CL • 00 • • cn • o • • 1— 1 • LO o 0 O 1 o H 1 O I 1 Ph to II II II II II II II II 0 Pi >1 >1 >1 >i >i >1 >1 >1 X 0 g ft g Pm g Pm g Pm g CO ro LO Cn Cn 0 0 p tn PI 01 Pi d p P P 0 £ d rH d rH d £ Pi p p •H 0 ■H 0 ■H ■H P P c n p & £ £ £ £ s D i p £ ft •H -rH 0 £ ft 0 0 0 a 0 g 0 • X 0 0) H d O 0 0 0 O p w H d 0 <0 0 d 0 Pi d td td ftP! ftrQ ftP 0 ft td id rH rH 0 0 0 0 0 tj> U ■H rH rH 0 0 u IH !H Cl — 1 p i 0 CD P PI 0 P 0 P 0 0 01 £1 £ U 0 0 0 0 O O H £ O U • 0 p p P P! P Pi P 01 OL •H P P PI PI 5h P PH P Ph -H P p p tr> to 0 d 0 d o c 46 Table 10. Covariance analysis of morphometric data from male and female Bathynectes superbus. Significance of F ratios determined at the .05 level. •H ■H Td n ■ *0 ■H P fa fa •H fa Q Td x a rH W fa x fa x p fa td e b b (D tn b 01 01 0 b td 01 0 > td b t b 0 0 b a X! o a) b 01 fi 0 \ eHe He •H 1 rts ! 0 X! r-~ ■—i He He He 0 0 rH \ VO O O00 VO 00 o 00 n i VO X -p 47 CN CN i—i He n i i—I n c n i n i o r n t — fd O a td >i b Td 0 (11 0 cn (D >. X • 1 Td x a H *H -H \ Td Td X! X! b X i CO -P X n i n i n i \ VO oo 1—1 o i—1 1—1 o NCO CN o r o o r P +> -P \ ■—1 He n c o r n c 00 - r ii —i i 00 o r o r •ej* n i o b b td 0 0 X X cn m o td o d X td td H XI - a td 1 0 b b CD rH > £ — • • • • • • ■ I x O >i o •H \ \ Td x* 0 X b X CN ■ej* CN i—i - r i—1 i n i X X X 00 CN - £"• 00 rH \ \ CN o r — - r LO o r o r n c O VO rH n c — Cd X dtd H 0 •H He \ d b Td x X x X VO CN LO - r \ 0 0 1 X X 00 CN CN CN n c O — o r n c He rH CN \ x a 0 O d x td td 0 b o X b tn o n b cn 0 O rH |5 X d X td O 0 O n c 1—1 1 Td X a •rH left cheliped length 1.6663* 136/145 16.9233* 1/281 9.8252* 1/282 left chela length CN O O rH pLf r ~ 0 0 H 1 H 1 Q CN CN r o r o \ \ \ \ H i—1 i—1 i—1 G td CD He He g 'SF r o "SF r - CN c n r o d r - c n r4 c n CD ID r - o +J •••• Hi co o -sF G rH - r o i n i n d (0 Pl, i—I t n c n o r - r o 'sF h CN CN r o r o Q \ \\ \ H (—1 rH rH G 0 -H He He cn i n rH c n O cn i—1 c n r o rH CL) r o LO "sF 0 0 M n > H* H 1 H 1 t n ••»• a ) 1—1 o CO r * U r o CN "SF *e f Fh o o c o t -" CO H 1 r - r ' i—1 r—1 i—1 t—i \ \ \ \ Q rH t n r o r o -SF r o ID H r—1 i—i i—i a) o G HC He He fd H 1 CN t—1 0 0 ■H CO O •Sf -sF M iH H 1 i n i n td ^ ji i—1 r - c n > •••• i n i—1 CN CN I n 42 42 d ■P 44 cd 42 ■ f d iH f d H G 42 -P 42 • h fd •H fd ■H -P 42 t n P £ G £ G e t n - P G & •rH *r4 td g a , CD 0) CD g CD g 4J X a) a; t— 1 d 0 0 0 O G w rH fd fd f d td f d O td fd CkX) 0.42 U cu td td rH i—1 td fd fd fd ■H rH i—1 CD CL) in u 42 a) a) 42 42 fd 4 4 fd 4 4 tn 42 42 O 0 O 0 U 0 G o o t o ID 0 4-> 4-> 44 42 - P 4 2 ■H ■P V 42 42 U 4-> • U 4-) -P 44 44 t n t n 0 n2 tn 0 d td a) a ■H -H 42 -H CD 42 -H i—i H rH 54 U cn ^ tn tn & 4 2 tunt Pi . t . . • • .. •• •• • • *i EH X > 1 X > i X > i X 48 Figure 6 The morphometric relationships of carapace measurements for male and female Bathynectes superbus. Male and female scales are staggered. a. The relationship of short carapace length to short carapace width. N = 172 males; 171 females. b. The relationship of long carapace length to short carapace length. N = 172 males; 183 females. c. The relationship of long carapace width to short carapace width. N = 163 males; 160 females. d. The relationship of body depth to short carapace width. N = 170 males; 182 females. e. The relationship of fronto-orbital width to short carapace width. N = 170 males; 183 females. f. The relationship of frontal width to short carapace width. N = 173 males; 184 females. 49 FEMALE CARAPACE LENGTH (mm) - 0 4 - 0 8 - 0 6 - 0 4 0 2 - 0 6 80 20 - - 0 . LONG CARAPACE LENGTHb. l SOT AAAE LENGTH CARAPACE SHORT Cl. 10 1 ------X ' ----- »*x 20 1 ------HR CRPC WDH (mm) WIDTH CARAPACE SHORT xx 1 ----- .V*Y 30 ( ------1 ----- • i • . y * * * $ 0 4 1 ------• 1------1 .. 50 17 ------•v** * * 9 " . ' ------.• f V* cf ••• • • • • 1 ------v . 1------1 A* * A X TO ------t 60 6 * r *x 8060 0 4 - -20 r r 80 - - -20 0 4 - 0 6 - 0 MALE CARAPACE LENGTH (mm) FEMALE BODY DEPTH (mm) FEMALE LONG CARAPACE WIDTH (mm) 100 80 - 0 4 0 2 - 0 6 - 0 3 - 0 4 20 10 - - - - . OY DEPTHd. BODY . OG AAAE WIDTH C. CARAPACE LONG 20 HR CRPC WDH (mm) WIDTH CARAPACE SHORT 0 40 30 ••« * • * « • • • 0705060 80 - 40 r- 100 r - 30 - - 40 - 80 - 20 20 10 60 MALE BODY DEPTH (mm) . MALE LONG CARAPACE WIDTH (mm) FRONTAL WIDTH (mm) FEMALE FRONTO—ORBITAL WIDTH (mm) - 0 3 20 20-1 10 - 5 1 10 - 5 - - - . RNOOBTL WIDTH FRONTO-ORBITALe. . RNA WIDTH FRONTAL f. x» 20 HR CRPC WDH (mm) CARAPACE WIDTHSHORT *»• 0 4 0 3 x fr •r x*. 0 5 0 6 • # % • 080 70 x - - 30 - 20 10 MALE FRONTO—ORBITAL WIDTH (mm) 50 For a given size, the chelipeds of adult males were generally larger than those of females (Figure 7 a, b ) . In both sexes, the mean size of the right cheliped was significantly larger than that of the left (t = 2.44, p <.02). There were significant sexual differences in the regressions of chela propodus length on cheliped length (Figure 7 c, d). Analysis of covariance revealed sig.hificant differences in variability but no difference in slope or adjusted mean for the relationship of left chela propodus length and left chela depth (Figure 7 e; Table 10). Male crabs were more variable than females. Since the relationship between right chela propodus length and right chela depth was similar for males and females, the data were pooled (Figure 7 f , Table 9). The most obvious sexually dimorphic character among brachyurans is the shape of the abdomen. Male B. superbus characteristically have a triangular shaped abdomen both as juveniles and adults (Figure 8). Female crabs have the same triangular shaped abdomen when very young but the ab domen broadens and becomes semi-circular at maturation (Figure 9). In addition, abdominal segments 3-5 are fused in the male but are separate in the female. The relation ship of abdominal segment 3 to short carapace width was linear (Figure 10 a) with distinct sexual differences in slope, adjusted mean and variance (Table 10). The relation between short carapace width and abdominal segment 6 was statistically linear for males and females but a distinct change in slope was apparent for females near 30 mm short Figure 7 The morphometric relationships of appendage measurements for male and female Bathynectes superbus. Male and female scales are staggered. a. The relationship of right cheliped length to short carapace width. N = 150 males; 144 females. b. The relationship of left cheliped length to short carapace width. N = 138 males; 145 females. c. The relationship of right chela propodus length to right cheliped length. N = 149 males; 14 6 females. d. The relationship of left chela propodus length to left cheliped length. N - 138 males; 147 females. e. The relationship of left chela depth to left chela propodus length. N = 132 males; 143 females. f. The relationship of right chela depth to right chela propodus length. N = 142 males; 141 females. 51 140 x x , xx * x Q. RIGHT CHELIPED LENGTH XX 120 o ' * 100 *? X X xx$*X X * % • *• 80 xw* * , .-*?• *x **♦ .• . v 60 x>c* •• x «**.' . • • 4 xx .»> • I*7' 4 0 •5 • 20 -I— ~1 ----- i 1 i |----- 1----- [----- r 140 b. LEFT CHELIPED LENGTH # 100 ¥ < V * • 80 ¥ xxx Q x • • ~A . •• 60 X X ,.!.*! x x Av X x I • • • *♦ X .*, 40 X j * . X 20 — , | ,---- !---- 1---- [---- 1---- 1---- 1---- 1---- 1 I I I r— ) 10 20 3 0 40 50 60 70 80 SHORT CARAPACE WIDTH (mm) FEMALE RIGHT CHELA LENGTH (mm) ro -p> CD 00 O o o O O o X o X 8 “ H o X m OJ o xx r* m z CD - t X x 2 Ol _ x o O X x H O X m CD. >k>x r O *o m o r O &• m X z o ••• H 2 X 00 O' HO Q. O. o o o x< X >*< IN). o "1 ro CD oo o o o MALE RIGHT CHELA LENGTH (mm) FEMALE LEFT CHELA LENGTH (mm) ro -t* m oo o o o o o o o a. -n m o r m z CD H X HO »< 3 3 oo MALE LEFT CHELA LENGTH (mm) 6 0 - 30 x e. LEFT CHELA DEPTH . * 4 * 1 50- x / X* Kx 1-20 m m i x d X X X X ^ 4 0 - x** xxk *“ x HO x x * X 30- - 0 MALE CHELA DEPTH (mm) 2 0 - 1 • .,«r» * • • p v 10- ,-.ii >f- V »v 0- 1------1------1------1-----1------1------1------1------1------1------r------1------1------r 30 f. RIGHT CHELA DEPTH ♦ • •« • • • • * 20 •X 10 c f . . . ? # ' 9 * * 0 1 ! , | i | i | i | i | i p 10 20 30 40 50 60 70 CHELA LENGTH (mm) Figure 8 Ventral series of male Bathynectes superbus; measure ments indicate short carapace width. Upper row, left to right: 74.7 mm, well-developed gonad 63.7 mm, well-developed gonad 55.2 mm, well-developed gonad Lower row, left to right: 50.3 mm, well-developed gonad 42.2 mm, moderately developed gonad 27.7 mm, undeveloped gonad 52 rj—w 5 03 Figure 9 Ventral series of female Bathynectes superbus showing abdominal shape change with size (short carapace width) and gonad development. The abdomen shape changes from triangular to semi-circular in crabs approximately 30 mm in width. Upper row, left to right: 62.2 mm, very well-developed gonad 61.0 mm, moderately developed gonad 54.5 mm, moderately developed gonad Lower row, left to right: 47.8 mm, very well-developed gonad 41.2 mm, very well-developed gonad 31.3 mm, slightly developed gonad 53 Figure 10 The morphometric relationships of abdominal segment measurements for male and female Bathynectes superbus. Male and female scales are staggered. a. The relationship of the width of abdominal segment 3 to short carapace width. N = 165 males, 178 fe males. b. The relationship of the width of abdominal segment 6 to short carapace width. N = 165 males; 179 fe males . 54 FEMALE ABDOMINAL SEGMENT WIDTH (mm) 0 3 0 4 50 0 2 30- 20 25- 15- 10- — 5 - . BOIA SGET 3 SEGMENTQ. ABDOMINAL . BOIA SGET 6 SEGMENT ABDOMINAL b. ,•/x 20 xx HR CRPC WDH (mm) WIDTH CARAPACE SHORT 0 0 50 40 30 ;xx tr 60 70 xx 80 - 15 - -20 0 -3 -10 -10 25 30 20 MALE ABDOMINAL SEGMENT WIDTH (mm) 55 carapace width (Figure 10 b; Table 10). The relationship between short carapace width and penis length was linear over the size range examined (Table 9) but variability was high (Figure 11). A plot of lateral spine length on short carapace width was too variable to merit formulation of a regression equation. The relationship between lateral spine length and body size is therefore not clear. The variability is apparent in Figures 12 and 13. Very small crabs were observed to have longer spines relative to overall body size than larger crabs. Although allometric growth may be present, the variability observed in spine length could be related to regeneration of the spines after damage. Figure 11 The relationship of penis length to short carapace width for Bathynectes superbus. N : 52 males. 56 LENGTH (mm) PENIS SHORT CARAPACE WIDTH (mm Figure 12 Dorsal series of female Bathynectes superbus showing fifth anterolateral spine variation with size. This spine is usually longer relative to body size in young er crabs. Measurements indicated refer to short carapace width. Upper row.,- left to right: 62.2 mm, very well-developed gonad 61.0 mm, moderately developed gonad 54.5 mm, moderately developed gonad Lower row, left to right: 47.8 mm, very well-developed gonad 41.2 mm, very well-developed gonad 31.3 mm, very slightly developed gonad 57 Figure 13 Male dorsal series of Bathynectes superbus. Measure- ments indicate short carapace width. Upper row, left to right: 74.4 mm, well-developed gonad 63.7 mm, well-developed gonad 55.2 mm, well-developed gonad Lower row, left to right: 50.3 mm, well-developed gonad 42.2 mm, moderately developed gonad 27.7 mm, undeveloped gonad 58 59 Discussion All of the morphometric relationships were statistically linear, but the growth of all body parts was not simply iso metric over the entire range of sizes considered. Allometry describes the growth of a measured body part that is found to be more or less rapid than that of the reference body part whereas isometry describes equal growth rates between the measured body part and the reference body part (Simpson, Roe, and Lewontin, 1950) . In this study, I have considered re lationships to be isometric if they are described by a slope value equal to one, positively allometric if the slope value is greater than one and negatively allometric if the slope value is less than one. Although regressions may be con sidered statistically linear, slope changes may be present which suggest some deviation from simple allometry. In this case, there is allometry of growth rate,I.e. , the two body parts grow at different rates and the rate of growth of at least one part changes with size of the organism. In the morphometric analysis of B. superbus, it was found that the relationship.between:;short,carapace width, the reference measurement, and long carapace width was nearly isometric, whereas the relationship with short carapace length and long carapace length showed slight negative allometry. The growth of body depth, fronto-orbital width and frontal width were negatively allometric. This indicates that the ratio of the dimensions of the carapace, except for 60 long carapace width are not constant over the size range examined. There were sexual differences in the growth of the carapace of B. superbus but no indication that there were puberty-related changes during growth of the crab, as exemplified by an abrupt change in slope for the size range examined. Among the Portunidae, Newcombe et al. (1949) described an allometric pattern of growth for the carapace dimensions of Callinectes sapidus but the authors lacked data for the middle size range of the growth curve. Allo metric growth of the fifth antero-lateral spines with size and age of B. superbus has been reported (Bouvier, 1940; Capart, 1951). In young specimens, these spines are much longer relative to carapace width than in older specimens. It has also been reported that length of the frontal teeth increase with size increases of crabs (Christiansen, 1969; Norman, 1891). The frontal teeth become more distinct and more acute with age. These growth patterns do occur but are apparently not related to sexual maturity or a molt of pub erty. In this study, carapace growth is simply allometric and no discernible discontinuity in growth rate occurs. The male cheliped lengths show positive allometric growth and there is a slight change in slope for the linear graph of both chelipeds at approximately 50 mm short cara pace width. Both chelipeds of female crabs grow allometrically, and the left-Gfieliped^showslsomeievidence^Qf^a^slope _ change at approximately 55 mm short carapace width. The growth of the chela propodus is negatively allometric for the size ranges examined. Since differences m the regression coefficients were present for males end females, there are sexual differences in cheliped size, for a designated carapace width. Shape, which is determined by chela propodus depth does not change with sex for the right chela depth and is only significantly different in variability for the left chela dapth. Both chela depth growth rates were negatively allo metric. The alteration in size but not shape has been re ported for other Brachyura (Hiatt, 1948; Knudsen, 1960), the xanthid Neopanope texana sayi (Swartz, 1972) , Ocypode ceratophthalmus (Barrass, 1963; Haley, 1973), Cancer pagurus and C. magister (MacKay, 1943; Weymouth and MacKay, 1936). It has been suggested that dimorphism of the cheliped is adaptive for use in agonistic displays (Swartz, 1972), but this has not been observed in B. superbus. The growth of abdominal segments 3 and 6 demonstrates true sexual dimorphism. Abdominal segment 3 in females was larger than that of male crabs for any given size (carapace width) above 30 mm. Abdominal segment 6 was totally different for males and females throughout:the entire size range examined. This segment showed a strong slope change in females whereas the slope of the line for male abdominal growth was constant throughout the size range examined and demonstrated negative allometric growth. There was a dis tinct pubertal change in the width of abdominal segment 6 for females. Abdominal growth in female crabs <30 mm short carapace width is apparently allometric, as is growth above 62 the slope change at > 30 mm. The change at a width of 30 mm correlated well with the size at which sperm id first observed in the spermathecae. An alteration in growth of the female abdomen at 30 mm is associated with increased setation of the pleopods and widening of the vulvae. These abdominal changes are adaptive since egg production could not occur without alteration in pleopod structure and the shape of the abdomen and vulvae. Swartz (1972) also noted that since fecundity is exponentially related to carapace width in Neopanope texana sayi, positive allometric growth of the abdomen is adaptive. There is some indication that this adaptation occurs in B. superbus. There is no distinct abdominal shape change in males, and growth with maturity is much less distinct in males. In males, the growth of seg ment 6 may be associated with sexual maturity since a slight slope change was observed at 30 mm, the size at which sperm and spermatophores were present in the reproductive system. Therefore, growth of segment 6 in males and, expecially in females, suggests a pubertal molt in which a discontinuity in slope occurs and growth continues allometrically after the discontinuity. The pubertal molt has been postulated for Macropodia rostrata (Perez, 1929), Ocypode quadrata (Haley, 1969) and Portunus sanguinolentus (Ryan, 1967 a). SECTION III REPRODUCTIVE BIOLOGY OF Bathynectes superbus(Costa) Introduction Knowledge of reproductive anatomy and histology are essential to the understanding of certain aspects of brachyuran life history, such as age of sexual maturity and reproductive potential. Gross and histological anatomy of reproductive organs has been described for certain species of Brachyura (Fasten, 1915, 1917, 1918, 1926; Williamson, 1904; Pearson, 1908; Binford, 1913; Harvey, 1929; Shen, 1935; Broehhuysen, 1936; Spalding, 1942; Demeusy, 1958; Haley, 1967). More recent investigations have considered the relationship between relative growth and reproductive development as an indication of sexual maturity (Haley, 1969, 1973). Anatomical and histological studies on the Portunidae have been confined to Callinectes sapidus and Portunus sanguinolentus. The gross morphology of the male reproductive system in C. sapidus was originally described by Brooks (1882) with subsequent accounts by Churchill (1919), Beaven (1932), Truitt (1939) and Newcombe (1945). The gross anatomy of the penis and male intromittent organs were described in detail by Snodgrass (1936) and Cochran (1935). Cronin (1947) pub- 63 64 lished the first detailed histological study on the entire male system in C, sapidus, The gross anatomy of the female system was presented by Churchill (1919) and Cronin (1942) described the histological development of the female system in immature crabs. Hard (1942) studied ovarian growth and ovulation in the mature blue crab and developed a method for determining the stage in the reproductive cycle from the gross ovarian appearance. Ryan (1967 a, b) described the histology and function of the male and female system of Portunus sanguinolentus. This study describes the reproductive anatomy, histology, reproductive cycle, and relationships between morphometric changes and onset of sexual maturity in Bathynectes superbus. Materials and Methods The crabs used in the reproductive biology study were obtained from several sources (Appendix 2): 138 specimens from June 1973? 6 specimens from November and December 1973; 12 specimens from March 1974; and 14 specimens from April 1974. Gross internal and external anatomy and the gonadal histology of these specimens were examined. Crabs were preserved in 10% sea water-formalin and later transferred to 5% glycerin in 70% ethanol. Specimens were initially examined for pleopod condition, presence of eggs, penis length, condition of the vulvae and for evidence of sexual dimorphism. The internal organs were then exposed 65 by removal of the carapace. The stage of gonad development was estimated by in situ examination according to the scheme outlined in Table 11. Fecundity estimates were made on ovigerous individuals. Abdomens with attached eggs were removed and placed in Gilson's solution (Bagenal, 1967) for at least one month to harden the eggs. The eggs were stripped from the pleopods with for ceps, blotted and allowed to dry for 24 hours. The egg mass was then weighed to the nearest 0.0001 g on a Sartorius analytical balance. Three subsamples were taken and weighed. Each was placed in a Petri dish to which was added enough glycerin to cover the eggs. The eggs were then spread out in the dish and the number of eggs in each subsample was counted. The fecundity index was then calculated by: Total Wt. of Egg Mass x x no. eggs x wt. of subsamples The degree of embryological development within the eggs of each ovigerous crab was observed and assigned to a stage based on the scheme of Meredith (1952). Twenty eggs from each subsample were randomly selected and the longest diameter measured with an ocular micrometer at 40 X. These diameters represent the size of eggs that have been blotted dry and placed in glycerin. Portions of the testes, anterior, median and posterior vasa deferentia were resected. A hanging drop preparation of the excised tissue was viewed with a light microscope. The presence or absence of sperm or spermatophores was noted. Table 11. Stages of gonadal development of Bathynectes superbus(Costa) ■rH ■H g Q o a. G 0 P 0 0 rH G G Q W G £ 0 ft G 0 0 ft 0 > O 0 0 tn ■H •H -H G P o g G P X G G is O 0 o G G G 1— 1 6 0 tn p G 0 P P 0 0 o 0 o p 0 p H P -P p g H r U C G G P P P G (0 O-HO O O tn 0 G E 0 0 0 > 0 G 0 ,, G G 0 "i-L G G rH s G ■H 0 0 G P rH G P P G o G G P 0 0 0 •H > G 0 0 0 P rH 0 G p 0 G 0 0 O 0 1 0 0 0 G G 0 P 0 i— 1 0 G 0 G 0 — tn p 0 > >1 0 1 p G0 rG ■H H 0 •H •H -P G 0 0 o 0 0 P E 0 ft E 0 ft 0 0 P O P 0 0 O £ 0 G U o P i P G H P I -P rH rH rH rH ftG G — 0 P £ G 0 P o 0 P 0 1 X O 0 •H ■H ■H G G G 0 0 0 0 O O G G E tn is 0 p 0 ■H •H i—I 0 E G p 0 0 P o 0 t f 0 G 0 u •H rQ H p •H rH X! ■H p P P P •» G G G o -P 0 G H P •H H G W I p XI q o LO 0 ^ 0 ro O oq H P 15 p 0 -P 0 G 0 £ p > o 0 0 0 > 0 >1p O P I rd 0 0 0 -H 0 0 0 G 0 0 0 0 0 0 0 0 0 G rH 0 0 0 0 X ft GG -H G 0 0 0 G 0 G 0 G 0 0 G 0 G 0 g > £ g 0 t n 0 H H 0 tn ■H M P ^4 -H p E M 0 0 G 0 0 0 0 G 0 P G O P GG h • G G 66 rH rH P 0 G -P G G 0 0 P 0 0 0 P 0 0 0 o G 0 G 0 0 15 H00 0 ■H £ 0 JT* ■H G G rH G P O G E * 0 A*G 0 0 G > £ 0 0 P 0 P 0 P G G 0 0 > 0 P >i G 0 0 0 G 0 0 0 0 G G > 0) CO 0 0 0 rH > 0 h O 0 ■H 0 rH rH G G G G P P 0 0 0 0 O G X E i 0 >i 0 0 G ft 0 G 0 ft t 0 ft 0 O 0 b1 0 ft tn P • O O00O0 O O0 •rl G ■H G G rH G G rH u G p p P 0 0 S 0 G P 0 0 0 0 t f 0 > E G 0 & tn ft 0 o 0 0 0 0 P •H rH G G G G 0 0 p 0 tn nP tn rQ 0 G 0 G 0 . i—1 G G P 1—1 G O G 0 0 G G 0) p 0 G rH 0 P >iG 0 0 0 0 > 0 P 0 tr P 0 0 p £ ft 0 0 ft G b1 p 0 0 P O 0 i—I 0 0 ft H H 0 - > rH 0 P G 0 ft ft O 0 G 0 0 0 O 0 0 t f p 0 0 0 ■» ■rl rH G G p 0 0 0 0 0 G 0 tJ» G 0 P tn tn 0 P 0 g tn ■H G 0 ■H 0 G -H G G H 0 0 -H -H G 0 >i 0 >i 0 G > 0 H -H r p 0 p rH P rH •H P G 0 0 rH 0 0 G > p > H i— p o G > > G I—I G 0 G G G G SP G P ISP 0 £ 0 0 0 P 0 0 0 tn 0 0 >i 0 0 0 0 0 0 0 1 0 &P 0 G 0 0 E G 0 G 0 0 E 0 1 0 G P 0 0 P 0 G P ft > 0 0 >1 P i 0 0 O - p ■H G 0 G 0 G 0 0 0 0 0 GG 0 0 t f 0 0 G 0 0 0 0 P 0 0 > 0 0 tr> G h i—1 0 p O p tn 0 tn • • -h. 0 £ tr* -rlG G 0 P 0 G G 0 P ft 0 p G 67 The widest diameter of 15 spermatophores was measured with an ocular micrometer. Sperm from testicular material was also examined by interference microscopy to determine its morphology in relation to that of other decapods. The ovary and spermathecae of female crabs were handled in a similar manner. The spermathecae were examined for the presence of sperm and 15 ova from each ovary were measured at the widest diameter. The maximum width of the anterior ovarian horns was measured with vernier calipers to the nearest 0.1 mm. Following in situ examination, gonadal tissue was re sected and prepared for routine histological sectioning by dehydration in alcohol and dioxane and infiltration with Paraplast (Fisher Scientific Co. ). Since fragile but hard tissues often crumble when processed by the paraffin method, it was necessary to double embed ovarian tissues with methyl benzoate, celloidin and Paraplast (Humason, 1962). Sperma thecae and mature eggs were treated with a decalcifying agent (Decal, Decal Corp.) in order to prevent fracture of the tissue during sectioning. Tissues were sectioned at 7 jx on a rotary microtome, stained with Harris hematoxylin and counter-stained with eosin. All histological procedures ex cept embedding and sectioning were performed with an Autotechnicon (Technicon Corp.). Prepared tissues were ex amined at 10 and 45 X magnification. Maturity on all specimens was checked by examination of the prepared slides of the reproductive organs. 68 Results Gross Reproductive Anatomy In male B. superbus, the first two pair of pleopods are modified, as in other portunids, to function as intromittent organs in copulation. Figure 14 a shows the insertion of the penis into the endopodite of the curved, wandlike first pleopod. Although copulation has not been observed in B. superbus, the morphology suggests that the first pleopod receives spermatophores and semen from the penis and acts as a plunger to insert these sexual products into the oviducts of the female. The first pleopod is grooved to form a tube like extension in which the sexual products are probably transported. No quantitative data are available on the change in pleopod size relative to age. The pleopods of smaller individuals were, however, less conspicuous than those of adult males. Both the first and second pleopods are fringed with setae. The shaft or telopodite (Cronin, 1947) of the first pleopod has spines along its tip. The penis is a white, flexible organ that originates in the ventral part of the coxopodite of the fifth pereiopod. In all the specimens of B. superbus examined, the penis was always found to be inserted in the proximal foramen of the first pleopod (Figure 14 a ) . When the abdomen is flexed, the penis is protected by the second, third and fourth abdominal segments. The testis of B. superbus is a paired convoluted organ 69 that lies along the anterolateral border between the hypo- dermis and the hepatopancreas, The testis joins the anterior vas deferens lateral to the stomach. The color of the testis varies from colorless in immature crabs to creamy white in mature specimens. The vas deferens consists of three parts. The anterior portion is a mass of small diameter coils located latero- ventral to the stomach. The median part consists of rel atively large white coils (Figure 14 b) situated ventral to the pericardial sac. The posterior vas deferens is a trans lucent white extension of the median vas deferens and is located beneath the heart. The pleopods of female crabs are located on the second through the fifth abdominal segments (Figures 14 c, d ) . Each pleopod is biramous, consisting of an endopodite and exo- podite arising from a basal protopodite. Both the endopod and exopod bear setae, the amount of which varies with stage of maturity. Pleopods of immature crabs are not fully setose whereas those of mature crabs are heavily setose. The ovary is a paired lobate organ located along the anterolateral regions of the carapace, uniting beneath the stomach and continuing ventro-posteriorly to the peri cardium. The coloration and size of the ovary varies according to the stage of development (Table 11, Figures 15 a, b ) . The seminal receptacle or spermatheca is joined to the Figure 14 Gross anatomy of the male and female reproductive system of Bathynectes superbus. a. Male intromittent organs. Note insertion of penis (p) into endopodite (end) of first pleopod. Penis originates from the coxopodite (cox) of the fifth pereiopod. Short carapace width 54.2 mm. b. Coils of the median vas deferens (mvd) in situ. c. Heavily setated pleopods and oval vulvae (v) of a mature female, 49.9 mm short carapace width. d. Non-setose pleopods and slitlike vulvae (v) of an immature female, 38.1 mm short carapace width. 70 Figure 15 Gross reproductive anatomy of ovaries from female Bathynectes superbus. a. Moderately developed ovary (ov) of a 48 mm crab. Note position of ovary relative to stomach (sto). b. Very well developed ovary (ov) of a 50 mm crab. The ovary covers the stomach in this stage of development. 71 72 ovary and oviduct. In the post'-copulatory condition, the seminal receptacle is distended with sperm and the oviduct is often blocked by a sperm plug. In intercopulatory mature crabs, the seminal receptacle is flat and is difficult to distinguish from the pereiopodal musculature. The vulvae located on the sixth thoracic segment, undergo shape changes with onset of sexual maturity. In immature crabs, the openings are slitlike (Figure 14 d) and are difficult to see with the naked eye. In mature forms, the vulvae are much wider and their margins are thickened (Figure 14 c). Histology of the Reproductive Organs Longitudinal and transverse sections of the penis are shown in Figures 15 c and 16 a, b. The gonoduct is con spicuous as the ejaculatory duct which extends the length of the penis. Columnar epithelial cells with an apparent brush border line the gonoduct and form a typhlosole which projects into the lumen (Figure 16 a). External to these columnar cells is a layer of circular muscle tissue. The space between the external cuticle of the penis and the muscle layers is apparently occupied by connective tissue and blood sinuses. The histological structure .of-the testisiLis^shownlin Figure 16 c. Numerous lobules are present and are filled with cells in various stages of spermatogenesis. The walls of the testicular lobes are thin and indistinct but appear to be similar to those described by Ryan (1967 c). Numerous Figure 16 Reproductive histology of male Bathynectes superbus, a. Cross section of penis with gonoduct (g) partly occluded by typhlosole (t), 41 X. b. Cross section of penis. Note muscle layer (m) around gonoduct and columnar epithelial (ce) lining of gonoduct. 41 X. c. Testicular lobules. Sperm appear in area designated by arrow. Granular substance in lumen is likely developing spermatocytes (ds) . Spermatogonial cells are large cells found along the wall of the lobules. 41 X. d. Anterior vas deferens. Developing spermato- phores (dsp) appear in lumen. 41 X. 73 • . ,• ' * . > r * ys -jwa & A ’VSBsetfSte 74 large cells which may he spermatogonia are present along the walls of the lobules. Ryan ( 1967 c) describes two portions of the anterior vas deferens. Only the second portion was effectively dis cerned in B. superbus. Sperm, developing spermatophores and completed spermatophores were present in the lumen (Figure 16 d). Developing spermatophores consisted of clumped sperm surrounded by a thin membrane; completed spermatophores were encapsulated by an eosinophilic membrane. The epithelial lining of the anterior vas deferens appears to be columnar. In sections which contain mature spermatophores, a basophilic substance appears in the lumen around the spermatophores. The median vas deferens is surrounded by a thick mem brane and contains mature spermatophores (Figure 17 a). Epithelium of the median vas deferens is low, perhaps cuboidal. The spermatophores are more numerous than in the anterior vas deferens and are covered with a heavy eosino philic coat. Numerous basophilic granules are present among the spermatophores. The posterior vas deferens is divided into two histo logically different parts. The anterior portion consists of high columnar epithelium surrounding a central lumen filled with basophilic granules (Figure 17 b). The more posterior section, which may be what Ryan (1967 c) described as the ejaculatory duct is typified by low columnar epithelial cells around a lumen containing a small amount of eosino philic and basophilic material. Figure 17 Reproductive histology of male Bathynectes superbus. a. Transverse section of coils of the median vas deferens, Mature spermatophores are present in lumen (1). 41 X. b. Section of posterior vas deferens with chromato- genic material (cm) in lumen (1), 41 X. c. Spermatophores from median vas deferens of well developed crab, short carapace width 69.1 mm. Sperm (s) from broken spermatophores appear as smaller entities among larger spermatophores (sp). Hanging drop preparation, 40 X. d. Sperm from the testis. Note radial processes extending from nuclear cup and the acrosome which appears as a central depression in the cup. Interference microscopy, 1300 X. 75 in top) 76 The male sexual products, the spermatophores and sperm appear in Figures 17 c and 17 d respectively. Spermatophores ranged in size from 8.8 to 57.2 /l. Size differences were most obvious between developing spermatophores of the anterior vas deferens and mature encapsulated spermatophores from the median vas deferens. Mature spermatophores, as shown in Figure 17c, are surrounded with a distinct membrane which is strongly acidophilic in histological preparations. Sperm are identical to the general decapod stellate type described by Brown (1966). The acrosome appears as a central depression within the nuclear cup which is surrounded by 2- 3 radial extensions. It is not known whether the depression in the acrosome is an artifact of preservation or a natural feature of decapod sperm. Histological sections of a moderately developed ovary are shown in Figures 18 a, b. Although most sections were fractured due to the presence of yolk, it was possible to distinguish a few anatomical features. Developing oocytes are present around the germinative area, A central lumen is present within each ovary, and germinative zones, which give rise to oocytes, are present along the sides of the central lumen. The oocytes and follicular cells described by Ryan (1967 b) are also present in B. superbus. The oo cytes appear as the large cells with a distinct large nucleus. The follicular cells are much smaller cells and are in close proximity to the oocytes, often surrounding them. A histological section of the seminal receptacle is Figure 18 Reproductive histology of female Bathynectes superbus. a. Moderately developed ovary with interlobular septa. Follicular cells (fc) appear within septa. The follicular cells have oval nuclei which are prominently stained. 40 X. b. Longitudinal section of a moderately developed ovary. Note central lumen (cl) with germinative zone containing small developing oocytes. 8.3 X. c. Cross section of spermatheca showing sperm within the lumen. 40 X. f e w W&B!** -IS i t i / v ^ 9 t ' J shown in Figure 18 ct The presence of the sperm plug makes sectioning difficult and most sections were fractured. Numerous spermatozoa are present within the lumen which is surrounded by fibrous connective tissue and blood sinuses. An outer, apparently stratified epithelial layer is overlaid by a cuticle. Hanging drop preparations of ova from a slightly developed and a very well developed ovary are shown in Fig ures 19 af b. The slightly developed ova are much smaller and do not appear to possess a distinct membrane; vitello- genesis is not as advanced as in the well developed female. Size differences are probably a result of variation in yolk development. Figure 19 Ova from female Bathynectes superbus at two stages of gonad development. Hanging drop preparations, 10 X. a. Ova from a slightly developed gonad, 37.7 mm short carapace width crab. b. Ova from a very well developed gonad, 41.1 mm short carapace width crab. 79 2 0 0 JU 200 n 80 Sexual Dimorphism The most obvious means of sexual distinction in larger individuals is by the shape of the abdomen. Male crabs have a triangular shaped abdomen with segments 3-5 fused (Figure 20). In small females, the abdomen is of the same shape but the abdominal segments are not fused and articulate freely (Figure 21). In adult females, the abdomen is semi-circular (Figure 21). In addition, the vulvae of the oviduct and the setated pleopods of females distinguish the sexes. Sexual differences were also noted in the size of the chelipeds for larger individuals. Adult males have larger chelipeds generally than females for a given size. Sexual differences in morphometry are described in Section II, No significant sexual differences in color of the carapace or chelipeds was noted. Gonad Development The gonad development of 70 male and 62 female B. superbus collected in June 1973 varied with size of the crab (Figure 22). No undeveloped or very slightly developed gonads were found in male crabs > 30 mm short carapace width. The gonads of most males > 40 mm were either moderate or well developed. Males: :< 40 mm were in the earlier stages of development, either undeveloped, very slight or slight. The reproductive system of male crabs was never observed to be the dominant internal organ system. This suggests that males in the well developed category (Table 11) are Figure 20 Ventral view of mature and immature male Bathynectes superbus. Upper; short carapace width 74.7 mm, well developed gonad. Lower: short carapace width 27.7 mm, undeveloped gonad. 81 Figure 21 Ventral view of female Bathynectes superbus showing relative change in abdomen shape with size and gonad development. Upper: short carapace width 62.2 ram, very well developed gonad. Lower: short carapace width 31.3 mm, very slight gonad development. 82 c n l 111 III III 1) III 111 111 11 111 11 1 1 1) 1 2 3 SPEC.______DATP 9866 [ * i * 4 ’ 3 SPEC.., -PATT i . ■ Figure 22 Cumulative percentage occurrence of various stages of gonad development in relation to size of male and fe male Bathynectes superbus captured in June 1973. 83 OCCURRENCE OF :GONAD STAGES ( CUMULATIVE PERCENT) - 0 5 50 25 - 0 5 - 5 2 75-1 75~| - 5 2 75-j 0 SLIGHT EY SLIGHT VERY UNDEVELOPED 20 d 060 40 AAAE IT (mm) WIDTHCARAPACE EL DEVELOPEDWELL EY WELLVERY DEVELOPED MODERATE 1 0 060 40 20 — r l ~ - - f % S ■ I --- - - 84 comparable in maturity to very well developed females. Male crabs captured in November and December 1973, March 1974 and April 1974 showed essentially the same pattern (Table 12). The majority of June 1973 females > 40 mm short carapace width had either moderately or very well developed ovaries; none were well developed (Figure 22). The ovaries of most females < 40 mm were either slight or very slight in development; none were undeveloped. The variable pattern exhibited by females captured in November and December 1973, and March and April 1974 was probably due to the presence of more ovigerous and/or spawned out individuals (Table 13). The majority of well developed and very well' developed females were >50 mm. Most crabs in early ovarian development were <50 mm except four individuals taken in November 1973 in the 51-70 mm range that were slightly developed and one undeveloped 54 mm crab captured in March 1974. The three male crabs with undeveloped or very slightly developed gonads captured in June 1973 had recently molted (A^-I^) (Table 13), whereas the majority of male crabs in advanced gonadal development were hardshell (C^-C^): 54% in moderate development and 60% well-developed. Crabs with slightly developed gonads were equally distributed between hard and soft-papershell molt stages. Female crabs exhibited a different pattern. The in cidence of very slightly developed and slightly developed Table 12. Number of Bathynectes superbus from November 1973, March 1974 and April MH & & & hpp mh 'O P i—i -p -P H Pi rH CD ^ ■P CTir- O td -p o u a) w> U 0 o a e rtf rtf o U O CD -P ) O Q H H CD iH P4 £ £ cd 0 0 £ > td CDH td a mh m CO i—1 -P CO -P ■P +J rH g CD g t'-'00 rd CD CM 0 co S 0 Di cd 0 td pi rH 0 > > 0 0 CO Pi a C 0 £ 0 £ CD N > 0 0 *H •0 o in in o i ■ i — n n r- oo O in - r '31 O in O in O ■ — I I I I I 1 i 1 0 (H 00 — i rH id i— i r~ i —i 8 5 CD00 H rH Q CM Pn £ £ CD P CD £ d > td CDiH 0 0 —. — O LO Or 0 rH 00 rH CO rH in o — N g 1 i 1 r—1 n i - oo r-' in in O O O O H H H H rH rH rH I i i I I i— H i rH O D O'- CD UO — i i 1 — 1 i— i LO 1—1 -M C 'sf CD e & o CD rW i—1 CD co CN td > 6 o LO 1| +> c CD £ &CO CN CN O tH CL) > CL) CD H Q CN CN CN rd S M-l 0 CD O'11—1 CN rd ■P • CO +J C 0 o o '— r-' CM CTi 1—1 i—l g h £ CD O O O O i—1 CO LO LO A Qj N 1 t 1 1 td r—1 rH i—1 pH Eh ^ CO c n ro l o 86 m G X CM P pq O rH O 0- 87 88 ovaries in hardshell and soft^papershell females was equal. The majority (55%) of females with very well developed ovaries were nearly equally distributed between hard (53%) and soft-papershell (47%), Specimens from November, December, March and April showed variable patterns which may be due to seasonal changes (Table 14). Since so few in dividuals were captured, however, a definitive conclusion concerning relations between molt stage and gonad develop ment are difficult to make. The condition of the spermathecae relative to size of 62 non-ovigerous B. superbus is found in Table 15, Crabs with undeveloped spermathecae ranged in size from 21-50 mm. Seminal receptacles were also detectable but uninflated in crabs throughout the entire size range. Only crabs > 31 mm had swollen spermathecae, an indication of recent copulation (Terretta, 1973). The relation between the stage of gonad development and the condition of the seminal receptacle is summarized in Table 16. The ovaries of most (7 5%) crabs with undeveloped seminal receptacles were only very slightly developed; none were more than moderately developed. Crabs with flat spermathecae, which may indicate individuals that are mature and have not copulated or have not copulated recently, were associated with every stage of gonadal development except undeveloped. Flat spermathecae were most often associated with ovaries in slight and very well developed stages. Swollen spermathecae were most frequently associated with moderately developed ovaries. Table 14. Relationship of intermolt stage to gonad development in Bathynectes superbus captured during November 1973, March 1974 and April 1974. H CM 91 o\° ^— 0 £ 03 G 0 1 1 1 0 G i—1 1 1 VO CT> 1 ro o G |Q> i—I 1 1 CM in 1 i—! CO O 0 tn G 0 £ ■H 0 03 O 03 0 ■P 0 0 P rG 0 G •p > -P 0 ^ ■H g <*P -P a p 1 LO LD CTi o r - G\ 0 0 0 -p 1 CM VO CM o 00 in H 0 a 0 1 i—i 0 C/3 03 rH P G a .G m 0 P 0 0 0 O Qa G 0 G o — rG w •H o\P -P ■P — 0 c/3 ■H •H >0 e 0 P 03 0 P 3-i a g a 0 0 0 o 1 I 1 CU 0 U H 1 LO < T \ CM I 1 H m G nO 0 1 i—1 i 1 i—1 G > 0 0 0 rG 03 ■P G D m ov O G rG O 0 w ■H p rQ -P 0> 0 0 -H -H S P *0 > u G 0 - | O I ro MH 1 ^ CO t-" CN in i—1 0 G ["■ 0 1 CN H i—i 0 c-\ 0 G rH rG 0 -P ip s 0 MH O ■P G 0 0 0 g 03 G a s 0 0 o a 03 i—i *»■ 0 ■H 0 r o rH 0 > i> 03 03 0 G 0 o\ 0 0 > H 03 rH G a 0 0 -P 03 4-1 0 Q 0 G 0 JG rH 0 a 0 rH VO «h g 0 -H 0 > rH i—) o a rH r-l -P 0 0 0 0 C/3 -P 0 Q !S 0 i-t > rG P iH 0> 0 0 N tn 0 t— 1 >1 (tf rQ 0 > 03 P •H 03 rH P +3 0 -P 0 G 0 i—1 0 92 93 The condition of the seminal receptacle was associated with intermolt stage (Table 17). Crabs with undeveloped seminal receptacles were associated with every molt stage except soft. However, only one soft crab was captured and the seminal receptacle was flat. Flattened seminal receptacles were observed in crabs in every molt stage but were most frequently associated with the hard (C-^-C^) and peeler (D2-D4) stages. Most (63%) of the papershell (A2-B2 ) crabs had swollen spermathecae. The seminal receptacles of non-ovigerous crabs captured during June, March, November, and April were flat in appearance. The gonadal condition of ovigerous females ranged from undeveloped to moderate and all the females were hard. The size of the ovigerous females ranged from 41-63 nun short carapace width (Table 19). Spermatophores were not found in males with very slightly developed testes and vasa deferentia (Table 19). Crabs with spermatophores in the vasa deferentia ranged from 28-75 mm short carapace width. There was a higher incidence of sperm associated with larger and more developed crabs. The relation between spermatophore size and gonadal development is sum marized in Figure 23. Crabs with slightly developed gonads had spermatophores that ranged from 30-36 ji. Spermatophores from moderately developed crabs ranged from 10-42 )i whereas well developed vasa deferentia contained spermatophores that ranged from 28 to >48 /a* Analysis of variance showed significant differences in spermatophore diameters (F = 4.8; 0 tn rd o\° P — CO d 0 1 1 P 00 p cn i i oo p r—1 p 1 I CD 00 0 cn 0 g H 0 £ td cn 0 (D o P d 0 d P! 0 173 P > rd ■H CO g o\° P £ Sh — CN CH O CD O ftf -H 0 P 00 cn O CN CD i—1 P a rd 1—1 0 d CO P shd fa p 0 d 0 rd 0 o u d 0 d 0 — pi p -H o\° p & • P rd rd ■H d g 0 d 0 d CO d Qi 0 COi—l 0 0 1 di d U P cn oo i p cn CO Pi I—1 0 1 rH U •H > 0 0 u 0 Pi Ot a d p d c d CO D p d 0 CO d 0 rd d p 0 d CO •H 0 t-" p +»:c CO td •H >irH SH d x: u d P Pi O rd o p oo oo p ^ O PQ u 0 CN P CN VD rd 0 co a • Pi d 0 P 0 ** 53 p 00 p 0 l> 0 tom ■H ■—f 0 > P 0 0 u P d i 0 0 0 d jQ g d o g Sh •h d 0 0 u > P d p 0 d H 0 3 H d • rd d p d >i 0 Pi 0 ^ ^ CN p a 1—1 o to c D B PJ rd rd i t i p rd U P P CN P CN 0 c D A T EH pq cn A 94 I > Mh £ o 0 £ 0 ft + + tn o W PQ PQ 0 i—1 + I I I ■p 0 < P < ! < ! < ! PQ PQ c/1 > 0 p p 0 -p 0 £ 0 . m cn ro o r- CO H 4 cn lx> ■rH m -p P s .Q P 0 ^ ro ro 1 1 c-» r- i—1 i—I i—1 0 tn tn • • 1 1 • •«•• ft CD CN 1 1 CD o in CO rH 3 0 iH i—1 01 a Ol 01 0 -P O H 4 CL) tp 0 ro r- ro r- O CO ro ro CO fi O P e' cn r- cn cn r- r- e' e' >i p en rH cn i— l i— l Short Carapace Gonad Development Width Slight Moderate Well Developed 28-30 3 31-33 2 34-36 1 37-39 3 40-42 5 43-45 1 2 46-48 1 6 1 49-51 1 7 1 52-54 1 15 2 55-57 4 1 58-60 1 2 61-63 3 2 64-66 2 3 67-69 7 70-72 73-75 1 96 Figure 23 The incidence of Bathynectes superbus in each stage of gonad development in relation to the mean diameter of spermatophores (from hanging drop preparations) observed at each stage of gonad development. 97 SLIGHT 3 - 2- 12- co MODERATE < ,0“ z> 9 8 - > o 6- z 0 * 2 " iLl 1 o-J Z> 5-i WELL DEVELOPED 4 - 12 24 4836 SIZE OF SPERMATOPHORES (/<) 98 2/58 df) between slight, moderate and well developed vasa deferentia (Appendix 4). The width of the anterior ovarian horns was directly proportional to the stage of gonad development (Figure 24). Analysis of variance showed significant differences between the width of the anterior horn and each stage of gonad development observed (F = 44.8; 4/64 df) (Appendix 5). All females with horn widths greater than 8 mm were very well developed. Under 8 mm, some overlap of sizes did occur. The mean size of ova are directly proportional to the stage of gonad development (Figure 25). Crabs with very slight gonad development had ova that ranged from 30-120 ji. Slightly developed gonads contained ova from 50-170 yx whereas ova of 100-180 ju were predominant in moderately developed ovaries. Ova from well developed gonads were within the 230- 240 range. Very well developed gonads had the largest ova (200- >360 M) • Ova were found to show highly significant size differences between the stages of development when tested by analysis of variance (F = 75.6; 4/59 df) (Appendix 6). The size of 29 non-ovigerous B. superbus (pooled from all cruises) with sperm plugs present in the vulvae and ovi ducts ranged from 33 to 68 mm. Data were pooled because there were an insufficient number of crabs collected to per mit analysis on the basis of seasonality. Histological examination of seminal receptacles revealed that spermatozoa were present in 17 females that ranged from 28-61 mm. In several cases, sperm plugs were present in the vulvae but the Figure 24 The maximum width of the anterior ovarian horns and the incidence of female Bathynectes superbus in each stage of gonad development. 99 VERY SLIGHT 10 SLIGHT 8 - 4 - tn _ < 2 - 3 O — > o z MODERATE 8 - u. O cc 6 - UJ CD 2 3 Z 2 - WELL DEVELOPED VERY WELL DEVELOPED 2 - 2 4 6 8 10 12 14 16 18 ANTERIOR HORN WIDTH (mm) Figure 25 The incidence of Bathynectes superbus in each stage of gonad development in relation to the mean diameter of ova at each stage. 100 CM — O — CM r r VERY SLIGHTM 1 - P - O inCO s"ivna ro m c — iaiqni o UJ Q LU jo awnN n w ja a O J o — OJ r r o L 5 L U O CL Q LU _1 > l l I I 1 o cm ro I I LJ LU O LJ LJ L > > > 3 CL Q 01 l I O — b-CM .o 'M- •o .o ■ID o CM o o o CM O CM O CD O O CM O CM oo CO O CM o MEAN DIAMETER OF OVA ( ^ ) 1 0 1 seminal receptacles were not detected during dissection. This was probably due to poor preservation of the crabs. Fecundity The majority (5) of ovigerous females were captured during November and December 1973, One ovigerous female was captured in June 1973, One was captured in March and two in April 1974. Fecundity estimates of the nine ovigerous females ranged from 20,924 to 192,250 eggs (Table 19). Regression analysis of the relation between short carapace width and fecundity indicated that the slope was not signif icantly different from zero at the 0,05 significance level (F = 5.1884; 1/6 df). The ovigerous females from June, November and December 1973 and March 1974 were carrying eggs in the early stages (A-B) of development. The two females captured in April 1974 carried eggs which were advanced in development (D-E). Eggs from ovigerous females were in six stages of development: A+, A, B-B+, D, D-E (Table 19). Analysis of variance showed a significant difference between egg diameters at each stage of development observed (F = 3.8 997; 2/24 df) (Appendix 7). The frequency distribution of eggs in both the A-A+, B-B+ stages were bimodal (Figure 26). Eggs in the A-A+ stage of development were most abundant in the 370-380 p and 450-480 p ranges. Modes of 390-400 p and 490-500 p were apparent in the B-B+ stage of development. The D-E stage eggs were most numerous in the 350-360 ju range. Eggs in the D-E stages were frequently fragmented with only the egg mem- Figure 26 Size frequency distribution of eggs at various stages of development (Meredith, 1952). Egg sizes are grouped by 20 )i intervals. 102 PERCENT OCCURRENCE OF EGGS 20 20 n 28-1 20 24- 12 16- 12 12 8 - 4 8 - 4 8 350 330 - 3 7 650 370 330 ------7 610 370 390 410 G DAEE (^) ^ ( DIAMETER EGG 450 7 9 . 630430 . 590 470 490 9 530 490 1 550 510 530 570 570 - BB- + A-A + 610450 670 650 membrane left, indicating that some eggs had hatched. This could account for the smaller size of countable eggs in the more advanced stages of development. Twelve females ranging from 40-68 mm short carapace width had egg remnants on the pleopods, evidence of a recent spawn. Two of these crabs were in the slight and very well developed gonad condition. 104 Discussion Reproductive Anatomy The male and female reproductive systems of B, superbus are morphologically and functionally similar to those of Callinectes sapidus and Portunus sanguinolentus, Spermato phores are formed in the anterior vas deferens and are fully developed by the time they reach the median vas deferens. The posterior vas deferens in Portunus sanguinolentus produces seminal fluid. Histological sections of B. superbus show that the lumen of the posterior vas deferens is filled with a granular basophilic substance which is apparently different in constituency from the eosinophilic substance noted by Ryan (1967 c ) . Ova increase in size through the accumulation of yolk in vitellogenesis. The seminal receptacle functions as a storage organ for sperm following copulation with the male. Sperm plugs which are formed from secretions by the male are found within the oviducts and extend into the seminal receptacles. The mature sexual products of Bathynectes, spermatozoa, spermatophores and ova are indicators of development and stage of sexual maturity. The sperm of B. superbus resembles that of other portunids which differs from the sperm shape found in other crustaceans. It has been recognized that crustacean sperm does not resemble other invertebrate or mammalian sperm types (Kolzoff, 1906). Modification and specialization of the spermatozoan components are generally 105 found in Crustacea. The amoeboid sperm of the cladoceran, Polyphemus: pediculus (Zachariasf 1884), the motile sperm of the barnacles (Kolzoff, 1906), the non-motile filament-like sperm of isopods (Reger, 1964), the non-motile "Strahlenzellen" (Brown, 1966) forms of the decapods (Bloch, 1935) and the elongate forms of the ostracods (Loundes, 1953) are examples. In general, decapod sperm are rounded and possess several radial processes. Bathynectes superbus sperm does not differ from the general decapod type and consists of a cen tral nuclear cup surrounded by 2-3 radial processes. Sexual Dimorphism Sexual dimorphism of B, superbus is similar to that of other crabs. The shape of the abdomen is the most obvious means of distinguishing the sexes. In addition, the di morphism in pleopods and the presence of oviducts on the female thoracic segments differentiate the females. Sub stantial changes in these characters occur with maturity in the female. Abdominal shape changes from triangular in juveniles to semi-circular in mature females. The pleopods become heavily setose in mature specimens and the vulvae of the oviducts change from barely noticeable slits in immature crabs to large ovals in mature crabs. Although cheliped sizes are slightly different between the sexes, the lack of a marked difference in the color pattern of the chelipeds suggests that no social display is associated with the chelipeds, 106 Gonad Development The collections of ovigerous females suggests a long spawning season for B. superbus with some element of the population being ovigerous year round. Most of the ovigerous females were captured during November and December whereas only one was captured in each of the months March and June. The extruded eggs of these crabs were in early stages of development. Large collections of B. superbus made in November 197 4 on the R/V Gilliss ,-but not included in this study, contained females which were ovigerous or which had very well developed gonads and were perhaps just ready to extrude the ova. This collection supports the hypothesis that the primary period of egg extrusion occurs in November and December, Two ovigerous females captured in April carried eggs in advanced stages of development. Roberts (1969) reared zoea from an ovigerous female of advanced egg development that was captured in February 1968. This suggests that crabs within the population may hatch eggs at different times or that females may release ova more than once. This latter hypothesis is supported by the presence of egg remnants on the pleopods of females with very well developed gonads and the presence of sperm in the seminal receptacle of very well developed females( which may also be true of females that have not extruded the first time). Sanders and Hessler (1969) have produced evidence that the isopod Tlyarchna sp. and the bivalve Nucula cancellata reproduce syschronously. This is apparently true also for 107 B. superbus since there was an indication of copulation for every season. Maturity within female crabs apparently occurs at > 28 ram short carapace width, based on the presence of sperm within the seminal receptacle. The presence of the sperm plug, a crystalline body composed of secretions by the male (Hartnoll, 1969; Ryan 1967 b) and which intrudes from the vulvae into the seminal receptacle, indicates that copulation has occurred. Sperm plugs were not found in females < 3 3 mm short carapace width. The marked change in abdominal shape at approximately 30 mm suggests a molt of puberty and substantiates the idea that maturity occurs at this size. Analysis of maturity of females, based on gonad size and color, is difficult and perhaps invalid since females may spawn more than once. There is also evidence that vitello- genesis does not occur immediately after mating. Crabs with swollen spermathecae, an indication of copulation, generally had moderately developed gonads although several crabs were slightly and very well developed. This suggests a period of ovarian growth and development after copulation. Females with slightly developed gonads may be undergoing a second period of yolk development following a previous spawn. The amount of yolk within the egg obviously increases with the advancement in gonad development since egg diameter increases. Similarly, the width of the anterior horn increases due to proliferation and enlargement of the ova by vitellogenesis. Very few data on seasonal gonad changes are available at 108 present. In June 1973, the ovaries of most females were slightly developed and were <45 mm short carapace width. In November and December 1973, the females were in both very slight and slight stages of gonad development. In March and April, crabs were in various stages of development from slight to very well developed. The wide range of gonad conditions observed are probably due to the presence of ovigerous or spawned out individuals within the population. Mating apparently occurs when the female is soft since sperm plugs were found only in soft and papershell crabs. This is similar to spawning behavior in Cancer irroratus (Terretta, 1973), Callinectes sapidus (Rathbun, 1896) and Portunus sanguino1entus (Ryan, 1967 b). Year-round sampling is needed to determine when the majority of soft-papershell individuals occur and when copulation is most frequent. Male crabs reach sexual maturity at approximately the same size as female crabs. The concept of maturity in males is based on the premise that a crab is usually mature when it enters the intermolt stage during which it is able to copulate successfully (Hartnoll, 1969). Among male crabs, the presence of spermatophores or sperm within the reproduct ive system is a useful criterion for maturity, Male crabs > 28 ram were considered mature; these had slight, moderate or well developed reproductive organs. The majority of sexually mature male crabs were hard crabs with the exception of equal numbers of hard and soft-papershell at the slight stage of development. 109 Mature male crabs are present in the population all year long. In June, most males were mature with either mod erate of well developed gonads. Similarly, most crabs in November, December, March and April were mature. There may be a relationship between stage of gonad development and the size of spermatophores, However, spermatophore size varies according to site, i. e. developing smaller spermatophores are found in the anterior vas deferens whereas mature large spermatophores are found in the median vas deferens. The percentage of mature spermatophores may be higher in well developed males than in those with slight development. Fecundity The fecundity estimates for ovigerous B. superbus suggest a high variability in, and a generally lower number of, the number of eggs per sponge in comparison with that of near shore species such as Callinectes sapidus and Portunus sanguinoientus. Callinectes sapidus has been estimated to carry 2 million eggs (Davis, 1965); the number of eggs carried by P_. sanguinoientus ranges from 0.96 to 2.25 million (Ryan, 1967 b). Because variability in egg numbers between female B. superbus was considerable, there is no indication that larger individuals produce more eggs. Davis (1965) postulated that increase in egg size was due to osmotic up take of water. Some of the variability in egg size of B. superbus with stage of development can perhaps be explained by the method of drying eggs prior to weighing. Although n o all eggs were dryed the same length of time (24 hrs,) before weighing, moisture content undoubtedly varies from sponge to sponge and could account for the unusual weight differences. Shrinkage of the eggs would occur due to loss of water. Loss of the eggs during capture and processing also increased variability of egg number. The difference in egg production between portunid species may reflect habitat differences or size differences but it is most probably a result of both. The smaller size of female B. superbus in comparison with Callinectes and Portunus would result in reduced egg number. Abdomens are smaller and the visceral cavity is reduced in this species relative to Callinectes and Portunus. If indeed increased size is positively related with increased fecundity, then lower relative fecundity for B. superbus is explainable. However, both C. sapidus and P. sanguinoientus occur in neritic zones that are subject to temperature and salinity changes. Since hatching in C. sapidus and other estuarine decapod crustaceans is accomplished by osmotic swelling (Davis, 1965), it is likely that alterations in salinity and temperature could lower reproductive potential in the species. Increased fecundity would be one adaptive solution to this problem. The shelf edge-continental slope environ ment of B. superbus represents a more stable thermal and haline environment. If maximum numbers of viable individuals hatch, the energy conservation of the female with regard to egg production is obvious. Ill Generally, crustacean species which produce large eggs that are few in number have a direct development in which the larvae do not undergo metamorphosis to attain the adult form or the eggs are brooded. Extruded egg sizes in B. superbus are larger than those reported by Davis (1965) for C. sapidus and Ryan (1967 b) for P, sanguinoientus. Bathynectes eggs ranged from 350-500 p depending on the stage of development, whereas C. sapidus eggs averaged 273 p x 263 p in early development and 320 p x 27 8 p just prior to hatching. P. sanguinoientus eggs were 280 p , Among other decapod crustaceans, Marshall(1954) mentions that the deep- sea shrimp Giyphocrangon sculptus, G. longirostris and Sclerocrangon (Metacrangon) agassizii produce large eggs ( 0.25-0.32 cm in diameter), By contrast, the estuarine carideans Crangon septemspinosa and Palaemonetes pugio produce much smaller and more numerous eggs. Yonge (1955) noted that the bathypelagic shrimp, Pasiphaea sirado, pro duced eggs that were few and very large. The portunids, in contrast to the shrimp mentioned, have larvae that meta morphose. Bathynectes superbus has 5 zoeal stages (Roberts, 1969) and does not undergo direct development. Compared to other portunid species studied, slightly larger egg sizes and lower fecundity in B. superbus could reflect an adaptation to life in a more stable environment. Larger eggs may also reflect increased yolk and lipid content which may serve to buoy eggs upwards in the water column as Herring (197 4) noted for some meso-and bathypelagic shrimp. SECTION IV FEEDING HABITS OF Bathynectes superbus (Costa) Introduction Studies of feeding habits in marine decapod crustaceans have been limited to commercially important species (Tagatz, 1968; Belding, 1930; Mead and Barnes, 1904; Scarratt and Lowe, 1972; Ropes, 1968; Doflein, 1904; Gray, 1969?; and Herrick, 1896). Relatively little work has been done on offshore, demersal species (Marshall, 1954; Menzies, 1962; Barnard, 1962). This qualitative analysis of the stomach contents of Bathynectes superbus provides information con cerning its nutritional adaptations for life in an archi- benthic environment. Materials and Methods Specimens for stomach analysis were obtained from several sources: 132 crabs from the R/V Columbus Iselin cruise in June 1973; 6 from the R/V Dan Moore cruise in November and December 1973; 12 from the R/V Albatross IV cruise in March 1974; and 14 from the R/V Eastward cruise in April 1974 (see Appendix 2 for station data). All crabs were initially preserved in 10% sea-water 112 113: formalin and later transferred to 7 0% ethanol and 5% glycerin. Sex and molt stage were determined for each crab. The stomachs of the crabs were excised and their contents re moved, examined and sorted under a dissecting microscope. Forage organisms were identified to species whenever possible. The number of stomachs with a particular food item was noted and a visual estimate of food volume in relation to total stomach volume was made. Results Of the 132 stomachs of the June 1973 sample examined, 106 contained food. Most of the hardshell crabs (91%) and papershell (A2-B2) crabs (88%) were feeding. The stomachs contained variable amounts of food ranging from empty to full. The greatest percentage (11%) of empty stomachs was found among peeler (D^-D^) crabs. When food was present, the sto machs contained only trace amounts of food. None of the soft crab (A^) stomachs contained food. Stomach contents were frequently in a semi-digested amorphous state and individual prey items were hard to identify. Amorphous matter predominated, since it included all unidentifiable triturated material (Table 20). When species identification was not possible, items were assigned to a general taxonomic group. The principal prey were poly- chaetes and arthropods (Table 20). Mollusk shell fragments, foraminifera, sand grains and compacted mud were present in t" o r" Cl in ro 00 ID o\o i • • •• • • • •• 0 CD ID 1—1 pH in 00 CM ro ro Q XI 1—I ro ID CM 1—1 rH I 0 i—I P Q . U —'ro f" rH • CM ID 00 rH i— 1 uo ID ro in 00 P c i—I O ro CM CO ro CM pH CD rH < 2 i—I 1-1 CD 0 CD G a P ■3 O ID ID -a* O Cl CO c ••• • *• • • • • g 00 H i—1 in 00 ■3* ID ^ 0 ID ro rH H o P 1 m CM r ~ CQ u 0 I • 3 CM O c in in i—1 o ■—I rH ro ro ro O X! < 2 m CM CM i—1 i—1 Td P p 0 td a 43 3 0 o M-t o 0 m 0 cn ■p ■P o g 0 CD G Three of the four stoamchs of hard crabs captured in November 1973 contained amorphous semi-digested material, arthropod fragments and foraminifera (Table 21). The one papershell crab examined contained unidentified arthropod parts. The volume of food in the stomachs ranged from 1% of capacity to full. Six of the 12 stomachs examined from the March 1974 sample contained amorphous paste, arthropod parts and sand grains (Table 22). Stomachs from two of the three peeler crabs contained food. The volume of food in the stomachs ranged from empty to full. Five of the 11 stomachs examined from the April 1974 sample contained either amorphous matter, polychaete or asteroid fragments (Table 23). The latter were identifiable by bits of pedicellariae and tube feet. All of the stomachs contained only trace amounts of food, less than 5% of the total stomach volume. 01 a) -M • 1 o o o o o *3 O ro o\° 1 CM CM CM p a) f" rd G 116 co r- co o\° 1 ... 15 * | O CO VD 00 S3 ^ in co H td — ■. r-1 03 U 43 42 I o td i—1 u P o «d O ■H • 13 ** H 0 CM <45 ^ CM iH P • sc s rH td 42 43 H < MH o n 0 03 P p mh CM c ffl CD 03 1 • -P 0 CM 0 1 I 1 S3 42 < s rH 1 1 >H | 0 P 0 CD £2j 43 3 0 03 td e m o 1 43 CD P o 117 , p ^p a) t"- iH 118 119 Discussion Bathynectes superbus, among other Portunidae (Marshall and Orr, 1960; Tagatz, 1968; Ropes, 1968), feeds on a variety of forms and is opportunistic in its feeding habits, The presence of mollusk, arthropod, echinoderm and polychaete fragments as well as sediment and foraminifera suggest that the crabs feed upon whatever foods are available at the depths sampled. Euryphagous food habits have been noted in other archibenthic crabs such as Geryon sp,, Scyramathia sp. and Platymaia sp. (Doflein, 1904). The frequent occurrence of foraminifera and sediments in the stomachs of B, superbus may be an artifact. Crabs were frequently trawled up covered with mud and with the branchial chamber filled with mud. It is possible that mud could have been forced into the stomach or that the crabs could be ingesting the substrate along with regular food. The substrate itself could be of nutritional value. Organic compounds and bacteria are often adsorbed on the surface of sand grains and serve as nutrients (Odum, 1971), Doflein (1904) noted that guts of two benthic crabs, Scyramathia sp. and Platymaia sp. contained ooze composed of foraminifera, sponge spicules, shell fragments, pieces of sponges and parts of crustacean skeleton and muscle. It was not dis cerned whether these species were substrate consumers or had consumed substrate after they entered the trawl. Garstang (1897) reports that B. longipes burrows in sand with only the 120 eyes and buccal frame exposed. Although B. superbus has not been observed to burrow, it is probable that it does settle into the substrate. Intake of bottom sediments in addition to regular prey would be likely. The high percentage of empty stomachs found in peeler stage B. superbus is not unusual, Passano (1960) reports that peeler crabs cease eating during the major portion of skeletal resorption prior to molting (D^-D^). In the peeler stage, feeding is reduced. This could account for the presence of food in a few of the peeler stage crab stomachs. The presence of B. superbus carapace fragments in the stomach of the crabs is not likely due to cannibalism but rather resulted from consumption of the cast exo-skeleton, I have observed this behavior in the laboratory. SECTION V EPIZOITES ASSOCIATED WITH Bathynectes superbus Introduction The only known documentation of epizoites on Bathynectes superbus is that of Capart (1951), who noted a lepadid barnacle, Scalpellum sp, on specimens from the South Atlantic coast of Africa. This study documents the occurrence of fouling organisms with regard to molt stage of Bathynectes superbus in the Chesapeake Bight of the western Atlantic Ocean. Materials and Methods Bathynectes superbus were collected from the R/V Columbus Iselin (73-10) cruise in June 1973, the R/V Dan Moore (030) in November and December 1973, the R/V Albatross IV (74-4) cruise in March 1974, and the R/V Eastward (2-74) in April 1974 (Appendix 2). These specimens were examined for molt stage and presence of fouling organisms. The crabs were ex amined externally and the dorsal carapace was removed to per mit examination of the gills and branchial chamber. In addition, U, S. National Museum (USNM) specimens were also examined for molt stage and externally for epizoites. Corn- 121 122 mensals were identified to species whenever possible and their location and incidence on the crabs were noted, No attempt was made to distinguish between early, mid and late paper stages of the crabs examined. Results In June 1973, crabs were captured in depths of 252 to 390 m in the Norfolk Canyon and the adjacent continental slope. Of the 184 crabs examined, 56% of the soft (A^) and papershell (A2-B2 ) and 90% of the hard and peeler (D2-D4 ) crabs possessed commensals (Table 24). In November and December 1973, the one papershell crab and all but one hard crab were fouled (Table 24). These crabs were captured in depths of 122-232 m. In March 1974, 12 crabs were cap tured at depths of 236-300 m on the continental slope. Commensals were found on 55% of the hard or peeler crabs (Table 24). No fouling organisms were observed on the only papershell crab examined. In April 1974, 11 crabs were captured in 280-350 m in Norfolk Canyon. Commensals were ob served on 25% of the hard and peeler crabs? 38% of the paper shell and soft crabs were fouled (Table 24). Among hard and peeler crabs in the USNM collections examined, 32% were fouled (Table 24). Commensals were found on 7% of the paper shell and soft crabs. Crabs were most heavily fouled with Poecilasma inaegui- 1aterale Pilsbry, a barnacle of the Family Lepadidae and a Table 24. Incidence and type of fouling organisms observed on Bathynectes superbus from June 1973, November and December 1973 (DM-030), March 1974 (Alb. IV-74-4), April 1974 (E-2-74) and U. S. National Museum specimens ■H U CM < k C 04 0 G X rH rH rH 3 3 13 13 o n rH G 13 I G III k ( -— - rH P 03 03 o\° P U P CU U CU (U G G fH • G 0 CU — G P G CU (UrH p 3 ro 0) P G G O t3 O 0 _ ■H — — ■H rH ■rH rH O o\° i EH _ P P U E P _ — P O G 0 G G ro e ro G tn G tn G 0 G CO _ O p CU G 0 m m m h • 1 v _ ■rH ■H ■H 1 CM 1 i—4 <—I rH CO CM i rH 1 OS r''CO -p CM u e CO — — l"3 — — ro o 0 o G p G G 1 h 0 G G 0 e G G 0) 0 • • • • 1 1 1 1 •H *rH CO "H vo 00 as CO M 00 : i in CM t f — 0 p 0 G £ m G o On 1 1 u U jg 0 H P "H i ■H CM + 03 P —1 0 a 03 CU O 03 tn o o e j 123 •rH VO o CO < p vo G o e G G O G CU o3 03 • . • oo av vo CM oo 3 O (U G H ij1 -H i-H U H £ rH oo co as o VO E P Ptl co m vo in rH 0 G G -H X G G e G -H 0 h n as -H CO i—i o u C s -P ■—i o o 0 0 > cu s Total examined m Table 24. (Cont. rH i—rH I b b b MH Pi 0 .3 rH i— i as PH pL, -p C/1 sh 0 iH b 0 3 0 3 3 ~ rd d) O, fd0 M 0 b 0 to 0 3 3 • ■H •H — ■H i— 1 i rH 0\0 rH EH E O 2 o\° •P 2 (H U ■p P — 0 0 o in £ 0 o fd ■rH ■H -H i n HrH rH i cn O' 00 n 3 P-P -P PM g — — 0 0 0 3 0 b 3 £ d0 0 P fd 0 O 0 0 3 0 fd H O • • • • 1 1 H•H ■H 04 00 o> rH ■P < 0 o 3 0 E 0 0 0 0 0 •H £ ■H £ X 04 1— 1 00 rH 04 PM 0 >3 0 £ 3 3 U 0 tn 1 H-P ■H b b 3 u >1 0 0 n in g o rH o 0 0 0 3 0 •rH rH rH i—1 E w 3 X 0 E 0 •H 0 124'r h ■H ■H •H H CU • S i 0 w S CM 0 0 p 0 0 CO 0 ) r H Q< (0 (d - P c u 0 EH ____ 0\0 —' r * r o P P • • (U CU r - c m r H ,-H v o r o CU P CU 0 P , (x . P r - v fd • 0 p a i n c o p — ' c o cr> td H m w P B CU CO P ■rH P e fd p p i X p w O r H p i - p p o o CU W •H CM p p CU u rH g St a fd CO Eh D 125 126 hydroid of the nominal genus, Perigonimus. Poecilasma inaequiiaterale was found on all exposed parts of the crab's carapace. A hard crab from the eastern Atlantic in the USNM collections (Geronimo-2-203,* Acc. No. 255091) was found with approximately 100 P, ihaeguilatera1e on the dorsal carapace, pereiopods, eyes and mouthparts. Perigonimus was found predominantly along the ventral anterolateral border and on the ecdysial suture line. Anomia aculeata (Pelecypoda) was also relatively abundant and frequently occurred in in dentations of the dorsal carapace and on the carinae of the pereiopods. Other fouling organisms encountered were cal careous tubes of an unidentified worm, an unidentified hydroid, and Stegopoma plicatile, a thecate hydroid. .Discussion The fouling organisms associated with B. superbus have been previously documented as inhabitants of the shelf and slope. Poecilasma inaequilaterale has a known range along the western Atlantic from Martha's Vineyard to Key West, having been recorded at depths from 21.6 to 1733 m , chiefly on the carapace of Geryon quinquedens (Pilsbry, 1907), In June 1973, and April 1974, P. iriaequilaterale was also found on Geryon quinquedens and Cancer borealis, species known to be associated with B. superbus. Anomia aculeata has been recorded from the Arctic Ocean to Cape Hatteras within a bathymetric range of 1.8 to 144 m 127 (Smith, 1937). The stations at which Anomia occurred on B. superbus were in 122 to 335 m« Perigonimus is a representative of a poorly known group of hydroids which need systematic revision (D. Calder, per sonal communication). Such hydroids actually belong to about six different families and the type species of the genus was based on a Bougainvillia(Calder, personal communication). One species, P. repens has a recorded range from Vineyard Sound to Eastern Point, Massachusetts in 25-83 m (Fraser, 1944). Stegopoma plicatile, a thecate hydroid, is common in deeper waters along the East coast. It extends geographically from Hudson Bay to Cape Hatteras with a bathymetric range of 45-1733 m. Sessile, commensal organisms are useful as indicators of the molting history of crabs (Van Engel, 1958; Terretta, 1973). In general, intermolt and premolt crabs have higher incidences of epibionts than postmolt crabs. This is a reflection of the long setting period for such organisms during intermolt and premolt, followed by shedding of the old exoskeleton. The presence of certain commensals may also vary with bathymetric distribution of the host and indicate migrational patterns of a species (Haefner, personal communication). No attempt was made to distinguish between early (B^), and late paper (B2) stages, but it is likely that the majority of crabs fouled in the papershell and soft category were in late postecdysis. The setting time for Poecilasma inaequi- laterale and Anomia aculeata would also indicate this, since 128 it is assumed that a time lapse must occur from set to attainment of macroscopic characters. SECTION VI EXOSKELETAL DEFORMITIES OF Bathynectes superbus Introduction Accounts of morphological malformations are numerous among the Portunidae. For Callinectes sapidus, abnormal chelipeds, pleopods and pereiopods have been reported (Shuster, Ulmer and Van Engel, 1963; Reinhard, 1950, 1956; Lawler and Van Engel, 1970). Faxon (1881) had previously reported a malformation of the carapace with a deficiency in lateral spines. Abnormal chelae (Perkins, 1924, 1925 a,b; Abeloos, 1933) have been reported for Carcinus maehas and Portunus puber (Andre, 1946), Sankarankutty and Thomas (1963) noted abnormalities of the carapace lobes in Lissocarcinus orbicularis. Various external deformities have been observed on several specimens of Bathynectes superbus collected along the Atlantic continental shelf. These abnormalities are docu mented in this report. Materials and Methods A total of 164 crabs were captured during trawling efforts on cruises CI-73-10 in June 1973; DM-030 in November and 129 130 December 1973; Alb. IV-74-4 in March 1974 and E-2-74 in April 1974. Tabulated station data are found in Appendix 2. All specimens were examined externally for morphological abnormalities. Short carapace width was measured and gonadal development was determined as described in Table 11. Results and Discussion Malformed abdominal segments were found in one female and two male crabs (Figure 27). Segments appeared either to be compressed and wrinkled (Figure 27 a,c) or twisted and slightly asymmetrical (Figure 27 b ) , Several types of abnormalities were associated with the frontal area of the carapace. In normal crabs, the two median frontal teeth are separated by a slight gap or dia stema (Figure 28). Abnormalities included one crab with three median frontal teeth (Figure 29 a) and two crabs with abnormal diastemata: the near fusion of the two median frontal teeth (Figure 29 b) and the wide separation of the two teeth (Figure 29 c). Comparisons with other specimens showed that these differences were sufficient to be considered abnormal. The amount of gaping has been found to vary with age in B, superbus (Christiansen, 1969). Young (small) crabs generally have a narrow diastema and shallow incision while older (larger) crabs have a much more noticeable gap. Morphometric analysis of short carapace length revealed that this alter ation in gap size is not associated with sexual maturity Figure 27 Malformations of abdominal segments in Bathynectes superbus. Measurements indicate short carapace width. a. Female, 49,1 mm. CI-73-10, Sta. 41. b. Male, 53,9 mm. CI-73-10, Sta. 47. c. Male, 61,7 mm. CI-73-10, Sta. 76. 131 Figure 28 Dorsal view of female Bathynectes superbus (CI-73-10) Upper; 62,2 mm. Very well developed gonad. Lower; 31.3 mm. Very slightly developed gonad. Note relative lengths of fifth anterolateral spines and wide diastema between median frontal teeth in larger female. 132 cm|llil|llll|lill|M II|llll|IIM | 1 1 2 3 nlTP -- (ll.|im|iiii|iiii|iiii^iii|iiij| SPEC------DATE______Figure 29 Abnormalities of the frontal region of the carapace of Bathynectes superbus. Measurements indicate short carapace width. Arrows denote abnormal regions. a, A 53.7 mm male with three median frontal teeth; Center tooth is supernumerary. CI-73-10, Sta. 46. b, A 67.6 mm male with reduced diastema between the madian frontal teeth, CI-73-10, Sta. 47. c, A 64,5 mm male with unusually wide diastema between the median frontal teeth. CI-73-10, Sta. 47. 133 134 (Section II). Other carapace abnormalities observed were associated with the orbital area, the lateral spines and the dorsal epicuticle itself (Figure 30), Generally, the orbital region of the carapace is notched twice directly beneath the eye. One crab was found to lack one orbital notch (Figure 30 a). The indentation present on the carapace of a female crab (Figure 30 b) and the abbreviated fifth anterolateral spine on a male crab were probably due to injuries (Figure 30 c). The fifth spine is normally much longer and more conspicuous than the other anterolateral spines (Figure 28). Most of these abnormalities that have been observed in crabs are thought to result from injury or parasitism (Lawler and Van Engel, 197 0) but there are also examples of real duplication (Faxon, 1881; Shuster, et al., 1963). In addition, deformities have generally been observed on those parts of the body which have the greatest capacity for regeneration (Shuster, el., 1963). It is likely that abdominal abnormal ities in B. superbus are also associated with injuries sus tained when the crab was in a vulnerable molt stage such as soft or papershell. Similarlyf the shortness of the fifth anterolateral spine and the indentation of the dorsal cara pace near the fifth pereiopod are likely a result of injuries. The lack of a notch along the orbital area of the carapace in B. superbus is a similar abnormality to that described by Sankarankutty and Thomas (1963) for Lissocarcinus orbicularis. They attributed alterations in the number and size of the Figure 30 Carapace abnormalities of Bathynectes superbus, Measure ment indicates short carapace width. a. Lack of outer orbital notch on a 52.2 mm male. CI-73-10 r Sta. 44. b. Indented area of carapace on a 38.1 mm female, CI-73-10, Sta. 44. c. Abbreviated fifth anterolateral spine of a 58.7 mm male. CI-73-10, Sta. 47. 135 136 anterolateral lobes of the carapace to abnormality rather than variation within the species. The abnormalities in frontal spines were only observed in two specimens of B „ superbus so it is doubtful if this could be attributed to species variation. SECTION VII SETAL VARIATION IN Bathynectes superbus(Costa) Introduction The setae of crustaceans are thought to have an important role in their bodily functions (Balss, 1944). Setae may function in respiration (Thomas, 197 0), reproduction (Lloyd and Yonge, 1940; Thomas, 1970), feeding (Fryer, 1960; Thomas, 1970), and to increase surface area (Borradaile, 1926; Dennell, 1933; Malagzuska-Suchzitz, 1956; Thomas, 1970) or sensory perception (Gulland, 1886; Holmes and Homuth, 1910; Thomas, 1970). In addition to the functional variation in setae, crustaceans possess a variety of setal morphological types with setae of the same form generally found in a species specific location (Thomas, 1970). Since setal types have been found to be consistently located within a species, Menzies (1960) has suggested that setal structure should aid in taxonomic and genetical studies in arthropods. Despite the homogeneity in setal type and location for other crustacean species, Bathynectes superbus exhibits intraspecific variation, Among allopatric populations of B. superbus, two patterns of setal presence were observed. This paper illustrates the major setal types and the variation in setal presence as it occurs in B, superbus. 137 Materials and Methods This study was based on 132 crabs from the R/V Columbus Iselin cruise in June 1973; 6 crabs from the R/V Dan Moore cruise in November and December 1973; 12 crabs from the R/V Albatross IV cruise in March 1974; and 14 crabs from the R/V Eastward cruise in April 1974 (Appendix 2). United States National Museum specimens were also examined for presence and location of setae (Appendix 3), Sections of the carapace of 3 crabs were removed and trimmed to 3 cm squares. These sections were washed in dis tilled water and dried in a vacuum chamber. They were then coated with a 60% palladium-40% gold alloy. Sections were examined and photomicrographs were made by scanning electron microscopy. Results Two patterns of setation were found in the specimens examined. Pattern I, in which pubescence covered the entire body, was only found in specimens from stations off Liberia, Gulf of Guinea, southern Florida and Georgia (Table 25; Figure 31 a). The incidence of pubescent specimens was greater from the eastern Atlantic (100%) than from the western Atlantic (18.4%), The other western Atlantic specimens possessed setae that were mainly confined to anterolateral areas of the ven- 138 Table 25. Geographic locations at which specimens of Bathynectes superbus with pubescent covering were collected. XI -P o co Q - -P E D ' -iH •H •H Ti h! *3 -P U pi PI -p CO -P i p •p ■P fd 0 P o o 3 CD g 3 3 fd o C o CP CD g 3 O O •H p o Cl td m p i—I ■H P C CM oo VO VO O'* PI £ CD -P o P ■— C C -H k00 0 Pk CD -H w n P o td p 3 cn oo I 2 i p •H i CM — P P P 0 DCD CD t"- 0 g g g g g g g 1 1 1 o 0 0 “ p I CM 00 00 < M00 vo CM 00 H■ H•H ■H ■H ■H Op LO CO 0c 00 CM CM P H i — 0 P P i 0 g 1 1 CM oo — i0 C 0 Ci 0 1 13 9 0 p — VO m p H 0 000 00 CO CM w CM 0 3 3 0 id P P P 0 CD g cri 1 1 . 0 H g CD O 0 - 1—1 CM CM p 00 CO M 1 CM | i C - r CM MP CM o 0 0 g g P DC DCD CD CD CD P rP 1 1 • 5 1E CM 0 0 P p o g 1 1 CM r- CM in in P 0 0 0 g CM 1 c t" I I 1 i CM o oo o CM 0 0 CM 0 0 P CO m M ■P CM 0 00 p C o g g 1 1 p oo O o •H 3 t .P 00 o Ci P4 fp r"- Cl 0 0 00 VO 00 P rQ P CM rP CD |H g rd Ci g < CM -P c CM O W P 3 3 Cfl VO ts VO 0 0 0 O A cm co P 3 0 p U1 u h 0 3 A 4-> (8 ro id O P tH U 0 0 3 0 0 0 o o -H P H P VO P p r-~ p h rd 3 NO 3 ^ -P PI CM Pi CM W H H rfj rfj X! p ^-N ■=j< r- o* e o on 0 w in ^ Q 0 ts 3 12 ? p cm r- -H CM o &> 3 O 0 0 ON ON Pi p- r- 0 !3 S T3 - •» 3 O ID P CM CM ■H p 0 0 3 i—I i™H PI CO CO p 3 O U p p • 0 0 0 3 3 3 H 3 W Pi W 0 3 0 in ■H '• S “ H CM P 3 3 3 0 3 ■HP H &i 0 O On H rQ O' 3 I—I 0 P C S P W PI PI 0 3 0 3 0 4-1 0 0 P E-t U 0 W U O 140 141 tral carapace (Pattern II) (Figure 31 b ) , Some setation was found on the legs and dorsal carapace but the type of setation was markedly different from Pattern I crabs. The dorsal carapace region was naked, Indentations, possible representing empty setal sockets (Figure 31 c) were present on all specimens examined from Chesapeake Bight. The external morphology of the setae also varied intra- specifically. Setae from Pattern II crabs arise from well- developed, rimmed invaginations of the cuticle and appear to be moveable (Figure 31 d ) . Both slim and heavy acuminate setae (Thomas, 1970) are found on the ventral anterolateral edge of the carapace. Setae of Pattern I crabs were located in shallow depressions of the carapace (Figure 31 e,f). These setae appear to be acuminate but are not articulated to the cuticle in the elaborate sockets that are found on the other crabs. Several of the setae appear to be surrounded by or tufted with a hydroid-like growth (Figure 31 a), The sheath surrounding the setae may be an artifact associated with preparation of the specimens, since no attempts were made to remove adhering materials prior to desiccation of the specimens (Perkins, personal communication). 1-42 Discussion Bouvier(1940) and Capart (1951) noted the presence of pubescence on crabs from the coasts of Africa and France. Rathbun (1930) did not indicate where the pubescent crabs were collected but included pubescence as one of the general descriptive characters. I examined the crabs mentioned by Rathbun and found pubescence on only those specimens from Florida and Georgia. The setae found on the carapace of B. superbus are simi lar to those acuminate setae described by Thomas (197 0), for the crayfish, Austropotamobius pallipes. He designates that forms of this type function as sense organs and extend the surface area of the organism, Simialr conclusions may apply to B. superbus on the basis of the gross setal structure and locations. Williams (personal communication) suggests that there may be an association of pubescence with bottom type, the pubescent specimens coming from softer substrates. If setal growth could be linked to substrate habitat, it is possible that extensive setation could prevent the carapace from being covered with mud in areas where this substrate is predominant. Further behavioral observations and histological observations would better define the functional significance of the setae. 143 Figure 31 Scanning electron photomicrographs of selected regions of the carapace of Bathynectes superbus, a. Posterolateral carapace of papershell male crab from the coast of Africa (4°40’N; 9o20*W) (USN.M No, 126161), 73 mm short carapace width. Granular mounds are seen among num erous setae. Setae appear to have tufts which may be epi- zoites or an artifact, 90 x. b. Ventral anterolateral carapace of hardshell ovigerous female crab, 64 mm short width, from western Atlantic (39°55'N and 69°30'W) (Albatross IV 74-4). Granular hill ocks are evident as well as setae. 95 x, c. Right dorsal posterolateral carapace of hardshell male crab, 62 mm short width, from 39°55TN and 69°30'W. Granular hillocks are evident as well as craterlike in dentations which may be empty setal sockets, 110 x, d. Ventral anterolateral edge of carapace of hardshell ovigerous female, 64 mm short width from 39°55'N and 69°30'W. Elaborate setal socket is present and seta appears to be moveable. 450 x. e. Right dorsal posterolateral edge of carapace of paper shell male, 73 mm short width, from 4°40'N and 9°20'W. Note shallow sockets in which setae are located, 180 x. f. Enlargement of setal socket from e. 450 x, SUMMARY Bathynectes superbus is contagiously distributed within the lower shelf-upper slope habitat of the Chesapeake Bight. Bathymetric range of B. superbus may vary with season, since crabs in November were found at shallower depths (122-223 m) than those in June (252-390 m) . Distribution and abundance was associated with temperature. Crabs were found within 5.1-11.3 C in June and 8.7-10.1 C in March. Maximum abundance occurred in the 8.0-11.9 C range. Male crabs ranged from 21-70 mm short carapace width; females from 28-62 mm. Width frequency plots indicate that male B. superbus achieve a larger size than females. Four modal size groups of male and female crabs were noted: ^35 mm, 36-45 mm, 46-57 mm and ^58 mm. Some element of the population, particularly crabs >45 mm, is molting throughout the year. In June, hardshell (C-^— ) and peeler (D^— ) crabs were more abundant on the slope whereas soft-papershell (A^-B2 ) individuals were more abundant in the canyon, suggesting that the canyon may provide shelter during the most vulnerable part of the molt cycle. 145 6 . Cole's coefficient of species association (C^) indicated that B, superbus co-occurred with Cancer borealis, Homarus americanus and Geryon quinquedens in June, Low positive associations were indicated between B, s superbus and G . quinquedens and H, americanus in March. 7. The relative growth of all body parts displayed simple allometric growth, except the isometric relation between short carapace width and long carapace width. Puberty- related changes in allometric growth of the width of abdominal segment 6 were observed in females. 8 . Sexual dimorphism of B. superbus is similar to that of other portunids. Abdominal shape, pleopod number and size and presence or absence of vulvae distinguish males from females. No color differences were observed, 9. Gross and histological anatomy of B. superbus reproductive organ systems is also similar to those described for other portunid crabs. 10. Synchronous reproduction is suggested for B. superbus. Some element of the population is ovigerous year round but the primary period of egg extrusion probably occurs in November and December. Maturation and release of the eggs occurs in late spring. 11, Maturity within female crabs apparently occurs at — 28 mm short carapace width based on the presence of sperm with in the seminal receptable. Male crabs > 28 mm were also considered mature on the basis of sperm present in the testis, 146 12. Fecundity estimates indicate that B, superbus produces fewer, but slightly larger eggs than inshore portunids. The comparative lower fecundity is probably due to the relatively smaller size of B. superbus. 13. Stomach content analysis of B, superbus indicates that the crabs are euryphagous, feeding on mollusks, arthro pods, echinoderms, and polychaetes, 14. Poecilasma inaequilaterale (Cirripedia; Lepadidae), Anomia aculeata (Pelecypoda) and the hydroids Stegopoma plicatile and Perigonimus sp, were epizoitic on B. superbus. 15. Exoskeletal malformations were associated with the carapace and abdomen. Most malformations were attributable to injury of the crab. 16. Intraspecific setal variation among two allopatric populations of B. superbus was noted. The major setal types and variation in setal presence of eastern and western Atlantic populations is illustrated and discussed. Appendix 1. Documented accounts of the distribution of Bathynectes superbus prior to this study. •H ■rH CO m "H x: o 0 P P CO 0 a 0 P u cn e SH 0 p a 0 O Co o Xi a a XI -P IO0 O XI ■H Q ■p •-3 - PI cn 0 p. p O CO p P o p p > £ 0 p p o 0 p 0 • , • * • • rp +J a H H 4H h m p td td P £ 0 & d> a XI XI o td 0 p t p t O CM — rH CM CO o o r- CM VO 03 o O o O'- MC I i— 1 IrH CM CM •H 'd -P Xi X! a !2 12 <33 CO O Pi > p P p >i £ 0 p P l 1 l 1 • • • • I , • , • • • • • • • • • • • • • • • • • • • • 0 o CM - TP 00 CO 00 m m ["• m o 03 a 0 0 0 0 CO CM 303 03 in CO o in o r'' CM O o r- a a & m o o i • 0 0 TP in p t CO 03 p t o o o in o m a 1 1 1 1 • • • • • • • • * • • • • — o s *TP •si* o o o t"- o ■— i o in CO a a 1 <4H MH : x >i a > O p - 14 7 P P p P 0 0 0 0 0 o 0 0 O — -H 'd O o o CM CM CM CO CO O 03 t-" ■H CM O t" o MCM CM a a a a a xn X3 i—I 00 00 CO •P p 0 p £ e PT CO TP TP o VO in 03 m i— I o o o 00 in TP a 1 ■» o o o TP O'- o o rH CO o i o in in in a a a a a a a a a — 1 1 ■» 03 TP TP 03 [■" in o in O rH 1 0 0 CO ■» MCM CM 303 03 in in TP rH VO o TP CM in 03 in rH rH a 1 0 0 O O o o OCO CO »■» ■» 00 C'' o o o o o VO o VO o o CM 00 VO Hr rH rH rH a a a a a a 1 03 in TP TP TP VO CM 03 O 03 TP TP 00 1 1 m 00 — — CO 03 in MCM CM OTP CO o o O • X! MH Mh •» O > O >3 a p ■ p p P 0 0 0 0 0 0 O -H 'd OCO CO TP 03 in TP OVO VO 03 03 TP o o 00 PCM TP i—1 XI XI -P PJ <33 CO O p p 0 £ P P P 1 1 1 rH 1 I • 03 TP VO i— i m TP in in MCM CM i— 1 rH • — in 00 o CO 03 CM o o o rH VO 00 1 • Off Nantucket Shoals 39° 57.5'N 69° 41.7'W 140 Appendix 1. 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O O o o - r - r *51* r~ MCO ■— i CM VO r*- in 2 IS i l 0 0 0 0 0 0 0 0 -H 5* 5*00 0 *51* *51* 0 0 0 0 n i o o — — — — — P ■H *5i* o o o o rH * r' o o r-' l*- O O *sj* *53* m c P i--vo oo in r- r"-r- r- r- P 2 2 2 2 03 CM i—CM 1i— 1CM H 0 ■H -p u IS IS IS IS X X 0 0 tn 0 01 0 Q 0 O > n. 00 0 P 1 = = t f 01 u o 0 0 P P o i o in o oo a in o o o in vo in vo o 0 -. cm cm H p •H o O 0 C W H M pi• • O 0 0 O O £ o HrH rH CM CTi £ H VO o r- p p m *51* o Cl VO 0 H m m rfj on 2 2 IS CM P X rH —' oo t f o 4-1 oo 0 -P 0 O 01 .-P L C p p • • P 0 0 o 0 f" 0 0 0 CM O') CM o O CM *5p in o o IS 0 -P H-> -p G — G _ i—1 o 0 0 0 I*- O o [> 00 £ £ £ 00 00 00 00 00 cn ov 00 H cn G cn G 0 ■—1 • VO • CN • VO '—' rH i—I — - ' _ _ ' o 0 O m oo tn c m —** p G rH G o G O G G G G G a h a rH a >H 0 G O G 0 0 G 0 G W G 0 CP £ a £ a £ a £ p’ -P p -p P -P g £ g £ g •p 0 tn 0 tn 0 m •H -P r ■H ■P -H 0 " pj g -H -P 0 ■P 0 -P PS •S3 w < £ < w CP PS CP PS CP > G rH 00 I I 1 I 1 I 1-0 CO d d d G 0 0 0 0 • X X X g a 1 ■— 11— i 1 1—1 CN 0 1 rH 0 G 0 G rH 0 w 0 O GG G 0 p D £ a • cp a 1 r- rH i— 1 1 rH rH t rH 1 G o o a N1 VO VO CM ID oo o o O O VO O CD 1■ i rG CN oo a tD ID ID o o CN VO 00 O -P OV 00 CN i—1 as rH 00 00 VO rH CN r^~ a C^ VO LD 00 00 rH rH rH CNCN CN 0 Q £ ? ? !S O ID LD o •• tjs VO 00 00 I I | 1 1 1 1 VO 1 G CN ro oo I i i 1 1 1 1 ro 1 0 I I 1 1I 1 1| 1 1 0 G 0 O O O tn OO a as i—f G r-~ r» r - 00 ■rl asP 0 CQ £ £ S £ • o 0 LD VO -P • 0 0 1 00 | | | 1 1 1 J 00 J G O ID I | | 1 1 1 1 I—1 1 1■ I 1 1 1 1 o 0 o G O G 00 00 00 s§ CN £ -P >1 1—1 £ 0 0 » d £ 0 -p G "H rG G 0 0 G d CP 0 O H G G U ■H G P •H >i >i 0 -P 4J G £ 0 0 d o G 0 0 CP 0 g 0 0 a -H G G 0 £ £ 0 0 0 o 0 -P 0 ft* • 0 0 -P & 0 IS ■rH P 0 G G a P G d d 15 P P i—1 tn O U P g rH 0 0 O G G £ >1 -P >1 T3 0 0 0 0 0 0 £ 0 CP 0 g X 0 G G a a Q ffl tn tn • -P £ £ ■H 0 0 0 0 d cp 13 •rH rH m g MH <4H <4H MH a tn MH G V|H 4H G tn . co m m m m uh m •H m G MH m 0) P W H o o o o ID G O a 0 o o o 0 O a 0 c 0 149 Appendix 1. (Cont. ■H -rH H £ ■H PQ CO X3 O o 0 P £ 0 0 CT> Pi o 0 g 0 U £ Pt-rH o o 0 P 0 tn O -P 0 P) h •P Hi -P Q [x, CO 0 0 O 0 P. pi 0 O £ p o 0 ! H MH ■H 1—1 3 m 73 C 0 0 o p m H cn (N p CD •H ■w rH CO 00 - r rH -P P 0 0 0 P1 0 0 H 0 £ g 0 i P 0 £ 1 • • £ 0 p 0 >i4H mh i CN rH — D00 rH CD O n t CO p - CQ H- 37 P 73 73 -H rH o CD CO CO CO — 0 0 £ 0 £ P n t n t 0 P 0 t 1 - u O CO CO ■P P • ■P 0 P MH 0 0 0 £ 0 0 0 tn in d c rH t- o o CN i £ 00 o rH H CD ■H A — u —1 ■P 0 u 0 0 .v 0 fl 0 £ I 1 H cn \ y < 150 rH 73 MH IS o H CO ro '—' 00 O'! 1—1 1 n- £ O P £ 0 0 — 1 0 P £ g 0 1 730 rHI CO -P 0 o 0 fl SS ■H MH i 73 CO rH CO 373 73 -P —1 13 P -p Hi 0 0 £ 0 X Oi 0 0 0 0 LO 0 g £ O H 0 £ I—1 1 1 1 • h ~' “~ — £ £ — CO o 0 3u 73 MH X! CD i— CN rH CN r-i CD CN -P CO IS £ £ rH 00 1—1 i ,— . Cn S2i 0 £ 0 0 — 0 p g 1 0 1 i 1 rH MH I C §: D — i o ■P — Hi 0 0 £ tn tn £ 0 0 X CTi 0 0 m 0 i—1 0 m O g 1 1 £ 1 1 1 CD CQ P 0 Qj MH0 UPi U PQ O 0 S £ mm£ r£0 C rH •P 0 CN ■H , ' p 0 CQ t u o o cn — 0 P £ o o 0 P _ 0 m £ 0 > 1 1 H 0 0 „ *H i X MH PP -P -P 0 0 K •H H i SI ■P ■p — P 0 0 0 £ 0 0 £ tn £ p 0 0 1 rH 0 CQ 0 0 P 0 0 0 0 0 £ 0 £ -H - 0 i—1 i—1 > P rH I I I I I I I I I I I I I I I I I I Cfl P oo • Cn tn tn tr* CD i—l &i *H -H r-4 -r4 -H S IP I I CN I H rl i—I *• -H I i—I CN **3* i—I > r4 > CN | - > > ■H CN > i—I O O H 0 O O i—I o CD ro ft C/3 I r—I I rl 1—1 CN 1 t—i H 1 I 0 0 i n 1 1 1— 1 1 CO rH 1 CN m o o O o o o o o o O O o o o CO o i—i o 0 0 CN 0 0 0 0 LO c n CN 0 0 m CO i n m i n 0 0 H 1 N* 0 0 oo 0 0 i ■ 1 I I 1 1 1 1 03 i 1 i 1 1 1 i 1 1 1 1 1 I JG N* I i I o O o o i n o o o o o O O o O o o -P CD I i i i n O i n r - 0 0 o o 0 0 o 1—1 0 0 O in I" c n c n ft in i i i ■—i 0 0 CN CN 0 0 H 1 H* 0 0 0 0 N* 0 0 H* CN (N CD Q W W W pq £ H £] W pq W w Cn llll illl I oo oo o o cn cn h1 m cn P t i l l l l l l 1 CN CN CN O rH O CN CN CN 00 r- 0 llll llll 1 03 PI 0 0 0 0 O O 0 0 0 0 Cn 00 CN 00 00 LO CN iH i—1 i—1 i—1 0 P i—1 i—1 i—1 i—1 i—1 i—1 i—1 i—! i—1 i—1 00 ■H p fd CD C/3 m in CO C/3 CQ CQ C/3 -p S C/3 C/3 C/3 C/3 C/3 C/3 fd t i l l CO I I I oo oo m in pi i i i i H1 I I I m m m h * in co cn co O'* oo in i i i I I I I O r4 CN o 00 CN r4 o 0 0 0 0 HHHO 0 0 0 0 0 0 o rH I—I rH rH t—I 00 I CO VO IT) LO o p w *r4 CD fd P *H ft■H >i rH PS td 0 cd i—i td fd 0 ft id 4-i S 0 CD CD fd cn 4-» H 01 rH p p -P td U3 T3 □ P H CD< fd p 4-> 0 P rH *H C CD "H >i 0 0 o 0 Q) CD fd -H •H PJ O 'd fd - Cfl -o 44 rj 4J +j 4J g N 03 4-1 P3 fd 4-1 ft-H P Cfl CD fd 0 0 -H *H *H ■H -H fd P P fd -M fd CD H 6 Cn Cn Cn CD P td 0 0 fd CD 54 fd -P ih a Cfl iH • CD W H H ft P3 -P S PS 0 Cn O p ft fd rH CD < 4-> rs fd S -H • P O O fd fd s fd■H f t 0 4-1 PI S S S 0 fd i >i kH 44 P 44 >) >1 0 CD 44 44 44 p B . fd Cfl P3 JQ P3 o •H 0 IS PI PI IS ft) £1)4444 44 (UH g CD 0 P C/3 C/3 O O o o C/3 < J3 53 IP |S |S |3 |S 0 |S IS IS 5 IS 13 P53 M 151 CO 03 P P 0 0 0 0 0 St -H P nc5 03 > G ■H W G O 0 — 1 0 — • CQ n in 0 CQ ^ 0 in G cm 0 cm H 03 oo •Hi—1 "H G rH q w a 0 — , > G 1 1-3 (0 03 G 0 a) • X g P4 0 iH •H W OG 0 D Q* • co a CO o a o in w I o o -P o o D j CO 00 0 Q C7> I l G I I 0 I I W (-1 Cn G *r-f p (0 0 cq » •p 0 I I PI I I I I -p G o o o *H G X! 0 0 C^'H G 0 -P 0 P 0 •H Cn O ■H 0 0 cm X 0 PI ■H | •H O 0 *P I 03 •H W | G P 0 1 0 0 U 1 01 Cm i i—1 | 3* < i 152 Appendix 2. Collection log of specimens collected during CI-73-10, Alb. IV-74- DM-030 and E-2-74 cruises. ■H xi p ■p PQ cn CD X cua p c CD fd u P D 4-J -p +J p C 0 td CD nO tn fd £ a CD fd o u 0 0 rH o p- i— 1 CN 00 O CN CTi CO p- i «sP IS ro p m H rH a\ H H H !X! X! H H CO co CO ~p- p~ H — S3 > > > 1 i 1 l 1 l . • , 1 o O — — ■ 'Sp CO CO -LO p- CO i co <— 1 p- co P^p ^P » IS P p -vP H co CO m VO rtj — 3 H — | I 1 1 1 1 i 1 O O — i— 1 oi CN ro ro CN rH 1— CO CO p- LO IS rH CO CT\ H X p- ■>5P "sP H S3 < H 1 1 ■ 1 1 1 1 0 0 » S 1 *SP CO m 00 o 00 00 p rH i in 00 CO in CO Cl P- p p ^ VO CN CP IS P> H ■^P > — M — 1 1 1 1 . - i 1 0 0 — CO m. H VD VD P >p- p> CN CO VO p P in CN 153 IS 13 CN p- ro ro <7\ X o o CO o H Q a 1 1 1 1 I 1 0 0 — CN ■sp 00 •O1 NCN CN m ro co VP CN 3 op ro p- ro IS P ro o NCN CN X ro ro H o O Q Q a 1 1 1 1 1 1 I . . . 0 0 0 0 0 0 0 00 i p p ^ i CN CO P CN -?P —1CN 1— 1 ro 3 IS — ro ro X p* —1 o"P-5P "?P ro ro H H p- o a 1 i I ■ 1 1 1 1 . • . . . 1 '•vP N' Nro CN o\ p CO CO CO o CO DV VP VD VD 00 00 CO 00 VP p 3 IS > —Iro 1— I O H ro U rH o H I I 1 1 0 NH CN 1— p ncn in co o o i p OC CO CO CO 00 Nr CO rH CN (T\ Is p 3 ro > H z z z I 1 0 ■» ■- ro P oi CN Oco CO p 1 N P CO 00 IS ro p 3 3 > H S — 1 1 1 1 1 0 1— 1 ■>p p ^ m. o in p p VP rH 00 o is 00 LO ro p p > <=P ^ in ■^P E Z H H i ■ . • * . 0 0 0 co 00 CN oO CO oo CN VO *?P •3* o CO CO VD OO 13 3 ro > < r«sp ■<=r £ - E 1 i 0 » NiH CN "p p p- p CO in CO VO ro IS rH 3 OO 000 00 CO > ro p p VO H H E 1 t 1 • 0 0 rH o> *3* Oi CO in P CO VD o IS o co VO > ro 3 r- i ■ 1 . 0 o 0 rH Sh 1 £ P ro 0 C/3 1 £ H O 1 CO CO 'sr VD 00 OV fd r-* r- r- r* -p w CO CO r- CO CO co II 0 r-> r'- o 0 0 -p I l 1 G G fd H HH PP Q > > > 1-3 h) i■ ft i■ i i O'! rH rH CN CN i—1 I— 1 rH i—1 U g o 0 w 1—1 VO CO CO CO -p •••••• -p • ID o o CO 00 rH o a rH 1—1 rH m g o Er> VD in -P • t &S O VD cr> o CN CN 0 X in VO r- r- in a rH CN IS IS IS is IS IS c\ VD CO r- CN • • ••• » • CO CN o *^F VO 00 G co •vT •sF CO CO CO O P! 0 0 O 0 o o -p ^F ">F ■^F ^F G 10 t-* I" r* r- r-' o CT> u G ■H M fd 0 2 2 CN « - 2 2 2 LO in — X • r-' CO CN 2 r-' •H -P ••• * ’’C td ^F VD av o CO g CO O O o o o 0 CL. 0 0 0 0 0 0 ft VD i"* c- r'- r- < CO co CO co CO CO 154 Appendix 3. USNM specimens of Bathynectes superbus examined with data on their collection. "H •H -H -P ! -P X! PQ 0 PI 0 fd p &> co i-q < G CM ni -vf o in in O0 o 00 CO 00 - rH 00 i P H P CM ,X m rH CO H o i X MH 00 p m i n m i n i p n i p p h p cm p cn p in m tp p ^ r i f f ^ r i f f o c i f f -p a — o QCO CQ (fimfdcsidimoofljcoonO'oO^OHOn 5 rd fd p . • . ■ . • • . • • 1 0 0 P T O+JHP^+J^+JlD+llOOn I n O O l l + D l J + ^ J + ^ P H J + O t s S fd (-p > td P CD P P P 0 — - - r nocn o cn o in i s cn cn — Mo CM i—1 m sj1 rH CO ro o mh rH CO & £ or ro ro ro tn 1 £ " i X o 1 a <1 a H H — CO £ fd —I H i i i i i p i— | i— I i— I i— I i— | -H I i— H cor? 1 1 1 ■ rC3 - i-P > fd fd P h p P d) a o rH *rP m jd in MH MH O C cn m M00 CM O C O C O C > rH 1— & £ cn O H H CO CO td cd 1 1 1 rl H f d C M f C C M f d C M f d C M f d C M d - rd fd p +> i+ > (d d CD fd 0 ■» » X r" i i r n n n T3 T3 0 -=p » r- tp - o o c O ID 1 i—l i> — 1 i—! > i i> - 00 rH & in O C £ Hi— ro rH •TP cn o c < c < cn t t 1 P P H -P •H rHCO u d u fd o P fd o d o td MH mh 1| | in - MH 0 h g (d O rH P CD -P tn 1 ■H U P -H P ’U 0 0 CD -P o fd p tjl fd a MH h a Mh o O CD P p P P ■*03 13 T3 o a 0 ro cr\ cn MH ro in o oin ro ■ ocn ro MH 1 no in > 15 £ O'* CM o o i— 1 — — in in ro d fd P 1 1 1 1 0 0 m 00 o rH tp -rP M• CM *rP r"- o in in 5 £ H > w H CM cn CM i— 1 r- cn cd tr>H I Hi IH a nd CM 0 a rH ID O a -TP in rH CO fd p CD in 1 1 1 Off Key West, Fla. 21-VI-1893 Bahama - 1 77116 Exped. VO 00 © o rH CN O CO LOVO 00 cn VO r*- r- r- 00 00 00 00 00 00 00 00 in in +3 0 \ 00 00 CO 00 00 00 00 00 00 00 1 rH H 0 © VD VO VD VO VO VO VO VD VO VD 1 r*- u VO CO CO CO 00 CO CO COCOCO CO 1 r-'- 77108 rH rH rH rH rH rH 1—1 rH rH i—1 w 3 o a H 1 rH i—1 CN CNCO 1— 1 CN 1 1 rH 0 S 1 rH 1 1 rH CN i—1 1 1 rH CN 1 i CO 1 a cn 0 i—I 0 W O ■P CO 0 0 0 0 0 0 0 0 0 0 cn XI r^. fd >d fd fd ' 3* fd ro fd fd ^ fd cn i 0 p p P p p P 00 P rH P P VD p CO I 0 0 0 0 0 0 -31 0 <3* 0 0 si* 0 I a 0 0 0 0 0 0 0 0 0 0 9 6 9 6 CN cn CN 6 2 6 3 6 5 6 4 0 O 6 7 in in I CO rH ■P I i i i i i i i VO VD H I CO -32 fd H H > > HH H IX II I H H CTi Q H H H H > > > H 1 H H > H rH H > 1 1 1 1 1 1 1 I 1 I > I 1 VO 00 rH co C^ CN in VO CN I -VII '53* CNCN CN CNCN CN CNCN CN CN CN 2 00 00 in O O OO o in in <7\ cn a a a a a a a a a a x! cn CO rH in 00 m CO cn CO in cn i -p i—I o in rH in o 00 © VD o r - i a CN CNCNCNCN 1—1 rH cn CN CN I 261.9 0 Q Mi 0 s £ 5 1 W !S S S 2 £ —• a —«- in w — tn 00 o VO CN 1— 1 CO a 1—1 00 - 1 £ m CN in O CNCN o rH LO O 0 kI 0 0 0 0 0 0 0 0 0 0 tn o CNCN o o o o OCN o 3 00 00 00 00 00 00 00 00 00 00 p 0 0 2 CQ 2 2 2 2 2 2 2 2 2 in — *» •> — *» — — • -P m o o rH rH 1—\ CN 00 —1 0 CN CN CN CO CN O rH CO 31 0 0 0 0 0 O 0 0 0 0 <3< * 3 * in *3* *3* VO LO O' +J CN CNCN CN CNCN CN CN CNCN 3 O 0 U &4 r—I F l a . 3 ■P O O W 0 0 0 0 0 0 0 0 0 a 0 Oh 0 co •H "H 0 i—1 rH i—1 rH rH i—1 i—1 i—1 i— 1 0 0 X l +» Es Cm 1 Cm Cm Cm Cm Cm a a a dv C7> X a 0 1 0 3 0 3 -H 0 o m h MH MH m h MH MH MH MH MH 0 +> 0 -P p o 0 o 0 0 0 o o 0 0 o tn p tr> P C a pi 2 3 o 3 0 0 •P Eh -P Eh a m h 0 0 0 0 0 0 0 0 0 P P Qa m h ■p -P -P ■P •p -p -p -P -p 0 0 < O cn cn cn cn cn cn cn cn cn EH cn Eh cn 156 3 ■P G G G G G -P 1 CN l 1 1 3 cy ( Of^-OcoOvooo 0 CO CO t ro i 1 1 & LO 1 ty o ty oo ty cn tji cy ty 1 l 1 1 a cn i 0O 0cN 0r^0io 0 CN 0 G H G H G G G u O O O OO CN i—I H rH LO CN ro co CO CO O' CN ^ LO LO CN CD cy 1 1 1 1 LO CO LO 1 1 LO rH +J ro MHH H 1 1 I H H 1 LO 3 1 H H H H HH > H HH cy P H > > > > HH H H X > rH > 1 1 1 1 H > 1 1 1 1 1 CN CN CN CN 1 1 CO CO CO 1 I 1 I o 1 0 1 1 O w P O 0 0 ty CO CO 00 VD VD G 00 00 00 00 00 ■H G rd 0 a a CQ - £ ss a cy CN -P* 1 1 1 l 1 cy 1 VD VO i—i O 1 fd 1 1 1 l 1 CN 1 CN O SF *3* 1 P 1 1 1 l 1 1 1 .—- 0 0 0 0 0 • ^ cy CN CN -P CN CN CN CN CN G H 0 > ) U • 0 0 '— 0 (d 3 • 3 • • 0 0 0 0 0 3 iH CQ 3 PQ 3 3 0 o OO 0 G ty Ph i—! rH H cn *H P •H •HP • O 0 G • MH - U Eh *3 Ph Eh 3 X X X X X ro ■H *H -P 0 0 " 0 0 0 ty 0 0 0 0 0 ,g -p G i—1 • > i (d f t - OS * •» 3 £ £ £ £ £ X O-i 3 0 El. cn co o ty cn cn cn -P •H td o EH cd • 3 3 CN fd CN 3 3 G MH MH MH MH MH G 0 ■v ty ■H CQ -P =#= ty =w= ty ty 0 0 0 0 0 0 G 0 p MH +J 3 £ G 3 3 3 Eh CD 0 X i -p 'O O MH -P MH -P -p MH MH MH MH MH G. ty G A" 0 Eh 0 G 0 G G MH rH rH rH i—1 rH Q < • -H O r - f t • • 0 • 0 o MH 3 3 3 3 3 < CQ P Eh rH =«= co CQ Eh CQ Eh Eh O 0 0 0 0 0 157 Appendix 3. (Cont. ■rl •rl -rl -r| PQ m -P P P O 0 cd P o p to G Cn Ck 0 U to 0 B G o u o cd cd Qt 0 g O Ph P *G ■P cn •P Q o p ■P -p p - cd o G a co cd CD G cn 0 0 0 0 H -H ■H N CN 1 CN i • o o mm . i— o D I VD 00 n cn in h D CO VD i • i— ip 0 0 • ropq P o o P P P G, p1 -p vd w D tJ w H 0cn 00 p 0 G 1 0 0 cd 6 cd 1 1 I I • • W X Q» 0 - 0 ■sr CN in 00 CN o cn r- VO 00 53 VD o IS p o H > in ro P CN Q o X 0 P 1 1 1 1 1 1 1 1 1 • a rH CP. r-» 0 H H P O H o P - 158 i—I cn r-'- > H O H H O O P CN H H cn O o P Cn lo G cn VD o VD CP. r- O H H CN ro > H P O H H o P 0 '=* 0 o I vo 0 G Cn i 1 1 d | vd 1 o 0 o in 00 CN o r (N vn ■sr w n i o cn LO •r| W VD M X o oCN ro P 0 o0 o P 0 P P G 0 B I I 1 CN ^ i—i > H CP. r- H H 0 O CN H H r- r O r~ 0 G P n t 1 p ■H a O ■— i P P g 0 - r 0 0 0 cn G i •H CN P 0 P P 0 r-G - r G 6 1 CN ro VD -r| H X U ro P P 0 o 0 B I I CN ro Appendix 3. (Cont. •H -H ■H -H ■H ffl cn ^ P 0 0 fd P p co CU s 0 U £ t f g G CO 0 r* tf 1 1 1 1 o fd fd ft o o 1 0 P '—■ ,£ cn P -p u • -p a ■p t f a P -p -p fd fd o 0 ft fd CO fd £ tr> p £ 0 • . , . M1 1— 1 1 CM CM P r- P CM 1— 1 CM in CO CO CD 1 H H H 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 i1 i 1 1 1 1 1 1 1 1 1 1 1 1 1fd 1 1 1 —1 1 1— 1 o in co CM CM CO O ^ O 00 > HH rH H Uo I (Uo £ tC | ft CO 1 0 I 1 ^ 1 1 1 1 c 1 co 1 1 1 1 1 1 1 1 1 1 1-H 1 1 1 1 1 1 1 1 1 1 1 1 cn 1— 1 00 1 •H M1 CM 00 CM 00 CM cn p p CO O CD cn LO CO pH 1— 1 4 EH cn o o -P T >1 TS P p fd dP£ > P 160 d td 0 * 0 0 -P 0 * 0 d 0 O'! £ o H p t" P tJV 0 0 161. -P He rt rt He 0 0 0 00 0 g ■H 162 3 * 0) 0 * 0 P ■H r-* s 3 P cn p 0 d cn a) e os CO Xi P d Eh cn CO 0 CD p 0 Eh • 3 „—. 0 CD CM P cn P • in d m CO cd 'a 163. LITERATURE CITED Abeloos, M. 1933. Sur quelques anomalies des pinces de Carcinus maenas Pen. Bull, Soc. Lin, Normandie Caen, Series 8 , 5: 15-18;26-27. Adensamer, T, 1898. 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VITA Elizabeth Gayle Lewis The author was born in Newport News, Virginia on January 17, 1950. She worked during the summers of 1966, 1967 and 1968 at the Virginia Institute of Marine Science in the departments of Microbiology, Geology and Photography. In June 1972, she was graduated cum laude from Mary Washington College of the University of Virginia with a B. S. in Biology. In September 1972, the author entered the College of William and Mary and held a teaching assistantship in that department. Upon transferal to the School of Marine Science in January 1973, the author began work as a graduate assistant in the department of Crustaceology. 176