J. Exp. Mar. Biol. Ecol., 1989, Vol. 131, pp.

215-222 215
Elsevier

JEMBE 01314

The effects of holding space and diet on the growth of the

West Indian spider crab ~~t~~ux ~p~~os~s~~~u~ (Lamarck)

Dara H. Wilber and T. Payson Wilber, Jr.

Smithsonian Laboratory, Grand Turk, Turks and Caicos Islands, British West Indies

(Received 20 March 1989; revision received 27 June 1989; accepted 6 July 1989)

Abstract: The effects of holding space and diet on molt increments and intervals were examined in the West
Indian spider crab Mithrax spinosissimus (Lamarck). Juvenile crabs were held individually for 4 months in
either 15 or 7.5 cm diameter rings and fed either macroalgae and conch mantle, macroalgae alone or
filamentous algae (algal turf). Both holding space and diet affected molt intervals more than molt increments
and holding space had a greater influence on both growth parameters than diet. Crabs held in the large rings
had shorter molt intervals and greater molt increments than those in small rings. The relative effects of diet
on growth were macroalgae and conch mantle > algal turf > macroalgae alone. For some growth
parameters, coefficients of variations were larger for treatments that reduced growth than treatments that
enhanced growth. After 2 months, some crabs in treatments that adversely affected growth (small ring and
no meat supplements) were transferred to large rings and given conch mantle. Growth rates improved for
these crabs primarily via reduced molt intervals.

Key words: Aquaculture; Diet; Growth; Space; Spider crab

Crustacean growth rates have two components, the size increase at a molt (molt
increment) and the time elapsed between molts (molt interval). ~oit-~crement data can
be easily obtained from newly molted individuals and their former exoskeletons. How-
ever, molt-interval data require maintaining individuals through successive molts and,
therefore, are rare (Botsford, 1985). In general, molt increments decrease and molt
intervals increase as body size increases and molt intervals and temperature are
inversely related (Mauchline, 1976). Other environmental effects on crustacean growth,
however, are not as easily generalized. Additional data, especially for molt intervals, are
required to understand the variability of molt increments and intervals. Aquaculture
projects are a good source of growth information because large numbers of individuals
are reared under controlled conditions for long time periods.
We tested the effects of holding space and diet on the growth and survival of the West

Correspondence address: D. H. Wilber, Biology Unit I, Florida State University, Tallahassee, FL 32306,
U.S.A.

0022-098 l/89/%03.50 0 1989 Elsevier Science Publishers B.V. (Biomedical Division)

216 D.H. WILBER ANDT.P.WTLBER,JR.

Indian spider crab ~~t~~u~ sp~nosjssim~s(Lamarck). Holding space affects the growth
of homarid lobsters (Shleser, 1974; Aiken & Waddy, 1979; Van Olst & Carlberg, 1979)
and may be important in the aquaculture of M. spinosissimus if aggressive behaviors
necessitate ~di~du~ rearing. The natural diet of M. spi~osissim~ is unclear, this
species has been described as an omnivore (Hazlett & Rittschof, 19’75) and as an
herbivore (Adey, 1987). We did not determine the natural diet of M. spinosissimus but
examined how diet affected growth. These data may be useful in determining whether
the availability of animal matter to M. spi~osissim~s under natural conditions affects
growth rates as in the lobster Punulirw cygnus (Jo11& Phillips, 1984).

MATERIALS AND METHODS

Two cohorts of&$. spi~os~ssimusthat hatched on the same day were reared in separate
field cages anchored to the bottom of a lagoon along the east side of Grand Turk, Turks
and Caicos, British West Indies. When the crabs were 90 days old, they were transferred
to the laboratory and kept in an outdoor flow-through seawater system. The experiment
was divided into two parts that correspond to age 90-157 days and age 158-217 days;
these will be referred to as Part I and Part II, respectively. During Part I, crabs were
reared individu~ly in either 7.5 or 15 cm diameter rings 6 cm high that were made of
7 x 7 -mm plastic mesh. Crabs were fed ad Iibitum either filamentous algae, macroalgae
or macroalgae supplemented with 1 g wet wt of conch mantle, the tissue of Strombus
gigas (Linnaeus) that is discarded during processing, The lilamentous algae were
cultured on O.&m* screens in a lagoon and consisted of many species that are collectively
called algal turf (Adey, 1987). Crabs fed algal turf were held in bottomless rings placed
on top of the algal screens at a density of 25 ind . m - ’ and the rings were moved daily
to ungrazed portions of the screens. Macroalgae were mixed species clumps (50 ml)
consisting primarily of Padi~apavonica, ffa~imeda opuntia, H. incrassata and Dasia sp.,
species that were abundant on local patch reefs and eaten by juvenile crabs (pers. obs.).
Uneaten food was removed from the rings and replaced with fresh food every 4 days.
The crabs were exposed to ambient light and water temperature ranged from 28 to
30 “C. Crabs were randomly assigned to the six treatment combinations, 22
ind . treatment - ‘, and each treatment was equally represented in two watertables,
Crabs were checked at 0800 and 2000 for deaths and recently molted indi~duals.
Deaths associated with molting were defined as crabs that died either during ecdysis,
within 3 days of ecdysis (post molts) or with a pigmented epicuticle beneath their present
exoskeleton (premolts) which was determined by dissection.
At the end of Part I, growth differences between treatments were tested by ANOVA
or, if the growth parameter was correlated with carapace length (CL),ANCOVA. The
following growth p~ameters were examined: molt increments (the difference between
old and new CL divided by the old CL),molt intervals, number of molts * crab- ‘, final
CL and difference between initial and final CL (delta). Preliminary analyses of both Part
 EFFECTS OF SPACE AND DIET ON SPIDER CRAB GROWTH 217

I and Part II data showed no block effects (i.e., differences in growth between the two
water-tables) or differences between hatches, thus, a two-factor ANOVA or ANCOVA
that used diet and ring size as factors was done unless otherwise specified. Analysis of
molt increments did not include the first molt in the experiment because it may have
reflected the prior rearing conditions of the crabs. In cases where a crab molted three
times in Part I (16% of the cases), the mean molt increment of the second and third
molts was used in the analysis. Molt-increment data were arcsine-square-root trans-
formed prior to analysis. Coefficients of variation (SD * X- ’ x 100%) were ranked from
high to low by treatment for the following growth parameters: size at the end of Part
I, molt increment, delta and molt interval. These rankings were compared to the
rankings of the treatment’s effect on growth enhancement by Spearman’s rank correla-
tion test. A three-factor G test that used ring size, diet and state (alive/dead) as factors
tested for treatment effects on survival at the end of Part I.
Part II of the experiment tested whether the effects of treatments that slowed growth
rates during Part I could be reversed. Half of the crabs in small rings were transferred
to large rings and half of the crabs on the algal turf diet were given a conch mantle
supplement. Otherwise, crabs were treated the same as in Part I. Thus, crabs were
divided into three exclusive groups. The first group consisted of crabs whose living
conditions did not differ between Parts I and II, these crabs were in large rings and fed
either macroalgae or macroalgae/conch mantle. The second group consisted of crabs
that were in large rings and fed algal turf during Part I but may or may not have been
given conch mantle during Part II. The last group consisted of crabs that were in small
rings during Part I and were divided into the eight possible combinations of ring size
(small or large), algae (macro or turf) and conch mantle (present or absent). These
divisions required separate analyses for the effects of living conditions on growth.
Treatment differences in the first two groups were examined via inspection or t tests,
differences in the last group were examined via three-factor ANOVA or ANCOVA if
the growth parameter was correlated with age 158 CL. Small sample sizes in several
treatments precluded statistical analysis of survival during Part II.

RESULTS

PART I: AGE 90-157 DAYS

Ring size significantly affected molt interval, final CL, number of molts and delta; molt
increment was marginally affected by this variable (Table I). Diet significantly affected
final CL, number of molts and delta and had a marginal effect on molt interval (Table I).
Growth was faster for crabs in large rings and, within each ring size, relative growth
rates for the diets were macroalgae/conch mantle> algal turf>macroalgae for all
growth parameters except molt increments (Table II, Fig. 1). The only apparent
synergistic effect between diet and ring size was for delta (Table I), reflecting a greater
size change for crabs held in the large rings and fed the macroalgae/conch mantle diet;
218 D. H. WILBER AND T. P. WILBER, JR

TABLE I
Results of two-factor ANOVAs that examine effects of diet and ring size on growth variables for Part I (age
90-157 days). Delta is difference between initial and final CL.

Growth variables Expt. factor F ratio P

Diet 2.66 0.070

Molt interval Ring 12.19 0.001
Diet * Ring 0.07 0.930
Diet 8.83 <O.OOl
Final CL Ring 17.34 <O.OOl
Diet * Ring 0.80 0.450
Diet 5.61 0.005
Number of molts Ring 12.09 0.001
Diet * Ring 1.09 0.342
Diet 10.36 <O.OOl
Delta Ring 18.20 < 0.001
Diet * Ring 2.63 0.077
Diet 0.69 0.505
Molt increment Ring 3.73 0.057
Diet * Ring 0.25 0.781

TABLE II

Means of growth variables and survival rate for Part I (age 90-157 days). Diets are algal turf screens (turf),
macroalgae (macro) and macroalgae/conch mantle (M-C); ring diameters are 15 cm (Lg) and 7.5 cm (Sm).
Delta is difference between intitial and final CL. Density indices (DI) are ratio of container-floor surface area
to individual’s squared CL. Mean initial crab size was 12.1 + 0.5 mm CL and did not differ between
treatments.

Treatment Final size Molt increment Number of Delta Molt Survival Final
(mm CL) (%) molts (mm CL) interval (%) Dl
(days)

Lg M-C 23.0 31.8 2.4 11.3 26.3 77 33.4

Turf 20.4 29.6 1.9 7.5 29.7 73 42.5
Macro 18.6 32.1 1.8 6.7 31.6 64 51.1
Sm M-C 19.2 29.5 1.8 6.8 33.2 73 12.0
Turf 17.9 28.3 1.7 6.1 37.1 86 13.8
Macro 16.8 28.5 1.4 4.8 40.6 73 15.7

however, the statistical significance of this interaction was marginal (F = 2.63,

P = 0.077). Pratios for tests concerning molt intervals and number of molts were larger
than those for molt increments (Table I). In general, mean molt intervals for crabs held
in large rings were 7.8 days shorter and mean molt increments were 2.4% greater than
those of crabs given the same diets but held in small rings. Adding conch mantle to the
macroalgal diet decreased mean molt intervals by 7.4 and 5.3 days in the small and large
rings, respectively. Molting was not synchronous, either within or between tables, with
the exception of a high incidence of molting during the 1st wk when 34% of the crabs
 EFFECTS OF SPACE AND DIET ON SPIDER CRAB GROWTH 219

30

zi
0

$ 2o
LARGE
ii SMALL
m 10
m
d
u

0
M/C T M
DIET
Fig. 1. Crab size at end of Part I for crabs in large (solid bars) and small (open bars) rings. M/C, macroalgae
with conch mantle; T, algal turf; M, macroalgae alone. Error bars depict + SD.

molted. Finally, molting occurred at equivalent frequencies during the day (54%) and
night (46%).
Coefficients of variation for delta were larger in the treatments in which delta was
small, e.g., the small ring x algal turf and small ring x macroalgae treatments
(Table III). Treatments with larger deltas, i.e., large rings x macroalgae/conch mantle
and large rings x algal turf, had smaller coefficients of variation.
There were no significant treatment effects on survival during Part I (all P
values > 0.4), overall survival was 74% (Table II). 39% of the deaths that occurred in
all treatments during Part I occurred during the first 5 days of the experiment and 12%
were associated with molting.

PART II: AGE 158-217 DAYS

Crabs kept in large rings for the entire experiment and given the macroalgae/conch
mantle diet continued to exhibit faster growth than crabs fed only macroalgae

TABLE III

Coefficients of variation (SD. X- ’ x 100%) for each treatment and tests by Spearman’s rank correlation
coefficient (I,). Treatment ranks for each of four growth parameters were based on X values in Table II.

Treatments Growth parameters

Final size (mm CL) Molt increment Delta (mm CL) Molt interval (days)

Large/M-C 17 19 34 24
Large/Turf 11 10 25 32
Large/Macro 15 20 36 29
Small/M-C 20 22 50 20
Small/Turf 18 17 49 31
Small/Macro 17 23 54 29
Ts - 0.24 - 0.09 - 0.77 - 0.24
P > 0.32 > 0.46 = 0.05 > 0.32
220 D. H. WILBER AND T. P. WILBER. JR.

TABLE IV

Results from Part II (age 157-217 days). Sample size (n) indicates number of crabs that molted in each
treatment combination during this 60-day period. Ring sizes are crabs kept in large rings during both Part I
and Part II (OL), crabs in small rings during both parts [OS) and crabs transferred to large rings for Part II
(NL).

Ring Diet n Increment Interval Delta Final size Nonmolting

size (%) (days) (mm CL) (mm cr_) (%)

OL Turf 5 29.6 50.8 9.5 30.7 38

OL T-C 3 36.9 36.7 13.7 34.7 57
OL Macro 9 21.8 51.7 6.1 26.2 40
OL M-C 8 31.2 385 7.1 31.4 43
NL Turf 4 33.6 42.5 5.8 22.8 0
NL T-C 6 29.8 40.8 7.9 28.5 0
NL Macro 2 29.8 40.5 6.6 22.5 75
NL M-C 3 28.0 41.8 6.2 25.8 62
OS Turf 2 25.9 65.5 3.8 19.6 60
OS T-C 3 31.0 56.3 6.3 25.2 25
OS Macro 2 16.4 51.0 2.8 19.8 50
OS M-C 4 31.2 43.3 8.8 29.4 33

(Table IV). Of the crabs originally kept in large rings on an algal turf diet, those that
were given conch mantle supplements had larger molt increments (t = 5.07, P < 0.0005)
and shorter molt intervals (t = 6.48, P < 0.0005) than their counterparts without conch
supplements (Table IV). The combined effects of these variables resulted in larger deltas
for crabs fed algal turf with conch mantle (t = 1.79, P < 0.05) and larger final CL
(t = 3.16, P (0.005) compared to that of crabs fed algal turf alone.
The three-way ANOVAs on molt interval, delta and molt increment showed the
following significant effects. Transferring crabs from small to large rings shortened molt
intervals by 12.6 days (Table IV; F = 5.8, P = 0.027) and conch-mantle supplements
increased delta (F = 4.55, P = 0.047). In addition, an interaction between ring size and
the presence of conch mantle for molt increments (F = 5.11, P = 0.036) reflected higher
molt increments for crabs in the conch mantle x small ring trials. A three-factor
ANCOVA showed final CL (F = 4.96, P = 0.04) was greater for crabs that received the
conch-mantle supplements.
A mortality peak (23 y0 of all Part II deaths) occurred when the crabs were 169 days
old and was associated with a pump failure that left the crabs without running seawater
for 16 h. Molt-related mortality accounted for 11% of all Part II deaths. 13 % of all
deaths in the entire experiment involved crabs that survived for at least 60 days without
molting and eventually died. These crabs often hung upside-down on the wails of the
rings and did not respond when food was added to their ring. Causes of the remainder
of deaths in the experiment (40.8 %) were unknown. Mortality was equally common in
both watertables and among treatments and, with the exception of initial mortality and
that associated with the pump failure, it showed no temporal pattern.
 EFFECTSOF SPACEANDDIETONSPIDERCRAB GROWTH 221

DISCUSSION

In M. spinosissimus, as in other crustaceans (Botsford, 1985), molt intervals were

influenced more by environmental factors than were molt increments. Social conditions
(Bliss & Boyer, 1964; Cobb & Tamm, 1974) and temperature (Kurata, 1962, as cited
in Mauchline, 1976) also affect molt intervals in crustaceans. Molt interval durations
appear to be affected by altering the D, (first premolt) stage of the molt cycle (Cobb
et al., 1982; Anger, 1987). Holding space more strongly influenced growth of
M. spinosissimus than diet as evidenced by the greater F ratios for this variable. Addi-
tional evidence that holding space affects molt intervals more than molt increments was
that transferring crabs from small to large rings increased molt frequencies, however,
molt increments were not affected. A density index (the ratio of the container-floor
surface area to the individual’s squared CL) was developed to standardize holding space
in lobsters. Density indices of 75 (cL)~ (Aiken & Waddy, 1979) to 37 (cL)~(Van Olst
& Carlsberg, 1979) are adequate for unrestrained lobster growth. In our study, initial
density indices were 30 and 125 (CL)'in the small and large rings, respectively. By the
end of Part II, density indices ranged from 14.7 to 25.7 (CL)*for crabs held in large rings
throughout the experiment, therefore, the large rings may have limited growth in the
latter portion of the experiment. Mean growth rates of communally reared
M. spinosissimus exceed those obtained in this study (unpubl. data), possibly because
of larger holding space.
Animal protein (conch mantle) enhanced growth rates of crabs on both macroalgal
and algal turf diets. M. spinosissimus fed various types of commercial pellets often grew
faster when fed pellets with a high protein content (W. Lellis, pers. comm.). Animal
matter associated with algae enhanced the growth and survival of the majid crab
Microphrys bicornutus (Kilar & Lou, 1986) and may affect algal quality as a food
resource for M. spinosissimus under natural conditions. Macroalgae in our experiment
were not intentionally rinsed of epibionts but appeared free of commensal animals.
Likewise, algal turfs have an associated commensal fauna consisting largely of
nematodes, polychaetes and microcrustaceans. Crabs on either algal diet, therefore,
may have ingested animal matter as well. Crabs fed almost exclusively on the edges of
the macroalgae which may have a greater nutrient value than interior areas (pers. comm.
from A.R.O. Chapman as cited in Kilar & Lou, 1986). Regardless of ring size,
M. spinosissimus grew faster when fed algal turf than when fed macroalgae. The effects
of algal type on crab growth were less clear, however, when conch mantle was added
to the diet. Results of Part II indicate growth rates improve for individuals if conditions
that inhibit growth are removed.
Nonuniform growth rates were recognized in earlier M. spinosissimus aquaculture
efforts (W. Lellis, pers. comm.) in which nine-fold differences in CL existed for
6-month-old crabs from three hatches reared together. Genetic factors were not
addressed in this study but may have accounted for some of the variation in growth.
In our study, variation in growth parameters was commonly less than variation in
222 D. H. WILBER AND T. P. WILBER, JR.

homarid lobsters (Hedgecock et al., 1976) and equivalent to that of palaemonid shrimp
(Sandifer & Smith, 1975). The variation observed in shrimp growth was not attributable
to stocking density and may reflect the influence of genetic factors (Sandifer & Smith,
1975). Variance for the growth parameter delta in M. spinosissimus was smaller in
treatments with better growth which may reflect less variable treatment conditions. For
example, the nutritive value of different pieces of conch mantle may be less variable than
that of different assortments of macroalgae. Alternatively, environmental conditions
that favor growth may reduce variance whereas stressful conditions may cause more
variable growth rates. Variance was typically greater for molt intervals than for molt
increments (Table III) which also indicates that molting frequency is a more plastic
growth parameter.
Juvenile M. spinosissimus growth was influenced both by holding space and diet
mainly because of changes in molting frequencies. Although molt increments were
significantly affected by treatment conditions, molt intervals were more strongly affected
and showed greater variance. Although M. spinosissimus survive and grow on algal diets,
animal-matter supplements accelerate growth rates. The amount of animal matter
ingested by crabs grazing in the field and possible correlations between diet and
population differences in size distribution (W. Adey, pers. comm.) need to be addressed.

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