Marine Biology (2003) 142: 77–92
DOI 10.1007/s00227-002-0920-8
C. Thurman
Osmoregulation in fiddler crabs (Uca) from temperate Atlantic
and Gulf of Mexico coasts of North America
Received: 9 February 2002 / Accepted: 16 July 2002 / Published online: 14 September 2002

Springer-Verlag 2002
Abstract Fiddler crabs (Crustacea: Decapoda; genus
Uca) were collected from 16 locations along the tem-
perate Atlantic and Gulf of Mexico coasts of eastern
North America for osmoregulation studies. Three spe-
cies, U. pugilator, U. pugnax and U. minax, were taken
from habitats between Cape Cod, MA, and St. Mary’s
River, Nassau Co., GA Seven species, U. panacea, U.
speciosa, U. spinicarpa, U. longisignalis, U. rapax, U.
pugilator and U. minax, were collected in habitats be-
tween the Ochlochonee River, Liberty Co., and
Thompson’s Bayou, Escambia Co., along the Gulf coast
of northwest Florida. To examine differences in osm-
oregulatory capabilities among the species, specimens
were placed in various concentrations of artificial sea-
water (0–3450 mOsm) for 5 days. The oligohaline or
‘‘freshwater’’ species, U. minax, U. spinicarpa and U.
longisignalis, possess the lowest average hemolymph
osmolality. They are unable to control hemolymph os-
molality above 2000 mOsm. On the other hand, the
euryhaline species, U. speciosa, U. panacea, U. pugilator
and U. pugnax, have much higher average hemolymph
osmolality. They are able to withstand an osmotic
challenge of 2200 mOsm or greater. Among the eight
species, U. panacea and U. pugilator are able to os-
moregulate across the broadest range of seawater con-
centrations (0–2800 mOsm). After an examination of the
osmoregulation in several populations of U. minax, U.
pugilator and U. pugnax, physiological adaptation is
apparent in the two former species, but not the latter.
Experimental evidence for capacity adaptation was ex-
amined in the freshwater species, U. minax. If slowly
Communicated by P.W. Sammarco, Chauvin
C. Thurman
Department of Biology, University of Northern Iowa,
Cedar Falls, IA 50614-0421, USA
E-mail: Thurman@uni.edu
Tel.: +1-319-2732276
Fax: +1-319-2732276
adapted to hypertonic seawater (1800 mOsm) for
14 days, this species is able to withstand much higher
osmotic pressures than unadapted crabs. Although he-
molymph isosmotic concentration remains the same, the
limits for regulation are extended to higher osmolality.
Consequently, as a result of studying osmoregulation in
several species of Uca captured at various locations
across their geographic range in the temperate zone, a
clearer view of water-balance physiology is available for
the genus. The Uca spp. are not uniform in osmoregu-
latory abilities. There is considerable inter- and intra-
specific physiological variation associated with the
ecological distribution of each species, respectively.
Introduction
Fiddler crabs in the genus Uca are circumtropical and
found in most temperate regions (Crane 1975). The
temperate Atlantic and Gulf of Mexico coasts of the
eastern United States are home to 12 species of fiddler
crabs (Barnwell and Thurman 1984). Bisected by Flori-
da, the two ‘‘temperate’’ coastlines differ dramatically in
their species composition. Between southern Massa-
chusetts and northern Florida, the Atlantic region has
three species: Uca pugilator, U. pugnax and U. minax.
The temperate gulf region, extending from Tampa Bay,
FL, to the Rio Grande, TX, hosts nine species, which are
considered either temperate, endemic, or tropical:
U. pugilator, U. minax. U. panacea, U. longisignalis,
U. spinicarpa, U. subcylindrica, U. rapax, U. speciosa and
U. vocator. On peninsular Florida, there are five tropical
and one temperate species: U. pugilator, U. rapax,
U. speciosa, U. burgersi, U. thayeri and U. leptodactyla.
Undoubtedly a large number of abiotic and biotic fac-
tors has influenced species diversity in the disjunct
temperate regions. One reason for the species disparity
between the two areas is that the temperate Gulf
region stretches for about 1100 km, paralleling the
30N latitude, while the eastern Atlantic seaboard is
78

perpendicular to the tropics. The temperate Atlantic The present study examines the osmoregulatory ca-
coast extends approximately 1600 km north from Flor- pabilities in seven species of Uca from different locations
ida to southern Cape Cod, covering 12 of latitude. along the temperate Atlantic and Gulf of Mexico coasts
For ectothermic invertebrates, climatic temperature is of the United States. The three temperate Uca from
a primary factor controlling biogeographic distribution. Atlantic and Gulf populations were challenged with
However, for marine and aquatic benthic species, sa- artificial seawater (ASW) varying from 30 to
linity and osmotic gradients are also highly significant 3400 mOsm. In addition, four species of fiddler crabs
determinants. Given the primary selective influence of from the temperate zone in northwestern Florida were
temperature, habitat selection among different Uca spp. also examined. Both survivorship and hemolymph os-
within a specific area can be linked to osmotic regulation molality are reported over the osmotic regime. Since
and salinity preference (Teal 1958). To partition the collections were made across the range for several spe-
habitat ecologically, crabs dig burrows among the veg- cies, this investigation demonstrates differences in both
etation in brackish marshes or above the high-tide mark interspecific as well as intraspecific physiology among
along the banks of rivers. Some species are known to live Uca.
in freshwater, while others live on arid beaches near
ephemeral hypersaline lagoons (Thurman 1984). To in-
habit such a wide variety of environments, ionic and Materials and methods
osmotic regulation is expected to be a key physiological
characteristic determining habitat selection among dif- Collections
ferent species. Generally, semiterrestrial and terrestrial
Fiddler crabs were collected from 16 locations along the east coast
crabs are excellent regulators under extreme osmotic of the United States (Fig. 1). Only non-ovigerous, intermolt adult
stress (Jones 1941). However, of the 90–100 species specimens with carapace width >10 mm were used in experiments.
within the genus Uca, osmoregulation has been studied In Massachusetts, Uca were collected between 13 and 16 July 2001
in only a few. Consequently very little is known about
the differences in physiological responses expressed by
species when they are challenged osmotically.
Several investigations have studied the osmotic regu-
lating capabilities of Uca from the east coast of the United
States. Green et al. (1959) examined iono- and osmore-
gulation in the temperate Atlantic species found around
Woods Hole, MA. When U. pugilator and U. pugnax were
adapted to 175% seawater, they were observed to be
excellent hypo-osmotic regulators. Unfortunately, sam-
ples of body fluids from each species were pooled for
the measurements. Baldwin and Kirschner (1976a,b)
examined Na+ and Cl– regulation in U. pugilator and
U. pugnax adapted to either 10% or 175% seawater. Since
the two species were assumed to have identical osmore-
gulation capabilities, the investigators combined hemol-
ymph samples for their analysis. As the medium varied
from 10% to 175% seawater, hemolymph osmolality
varied from 410 to 630 mOsm. They reported ‘‘Uca’’ to
maintain a hemolymph osmotic pressure around
500 mOsm. Although Wright et al. (1984) studied each
species separately, they found only ‘‘slight differences’’ in
the osmoregulatory capabilities of U. pugilator, U. minax
and U. pugnax as the medium was varied from 5% to
150% seawater. Holliday (1985) reported U. pugnax to be
a good osmotic regulator in media with osmolality up to
2000 mOsm. Rabalais and Cameron (1985) examined
two temperate-Gulf fiddler crabs, U. subcyindrica and
U. longisignalis, and found both to be exceptional regu-
lators in medium of up to 3000 mOsm. Consequently, our Fig. 1 Uca collection sites along the temperate Atlantic and Gulf
understanding of the ecological significance of osmore- of Mexico coasts of the United States (A Waquoit Bay; B Wild
gulation in this taxon is based, for the most part, on the Harbor River; C Sippican River; D Slocum River; E Taunton
assumption that the different species have identical River; F Mt. Hope Bay; G Colt State Park; H Isle of Wight; I Point
Lookout State Park; J Crooked River; K St. Mary’s River; L
capabilities. Thus, we have no clear description drawing a Ochlochonee River; M Alligator Point; N Money Bayou; O St.
relationship between physiological capability and eco- Joseph Peninsula; P Thompson’s Bayou). Scale bar: 200 miles
logical diversity within the genus. (322.6 km)

at: (1) Waquoit Bay (790 mOsm) near State Highway 28, East
Falmouth (4133¢N; 7029¢W) and (2) Wild Harbor River
(874 mOsm), North Falmouth (4138¢N; 7038¢W), in Barnstable
Co.; (3) Sippican River (178 mOsm), Wareham, in Plymouth Co.
(4145¢N;7043¢W); (4) the Lloyd Center and Demarst Lloyd State
Park (882/887 mOsm) on the Slocum River (4131¢N;7056¢W); (5)
Taunton River (636 mOsm), Somerset (4144¢N; 7108¢W); and (6)
Mt. Hope Bay (777 mOsm), Swansea (4143¢N;7057¢W), in Bristol
Co. In Rhode Island, crabs were taken at: (1) Colt State Park
(777 mOsm), Bristol Point (4139¢N; 7116¢W), Narragansett Bay,
in Bristol Co. In Maryland, fiddler crabs were collected from (1) the
Isle of Wight (863 mOsm) near the US 50 bridge to Ocean City in
Worcester Co. (3820¢N; 7505¢W) and from (2) Point Lookout
State Park (382/389 mOsm) near Scotland in St. Mary’s Co.
(3802¢N; 7619¢W) on 20 and 22 May 2001, respectively. In
southern Georgia (3043¢N; 8129¢W) and northwestern Florida
(2943¢N; 8450¢W), crab colonies were sampled between 13 and 16
March 2001 at: (1) St. Mary’s River (380 mOsm) at US 17 bridge
near Woodbine, Nassau Co., FL.; (2) Crooked River (885/
935 mOsm), at Elliot’s Bluff, Camden Co., GA; (3) Alligator Point
(722 mOsm) near St. Teresa, Franklin Co., FL; (4) Ochlochonee
River (14 mOsm) near US 319, Liberty Co., FL; (5) Money Bayou
(32 mOsm) at SR 30 bridge near Indian Pass, Gulf Co., FL; and (6)
St. Joseph Peninsula (987 mOsm), Gulf Co., FL. On 8 October
2000, crabs were collected from Thompson’s Bayou (70 mOsm)
near Pensacola, Escambia Co, FL (3031¢N; 8710¢W).
Crabs were transported to the laboratory in plastic boxes
containing water from the collection site. Within 24 h they were
submitted to laboratory conditioning at 23C. The collections were
divided into three groups. The first consisted of five crabs of each
species held in habitat water. For collections of more than 80
specimens, a second group was submitted to the full regime of
osmotic challenge. A third group consisting of 25 or more speci-
mens was kept in habitat water for 30 days before experimentation.
Water was replaced every 3-4 days. In every case, crabs were able
to leave the medium at will. For long-term maintenance, Uca were
fed goldfish food (Tetra; Melle, Germany) and frozen brine shrimp.
Osmotic solutions
To obtain different concentrations of ASW, Instant Ocean
(Aquarium Systems, Mentor, Ohio) was mixed with distilled H2O
to the desired density and the pH adjusted to 8.0 with HCl or
NaOH. Groups of four to six crabs were placed in covered bowls
(13·5 cm) containing 50 ml of ASW for 5 days. To eliminate waste
and maintain a constant osmolality, ASW solutions were replaced
on day 2 and day 4 of each experiment. Uca were not fed 3 days
prior to or during an experiment. Survivorship records were kept
during the exposure period. A crab was considered deceased if it
could no longer right itself. Only living Uca were used for osmotic
measurements.
Osmolality
Following osmotic challenge, hemolymph was withdrawn from the
ventral hemocoel via puncture of the arthrodial membrane between
the basis and coxa of the fifth pereiopods using a chilled 1.0 ml
tuberculin syringe (27 gage needle). The osmolality (mmol kg–1) of
each 10 ll hemolymph sample was determined immediately using a
Wescor 5520 vapor pressure osmometer (Logan, Utah) by com-
parison to commercial standards. Water samples from finger bowls
and habitats were also measured. Standard concentrations and
osmolality measurements were identical between 60 and
3200 mOsm. However, between 1 and 55 mmol kg–1, the actual
osmolality was determined using a standard curve constructed from
serial dilutions of a 100 mOsm standard. The osmolality of double-
glass distilled, deionized H2O was between 2 and 7 mOsm.
Fluid osmolality was taken as the average of all hemolymph
samples from each crab or water sample. For each measurement,
an average for all crabs is reported (mean±standard error). Sta-
tistical significance was assessed using Students’ t-test with P<0.05
79
significant and P<0.01 highly significant. In general, ‘‘N’’ refers to
the number of crabs unless otherwise stated. Probit analysis (Fin-
ney 1947) was used to estimate the 50% lethal osmotic concen-
tration (LC50), while the isosmotic medium concentration was
calculated using linear regression.
Results
Field studies
Collections
Between Cape Cod, MA, and Pensacola, FL (Fig. 1),
eight species of Uca were collected from an array of
temperate habitats (Table 1). As a generalization, these
species can be classified by habitat and taxonomic group.
Habitats can be classified into one of three saline envi-
ronments based on osmolality. Freshwater (FW) or oli-
gohaline habitats are considered to have an osmolality
between 0 and 380 mOsm, brackish (BW) or mesohaline
between 381 and 630 mOsm and hypersaline or euryha-
line (EH) >631 mOsm. Likewise, the eight species of
fiddler crabs can be categorized into two of the subgenera
proposed by Crane (1975) (see also Barnwell and Thur-
man 1984). The four species in the Celuca subgenus are U.
spinicarpa, U. speciosa, U. panacea and U. pugilator. The
four species considered to be members of the Minuca
subgenus are U. minax, U. longisignalis, U. pugnax and U.
rapax. As shown in the following sections, each subgen-
eric grouping is represented among the three environ-
mental osmotic regimes (i.e. FW, BW and EH).
Among the eight species of fiddler crabs found, three,
U. spinicarpa, U. speciosa and U. longisignalis, were
captured at only one location during this study. Further,
collections of U. panacea and U. rapax were limited to
two locations. The remaining three species, U. pugilator,
U. pugnax and U. minax, were taken throughout the
study area. Two locations are of particular interest.
Point Lookout, Md. (385 mOsm), is the only site where
all three Atlantic species, U. pugilator, U. pugnax and U.
minax, were encountered simultaneously. From an eco-
physiological perspective, environment conditions at this
location are permissive for their rare coexistence. On the
Gulf coast, Money Bayou (32 mOsm) is a habitat with
an unusually high Uca species diversity. Five species of
fiddler crabs were collected from the tidal creek: U.
spinicarpa, U. panacea, U. minax, U. longisignalis and U.
rapax. As pointed out previously (Barnwell and Thur-
man 1984), the region between Tampa Bay and Cape
San Blas, FL, which includes Money Bayou, forms an
ecotone between temperate and tropical biomes. Typi-
cally the ‘‘edge-effect’’ in such an area maintains high
species diversity. Uca spp. from collection sites in Mas-
sachusetts, southern Georgia and western Florida live
close to the edge of their geographic ranges.
As illustrated in Table 1, U. minax, U. spinicarpa and
U. longisignalis are inhabitants of FW sites. During the
present study, U. spinicarpa and U. longisignalis were
collected in a habitat with an osmolality of 32 mOsm.
80

Table 1 Uca spp. Relation between habitat and hemolymph osmolality for Uca from the eastern United States. For each species, N=5
specimens, except where noted in parentheses (FW freshwater; BW brackish water; EH euryhaline; df degrees of freedom)
Subgenus, Habitat Collection Habitat Habitat Hemolymph Hemolymph df P
species preference site osmolality salinity (&) osmolality osmolality
(mOsm) (mOsm) after 30 days
(mOsm)

Celuca
U. spinicarpa FW Money Bayou, FL 32 1 605±46
U. speciosa EH Alligator Point, FL 722 25 849±70
U. panacea BW/EH Money Bayou, FL 32 1 709±34
Alligator Point, FL 722 25 845±8 862±11 8 NS
U. pugilator BW/EH Sippican River, MA 178 6 791±31
Point Lookout, MD 385 13 809±18 822±10 8 NS
Taunton River, MA 636 22 817±27
Alligator Point, FL 722 25 871±24 881±22 8 NS
Mt. Hope Bay, MA 777 27 859±16
Colt State Park, R.I. 777 27 869±25
Waquoit Bay, MA 790 27 893±47
Isle of Wight, MD 863 27 879 (1)
Wild Harbor 874 30 901±13
River, MA
Crooked River, GA 935 34 917±35 864±20 8 NS
St. Joseph Pen., FL 987 34 1043±36 920±22 8 0.1>
Minuca
U. minax FW Ochlochonee 14 <1 560±30
River, FL
Money Bayou, FL 32 1 636±52 621±29 8 NS
Thompson’s 70 2 600±25
Bayou, FL
Sippican River, MA 178 6 621±15
St. Mary’s River, GA 380 13 659±28 640±5 8 NS
Point Lookout, MD 385 13 624±11 608±11 8 NS
U. longisignalis FW/BW Money Bayou, FL 32 1 640±15 692±24 8 NS
U. pugnax BW/EH Point Lookout, MD 385 13 749±13 783±17 8 NS
Taunton River, MA 636 22 738±33
Colt State Park, R.I. 777 27 798±7
Mt. Hope Bay, MA 777 27 784±14
Waquoit Bay, MA 790 27 793±23
Isle of Wight, MD 860 29 843±4 836±9 8 NS
Wild Harbor 874 30 877±30
River, MA
Crooked River, GA 885 31 841±39 831±16 8 NS
Slocum River, MA 885 31 822±16
U. rapax EH Money Bayou, FL 32 1 668 (1)
Alligator Point, FL 722 25 814±35 807+34 8 NS

U. minax were taken from several habitats with osmol-
ality ranging between 14 and 385 mOsm. Species typi-
cally found in EH habitats are U. speciosa, U. panacea,
U. rapax, U. pugilator and U. pugnax. During this study,
U. speciosa, U. panacea and U. rapax were collected in
722 mOsm habitats. U. pugilator were taken from hab-
itats ranging in osmolality from 722 to 987 mOsm, while
U. pugnax were collected in habitats ranging from 777 to
885 mOsm. Based on the current as well as previous
collections (Barnwell and Thurman 1984; Thurman
1984), U. spinicarpa, U. panacea, U. pugilator, U. longi-
signalis and U. pugnax are known to also occupy BW
habitats. In the present study, both U. pugilator and U.
pugnax were common in BW habitats and occasionally
occurred in FW habitats.
Across temperate coasts in eastern North America,
fiddler crab species from both subgeneric taxa, the Cel-
uca and the Minuca, have representatives occupying all
major three environmental salinity regimens. For the
Celuca subgenus, U. spinicarpa are typically FW and
occasionally BW, both U. panacea and U. pugilator
range from BW to EH, and U. speciosa are typically
limited to EH habitats. For the Minuca subgenus, U.
minax are typically FW, U. longisignalis are typically
FW and BW, U. pugnax are both BW and EH, and U.
rapax are usually EH. In the western Gulf, other pop-
ulations of endemic fiddler crab species are known to
occupy similar regimes (Thurman 1984, 1998).
Hemolymph osmolality
Hemolymph osmolality was measured in samples of
crabs from each environment (Table 1). A distinct
species–environment relationship is evident. In the
following discussion, ‘‘N’’ is the number of popula-
tions sampled. The FW species, U. spinicarpa, U. minax
and U. longisignalis, have hemolymph osmolality

<700 mOsm (618±28, N=8). On the other hand, the
EH species, U. speciosa, U. panacea, U. pugilator,
U. pugnax and U. rapax, have hemolymph osmolality
>800 mOsm (862±59 mOsm, N=17). As an indication
of its rare occurrence in FW, the individual U. rapax
collected at Money Creek, FL, had a hemolymph
osmolality of 668 mOsm, which is near the lower limits
tolerated by the species (Zanders and Rojas 1996). BW
populations of U. pugilator have hemolymph osmolality
of 813±4 mOsm (N=2), while U. pugnax have
744±6 mOsm (N=2) and U. minax have 642±
18 mOsm (N=2). It is interesting that at Point Lookout,
MD, where U. pugilator, U. pugnax and U. minax were
collected in sympatry (385 mOsm), all have significantly
different hemolymph osmolality (809±18, 749±13,
624±11, respectively, df=12, P<0.001). It is clear that
there are intraspecific as well as interspecific differences
in hemolymph osmolality.
U. pugilator, U. minax and U. pugnax were collected
from a sufficient number of habitats to allow assessment
of intraspecific variation in blood osmolality. Hemol-
ymph osmolality in U. pugilator varied from 800 to
1000 mOsm across sites, with salinity ranging from 178
to 987 mOsm. The regression relationship between
habitat (X) and hemolymph osmolality (Y) for U. pugi-
lator is Y=0.2234X+716 mOsm (N=11, r2=0.6356,
P<0.005). In habitats (X) where the salinity varied from
385 to 885 mOsm, hemolymph osmolality (Y) in
U. pugnax changed from 750 to 850 mOsm, with a linear
relationship of Y=0.2226X+635 mOsm (N=9,
r2=0.6365, P<0.005). In all habitats where the two are
sympatric, U. pugnax consistently maintained hemol-
ymph osmolality below that of U. pugilator. Although
U. minax were not collected in habitats with salinity
>385 mOsm, the hemolymph osmolality in this species
also varied between 560 and 650 mOsm . For this species
the regression of habitat (X) to hemolymph (Y) osmol-
ality is Y=0.1278X+594 mOsm (N=6, r2=0.4092,
P<0.1). Consequently, based on field observations,
there is significant intraspecific variation in hemolymph
osmolality among the three Atlantic species, U. pugila-
tor, U. minax and U. pugnax, over their habitats.
The relationship between the two Celuca species,
U. speciosa and U. spinicarpa, is very interesting. Crane
(1975) considered the two to be subspecies. However,
Salmon et al. (1979) collected U. spinicarpa within 50 km
of U. speciosa near Apalachicola Bay. After considering
their morphology, distributions, behavior and elec-
trophoretic patterns, the two have been reinstated to
their former status as full species (Barnwell and Thur-
man 1984). In the present study, the two were collected
within 100 km of each other. Since U. spinicarpa from a
FW habitat (32 mOsm) maintained a hemolymph os-
molality of 605±46 mOsm, while U. speciosa from a EH
habitat (722 mOsm) were 849±70 mOsm, the two ap-
pear to differ based on field measurements (df=8,
P<0.001). Further, as shown in the subsequent sections,
the two illustrate physiological divergence under labo-
ratory conditions.
81
Laboratory studies
After capture, Uca spp. from various locations were
returned to the laboratory for experimentation. To de-
termine the stability of hemolymph values, crabs from
some populations were maintained in water from their
respective habitat for 30 days. Hemolymph samples
taken afterwards were not significantly different from
those taken following capture (Table 1). Only the U.
pugilator collected at Port St. Joseph Bay, FL, appeared
to show any indication of change. These crabs decreased
hemolymph osmolality to match the osmotic pressure of
their surrounding water.
Survivorship
Seven of the eight species were collected in sufficient
numbers to examine their ability to withstand osmotic
stress over a 5-day period (Figs. 2, 3, 4). Fiddler crabs can
survive a wide range of osmotic stress. Survivorship for
three species in the Minuca group exposed to ASW
ranging in osmolality from 50 to 3450 mOsm is shown in
Fig. 2. Both U. longisignalis and U. minax survived oli-
gohaline conditions very well. There was no significant
difference in survivorship between Gulf and Atlantic
populations of U. minax. On the other hand, on the whole,
the U. pugnax populations were susceptible to habitat
osmolality <150 mOsm. Further, specimens from
Georgia populations did not withstand hypotonic ASW
as well as those from Massachusetts and Maryland. There
are differences in the ability of each species in the Minuca
group to survive in hypertonic ASW. The upper limit of
tolerated osmotic stress was calculated as the LC50 for
each species (Table 2). It is obvious that among the FW/
BW species, U. longisignalis did not survive well in ASW
with osmolality >1850 mOsm. The FW species U. minax
was susceptible to ASW >1950 mOsm, and the BW/EH
species U. pugnax to ASW >2700 mOsm.
Survivorship for the four species of the Celuca group-
ing in ASW are shown in Figs. 3 and 4. Both species
shown in Fig. 3 were susceptible to low osmolality.
U. spinicarpa did not survive well in ASW <100 mOsm,
while U. speciosa did not survive well in ASW
<400 mOsm. By contrast, the FW species U. spinicarpa
did not survive well above 1500 mOsm, and the EH spe-
cies U. speciosa above 2550 mOsm. Survivorship of the
sibling species U. pugilator and U. panacea is shown in
Fig. 4. Both were susceptible to ASW with osmolality
<100 mOsm. However, U. pugilator can withstood
50 mOsm ASW, but U. panacea could not. Also, speci-
mens of U. pugilator from Atlantic populations were
susceptible to ASW <200 mOsm, while those from the
Gulf of Mexico were not. On the other hand, in hypertonic
ASW, U. pugilator regardless of origin did not survive well
above 2900 mOsm, while U. panacea survived well up to
3350 mOsm. Consequently, from these observations,
there are significant differences in the ability of both
temperate Atlantic and Gulf species to survive osmotic
stress.
82

Fig. 2 Uca spp. Percent sur-

vival of species from the Minuca
subgenus under osmotic stress
for 5 days. ‘‘(N)’’ indicates total
number of specimens. Solute
concentration in osmolality
(mOsm)

Fig. 3 Uca spp. Percent sur-
vival for species in the Celuca
subgenus (U. spinicarpa and U.
speciosa) under osmotic stress
for 5 days. ‘‘(N)’’ indicates total
number of specimens. Solute
concentration in osmolality
(mOsm)
Regulation of hemolymph osmotic pressure
The hemolymph osmotic response of species of crab to
changing external osmolality from 0 to 3450 mOsm is
shown in Figs. 5, 6 and 7. As indicated by the graphs,
each species of fiddler crab is an excellent osmoregula-
tor. The patterns of regulation are best described as
type IV: hyper- and hyporegulators (Vernberg and
Vernberg 1972). Below 600 mOsm each was able to
maintain hemolymph osmotic concentration above the
concentration of the medium. Above 800 mOsm signif-
icant variation was seen among the species in their
ability to regulate hemolymph below medium osmolali-
ty. The patterns of osmoregulation for three species in
the Minuca grouping are shown in Fig. 5, while those for
the four species in the Celuca grouping are shown in
Figs. 6 and 7. In the discussion below, ‘‘N’’ is the
number of sample means from each population.
Among the Minuca, the two FW species, U. longi-
signalis and U. minax, were able to maintain a constant
hemolymph osmolality at 646±48 and 618±36 mOsm,
respectively, as the medium was changed from 0 to
83
600 mOsm. As the medium was changed from 0 to
300 mOsm for U. pugnax, hemolymph osmolality in-
creased significantly (N=7, r2=0.8860, P<0.005). For
U. longisignalis hemolymph osmolality (average
696±47 mOsm ) exhibited a linear correlation with the
media between 300 and 1400 mOsm (Y=0.1189X+
576 mOsm, N=5, r2=0.9101, P<0.01). The isosmotic
medium concentration was 653 mOsm, as shown in
Table 2. This relationship is similar for U. minax in
media with osmolality between 200 and 1000 mOsm.
Over this range, hemolymph had an average osmolality
of 671±43 mOsm. The correlation equation of medium
with hemolymph osmolality is Y=0.138X+580 mOsm
(N=9, r2=0.8455, P<0.01), with an isosmotic medium
concentration of 673 mOsm. For U. pugnax, hemol-
ymph osmolality was constant (901±89 mOsm) in me-
dia between 300 and 1900 mOsm (Y=0.0935X+
797 mOsm, N=19, r2=0.5122, P<0.02), with the isos-
motic concentration being 879 mOsm. All species be-
came less effective in controlling hemolymph osmolality
as the osmotic concentration of the medium was in-
creased. For U. longisignalis in media with osmolality
84
Fig. 4 Uca spp. Percent sur-
vival for species in the Celuca
subgenus (U. pugilator and
U. panacea) under osmotic
stress for 5 days. ‘‘(N)’’ indi-
cates total number of speci-
mens. Solute concentration in
osmolality (mOsm)
>1400 mOsm, the crabs lost the ability to control he-
molymph osmolality. In the cases of U. minax and U.
pugnax, they became less effective regulators in media
above 1000 and 1900 mOsm, respectively. During these
experiments, U. minax and U. pugnax were able to tol-
erate hemolymph osmolality as high as 1828 and
2118 mOsm, respectively. It is most interesting that the
hemolymph–medium curve for U. minax from the
Ochlochonee River (14 mOsm) lies to the left of other U.
minax populations from more saline habitats (32–
385 mOsm). A similar shift is not evident in the data for
other Minuca.
The hemolymph responses of the four species of the
Celuca grouping to the osmotic regime are illustrated in
Figs. 6 and 7, as well as Table 2. As shown in Fig. 6,
between 0 and 300 mOsm, hemolymph osmolality in-
creased from 605 to 700 mOsm in the FW species U.
spinicarpa (Y=0.314+605 mOsm, N=3, r2=0.7545,
P>0.10). In media between 300 and 1200 mOsm, the
osmolality of the hemolymph was controlled fairly
well (mean=745±29 mOsm; Y=0.0802X+678 mOsm,
N=3, r2=0.8584, P<0.01), and the isosmotic medium
concentration was 737 mOsm. Above 1200 mOsm,
U. spinicarpa did not control hemolymph osmolality
very well. For the EH species U. speciosa, as the medium
was increased from 100 to 2600 mOsm, there was a
significant linear change in hemolymph osmolality (Y=
0.3455X+624 mOsm, N=10, r2=0.9147, P<0.001).
For U. speciosa, the isosmotic medium concentration
was 953 mOsm, which contrasts sharply with U. spini-
carpa (737 mOsm).
The responses of the sibling species pair U. pugilator
and U. panacea are shown in Fig. 7 and Table 2. The
hemolymph response (Y) of U. panacea to increasing
medium osmolality (X) between 0 and 2000 mOsm is
described by the linear equation Y=0.0939X+
798 mOsm (N=8, r2=0.9426, P<0.001). The isosmotic
medium concentration for U. panacea was 880 mOsm.
The highest hemolymph concentration recorded for this
species was 1860 mOsm. For U. pugilator, hemolymph
responses to changes in medium osmolality appeared
more complex due to the number of populations
sampled. For medium osmolality between 26 and
300 mOsm, there was an increase in hemolymph from

Table 2 Uca spp. Summary of osmoregulation (in mOsm). LC50 is upper limit for survival of osmotic stress calculated by probit analysis
(FW freshwater; BW brackish water; EH euryhaline)
Subgenus Species Habitat
Celuca U. spinicarpa FW
U. speciosa EH
U. pugilator BW/EH
U. panacea BW/EH
Minuca U. longisignalis FW/BW
U. minax FW
U. pugnax BW/EH
753 to 850 mOsm (Y=0.3525X+744 mOsm, N=10,
r2=0.4363, P<0.05). In media between 300 and
2600 mOsm, hemolymph osmolality increased from 780
to 1156 mOsm (Y=0.1619X+735, N=33, r2=0.7769,
P<0.001), with an isosmotic medium concentration of
876 mOsm. In media >2600 mOsm the crabs did not
control hemolymph osmolality well. The highest toler-
ated hemolymph osmotic concentration was
1738 mOsm. Above 2600 mOsm, hemolymph samples
from U. pugilator at Waquoit Bay diverged from those
of other populations. As medium increased to
3380 mOsm, hemolymph osmolality in these crabs
increased to 1340±53 mOsm, while populations
from Crooked River (1735±64 mOsm), Alligator
Point (1664 mOsm) and St. Joseph Peninsula
(1675±13 mOsm) had significantly higher osmolality
(P<0.05) in media with lower osmotic pressure (2700–
3000 mOsm). Thus, the Waquoit Bay sample is more
resistant to higher osmotic pressure than other U. pug-
ilator populations.
To examine osmoregulation under laboratory con-
ditions, crabs kept in habitat water for 30 days were
submitted to an abbreviated regime of osmotic stress. In
these studies U. pugilator, U. panacea and U. pugnax
were exposed to 250, 400, 900 and 1650 mOsm ASW,
while U. minax were exposed to 250, 400, 900 and
1450 mOsm ASW, and U. longisignalis to 250, 780 and
1450 mOsm ASW. In all cases, there was 100% survival
after 5 days of exposure to the ASW. Hemolymph os-
molality for crabs from each population is shown in
Table 3. Neither the individual populations nor the
grand mean for a species were strikingly different from
the 30-day values (Table 1) or the data shown in Figs. 5,
6 and 7. Thus, under this limited regime, it appears that
living in captivity for a month does not alter significantly
the osmoregulatory capability of the five species of
fiddler crabs.
Variation of the internal state as a function of envi-
ronment conditions is considered capacity adaptation
(Prosser 1991). Since evidence for this is strongest in FW
and BW populations of U. minax, crabs from Point
Lookout, MD (385 mOsm), were examined for regu-
lated capacity adaptation. In this case, fiddler crabs were
adapted to a concentration of ASW that they would not
likely encounter in their habitat (1800 mOsm). First,
specimens were kept in habitat water for 10 days,
then transferred to 940 mOsm ASW for 14 days.
Afterwards, they were transferred to 1800 mOsm for
85
LC50(conc.) Isosmotic conc. Max. medium conc.
1500 737 1511
2550 953 2623
2900 876 3379
3350 880 3421
1850 653 1746
1950 673 2098
2700 879 3414
2 weeks. After the 14 days, crabs in the 1800 mOsm
medium had a hemolymph osmolality of 910±28 mOsm.
This is significantly >608±11 mOsm for the specimens
(N=5) held in habitat water for the same length of time
(P<0.001). Hypertonic-adapted U. minax were exposed
to a medium regime ranging from 0 to 2544 mOsm for
5 days. Survivorship and the hemolymph–medium os-
molality curve are shown in Fig. 8. After hypertonic
adaptation, the LC50 was shifted from 1950 to
2600 mOsm. In media between 0 and 1600 mOsm, the
hemolymph osmolality was similar to that reported in
Fig. 5. In ASW between 0 and 100 mOsm, the hemol-
ymph osmolality was 518±11 mOsm. Hemolymph (Y)
and media (X) osmolality are related by
Y=0.3432X+560 mOsm (N=3, r2=0.9147, P<0.10).
Between 200 and 1600 mOsm ASW, hemolymph os-
molality was 716±75, and is related to the media by
Y=0.1658X+569 mOsm (N=6, r2=0.9486, P<0.01).
The isosmotic medium concentration for adapted crabs
was 681 mOsm, which was not appreciably different
from other U. minax populations (673 mOsm; Table 2).
In extremely hypertonic ASW, hemolymph osmolality
may rise as high as 1475 mOsm in the adapted crabs.
The most dramatic effect of adaptation is shown in
Fig. 8. Hypertonic-adapted crabs are better able to
regulate and survive media above 2000 mOsm. The ex-
tension of the hemolymph–medium curve to the right is
evidence for regulated capacity adaptation in osmotic
balance by U. minax.
Discussion
Among Crustacea, patterns of ionic and osmotic reg-
ulation are well established for a number of species.
Excellent reviews on the subject have been written by
Mantel and Farmer (1983) and Greenaway (1988).
More recent work by Wolcott (1991) and Ahearn et al.
(1999) focus on cellular aspects of ion transport and
water balance throughout the class. All together these
reveal a vast knowledge of salt and water balance for
crustaceans. However, from a perspective of compar-
ative ecophysiology, most studies document similarities
and differences in species which are phylogenetically
and ecologically unrelated. Few studies deal with
closely related species living in sympatry. In general,
most studies with fiddler crabs are limited to a single
species (Table 4). In cases where more than one
86

Fig. 5 Uca spp. Relationship

between medium and hemol-
ymph solute concentration in
species from the Minuca subge-
nus: U. longisiganlis (N=31), U.
minax (N=278), U. pugnax
(N=219). Solid diagonal line is
isosmotic. Solute concentration
in osmolality (mOsm). Dura-
tion of exposure: 5 days

species has been used, the data are presented in such a 16 different locations. With this approach, it is clear
way that few, if any, distinctions between congeners that significant physiological differences do occur
can be made. The present study examines osmoregu- among populations, as well as among closely related
lation in eight species from the same genus taken at species.

Fig. 6 Uca spp. Relationship
between medium and hemol-
ymph solute concentration in
species from the Celuca subge-
nus: U. spinicarpa (N=34),
U. speciosa (N=63). Solid
diagonal line is isosmotic. Solute
concentration in osmolality
(mOsm). Duration of exposure:
5 days
Previous studies
Uca spp. have long been important subjects for studies
of ionic and osmotic regulation in crustaceans. Seminal
works by Jones (1941) and Gross (1964) demonstrated
the excellent osmoregulatory capabilities of fiddler crabs
when compared to other aquatic and terrestrial genera.
After studying the local distribution of fiddler crabs in a
Georgia salt marsh, Teal (1958) concluded that different
configurations of substrate, food, vegetation and salinity
were the most influential determinants for the presence
of U. minax, U. pugnax and U. pugilator. Thus, osmotic
stress is an integral factor regulating the ecological
distribution of the three Atlantic fiddler crabs.
Despite these findings, Green et al. (1959) considered
the three species to have equal physiological capabilities,
and consequently pooled their blood samples. In their
experiments, crabs adapted to 100% (1050 mOsm) and
87
175% (1850 mOsm) seawater have hemolymph osmol-
ality of 497±12 mOsm and 758±36 mOsm, respec-
tively. Later, Baldwin and Kirschner (1976a,b), studying
a mixture of U. pugnax and U. pugilator, found hemo-
lymph osmolality to vary from 410 to 650 mOsm as the
medium was changed from 10% (105 mOsm) to 100%
(1050 mOsm) seawater. More recently, Holliday (1985)
examined the hemolymph osmolality of U. pugnax
adapted to 10% (105 mOsm), 50% (520 mOsm), 100%
(1050 mOsm), 150% (1575 mOsm) and 200%
(2100 mOsm) seawater. At the same time, average blood
osmolality varies in a linear fashion from 650 to
1000 mOsm. Similar studies with U. pugilator and U.
minax by Wright et al. (1984) found the two to be good
hyper-osmotic regulators from 5% (50 mOsm) to 75%
(790 mOsm) seawater as hemolymph osmolality
increases from 550 to 700 mOsm. At higher
medium concentrations, they lose the ability to regulate
88
Fig. 7 Uca spp. Relationship
between medium and hemol-
ymph solute concentration in
species from the Celuca subge-
nus: U. pugilator (N=321),
U. panacea (N=80). Solid
diagonal line is isosmotic. Solute
concentration in osmolality
(mOsm). Duration of exposure:
5 days
hemolymph below medium osmolality. However, they
note that maintenance of hemolymph below medium
osmolality was more pronounced in U. pugilator than in
U. minax. D’Orazio and Holliday (1985) also found U.
pugilator from Florida to be an excellent regulation be-
tween 50% and 200% seawater. Consequently, from the
literature one is left with the impression of physiological
uniformity among North American species of Uca.
Other studies have examined species from Africa and
the Caribbean. After studying both U. pugilator and U.
tangeri, Graszynski and Bigalke (1986) found African
species to be the better osmoregulators between 33%
(350 mOsm) and 150% (1560 mOsm) seawater. Their
blood osmolality does not rise above 651±84 mOsm as
medium osmolality increases to 3120 mOsm. Spaarga-
ren (1977) reported that U. inversa from the Red Sea can
tolerate osmolality as high as 1235 mOsm. Hemolymph
osmolality in this species varies from 790 to 1016 mOsm.
In a Caribbean study, Schmidt-Nielsen et al. (1968)
exposed U. burgersi from Jamaica (Barnwell 1986) to
50% (525 mOsm), 100% (1050 mOsm) and 200%
(2100 mOsm) seawater, and found hemolymph osmol-
ality to vary from 784±56 to 1000±90 mOsm. Conse-
quently, this hypersaline species is a fairly good
osmoregulator, but hemolymph concentrations are
consistently higher than those of its relatives on the
mainland. Finally, Spaargaren (1975) studied the Ca-
ribbean Celuca, U. speciosa, on Curacao. As the external
medium was changed from 270 mOsm (25%) to
1240 mOsm (120%), the osmolality of the hemolymph
increased from 560 to 900 mOsm. In media with higher
osmolality, the crab lost its ability to control hemol-
ymph osmolality.
Although these early studies are informative, most
are limited to examining hemolymph changes between
5% and 150% or 200% seawater. In addition, most do
not present statistics on survivorship. Consequently, an
incomplete view of osmoregulation in Uca spp. has
evolved due to the restricted experimental designs. The
efficiency and elegance of a physiological mechanism
may not be fully appreciated until the organism is forced
to function near its physiological limits. Thus, it is not
surprising that the osmoregulatory abilities of different
species were viewed as uniform. In their examination of
U. subcylindrica and U. longisignalis, Rabalais and
Cameron (1985) exposed crabs to a 3400 mOsm range of
 89

Table 3 Uca spp. Osmoregulation after 30 days in habitat water (N=16 crabs from each collection site)
Subgenus, Location (mOsm) Medium osmolality (mOsm):
species
250 400 780 900 1450 1650

Celuca
U. pugilator Alligator 836±24 815±18 – 898±23 – 1008±60
Point (722)
Pt. St. 845±14 795±4 – 881±21 – 1052±29
Joseph (987)
Crooked 769+26 815+33 – 865+19 – 1128+38
River (935)
Mean 816±34 808±9 881±13 1063±50
U. panacea Alligator 815±10 810±5 – 880±44 – 989±27
Point (722)
(Minuca)
U. minax Money 620±30 632±18 – 729±47 999±28 –
Bayou (32)
St. Mary’s 640±5 627±6 – 672±24 863±33 –
R (380)
Lookout 630+31 616+18 – 675+21 967+37 –
Point (385)
Mean 630±8 625±7 – 692±26 943±58
U. longisignalis Money 692±24 – 694±17 – 779±22 –
Bayou (32)
U. pugnax Isle of 720±31 713±26 – 826±3 – 1259±69
Wight (860)
Crooked 668+4 825+15 – 830+17 – 1034+58
River (885)
Mean 694±26 769±56 – 828±2 – 1146±112

stress. This revealed physiological differences in the two
species. Although both are excellent regulators, U. lon-
gisignalis were better able to survive in hypertonic media
than U. subcylindrica. Using a similar osmolality regime
(Zanders and Rojas 1996), U. rapax from Venezuela was
demonstrated to be an exceptional regulator. In 5–300%
ASW (3120 mOsm), the crabs’ hemolymph increases
from 670 to 950 mOsm, and to as high as 1500 mOsm in
400% ASW (4200 mOsm). Only as a result of applying
extreme osmotic pressure under laboratory conditions
are differences in the osmoregulatory abilities among
species in this genus becoming obvious.
Present study
Uca spp. were collected from a variety of temperate
habitats between Cape Cod, MA, and Pensacola, FL.
The average osmolality of hemolymph was determined
for each population in the study (Table 1). Where pos-
sible, samples from these populations were kept in
habitat water for 30 days, and the hemolymph osmol-
ality measured a second time (Table 3). In general, there
was no significant change in either hemolymph osmotic
concentration or the ability of the crabs to osmoregulate
after this period in captivity. Consequently, the variation
noted among the populations for each species appears to
be a reasonable assessment of their state in nature.
Unsurprisingly, habitat has greater impact on os-
moregulation among Uca spp. than taxonomic relation-
ships. The two subgenera, Celuca and Minuca, each have
four species occupying different portions of the salinity
array (oligo- to euryhaline). Among the two subgenera,
U. minax and U. spinicarpa are typically freshwater, and
both have average hemolymph around 600 mOsm in na-
ture. In brackish water U. longisignalis and U. pugnax
have hemolymph osmolality between 690 and 750 mOsm,
respectively. When U. panacea are taken from brackish
water habitats, their hemolymph osmolality is near
710 mOsm. All the euryhaline specimens have hemol-
ymph osmolality >790 mOsm. This ecological cohort
includes U. speciosa, U. panacea, U. pugilator, U pugnax
and U. rapax. Based on field measurements taken during
the present study, Uca spp. in oligohaline environments
are expected to possess a hemolymph osmolality around
600 mOsm. Those living in brackish habitats should have
hemolymph with osmolality somewhat >700 mOsm
while those in euryhaline habitats should possess a
hemolymph of 790 mOsm or greater.
Species are not uniform in their osmoregulatory ca-
pabilities. If hemolymph–medium osmolality curves
(Figs. 5, 6, 7) are compared, considerable interspecific
variation is obvious. Virtually every species is able to
maintain constant hemolymph osmolality between 0 and
1400 mOsm (0–125% seawater). To be sure, some spe-
cies such as U. pugnax and U. pugilator do not seem to
tolerate low osmotic concentrations as well as U. minax,
U. longisignalis and U. panacea. As medium concentra-
tion is increased above 125% seawater, U. longisignalis,
U. minax and U. spinicarpa succumb to the osmotic
pressure. U. pugnax, U. speciosa, U. pugilator and U.
panacea can withstand medium of 2200 mOsm (200%
seawater) or greater. Overall, the sibling species U.
pugilator and U. panacea seem to regulate hemolymph
90

Fig. 8 Uca minax. Survivorship

and osmoregulation in U. min-
ax acclimated to 1800 mOsm
ASW. Upper graph: survivor-
ship (N=42); lower graph: rela-
tionship between medium and
hemolymph osmolality (N=39,
curve with open data points)
[solid diagonal line isosmotic;
dotted curve data from Och-
lockonee River population
(14 mOsm; ppt<1); continuous
curve data from Thompson
Bayou, Point Lookout and St.
Mary’s River (32–385 mOsm)]

osmolality over a wider medium osmotic range than
other species.
In crustaceans, the fundamental mechanisms for fluid
and ion balance are consumption, elimination and
transport across epithelial barriers (Prosser 1991). His-
torically, Uca spp. are known for their ability to regulate
their internal milieu using these mechanisms (Jones
1941; Gross 1964). The concentration of water and ions
in hemolymph remains somewhat constant as the ex-
ternal medium changes over a wide range. As the in-
tensity of the challenge increases beyond the regulated
range, internal osmotic concentration begins to waiver,
and the relationship becomes one of conformity and
tolerance. At the extremes of the range, the organism
cannot tolerate the pressure, and eventually dies. If it is
important for a species to maintain physiological
flexibility for survival in various habitats, capacity
adaptation allows the organism to modify its ability to
hyper- and hyporegulate economically in different
media. As the organism is gradually adapted to a new
osmotic environment, the curve of regulation is modified
depending on the external condition. The central portion
of the curve, representing a physiological optimum,
remains constant, while the terminal portions are shifted
either to the right or the left. As crabs adapt to media of
increasing osmolality, there is a concomitant decrease in
Na/K-ATPase activity in the posterior gills (Wanson
et al. 1984; Holliday 1985). On the other hand, when
assaulted with a hypotonic medium, specific Na/K-AT-
Pase will increase and water permeability will decrease.
These changes begin within 24–72 h. Consequently, ad-
aptation producing a new osmoregulatory capacity takes
only a few days (Rabalais and Cameron 1985; Zanders
and Rojas 1996).
However, from an intraspecific perspective, descrip-
tions of differences in volume regulation, ionoregulation
and osmoregulation among populations of grapsid spe-
cies are not common. Ferraris and Norenburg (1997)
documented differences among U. rapax rapax popula-
tions in Florida, Belize and Panama. Stanton and Felder
(1992) have also reported variation in osmoregulation
among populations of Sesarma reticulatum from Georgia
 91

Table 4 Uca spp. Habitat preference and osmoregulatory capabilities (FW freshwater; BW brackish water; EH euryhaline)
Subgenus, Reference Habitat Isosmotic Minimum hemolymph Maximum
species conc. osmolality (mOsm) tolerated medium
osmolality (mOsm)

Amphiuca
U. inversa Spaargaren (1977) BW/HS 1016 790 1430
Celuca
U. crenulata Jones (1941) BW 860 750 1775
U. pugilator Baldwin and HS 800 700 1040
Kirschner (1976a)
Wright et al. (1984) HS 750 600 1500
D’Orazio and HS 990 850 1900
Holliday (1985)
U. speciosa Spaargaren (1975) HS 960 700 1660
U. subcylindrica Rabalais and BW/HS 850 525 2600
Cameron (1985)
U. spinicarpa Present study FW 737 583 1511
U. pugilator Present study BW/HS 876 632 3379
U. panacea Present study BW/HS 880 835 3421
U. speciosa Present study HS 953 691 2623
Minuca
U. minax Wright et al. (1984) FW 790 550 1600
U. pugnax Holliday (1985) BW 1000 700 1800
U. longisignalis Rabalais and FW/BW 700 500 3400
Cameron (1985)
U. burgersi Schmidt-Nielsen HS 900 784 2000
et al. (1968)
U. rapax Zander and HS 750 670 4400
Rojas (1996)
U. minax Present study FW 673 542 2098
U. longisignalis Present study FW/BW 653 570 1746
U. pugnax Present study BW/HS 879 581 3414

and Louisiana. In the present study population
differences were noted in two species: U. minax and
U. pugilator. Variation in osmoregulation between pop-
ulations was most obvious in U. minax. The hemolymph–
medium curves for populations from the Atlantic and
Gulf of Mexico coasts are similar, except the specimens
from the Ochlochonee River. A key observation is that the
isosmotic concentration for crabs adapted to different
osmotic conditions is virtually the same. Since there is
little genetic divergence among U. minax populations in
this region (Felder and Stanton 1994; Mangum 1996), the
physiological change is most likely due to adaptive vari-
ation in gene expression. The expression of enzymes as-
sociated with ion transport varies during adaptation of U.
minax to extreme osmotic concentrations (Wanson et al.
1984). Data from natural and laboratory populations for
U. minax (Fig. 8) are best explained by the notion of
regulated capacity adaptation. This also explains the di-
vergence of U. pugilator at Waquoit, Mass., from other
populations (Fig. 7). It is curious that similar physiolog-
ical behavior was not observed in the populations of U.
pugnax from the Atlantic coast; it may simply be that too
few specimens of this species were collected in oligohaline
or brackish habitats to permit the observation.
In summary, osmoregulation has been observed in
seven species of Uca from the temperate Atlantic and
Gulf of Mexico coasts of eastern North America. All are
excellent hyper- and hyporegulators across osmolality
ranges of 30–1400 mOsm. However, beyond this range,
both inter- and intraspecific physiological variations are
common. Based on environmental water and hemo-
lymph measurements, Uca are readily classified as
oligohaline, brackish or euryhaline species. Since habitat
salinity varies along the coast, populations of each spe-
cies can differ in average hemolymph osmolality. Under
laboratory conditions, the euryhaline species were found
to osmoregulate over a greater medium range than
species from freshwater habitats. The capability for
regulating under osmotic challenge can be adjusted by
acclimation. This has been demonstrated in the fresh-
water species U. minax. Although broad capabilities of
osmoregulation exist in many species, there is consider-
able physiological flexibility permitting adaptation.
Acknowledgements The author wishes to thank the Graduate
College at the University of Northern Iowa for financial support of
this study. Help in collecting and transporting Uca specimens from
Georgia and Florida was generously provided by Chad Thurman.
J. Makepeace graciously provided transportation and help with
collections in Maryland. K. Flaherty provided crabs from
Thompson’s Bayou. Accommodations in Massachusetts were
provided by C. Weidmann, Waquoit Bay National Estuarine
Research Reserve.
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