(DRAFT) - Taxonomy
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



TAXONOMY

NAME - crab, Atlantic marsh fiddler OTHER COMMON NAMES - calling crab, fiddler crab, fiddler, mud fiddler(Williams 1984), mud fiddler crab(Wheeler 1978) and marsh fiddler crab(Ward et al. 1976) ELEMENT CODE - CATEGORY - Aquatic Crustaceans PHYLUM AND SUBPHYLUM - Arthropoda, Mandibulata CLASS AND SUBCLASS - Crustacea, Malacostraca ORDER AND SUBORDER - Decapoda, Reptantia FAMILY AND SUBFAMILY - Ocypodidae, GENUS AND SUBGENUS - Uca, SPECIES AND SSP - pugnax, SCIENTIFIC NAME - Uca pugnax AUTHORITY - Smith TAXONOMY REFERENCES - 206 Taxonomy - 1
                                  (DRAFT) - Status
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



STATUS

Coded Status See Comments REFERENCES FOR STATUS - 206 COMMENTS ON STATUS - Fiddler crabs are the most abundant and conspicuous invertebrates in many slat marshes. They are probably the most thoroughly studied of the shore crabs in North America. Although there are 15 species along the North American coast, the Atlantic marsh fiddler is the only one endemic to the temperate Mid-Atlantic coast of the United States. The Atlantic marsh fiddler, red-jointed fiddler, and the sand fiddler are the three major species on this coast and make up the greatest animal invertebrate biomass in the salt marsh intertidal zone. The Atlantic marsh fiddler is the most abundant of these *206*. The ecological influence of fiddler crabs in the salt marsh is large. Their activities and byproducts can significantly influence the transfer of energy and nutrients within the marsh ecosystem. The sensitivity of these crabs to pollutants and their role in the balance of the salt marsh ecosystem, are major reasons for the inclusion of the Atlantic marsh fiddler in this series *206*. Status - 1
                               (DRAFT) - Distribution
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



DISTRIBUTION

Distribution - 1
     

HABITAT ASSOCIATIONS

NATIONAL WETLAND INVENTORY CODES NWI NWICLS NWIMOD NWISPEC Estuarine, intertidal UB3 2 Habitat Associations - 1
                                (DRAFT) - Food Habits
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



FOOD HABITS

TROPHIC LEVEL - OMNIVORE REFERENCES FOR TROPHIC LEVEL - 206 LIFESTAGE FOOD FOOD PART General Plankton Not Specified General Tracheophyta Not Specified General Benthos Not Specified General Detritus - Organic Not Specified General Microorganisms Not Applicable General Crustaceans Not Specified General Annelids Not Specified General See Comments; Food REFERENCES FOR GENERAL FOOD - 206 REFERENCES FOR LARVAE FOOD - 206 COMMENTS ON FOOD - Fiddler crabs emerge in great numbers from their burrows at low tides and feed. The Atlantic marsh fiddler feeds by scrubbing muddy particles of substratum. Its mouthparts are structured to manipulate and feed on particulate organic matter in muddy substrates. Male fiddlers are reported to spend more time feeding than females, probably compensating for the fact that males have only one chela (the smaller claw) that is functional in feeding. Some Atlantic marsh fiddlers forage far from their burrows. Some adults have been seen to feed underwater *206*. The crabs significantly reduce the abundance of meiofauna (crustaceans, nematodes, and segmented worms), probably by feeding on them *206*. In foraging, fiddlers actively sort out indigestible from digestible items and place indigestible material aside in the form of balls which break up and return to the ecosystem beginning on the next tide. Undigested material passing through the gut of the fiddler is deposited as fecal pellets. These pellets often blanket the marsh, and can persist through successive tidal cycles *206*. COMMENTS ON LARVAE FOOD - The zoeae of all three Atlantic species of fiddler crabs are carnivorous *206*. Food Habits - 1
                         (DRAFT) - Environment Associations
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



ENVIRONMENTAL ASSOCIATIONS

G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS G Water Temperature: Specified in Comments G G Relation to Substrate: Occurs in substrate [penetrating] G Bottom Type [Aquatic]: Mud or silt G Stability of Bottom: Stable G Aquatic Habitat Zonation: Shallows with emergent vegetation [littoral zone] G Coastal Features: Sand beaches REFERENCES FOR ENVIRONMENTAL ASSOC_ - 206 COMMENTS ON ENVIRONMENTAL ASSOCIATIONS - Temperature. The range of the Atlantic marsh fiddleris limited by temperature extremes. Researchers found that molting in adults is blockedby temperatures less than 20 degrees C. This temperature is the same as the summer water temperature at Cape Cod (the upper limit of the geographic range of the Atlantic marsh fiddler). It was hypothesized that the air temperature may be more influential than water temperature in semiterrestrial crabs, and that the northern geographical limit of the Atlantic marsh fiddler may be determined by the effect of water temperature on the planktonic larvae. The Atlantic marsh fiddler, in particular, can eventually acclimate to lower temperatures, but it dies at 2-3 degrees C. Hibernating crabs dug from New Jersey marshes when air temperatures were from 1.7 to 5.5 degrees C were revived. To raise their body temperatures a few degrees, some fiddler species orient themselves to the sun during low-temperature periods. Death from thermal exposure occurs at 40 degrees C in humid air and at 45 degrees C in dry air. Lethal temperatures can occur in nature. For example, in Georgia marshes were reported to reach lethal air temperatures of 40 degrees C during the summer. To avoid lethal temperatures and subsequent desiccation, fiddler crabs move to their burrows or shady parts of the marsh during periods of high temperatures *206*. Environment Associations - 1
                               (DRAFT) - Life History
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



LIFE HISTORY

PHYSICAL DESCRIPTION: Crabs belonging to the genus Uca are moderate to large in size. The three species common to the Mid-Atlantic region are Uca pugnax (Atlantic marsh fiddler), U. minax (red-jointed fiddler), and U. pugilator (sand fiddler). The carapace of the Atlantic marsh fiddler male averages 15 mm long x 23 mm wide; that of the female averages 13 mm long x 18 mm wide *206*. In life, the Atlantic marsh fiddler is mostly brown (with some pale gray in the gill regions). The anterior part of the carapace and eyestalks range from blue to blue-green. There are no purple or red spots as are found on other fiddlers. The frontal region is about two-sevenths of the body width, with slender eyestalks. The dorsal carapace is flattened rather than convex as in other crabs. At the intersection of the front and lateral edges of the dorsal carapace, there is a sharp angle *206*. Specific identification characteristics are usually descriptive of the male and often refer to the major cheliped (large claw). In the Atlantic marsh fiddler, the major cheliped ranges from a dull yellowish orange to yellow-white. In males, joints of the major cheliped have a yellow or yellow-brown border. In the northern part of the range, fingers of the major cheliped are nearly always white. These structures in females are colored less strongly than males *206*. The minor (smaller) chela is white. The other appendages--the walking legs--are usually dark and may be banded. A patch of rows or paired rows of dense velvety pubescence, as well as sparse rows of stiff hairs, are on the ventral surface of the merus (the long section of the appendage closest to the body) of the second and third walking legs. The meral surface of the second maxilliped usually has 0-75 spoon-tipped hairs *206*. The large claw of the males occurs about equally on the right or on the left of the Atlantic marsh fiddler. The weight relationship of the large cheliped ranges from 2% to 65% of the body weight. Females have two small chela that are equal in size and are colored similarly to males, but the color is less vivid *206*. REPRODUCTION: Courtship of the Atlantic marsh fiddler consists of a series of visual and acoustical displays. Visual and acoustical signals replace the standard chemical communications of most aquatic crustaceans. Usually in precopulatory behavior, male fiddler crabs display a high intensity waving of the major cheliped and produce acoustical signals; however, male Atlantic marsh fiddlers often have lethargic displays. Acoustical displays by males are produced by vibrating and stamping of the walking legs on the substrate. The waving of the large claw can be seen at considerable distances and is weakly circular and very jerky. Displaying males are not bleached in color during mating to the extent shown by males of other species of fiddlers, but the major cheliped lightens to light brown or yellow. Nocturnal reproductive behavior, including acoustical displays, of the Atlantic marsh fiddler in the southern part of its geographic range, have been reported. After these courtship displays, the female usually follows the male to his burrow to mate. Although herding (the male physically maneuvering the female to his burrow) is found in other species of fiddler crabs, it is seldom seen in the Atlantic marsh fiddler. Copulation usually takes place Life History - 1 (DRAFT) - Life History Species crab, Atlantic marsh fiddler Species Id M070000 Date 26 AUG 96 in the burrow, but has been observed on the surface of the marsh in nature; in captivity, mating has been observed underwater. Unlike mating in some other crabs, fiddlers mate while the exoskeleton of the female is in a hardened state *206*. EGGS. The fertilized eggs are carried on the abdomen of female intermolt Atlantic marsh fiddlers until they hatch and are released. Clutches of 1,500 to 94,000 eggs were found in one study. Other reports of the Atlantic marsh fiddler clutches range from 4,500 to 23,700. The size of the clutch, commonly known as the sponge, is probably related to the size of the female. Researchers have found that eggs hatched over a period of 2 hours. The larvae were released (with the aid of abdominal contractions) in phase with the nocturnal high tide. In laboratory experiments, isolated females released their larvae in synchrony with those females in the wild. Researchers found that larval release coincided with the lunar cycle in the Delaware Bay, and that the Atlantic marsh fiddler released larvae during the spring and neap tides. It is hypothesized that the synchrony with the nocturnal high tide maximum allows minimal exposure of the ovigerous females to predation and provides the zoeae with a favorable tidal current upon which to be swept from the marsh into the coastal waters. Researchers concluded that the timing of the release of zoeae probably is a response to selective pressures that cause larval mortality such as lethal high temperature, low salinity, and predation by planktivores *206*. LARVAE. After hatching, the planktonic larvae of the Atlantic marsh fiddler pass through five zoeal stages (each lasting from 7 days to a month) and one megalops stage of 4 days to a month. Most published information on larval and postlarval stages of fiddler crabs is on sand fiddlers; however, the Atlantic marsh fiddler larvae are similar in most respects to those of sand fiddler larvae though they are smaller. The zoeae of all three Atlantic species fiddler crabs are carnivorous. The zoeae of the three common Mid-Atlantic fiddlers make up a significant portion of the estuarine plankton. The distribution of zoeae appeared to be stratified in the water column, and surface waters were preferred by first and second stage zoeae. Third stage zoeae were found in intermediate depths, and fourth and fifth stages were in greater depths *206*. METAMORPHOSIS AND JUVENILES. Megalops larvae of the Atlantic marsh fiddler metamorphose into the first crab stage (lasting from 3 to 4 days) and settle to the substratum. The second crab stage lasts 4 to 5 days and the third stage lasts 7 days. While in these stages the crabs are weak, cling to objects, and are not capable of burrowing *206*. LIMITING FACTORS: Predators. Fiddler crabs, including the Atlantic marsh fiddler, are a food source for many other animals. Fiddler crab parts have been found in stomach contents of adult pigfish (Orthopristis chrysoptera), channel bass (Sciaenops ocellata), and bighead searobin (Prionotus tribulus). Fiddler crabs are an integral part of the diet of the white catfish (Ictalurus catus). Some wading shorebirds and marsh birds take fiddlers as part of their diet, including the white ibis (Eudocimus albus) and clapper rail (Rallus longirostris). Researchers have reported fiddler crabs in the diet of egrets, ibis, and herons. Cattle egrets (Bubulcus ibis) and snowy egrets (Egretta thula) also prey upon fiddlers. Some species of crustaceans, e.g., red-jointed fiddlers and the blue crab, Life History - 2 (DRAFT) - Life History Species crab, Atlantic marsh fiddler Species Id M070000 Date 26 AUG 96 (Callinectes sapidus) prey upon fiddlers. According to one report, raccoons (Procyon lotor) in Georgia salt marshes and elsewhere take fiddler crabs as a normal part of their diet *206*. Parasites and Commensals. The Atlantic marsh fiddler has not been extensively studied for its parasites and commensal organisms. Researchers have reported on trematodes and cestodes that parasitize fiddler crabs. One researcher noted apostome cysts on the gills and in the molts of the crabs from Woods Hole, MA. Another report extensively examined the ciliate commensals on the gills of North Carolina crustaceans and described a new species of apostome ciliate found as a small oval resting stage in the bases of gills of the Atlantic marsh fiddler. These commensals can reach great numbers (greater than 100) on the gills and may clog the gills to some extent, reducing the oxygen absorbing capabilities of the crab *206*. POPULATION ATTRIBUTES: Growth and Molting Characteristics. Little is known about the molting habits of the Atlantic marsh fiddler except that molting in adult crabs occurs 1-2 times per year. Researchers have hypothesized that the burrow is the likely place for routine molting because it offers protection from desiccation and predators; available water; and lack of disturbance. During an extensive two year study in North Carolina, researchers observed no molts on the marsh surface. Molting in adult Atlantic marsh fiddlers is temperature dependent and is completely inhibited at 20 degrees C and below. Researchers determined that other factors that negatively influence molting are light, extremes in salinity, and mechanical disturbances. Many physiological changes occur during the molting cycle of fiddler crabs *206*. REFERENCES FOR LIFE HISTORY- 206 Life History - 3
                           (DRAFT) - Management Practices
                        Species crab, Atlantic marsh fiddler
                                 Species Id M070000
                                   Date 26 AUG 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Adverse Applying insecticides Adverse Controlling pollution [thermal, chemical, physical] Adverse Other management practices [specified in comments] REFERENCES FOR ADVERSE MANAGEMENT PRACTICES - 206 COMMENTS ON MANAGEMENT PRACTICES - The effects of contaminants. Different response patterns to the effects of radiation (using a cobalt source) were found for the Atlantic marsh, red- jointed, and sand fiddler crabs along the Mid-Atlantic coast. In the experiments with male fiddler crabs, natural conditions (e.g., temperature, salinity) were kept constant. The Atlantic marsh fiddler had a slightly higher projected LD-50 (16,000 rad) than the other two common sympatric species. Additionally, the mortality response pattern of the Atlantic marsh fiddler differed from patterns of other species of fiddler crabs. In other species, mortality was proportional to dose; however, in the Atlantic marsh fiddler at dosages from 4,000 to 16,000 rad, mortality was independent of dose. In experiments in a New Jersey marsh, it was found that the application of Temefos (Abate), an organophosphate insecticide, at normal-use levels significantly reduced the population of the Atlantic marsh fiddler. Temefos is commonly applied as a granular larvicide for control of salt-marsh mosquitoes. In other experiments with caged populations of the Atlantic marsh fiddler, researchers determined lethal and sublethal effects of Temefos. Further, they found that sublethal doses reduced populations of crabs in open test plots, but not in closed cages, and hypothesized that the insecticide propbably impairs the fiddler's escape response, leading to increased predation. In an 11-day laboratory experiment, researchers fed Atlantic marsh fiddlers natural detritus, contaminated with DDT residues (10 ppm) from Long Island Sound, New York. After 5 days all crabs showed a loss of coordinated avoidance reaction, which was hypothesized to negatively affect predator avoidance in natural populations. At the end of the experiment, crab muscle from the large claw showed a three-fold increase in concentration of DDT over background level. Other workers have found a three-fold concentration of DDE (a degradtion product of DDT) in natural populations of fiddlers. Researchers found other contaminants which affect fiddler crabs (including the Atlantic marsh fiddler) are PCB's, and the insecticides in fertilizers, Aldrin and Dieldrin, which are found in agricultural runoff. Though they found no PCB's in marsh sediments, researchers did find measurable levels in crabs and hypothesized that fiddler crabs concentrate these chlorinated hydrocrabons from seawater or from food. Dieldrin was concentrated in crab tissues and impaired locomotion, killing crabs at the higher concentrations. Chemical contaminants were responsible for the drastically reduced populations of fiddler crabs in the marsh. Other species of fiddler crabs have been found to be sensitive to pollutants and contaminants. This sensitiviity has been shown for mercury, PCB'c, and cadmium *206*. Management Practices - 1
                                   (DRAFT) - References
                           Species crab, Atlantic marsh fiddler
                                    Species Id M070000
                                      Date 26 AUG 96



     

References

206 * Grimes, Barbara, Melvin Huish, J. Howard Kerby, David Moran. 1989. Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Mid-Atlantic) --Atlantic Marsh Fiddler. Species Profile Series 82(11.114) (ed.). U.S. Fish and Wildlife Service, U.S. Army Corps of Engineers Washington, DC:18. References - 1