Hydrocarbon metabolism in the intertidal fiddler crab Uca pugnax

The fiddler crab Uca pugnax was examined for its ability to metabolize foreign hydrocarbons. The microsomal mixed function oxidase system was identified in U. pugnax tissues using Aldrin epoxidation rates as the assay. Rates were slow: 96 pM Dieldrin per mg microsomal protein per hour in the hepatopancreas, 438 pM mg^-1 h^-1 in the gill, and 228 pM mg^-1 h^-1 in claw-muscle microsomes. Using standard methods, no difference in rates could be detected between crabs living in clean areas and those living in environments highly contaminated with foreign hydrocarbons. In vivo rates of naphthalene oxidation were measured and used to calculate a clearance time for U. pugnax body tissues based on the aromatic hydrocarbon content of crabs collected from an oil-polluted salt marsh. Calculated clearance time was beyond the life span of the crab. It is concluded that this minimal ability of U. pugnax to metabolize foreign hydrocarbons partially accounts for its sensitivity to oil pollution in the environment.     

Adaptive droving behavior observed in the fiddler crab Uca vocans vocans

Droving is conspicuous in Uca vocans vocans in summer. The crabs burrowing on the upper habitat have a higher tendency to wander compared to the ones burrowing on the lower habitat. Most of the wandering crabs captured on the low tide levels are relatively large and male. Larger crabs and males prefer to burrow on the upper zones of U. vocans habitat, but the smaller ones and females prefer the lower habitat. The upper level of the U. vocans habitat has relatively low N-content compared to the water's edge. Therefore droving is advantageous in crabs that have burrows on the upper level. Female and smaller resident crabs have faster feeding motions than male and larger ones, and can satisfy their feeding demands more rapidly. Therefore, for males and larger crabs it is advantageous to move away from the burrow area and forage in areas of higher food content near the low tide level where the number of feeding motions increases.     

Control of distal retinal pigment migration in the fiddler crab Uca pugilator

Several concentrations of extracts prepared from the eyestalks of a specimen of Uca pugilator were injected into other U. pugilator individuals. The distal pigment of the eyes first became light adapted and then dark adapted, the whole process lasting 6 h. The mean integrated response for light adaptation increased progressively up to the highest tested extract (3 eyestalk equivalents/dose), but with the darkadapting response the maximal effect was produced by the extract containing 2 eyestalk equivalents/dose. Gel filtration of eyestalk extracts in Sephadex G-50 showed that the fractions associated with greatest light adaptation were also associated with greatest pigment dispersion in the melanophores. Almost no light or dark adaptation of the retinal pigment resulted from injections of eyestalk extracts treated with -chymotrypsin which supports the interpretation that these substances are polypeptides of neurosecretory origin.Supported by Grant GB-7595 X from the National Science Foundation.     

Chitobiase activity in the epidermis and hepatopancreas of the fiddler crab Uca pugilator during the molting cycle

The activity of chitobiase, also known as N-acetyl-β-glucosaminidase, in the epidermis and hepatopancreas of the fiddler crab Uca pugilator (Bosc, 1802), during the molting cycle, was investigated. A pH optimum of 5 to 6 was found for the enzymatic activity in both the epidermis and hepatopancreas. The temperature optimum for epidermal and hepatopancreatic chitobiase activities was 50 to 60 °C. The K _m values for epidermal and hepatopancreatic chitobiase activities at 19 °C were 0.190 ± 0.027 and 0.203 ± 0.016 mM 4-methylumbelliferyl-N-acetyl-β-glucosaminide, respectively. Hepatopancreatic chitobiase activity was significantly higher than epidermal enzymatic activity in all the molt cycle stages tested except Postmolt Stage A-B. Chitobiase activity varied significantly during the molting cycle, with the epidermal enzymatic activity in Premolt Stage D_3–4 significantly higher than in Stage C (intermolt) and Premolt Stage D₀, whereas hepatopancreatic chitobiase activity in Premolt Stage D_3–4 was significantly higher than in all other molt stages tested. The patterns of chitobiase activity in the epidermis and hepatopancreas correlate well with the hemolymph titer of ecdysteroids in U. pugilator during the molting cycle; this suggests that chitobiase activity in both tissues is regulated at least in part by the steroid molting hormones. Received: 6 May 1998 / Accepted: 12 September 1998     

A bacterial infection in the common shore crab Carcinus maenas and the fiddler crab Uca pugilator

A bacterial infection in Carcinus maenas and in Uca pugilator is described. The percentage of infections recorded in the field was between 0 and 10%, depending on season. Infected crabs remain about 3 times longer in Intermolt Stages C3 and C4 than do healthy crabs. In infected specimens of both species, the glucose level in the hemolymph is increased and protein, copper and lipid concentrations in the hemolymph as well as the glycogen content of various tissues are diminished. The coagulation process is disturbed in infected crabs. The hemocytes aggregate, but the formation of fibrin-like fibers, which can be observed in healthy crabs, does not occur.This paper forms part of a doctoral thesis presented at the University of Ulm (FRG).     

Rhythmic activity responses of the fiddler crab Uca crenulata to artificial tides and artificial light

The fiddler crab Uca crenulata, from California (USA), was exposed to artificial tides in order to differentiate between the influence of a tidal cycle from that of a light-dark cycle on its locomotor activity. Most crabs could be well synchronized by tides, but the activity patterns of other crabs was merely exogenously reinforced. Under constant conditions, after tidal treatment, crabs showed bimodal or unimodal activity patterns. When a 24 h light-dark cycle and a 12.35 h tidal cycle acted as concurrent stimuli, their effectiveness in synchronizing the crabs' activity was variable, depending on the undividual. It is likely that crabs which tend to exhibit a unimodal activity pattern are more sensitive to a light stimulus, whereas crabs with a bimodal activity pattern respond preferably to the tides. Thus, responses of endogenous activity of U. crenulata to tides and light-dark cycles are similar, suggesting that the basic oscillarory mechanism for circadian and tidal activity may be the same.     

Rhythmic motor activity responses of the California fiddler crab Uca crenulata to artificial light conditions

Uca crenulata, the California species of fiddler crab, was exposed to artificial light conditions to separate the influence of the light cycle from that of the tidal input on its rhythmic motor activity. Under both constant light and light-dark cycles, rhythmic activity was demonstrated in only 50% of the experimental crabs; the activity of the remaining 50% was random. Individuals exposed to constant light conditions after having been subjected to 24 h light-dark cycles demonstrated no significant difference in period length of their rhythmic activity from crabs investigated in constant light immediately after field collection. The mean period did not differ significantly from the tidal period of 24.8 h, but the variation was considerable. In artificial light-dark cycles, the activity rhythms were usually masked but, in some cases, synchronized. The results indicate that U. crenulata has an endogenous rhythm with a period close to the tidal cycle which may be synchronized by light as well as by tidal cues. The display of this endogenous rhythm, however, is poor.     

Osmotic and ionic regulation in the fiddler crab Uca rapax acclimated to dilute and hypersaline seawater

Adult male Uca rapax, collected from the central coast of Venezuela in early 1994 were gradually acclimated to salinities ranging from 1.7 to 139S. The hemolymph osmolality (791±15 mOsmol in crabs from 26S) changed less than three-fold over the entire range of concentrations tested. The urine was isosmotic with the hemolymph in crabs exposed to dilutions <26S, and significantly hyperosmotic in those exposed to media >34.8S. The hemolymph levels of Na⁺, Cl^–, K⁺, Ca²⁺ and Mg²⁺ (320±13, 405±17, 7.8±0.7, 7.2±0.1 and 31±2.2 mmol l^–1, respectively, in crabs acclimated to 26S) were maintained fairly constant over the range from 8.7 to 99S, decreasing by 15% in the more dilute media or increasing sharply to about twice those values in crabs from 139S. The excretory organs contributed to the osmoionic regulation of the hemolymph in crabs exposed to <3.5 or to >34.8S, by means of the partial reabsorption or excretion, respectively, of salts from or into the urine. The results described place U. rapax among the most powerful hypo/hyper-regulating crustaceans known.     

Pillar building in the fiddler crab Uca beebei: evidence for a condition-dependent ornament

In the fiddler crab (Uca beebei) males build a small mud pillar next to their burrow which increases their attractiveness to females. Three hypotheses were tested to explain inter-male variation in pillar-building. (1) The benefits of pillar-building are density dependent. The experimental addition of vertical structures did not support this hypothesis as there was no change in the level of pillar-building. (2) There are two classes of males (pillar-builders and non-pillar-builders). This could either be due to an alternative mating strategy, or because pillar building is age or size-dependent. There was also no support for this hypothesis. (3) Pillar-building is an honest signal of male quality dependent on body condition. A food supplementation experiment was performed. Addition of food affected several aspects of male behaviour and resulted in a two fold increase in the number of pillars built between control and food treatments (P < 0.001). However, the percentage of males building pillars did not increase significantly. Pillar building in this species has been attributed to sensory exploitation. Our results indicate that a trait which may well have evolved through sensory exploitation also appears to be condition-dependent. We emphasise that showing that an ornament or behaviour is condition-dependent does not necessarily mean that it evolved through good gene processes. However, in terms of its current selective value, pillar building may be maintained through female choice because it acts as a signal of male condition.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

Paternity analysis of two male mating tactics in the fiddler crab, Uca mjoebergi

The fiddler crab Uca mjoebergi mates both underground in male-defended burrows and on the surface near female-defended burrows. The reproductive tract of Uca species facilitates last-male precedence, suggesting that males that do not guard-mated females are likely to gain very little paternity if the female re-mates with another male. Here, we test the reproductive success of burrow and surface matings using paternity analysis. We found that 100?% of the females that mated in burrows extruded a clutch of eggs. Furthermore, we show conclusively, for the first time in a fiddler crab species, that last-male sperm precedence results in the majority of the female’s eggs being fertilised by the burrow-mated male. In contrast, surface matings resulted in significantly fewer females extruding eggs (5.6?%). Paternity analysis also revealed that more than half of the clutches from burrow-mated females showed low levels of extra-pair paternity from previous matings. Although multiple matings appear common in U. mjoebergi, burrow-mated males that guard females are guaranteed a successful mating with extremely high rates of assured paternity. Surface matings therefore appear to be an opportunistic tactic that may increase male reproductive success in a highly competitive environment.     

A conspicuous courtship signal in the fiddler crab Uca perplexa: female choice based on display structure

The fiddler crab Uca perplexa has a conspicuous male courtship signal that is directed at females to attract them to the male’s burrow for mating. The signal involves waving the unflexed large claw up and down. To determine whether the spatiotemporal structure of the wave is under selection by female choice, we examined whether females had a preference for any particular features of the wave. Females respond to a waving display by either visiting the male’s burrow entrance or by electing to pass without visiting the burrow. We filmed mate-searching females and the waving males that they visited or passed. We documented the wave structure of these males using frame-by-frame analysis. Males produce a two-part wave with component A preceding component B. Both components have an upstroke, a pause at the apex and a downstroke. The tip of the claw was raised much higher in B than in A. Visited males had a shorter delay between the two wave components than did males that the females passed without visiting. Visited males also produced component B waves that had a slower upstroke than those of passed males. There was a significant correlation between the relative height of the raised claw and the duration of the upstroke of component B. Females were selecting males that raised their major claw to the highest position (two to three times as high as the carapace width). Passed males brought down their major claw earlier and from a lower position than did visited males. The data suggests that wave structure has evolved through female choice. Male display rate and body size were not female choice cues. An earlier study showed that display duration was also not used by females in selecting mates.     

Influence of the polychlorinated biphenyl preparation Aroclor 1242® on color changes of the fiddler crab Uca pugilator

The melanin in the melanophores of specimens of the fiddler crab, Uca pugilator, exposed to 2, 4, and 8 ppm solutions of the polychlorinated biphenyl (PCB) preparation, Aroclor 1242^®, became less dispersed than in untreated specimens. This effect was dose-dependent. It was probably due to the PCB itself and not to polychlorinated dibenzofurans which are contaminants of commercial PCBs. The decreased melanin dispersion appeared to be related to a decrease in the rate of release of melanin-dispersing hormone from eyestalk neuroendocrine cells into the hemolymph. Eyestalks of crabs kept for 4 days in Aroclor 1242 contained 4 times as much melanin-dispersing hormone as did control crabs. When injected into isolated legs, did hemolymph from control crabs. Aroclor 1242 did not appear to have a direct effect on the pigment-dispersing mechanism of the melanophores.     

Time constraints and multiple choice criteria in the sampling behaviour and mate choice of the fiddler crab,Uca annulipes

Active female sampling occurs in the fiddler crab Uca annulipes. Females sample the burrows of several males before remaining to mate in the burrow of the chosen partner. Females time larval release to coincide with the following nocturnal spring tide and must therefore leave sufficient time for embryonic development after mating. Here we show how this temporal constraint on search time affects female choosiness. We found that, at the start of the sampling period (when time constraints are minimal), females selectively sample the larger males in the population. Towards the end of the sampling period (when the temporal constraints increase the costs of sampling), females are less selective. Furthermore, we suggest that the number of males sampled (and other indices of ‘‘sampling effort’’) may not be reliable indicators of female choosiness and may not reflect the strength of female mating preferences under certain conditions. Burrow quality also emerged as an important criterion in final mate choice. Burrow structure potentially influences reproductive success, and mate acceptance based on burrow structure appears to involve a relatively invariant threshold criterion. Since there is no relationship between male size and burrow quality, females are using at least two independent criteria when choosing potential mates. We envisage mate choice as a two-stage process. First, females select which males to sample based on male size. They then decide whether or not to mate with a male based on burrow features. This sampling process explains how two unrelated variables can both predict male mating success. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Interactions of methylmercury,cadmium, and salinity on regeneration in the fiddler crabs Uca pugilator,U. pugnax and U. minax

After multiple autotomy in the fiddler crabs Uca pugilator, U. pugnax and U. minax, both methylmercury and cadmium retard limb regeneration and ecdysis. When crabs in sea water are exposed to a mixture of both metals, the effect is increased, indicating that the two are interacting in an additive way. In 50% sea water (=15% S), the effects of cadmium are greatly intensified so that growth of limb buds is extremely slow, if it occurs at all. When methylmercury is present in the water at the same time, the severe effects of cadmium are somewhat ameliorated, indicating an antogonistic interaction of the two metals under these conditions. Adding additional calcium to the 50% sea water also decreased the severity of the cadmium effect, thus supporting the idea of a calcium-cadmium competition.     

Changes in lipid class and fatty acid contents in the ovary and midgut gland of the female fiddler crab Uca tangeri (Decapoda,Ocypodiadae) during maturation

Changes in biochemical composition, lipid class and fatty acid contents were studied in the ovaries and midgut glands of the fiddler crabs Uca tangeri Eydoux during maturation. Wild females were caught during spring and early summer of 1992 in the Bay of Cádiz (southwest Spain), near the mouth of the San Pedro river. Protein and total lipid contents in the ovaries increased significantly from Stages III to IV, at the expense of total carbohydrate, which showed a large decrease during the same period. In the midgut gland, the protein content did not present any significant variation, whereas total lipids and total carbohydrates presented opposite up and down trends during maturation. In the ovary, total polar lipids increased significantly during the final phase of maturation (Stages III to IV), mainly due to the significant contribution of the phosphatidylcholine and phosphatidylethanolamine fractions. In contrast, total neutral lipids showed an upward trend throughout the whole maturation period, mainly due to significant increases of the triacylglycerol fraction. In the midgut gland, total polar lipids (mainly phosphatidylcholine) and total neutral lipids (mainly triacylglycerol) presented significant decreases from Stages II to III, the phase which preceded major increases in both polar and neutral lipids in the ovaries. Cholesterol content did not vary during maturation in either organ, in the ovary or midgut gland. Major fatty acids in the ovaries [16:0, 16:1 (n-7), 18:1 (n-9), 18:1 (n-7), 18:2 (n-6), 18:3 (n-3), 20:4 (n-6), 20:5 (n-3) and 22:6 (n-3)] did, however, accumulate significantly at later stages of maturation. It is noteworthy that arachidonic acid [20:4 (n-6)] content remained constant during all stages of maturation but decreased significantly in total polar lipids in the later phases of maturation. In contrast, eicosapentaenoic acid [20:5 (n-3)] increased significantly in all lipid fractions in the later stages, and docosahexaenoic acid [22:6 (n-3)] remained constant in the polar lipids and increased during later stages in the triacylglycerol fraction. Major fatty acids in the midgut gland lipids showed significant decreases from Stages II to III, just before the final period of maturation.     

Lack of population structure in the fiddler crab Uca annulipes along an East African latitudinal gradient: genetic and morphometric evidence

Phylogeographic patterns in Uca annulipes sampled from 30 locations across an East African latitudinal gradient were investigated using mitochondrial DNA cytochrome oxidase I sequences and analysed together with patterns of morphometric differentiation. Four hundred and four specimens along the east African coast were sampled, and 18 haplotypes were encountered. Haplotype and nucleotide diversity values were very low and the phylogenetic analysis did not reveal any clear phylogeographic structure. Furthermore, the analysis of molecular variance and pairwise Φ _ST values showed no significant spatial population differentiation. Mismatch analyses and tests of neutrality supported the hypothesis that this species has undergone a fairly recent demographic expansion. Our results, therefore, failed to demonstrate significant geographical structure in the pattern of genetic variation, indicating that populations of U. annulipes are capable of extensive gene flow among mangroves along the coast. The genetic structure of this species could be panmictic due to a high amount of gene flow along the geographical gradient in study during the planktonic larval phase, when larvae are carried along stream by the dominant currents. Moreover, the morphometric analysis performed did not reveal differences of shape differentiation according to a geographical pattern, although significant differences among the sampling areas were found.     

Interactive effects of predation pressure and insecticide (temefos) toxicity on populations of the marsh fiddler crab Uca pugnax

The interaction of predation pressure with lethal and sublethal effects of temefos (Abate^®) an organophosphorous insecticide, were studied in field populations of Uca pugnax (S.I. Smith). Changes in fiddler crab population densities were followed in open-marsh temefos-treated and untreated test plots and in treated and untreated plots which were caged over to reduce predation by marsh birds. Temefos significantly reduced the population density of U. pugnax in the open test plots but not in the caged plots. These results indicate that temefos has a primarily sublethal effect on the crabs, the effect becoming lethal only after interaction with avian predation. Evidence from the presence of a time-lag effect in the population decrease, from a calculated predation index, and from laboratory studies reported elsewhere of behavioral alteration by temefos also supports the conclusion that temefos primarily impairs the escape response of U. pugnax: this leads to increased predation and subsequently to a decreased fiddler crab population. Such studies of sublethal effects of toxicants and field studies of interactions of lethal and sublethal effects of such compounds with natural population dynamics of affected species are necessary to evaluate possible effects of toxicants on populations.     

Spatial and temporal patterns in the feeding behavior of a fiddler crab

Fiddler crabs consume the surficial microphytobenthos around their burrows during low tide. We studied the spatial and temporal feeding patterns in the species Uca uruguayensis by using sequences of digital pictures of feeding pellets accumulation. Data from 61 crabs, feeding without the interference of neighbors, were fitted to different models using Maximum Likelihood and Bayesian approaches. Initial feeding location was independent from the emergence location, and then, crabs continued holding a main feeding direction (clockwise or counterclockwise), suggesting a systematic mechanism that may avoid feeding over already processed sediment. Crabs used at least half of their potential feeding area, but these areas were heterogeneous. Both sexes developed similar feeding areas; however, females were faster and needed less time to feed than males, suggesting that males are time restricted. Our work also highlights the importance of incorporating other underlying mechanisms of the behavior of species into the study of feeding strategies.