The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.     

Freshwater crab origins—Laying Gondwana to rest

We review the available data on the phylogeny, palaeontology and divergence time estimation of primary freshwater crabs in relation to a hypothesized Gondwanan origin of these brachyurans, as postulated by some workers in recent decades. Known phylogenetic relationships within the Old World freshwater crabs do not correspond to the successive fragmentation of the Gondwana continent. This is strong evidence against an ancestral Gondwanan distribution of Afrotropical Potamonautidae and Asian-Australian Gecarcinucidae. The fossil record of freshwater crabs (no older than the Oligocene) and heterotreme brachyurans also postdate the initial break up of Gondwana. Molecular-clock based time estimates for the most common recent ancestor of freshwater crab families differ profoundly, depending on the method of calibration used, and whether freshwater or marine brachyuran fossils are used as calibration points. As such, molecular clock estimates calibrated on freshwater crab fossils favour a post-Gondwanan evolution of freshwater crabs whereas calibration based on marine brachyuran fossils date their last common ancestor before the fragmentation of Gondwana.     

Ionoregulatory and urinary responses to emersion in the mangrove crab Ucides cordatus and the intertidal crab Carcinus maenas

The mangrove crab Ucides cordatus showed a significant reduction in its output of isosmotic urine (to 60% of submerged rates) in humid, emersed conditions when in apparent water balance. A similar, but lesser, reduction was observed in the intertidal temperature species Carcinus maenas. Some of the decrease is effected by increased water withdrawal of an isosmotic fluid in the antennal gland (inulin urine to haemolymph concentration ratio=1.27) in Ucides but most is due to the reduced filtration rate which decreases from 3.41 to 2.19 ml·100 g^-1·day^-1 (in Carcinus from 7.37 to 5.88). In Carcinus, inulin urine to haemolymph concentration ratios are within the range 1.34–1.41 and show significant change on emersion. Comparisons of ion and inulin clearance ratios revealed interspecific differences in renal handling of Na⁺ and Cl^-. During emersion total Na⁺ efflux is markedly reduced in both species. Urinary Na⁺ losses are responsible for most of the efflux in Carcinus but represent only 50% of the total in Ucides. Significant Na⁺ uptake from interstitial water occurs in Ucides. REductions in ion and water loss appear to be mainly dependent on decreases in urine output (clearance rate). However, some released urine was found to accumulate in the branchial chambers in both species. Whether or not extrarenal reabsorption of ions from released urine occurs in the gill chambers in either crab species remains to be proved.Abbreviations CPM counts per minute - ECF extracellular fluid - k rate constant for inulin clearance - k ^Na rate constant for sodium efflux - J _out ^Na sodium efflux rate - Q _tot total radioactivity injected - SEM standard error of mean - SW sea water - 9-SW sea water, salinity 9 mg·ml^-1 (ppt) - 26-SW sea water, salinity 26 mg·ml^-1 (ppt) - U/B ratio of urine (U) to haemolymph (B) concentrations - V _c1 clearance rate - V _u urine output - V _a inulin space     

Entrainment of juvenile horseshoe crab activity to artificial tides

Juvenile American horseshoe crabs, Limulus polyphemus, express both daily and tidal rhythms. To determine if, and how, tidal cues influence the expression of these rhythms, we exposed 25 animals to artificial tides, and 17 to artificial tides with inundation, both with a 12:12 LD cycle. In the first experiment, 24% expressed daily rhythms of activity, 24% tidal rhythms, 12% a combination of the two, and the rest were arrhythmic. Under subsequent atidal conditions some expressed daily rhythms, but more were circatidal. In the second experiment, 6% expressed daily rhythms, 71% tidal, 12% a combination, and 12% were arrhythmic. Those expressing tidal rhythms were more active during flood/high tide, while daily animals tended to be nocturnal. Under subsequent constant conditions, the majority exhibited circatidal activity, with some expressing one activity bout per day. We conclude that juvenile horseshoe crabs entrain to artificial tides, with inundation cycles providing stronger cues than water depth changes.     

A common precursor to two major crab neurosecretory peptides

The biosynthesis of proteins by the X-organ sinus gland (XOSG) neurosecretory system of the crab, Cardisoma carnifex was studied using the pulse-chase technique. Analysis of radioactive proteins following 2D-PAGE showed that during pulse incubations of less than or equal to 30 min a single predominant 14Kd prohormone was synthesized. With chase less than or equal to 3 hr the primary 14Kd protein was found to undergo differential and/or multiple post-translational modifications prior to its proteolytic cleavage. Increasing the chase to greater than 3 hr showed a shift in labeling from the 14Kd forms to 3 separate 6Kd proteins. Two of the 6Kd proteins were identified as crustacean hyperglycemic peptides (CHH). Similarity in protein labeling using [3H]leucine and [35S]cysteine suggest a second major peptide group, the H peptide, known to lack cysteine, is also contained within the 14Kd precursor. Peptide mapping of the 14Kd proteins and of unlabeled CHH and peptide H provide substantive evidence for this biosynthetic scheme. Thus, both the CHH and H peptide groups, which together constitute greater than 90% of the XOSG peptide content, in this species, arise from a common 14Kd precursor molecule.     

Structural-functional differences of a crab motoneuron to four stomach muscles

A motor unit in the stomach of the blue crab, Callinectes sapidus, consists of four separate muscles involved in different aspects of the trituration and filtering of food. Motor nerve terminals to two of the muscles (CPV7a and GM5) release small amounts of transmitter (low-output) while those to the other two muscles (CV2 and CV3) release between three and five-fold greater amounts (high-output). Structural features underlying the disparity in synaptic strength were analysed with thin serial-section electron microscopy. Nerve terminals were similar in their volume percent of mitochondria, clear vesicles and dense core vesicles among the four muscles. This was also the case for the number and size of synaptic contacts. However, presynaptic dense bars representing active zones were longer and occurred more frequently at high-output synapses than at low-output ones. High-output synapses were also characterized by the close spacing of adjacent dense bars. The longer and more closely spaced dense bars at high-output synapses would be factors in the generation of larger synaptic potentials in these terminals compared to their low-output counterparts. Other factors, however, need to be considered to fully account for the physiological differences in synaptic strength among the four muscles.     

Responses to agonistic postures by the spider crab Microphrys bicornutus †

Models were presented to 102 specimens of the spider crab Microphrys bicornutus to test the importance of several appendage and body postures, which occur during agonistic interactions. The double first ambulatory raise posture and the cheliped spread posture both elicited a significant number of agonistic responses. Multiple ambulatory raise postures and raised or lowered body postures did not elicit significantly different response spectra. Size and sex of test animals seemed to have little effect on their responses.     

Number of ghost crab burrows does not correspond to population size

Ghost crabs are distributed worldwide on sandy beaches, and several studies have associated the number of ghost crab burrows with the levels of anthropogenic impacts on the beaches under study. However, our results show that the use of ghost crab Ocypode quadrata burrows to assess levels of anthropogenic impacts on sand beaches may not be accurate, as previously thought, because the number of burrows does not represent an estimate of the population size. In addition, we propose three hypotheses to explain the extremely low number of individuals/number of burrows ratio: the “secret chamber”, the “multiple openings”, and the “one crab, several burrows” hypotheses. We also observed an unusual sex ratio.     

Contributions made to chronobiology by studies of fiddler crab rhythms.

One of the classic organisms used in chronobiological research is the fiddler crab (genus Uca), an animal unique in that it displays both circadian and tidal (i.e., circalunidian) rhythms. The pioneering work on this animal helped produce the early evidence for many of the standard properties now recognized for all circadian rhythms: near temperature independence of the period, phase lability and setability, the light and temperature sensitivity rhythms expressed by phase response curves, and the persistence of rhythms in organs isolated from a multicellular animal. Importantly, results arising from studies of this crab--and a few other organisms--resulted in the development of the exogenous timing hypothesis. While philosophically sound, the lack of supporting evidence for this hypothesis has resulted in it being discarded by most chronobiologists; but while still in its prime, it drew great interest, and therefore grant support, to the field in general, stimulated a great deal of research that otherwise might not have been performed, and resulted in the discovery of environmental stimuli previously unsuspected to influence organisms. As could be expected, continuing work with this crab, using modern approaches and statistical techniques, has modified earlier findings and interpretations, has revealed new properties, and has resulted in the creation of new hypotheses. The review and update is a synthesis of 45 years of this work.     

Functional conversion of hemocyanin to phenoloxidase by horseshoe crab antimicrobial peptides

Arthropod hemocyanins and phenoloxidases serve different physiological functions as oxygen transporters and enzymes involved in defense reactions, respectively. However, they are equipped with a structurally similar oxygen-binding center. We have shown that the clotting enzyme of the horseshoe crab, Tachypleus tridentatus, functionally converts hemocyanin to phenoloxidase by forming a complex without proteolytic cleavage (Nagai, T., and Kawabata, S. (2000) J. Biol. Chem. 275, 35297-35301). Here we show that chitin-binding antimicrobial peptides of the horseshoe crab induce the intrinsic phenoloxidase activity of hemocyanin. Tachyplesin, a major Tachypleus antimicrobial peptide with an amphiphilic structure, converted the hemocyanin to phenoloxidase. Surface plasmon resonance analysis revealed the specific interaction of tachyplesin with hemocyanin at K(d) = 3.4 x 10(-)6 m. The chemical modification of Trp or Tyr in tachyplesin, but not Lys or Arg, dramatically reduced the affinity to hemocyanin, suggesting that the binding site is located in the hydrophobic face of tachyplesin. Hemocyanin has no affinity with chitin, but it significantly binds to tachyplesin-coated chitin, leading to the expression of phenoloxidase activity. The chitin coated with antimicrobial peptides may serve as a scaffold for the binding of hemocyanin, and the resulting phenoloxidase activity appears to function as a trigger of exoskeleton wound healing.     

A factor in crab amebocytes which stimulates in vitro clotting of crab blood

Cellular blood clotting in the crab which is initiated on contact with glass is rapidly accelerated in vitro by the addition of a crude amebocyte extract to the syringe and needle with which the blood is obtained. This accelerating action is destroyed by boiling, but is not destroyed by exposure to N-ethylmaleimide.     

Modelling the lethargic crab disease

The lethargic crab disease (LCD) is an emergent infirmity that has decimated native populations of the mangrove land crab (Ucides cordatus, Decapoda: Ocypodidae) along the Brazilian coast. Several potential etiological agents have been linked with LCD, but only in 2005 was it proved that it is caused by an ascomycete fungus. This is the first attempt to develop a mathematical model to describe the epidemiological dynamics of LCD. The model presents four possible scenarios, namely, the trivial equilibrium, the disease-free equilibrium, endemic equilibrium, and limit cycles arising from a Hopf bifurcation. The threshold values depend on the basic reproductive number of crabs and fungi, and on the infection rate. These scenarios depend on both the biological assumptions and the temporal evolution of the disease. Numerical simulations corroborate the analytical results and illustrate the different temporal dynamics of the crab and fungus populations.     

Variation in dog-whelk shells in relation to wave action and crab predation

In Nucella lapillus there is usually a progressive gradation in mean shell shape from short squat individuals on exposed headlands to more elongated ones in shelter. This pattern has been attributed to the differential selective effects of wave action and crab predation, although it would almost certainly be an oversimplification to imagine these as the only significant forces involved, especially near the limits of the species' distribution.
However, whilst this pattern of shell-shape variation with exposure is generally true over most of the animal's range in both Europe and North America, there are exceptions. There are occasional enclaves with much more elongated shells than would normally have been expected. Sometimes these are so abundant, as in the Severn Estuary, that all dog-whelk enclaves are unusually elongated, regardless of the exposure of their habitat.
In other areas, such as south-eastern England, the Solway Firth and the area around the mouth of the Conway the shells are not unusually elongated, but the enclaves show very little variation: a normal sheltered-shore form is seen wherever the species occurs. Shetland samples appear to be a combination of this pattern (predominating) with the 'normal' one showing the full range of shellshape variation.
Nucella appears in the Atlantic fossil record at the end of the Pliocene without any antecedent forms. It is assumed to have colonized from the Pacific and exploited the vacant niche before the onset of the Ice Age. Successive advances and retreats of the ice, with attendant changes in sea level, will have served to break up and recombine those dog-whelk populations that were not eliminated. It is postulated that the modern Nucella lapillus is a combination of several genetically distinct populations, some of which show the full range of variation and may respond to the selective effects of crabs and waves, and some of which do not and show little variation.