Relative growth,morphological sexual maturity,heterochely, and handedness in Panopeus occidentalis (Brachyura,Panopeidae)

Studies on relative growth and sexual maturity are important to understand the reproductive biology of a species. The aims of this study were to determine the relative growth and to estimate the size of Panopeus occidentalis at morphological sexual maturity, as well as to confirm whether this species demonstrates heterochely or handedness. Individuals were collected every two months from March 2013 to July 2014 in the intertidal estuarine zone of Cananeia, São Paulo, Brazil. The following measurements were taken: carapace width (CW), carapace length (CL), right and left cheliped propodus length (CPL), right and left cheliped propodus height (CPH), right and left cheliped propodus width (CPW), abdomen width (AW), and first pleopod length (FPL). The morphometric relationship used to estimate the size at morphological sexual maturity were CW vs. AW for females and CW vs. FPL for males; these comparisons yielded estimated CW values of 15.60 mm and 16.67 mm, respectively. Heterochely was observed but handedness was not present. The species has a major cheliped on one side, but the side is not constant. This study provides the first insights on the relative growth, sexual maturity, heterochely, and handedness on a population of P. occidentalis in a conserved area.     

A Mathematical Modelling for the Cheliped Regeneration with Handedness in Fiddler Crab

An enormously developed giant cheliped with the other small one characterizes the adult male fiddler crab. Some experiments with artificial severances of cheliped indicate that such a handedness in the cheliped size is maintained even after the regeneration of severed cheliped. Other experimental researches give some results about an unknown physiological system which controls the emergence and the regeneration of the handedness in the cheliped size. In this paper, with two hypothesized factors relevant to the regeneration of a severed cheliped, we propose a simple mathematical model to describe the experimental result about the cheliped regeneration with a handedness after the cheliped severance for the fiddler crab. Our model gives a suggestion about an underlying system for the cheliped regeneration in the fiddler crab or some other crustacean species.     

Seasonal change in sexual size dimorphism of the major cheliped in the hermit crab <Emphasis Type="Italic">Pagurus minutus</Emphasis>

Sexual size dimorphism is a common phenomenon in the animal kingdom, and its seasonal change has been reported in some species that possess traits dimorphic only in males and specialized for male mating success. However, few studies have examined seasonal change in sexual dimorphism of traits possessed by both sexes. Here, we examined the reproductive biology of the hermit crab Pagurus minutus, at a sandflat in the Waka River estuary, Japan, with special reference to seasonal changes in sexual dimorphism of the large claw (major cheliped) size by conducting population and precopulatory guarding-pair sampling. Previous investigation demonstrated that the major cheliped is used as a weapon, and its size, more than body size, determines the winner in male–male contests of this species. We found ovigerous females from November to April, peaking in January, when 80% of females were ovigerous. Sexual size dimorphism of the major cheliped was observed; the degree of dimorphism increased in the reproductive season, when only males possessed an enlarged major cheliped. In addition, in the reproductive season, precopulatory guarding males had a larger body and larger relative size of the major cheliped than did solitary males, although the major cheliped size in guarding males seemed to reach an upper limit. These results suggest that seasonal change in sexual dimorphism of the major cheliped size in P. minutus strongly reflects sexual selection favoring the development of this natural weaponry, and that the degree of the dimorphism might be limited through natural selection.     

Biomechanics of chelipeds in some decapod crustaceans

The major chelipeds of five species of decapod crustaceans were studied with reference to lever system mechanical advantage, pattern of occlusive geometry, and force/pressure developed during cheliped closure by intact animals. Every cheliped type was found to possess a linear array of two to four distinctive regions of occlusion. The factors responsible for the differences in occlusive design are discussed. It is suggested that crustacean major chelipeds must be regarded as regionally-specialized, multifunctional appendages.     

Patterns of energy metabolism in the stone crab, Menippe mercenaria, during severe hypoxia and subsequent recovery

Specimens of the stone crab, Menippe mercenaria, survived severe hypoxia (PO2 less than 8mm Hg) for at least 12 hr at 28-30 degrees C. During the time course of 12 hr of hypoxia, hemolymph L-lactate levels rose to 30-50 mumoles/g wet wt. There was a slight elevation of L-alanine levels, whereas succinate was found in only trace quantities in the hemolymph. Pronounced metabolic changes took place in the heart, cheliped closer, and leg socket muscles during severe hypoxia. L-lactate accumulated to levels ranging from 16-20 mumoles/g wet wt. There were pronounced changes in high-energy phosphate levels in the cheliped closer and leg socket muscles. Taking into account expected intra- and extracellular water content, the calculated intracellular lactate content in the three muscles investigated is substantially less than the hemolymph lactate concentrations. Part of this reverse concentration gradient may be accounted for by the reduction in lactate activity due to cation-lactate complex formation. Hemolymph calcium and magnesium concentrations rose considerably during severe hypoxia. During recovery from severe hypoxia, approximately 50% of the accumulated lactate in the hemolymph was cleared in 6 hr. Hemolymph lactate and alanine levels returned to near control levels after 24 hr of recovery. This study shows that the stone crab, M. mercenaria, survives severe hypoxia by a reliance on glycogen fermentation to lactate. This species is capable of tolerating high levels of accumulated lactate.     

The enigma of reversed asymmetry in lithodid crabs: absence of evidence for heritability or induction of morphological handedness in Lopholithodes foraminatus

SUMMARY Mutations or environmental factors that result in reversal of conspicuous left–right asymmetries provide an opportunity to study developmental mechanisms. They may also provide insight into evolutionary changes in asymmetry states within and between species. King crabs (family Lithodidae) have a larger right claw and females typically exhibit a dextrally offset abdomen. Nevertheless, I observed a high incidence of left handedness in laboratory reared box crabs ( Lopholithodes foraminatus ) and captured the first known egg-bearing female lithodid to exhibit reversed asymmetry. This provided a unique opportunity to characterize the reversed phenotype and to compare the incidence of reversed asymmetry in the offspring of normal and reversed females. Asymmetry of the chelae became apparent in the first postzoeal stage (glaucothoe) and handedness was maintained through subsequent instars. Females with larger left claws developed reversed abdominal asymmetry by the fourth crab stage. No reversed asymmetry was observed in the mandibles of zoea larvae or juveniles of either handedness. The incidence of reversed asymmetry in glaucothoe reared from one reversed and three normal females was high (between 20% and 30%), and independent of maternity ( P =0.67). Removal of the right cheliped of fourth stage zoeae, and the major cheliped of glaucothoe, did not reverse the direction of asymmetry. Elevated larval rearing temperature also did not affect the frequency of reversed individuals. This lack of evidence for either heritability or induction of handedness is enigmatic. Further investigation of reversed asymmetry in lithodid crabs may provide valuable insights into the development and evolution of bilateral asymmetries.     

Do weaponless males of the hermit crab <Emphasis Type="Italic">Pagurus minutus</Emphasis> give up contests without escalation? Behavior of intruders that lack their major cheliped in male–male contests

In dyadic contests, theoretical studies have predicted that weaker contestants are less likely to engage in fights to minimize the cost of aggression. Since the major cheliped of decapod crustaceans is critically important as a weapon, contestants without a major cheliped should be more likely to give up the contests. We therefore examined whether loss of the major cheliped by the hermit crab Pagurus minutus would affect their decision to escalate male–male contests over guarded females. Intruders without a major cheliped showed no difference in the frequency of escalation compared with intact intruders, and the decision to give up was affected by the body size difference between the contestants. After escalation, compared with intact intruders, intruders without a major cheliped had significantly decreased success of takeover of a female from opponents, suggesting a strong disadvantage of losing their major cheliped. Although the decision of weaponless intruders to escalate seems irrational, several factors, such as poor accuracy of resource holding potential assessment, the influence of body size, and a high benefit to cost ratio of male–male contests, may have affected their behavior.     

Relative growth and morphological sexual maturity of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) (Decapoda,Brachyura, Sesarmidae) on the southern coast of the state of São Paulo,Brazil

Growth in crustaceans is characterized by ontogenetic differentiation during the development of their body structures, so studies on relative growth are widely applied in this group. In this study, the growth pattern of the body structures was verified through the analysis of relative growth, and then, morphological sexual maturity of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) was estimated. The carapace width (CW), cheliped propodus length (PL), cheliped propodus height (PH), propodus width (PW), abdomen width (AW), and first pleopod length (FPL) of the crabs were measured. The relationships that best showed changes in the allometric coefficient among demographic categories were FPL vs. CW for males and AW vs. CW for females. This study verified the increased size of the cheliped in terms of length, width, and height, which occurred mainly in adult males. This increase reflects the importance of this structure in the reproductive processes of A. pisonii. For females, the increase in abdomen growth reflects their reproductive potential, since it is a structure that provides protection for eggs.     

The timing of decisions during shell investigation by the hermit crab, Pagurus bernhardus

Factors which influence decisions by hermit crabs concerning whether to approach, investigate and enter another shell have been investigated by systematically varying the size of the shell in possession and the size of the shell being offered. The probability that a crab will approach or enter the shell depends on both variables. Investigation after contact, however, depends only on the size of the shell in possession. Durations of each stage of the sequence are negatively correlated with the numbers of crabs in each experimental group performing the next stage. For the final sequence of aperture investigatory activities, the fewer crabs entering the offered shell, the greater the number of investigatory acts performed by those crabs. In addition to variation in number of acts and duration of investigation, specific use of appendages varies according to the experimental situation. Thus naked crabs use the minor cheliped to investigate shells in preference to the major cheliped, which is normally used by housed crabs, with the major cheliped being reserved for defence by naked crabs. These data are discussed in terms of information collection and decisions made on the basis of that information.     

Shell fights in the hermit crabPagurus geminus: Effect of cheliped use and shell rapping

In shell fights of the hermit crab,Pagurus geminus, frequently it is observed that large crabs (attackers) grasp the thoracic appendage of small crabs (defenders) with the major cheliped and pull the smaller crabs out of their shells. If this is a standard occurrence and result, then the interaction should not be called a “negotiation” (Hazlett 1978). The role of cheliped use by the attckers in the eviction of defenders was therefore studied using crabs with tubes on their chelipeds, and the effect of shell rapping, which is thought to be necessary for eviction, was studied using crabs without shells. The experimental crabs evicted the defenders but fighting was significantly prolonged. Therefore, the negotiation model cannot be rejected. Specific aspects of shell fights in hermit crabs are discussed.     

Structure,composition and properties of naturally occurring non-calcified crustacean cuticle

Crustaceans are known for their hard, calcified exoskeleton; however some regions appear different in colour and opacity. These include leg and cheliped tips in the grapsid crab, Metopograpsus frontalis. The chelipeds and leg tips contain only trace levels of calcium but a significant mass of the halogens, chlorine (Cl) and bromine (Br). In contrast, the carapace is heavily calcified and contains only a trace mass of Cl and no Br. In transverse section across the non-calcified tip regions of cheliped and leg the mass percent of halogens varies with position. As such, the exoskeleton of M. frontalis provides a useful model to examine a possible correlation of halogen concentration with the physical properties of hardness (H) and reduced elastic modulus (E_r), within a chitin-based matrix. Previously published work suggests a correlation exists between Cl concentration and hardness in similar tissues that contain a metal (e.g. zinc). However, in M. frontalis H and E_r did not vary significantly across cheliped or leg tip despite differences in halogen concentration. The non-calcified regions of M. frontalis are less hard and less stiff than the carapace but equivalent to values found for insect cuticle lacking metals. Cheliped tips showed a complex morphological layering that differed from leg tips.     

Relative growth and morphological sexual maturity of the freshwater crab Trichodactylus fluviatilis Latreille 1828 (Decapoda,Trichodactylidae) from west central São Paulo State,Brazil

An analysis of relative growth was performed to determine the size at morphological sexual maturity of Trichodactylus fluviatilis Latreille 1828. Samples were collected by trapping at night from January 2010 through June 2011 from the Barreiro River (22°34′42″S; 49°11′92″W), west central São Paulo State, Brazil. The following measurements were obtained (mm): carapace width (CW) (independent variable), cheliped propodus length (PL), cheliped propodus height (PH), cheliped propodus width (PW), first pleopod length, and abdomen width (AW). The males showed positive allometry in the juvenile and adult stages for the following relationships: PL vs. CW, PH vs. CW, and PW vs. CW. In the females, a positive allometry was observed for AW vs. CW in the juvenile stage, and a negative allometry was observed for this relationship in the adult stage. The differential growth of the chelipeds in males could be related to territorial conflict and courtship, whereas the changes in the growth of the female abdomen most likely favor higher reproductive success.     

Does the handedness of the pebble crab Eriphia smithii influence its attack success on two dextral snail species?

The predaceous crab Eriphia smithii (Xanthidae) has one larger claw with molar teeth on either the right or the left cheliped, which it uses to crush the shell of prey. Whether the handedness of crabs affected successful predation on two snail species, Nerita albicilla (Neritidae) and Planaxis sulcatus (Planaxidae) was experimentally investigated. The fate of snails of each species was analysed by multiple logistic regression with three explanatory variables: handedness, shell-size index and individuality of crabs. No effect of handedness was detected in attacks on N. albicilla , probably as a result of the spherical and more symmetrical shell morphology of this species. In contrast, right-handedness contributed to greater attack success on P. sulcatus , which has more conical shells. Further investigation of how snail shells were broken revealed that left-handed crabs had more difficulty breaking the aperture of larger P. sulcatus , which was thought to cause the difference in attack success between right- and left-handed crabs. The advantages conferred by handedness are discussed.     

Relative Growth and Sexual Maturity of the Hermit Crab Paguristes erythrops (Anomura, Diogenidae) from South Atlantic

Some morphometric relationships were studied in the hermit crab Paguristes erythrops to describe its relative growth and the size at which sexual maturity is reached, as well as the shell influence on this process. Individuals were collected monthly from January to December/1999, by SCUBA diving and 12 different characters were measured on males and females. Sexual size dimorphism was observed in length and width of left cheliped. Size at sexual maturity was estimated between 2.8 and 3.5 mm shield length based on the left cheliped relations of the males and on the endopod and exopod of the second pleopod of the females. Differences in the allometry of individuals occupying different shell species were observed. The present work demonstrated the importance of characterizing the type of growth of some parts (pleopods) not yet used in other studies on relative growth of hermit crabs that may correspond to indicators of the size at sexual maturity.     

Differences in the parasitic effects of a bopyrid isopod and rhizocephalan barnacle on the portunid crab,Charybdis bimaculata

Bopyrid isopods and rhizocephalan barnacles are obligate parasite crustaceans which harm their decapod hosts. However, to the best of our knowledge, studies have not compared which of these parasites has a greater parasitic effect on its hosts. Here, the parasitic effect of the bopyrid isopod, Allokepon hendersoni, and an unidentified sacculinid rhizocephalan species, infesting the same population of portunid crabs, Charybdis bimaculata, was investigated and compared for the first time. Samples were collected from the bycatch of a trawl fishery in Tosa Bay, Japan. A total of 2601 crabs were collected, of which 14 (0.55%) were parasitized by the bopyrid and 21 (0.82%) by the rhizocephalan. One of the two female crabs parasitized by the bopyrid was ovigerous (with much fewer eggs than unparasitized females). No ovigerous crab was found from the eight females parasitized by the rhizocephalan. Because only two female crabs were parasitized by the bopyrid, the following analyses were made using the male crabs. Both parasites reduced the wet weight (crab condition) and the cheliped size (secondary growth) of C. bimaculata, but the impact of the parasitism did not differ between the parasite species. The size of the abdominal flap of male hosts was reduced by the bopyrid infestation; however, rhizocephalan infestation caused enlargement of the abdominal flap, which is an indication of feminization. The present study provides information on how the effect of these two parasitic castrators on the same host crab varies. A moderate decrease in crab condition and cheliped development was common among the parasites, suggesting that the degree of impact might be favorable for the survival of the two parasites species.     

Population structure,relative growth and morphological sexual maturity of the snapping shrimp Alpheus brasileiro Anker, 2012 (Caridea: Alpheidae) from the south-eastern coast of Brazil

The present study investigated the population structure of the snapping shrimp Alpheus brasileiro. We tested the hypotheses that the sex ratio of the population differs from the expected 1:1 and that the growth pattern differs between adults and juveniles and between sexes. Reproductive period, juvenile recruitment and morphological sexual maturity were estimated. Samples were collected in two periods. The first sampling occurred bimonthly from March 2013 to January 2014. The second sampling occurred monthly from April 2015 to March 2016. The population structure was determined based on the size-frequency distribution of different demographic categories. To analyse the relative growth, we measured the second pleuron length, major cheliped propodus length, major cheliped propodus width, major cheliped propodus height, appendix interna length, appendix masculina length. The results showed that the sex ratio did not differ from 1:1 (male:female). Reproduction and juvenile recruitment were continuous throughout the study period. The analysis of relative growth confirmed the hypothesis that males and females, and juveniles and adults, have different growth patterns, indicating distinct strategies of energy allocation. The estimated carapace length at the onset of morphological sexual maturity was 4.9?mm in males and 4.7?mm in females. The results obtained in this study are probably related to the monogamous social behaviour of A. brasileiro. Monogamy can influence demographic parameters such as sex ratio and the size at which males and females reach morphological sexual maturity.     

Relative growth and onset of morphological sexual maturity of the freshwater crab Sylviocarcinus pictus in a river of the Caatinga of northeastern Brazil

The aim of this study was to investigate the relative growth and onset of morphological sexual maturity of the freshwater crab Sylviocarcinus pictus. Specimens were collected every month from October 2013 to September 2014, at night, on a stretch of the river Guaribas, Piauí, Brazil. Crabs were sexed and their carapace width (CW, independent variable), carapace length (CL), cheliped propodus length (PL) and height (PH), gonopod length (GL), and abdomen width (AW) (dependent variables) were measured. These measurements were related to characterise relative growth and possible sex differences. On average males were larger than females (p = 0.0001). Size at the onset of morphological sexual maturity was defined by relating CL vs. PL for males (30.82 mm) and CL vs. AW for females (28.63 mm). These are considered secondary sexual characters and reflect initial size at morphological sexual maturity with greater precision. The differential cheliped growth of males may be related to courting and disputes with other males, while the allometric growth of the abdomen of females indicates increased probability of reproductive success.     

Relative growth and morphological sexual maturity of the caridean shrimp Nematopalaemon schmitti (Decapoda: Caridea: Palaemonidae) in an upwelling region in the Western Atlantic*

In crustaceans, successful reproductive processes, such as the transition from juvenile to adult, exhibit important morphological changes that can be detected by analyzing relative growth. This study describes the relative growth of body structures in Nematopalaemon schmitti and its secondary sexual characteristics, and also estimates the morphological sexual maturity of this species in a region influenced by upwelling. The carapace length (CL), second pleuron length (PlL), cheliped carpus length (CaL), cheliped propodus length (PrL) and the length of appendix masculina (AML) of the shrimp were measured. The relationships that best demonstrated the changes in allometric coefficient between demographic categories were AML vs. CL for males, and PlL vs. CL for females. The estimated CL for morphological sexual maturity in males was 8.51 mm and 9.30 in females. Our results showed the appendix masculine and the second pleuron were secondary sexual characteristics that play roles in reaching the morphological sexual maturity necessary for reproductive success and to assure the life cycle of this species.     

A new species of the ghost shrimp genus Neocallichirus Sakai, 1988 (Decapoda: Axiidea: Callianassidae) from the southwestern Atlantic

ABSTRACT

A new species of ghost shrimp, Neocallichirus pinheiroi, is described based on specimens collected in subtidal habitats of Saquaira beach, northeastern coast of Brazil. This new species resembles N. lemaitrei Manning, 1993, but differs from it primarily in having a third maxilliped propodus expanded ventrally, and in the case of males, a large subtriangular tooth on the cutting edge of the major cheliped dactylus. Diagnostic features of the new species include: (i) frontal margin of carapace with rounded lateral projections not exceeding rostrum distal margin, (ii) third maxilliped carpus with row of sclerotized setae on median internal surface, (iii) minor cheliped with cutting edge of dactylus microscopically pectinate, (iv) endopod of male second pleopod with appendices interna and masculina. This is the seventh described species of the genus Neocallichirus from the western Atlantic and the fourth report of this genus in Brazil. A key for identification of western Atlantic species belonging to the genus is provided.     

Histochemical localization of glycogen and some representative enzymes of glycolytic cycle in the cheliped muscle of Scylla serrata (Forskal) (Brachyura: Decapoda)

Glycogen, phosphorylase, aldolase, a-glycerophosphate dehydrogenase, lactic dehydrogenase and glucose-6-phosphate dehydrogenase were localized histochemically in the cheliped muscle of Scylla serrata (Forskal) in normal, destalked, destalked receiving eyestalk extract injection and intact injected with eyestalk extract.