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.     

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.     

Size at the onset of sexual maturity in the anomuran crab, Aegla uruguayana (Aeglidae)

The size at maturity was studied in the crab Aegla uruguayana from the Areco River (31°14′ S, 59°28′ W), Argentina. Size at sexual maturity was determined according to three criteria: morphometric (change in the relative growth of reproductive characters), histological (first maturation of gonads) and functional (capability to mate and carry eggs). Regarding females, morphometric maturity occurred at a carapace length (CL) of 11.50 mm, considering abdomen width as a reproductive character. Gonad maturity of females could be observed at a minimum size ranging from 15 to 17 mm CL. The smallest ovigerous female observed in the field was 15.60 mm CL, although a relevant population incidence of ovigerous females (86.6%) has just been observed at values higher than 17 mm CL. As for males, the relative growth of the left chela length changed at a value of 15.40 mm CL, while morphological changes in sexual tube occurred between CL of 14 and 16 mm. Testicular maturation occurred at a CL ranging from 17 to 19 mm. The smallest size of males having spermatozoids in their vasa deferentia was 18.70 mm CL. The results obtained indicated that, in both sexes, functional maturity occurred after morphometric maturity and at a size similar to that of gonad maturity. Comparing sexes, females acquired sexual maturity (morphometric, gonad and functional maturity) at sizes statistically smaller than those of males.     

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.     

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.     

Size distribution,length–weight relationship and size at the onset of sexual maturity of the orange mud crab,Scylla olivacea,in Malaysian waters

The size distribution, length–weight relationship and size at the onset of sexual maturity of the orange mud crab (Scylla olivacea) from four geographically distinct locations (Taiping, Setiu, Kota Marudu and Lundu) representing Malaysian waters were analysed and estimated. Scylla olivacea was found in the size range of 47–134?mm carapace width. Males were significantly smaller in size but heavier than females. Geographical variation in carapace width and body weight were significant, but no interaction was found between sexes and locations. As shown by the length–weight relationships of S. olivacea, the males exhibited positive growth allometry whereas the females exhibited negative growth allometry. Males mature physiologically prior to attaining morphometric sexual maturity. Females, however, achieve physiological and morphometric sexual maturity in synchrony. No significant variation was found in the estimates of size at the onset of sexual maturity of males and females among different locations. We recommend the use of the third right walking leg merus length and carapace width to estimate the size at the onset of sexual maturity (morphometric maturity) for S. olivacea. Data obtained in this study serve as important baseline data for future mud crab resource management in Malaysia and were used to recommend minimum landing sizes for S. olivacea in each respective location based on the largest size at the onset of sexual maturity estimates were suggested.     

Relative growth,morphological sexual maturity,and size of Macrobrachium amazonicum (Heller 1862) (Crustacea,Decapoda, Palaemonidae) in a population with an entirely freshwater life cycle

The objective of this study was to estimate the size of morphological sexual maturity based on the study of relative growth, to determine the maximum size of individuals, and to determine if there are different morphotypes of males in a population of Macrobrachium amazonicum with an entirely freshwater life cycle. Collections were made monthly, with the use of a net, from September 2006 through August 2007. In each individual, the following structures were measured: carapace length (CL, in mm), width of the second pleuron (PlL, mm), length of the carpus (CaL, mm), and length of the propodus (PrL, mm). Relative growth was analyzed by observing the change in growth patterns of certain parts of the body in relation to the independent variable CL. The maximum sizes found were 8.5 and 11.4?mm CL for males and females, respectively. The morphometric variables: length of the carpus (CL?×?CaL) for males, and width of the second pleuron (CL?×?PlL) for females gave the best estimates for the size at maturation, which was 4.26?mm CL for males and 5.39?mm CL for females. The growth pattern in the different stages and the beginning of differential growth seemed to be closely related to reproductive aspects. No indices were found that separated the males into four different morphotypes, as proposed in the literature for coastal or artificially farmed populations. Only the male morphotype termed translucent claw was found in this population. The different morphological patterns in different regions are probably explained by ecological differences in the environments inhabited by these groups, principally in the availability of nutrients and the salinity in which the larvae develop.     

Sex‐specific differences in gonopore and gonadal growth trajectories in the brooding sea urchin,Abatus cavernosus (Spatangoida)

The heart urchin Abatus cavernosus shows sexual dimorphism characterized by the development of external brood pouches and the enlargement of gonopores in brooding females. Relationships between body size, gonopore size, and gonadal maturation in each sex were examined for inflection points using piecewise regression models (PRM). Opening of the gonopore occurred at 15.5 mm test length. Inflection points in the gonadal growth and gonopore diameter trajectories were clustered at smaller sizes in males (23 and 24.2 mm, respectively) than in females (25.1 and 25.9 mm), indicating sex‐specific differences in sexual maturation. Gonopore growth showed positive allometry at pre‐adult stages of development in both sexes, but isometry and negative allometry in adult females and males, respectively. Gonadal growth was initiated at smaller sizes and proceeded at a higher rate with increasing body size in males than in females. Identification of inflection points in gonopore and gonadal growth trajectories, using objective PRM, allows the determination of life stages and sexual maturation for individuals, thus providing a complementary tool for population studies.     

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.     

Insights of the life history in the porcellanid crab Petrolisthes armatus (Gibbes, 1850) (Crustacea: Anomura: Porcellanidae) from the Southwestern Atlantic coast

Petrolisthes armatus (Gibbes, 1850) life history was evaluated based on growth and reproduction. Specimens were captured at Ubatuba (Brazil), identified, sexed and measured: carapace (CL, length; CW, width), larger cheliped propodus (PL, length; PW, width; and PH, height), and 2th and 5th abdominal somite (AW₂ and AW₅, width). Relative growth was studied using CL as independent variable and its relation with other variables (dependents), to estimate puberty size and morphological maturity. Fecundity was represented by number of eggs vs. body size CL, with a better fit using a power function. Considering the 257 specimens analyzed, males were larger than females, and the overall sex ratio was 1:1, with some differences among size classes. Maturity (puberty size) was revealed by: AW₂ × CL (males: 8.6 mm CL; females: 7.6 mm CL); and AW₅ × CL (7.1 and 7.6 mm CL, respectively). Reproduction was continuous, but more intense in rainy season and recruitment occurring in dry season. Fecundity of this species was 228 ± 163 eggs, with better fit by a power function (R² = 0.72). Maturity size and growth differed when a pristine area (Ubatuba) was compared with a polluted area (São Sebastião), in the same Brazilian area (São Paulo State).     

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.     

SEX‐SPECIFIC PATTERNS OF MORPHOLOGICAL DIVERSIFICATION: EVOLUTION OF REACTION NORMS AND STATIC ALLOMETRIES IN NERIID FLIES

The consequences of sex‐specific selection for patterns of diversification remain poorly known. Because male secondary sexual traits are typically costly to express, and both costs and benefits are likely to depend on ambient environment and individual condition, such traits may be expected to diversify via changes in reaction norms as well as the scaling of trait size with body size (static allometry). We investigated morphological diversification within two species of Australian neriid flies (Telostylinus angusticollis, Telostylinus lineolatus) by rearing larvae from several populations on larval diets varying sixfold in nutrient concentration. Mean body size varied among populations of T. angusticollis, but body size reaction norms did not vary within either species. However, we detected diversification of reaction norms for body shape in males and females within both species. Moreover, unlike females, males also diversified in static allometry slope and reaction norms for static allometry slope of sexual and nonsexual traits. Our findings reveal qualitative sex differences in patterns of morphological diversification, whereby shape–size relationships diversify extensively in males, but remain conserved in females despite extensive evolution of trait means. Our results highlight the importance of incorporating plasticity and allometry in studies of adaptation and diversification.     

Gonadosomatic index‐based size at first sexual maturity of the catfish Eutropiichthys vacha (Hamilton, 1822) in the Ganges River (NW Bangladesh)

This study describes the size at first sexual maturity, length–weight relationships (LWR) in relation to size at first sexual maturity, and Fulton’s condition factor (K_F) of Eutropiichthys vacha in the Ganges River, northwestern Bangladesh. Sampling was done using traditional fishing gear including cast nets, square lift nets and conical traps during January and April, and July to December 2010. For each individual, total length (TL) was measured to the nearest 0.01 cm, and total weight (BW) was determined to the nearest 0.01 g. The gonadosomatic index (GSI) was calculated by the equation, GSI (%) = (Gonad weight in g/BW) × 100. The size at first sexual maturity of males and females was estimated by the relationship between gonadosomatic index and total length. A total of 583 specimens (289 males; 294 females) ranging from 8.30 to 27.00 cm TL and 3.16 to 159.50 g BW were analyzed. Sizes at first sexual maturity for male and female E. vacha were 13.15 and 14.00 cm TL, respectively. The analysis of covariance (ancova ) revealed significant differences in slope and intercept between early and late phases for males (F = 4.532, P < 0.001) and females (F = 21.984, P < 0.001). The K_F was not significantly correlated with TL for males (r_s = 0.052; P = 0.378), but was highly correlated for females (r_s = ?0.165; P = 0.005). This study establishes a strong base for monitoring changes in length at first sexual maturity attributable to high fishing pressures or other reasons within the Ganges and associated river ecosystems.     

Length‐weight relationships,condition factor,gonadosomatic index‐based size at first sexual maturity,spawning season and fecundity of Aspidoparia morar (Cyprinidae) in the Jamuna River (Brahmaputra River distributary), northern Bangladesh

The present study describes the length‐weight relationships (LWRs), length‐length relationships (LLRs), Fulton's condition factor (K_F), size at first sexual maturity, spawning season, sex ratio and fecundity of the Morari Aspidoparia morar (Hamilton, 1822) (Cyprinidae). Sampling was done using traditional fishing gear jhaki jal (cast net) from July 2010 to June 2011. Total length (TL), fork length (FL) and standard length (SL) were measured with digital slide calipers. Individual body weight (BW) and gonad weight (GW) were determined to an accuracy of 0.01 g for all specimens. The gonadosomatic index (GSI) was calculated and size at first maturity for males and females estimated using GSI and TL as indicators. Female ≥ size at first maturity was used to determine fecundity. A total of 1200 specimens (males = 552, females = 648) ranging from 4.06–12.84 cm TL and 0.53–16.75 g BW were analyzed. The overall coefficient b for the LWR indicated positive allometric growth (>3.00) in males and isometric growth in females (~ 3.00). ancova (analysis of covariance) revealed significant differences between males and females (P < 0.001). All LLRs were highly correlated (r² > 0.973, P < 0.001). Sizes at first sexual maturity for males and females were 6.0 and 7.0 cm TL, respectively. K_F changed little throughout the year and GSI peaked in November to April, indicating the spawning season (GSImax = 15.0 in females, 2.0 in males). Mature females were dominant during the entire spawning season except in April. Mean total fecundity was 6700 ± 3500, ranging from 1860 to 19680. In addition, relative fecundity ranged from 190 to 1200 (mean 560 ± 235) in the Jamuna River. To ensure sustainable management of this species, the protection of mature individuals during the peak spawning season is highly recommended.     

Annual reproductive cycle and sexual dimorphism in the dragonet,repomucenus valenciennei, in tokyo bay, japan

The annual reproductive cycle of the dragonet,Repomucenus valenciennei, from Tokyo Bay, Japan, was studied by analysis of seasonal trends in gonadosomatic indices and histological observations of gonads. Sexual dimorphism in the growth of several body parts relative to standard length (SL) and changes in color pattern of the first dorsal fin with growth were also investigated. The spawning season lasted from spring (April [1991] or February [1992]) to autumn (October) with two spawning peaks, in spring and autumn. In spring, only one-year-old (age 1+) fish spawned, age 0+ females not spawning until autumn, at which time they had reached age 1+. Likewise, histological observations of males indicated that testes had reached full maturity by 80 mm SL (age 1+). The minimum mature size of females was estimated as 60 mm SL. In males 45–80 mm SL, the first spine of the first dorsal fin, last ray of the second dorsal fin, last ray of the anal fin, and caudal fin ray showed strongly positive allometry, indicating rapid growth of these structures relative to SL. Subsequently, their growth returned to an isometric pattern in males>80 mm SL. In females, on the other hand, the same body parts showed slightly positive allometry throughout their growth. The color pattern on the first dorsal fin also changed in males 45–80 mm SL.     

Sexual size dimorphism,growth, and maturity of the fluvial eight-barbel loach in the Kako River,Japan

The maturation and growth pattern of the fluvial eight-barbel loach Lefua sp. (Japanese name: nagare-hotoke-dojo), an endangered species, was investigated using an individual identification-recapture method from 1995 to 1998 in an upper reach of a headwater tributary of the Kako River, Hyogo Prefecture, Japan. Based on observations of the gonads through the abdominal skin, the loach was estimated to breed mostly from May to July. All the males matured by age 1⁺, and all the females matured by age 2⁺. Gamete release in all individuals of both males and females was predicted from recaptured loaches during each breeding season. The standard length of mature females was significantly larger than that of males, showing sexual size dimorphism (SSD). The maximum sizes recorded were 75.4 mm SL for females and 61.2 mm SL for males. Both males and females of immature specimens grew mainly from May to November, including the breeding season, with no significant differences in growth rates between them. After sexual maturity, both males and females grew mainly from July to October (or November), after the breeding season, and the females exhibited higher growth rates than males. Therefore, SSD of the species seems to be attributable to the different growth rates after maturity. The longevity of the loach was estimated to exceed ten years based on individual growth patterns of various sizes during the survey period. It is likely that the loach has an iteroparous life history, breeding every year, and moderate growth rates after maturity.     

Population structure and reproduction of the longtail grass shrimp Urocaris longicaudata (Decapoda,Caridea, Pontoniinae) in Laguna de Términos,SW Gulf of Mexico

The purpose of this study was to describe the population structure and some reproductive traits of Urocaris longicaudata collected in Laguna de Términos, Campeche, Mexico in October and December 2009, and February and April 2010. A total of 2977 specimens were collected of which 1596 were females (including 276 ovigerous) and 1381 were males. Females were larger than males (2.03 ± 0.63 vs. 1.97 ± 0.44 mm carapace length (CL), respectively). The overall sex-ratio differed significantly from 1:1. The reproductive period was apparently continuous, with most ovigerous females recorded from December 2009 to April 2010. The estimated size at sexual maturity in females was 2.90 ± 0.64 mm CL. A significant and positive linear relationship between CL and realised fecundity was estimated. The number of embryos ranged from 33 to 320. The average volume of newly spawned embryos was 0.038 ± 0.009 mm³ and that of those near hatching was 0.062 ± 0.012 mm³, representing a 63.15% increase throughout embryogenesis. The females lost on average 27.6% of the initially produced embryos during the incubation period, which favoured the survival of the remaining embryos. This is the first report on any biological trait of U. longicaudata along the southwestern Gulf of Mexico.     

Notes on the biology of Cancer bellianus (Brachyura, Cancridae) around the Canary Islands

Information on biometric and biological parameters of Cancer bellianus Johnson, 1861 (Crustacea, Decapoda, Brachyura, Cancridae) off the Canary Islands is given. Crabs examined were collected during experimental fishing surveys during 1974–1998. Carapace length, carapace width, total wet weight, sex and ovigerous condition were determined. This species was caught at depths from 153 to 750 m, the deepest ever recorded. Size frequency distributions were assembled and size-weight relationships were estimated by sex. Sex-ratio as a function of size and depth was determined. The size at first maturity was calculated by analysing the relative growth between the carapace length and the left chela width: 103.5 mm CL in males, 101.2 mm CL in females. Ovigerous females, egg size and fecundity estimates are reported apparently for the first time.     

Use of pleopod morphology to determine sexual dimorphism and maturity in hermit crabs: <Emphasis Type="Italic">Isocheles sawayai</Emphasis> as a model

In the Anomura, studies on growth patterns are infrequent, possibly because the heterogeneity of the group, especially in terms of morphology, makes it difficult to construct generalized growth models. Particularly hermit crabs are an interesting group to evaluate aspects of growth, because of their unique body. Isocheles sawayai, a hermit crab found only in the western Atlantic Ocean, poorly known with respect to its sexual dimorphism and maturity, was investigated here based on morphometry. Monthly collections (July 2001 through June 2003) were made from a shrimp fishing boat in the Caraguatatuba region on the northern coast of the state of São Paulo, Brazil. The specimens were measured and weighed, and had their sex checked. Throughout the sampling period, 374 specimens of I. sawayai were collected (11.23% nonovigerous females, 6.69% ovigerous females, 79.41% males and 2.67% intersexes). The size at which morphological sexual maturity was reached by both sexes ranged from 4.0 to 4.3 mm shield length, according to the relative growth and the size of the smallest ovigerous female. Sexual dimorphism was shown by males, which were significantly larger than females, and by differences in growth pattern between the sexes, especially for relationships that involved the pleopods, which is related to their different functions in males and females. The present study is one of the first to use pleopod morphometry to determine sexual maturity and dimorphism in hermit crabs, especially for species with intersexuality such as I. sawayai.