Patterns of distribution of the hermit crab Loxopagurus loxochelis (Moreira, 1901) (Decapoda, Diogenidae) in two coastal areas of southern Brazil

The present study determined the distribution pattern of the hermit crab Loxopagurus loxochelis by a comparison of catch, depth and environmental factors at two separate bays (Caraguatatuba and Ubatuba) of Sao Paulo State, Brazil. The influence of these parameters on the distribution of males, non- ovigerous females and ovigerous females was also evaluated. Crabs were collected monthly, over a period of one year (from July/2002 to June/2003), in seven depths, from 5 to 35 m. Abiotic factors were monitored as follows: superficial and bottom salinity (psu), superficial and bottom temperature (C), organic matter content (%) and sediment composition (%). In total, 366 hermit crabs were sampled in Caraguatatuba and 126 in Ubatuba. The highest frequency of occurrence was verified at 20 m during winter (July) in Caraguatatuba and 25 m during summer (January) in Ubatuba. The highest occurrences were recorded in the regions with bottom salinities ranging from 34 to 36 psu, bottom temperatures from 18 to 24 C and, low percentages of organic matter, gravel and mud; and large proportion of sand in the substrate. There was no significant correlation between the total frequency of organisms and the environmental factors analyzed in both regions. This evidence suggests that other variables as biotic interactions can influence the pattern of distribution of L. loxochelis in the analyzed region, which is considered the limit of the northern distribution of this species.

Comparative analysis of population structure of endemic Western Atlantic hermit crab Loxopagurus loxochelis (Decapoda, Anomura) in two regions of state of Sao Paulo, Brazil

This work characterized the population structure of the hermit crab Loxopagurus loxochelis (Moreira, 1901) in terms of size frequency distribution and sex ratio. Specimens were collected monthly, over a period of one year (from July 2002 to June 2003), in seven transects (from 5 to 35 m of depth) using fishing boat equipped with two double-rig trawl nets, in Caraguatatuba and Ubatuba regions (state of Sao Paulo, Brazil). A total of 366 hermit crabs were collected in Caraguatatuba [222 males (60.65%), 114 non-ovigerous females (31.15%) and 30 ovigerous females (8.20%)] and 126 hermit crabs in Ubatuba [81 males (64.28%), 38 non-ovigerous females (30.16%) and seven ovigerous females (5.56%)]. In Caraguatatuba the highest incidence of ovigerous females occurred during winter (July 2002), whereas in Ubatuba, the number was incipient. The cephalothoracic shield length ranged from 2.0 to 7.9mm (5.29 +/- 0.96mm) in Caraguatatuba, and from 2.7 to 7.5mm (5.32 +/- 0.95mm) in Ubatuba. The mean size of males was significantly larger than the mean size of females in both regions. Overall sex ratio was in favor of males (1.54:1 in Caraguatatuba and 1.9:1 in Ubatuba). Sexual dimorphism was recorded to L. loxochelis by the presence of males in the largest size classes, following the standard pattern observed in Decapoda. There was an unimodal size distribution for both sexes, with normal distributions in both regions. The higher number of males in relation to females may indicate the existence of different growth and mortality rates between the sexes. Despite of the different geomorphologic characteristics between Caraguatatuba and Ubatuba regions, the dynamics of development was similar for both populations.

Uncommon mechanism of egg incubation in the endemic Southern hermit crab Loxopagurus loxochelis: how is this phenomenon related to egg production?

The aim of the present work was to characterize the egg production of Loxopagurus loxochelis. A total of 71 ovigerous females were obtained, 28% of which were simultaneously incubating eggs at different developmental stages. This phenomenon can be the result of incomplete fertilization, or may represent a rapid gonadal cycle by this species in this area, which is, to our understanding, the best explanation of this phenomenon. Egg volume decreased 25.6% during the incubation period. The reproductive output based on dry and wet weight was 6.8 and 19.3%, respectively. Water was the prevailing component of the eggs, representing 86.0% of the total weight at initial stage, increasing to 95.1% at the final stage. Ash content increased at the same time as a decrease in the organic content occurred, indicating the consumption of yolk and absorption of salts from the water medium. In conclusion, we hypothesized that this population has the capacity to copulate and deposit another brood even before the release of the larvae from the previous one, intensifying the reproductive effort of these hermit crabs as a strategy of adaptation in a region considered the northern limit of the geographical distribution of this species.

Análise comparativa da estrutura populacional do ermitão endêmico do Atlântico Ocidental Loxopagurus loxochelis (Decapoda, Anomura) em duas regiões do estado de São Paulo, Brasil

A estrutura populacional do ermitão Loxopagurus loxochelis (Moreira, 1901) foi analisada por meio da distribuição de freqüência dos animais em classes de tamanho e da razão sexual. Espécimes foram coletados mensalmente no período de um ano (julho de 2002 a junho de 2003), em sete transectos (5 aos 35 m de profundidade), usando barco equipado com duas redes do tipo double-rig, nas enseadas de Caraguatatuba e de Ubatuba (Estado de São Paulo, Brasil). Em Caraguatatuba foram coletados 366 animais, sendo 222 machos (60,65%), 114 fêmeas não-ovígeras (31,15%) e 30 fêmeas ovígeras (8,20%) e, em Ubatuba, 126 ermitões, dos quais 81 machos (64,28%), 38 fêmeas não-ovígeras (30,16%) e sete fêmeas ovígeras (5,56%). Em Caraguatatuba, a alta incidência de fêmeas ovígeras ocorreu durante o inverno (julho de 2002), enquanto em Ubatuba, o número foi incipiente. O comprimento do escudo cefalotorácico variou de 2,0 a 7,9mm (5,29 ± 0,96mm) em Caraguatatuba, e em Ubatuba, de 2,7 a 7,5 mm (5,32 ± 0,95mm). A média de tamanho dos machos foi significativamente maior do que das fêmeas em ambas as áreas. A razão sexual total foi favorável aos machos (1,54:1 em Caraguatatuba e 1,9:1 em Ubatuba). Houve maior freqüência de machos nas últimas classes de tamanho, acompanhando os padrões observados em decápodes e evidenciando a existência de dimorfismo sexual em L. loxochelis. A distribuição de freqüência total foi unimodal para ambos os sexos, nas duas áreas. A presença de um número maior de machos em relação a fêmeas pode ser indício de que existam diferentes taxas de crescimento e de mortalidade entre os sexos, além de distribuição espacial diferencial entre estes. Apesar das diferentes características geomorfológicas entre as regiões de Caraguatatuba e Ubatuba, foi observada uma dinâmica de desenvolvimento similar para ambas as populações.

Ecological distribution and reproductive period of the hermit crab Loxopagurus loxochelis (Anomura, Diogenidae) on the northern coast of São Paulo State, Brazil

The present study aimed to determine the ecological distribution of Loxopagurus loxochelis as a function of selected environmental factors, as well as its reproductive period based on the combined analysis of the presence of ovigerous females and the development of the gonad. The collections were carried out monthly from January to December 2000 in the region of Ubatuba (SP), using a fishing boat equipped with two double rig nets, in isobaths of 5, 10, 15, 20, 25, 30, 35 and 40 m, at which depths samples of sediment and water were collected. The cephalothoracic shield length and sex of the animals were determined; the abdomen was dissected to verify gonad development stages. A total of 334 individuals was obtained ( 196 males, 48 ovigerous females and 90 non-ovigerous females) with sizes varying from 3.0 to 8.6 mm (5.7 +/- 1.0 mm). A greater abundance of L. loxochelis (95.2%) was observed at depths of 20 and 25 m. These sites mainly revealed a substratum stable with a very fine sand fraction, which facilitates the habit of embedding shown by this hermit. With regard to gonadal analysis, it was possible to classify four gonad development stages for each sex: immature, rudimentary, developing and developed. It was observed that in the winter months about 71% of the females had gonads either developing or developed and 67% were ovigerous. Therefore, the winter period can be considered the peak of reproduction for this species, characterizing a typical seasonal-continuous reproduction.

Population structure and breeding season of the South Atlantic hermit crab, Loxopagurus loxochelis (Anomura, Diogenidae) from the Ubatuba region, Brazil

A population of Loxopagurus loxochelis was studied in terms of seasonal abundance, size frequency distribution, sex ratio, and reproductive period (percentage of ovigerous females). Specimens were collected monthly over a period of two years (from September 1995 to August 1997) in the non-consolidated areas of the Ubatuba, Mar Virado, and Ubatumirim bays (northern coast of São Paulo State, Brazil) with a double-rig trawl net. A total of 1,084 individuals were analysed. Animal size (minimum, maximum, and mean +/- SD shield length) was 2.8, 9.1, and 6.88 +/- 1.13 mm for 625 males; 2.8, 8.2, and 5.78 +/- 0.98 mm for 236 non-ovigerous females; and 4.6, 8.0, and 6.24 +/- 0.68 mm for 223 ovigerous females, respectively. Sexual dimorphism was recorded by the presence of males in the largest size classes. The sex ratio was 1.4 : 1 in favour of males. The highest incidence of ovigerous females occurred during winter and spring (June to October), with a low percentage in summer and fall (November to May), indicating continuity in the reproductive cycle. This strategy of reproduction is related to temperature. The occurrence of L. loxochelis in the Ubatuba region represents the final point of northern distribution of this species as a function of water mass influence, and could be its real limit on the Atlantic coast of South America.

Temporal-spatial distribution of the hermit crab Loxopagurus loxochelis (Decapoda: Diogenidae) from Ubatuba Bay, São Paulo State, Brazil

The present work was conducted to determine the distribution of Loxopagurus loxochelis collected monthly, over a period of one year, in Ubatuba Bay (from September/95 to August/96). A total of 179 specimens were collected (138 males, 30 females and 11 ovigerous females). The greatest depth, predominance of very fine sand and highest mean value of organic matter contents of sediment, in combination with the low temperatures registered in winter (July and August), determined the presence of L. loxochelis in the subarea located at the Ubatuba Bay mouth, exposed to the open sea with high water current energy, important because this position insures that spawned larvae will enter into the oceanic circulation.

Shell occupation by the South Atlantic endemic hermit crab Loxopagurus loxochelis (Moreira, 1901) (Anomura: Diogenidae)

The evaluation of population characteristics, particularly those of endemic species, aids in population preservation and management. Hermit crabs present an innate behavior of occupying shells, which tends to individual needs and limits their distribution. This study characterized the pattern of occupation of gastropod shells by the hermit Loxopagurus loxochelis in three bays of the southwestern coast of Brazil. Monthly collections were made from January/1998 to December/1999 in the bays Ubatumirim (UBM), Ubatuba (UBA) and Mar Virado (MV) with a shrimping boat. Overall, ten species of gastropod shells were occupied by L. loxochelis. The shell of Olivancillaria urceus represented 66.8% of those occupied. Morphometric relationships demonstrated a differential occupation of the more abundant shells among demographic groups, where most of the males occupied O. urceus, non-ovigerous females occupied O. urceus and Buccinanops cochlidium, and ovigerous females occupied B. cochlidium and Stramonita haemastoma. Most of the individuals occupied the more abundant shells, considered adequate for the morphology of this hermit crab species. Thus, the studied bays seem to be stable and propitious environments for population perpetuation and the settlement of new individuals.

Comparative analysis of shell occupation by two southern populations of the hermit crab Loxopagurus loxochelis (Decapoda, Diogenidae)

The present study aimed to comparatively verify the relation between the hermit crabs and the shells they use in two populations of Loxopagurus loxochelis. Samples were collected monthly from July 2002 to June 2003, at Caraguatatuba and Ubatuba Bay, Sao Paulo, Brazil. The animals sampled had their sex identified, were weighed and measured; their shells were identified, measured and weighed, and their internal volume determined. To relate the hermit crab's characteristics and the shells' variables, principal component analysis (PCA) and a regression tree were used. According to the PCA analysis, the three gastropod shells most frequently used by L. loxochelis varied in size. The regression tree successfully explained the relationship between the hermit crab's characteristics and the internal volume of the inhabited shell. It can be inferred that the relationship between the morphometry of an individual hermit crab and its shell is not straightforward and it is impossible to explain only on the basis of direct correlations between the body's and the shell's attributes. Several factors (such as the morphometry and the availability of the shell, environmental conditions and inter- and intraspecific competition) interact and seem to be taken into consideration by the hermit crabs when they choose a shell, resulting in the diversified pattern of shell occupancy shown here and elsewhere.

Comparative analysis of shell occupation by two southern populations of the hermit crab Loxopagurus loxochelis (Decapoda, Diogenidae)

The present study aimed to comparatively verify the relation between the hermit crabs and the shells they use in two populations of Loxopagurus loxochelis. Samples were collected monthly from July 2002 to June 2003, at Caraguatatuba and Ubatuba Bay, São Paulo, Brazil. The animals sampled had their sex identified, were weighed and measured; their shells were identified, measured and weighed, and their internal volume determined. To relate the hermit crab's characteristics and the shells' variables, principal component analysis (PCA) and a regression tree were used. According to the PCA analysis, the three gastropod shells most frequently used by L. loxochelis varied in size. The regression tree successfully explained the relationship between the hermit crab's characteristics and the internal volume of the inhabited shell. It can be inferred that the relationship between the morphometry of an individual hermit crab and its shell is not straightforward and it is impossible to explain only on the basis of direct correlations between the body's and the shell's attributes. Several factors (such as the morphometry and the availability of the shell, environmental conditions and inter- and intraspecific competition) interact and seem to be taken into consideration by the hermit crabs when they choose a shell, resulting in the diversified pattern of shell occupancy shown here and elsewhere.

Comparative reproductive biology of two southwestern Atlantic populations of the hermit crab Pagurus exilis (Crustacea: Anomura: Paguridae)

Pagurus exilis (Benedict, 1892) is an endemic South Atlantic hermit crab with a distribution ranging from Rio de Janeiro State, Brazil, to Buenos Aires Province, Argentina. The present study analyzed the reproduction of two populations at the extremes of this geographical distribution, and compared their reproductive period, fecundity, and changes in egg size and egg volume during the incubation period in order to assess the variability over this latitudinal range. Hermit crabs were collected monthly over a 2-year period. In total, 108 females were analyzed for Brazil (44 non-ovigerous and 64 ovigerous), and 141 for Argentina (87 non-ovigerous and 54 ovigerous). Reproduction in Brazil occurs year-round, with peaks in the fall and winter seasons; in Argentina reproduction occurs only in spring and summer. The Brazilian ovigerous females were significantly larger than the Argentina ones (Brazil: SL = 5.33 +/- 1.45 mm; Argentina: SL = 4.15 +/- 0.52 mm; P < 0.001). The fecundity was 1447 +/- 831 eggs (317 to 2885) in Brazil and 987 +/- 711 eggs (114 to 2665) in Argentina, with a trend towards higher fecundity in Brazil. Eggs in the Argentina population were larger than those in Brazil for all the three stages investigated, and no changes in egg volume were found during egg development for both populations. The reproductive traits of the two populations showed some important differences, which may reflect adaptations to local environmental conditions, demonstrating a high plasticity of reproductive features of the species in Brazilian and Argentine waters. The strategy adopted by the Argentina population involves a lower production of larger eggs compared to the population in Brazil; this lower production is associated with reproduction in cold-water regions.

Morphological and morphometric appraisal of the spermatophore of the southern hermit crab Isocheles sawayai Forest and Saint Laurent, 1968 (Anomura: Diogenidae), with comments on gonopores in both sexes

The spermatophore morphology of the hermit crab Isocheles sawayai from Southwestern Atlantic (Brazil) is described. The spermatophores show similarities with those described for other members of the family Diogenidae, especially with the recently described Loxopagurus loxochelis. The spermatophore is composed of three major regions: a sperm filled head or ampulla, a columnar stalk and a foot or pedestal. The spermatophores show specific morphology in having a circular ampulla, and a constriction or neck between the ampulla (100 mu m) and the thin (27 mu m), long stalk (500 mu m). The stalk penetrates less than half way into the spermatophore head. Most spermatophores show one or the small posterior projections on the underside of the ampulla as being bigger than the other, making it asymmetrical. The size of the spermatophore is related to hermit crab size with direct relationships found between spermatophore ampulla width, total length, and peduncle length with shield length of the hermit crab. The morphological characteristics of the spermatophore of L sawayai. are species-specific distinguishing it from other members of the family, and are useful to infer further phylogenetic relationships. (C) 2008 Elsevier GmbH. All rights reserved.

Use of pleopod morphology to determine sexual dimorphism and maturity in hermit crabs: Isocheles sawayai 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 SA o pound 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.

Ocupação de conchas de gastrópodes por ermitões (Decapoda, Anomura) no litoral de Rio Grande, Rio Grande do Sul, Brasil

O presente estudo teve como objetivo caracterizar a ocupação de conchas por ermitões no litoral da cidade de Rio Grande, Estado do Rio Grande do Sul. Os animais foram amostrados em 14 radiais em Rio Grande, entre 12 e 50 metros de profundidade. Cada ermitão e sua respectiva concha foram identificados, pesados e medidos. Um total de 408 animais foi capturado, pertencentes às famílias Diogenidae e Paguridae; as duas espécies mais abundantes foram Dardanus insignis (de Saussure, 1858) e Loxopagurus loxochelis (Moreira, 1901). Os ermitões ocuparam conchas de 13 espécies de gastrópodes, principalmente de Buccinanops lamarckii (Kiener, 1834) e B. gradatum (Deshayes, 1844). Dardanus insignis utilizou 12 das 13 espécies de moluscos registradas; Loxopagurus loxochelis utilizou nove. De um modo geral, o padrão de ocupação de conchas apresenta uma correlação entre o tamanho do ermitão e o tamanho da concha; no caso das duas espécies de ermitões mais abundantes, a maior correlação foi entre peso/tamanho do animal e o tamanho da abertura da concha, evidenciando, que a ocupação de conchas se dá não apenas pela disponibilidade local das mesmas, mas também pelas relações entre as variáveis dos ermitões e das conchas de gastrópodes.

Ecological aspects of hermit crabs (Crustacea, Anomura, Paguroidea) off the northern coast of São Paulo State, Brazil

The occurrence of species of hermit crabs and their ecological distribution in soft bottoms off Ubatuba, São Paulo, Brazil were analyzed. To better understand the distribution of the species in relation to environmental factors, the similarity and species-diversity indexes were calculated. Paguroideans were sampled monthly from January through December 2000. The trawls were made with two otter-trawl nets at 13 different sites, at depths of 2-40 m. Water temperature, salinity, sediment texture, and organic matter content were measured. Gastropod shells occupied by hermit crabs were also assessed. A total of 1,238 specimens was collected, belonging to the families Diogenidae and Paguridae, comprising seven genera and thirteen species. The most abundant hermit crab species were Dardanus insignis (761 specimens) and Loxopagurus loxochelis (351 specimens). Phimochirus holthuisi is newly reported from the São Paulo coast. The highest diversity index was found for the shallower sites near rocky shores. The results of the grouping analysis for sites and species indicated three distinct groups for sites, and four groups for species. This suggests that the occurrence of these anomurans is associated with the environmental and biotic factors analyzed.