Macrocrustaceans associated with reefs of Phragmatopoma caudata Krøyer in Mörch, 1863 (Polychaeta: Sabellariidae) and rocky shore in the Northeastern Brazil

Abstract Ecosystem engineering species create, modify, and/or maintain the characteristics of the environment. The polychaete Phragmatopoma caudata builds large sand reefs in the intertidal region of the Brazilian coast with high structural complexity, favoring the increase of diversity and interactions among the species associated. However, there are no studies concerning the association of polychaetes with crustacean macrofauna in the northeastern Brazil ecoregion, leaving an information gap on baseline biodiversity. Our aim was to analyze the effect of P. caudata colonies (PC) on the local diversity of macrocrustaceans compared to the rocky shore (RS) microhabitat. Monthly collections were carried out in low tide from September 2015 to August 2016 on 10 × 10 m quadrants for fauna and environmental variables (temperature and salinity) samples. In each microhabitat, the capture effort was two hours by two researchers. We collected 3,390 individuals, 60% associated with the colonies of PC and 40% with the RS. The PC obtained higher Shannon diversity, Pielou evenness and species richness coupled with milder water temperature and salinity conditions (minor air exposure during tide), compared to the RS that obtained greater species dominance and more extreme abiotic conditions (major air exposure). The Porcellanidae family stood out because all its species were highly abundant and had high occurrence in the colonies. The tropical Brazil porcelain crab Pachycheles greeleyi was dominant in both microhabitats (major dominance in PC). The structural complexity in the reefs of PC promoted higher availability of niches for the species, as more shelter for the resident species and refugium for temporary species with preference for more complex microhabitats. Conservation managers should prioritize the health of these colonies and subsequent species that constitute important ecosystemic and fishery resources.

Macrocrustaceans associated with reefs of Phragmatopoma caudata Krøyer in Mörch, 1863 (Polychaeta: Sabellariidae) and rocky shore in the Northeastern Brazil

Abstract Ecosystem engineering species create, modify, and/or maintain the characteristics of the environment. The polychaete Phragmatopoma caudata builds large sand reefs in the intertidal region of the Brazilian coast with high structural complexity, favoring the increase of diversity and interactions among the species associated. However, there are no studies concerning the association of polychaetes with crustacean macrofauna in the northeastern Brazil ecoregion, leaving an information gap on baseline biodiversity. Our aim was to analyze the effect of P. caudata colonies (PC) on the local diversity of macrocrustaceans compared to the rocky shore (RS) microhabitat. Monthly collections were carried out in low tide from September 2015 to August 2016 on 10 × 10 m quadrants for fauna and environmental variables (temperature and salinity) samples. In each microhabitat, the capture effort was two hours by two researchers. We collected 3,390 individuals, 60% associated with the colonies of PC and 40% with the RS. The PC obtained higher Shannon diversity, Pielou evenness and species richness coupled with milder water temperature and salinity conditions (minor air exposure during tide), compared to the RS that obtained greater species dominance and more extreme abiotic conditions (major air exposure). The Porcellanidae family stood out because all its species were highly abundant and had high occurrence in the colonies. The tropical Brazil porcelain crab Pachycheles greeleyi was dominant in both microhabitats (major dominance in PC). The structural complexity in the reefs of PC promoted higher availability of niches for the species, as more shelter for the resident species and refugium for temporary species with preference for more complex microhabitats. Conservation managers should prioritize the health of these colonies and subsequent species that constitute important ecosystemic and fishery resources.

Macrocrustaceans associated with reefs of Phragmatopoma caudata Krøyer in Mörch, 1863 (Polychaeta: Sabellariidae) and rocky shore in the Northeastern Brazil

Ecosystem engineering species create, modify, and/or maintain the characteristics of the environment. The polychaete Phragmatopoma caudata builds large sand reefs in the intertidal region of the Brazilian coast with high structural complexity, favoring the increase of diversity and interactions among the species associated. However, there are no studies concerning the association of polychaetes with crustacean macrofauna in the northeastern Brazil ecoregion, leaving an information gap on baseline biodiversity. Our aim was to analyze the effect of P. caudata colonies (PC) on the local diversity of macrocrustaceans compared to the rocky shore (RS) microhabitat. Monthly collections were carried out in low tide from September 2015 to August 2016 on 10 × 10 m quadrants for fauna and environmental variables (temperature and salinity) samples. In each microhabitat, the capture effort was two hours by two researchers. We collected 3,390 individuals, 60% associated with the colonies of PC and 40% with the RS. The PC obtained higher Shannon diversity, Pielou evenness and species richness coupled with milder water temperature and salinity conditions (minor air exposure during tide), compared to the RS that obtained greater species dominance and more extreme abiotic conditions (major air exposure). The Porcellanidae family stood out because all its species were highly abundant and had high occurrence in the colonies. The tropical Brazil porcelain crab Pachycheles greeleyi was dominant in both microhabitats (major dominance in PC). The structural complexity in the reefs of PC promoted higher availability of niches for the species, as more shelter for the resident species and refugium for temporary species with preference for more complex microhabitats. Conservation managers should prioritize the health of these colonies and subsequent species that constitute important ecosystemic and fishery resources.(AU)

Pisidia longimana ( ), a junior synonym of P. bluteli ( ) (Crustacea: Decapoda: Anomura: Porcellanidae) and a species distinct from P. longicornis ( )

Abstract Pisidia longimana (Risso, 1816) and P. bluteli (Risso, 1816), both described from Nice, France, have been considered each others synonyms or have been validated depending on successive taxonomic opinions. The validity of both in respect to P. longicornis (Linnaeus, 1767) has also been contradicted a number of times. The current lack of clarity in the use of the names P. longicornis, P. longimana and P. bluteli has resulted in nomenclatural instability, but also in unreliability and miscommunication as regards the available ecological and distributional information. The validity of P. bluteli and P. longimana is revisited herein based on a large number of specimens (241 males, 190 females and 33 juveniles) from many different localities. The latter species is confirmed as a junior synonym of the former, whereas P. bluteli and P. longicornis are herein considered separate species. Diagnostic characters and morphological variations are discussed and illustrated.

Pisidia longimana (Risso, 1816), a junior synonym of P. bluteli (Risso, 1816) (Crustacea: Decapoda: Anomura: Porcellanidae) and a species distinct from P. longicornis (Linnaeus, 1767)

Pisidia longimana (Risso, 1816) and P. bluteli (Risso, 1816), both described from Nice, France, have been considered each other’s synonyms or have been validated depending on successive taxonomic opinions. The validity of both in respect to P. longicornis (Linnaeus, 1767) has also been contradicted a number of times. The current lack of clarity in the use of the names P. longicornis, P. longimana and P. bluteli has resulted in nomenclatural instability, but also in unreliability and miscommunication as regards the available ecological and distributional information. The validity of P. bluteli and P. longimana is revisited herein based on a large number of specimens (241 males, 190 females and 33 juveniles) from many different localities. The latter species is confirmed as a junior synonym of the former, whereas P. bluteli and P. longicornis are herein considered separate species. Diagnostic characters and morphological variations are discussed and illustrated.

Pisidia longimana (Risso, 1816), a junior synonym of P. bluteli (Risso, 1816) (Crustacea: Decapoda: Anomura: Porcellanidae) and a species distinct from P. longicornis (Linnaeus, 1767)

Pisidia longimana (Risso, 1816) and P. bluteli (Risso, 1816), both described from Nice, France, have been considered each others synonyms or have been validated depending on successive taxonomic opinions. The validity of both in respect to P. longicornis (Linnaeus, 1767) has also been contradicted a number of times. The current lack of clarity in the use of the names P. longicornis, P. longimana and P. bluteli has resulted in nomenclatural instability, but also in unreliability and miscommunication as regards the available ecological and distributional information. The validity of P. bluteli and P. longimana is revisited herein based on a large number of specimens (241 males, 190 females and 33 juveniles) from many different localities. The latter species is confirmed as a junior synonym of the former, whereas P. bluteli and P. longicornis are herein considered separate species. Diagnostic characters and morphological variations are discussed and illustrated.(AU)

On the identity of Porcellana paivacarvalhoi Rodrigues da Costa, 1968 (Crustacea: Anomura: Porcellanidae)

Porcellana paivacarvalhoi Rodrigues da Costa, 1968, described from southeastern Brazil (São Sebastião, São Paulo), is morphologically similar to the eastern Atlantic P. platycheles (Pennant, 1777), and also to the west African P. africana Chace, 1956. The synonymy between P. paivacarvalhoi and P. platycheles was proposed by previous authors without examination of any specimens of P. platycheles, nor did they considered the morphological similarities between P. paivacarvalhoi and P. africana. This synonymy has been implicitly accepted without further analysis. The recent discovery of one male and two females syntypes of P. paivacarvalhoi has prompted new investigations into the status of P. paivacarvalhoi as a junior synonym either of P. platycheles or P. africana. Porcellana paivacarvalhoi is redescribed and illustrated based upon its lectotype herein designated. It is confirmed to be a junior synonym of P. platycheles; all evidence suggests that P. africana is a distinct, separate species. All previous records of P. platycheles from the Brazilian coast are reviewed. Its presence in southeastern Brazil in the late 1960’s most probably represents an historical case of human-mediated introduction by shipping activities. Porcellana platycheles did not manage to establish itself in São Sebastião, nor has it been recorded subsequently elsewhere in Brazil.

On the identity of Porcellana paivacarvalhoi Rodrigues da Costa, 1968 (Crustacea: Anomura: Porcellanidae)

Porcellana paivacarvalhoi Rodrigues da Costa, 1968, described from southeastern Brazil (São Sebastião, São Paulo), is morphologically similar to the eastern Atlantic P. platycheles (Pennant, 1777), and also to the west African P. africana Chace, 1956. The synonymy between P. paivacarvalhoi and P. platycheles was proposed by previous authors without examination of any specimens of P. platycheles, nor did they considered the morphological similarities between P. paivacarvalhoi and P. africana. This synonymy has been implicitly accepted without further analysis. The recent discovery of one male and two females syntypes of P. paivacarvalhoi has prompted new investigations into the status of P. paivacarvalhoi as a junior synonym either of P. platycheles or P. africana. Porcellana paivacarvalhoi is redescribed and illustrated based upon its lectotype herein designated. It is confirmed to be a junior synonym of P. platycheles; all evidence suggests that P. africana is a distinct, separate species. All previous records of P. platycheles from the Brazilian coast are reviewed. Its presence in southeastern Brazil in the late 1960s most probably represents an historical case of human-mediated introduction by shipping activities. Porcellana platycheles did not manage to establish itself in São Sebastião, nor has it been recorded subsequently elsewhere in Brazil.(AU)

Chaetotaxy and setal diversity of grooming legs in species of porcelain crabs (Crustacea: Anomura: Porcellanidae)

The morphology of the fifth pereiopods was studied under scanning electron microscopy in ten species of porcelain crabs for chaetotaxy and setal diversity, namelly Megalobrachium pacificum, Megalobrachium roseum, Pachycheles grossimanus, Petrolisthes armatus, Petrolisthes tuberculatus, Pisidia brasiliensis, Pisidia longicornis, Polyonyx gibbesi, Porcellana platycheles and Porcellana sayana. Six setal types were identified: simple, pappose, sickle-shaped serrate, straight serrate, club-shaped and tooth-like cuspidate. Porcelain crabs can differ in the fifth pereiopod setal morphology, chaetotaxy and setal density, even among species within the same genus. The absence of sexual dimorphism of the grooming legs in porcelain crabs suggests that grooming eggs requires no particular grooming apparatus in females and that male and female are equal in grooming efficiency.

Chaetotaxy and setal diversity of grooming legs in species of porcelain crabs (Crustacea: Anomura: Porcellanidae)

The morphology of the fifth pereiopods was studied under scanning electron microscopy in ten species of porcelain crabs for chaetotaxy and setal diversity, namelly Megalobrachium pacificum, Megalobrachium roseum, Pachycheles grossimanus, Petrolisthes armatus, Petrolisthes tuberculatus, Pisidia brasiliensis, Pisidia longicornis, Polyonyx gibbesi, Porcellana platycheles and Porcellana sayana. Six setal types were identified: simple, pappose, sickle-shaped serrate, straight serrate, club-shaped and tooth-like cuspidate. Porcelain crabs can differ in the fifth pereiopod setal morphology, chaetotaxy and setal density, even among species within the same genus. The absence of sexual dimorphism of the grooming legs in porcelain crabs suggests that grooming eggs requires no particular grooming apparatus in females and that male and female are equal in grooming efficiency.(AU)

Hábitat y historia de vida de Pachychelles laevidactylus (Crustacea, Anomura, Porcellanidae) en el intermareal rocoso de Mar del Plata, Argentina

RESUMEN Se analizó la estructura del hábitat, el crecimiento relativo, la heteroquelia, la estructura poblacional, la madurez morfométrica y la fecundidad de Pachycheles laevidactylus Ortmann, 1892, durante Abril, Mayo y Junio del 2005, en sustratos naturales de Mar del Plata, Argentina (38°02'S, 57°31'W). La cobertura algal, la exposición al oleaje y el tiempo de inmersión varió de manera significativa entre estratos y entre meses. La morfometría de las quelas no manifestó diferencias entre sexos. Las hembras mostraron el abdomen más grande que los machos. Si bien existió la heteroquelia, no se detectó lateralidad. Las tallas (ancho de caparazón) estuvieron comprendidas entre 0.67 mm a 9.3 mm para los machos, y entre 1.07 mm a 10.65 mm, para las hembras. La densidad de organismos varió entre meses y entre los estratos. La estructura poblacional, para el periodo de estudio, se explicó en función de la estructura del hábitat. La madurez morfométrica de las hembras fue registrada a los 4.38 mm de ancho de caparazón. El número de huevos se correlacionó positivamente con la talla de las hembras ovígeras con un mínimo de 2 huevos y un máximo de 160, pero esa relación dependió del avance en el desarrollo de los embriones.

Abstract Habitat structure, relative growth, the heterochely, population structure, morphometric maturity and fecundity of Pachycheles laevidactylus Ortmann, 1892 were analyzed during April, May and June 2005, in natural substrates of Mar del Plata, Argentina (38°02'S, 57°31'W). Algal cover, wave exposure and immersion time varied significantly between strata and between months. Chelae morphometry showed no differences between sexes. Females showed the greatest abdomen. While there was the heteroquelia, laterality not detected. The size (carapace width) ranged from 0.67 mm to 9.3 mm for males and from 1.07 mm to 10.65 mm, for females. The density of organisms varied among months and between strata. The population structure for the period of study explained depended on habitat structure. The morphometric maturity of females was recorded at the 4.38 mm of carapace width. The number of eggs was positively correlated with the size of the gravid females with a minimum of 2 eggs and a maximum of 160, but that relationship depended on progress in the development of embryos.

Hábitat y historia de vida de Pachychelles laevidactylus (Crustacea, Anomura, Porcellanidae) en el intermareal rocoso de Mar del Plata, Argentina / Habitat and life's story of Pachychelles laevidactylus (Crustacea, Anomura, Porcellanidae) in the rocky shore of Mar del Plata, Argentina

Se analizó la estructura del hábitat, el crecimiento relativo, la heteroquelia, la estructura poblacional, la madurez morfométrica y la fecundidad de Pachycheles laevidactylus Ortmann, 1892, durante Abril, Mayo y Junio del 2005, en sustratos naturales de Mar del Plata, Argentina (38°02'S, 57°31'W). La cobertura algal, la exposición al oleaje y el tiempo de inmersión varió de manera significativa entre estratos y entre meses. La morfometría de las quelas no manifestó diferencias entre sexos. Las hembras mostraron el abdomen más grande que los machos. Si bien existió la heteroquelia, no se detectó lateralidad. Las tallas (ancho de caparazón) estuvieron comprendidas entre 0.67 mm a 9.3 mm para los machos, y entre 1.07 mm a 10.65 mm, para las hembras. La densidad de organismos varió entre meses y entre los estratos. La estructura poblacional, para el periodo de estudio, se explicó en función de la estructura del hábitat. La madurez morfométrica de las hembras fue registrada a los 4.38 mm de ancho de caparazón. El número de huevos se correlacionó positivamente con la talla de las hembras ovígeras con un mínimo de 2 huevos y un máximo de 160, pero esa relación dependió del avance en el desarrollo de los embriones.(AU)

Habitat structure, relative growth, the heterochely, population structure, morphometric maturity and fecundity of Pachycheles laevidactylus Ortmann, 1892 were analyzed during April, May and June 2005, in natural substrates of Mar del Plata, Argentina (38°02'S, 57°31'W). Algal cover, wave exposure and immersion time varied significantly between strata and between months. Chelae morphometry showed no differences between sexes. Females showed the greatest abdomen. While there was the heteroquelia, laterality not detected. The size (carapace width) ranged from 0.67 mm to 9.3 mm for males and from 1.07 mm to 10.65 mm, for females. The density of organisms varied among months and between strata. The population structure for the period of study explained depended on habitat structure. The morphometric maturity of females was recorded at the 4.38 mm of carapace width. The number of eggs was positively correlated with the size of the gravid females with a minimum of 2 eggs and a maximum of 160, but that relationship depended on progress in the development of embryos.(AU)

Hábitat y historia de vida de Pachychelles laevidactylus (Crustacea, Anomura, Porcellanidae) en el intermareal rocoso de Mar del Plata, Argentina / Habitat and life's story of Pachychelles laevidactylus (Crustacea, Anomura, Porcellanidae) in the rocky shore of Mar del Plata, Argentina

Se analizó la estructura del hábitat, el crecimiento relativo, la heteroquelia, la estructura poblacional, la madurez morfométrica y la fecundidad de Pachycheles laevidactylus Ortmann, 1892, durante Abril, Mayo y Junio del 2005, en sustratos naturales de Mar del Plata, Argentina (38°02'S, 57°31'W). La cobertura algal, la exposición al oleaje y el tiempo de inmersión varió de manera significativa entre estratos y entre meses. La morfometría de las quelas no manifestó diferencias entre sexos. Las hembras mostraron el abdomen más grande que los machos. Si bien existió la heteroquelia, no se detectó lateralidad. Las tallas (ancho de caparazón) estuvieron comprendidas entre 0.67 mm a 9.3 mm para los machos, y entre 1.07 mm a 10.65 mm, para las hembras. La densidad de organismos varió entre meses y entre los estratos. La estructura poblacional, para el periodo de estudio, se explicó en función de la estructura del hábitat. La madurez morfométrica de las hembras fue registrada a los 4.38 mm de ancho de caparazón. El número de huevos se correlacionó positivamente con la talla de las hembras ovígeras con un mínimo de 2 huevos y un máximo de 160, pero esa relación dependió del avance en el desarrollo de los embriones.

Habitat structure, relative growth, the heterochely, population structure, morphometric maturity and fecundity of Pachycheles laevidactylus Ortmann, 1892 were analyzed during April, May and June 2005, in natural substrates of Mar del Plata, Argentina (38°02'S, 57°31'W). Algal cover, wave exposure and immersion time varied significantly between strata and between months. Chelae morphometry showed no differences between sexes. Females showed the greatest abdomen. While there was the heteroquelia, laterality not detected. The size (carapace width) ranged from 0.67 mm to 9.3 mm for males and from 1.07 mm to 10.65 mm, for females. The density of organisms varied among months and between strata. The population structure for the period of study explained depended on habitat structure. The morphometric maturity of females was recorded at the 4.38 mm of carapace width. The number of eggs was positively correlated with the size of the gravid females with a minimum of 2 eggs and a maximum of 160, but that relationship depended on progress in the development of embryos.

Sistema reprodutor masculino em Porcellanidae: reconstrução do caráter ancestral com base na morfologia, histoquímica e ultraestrutura dos espermatóforos e espermatozoides.

Os caranguejos da família Porcellanidae tem seu sistema reprodutor masculino desconhecido do ponto de vista histológico e pouco estudado no aspecto ultraestrutural dos espermatozoides. Nesta dissertação, apresentamos o padrão anato-histológico do sistema reprodutor masculino, além da ultraestrutura dos espermatóforos e espermatozoides de Porcellanidae, buscando entender as relações de parentesco e a evolução do sistema reprodutor e suas células germinativas entre as superfamílias da infraordem Anomura. Foram coletados com auxílio de um barco de pesca camaroeira, além de coleta manual na região entre marés e por meio de mergulho livre e autônomo (3m), machos adultos dos gêneros Pachycheles, Petrolisthes, Pisidia, Porcellana e Megalobrachium em Ubatuba e São Vicente (SP), Brasil, e em Valdivia, Chile. As amostras foram processadas seguindo rotinas para histologia, microscopia eletrônica de trasmissão e varredura. Análises do fluido seminal, empacotamento dos espermatozoides e formação dos espermatóforos, apresentaram o mesmo padrão para todas as espécies estudadas, e semelhante a outros Anomuras já descritos na literatura, sendo simplificado e sem diferenciações macroscopícas. Os espermatóforos de Porcellanidae seguem o padrão tripartido pedunculado como descrito para a maioria dos Anomura, porém se diferenciam por apresentarem um pedúnculo diminuto, característica compartilhada com Galatheidae e Munidae. A ultraestrutura dos espermatozoides dos Porcellanidae é caracterizada por uma vesícula acrossomal saliente ao espermatozoide, possuindo camadas concêntricas e diferente dos outros Galatheoidea por apresentarem: opérculo centralmente perfurado, cone perforatorial eletrondenso, e anel perforatorial posterior. Porcellanidae possuem caracteristicas compartilhadas entre as espécies mais proximamente relacionadas e algumas exclusivas de cada gênero. Pisidia brasiliensis e Porcellana sayana, possuem um ombro na região apical do acrossoma. Porcellana sayana e Megalobrachium roseum o cone perforatorial se estende até a abertura opercular. No gênero Petrolisthes, ocorre uma estrutura excusiva, o anel tubular eletrondenso. Pisidia brasiliensis possui botão apical. Assim, a ultraestrutura do espermatozoide destas espécies de Porcellanidae representam um novo conjunto de caracteres os quais são claramente relacionados filogeneticamente, quando traçados por meio do uso da filogenia mais recente da infraordem.

The crabs of the Porcellanidae family have their male reproductive system unknown from the historical point of view and little studied in the ultrastructural aspect of sperm. In this dissertation, we present the histological pattern of the male reproductive system, in addition to the ultrastructure of sperm and sperm from porcellanids, seeking to understand how kinship relationships and the evolution of the reproductive system and its germ cells between the superfamilies of Anomura's infrastructure. Were collected with the aid of a camaro fishing boat, in addition to manual collection in the region between tides and free and autonomous diving medium (3m), adult males of the genera Pachycheles, petrolitic, pisis, porcellana and megalobrachium in Ubatuba and São Vicente (SP ), Brazil, and in Valdivia, Chile. As the samples were processed following routines for histology, transmission electron microscopy and scanning. Analysis of seminal fluid, sperm packaging and sperm formation, analysis or the same pattern for all studied species, and similar to other Anomuras already applied in the literature, being simplified and without macroscopic differences. Porcellanid spermatozoa follow the pedunculated tripartite pattern as described for most Anomura, but they differ by presenting a tiny peduncle, resources shared with Galatheidae and Munidae. An ultrastructure of porcelain spermatozoa is characterized by an acrosomal vesicle protruding from the sperm, having concentric layers and different from the other Galatheoidea in that they have: centrally perforated operculum, perforated electron- condensed cone and perforated posterior ring. Porcellanidae has characteristics shared between the closest species and some exclusive to each genus. Pisidia brasiliensis and Porcellana sayana, have a "shoulder" in the apical region of the acrosome. Porcellana sayana and Megalobrachium roseum or perforated cone extend to the opercular opening. In the Petrolisthes genus, there is an excusive structure or an electronic tubular ring. Pisidia brasiliensis has an apical bud. Thus, an ultrastructure of the sperm of these species of Porcellanidae represents a new set of characters, which are clearly related phylogenetically, when traced using the most recent phylogeny of the infraorder.

Intersexuality in the porcellanid crab Pisidia longicornis (Crustacea: Decapoda: Anomura: Porcellanidae)

Intersex specimens of Pisidia longicornis (Linnaeus, 1767) are recorded for the first time and their secondary sexual characters are analyzed. Of 145 specimens of P. longicornis examined in this study, six were identified as intersex individuals. Of these, three presented male secondary sexual characters, well-developed male gonopores and rudimentary female gonopores, whereas the other three had female secondary sexual characters, with female gonopores being more pronounced than the male ones. The present study provides the first record of intersex porcelain crabs. (AU)

Intersexuality in the porcellanid crab Pisidia longicornis (Crustacea: Decapoda: Anomura: Porcellanidae)

Intersex specimens of Pisidia longicornis (Linnaeus, 1767) are recorded for the first time and their secondary sexual characters are analyzed. Of 145 specimens of P. longicornis examined in this study, six were identified as intersex individuals. Of these, three presented male secondary sexual characters, well-developed male gonopores and rudimentary female gonopores, whereas the other three had female secondary sexual characters, with female gonopores being more pronounced than the male ones. The present study provides the first record of intersex porcelain crabs.

Larval and adult density of the porcellanid crab Petrolisthes armatus (Anomura: Porcellanidae) in an Amazon estuary, northern Brazil

Petrolisthes armatus (Gibbes, 1850) is a porcellanid crab with a wide geographical distribution. In the present study we analyzed variations in the abundance of P. armatus adults and larvae over an annual cycle in the Marapanim estuary of the Amazon coastal zone, in the northeastern portion of the state of Pará, Brazil. Particularly, we focused on the presence of ovigerous females and timing of larval release, with the aim of elucidating reproductive patterns in a tropical estuarine system. The mean density of P. armatus larvae (zoea I and II) correlated positively with the salinity of the shallow waters of the estuary, whereas the abundance of adults correlated with the salinity registered in water samples collected from the benthic environment. There was also a significant positive correlation between larval density (zoea I and II) and water temperature. Ovigerous females were captured throughout the study period, from August 2006 to July 2007, but were more abundant in June and less abundant during the rainy months, between February and May. Larvae were only present during the dry season and transition months (June to January), and were absent during the rainy season (February to May). Petrolisthes armatus reproduces throughout the year in the Marapanim estuary and all developmental stages of this species (zoeal stages I and II, megalopae and adults) are found in the estuary. The results indicate that the study area is an important environment for the reproduction of this decapod.

Larval and adult density of the porcellanid crab Petrolisthes armatus (Anomura: Porcellanidae) in an Amazon estuary, northern Brazil

Petrolisthes armatus (Gibbes, 1850) is a porcellanid crab with a wide geographical distribution. In the present study we analyzed variations in the abundance of P. armatus adults and larvae over an annual cycle in the Marapanim estuary of the Amazon coastal zone, in the northeastern portion of the state of Pará, Brazil. Particularly, we focused on the presence of ovigerous females and timing of larval release, with the aim of elucidating reproductive patterns in a tropical estuarine system. The mean density of P. armatus larvae (zoea I and II) correlated positively with the salinity of the shallow waters of the estuary, whereas the abundance of adults correlated with the salinity registered in water samples collected from the benthic environment. There was also a significant positive correlation between larval density (zoea I and II) and water temperature. Ovigerous females were captured throughout the study period, from August 2006 to July 2007, but were more abundant in June and less abundant during the rainy months, between February and May. Larvae were only present during the dry season and transition months (June to January), and were absent during the rainy season (February to May). Petrolisthes armatus reproduces throughout the year in the Marapanim estuary and all developmental stages of this species (zoeal stages I and II, megalopae and adults) are found in the estuary. The results indicate that the study area is an important environment for the reproduction of this decapod.

Larval and adult density of the porcellanid crab Petrolisthes armatus (Anomura: Porcellanidae) in an Amazon estuary, northern Brazil

Petrolisthes armatus (Gibbes, 1850) is a porcellanid crab with a wide geographical distribution. In the present study we analyzed variations in the abundance of P. armatus adults and larvae over an annual cycle in the Marapanim estuary of the Amazon coastal zone, in the northeastern portion of the state of Pará, Brazil. Particularly, we focused on the presence of ovigerous females and timing of larval release, with the aim of elucidating reproductive patterns in a tropical estuarine system. The mean density of P. armatus larvae (zoea I and II) correlated positively with the salinity of the shallow waters of the estuary, whereas the abundance of adults correlated with the salinity registered in water samples collected from the benthic environment. There was also a significant positive correlation between larval density (zoea I and II) and water temperature. Ovigerous females were captured throughout the study period, from August 2006 to July 2007, but were more abundant in June and less abundant during the rainy months, between February and May. Larvae were only present during the dry season and transition months (June to January), and were absent during the rainy season (February to May). Petrolisthes armatus reproduces throughout the year in the Marapanim estuary and all developmental stages of this species (zoeal stages I and II, megalopae and adults) are found in the estuary. The results indicate that the study area is an important environment for the reproduction of this decapod.

Larval and adult density of the porcellanid crab Petrolisthes armatus (Anomura: Porcellanidae) in an Amazon estuary, northern Brazil

Petrolisthes armatus (Gibbes, 1850) is a porcellanid crab with a wide geographical distribution. In the present study we analyzed variations in the abundance of P. armatus adults and larvae over an annual cycle in the Marapanim estuary of the Amazon coastal zone, in the northeastern portion of the state of Pará, Brazil. Particularly, we focused on the presence of ovigerous females and timing of larval release, with the aim of elucidating reproductive patterns in a tropical estuarine system. The mean density of P. armatus larvae (zoea I and II) correlated positively with the salinity of the shallow waters of the estuary, whereas the abundance of adults correlated with the salinity registered in water samples collected from the benthic environment. There was also a significant positive correlation between larval density (zoea I and II) and water temperature. Ovigerous females were captured throughout the study period, from August 2006 to July 2007, but were more abundant in June and less abundant during the rainy months, between February and May. Larvae were only present during the dry season and transition months (June to January), and were absent during the rainy season (February to May). Petrolisthes armatus reproduces throughout the year in the Marapanim estuary and all developmental stages of this species (zoeal stages I and II, megalopae and adults) are found in the estuary. The results indicate that the study area is an important environment for the reproduction of this decapod.