Male reproductive system of the freshwater prawn Macrobrachium brasiliense (Decapoda: Palaemonidae): Notes on spermatophore formation and sperm count.

Studies on the morphology of the reproductive system are essential for understanding the reproduction processes of species or even within genera or families. The present study aimed to describe the functional morphology of the male reproductive system, spermatophore formation, and sperm count of Macrobrachium brasiliense. The anatomy of the reproductive system consists of a pair of testes from which the vasa deferentia (VD) starts, extending to the fifth pair of pereopods. The VD is divided into three regions: proximal (PVD), middle (MVD), and distal (DVD). In the PVD, there is a prominent fold, the typhlosole, formed by columnar cells. The typhlosole disappears in the MVD, being incorporated into one of the faces of the VD wall, identified by its simple columnar epithelium while the remainder of the vessel wall is formed by squamous or simple cubic epithelium. Columnar cells produce type-II and III secretion. The epithelium in the DVD is made up only of cubic cells. Low sperm concentration was observed when compared to other species of the genus Macrobrachium. In conclusion, the typhlosole and columnar epithelium are responsible for the asymmetric spermatophore, which seems the pattern of Macrobrachium that is probably shared with other caridean shrimps.

Life History Traits of the Snapping Shrimp Alpheus carlae (Decapoda: Alpheidae) from the South-eastern Coast of Brazil.

The life history traits of the snapping shrimp Alpheus carlae were investigated. We evaluated the relative growth, morphological sexual maturity, sexual dimorphism, handedness, fecundity, and egg volume. Sampling was performed monthly in an estuarine region of Cananéia, in the south-eastern coast of Brazil. The following structures were measured to perform the relative growth analysis: length of carapace and second pleonal pleuron, length, width and height of major cheliped propodus, and length of appendix interna and appendix masculina. Juveniles and adults males and females have different growth patterns, indicating distinct strategies of energy allocation. The estimated carapace length at the onset of morphological sexual maturity of females and males was 5.6 mm and 6.2 mm, respectively. The sexual dimorphism of specific body structures was evident, mainly in the appendix interna of females and major cheliped of males. There was no evidence of handedness in females or males. The mean fecundity of females was 364 ± 204 eggs (mean ± SD) eggs and was positively correlated with the carapace length. The egg volume differed between developmental stages I and II, and I and III, with an overall volume increase of 73.23% and 95.45%, respectively. The results contribute to the knowledge about A. carlae and its life history in natural environment.

Checklist of decapod crustaceans from the coast of the São Paulo state (Brazil) supported by integrative molecular and morphological data: III. Infraorder Brachyura Latreille, 1802.

This checklist is the third part of a series derived from a long-term multidisciplinary project on the biodiversity of decapod crustaceans from marine and coastal environments of São Paulo state (Brazil). We integrated molecular techniques (DNA markers) and morphological analyses of adult specimens for accurate identifications. We compilated 185 species from the literature, but we confirmed the presence of 168 species: 130 of which we sampled, analyzed and obtained sequences (COI and/or 16S totalizing 113 sequences) and 38 that were not directly collected but were confirmed by analyses. In addition, 27 had their distribution removed from São Paulo due to uncertainties, and absence of material as voucher. Five species were reported for the first time on the coast of São Paulo (Calappa ocellata, Neohelice granulata, Teleophrys pococki, Teramnonotus monodi, Tetraxanthus rathbunae) and one on the Brazilian coast (Pseudomedaeus agassizi). Most of the non-sampled species previously reported on the coast of São Paulo might be considered doubtful records stablished in the past by inaccurate analyses, which assumed extended distribution to the area and/or misidentifications. At this time and based on our refined compilation, we can estimate the brachyuran diversity on the coast of São Paulo in 168 species. This detailed inventory contributes to the knowledge on the local decapod fauna by checking available dataset, adding new species records in São Paulo and new sequences to GenBank database. These data may serve as baseline for future identifications and studies on conservation, population genetics, biogeography and phylogenetics, which might flag species that deserve further investigations and concerns.

Validation of Xiphopenaeus dincao Carvalho-Batista, Terossi, Zara, Mantelatto amp; Costa and Xiphopenaeus baueri Carvalho-Batista, Terossi, Zara, Mantelatto amp; Costa (Decapoda: Penaeidae) from western Atlantic.

Two species of Penaeidae shrimp from western Atlantic were described by Carvalho-Batista et al. (2019) as Xiphopenaeus dincao and Xiphopenaeus baueri. The descriptions and figures presented by Carvalho-Batista et al. (2019) and the posterior author's correction (Carvalho-Batista et al., 2020) characterize the species. However, the journal issue in which the description was published is online only, and the article with the new names did not include a Zoobank registration number (LSID), required for validation of new names in electronic-only publications (ICZN 2012). Thus, the names Xiphopenaeus dincao and Xiphopenaeus baueri, as published in Scientific Reports 9: 15281, is not available according the International Code of Zoological Nomenclature (ICZN 1999, 2012). Therefore, the present note serves to validate the names Xiphopenaeus dincao and Xiphopenaeus baueri by fulfilling ICZN conditions for nomenclatural availability. Type specimens of both species are deposited in Museu de Zoologia da Universidade de São Paulo (MZUSP), São Paulo, Brazil.

Description of the first zoeal stage of Synalpheus apioceros Coutière, 1909 (Caridea: Alpheidae), including a comparative analysis with larval morphology from the genus Synalpheus Spence Bate, 1888.

Herein, the Zoea I of Synalpheus apioceros is described, followed by a comparative analysis of the first larval stage from Synalpheus. Larvae were obtained from two ovigerous females sampled in Ubatuba, São Paulo. The morphology of the Zoea I of S. apioceros was compared to five other Synalpheus species whose structures were previously described: S. biunguiculatus, S. minus, S. neomeris, S. pectiniger, and S. tumidomanus. A set of unique morphological characteristics was found for S. apioceros: exopod of antennule with 4 aesthetascs and 1 plumose seta; coxal endite of maxillule with 2 simple setae plus 2 plumose setae; basial endite of maxillule with 2 simple setae plus 2 short spines; endopod of maxillule with 1 plumose seta plus 2 simple setae; basial endite of maxilla bilobed with 2 (1 plumose and 1 simple) + 2 (1 plumose and 1 simple) setae; coxal endite of maxilla bilobed with 2 + 1 plumose setae; endopod of maxilla with 3 (1 plumose and 2 simple) terminal setae + 2 simple setae; endopod of first maxilliped unsegmented with 3 terminal simple setae; basis of first maxilliped with 5 spines; endopod of second maxilliped 5-segmented with 0, 0, 0, 1 plumodenticulate, 4 (1 serrate + 3 simple) setae; endopod of third maxilliped 5-segmented with 0, 0, 0, 1 simple, 3 simple setae; pereiopods 1st to 3rd birramous and 4th and 5th uniramous. S. apioceros shows higher morphological similarity with S. minus, followed by S. tumidomanus, S. neomeris and S. pectiniger, probably since these species present extended larval development. Besides the similar morphology among species, the unique attributes presented here are important for the genus' taxonomy, being fundamental for identifying the first larval stages of Synalpheus, as well as for subsidizing information for species identification keys.

New record of the sponge-dwelling shrimp Typton distinctus Chace, 1972 (Decapoda: Caridea: Palaemonidae) in São Paulo State, Brazil.

Among the Caridea Infraorder, the palaemonid shrimp from the genus Typton Costa, 1844 are commonly found in association with sponges, frequently feeding on the tissues of their hosts ((Duris et al. 2011; Almeida et al. 2014; Pachelle et al. 2015; Soledade et al. 2017). Typton is mostly characterized by morphological features related to their sponge-dwelling lifestyle, as a simple and compressed rostrum, carapace smooth and antennal spines present, antennae extremely reduced and scaphocerite rudimentary. Mandible without palp, incisor process normal, reduced or absent. Second legs unequal, asymmetrical, without molar process on major chela (Bruce, 1972).

Author Correction: A multigene and morphological analysis expands the diversity of the seabod shrimp Xiphopenaeus Smith, 1869 (Decapoda: Penaeidae), with descriptions of two new species.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A multigene and morphological analysis expands the diversity of the seabod shrimp Xiphopenaeus Smith, 1869 (Decapoda: Penaeidae), with descriptions of two new species.

After being stable for nearly a century, the taxonomic history of the genus Xiphopenaeus has been marked by many changes in the last three decades. The taxonomic status of the Atlantic species has a low resolution, and many species are still undefined and grouped as cryptic species. Here we employed an integrative approach to define the species of Xiphopenaeus and the morphological characters needed to differentiate them. We combined the analyses of two molecular markers (COI and 16 S rDNA), scanning electron microscopy and light microscopy. Based on specimens from 17 localities from the Atlantic and Pacific oceans, we detected five divergent genetic groups, three in the Atlantic (A1, A2, A3) and two in the Pacific (P1, P2). Male secondary sexual characters were able to differentiate four out of the five genetic groups. Group A1 corresponds to X. kroyeri, and A2 and A3 correspond to new species. We redescribed the genus and two new species are described and illustrated: Xiphopenaeus dincao nov. sp. (A2) and Xiphopenaeus baueri nov. sp. (A3). Since the holotype of X. riveti was missing and the specimen analysed from group P2 was a female, the status of the species of Xiphopenaeus from the Pacific remains unresolved.

Checklist of decapods (Crustacea) from the coast of São Paulo State (Brazil) supported by integrative molecular and morphological data: II. Infraorder Caridea: family Alpheidae.

This study is part of a series of checklists resulting from a long-term multidisciplinary project on the biodiversity of decapod crustaceans from the marine and coastal environments (including estuaries) of São Paulo State (Brazil). For that, we integrated molecular techniques (mitochondrial DNA markers) and morphological analyses of adult specimens for an accurate and detailed identification. The DNA markers were used when the morphological identification was doubtful, particularly in the recognition of cryptic species. This second manuscript presents a checklist of the Alpheidae caridean shrimps from the coast of São Paulo. We report the occurrence of Alpheus cf. paracrinitus and Synalpheus townsendi for the first time in the region. Based on our survey, 39 species of Alpheidae are known for this region: Alpheus (21 spp.), Athanas (2 spp.), Automate (2 spp.), Leptalpheus (1 spp.), Salmoneus (3 spp.), and Synalpheus (10 spp.). We collected 28 species and obtained cytochrome oxidase subunit I (barcode region) and/or 16S partial sequences of 26 of them. These sequences may be used for phylogenetic and populational analyses in further studies.

The influence of environmental variables in the reproductive performance of Macrobrachium amazonicum (Heller, 1862) (Caridea: Palaemonidae) females in a continental population.

Macrobrachium amazonicum is a commercially important freshwater prawn with a high degree of reproductive plasticity. The species is classified into two groups: coastal populations, with larger individuals exhibiting high fecundity and needing brackish water for larval development; and continental populations, with smaller specimens exhibiting low fecundities and completing metamorphosis in freshwater. The objective of this study was to investigate the influence of environmental factors in the fecundity, egg size and volume, and reproductive output in females of M. amazonicum from a continental population during a two-year period. We also compared our results with those obtained for other continental and coastal populations. Reproductive parameters differed markedly between continental and coastal populations in most cases. The continental population studied here, however, exhibited reproductive characteristics similar to those of coastal populations. The present study found a correlation between the reproductive parameters and the environmental variables analyzed. This result corroborates the hypothesis that wide variation in reproductive parameters in the geographical distribution of M. amazonicum is related to the environmental characteristics in which populations are inserted. We suggest that further studies could investigate the potential of continental populations for aquaculture, which could significantly reduce production costs.

Checklist of decapods (Crustacea) from the coast of the São Paulo state (Brazil) supported by integrative molecular and morphological data: I. Infraorder Caridea: families Hippolytidae, Lysmatidae, Ogyrididae, Processidae and Thoridae.

The current checklist is the result of a long-term multidisciplinary project which combined molecular techniques (mitochondrial DNA markers) and morphological analyses of adult specimens for an accurate and detailed identification of the total biodiversity of decapod crustaceans from marine and coastal (including estuaries) environments of São Paulo State (Brazil). This is the first of a series of reports and providing a checklist of caridean shrimps of the families Hippolytidae (5 spp.), Lysmatidae (6 spp.), Ogyrididae (2 spp.), Processidae (5 spp.) and Thoridae (1 sp.). We collected material of 13 species out of 19 recorded, with sequences of cytochrome oxidase subunit I - barcode region and 16S generated from 10 species. The previous record of Lysmata cf. intermedia for São Paulo is actually L. jundalini, as the first record in São Paulo/South Atlantic waters. The molecular data were helpful to confirm the identification of some species, as the occurrence of L. wurdemanni which is confirmed in the South Atlantic Ocean based on morphological, color pattern and molecular data.

Morphological description of early zoeal stages of Alpheus brasileiro Anker, 2012 reared in the laboratory, including a revision of the larval morphology of the first zoeal stage of the genus Alpheus Fabricius, 1798 (Caridea: Alpheidae).

The aims of this study were to describe and illustrate the early larval stages of the snapping shrimp Alpheus brasileiro Anker, 2012 and to review the larval morphology (Zoea I) of the genus Alpheus Fabricius, 1798. Larvae were obtained from two females with embryos collected in an intertidal estuary in Cananéia, São Paulo state, Brazil. The published descriptions of all available Alpheus zoea I (only 14 spp., 4.7%) were used for a comparison of larval morphology. The zoeae I of Alpheus species are very similar, but A. brasileiro can be separated from the other studied species by the following set of characteristics: antennal scale 5-segmented with 11 plumose setae and 2 simple setae; coxal endite of maxillule with 3 simple setae and 2 sparsely plumose setae; and coxal endite of maxilla with 1 simple seta and 1 sparsely plumose seta. The description of early larval development of A. brasileiro indicates consistent characteristics, which, when analyzed along with other morphological, molecular, and biogeographical aspects, can help to elucidate the complex phylogenetic relationships between the species in the group. However, this is the first zoeal description of a member of Alpheus       armillatus species complex and it therefore does not permit a comparison. Only with new additional descriptions we will be able to generate a clearer scenario in this field due the high similarity among the species.

Population dynamics of Rimapenaeus constrictus (Stimpson, 1874) (Penaeoidea) on the southeastern Brazilian coast: implications for shrimp fishing management from a 5-year study on a bycatch species

ABSTRACT This is the first study to evaluate in broad spatiotemporal scales the growth parameters and population structure of Rimapenaeus constrictus, a barely damaged species composing the bycatch from shrimp fishing in the Western Atlantic. The abundance and size-class frequency distribution, growth, longevity and sex ratio were evaluated from monthly samples obtained in the northern littoral of São Paulo state from Jan/1998 to Jun/2003. We measured 5,812 individuals in which the sex ratio was skewed toward females; this was more evident in size classes greater than 10 mm in CL (carapace length) (binomial test, p<0.05). We selected 16 growth cohorts of females, and 8 of males, the majority consisting of younger individual cohorts excluded from the fisheries closure period. Growth estimates resulted in a CL∞ of 17.42 mm, a growth coefficient of 0.008 and a longevity of 579 days (1.60 year) for females, as well as a CL∞ of 16.3 mm, a growth coefficient of 0.01 and a longevity of 425 days (1.17 year) for males. Our results provide information of incontestable relevance to our knowledge of fishing management. We therefore strongly recommend that the fisheries closure period be changed to protect this species' recruitment period and consequently its adult individuals.

Population dynamics of Rimapenaeus constrictus (Stimpson, 1874) (Penaeoidea) on the southeastern Brazilian coast: implications for shrimp fishing management from a 5-year study on a bycatch species.

This is the first study to evaluate in broad spatiotemporal scales the growth parameters and population structure of Rimapenaeus constrictus, a barely damaged species composing the bycatch from shrimp fishing in the Western Atlantic. The abundance and size-class frequency distribution, growth, longevity and sex ratio were evaluated from monthly samples obtained in the northern littoral of São Paulo state from Jan/1998 to Jun/2003. We measured 5,812 individuals in which the sex ratio was skewed toward females; this was more evident in size classes greater than 10 mm in CL (carapace length) (binomial test, p<0.05). We selected 16 growth cohorts of females, and 8 of males, the majority consisting of younger individual cohorts excluded from the fisheries closure period. Growth estimates resulted in a CL∞ of 17.42 mm, a growth coefficient of 0.008 and a longevity of 579 days (1.60 year) for females, as well as a CL∞ of 16.3 mm, a growth coefficient of 0.01 and a longevity of 425 days (1.17 year) for males. Our results provide information of incontestable relevance to our knowledge of fishing management. We therefore strongly recommend that the fisheries closure period be changed to protect this species' recruitment period and consequently its adult individuals.

Sperm ultrastructure of shrimps from the family Penaeidae (Crustacea: Dendrobranchiata) in a phylogenetic context.

We describe the sperm ultrastructure of six penaeid species, including at least one member of each tribe (Penaeini, Parapenaeini and Trachypenaeini). Fragments of the vas deferens of the Penaeidae Farfantepenaeus brasiliensis, Farfantepenaeus paulensis, Litopenaeus schmitti, Parapenaeus americanus, Rimapenaeus constrictus and Xiphopenaeus kroyeri were fixed and processed according to the routine for transmission electron microscopy. The morphological results were contextualized in an evolutionary perspective using molecular markers for the phylogenetic reconstruction of this group. A phylogram was proposed by Bayesian inference based on 1007 bp of 33 sequences of the combined genes (16S rDNA and COI mtDNA) from 27 dendrobranchiate specimens. Our findings show that morphological differences in the sperm ultrastructures of members among the tribes of Penaeidae can be used as a baseline to understand their evolutionary relationships. Individuals from the Penaeini tribe show plesiomorphic characteristics in the sperm ultrastructure compared to the Trachypenaeini tribe from which they were derived, such as shrimp from family Sicyoniidae. The morphological complexity of the sperm of the different penaeid members corroborated with the genetic phylogeny, which showed different clades for each tribe and the close relationship with Sicyoniidae. The sperm features of the selected species studied here reflected their evolutionary history. These features confirm the previous phylogenetic hypothesis and question the monophyly of Penaeidae, which should be verified in the future with a more complete set of representative members of each tribe.

Lifespan and population dynamics of the endemic South American shrimp Artemesia longinaris (Crustacea: Penaeidae) in southeastern Brazil.

The present study investigated the growth, longevity and reproductive dynamics of Artemesia longinaris in the southeastern coast of Brazil over a two-year period. Monthly collections were conducted in Ubatuba and Caraguatatuba using a shrimp fishing boat equipped with "double-rig" nets. Each region was divided into 7 sampling stations up to 35 m deep. Size frequency distributions, growth, longevity, sex ratio, and abundance of individuals in each demographic class, were compared. The relationship between abiotic factors and abundance of each demographic class was assessed using a Canonical Correlation Analysis. A total of 64,641 individuals were collected (6,928 measured) with an estimated longevity of 1.30 (Ubatuba) and 1.14 (Caraguatatuba) years for females and 1.03 years for males in both regions. There was a statistically significant bias in sex ratio toward females (Chi-squared test, p < 0.05) in both regions. The Canonical Correlation Analysis resulted in a canonical correlation coefficient of 0.31 (p = 0.00002). Salinity and temperature showed high correlation mainly with the presence of reproductive females. In general, this demographic class was most common in conditions of low temperature and high salinity. These findings, as well as other studies carried out in colder regions with the same species, are consistent with classical latitudinal paradigm.

Annual, seasonal and spatial abundance of the seabob shrimp Xiphopenaeus kroyeri (Decapoda, Penaeidae) off the Southeastern coast of Brazil.

We investigated the influence of environmental factors in spatial and temporal distribution of the seabob shrimp Xiphopenaeus kroyeri in Santos Bay and São Vicente Estuary, state of São Paulo, Brazil. Monthly samples were obtained, from May 2008 through April 2010, from four locations in the estuary and four in the bay. No individual was collected in the estuary and this was attributed to the low salinity means recorded in this environment. We collected 109,153 individuals in the bay and there was no difference in abundance between the two years comprised by the study period. The similarity in spatial distribution can be related to sediment grain size that in all sampling locations showed great amount of very fine sand. The largest amount of reproductive females was obtained in early 2010, when temperature was high, and this could have increased the juvenile recruitment in April 2010. According to our results, the distribution of X. kroyeri in the study area is influenced by temperature, which is related to reproduction, and salinity, limiting the entrance of individuals in the estuarine region.

Inferring population connectivity across the range of distribution of the stiletto shrimp Artemesialonginaris Spence Bate, 1888 (Decapoda, Penaeidae) from DNA barcoding: implications for fishery management.

Artemesialonginaris is a marine shrimp endemic to the southwestern Atlantic and distributed from Atafona, Rio de Janeiro (Brazil) to Rawson, Chubut (Argentina). In recent years, this species has become an important target of the commercial fishery as a consequence of the decline in the fishery of more traditional and profitable marine shrimps. In addition, phenotypic variations have been documented in populations along its distribution. Therefore, investigations on the genetics of the fishing stocks are necessary for the development of sustainable management strategies and for understanding the possible sources of these variations. The mitochondrial gene Cytochrome Oxidase I (COI) was used to search for evidence of genetic structure among the populations of Artemesialonginaris and to analyze the phylogenetic relationships among them. A total of 60 specimens were collected from seven different localities, covering its geographical range. The final alignment showed 53 haplotypes (48 individuals and 5 shared), with no biogeographical pattern. The low genetic divergence found, with a non-significant FST value, also suggests the absence of population structure for this gene. These findings indicate a continuous gene flow among the populations analyzed, suggesting that the phenotypic variation is a consequence of different environmental conditions among the localities.

Effect of environmental factors on the abundance of decapod crustaceans from soft bottoms off southeastern Brazil.

This study investigated the importance of variations in environmental factors affecting the abundance patterns of decapods on the southeastern Brazilian coast. Sampling was carried out monthly from January 1998 through December 1999 in Ubatumirim and Mar Virado, Ubatuba region, using a commercial shrimp fishing boat equipped with double-rig nets. Six areas adjacent to rocky shores were chosen. Bottom-water samples were collected using a Nansen bottle, to measure the temperature and salinity. Sediment samples were also obtained by means of a Van Veen grab, for determination of texture and organic-matter content. The association of environmental factors with species abundance was evaluated by Canonical Correspondence Analysis (α = 0.05). Forty-one species of Decapoda were used in the multivariate analysis. The analysis indicated that sediment texture (phi) and bottom temperature were the main factors correlated (p < 0.05) with the spatial and temporal abundance of the species. Considering the study region as faunal transition zone, including a mixture of species of both tropical and subantarctic origin, the species responded differently to environmental factors, mainly temperature. It is conceivable that the decapods adjust their distribution according to their intrinsic physiological limitations, possibly as a result of the available resources.

First zoeal stage of the partner shrimp Periclimenes paivai Chace, with remarks on the genus Periclimenes O.G. Costa (Caridea, Palaemonidae.

The morphology of the first zoeal stage of Periclimenes paivai Chace is described and illustrated for the first time. Larvae were obtained from three females with embryos, caught in the type locality (Cananéia, São Paulo state, Brazil). The morphological characters are detailed and compared with all previous descriptions of larvae in the genus (P. amethysteus, P. brevicarpalis, P. diversipes, P. pandionis, P. sagittifer and P. soror). The zoeae I of Periclimenes species are very similar, but P. paivai can be separated from the other six species by means of five characteristics: 8 plumose setae on the inner margin of the antennal scale, one spine on the endopod of the maxillule, one cuspidate seta on the basal endite of the maxilulle, one plumose seta on the single coxal endite of the maxilla, and one plumose seta on the endopod of the maxilla. Remarks from a comparative analysis of available descriptions of the genus are furnished.