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Astropecten marginatus Gray, 1840 Figures 6–7 Astropecten marginatus Gray, 1840: 181. Astropecten marginatus — Brito 1962: 3; 1968: 7, pl. 4, fig. 1; Nomura & Fausto Filho 1966: 19; Lima-Verde 1969: 11; Carrera- Rodriguez & Tommasi 1977: 88–89; Tommasi et al. 1988: 5; Clark & Downey 1992: 41–42, fig. 10e, pl. 11A–B; Gondim et al. 2008: 155; Ventura et al. 2008: 177–178; Lima & Fernandes 2009: 58; Magris & Deìstro 2010: 59; Xavier 2010: 75; Gondim et al. 2014: 22–24, figs. 6a–e, 12b; Guilherme & Rosa 2014; Fernandez et al. 2017; Bueno et al. 2018: 177, fig. 6; Gurjão & Lotufo 2018: 10; Miranda 2018: 14; Patrizzi & Dobrovolski 2018: 182; Torres & Torres 2019: 412; Turra et al. 2019. Astropecten richardi — Perrier 1875: 372. Material examined (21 specs, 10–50 mm R). BRAZIL. Bahia, Santa Cruz de Cabrália (16°14’– 16°15’S; 38°00’– 38°58’W)— 7 m, 16.xi.2010, 5 specs, R 11–25 mm (MZUSP 2100); 6 m, 16.xi.2010, 6 specs, R 10–28 mm (MZUSP 2101); 7 m, 16.xi.2010, 5 specs, R 30–50 mm (MZUSP 2100); 6 m, 16.xi.2010, 5 specs, R 30–40 mm (MZUSP 2101). Description (R 30–50 mm). Disc broad and flattened, arms short and flattened (Fig. 6A–C); R/r 3.3; 39 SM plates (R 50 mm); R/SM# 1.3. Abactinal region covered by paxillae as follows: proximal region of arms and disc with 10–12 central and 15 peripheral spinelets (Fig. 6D), midline region of arms with 5–8 central and nine peripheral spinelets, and distal region of arms with 1–3 central and seven peripheral spinelets. Madreporite large, not hidden by paxillae (Fig. 6E). Marginal plates horizontal and elongated. Superomarginal plates without spines and covered by granules: central granules robust, peripheral granules elongated, hyaline and thin. Inferomarginal plates projected beyond superomarginal plates. Fringe with two layers of spines, parallel, top layer with two spines of equal size, and a third proximal 3x smaller and thinner (Fig. 6G); bottom layer with 4–5 small spines, proximal spine smaller and thinner (Fig. 6I). Actinal region of inferomarginal plates lacking squamules; central region naked, sometimes with long spines similar to lateral fringe spines (Fig. 6J). Three adambulacral spines forming single line, central spine longer than adjacent ones (Fig. 6K). One large subambulacral spine in center of plate, largest spine in adambulacral plate, surrounded by acicular spinelets; spinelets between large spine and furrow spines. Oral spines flattened with blunt tip (Fig. 6F). Pedicellariae absent. Ontogenetic variation (R 10–28 mm). Average R/r 3.0; 22 SM plates (R 20 mm); R/SM# 1.1. Proximal paxillae with 3–5 central and 8–10 peripheral spinelets (Fig. 7C), distal paxillae with 0–1 central and 4–5 peripheral spinelets. Inferomarginal fringe with two layers as in larger specimen, but proximal spine in top layer 5x smaller than adjacent spines, and bottom layer with 3–4 small spines, proximal spine smallest (Fig. 7F). Three adambulacral spines forming single line, central spine largest in adambulacral plate (Fig. 7E). Second subambulacral row as in larger specimen, with central spine largest than adjacent spines, but not largest in adambulacral plate; third row with 2–4 sub-equal spines. Coloration. Abactinal surface greyish in center of disc and arms, margins orangish to creamy; inferomarginal spines fade from a bluish-grey base to creamy tips. Specimens in ethanol are pale brown. Distribution. Gulf of Mexico, Mexico, Dominican Republic, Puerto Rico, Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guyana (Walenkamp 1976; Clark & Downey 1992; Alvarado et al. 2008; Benavides- Serrato et al. 2005, 2011; Pawson et al. 2009, Alvarado & Solís-Marín 2013). BRAZIL: Amapá, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Bahia, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul (Bernasconi 1955; Tommasi 1958; Brito 1962; Lima-Verde 1969; Netto 2006; Gondim et al. 2008; Pelaes 2008; Lima & Fernandes 2009; Xavier 2010; Gondim et al. 2014; Bueno et al. 2018; Miranda 2018; Torres & Torres 2019; Turra et al. 2019). Depth. 1–130 m (Clark & Downey 1992). Biological notes. This species lives in muddy, sandy and rubble bottoms (Alvarado & Solís-Marín 2013) and most likely feeds during the day, preferably on gastropods, bivalves and crustaceans (Bitter 1984; Zoldan 2005; Guilherme & Rosa 2014; Fernandez et al. 2017). In Caraguatatuba Bay, São Paulo, A. marginatus lives in sympatry with L. senegalensis, where they partition food resources (Fernandez et al. 2017). The reproductive cycle of A. marginatus in this region is annual and spawning happens mainly during the rainy season (Turra et al. 2019). Netto (2006) found many specimens gathered in shallow pools at night with 2–5 arms facing upwards, possibly engaged in gas exchange as the exposure of the tube feet into the water column should increase its breathing surface (Farmanfarmaian 1966). Astropecten marginatus is classified as “Vulnerable” (baseline data indicate a reduction in population by at least 30% over the next 100 years) by the Ministry of the Environment (MMA 2018). According to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Holotype. Gray (1840, p. 178) mentioned that the specimens he described were deposited in the collections of the Zoological Society of London (especially South American specimens collected by Mr. Hugh Cuming) and of the British Museum, where he worked. However, Andrew Cabrinovic told us that the type has not been found at the NHM-UK (pers. comm. on 20 Feb 20). Type locality. Not reported by Gray (1840). Taxonomic remarks of the genus Astropecten As noted by Cobb et al. (2019), telling some of the Astropecten species apart is not trivial. We found several instances of misidentification in prior work from Northeastern Brazil and we hope that the present work will reduce errors in future studies. Phylogenetic relationships between some species of Brazilian Astropecten were addressed by Zulliger & Lessios (2010) and Cobb et al. (2019). Their molecular cladograms indicate that A. antillensis is more closely related to A. articulatus than to A. cingulatus. The relationship between these species and A. marginatus is uncertain as this species could be equally related to A. antillensis, A. articulatus and A. cingulatus or most closely related to A. cingulatus (Zulliger & Lessios, 2010). Astropencten brasiliensis was not included in these studies. Astropecten acutiradiatus differs from A. alligator by having an enlarged spine in the first subambulacral row (vs. three (sub)equal spines); from A. antillensis by having an enlarged distal spine in the first subambulacral row (vs. enlarged central spine); from A. articulatus by having clavate paxillar spinelets (vs. paxillar spinelets granulose); from A. brasiliensis brasiliensis and A. duplicatus by the absence of enlarged spines on the distal superomarginal plates (vs. presence of enlarged spines on the distal superomarginal plates); from A. cingulatus and A. marginatus by having spines on the proximal superomarginal plates (vs. superomarginal spines absent). Astropecten antillensis differs from A. acutiradiatus, A. articulatus, A. brasiliensis brasiliensis and A. duplicatus by having an enlarged central spine in the first subambulacral row (vs. enlarged distal spine); from A. alligator by having an enlarged spine in the first subambulacral row (vs. three (sub)equal spines); from A. cingulatus and A. marginatus by having spines on the superomarginal plates (vs. superomarginal spines absent). Astropecten antillensis also differs from A. brasiliensis brasiliensis because its inner superomarginal (not the first pair) spine is smaller than the outer spine (vs. inner spine larger than outer spine). Astropecten brasiliensis brasiliensis differs from A. marginatus and A. cingulatus by having superomarginal spines (vs. superomarginal spines absent); from A. articulatus and A. duplicatus by having subambulacral spines with blunt tips (vs. spines truncated in two), and inferomarginal fringe spines oblique (vs. inferomarginal fringe spines horizontal); and from A. alligator by having an enlarged distal spine in the first subambulacral row (vs. (sub)equal spines). Astropecten cingulatus differs from A. marginatus by having inferomarginal plates densely covered with overlapping squamules and spines (vs. inferomarginal plates bare or with sparse, widely spaced squamules), and these two species differ from the other Brazilian Astropecten by having no spines on the superomarginal plates (vs. spines present)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 15-18, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Nomura, H. & Fausto Filho, J. (1966) Shrimp survey in coastal and offshore waters of northeastern Brazil. Arquivos da Estacao de Biologia Marinha da Universidade Federal do Ceara, 6, 15 - 29.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do Nordeste brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Gondim, A., Lacouth, P., Alonso, C. & Manso, C. (2008) Echinodermata da praia do Cabo Branco, JoaTo Pessoa, Paraiiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Guilherme, P. D. & Rosa, L. C. (2014) Seasonal variation in body size and diet of the sea star Astropecten marginatus (Paxillosida, Astropectinidae) off coast of Parana, Southern Brazil. Revista de Biologia Tropical, 62, 59 - 68. https: // doi. org / 10.15517 / rbt. v 62 i 1.7888","Fernandez, W. S., Dias, G. M., Majer, A., Delboni, C., Denadai, M. R. & Turra, A. (2017) Resource partitioning between sympatric starfish from tropical unconsolidated substrate: implications for coexistence and top-down control on benthic prey. Estuarine, Coastal and Shelf Science, 196, 141 - 149. https: // doi. org / 10.1016 / j. ecss. 2017.06.039","Bueno, M. L., Alitto, R. A. S., Guilherme P. D. B., Domenico, M. D. & Borges, M. (2018) Guia ilustrado dos Echinodermata da porcao sul do Embaiamento Sul Brasileiro. Pesquisa e Ensino em Ciencias Exatas e da Natureza, 2 (2), 169 - 237. https: // doi. org / 10.29215 / pecen. v 2 i 2.1071","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Turra, A., Checon, H. H., Fernandez, W. S., Majer, A. P., Delboni, C. G., Denadai, M. R. & Dias, G. M. (2019) The importance of embayed coastal areas for the life cycle of two sympatric starfish species (Echinodermata, Asteroidea). Estuarine, Coastal and Shelf Science, 219, 128 - 138. https: // doi. org / 10.1016 / j. ecss. 2019.01.028","Perrier, E. (1875) Revision de la collection de stellerides du Museum d'Histoire Naturelle de Paris. Archives de Zoologie Experimentale et Generate, 4, 265 - 450.","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Pawson, D. L., Vance, D. J., Messing, C. G., Solis-Marin, F. A. & Mah, C. L. (2009) Echinodermata of the Gulf of Mexico. In: Felder, D. L. & Camp, D. K. (Eds.), Gulf of Mexico - Origins, Waters, and Biota. Texas A & M University Press, College Station, Texas, pp. 1177 - 1204.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Bernasconi, I. (1955) Equinoideos y asteroideos de la coleccion del instituto oceanografico de la Universidad de San Pablo. Primera contribucion. Boletim do Instituto Oceanografico, 6, 51 - 57. https: // doi. org / 10.1590 / S 0373 - 55241955000100002","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Netto, L. F. (2006) Echinodermata do Canal de Sao Sebastiao, Sao Sebastiao. Unpublished Master Thesis, Universidade de Sao Paulo, Sao Paulo, 201 pp.","Pelaes, M. A. L. (2008) Composicao, distribuicao espacial e temporal das especies de Echinodermata na plataforma rasa do sul da Bahia num trecho entre Itacare e Canavieiras. Unpublished Master Thesis, Universidade Estadual de Santa Cruz, Ilheus, Bahia, 97 pp.","Bitter, R. (1984) Composicion de la dieta de Astropecten marginatus (Echinodermata: Asteroidea). Golfo Triste, region Centro- Occidental. Boletin del Instituto Oceanografico, Universidad de Oriente, 23, 169 - 176.","Zoldan, L. V. (2005) Analise do conteudo estomacal de duas especies de estrela-do-mar: Astropecten marginatus (Gray, 1840) e Luidia senegalensis (Lamarck, 1816) na enseada de Caraguatatuba, SP, sudeste do Brasil. Unpublished Honors Thesis, Centro Universitario da Fundacao de Ensino Octavio Bastos, Sao Joao da Boa Vista, Sao Paulo, 41 pp.","Farmanfarmaian, A. (1966) The respiratory physiology of echinoderms. In: Boolootian, R. A. (Ed.), Physiology of Echinodermata. John Wiley & Sons, New York, pp. 245 - 265.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Cobb, J. C., Lawrence, J. M., Herrera, J. C. & Janies, D. (2019) A new species of Astropecten (Echinodermata: Asteroidea: Paxillosida: Astropectinidae) and a comparison of the Astropecten species from the Gulf of Mexico and the East Florida Shelf. Zootaxa, 4612 (3), 301 - 325. https: // doi. org / 10.11646 / zootaxa. 4612.3.1","Zulliger, D. E. & Lessios, H. A. (2010) Phylogenetic relationships in the genus Astropecten Gray (Paxillosida: Astropectinidae) on a global scale: molecular evidence for morphological convergence, species-complexes and possible cryptic speciation. Zootaxa, 2504 (1), 1 - 19. https: // doi. org / 10.11646 / zootaxa. 2504.1.1"]}

Astropecten marginatus Gray 1840 presenta una amplia distribución en el norte y este de sur América, encontrándose en fondos arenosos y fangosos de aguas someras y profundas. Se determinó su distribución, parámetros poblacionales tales como: talla, peso y dieta, de los ejemplares provenientes de los muestreos de megabentos de los proyectos LBAPD y CFI en las épocas de lluvia y sequía en la ecorregión Delta del Orinoco, Venezuela. Se dividió la ecorregión en tres sectores: Golfo de Paria, Boca de Serpiente y Plataforma Deltana. Las estrellas fueron medidas, pesadas y se extrajo su estómago para la determinación de las presas. Las mayores densidades se encontraron en época de sequía en Golfo de Paria (2006) y en época de lluvia en Plataforma Deltana (2005). El intervalo de tallas más frecuente fue 3.1-4.6cm. La mayor biomasa se encontró en Golfo de Paria en época de sequía. No se observaron relaciones entre el tamaño de la estrella y la profundidad de muestreo. Existe una dominancia de los moluscos en la dieta, a pesar de tener una alimentación heterogénea. Se observó que aparte de la depredación, también pueden presentar comportamientos carroñeros y detritívoros.Distribution, population parameters, and diet of Astropecten marginatus (Asteroidea: Astropectinidae) in the Venezuelan Atlantic coast. Astropecten marginatus is a sea star widely distributed in Northern and Eastern South America, found on sandy and muddy bottoms, in shallow and deep waters. To describe some of its ecological characteristics, we calculated it spatial-temporal distribution, population parameters (based on size and weight) and diet in the Orinoco Delta ecoregion (Venezuela). The ecoregion was divided in three sections: Golfo de Paria, Boca de Serpiente and Plataforma Deltana. Samples for the rainy and dry seasons came from megabenthos surveys of the "Línea Base Ambiental Plataforma Deltana (LBAPD)" and "Corocoro Fase I (CFI)" projects. The collected sea stars were measured, weighted and dissected by the oral side to extract their stomach and identify the preys consumed. A total of 570 sea stars were collected in LBAPD project and 306 in CFI one. The highest densities were found during the dry season in almost all sections. In LBAPD project the highest density was in "Plataforma Deltana" section (0.007±0.022ind/m² in dry season and 0.014±0.06ind/m² in rainy season) and in the CFI project the densities in "Golfo de Paria" section were 0.705±0.829ind/m² in rainy season and 1.027±1.107ind/m² in dry season. The most frequent size range was 3.1-4.6cm. The highest biomass was found in "Golfo de Paria" section (7.581±0.018mg/m² in dry season and 0.005±6.542x10-06mg/m² in rainy season for 2004-2005 and 3.979±4.024mg/m² in dry season; and 3.117±3.137mg/m² in rainy season for 2006). A linear relationship was found between the sea star size and its weight but no relationship was observed between its size and the depth where it was collected. Mollusks are dominant in the sea star diet (47.4% in abundance). The diet in any of the sections, seasons or between projects or size class was heterogeneous, using multivariate ordinations (MDS) and SIMPER analysis and there was no difference in the prey number or food elements that a sea star can eat. Although A. marginatus has been described as a predator, in this study were also inferred scavenger and detritivorous habits. Rev. Biol. Trop. 59 (1): 53-69. Epub 2011 March 01.

The sea star Astropecten marginatus has a neotropical distribution and is a highly abundant and frequent species in shrimp trawling by-catchin many places along the Brazilian coast. This has caused its threat to extinction and in addition, its bio-ecological aspects are poorly known. Thus, the main objective of this study was to analyze the seasonal variations of population length structure and feeding habits of the sea stars A. marginatus inhabiting off state of Paraná, Southern Brazil. The analyzed specimens were collected in February (summer), April (fall), June (winter) and October (spring) of 2008 from shrimp by-catch trawling. In the laboratory, each individual had its length measured and then weighed on an analytical scale. Afterwards, the stomach contents of 10 individuals of each of two most frequent length classes were seasonally analyzed. The relative frequency and abundance for each prey category was determined and, then combined into an index of alimentary importance. A total of 994 individuals of A. marginatus were collected with length ranging from 7.0 to 56.2mm, but most individuals were in the 20.1-25mm length classes. Individuals larger than 40mm were only collected in the spring while a few recruits (40mm) se encontraron solamente en la primavera, mientras aquellos con talla inferior a los 10mm se obtuvieron durante el otoño y el invierno. El peso de los individuos se mantuvo entre los 0.1-15.3g. La relación entre el peso y talla mostró un crecimiento alométrico negativo (b

The common Puerto-Rican keyhole urchin, Mellita lata H. L. Clark, and the sea-star, Astropecten marginatus Gray, live in the uppermost layers of the sand of shallow beaches. The color in life, and various locomotor activities of the two species are described.