Abstract
The shrimp genera Leptalpheus Williams, 1965 and Fenneralpheus Felder & Manning, 1986 are composed entirely of symbiotic species that co-inhabit burrows of infaunal macrocrustaceans. We report extensive collections of these genera from western Atlantic, eastern Pacific and Indo-West Pacific regions. Integrative taxonomy methods, including morphological comparisons and analysis of three mitochondrial genetic markers, are used to test species hypotheses and evolutionary relationships among members of these genera. Our molecular analysis failed to recover Leptalpheus or Fenneralpheus as monophyletic groups. Our results strongly supported the monophyly of three clades composed of species of Leptalpheus, loosely corresponding to previously proposed species groups. Three new species closely related to Leptalpheus forceps Williams, 1965, L. marginalis Anker, 2011, and L. mexicanus Ríos & Carvacho, 1983 are described. Leptalpheus ankeri n. sp., from the Caribbean Sea, Atlantic coast of Florida, and Gulf of Mexico, is a polymorphic species that exhibits two major cheliped morphotypes. Leptalpheus sibo n. sp., from the Pacific coast of Nicaragua, is morphologically very similar to L. ankeri n. sp., likely its transisthmian sister species, and shares its cheliped polymorphism. A reassessment of L. forceps concluded that records of this species from the Caribbean Sea and Brazil are not conspecific with L. forceps sensu stricto from the Atlantic coast of the USA and the Gulf of Mexico, and they are herein described as Leptalpheus degravei n. sp. Based on both molecular and morphological evidence, we found Leptalpheus bicristatus Anker, 2011 to be a junior synonym of L. mexicanus and Leptalpheus canterakintzi Anker & Lazarus, 2015 to be a junior synonym of Leptalpheus azuero Anker, 2011. First reports of Leptalpheus axianassae Dworschak & Coelho, 1999 in Texas and Mexico, Leptalpheus denticulatus Anker & Marin, 2009 in the Mariana Islands, Leptalpheus felderi Anker, Vera Caripe & Lira, 2006 and Leptalpheus lirai Vera Caripe, Pereda & Anker, 2021 in the USA, and Leptalpheus pereirai Anker & Vera Caripe, 2016 in Cuba are included.
References
- Almeida, A.O., Boehs, G., Araújo-Silva, C.L. & Bezerra, L.E.A. (2012) Shallow-water caridean shrimps from southern Bahia, Brazil, including the first record of Synalpheus ul (Ríos & Duffy, 2007) (Alpheidae) in the southwestern Atlantic Ocean. Zootaxa, 3347 (1), 1–35. https://doi.org/10.11646/zootaxa.3347.1.1
- Almeida, A.O., Costa-Souza, A.C., Cunha, A.M., Santos, P.S., Oliveira, M.V. & Soledade, G.O. (2013) Estuarine caridean shrimps (Crustacea: Decapoda) from Ilhéus, Bahia, Brazil: Updated checklist and a key for their identification. Check List, 9 (6), 1396–1405. https://doi.org/10.15560/9.6.1396
- Almeida, A.O., Terossi, M., Buranelli, R.C., Castilho, A.L., Costa, R.C., Zara, F.J. & Mantelatto, F.L. (2018) Checklist of decapods (Crustacea) from the coast of São Paulo State (Brazil) supported by integrative molecular and morphological data: II. Infraorder Caridea: family Alpheidae. Zootaxa, 4450 (3), 331–358. https://doi.org/10.11646/zootaxa.4450.3.2
- Álvarez, F., Villalobos, J.L., Rojas, Y. & Robles, R. (1999) Listas y comentarios sobre los crustáceos decápodos de Veracruz, México. Anales del Instituto de Biología, Universidad Nacional Autónoma de México (Serie Zoología), 70 (1), 1–27.
- Anker, A. (2001) Taxonomie et évolution des Alpheidae (Crustacea, Decapoda). Vols. 1 & 2. PhD Thesis. Muséum National d’Histoire Naturelle, Paris, 547 + 331 pp.
- Anker, A. (2003) Alpheid shrimps from the mangroves and mudflats of Singapore. Part I. Genera Salmoneus, Athanas and Potamalpheops, with the description of two new species (Crustacea: Decapoda: Caridea). Raffles Bulletin of Zoology, 51 (2), 283–314.
- Anker, A. (2008) The shrimp genus Leptalpheus Williams, 1965 in the southwestern Caribbean Sea, with description of one new species from Panama (Crustacea, Decapoda, Alpheidae). Zoosystema, 30 (4), 781–794.
- Anker, A. (2010) New findings of rare or little-known alpheid shrimp genera (Crustacea, Decapoda) in Moorea, French Polynesia. Zootaxa, 2403 (1), 23–41. https://doi.org/10.11646/zootaxa.2403.1.3
- Anker, A. (2011) Six new species and three new records of infaunal alpheid shrimps from the genera Leptalpheus Williams, 1965 and Fenneralpheus Felder & Manning, 1986 (Crustacea, Decapoda). Zootaxa, 3041 (1), 1–38. https://doi.org/10.11646/zootaxa.3041.1.1
- Anker, A., Ahyong, S.T., Noël, P.Y. & Palmer, A.R. (2006a) Morphological phylogeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw. Evolution, 60, 2507–2528. https://doi.org/10.1111/j.0014-3820.2006.tb01886.x
- Anker, A. & Ashrafi, H. (2019) Salmoneus durisi sp. nov., an infaunal alpheid shrimp probably associated with callianassid ghost shrimps in the tropical Indo-West Pacific (Malacostraca: Decapoda: Caridea). Zootaxa, 4651 (1), 64–74. https://doi.org/10.11646/zootaxa.4651.1.4
- Anker, A. & Dworschak, P.C. (2007) Jengalpheops rufus gen. nov., sp. nov., a new commensal alpheid shrimp from the Philippines (Crustacea: Decapoda). Zoological Studies, 46 (3), 290–302.
- Anker, A., Hurt, C. & Knowlton, N. (2007) Three transisthmian snapping shrimps (Crustacea: Decapoda: Alpheidae: Alpheus) associated with innkeeper worms (Echiura: Thalassematidae) in Panama. Zootaxa, 1626 (1), 1–23. https://doi.org/10.11646/zootaxa.1626.1.1
- Anker, A., Jeng, M. & Chan, T.-Y. (2001) Two unusual species of Alpheidae (Decapoda: Caridea) associated with upogebiid mudshrimps in the mudflats of Taiwan and Vietnam. Journal of Crustacean, Biology, 21, 1049–1061. https://doi.org/10.1163/20021975-99990197
- Anker, A. & Lazarus, J.F. (2015) Description of two new associated infaunal decapod crustaceans (Axianassidae and Alpheidae) from the tropical eastern Pacific. Papéis Avulsos de Zoologia, 55 (8), 115–129. https://doi.org/10.1590/0031-1049.2015.55.08
- Anker, A. & Marin, I.N. (2009) The alpheid shrimp genus Leptalpheus Williams, 1965, in the tropical western Pacific, with descriptions of two new species (Crustacea: Decapoda: Caridea). Raffles Bulletin of Zoology, 57 (1), 91–107.
- Anker, A., Pratama, I.S., Firdaus, M. & Rahayu, D.L. (2015) On some interesting marine decapod crustaceans (Alpheidae, Laomediidae, Strahlaxiidae) from Lombok, Indonesia. Zootaxa, 3911 (3), 301–342. https://doi.org/10.11646/zootaxa.3911.3.1
- Anker, A. & Vera Caripe, J. (2016) Leptalpheus pereirai sp. nov., a new alpheid shrimp from Panama and Venezuela (Decapoda: Caridea). Zootaxa, 4127 (1), 185–191. https://doi.org/10.11646/zootaxa.4127.1.11
- Anker, A., Vera Caripe, J. & Lira, C. (2006b) Description of a new species of commensal alpheid shrimp (Crustacea, Decapoda) from the southern Caribbean Sea. Zoosystema, 28 (3), 683–702.
- Banner, A.H. & Banner, D.M. (1974) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part XVII. Additional notes on the Hawaiian alpheids: new species, subspecies and some nomenclatorial changes. Pacific Science, 28 (4), 423–437.
- Banner, A.H. & Banner, D.M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l’ORSTOM, 158, 1–164.
- Botter-Carvalho, M.L., Costa, L.B., Gomes, L.L., Clemente, C.C.C. & Carvalho, P.V.V.C. (2015) Reproductive biology and population structure of Axianassa australis (Crustacea, Axianassidae) on a sand-mud flat in north-eastern Brazil. Journal of the Marine Biological Association of the United Kingdom, 95, 735–745. https://doi.org/10.1017/S002531541400174X
- Bracken, H.D., De Grave, S. & Felder, D.L. (2009) Phylogeny of the infraorder Caridea based on mitchondrial and nuclear genes (Crustacea: Decapoda). In: Martin, J.W., Crandall, K.A. & Felder, D.L. (Eds.), Decapod Crustacean Phylogenetics. Koenemann, S., Crustacean Issues. Vol. 18. CRC Press, Taylor & Francis Group, Boca Raton, London and New York, pp. 281–305. https://doi.org/10.1201/9781420092592-c14
- Bruce, A.J. (1976) Coral reef Caridea and “commensalism”. Micronesica, 12 (1), 83–98.
- Buhay, J.E., Moni, G., Mann, N. & Crandall, K.A. (2007) Molecular taxonomy in the dark: evolutionary history, phylogeography, and diversity of cave crayfish in the subgenus Aviticambarus, genus Cambarus. Molecular Phylogenetics and Evolution, 42 (2), 435–448. https://doi.org/10.1016/j.ympev.2006.07.014
- Campos, E., Félix-Pico, E.F. & García-Domínguez, F. (1995) Distribution and host for four symbiotic crustaceans of the Mexican Pacific (Stomatopoda and Decapoda). Bulletin ofthe Southern California Academy of Sciences, 94 (2), 176–178
- Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution, 17, 540–552. https://doi.org/10.1093/oxfordjournals.molbev.a026334
- Chace, F.A. Jr. (1972) The shrimps of the Smithsonian-Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology, 98, 1–179. https://doi.org/10.5479/si.00810282.98
- Chow, L.H., De Grave, S., Anker, A., Poon, K.K.Y., Ma, K.Y., Chu, K.H., Chan, T.Y. & Tsang, L.M. (2021) Distinct suites of pre- and post-adaptations indicate independent evolutionary pathways of snapping claws in the shrimp family Alpheidae (Decapoda: Caridea). Evolution, 75 (11), 2898–2910. https://doi.org/10.1111/evo.14351
- Christoffersen, M.L. (1998) Malacostraca. Eucarida. Caridea. Crangonoidea and Alpheoidea (except Glyphocrangonidae and Crangonidae). In: Young, P.S. (Ed.), Catalogue of Crustacea of Brazil. Museu Nacional, Rio de Janeiro, pp. 351–372.
- Coutière, H. (1897) Note sur quelques genres nouveaux ou peu connus d’Alphéidés, formant la sous famille des Alphéopsidés. Bulletin du Muséum d’Histoire naturelle, 2, 380–386.
- Crandall, K.A. & Fitzpatrick, J.F. Jr. (1996) Crayfish molecular systematics: using a combination of procedures to estimate phylogeny. Systematic Biology, 45 (1), 1–26. https://doi.org/10.1093/sysbio/45.1.1
- Dawson, C.E. (1967) Notice of the occurrence of the alpheid shrimp Leptalpheus forceps Williams in the northern Gulf of Mexico. Crustaceana, 12 (2), 224. https://doi.org/10.1163/156854067X00648
- De Grave, S. & Anker, A. (2017) An annotated checklist of marine caridean and stenopodidean shrimps (Malacostraca: Decapoda) of the Caribbean coast of Panama. Nauplius, 25, 1–40. https://doi.org/10.1590/2358-2936e2017015
- De Grave, S. & Fransen, C.H.J.M. (2011) Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, Leiden, 85, 195–589.
- De Kay, J.E. (1844) Crustacea. In: Zoology of New York, or the New York Fauna; Comprising detailed descriptions of all the animals hitherto observed within the state of New-York, with brief notices of those occasionally found near its borders, and accompanied by appropriate illustrations. Part VI. Carroll and Cook, Albany, ix + 72 pp., 13 pls.
- Dworschak, P.C. (2011) Redescription of Callianassa jousseaumei Nobili, 1904, a junior subjective synonym of Callianassa indica de Man, 1905 with description of a new species of Neocallichirus (Decapoda: Axiidea: Callianassidae). Zootaxa, 2746 (1), 1–19. https://doi.org/10.11646/zootaxa.2746.1.1
- Dworschak, P.C. & Coelho, V.R. (1999) On two alpheids from Araçá (São Paulo, Brazil) with a description of a new species of Leptalpheus (Decapoda: Caridea: Alpheidae). Annalen des Naturhistorischen Museums in Wien, 101B, 475–488.
- Felder, D.L. (2001) Diversity and ecological significance of deep-burrowing macrocrustaceans in coastal tropical waters of the Americas (Decapoda: Thalassinidea). Interciencia, 26 (10), 440–449.
- Felder, D.L. (2015) A new species of the ghost shrimp genus Lepidophthalmus (Crustacea: Decapoda: Axiidea) from the southwestern Gulf of Mexico. Zootaxa, 3985 (3), 409–420. https://doi.org/10.11646/zootaxa.3985.3.5
- Felder, D.L. & Manning, R.B. (1986) A new genus and two new species of alpheid shrimps (Decapoda: Caridea) from South Florida. Journal of Crustacean Biology, 6 (3), 497–508. https://doi.org/10.2307/1548189
- Felder, D.L. & Manning, R.B. (1997) Ghost shrimps of the genus Lepidophthalmus from the Caribbean region, with description of L. richardi, new species, from Belize (Decapoda: Thalassinidea: Callianassidae). Journal of Crustacean Biology, 17 (2), 309–331. https://doi.org/10.1163/193724097X00341
- Felder, D.L., Nates, S.F. & Robles, R.R. (2003) Hurricane Mitch: impacts of bioturbating crustaceans in shrimp ponds and adjacent estuaries of coastal Nicaragua. US Geological Survey Open File Report, 03-179, 1–70. https://doi.org/10.3133/ofr03179
- Felder, D.L. & Robles, R. (2015) Two new species of the genus Lepidophthalmus (Decapoda, Axiidea, Callianassidae) from coastal Pacific waters of Central America. Zootaxa, 4020 (3), 431–452. https://doi.org/10.11646/zootaxa.4020.3.2
- Felder, D.L. & Rodrigues, S.A. (1993) Reexamination of the ghost shrimp Lepidophthalmus louisianensis (Schmitt, 1935) from the northern Gulf of Mexico and comparison to L. siriboia, new species, from Brazil (Decapoda: Thalassinidea: Callianassidae). Journal of Crustacean Biology, 13 (2), 357–376. https://doi.org/10.1163/193724093X00138
- Felder, D.L. & Staton, J.L. (2000) Lepidophthalmus manningi, a new ghost shrimp from the southwestern Gulf of Mexico (Decapoda: Thalassinidea: Callianassidae). Journal of Crustacean Biology, 20 (Special Number 2), 170–181. https://doi.org/10.1163/1937240X-90000018
- Froglia, C. & Atkinson, R.J.A. (1998) Association between Athanas amazone (Decapoda: Alpheidae) and Squilla mantis (Stomatopoda: Squillidae). Journal of Crustacean Biology, 18, 529–532. https://doi.org/10.2307/1549417
- Geller, J., Meyer, C., Parker, M. & Hawk, H. (2013) Redesign of PCR primers for mitochondrial cytochrome c oxidase subunit I for marine invertebrates and application in all-taxa biotic surveys. Molecular Ecology Resources, 13 (5), 851–861. https://doi.org/10.1111/1755-0998.12138
- Gibbes, L.R. (1850) On the carcinological collections of the United States, and an enumeration of species contained in them, with notes on the most remarkable, and descriptions of new species. Proceedings of the American Association for the Advancement of Science, 3, 165–201.
- Gomes Corrêa, M.M. (1968) Sobre as espécies de Upogebia Leach do litoral brasileiro, com descrição de uma espécie nova (Decapoda, Callianassidae). Revista Brasileira de Zoologia, 28, 97–109.
- Hermoso-Salazar, M. (2001) Ampliación de ámbito de Leptalpheus forceps (Caridea: Alpheidae) al Golfo de México. International Journal of Tropical Biology and Conservation, 49, 1278.
- Holthuis, L.B. (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda). With an appendix on the order Amphionidacea. National Natuurhistorisch Museum, Leiden, 328 pp. https://doi.org/10.5962/bhl.title.152891
- Kaji, T., Anker, A., Wirkner, C.S. & Palmer, A.R. (2018) Parallel saltational evolution of ultrafast movements in snapping shrimp claws. Current Biology, 28, 106–113. https://doi.org/10.1016/j.cub.2017.11.044
- Kemp, S. (1915) The Fauna of the Chilka Lake, Crustacea, Decapoda. Memoirs of the Indian Museum, 5 (3), 199–325, pls. 12–13. https://doi.org/10.5962/bhl.title.10414
- Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35 (6), 1547. https://doi.org/10.1093/molbev/msy096
- Lazarus-Agudelo, J.F. & Cantera-Kintz, J.R. (2007) Crustáceos (Crustacea: Sessilia, Stomatopoda, Isopoda, Amphipoda, Decapoda) de Bahía Málaga, Valle del Cauca (Pacífico colombiano). Biota Colombiana, 8 (2), 221–239.
- Lemaitre, R. & Rodrigues, S.A. (1991) Lepidophthalmus sinuensis: a new species of ghost shrimp (Decapoda, Thalassinidea, Callianassidae) of importance to the commercial culture of penaeid shrimps on the Caribbean coast of Colombia, with observations on its ecology. Fishery Bulletin, 89, 623–630.
- Manning, R.B. (1975) Two methods for collecting decapods in shallow water. Crustaceana, 29 (3), 317–319. https://doi.org/10.1163/156854075X00450
- Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 gateway computing environments workshop (GCE), 2010, 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Milne-Edwards, A. (1870) Révision du genre Callianassa (Leach) et description de plusieurs espèces nouvelles de ce groupe faisant partie de la collection du Muséum. Nouvelles Archives du Muséum d’Histoire Naturelle de Paris, 6, 75–102, pls. I–II. https://doi.org/10.5962/bhl.title.10050
- Miya, Y. (1997) Stenalpheops anacanthus, new genus, new species (Crustacea, Decapoda, Alpheidae) from the Seto Inland Sea and the Sea of Ariake, south Japan. Bulletin of the Faculty of Liberal Arts, Natural Science, Nagasaki University, 38, 145–161.
- Moyo, R., Pillay, D. & Baeza, J.A. (2017) Symbiont-mediated shifts in sandprawn behaviour: implications for ecosystem functioning in marine soft-sediment ecosystems. Journal of Experimental Marine Biology and Ecology, 486, 296–304. https://doi.org/10.1016/j.jembe.2016.10.022
- Nobili, G. (1904) Diagnoses préliminaires de vingt-huit espèces nouvelles de stomatopodes et décapodes macroures de la Mer Rouge. Bulletin du Muséum d’Histoire naturelle, 10, 228–238.
- Oliveira, M.V., Costa-Souza, A.C., Guimarães, F.J., Almeida, A.O. & Baeza, J.A. (2015) Observations on the life history of a rare shrimp, Salmoneus carvachoi (Crustacea: Caridea: Alpheidae), a possible simultaneous hermaphrodite. Marine Biodiversity Records, 8, e141. https://doi.org/10.1017/S1755267215001219
- Pachelle, P.P.G., Anker, A., Mendes, C.B. & Bezerra, L.E.A. (2016) Decapod crustaceans from the state of Ceará, northeastern Brazil: an updated checklist of marine and estuarine species, with 23 new records. Zootaxa, 4131 (1), 1–63. https://doi.org/10.11646/zootaxa.4131.1.1
- Palumbi, S.R., Martin, A.P., McMillan, W.O., Romano, S.R., Grabowsky, G. & Stice, L. (1991) The simple fool’s guide to PCR. Version 2. Department of Zoology, University of Hawaii, Honolulu, 47 pp.
- Posey, M.H., Alphin, T.D., Banner, S., Vose, F. & Lindberg, W. (1998) Temporal variability, diversity and guild structure of a benthic community in the northeastern Gulf of Mexico. Bulletin of Marine Science, 63 (1), 143–155.
- Rambaut, A. (2007) FigTree, a graphical viewer of phylogenetic trees. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh. [program]
- Ramos, G.E. (1995) Nuevos registros de camarones alfeídeos (Crustacea, Decapoda, Alpheidae) para el Pacífico de Colombia. In: Cantera, J.R. & Restrepo, J.D. (Eds.), Delta del Río San Juan, Bahías de Málaga y Buenaventura, Pacífico Colombiano. Vol. 2. Colciencias. Universidad EAFIT, Medellín and Universidad del Valle, Cali, pp. 127–153.
- Ramos-Porto, M. & Souza, S.T. (1994) Descrição de Leptalpheus petronii sp. n. (Decapoda, Alpheidae) para o Brasil. In: Bonecker, S.L.C. (Ed.), Resumos do XX Congresso Brasileiro de Zoologia, 24–29 de julho 1994. Universidade Federal do Rio de Janeiro, Rio de Janeiro, 22 pp.
- Ríos, R. (1992) Camarones Carideos del Golfo de California VI. Alpheidae del estuario de Mulegé y de Bahía Concepción, Baja California Sur, México (Crustacea: Caridea). Proceedings of the San Diego Society of Natural History, 14, 1–13.
- Ríos, R. & Carvacho, A. (1983) Caridean shrimps of the Gulf of California. III. Leptalpheus mexicanus, new species (Crustacea, Decapoda, Alpheidae). Journal of Crustacean Biology, 3 (2), 306–313. https://doi.org/10.2307/1548265
- Ríos, R. & Duffy, J.E. (2007) A review of the sponge-dwelling snapping shrimp from Carrie Bow Cay, Belize, with description of Zuzalpheus, new genus, and six new species (Crustacea: Decapoda: Alpheidae). Zootaxa, 1602 (1), 1–94. https://doi.org/10.11646/zootaxa.1602.1.1
- Riul, P., Rodrigues, F.M.A., Xavier-Filho, E.S., Santos, R.G., Leonel, R.M.V. & Christoffersen, M.L. (2008) Macrocrustaceans from Ponta do Cabo Branco, João Pessoa, Paraíba, Brazil, the easternmost point of South America. Revista Nordestina de Biologia, 19, 3–13.
- Robles, R. (2005) Regional speciation among selected groups of marine nektonic, fossorial, and commensal decapod crustaceans (Portunoidea, Thalassinidea, and Alpheoidea). Ph.D. Thesis, University of Louisiana at Lafayette, Lafayette, 155 pp.
- Robles, R., Dworschak, P.C., Felder, D.L., Poore, G.C.B. & Mantelatto, F.L. (2020) A new molecular phylogeny of the Callianassoidea (Crustacea: Decapoda: Axiidea) with morphological support. Invertebrate Systematics, 34, 113–132. https://doi.org/10.1071/IS19021
- Rodrigues, S.A. & Shimizu, R.M. (1992) Description of a new Axianassa (Crustacea: Decapoda: Thalassinidea) from Brazil, and its first larval stage. Proceedings of the Biological Society of Washington, 105, 317–323.
- Rosário, T.N., Pires, M.A.B., Souza, A.S., Fernandes, M.E.B., Abrunhosa, F.A. & Simith, D.J.B. (2020) First known occurrence of the alpheid shrimps Leptalpheus forceps Williams, 1965 and L. marginalis Anker, 2011 (Decapoda: Caridea) in the state of Pará, Amazon region, Brazil. Zootaxa, 4766 (1), 61–85. https://doi.org/10.11646/zootaxa.4766.1.3
- Sakai, K. (2005) Callianassoidea of the world (Decapoda: Thalassinidea. Crustaceana Monographs, 4, 1–285. https://doi.org/10.1163/9789047416890
- Salgado-Barragán, J., Ayón-Parente, M. & Hendrickx, M.E. (2014) A new species of Leptalpheus Williams, 1965 and new records of L. mexicanus Ríos & Carvacho, 1983 and L. hendrickxi Anker, 2011 (Crustacea: Decapoda: Alpheidae) from the Pacific coast of Mexico. Zootaxa, 3835 (4), 573–582. https://doi.org/10.11646/zootaxa.3835.4.8
- Salgado-Barragán, J., Ayón-Parente, M. & Zamora-Tavares, P. (2017) New records and description of two new species of carideans shrimps from Bahía Santa María-La Reforma lagoon, Gulf of California, Mexico (Crustacea, Caridea, Alpheidae and Processidae). ZooKeys, 671, 131–153. https://doi.org/10.3897/zookeys.671.9081
- Saloman, C.H. (1971) The shrimp Leptalpheus forceps in Old Tampa Bay, Florida. Quarterly Journal of the Florida Academy of Sciences, 34 (1), 67–77.
- Say, T. (1818) An account of the Crustacea of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 1 (2), 235–253 + 313–319 + 374–401 + 423–441.
- Schmidt de Melo, G.A., Vezzani, R.M. & Campos, O. Jr. (2003) Type catalogue of the Crustacea Decapoda in the collections of the Museu de Zoologia da Universidade de São Paulo, Brazil. Proceedings of the Biological Society of Washington, 116 (2), 423–437.
- Schmitt, W.L. (1935) Mud shrimps of the Atlantic coast of North America. Smithsonian Miscellaneous Contributions, 93, 1–21.
- Schmitt, W.L. (1936) Zoologische Ergebnisse einer Reise nach Bonaire, Curaçao und Aruba im Jahre 1930. No. 16. Macruran and anomuran Crustacea from Bonaire, Curaçao und Aruba. Zoologische Jahrbücher, Abtheilung für Systematik, Geographie und Biologie der Thiere, 67, 363–378, pls. 11–13.
- Schubart, C.D., Cuesta, J.A. & Felder, D.L. (2002) Glyptograpsidae, a new brachyuran family from Central America: larval and adult morphology, and a molecular phylogeny of the Grapsoidea. Journal of Crustacean Biology, 22 (1), 28–44. https://doi.org/10.1163/20021975-99990206
- Scioli, J.A., Plouviez, S. & Felder, D.L. (2020) The complete mitochondrial genome of the symbiotic infaunal snapping shrimp Leptalpheus forceps (Decapoda, Alpheidae). Mitochondrial DNA, Part B, 5 (1), 629–630. https://doi.org/10.1080/23802359.2019.1704650
- Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30 (9), 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Stimpson, W. (1866) Descriptions of new genera and species of Macrurous Crustacea from the coasts of North America. Proceedings of the Chicago Academy of Sciences, 1, 46–48.
- Vera Caripe, J., Pereda, L. & Anker, A. (2021) A new species and two new records of symbiotic infaunal alpheid shrimps from the genera Leptalpheus Williams, 1965 and Fenneralpheus Felder & Manning, 1986 (Decapoda: Caridea) from Venezuela. Zootaxa, 5061 (1), 177–184. https://doi.org/10.11646/zootaxa.5061.1.10
- Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics, 27 (2), 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x
- Williams, A.B. (1965) A new genus and species of snapping shrimp (Decapoda, Alpheidae) from the southeastern United States. Crustaceana, 9, 192–198. https://doi.org/10.1163/156854065X00352
- Williams A.B. (1984) Shrimps, lobsters and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C., 550 pp.