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Richness and large-scale distribution of marine benthic caridean shrimps (Decapoda: Caridea) from the Eastern Tropical Pacific

Abstract

Based on an updated checklist of the benthic caridean shrimp fauna of the Eastern Tropical Pacific (ETP), we analyzed their large-scale species richness distribution patterns using a grid approximation. Caridean fauna is composed of 183 species belonging to 67 genera and 18 families. Alpheidae, Palaemonidae and Thoridae contributed largely to species richness. Alpheus and Synalpheus were the most diverse genera. Most species (24%) have a narrow distribution inhabiting a single grid, and as many as 135 are restricted to < 5 grids. Mexico has the largest richness (130) besides Ecuador (98) and Panama (75). Richness among grids varies largely; overall, it is larger around Galapagos and Panama, Colombia, Costa Rica and the Gulf of California. There was a meaningful correlation between a proxy of sampling intensity and richness, suggesting that along with habitat heterogeneity or large-scale evolutionary processes currently invoked as richness drivers, the pattern can also be related to differences in research efforts. Exponential fitting and Chao2 index suggest that the caridean faunal inventory from the ETP is still far from being complete. Overall, the results call for a more thoughtful and systematic sampling scheme in order to get information from poorly sampled environments and areas in the ETP.

Key Words
Checklist; faunal gap; species rarity; latitudinal patterns

INTRODUCTION

Shrimps included in the infraorder Caridea are the most diverse group of decapod shrimps (Gorny, 1999Gorny, M. 1999. On the biogeography and ecology of the southern ocean decapod fauna. Scientia Marina, 63: 367-382.; De Grave and Fransen, 2011De Grave, S. and Fransen, C. 2011. Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededeelingen, 85: 195-589.). Caridean shrimps are mostly benthic organisms inhabiting muddy to rocky bottoms from a wide diversity of habitats (e.g., tidal pools, seaweeds), either free or associated to other marine organisms (Wicksten, 1983Wicksten, M.K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico. Allan Hancock Monographs in Marine Biology, 13: 1-59.; Bauer, 2004Bauer, R.T. 2004. Remarkable shrimps-adaptations and natural history of the carideans. Oklahoma, Ed. University of Oklahoma Press, 316p.; Guzmán, 2008Guzmán, G.G. 2008. Camarones pelágicos (Crustacea, Decapoda) en aguas del Pacifico Sureste. p. 27-45. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 5. Mexico, ICMyL, UNAM. ). They can be found from intertidal to hydrothermal vents, and in freshwater, estuarine and marine habitats worldwide, from the equator to the poles but mainly inhabiting tropical waters (Carvacho and Ríos, 1982Carvacho, A. and Ríos, R. 1982. Los camarones carideos del Golfo de California II. Catálogo, claves de identificación y discusión biogeográfica. Anales del Instituto de Ciencias del Mar y Limnología, 9: 279-294.; Martin, 2003Martin, J.W. 2003. Oplophorid shrimp (Decapoda: Caridea) from an arctic hydrothermal vent. Crustaceana, 76: 871-878.; Anker et al., 2006aAnker, A.; Poddoubtchenko, D. and Wehrtmann, I.S. 2006a. Leslibetaeus coibita, n. gen., n. sp., a new alpheid shrimp from the Pacific coast of Panama (Crustacea: Decapoda). Zootaxa, 1183: 27-41.).

Caridean shrimp studies in the Eastern Tropical Pacific (ETP) started as early as 1869 in Panama (Smith, 1869Smith, S.I. 1869. Descriptions of new genera and species of Crustacea. p. 239-250. In: A.E. Verrill (ed), On the parasitic habits of some Crustacea. American Naturalist, Vol. 3. Chicago, University of Chicago Press.) and studies in the Pacific coast of Mexico and Ecuador contributed steadily via the discrete addition of new species to the regional fauna (Faxon, 1893Faxon, W. 1893. Reports on the dredging off the west coast of Central America to the Galapagos, to the west of Mexico, and in the gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Comission Steamer "Albatross”, during 1891, Lieut.-Commander Z.L. Tanner, U.S.N., Commanding. Bulletin of the Museum of Comparative Zoology, 24: 150-220.; 1896Faxon, W. 1896. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico and Caribbean Sea, and on the east coast of United States, 1877-1880, by the U.S. Coast Survey steamer "Blake” 37- Supplementary notes on the Crustacea. Bulletin of the Museum of Comparative Zoology, 30: 153-166.). This early trend was occasionally disrupted by large-scale works where the caridean taxonomy of large areas (Coutière, 1909Coutière, H. 1909. The american species of snapping shrimps of the genus Synalpheus. Proceedings of the United States Natural Museum, 36: 1-93.) or entire supraspecific groups (Holthuis, 1951Holthuis, L.B. 1951. A general revision of the Palaemonidae (Crustacea: Decapoda: Natantia) of the Americas I. The subfamilies Euryrhynchinae and Pontoniinae. Allan Hancock Foundation Occasional Papers, 11: 1-332.; 1952Holthuis, L.B. 1952. A general revision of the Palaemonidae (Crustacea: Decapoda: Natantia) of the Americas. II. The subfamily Palaemoninae. Allan Hancock Foundation Occasional Papers, 12: 1-396.; Kim and Abele, 1988Kim, W. and Abele, L.G. 1988. The snapping shrimp genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheida). Smithsonian Contribution to Zoology, 454: 1-119.) were addressed. Recently, as a result of studies conducted by local taxonomists (Abele and Kim, 1989Abele, L.G. and Kim, W. 1989. The decapods crustaceans of the Panama canal. Smithsonian Contributions in Zoology, 482: 1-50.; Wicksten, 1991Wicksten, M.K. 1991. Caridean and stenopodid shrimp of the Galapagos Islands. p. 147-256. In: H.J. James (ed), Galapagos marine Invertebrates: Taxonomy, biogeography and evolution Darwin’s islands. New York, Plenum Press. ; Li and Poupin, 2009Li, X. and Poupin, J. 2009. Report on some species of Palaemonidae (Decapoda: Caridea) from Clipperton island. Journal of Crustacean Biology, 29: 579-594.) and ecologists (Hernández et al., 2013), there has been an increase in the knowledge of the caridean shrimp fauna from the ETP.

The most recent checklist of the benthic caridean shrimp fauna from the ETP was published by Wicksten and Hendrickx (2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.). Since then, species descriptions (Wicksten and Martin, 2004Wicksten, M.K. and Martin, J.W. 2004. A new species of caridean shrimp of the family Stylodactylidae from the Eastern Pacific Ocean. Proceedings of the Biological Society of Washington, 117: 337-384.; Hermoso-Salazar and Álvarez, 2005Hermoso-Salazar, A.M. and Hendrickx, M.E. 2005. New records for Synalpheus peruvianus Rathbun, 1910 (Decapoda: Caridea: Alpheidae) in the East Pacific. Crustaceana, 78: 763-765.; Anker et al., 2006aAnker, A.; Poddoubtchenko, D. and Wehrtmann, I.S. 2006a. Leslibetaeus coibita, n. gen., n. sp., a new alpheid shrimp from the Pacific coast of Panama (Crustacea: Decapoda). Zootaxa, 1183: 27-41.; 2007Anker, A.; Hurt, C. and Knowlton, N. 2007. Revision of the Alpheus nuttingi (Schmitt) species complex (Crustacea: Decapoda: Alpheidae), with description of a new species from the Eastern Tropical Pacific. Zootaxa, 1577: 41-60.; 2009Anker, A.; Baeza, A.J. and De Grave, S. 2009. A new species of Lysmata (Crustacea: Decapoda: Hippolytidae) from the Pacific coast of Panama, with observations of its reproductive biology. Zoological Studies, 48: 682-692.; Hermoso-Salazar and Hendrickx, 2006Hermoso-Salazar, A.M. and Hendrickx, M.E. 2006. Two new species of Synalpheus Bate, 1888 (Decapoda: Caridea: Alpheidae) from the SE Gulf of California, Mexico. Crustaceana, 78: 1099-1116.; Li, 2006Li, X. 2006. Chacella mclaughlinae n. sp., a new pontoniine shrimp from Clipperton Island (Crustacea, Caridea, Palaemonidae). Zoosystema, 28: 359-366.; Marin and Anker, 2008Marin, I. and Anker, A. 2008. A new species of Pontonia Latreille, 1829 (Crustacea: Decapoda: Palaemonidae) associated with sea squirts (Tunicata: Ascidiacea) from the Pacific coast of Panama. Zoosystema, 30: 501-515.; Li and Poupin, 2009Li, X. and Poupin, J. 2009. Report on some species of Palaemonidae (Decapoda: Caridea) from Clipperton island. Journal of Crustacean Biology, 29: 579-594.; Hendrickx, 2010Hendrickx, M.E. 2010. A new species of Glyphocrangon (Decapoda: Caridea: Glyphocrangonidae) from off the coast of western Mexico. Zootaxa, 2372: 358-366.), geographic range extension (Hermoso-Salazar and Hendrickx, 2005Hermoso-Salazar, A.M. and Hendrickx, M.E. 2005. New records for Synalpheus peruvianus Rathbun, 1910 (Decapoda: Caridea: Alpheidae) in the East Pacific. Crustaceana, 78: 763-765.), and changes in taxonomy and systematics (Marin, 2009Marin, I. 2009. Sandyella gen. nov., a new shrimp genus for the Eastern Pacific species Chacella tricornuta Hendrickx, 1990 and Chacella mclaughlinae Li, 2006 (Caridea: Palaemonidae: Pontoniinae). Crustaceana, 82: 913-918.; Bracken et al., 2010Bracken, H.D.; De Grave, S. and Felder, D.L. 2010. Phylogeny of the infraorder Caridea based on mithochondrial and nuclear genes (Crustacea: Decapoda). p. 281-305. In: S. Koenemann, S.; Martin, J.W.; Crandall, K.A, and Felder, D.L. (eds), Decapod crustacean phylogenetics. Crustacean Issues, Vol. 18. Florida, CRC Press. ) have substantially modified the previously known information concerning caridean shrimp faunal inventory and its distribution patterns in the area. Current distribution patterns of ETP caridean shrimps were known after studies focused on more inclusive groups such as macroinvertebrates, crustaceans or decapods (Arntz et al., 1999Arntz, W.E.; Wehrtmann, I.S.; Gorny, M.; Soto, R; Lardies, M. and Retamal, R. 1999. Species composition and distribution of decapod crustaceans in the waters off Patagonia and Tierra del Fuego, South America. Scientia Marina, 63: 303-314.; Blanchette et al., 2008Blanchette, C.A.; Miner, C.M.; Raimondi, P.T.; Lohse, D.; Heady, K.E.K. and Brotman, B.R. 2008. Biogeographical patterns of rocky intertidal communities along the Pacific coast of North America. Journal of Biogeography, 35: 1593-1607.) and at large-scale levels (i.e., Province) (Boschi, 2000Boschi, E.E. 2000. Species of decapods crustaceans and their distribution in the American zoogeographic provinces. Revista de Investigacion y Desarrollo Pesquero, 13: 7-136.; Wicksten and Hendrickx, 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.). For example, Boschi (2000Boschi, E.E. 2000. Species of decapods crustaceans and their distribution in the American zoogeographic provinces. Revista de Investigacion y Desarrollo Pesquero, 13: 7-136.) indicates that the so called Panamic province hosts the largest number of both planktonic and benthonic caridean species (161), followed by the Galapagos (67 species) and Cortes (50 species) provinces. Additionally, the same general pattern emerges when only benthonic caridean species are considered (Wicksten and Hendrickx, 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.). While there is an overall agreement among studies regarding richness patterns, the previously used spatial scale precludes direct comparisons among areas which obscure intraregional species richness patterns. Beside taxonomic bias, databases resulting from field exploration projects and/or previously published information allow for large-scale analyses that evaluate current species richness and distribution scenarios (Costello et al., 2010Costello, M.J.; Coll, M.; Danovaro, R.; Halpin, P.; Ojaveer, H. and Miloslavich, P. 2010. A Census of Marine Biodiversity Knowledge, Resources, and Future Challenges. PLoS ONE, 5: e12110.). In this regard, there is a need to integrate previously published findings such as spatially explicit checklist, to gain perspective on what is known and what are the major scientific gaps regarding benthic caridean shrimp species richness and distribution in the ETP.

In this paper, species composition and distribution of the benthic caridean shrimp fauna of the Eastern Tropical Pacific are analyzed. An updated (up to 2017) checklist of the benthic caridean shrimp fauna from the ETP, at a 1° of latitude by 1° longitude, was assembled and further analyzed to address a set of interrelated questions. What is the richness and composition of the ETP benthic caridean shrimp fauna? Has the accumulative historic species record been homogeneous over the entire region or do discrepancies exist among areas? Does species richness show any distribution pattern in the area? Are ETP benthic caridean shrimp species widely or narrowly distributed in the area? Finally, how complete is the benthic caridean shrimp fauna inventory?

MATERIAL AND METHODS

We compiled a checklist of the marine benthic caridean shrimps that have been collected and/or recorded in the coastal and shelf area of the Eastern Tropical Pacific. A database was generated from primary references that have been formally published as papers, monographs and books (mainly, but not restricted to Kingsley, 1878Kingsley, J.S. 1878. A synopsis of the north American species of the genus Alpheus. Bulletin of the United States Geological and Geographical Survey, 4: 189-199.; Faxon, 1893Faxon, W. 1893. Reports on the dredging off the west coast of Central America to the Galapagos, to the west of Mexico, and in the gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Comission Steamer "Albatross”, during 1891, Lieut.-Commander Z.L. Tanner, U.S.N., Commanding. Bulletin of the Museum of Comparative Zoology, 24: 150-220.; Rathbun, 1904Rathbun, M.J. 1904. Decapod crustaceans of the northwest coast of North America. p. 1-210. In: M.J. Rathbun, H. Richardson, S.J. Holmes and L.J. Cole (eds), Harriman Alaska Expedition with cooperation of Washington Academy of Sciences, Vol. 10. New York, Doubleday, Page and Co. ; Coutière, 1909Coutière, H. 1909. The american species of snapping shrimps of the genus Synalpheus. Proceedings of the United States Natural Museum, 36: 1-93.; Chace, 1937Chace, F.A. 1937. The Templeton Crocker Expedition. VII. Caridean decapods crustacean from the Gulf of California and the west coast of Lower California. Zoologica, 22: 109-138.; 1962Chace, F.A. 1962. The non-brachyuran decapods crustaceans of Clipperton Island. Proceedings of the United States Natural Museum, 113: 605-635.; Holthuis, 1951Holthuis, L.B. 1951. A general revision of the Palaemonidae (Crustacea: Decapoda: Natantia) of the Americas I. The subfamilies Euryrhynchinae and Pontoniinae. Allan Hancock Foundation Occasional Papers, 11: 1-332.; 1952Holthuis, L.B. 1952. A general revision of the Palaemonidae (Crustacea: Decapoda: Natantia) of the Americas. II. The subfamily Palaemoninae. Allan Hancock Foundation Occasional Papers, 12: 1-396.; Abele, 1975Abele, L.G. 1975. The macruran decapods crustaceans of Malpelo Island. Smithsonian Contributions in Zoology, 176: 69-85.; Abele and Patton, 1976Abele, L.G. and Patton, W.K. 1976. The size of coral heads and the community biology of associated decapods crustaceans. Journal of Biogeography, 3: 35-47.; Wicksten, 1983Wicksten, M.K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico. Allan Hancock Monographs in Marine Biology, 13: 1-59.; 1991Wicksten, M.K. 1991. Caridean and stenopodid shrimp of the Galapagos Islands. p. 147-256. In: H.J. James (ed), Galapagos marine Invertebrates: Taxonomy, biogeography and evolution Darwin’s islands. New York, Plenum Press. ; 2000Wicksten, M.K. 2000. A new species of Lysmata (Caridea, Hippolytidae) from the eastern Pacific. Crustaceana, 73: 207-213.; Kim and Abele, 1988Kim, W. and Abele, L.G. 1988. The snapping shrimp genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheida). Smithsonian Contribution to Zoology, 454: 1-119.; Abele and Kim, 1989Abele, L.G. and Kim, W. 1989. The decapods crustaceans of the Panama canal. Smithsonian Contributions in Zoology, 482: 1-50.; Lemaitre and Álvarez-León, 1992Lemaitre, R. and Álvarez-León, R. 1992. Crustáceos decápodos del Pacífico colombiano: lista de species y consideraciones zoogeográficas. Anales del Instituto de Investigaciones Marinas de Punta de Betin, 21: 33-76.; Wicksten and Hendrickx, 1992Wicksten, M.K. and Hendrickx, M.E. 1992. Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea: Caridea) from the Eastern Tropical Pacific. Proceedings of the San Diego Society of Natural History, 9: 1-11.; 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.; Burukovsky, 2001Burukovsky, R.N. 2001. Taxonomy of Nematocarcinus (Decapoda: Nematocarcinidae). Description of Nematocarcinus from waters of the American Continent. Journal of Zoology, 80: 1429-1443.; complete reference list is included in Tab. 1). In order to minimize taxonomic bias, only valid species were included, hence all taxa names were crosschecked in the World Register of Marine Species (WoRMS) and Integrated Taxonomic Information System (ITIS). Systematic arrangement of benthic caridean shrimps follows De Grave and Fransen (2011De Grave, S. and Fransen, C. 2011. Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededeelingen, 85: 195-589.). The taxa (genera and species) were arranged alphabetically (Tab. 1).

Table 1
Checklist of marine benthic caridean shrimps from the Tropical Eastern Pacific. Systematic arrangement follows De Grave and Fransen (2011De Grave, S. and Fransen, C. 2011. Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededeelingen, 85: 195-589.). Mexico-Revillagigedo: MEX, Clipperton: CLP, El Salvador: SAL, Nicaragua: NIC, Costa Rica: CR, Panama: PAN, Colombia-Malpelo: COL, Ecuador-Galapagos: ECU, Peru: PER, Central Pacific: PC, Indopacific: IP, Indian: IND, Atlantic: ATL, s: shallow.

After checking for species validity, historic species accumulation curves (species added by year of publication) were constructed for each country and for the entire ETP. Further, the data was fitted with an exponential function (larger r2).

The occurrence of each taxon was mapped on the ETP. When there was a lack of geographic information of the species record or the accuracy of the data was larger than 1° of latitude by 1° longitude, the information was discarded. Once the distribution of each species was mapped on the ETP, we traced a grid of 1° of latitude by 1° longitude and obtained the species richness per grid. Since not all records have accurate geographic data, the amount of information included in the following analysis is slightly reduced (~0.03%) than the published information. Nonetheless, mapping of species distribution was a methodological prerequisite to generate an accurate picture of the knowledge and gaps of the marine biodiversity of the benthic caridean shrimp species in the ETP. In addition, occurrence of spatially rare (inhabiting 1 to 2 grids) species in the area was also mapped. After mapping, we fitted species richness and species richness of rare species (inhabiting 1 to 2 grids) against latitude through a linear model. Additionally, we related the number of caridean species against the number of caridean references (proxy for sampling intensity) per grid with a Spearman Rank correlation.

We used the species occurrence per grid to generate sample-based rarefaction curves of observed species and calculated the species richness estimator Chao2 for the ETP (Colwell, 2009Colwell, R.K. 2009. EstimateS, Version 7.5: statistical estimation of species richness and shared species from samples. Available at Available at http://viceroy.eeb.uconn.edu/estimates . Accessed on: 16 December 2017.
http://viceroy.eeb.uconn.edu/estimates...
). While the former analysis indicates the observed species accumulation curve, the latter is a sample-based nonparametric estimator for addressing expected species richness of the benthic caridean shrimp in the Eastern Tropical Pacific. EstimateS (Colwell, 2009Colwell, R.K. 2009. EstimateS, Version 7.5: statistical estimation of species richness and shared species from samples. Available at Available at http://viceroy.eeb.uconn.edu/estimates . Accessed on: 16 December 2017.
http://viceroy.eeb.uconn.edu/estimates...
) were used to compute observed and expected rarefaction curves and their confidence intervals (95%) after 10,000 randomizations with replacement.

RESULTS

The present checklist of the marine benthic caridean shrimp of the Eastern Tropical Pacific includes 183 species considered as currently valid, belonging to 67 genera, 18 families and 9 superfamilies (Tab. 1). According to the data, families Alpheidae (66 spp.), Palaemonidae (51 spp.) and Thoridae (12 spp.) contribute to most of the species (>70%) (Fig. 1A), while Alpheus (41 species) and Synalpheus (16 species) were the most diverse genera of benthic caridean shrimps in the ETP (Fig. 1B).

Figure 1
Richness of marine benthic caridean shrimps in the Eastern Tropical Pacific. A, Species per family (n = 183). B, Species per genera of Alpheidae (n = 160).

México (including Revillagigedo Archipelago) is the country with the greatest number of benthic caridean species (130 species) (Fig. 2A), followed by Ecuador (including Galapagos Archipelago) (98 species) (Fig. 2B), Panama (75 species) (Fig. 2C), Colombia (including Malpelo Island) (65 species) (Fig. 2D), Costa Rica (including Cocos Island) (48 species) (Fig. 2E), Peru (20 species) (Fig. 2F) and France (Clipperton Atoll) (19 species) (Fig. 2G);. Meanwhile, there are only two records in El Salvador [Alpheus galapagensis Sivertsen, 1933 addressed in La Unión [Kim and Abele, 1988Kim, W. and Abele, L.G. 1988. The snapping shrimp genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheida). Smithsonian Contribution to Zoology, 454: 1-119.], and A. panamensisKingsley, 1878Kingsley, J.S. 1878. A synopsis of the north American species of the genus Alpheus. Bulletin of the United States Geological and Geographical Survey, 4: 189-199. founded in Acajutla [Wicksten, 1994Wicksten, M.K. 1994. On the identity of snapping shrimps described and identified by W. N. Lockington, 1878. Bulletin of the Southern California Academy of Science, 93: 118-126.]), and two more in Nicaragua (Palaemon gracilis (Smith, 1871) recorded in Realejo [Wicksten, 1983Wicksten, M.K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico. Allan Hancock Monographs in Marine Biology, 13: 1-59.] and Ogyrides tarazonaiWicksten and Mendez, 1988 Wicksten, M.K. and Méndez, M. 1988. New records for Ogyrides alphaerostris and a new species, Ogyrides tarazonai (Crustacea: Ogyrididae), from the eastern Pacific Ocean. Proceedings of the Biological Society of Washington, 101: 622-625.in San Juan del Sur (Wicksten and Mendez, 1988Wicksten, M.K. and Méndez, M. 1988. New records for Ogyrides alphaerostris and a new species, Ogyrides tarazonai (Crustacea: Ogyrididae), from the eastern Pacific Ocean. Proceedings of the Biological Society of Washington, 101: 622-625.)). To date, no marine benthic caridean species have ever been recorded in Guatemala or Honduras.

Figure 2
Temporal accumulation species curves of marine benthic caridean shrimps in the Eastern Tropical Pacific. A, Mexico (Mexico-Revillagigedo Archipelago). B, Clipperton Atoll. C, Ecuador (Ecuador-Galapagos Archipelago). D, Costa Rica (Costa Rica-Cocos Island). E, Panama. F, Colombia (Colombia-Malpelo Island). G, Peru. H, Tropical Eastern Pacific. Dotted line= observed values, continuous line= exponential fit.

Historic accumulation curves varied along the ETP, nevertheless, some results were similar among countries (Fig. 2). For example, for countries such as Mexico (Fig. 2A), Ecuador (Fig. 2B) and Panama (Fig. 2C), small but steady early increases were followed by sudden increments in species richness. Likewise, the same trend is recorded when data are pooled across the ETP (Fig. 2H). The observed trend was meaningfully fitted with an exponential function, indicating that during the last three decades many more species records were added to the benthic caridean fauna than those recorded in the ETP during the previous 110 years. Function fitting also indicates that the current increment of species records still not leveling off for any particular country neither for the entire ETP (Fig. 2).

According to species mapping, richness varies largely among grids in the ETP (Fig. 3). Except for a few empty spots in the northern Gulf of California, the Tropical Mexican Pacific, Guatemala and Honduras, marine benthic caridean shrimp have been recorded in the coastal margin of the entire ETP and its oceanic islands (i.e., Revillagigedo, Clipperton, Malpelo, Cocos, Galapagos). Species richness is concentrated in grids located around Galapagos and Panama (i.e., Central America), but relatively rich spots can also be observed in Colombia, Costa Rica and the Gulf of California, hence disrupting any meaningful trend between richness and latitude (r2 = 0.002, n = 37, p> 0.05).

Figure 3
Species richness of marine benthic caridean shrimps per grid (1° latitude by 1° longitude) in the Eastern Tropical Pacific. Notice color scale.

Data indicates that up to 44 species (24%) inhabit a single grid and hence may be considered spatially rare, but as many as 135 species (~74%) are restricted to less than five grids each. In contrast, just 22 species (12%) inhabit more than 10 grids and hence may be considered as spatially common. The species Processa peruviana Wicksten, 1983 (15 grids), Harpiliopsis depressa (Stimpson, 1860) (17 grids), Alpheus hebes Kim and Abele, 1988 and Palaemonella holmesi (Nobili, 1907) (18 grids), Brachycarpus biunguiculatus (Lucas, 1849) (19 grids), and Alpheus bellimanus Lockington, 1877 (21 grids) are the most widespread distributed species in the ETP.

Rare species, inhabiting one to two grids, concentrated around Central America (Costa Rica, Panama, Colombia and Galapagos Islands) and the entrance of the Gulf of California, Mexico; however, from northern Costa Rica (~11°N) up to the Central Mexican Pacific (~19°N) spatially rare species are lacking (Fig. 4). Overall, linear regression models indicates no latitudinal trend (r2 = 0.001-0.004) in the number of rare species (i.e., inhabiting 1 to 2 grids). Nevertheless, grids around Clipperton Atoll (10°N 109°W) (5 species), Galapagos Islands (0° 90°W), entrance of the Gulf of California (24°N 110°W) (7 species) and Panama (8°N 79°W) (10 species) have relatively large number of rare species.

Figure 4
Species richness of narrow-distributed (1-2 grids) marine benthic caridean shrimps per grid (1° latitude by 1° longitude) in the Eastern Tropical Pacific. Notice color scale.

Finally, the non-parametric Chao2 species estimator indicates that the observed species richness is lower than expected (Fig. 5), in fact with a 95% confidence the model suggests that the expected species richness of marine benthic caridean shrimp in the ETP may lay somewhere between 202-269 species, a number 11%-47% larger than the observed.

Figure 5
Species richness of marine benthic caridean shrimps in Eastern Tropical Pacific. Orange continuous line = observed richness, blue continuous line = expected richness, blue long-cut lines = confidence interval (95%). Expected species number and confidence intervals (95%) were generated after 10,000 randomizations with replacement using non-parametric Chao2 estimator.

DISCUSSION

As a result of recent species descriptions (Wicksten and Martin, 2004Wicksten, M.K. and Martin, J.W. 2004. A new species of caridean shrimp of the family Stylodactylidae from the Eastern Pacific Ocean. Proceedings of the Biological Society of Washington, 117: 337-384.; Hermoso-Salazar and Álvarez, 2005Hermoso-Salazar, A.M. and Hendrickx, M.E. 2005. New records for Synalpheus peruvianus Rathbun, 1910 (Decapoda: Caridea: Alpheidae) in the East Pacific. Crustaceana, 78: 763-765.; Anker et al., 2006aAnker, A.; Poddoubtchenko, D. and Wehrtmann, I.S. 2006a. Leslibetaeus coibita, n. gen., n. sp., a new alpheid shrimp from the Pacific coast of Panama (Crustacea: Decapoda). Zootaxa, 1183: 27-41.; 2007Anker, A.; Hurt, C. and Knowlton, N. 2007. Revision of the Alpheus nuttingi (Schmitt) species complex (Crustacea: Decapoda: Alpheidae), with description of a new species from the Eastern Tropical Pacific. Zootaxa, 1577: 41-60.; 2009Anker, A.; Baeza, A.J. and De Grave, S. 2009. A new species of Lysmata (Crustacea: Decapoda: Hippolytidae) from the Pacific coast of Panama, with observations of its reproductive biology. Zoological Studies, 48: 682-692.; Hermoso-Salazar and Hendrickx, 2006Hermoso-Salazar, A.M. and Hendrickx, M.E. 2006. Two new species of Synalpheus Bate, 1888 (Decapoda: Caridea: Alpheidae) from the SE Gulf of California, Mexico. Crustaceana, 78: 1099-1116.; Li, 2006Li, X. 2006. Chacella mclaughlinae n. sp., a new pontoniine shrimp from Clipperton Island (Crustacea, Caridea, Palaemonidae). Zoosystema, 28: 359-366.; Marin and Anker, 2008Marin, I. and Anker, A. 2008. A new species of Pontonia Latreille, 1829 (Crustacea: Decapoda: Palaemonidae) associated with sea squirts (Tunicata: Ascidiacea) from the Pacific coast of Panama. Zoosystema, 30: 501-515.; Li and Poupin, 2009Li, X. and Poupin, J. 2009. Report on some species of Palaemonidae (Decapoda: Caridea) from Clipperton island. Journal of Crustacean Biology, 29: 579-594.; Hendrickx, 2010Hendrickx, M.E. 2010. A new species of Glyphocrangon (Decapoda: Caridea: Glyphocrangonidae) from off the coast of western Mexico. Zootaxa, 2372: 358-366.), geographic range extensions (Martínez-Guerrero and Hendrickx, 2011Martínez-Guerrero, B. and Hendrickx, M. E. 2011. Four new records of caridean shrimps (Decapoda: Palaemonidae, Alpheidae) in Mexican Pacific waters. Crustaceana, 84: 1111-1115.) and new records of previously recorded species (Ríos et al., 2003Ríos, C.; Mutschke, E. and Morrison, E. 2003. Biodiversidad bentónica sublitoral en el estrecho de Magallanes, Chile. Revista de Biologia Marina y Oceanografia, 38: 1-12.; Wicksten and Hendrickx, 1992Wicksten, M.K. and Hendrickx, M.E. 1992. Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea: Caridea) from the Eastern Tropical Pacific. Proceedings of the San Diego Society of Natural History, 9: 1-11.; 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.; Poupin, 2008Poupin, J. 2008. Biogeography of the decapod and stomatopod crustacean of the Tropical Pacific: issues and prospects. Pacific Science, 62: 377-383.), we assembled a list of 183 species of caridean regarded as currently valid in the Eastern Tropical Pacific. The number of species is slightly smaller compared with similar but previous studies (188 species, Wicksten and Hendrickx, 1992Wicksten, M.K. and Hendrickx, M.E. 1992. Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea: Caridea) from the Eastern Tropical Pacific. Proceedings of the San Diego Society of Natural History, 9: 1-11.; 204 species, Wicksten and Hendrickx, 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.). The observed differences in species numbers among studies are mainly due to the exclusion of 30 species, either because they have a freshwater affinity (Hendrickx and Estrada-Navarrete, 1989Hendrickx, M.E. and Estrada-Navarrete, F.D. 1989. A check list of the species of pelagic shrimps (Penaeoidea and Caridea) from the Eastern Pacific, with notes on their geographic and depth distribution. California Cooperative Oceanic Fisheries Investigation Report, 30: 104-121.; Guzmán, 2008Guzmán, G.G. 2008. Camarones pelágicos (Crustacea, Decapoda) en aguas del Pacifico Sureste. p. 27-45. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 5. Mexico, ICMyL, UNAM. ), distribute outside of the ETP (Cardoso and Serejo, 2007Cardoso, I.A. and Serejo, C.S. 2007. Deep sea Caridea (Crustacea, Decapoda) from Campos Basin, R J, Brazil. Brazilian Journal of Oceanography, 55: 39-50.; Anker et al., 2008Anker, A.; Hurt, C. and Knowlton, N. 2008. Revision of the Alpheus cristulifrons species complex (Crustacea: Decapoda: Alpheidae), with description of a new species from the Tropical Eastern Atlantic.Journal of the Marine Biological Association of the United Kingdom, 88: 543-562.), or have an invalid taxonomic status (nomen dubium).

Data revealed that families Alpheidae and Palaemonidae, and genus Alpheus contributed to most of the species. Dominance of the former two families was previously recognized by Wicksten and Hendrickx (1992Wicksten, M.K. and Hendrickx, M.E. 1992. Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea: Caridea) from the Eastern Tropical Pacific. Proceedings of the San Diego Society of Natural History, 9: 1-11.) and Anker et al. (2006bAnker, A.; Ahyong, S.T.; Noël, P.Y. and Palmer, A.R. 2006b. Morphological phylogeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw. Evolution, 60: 2507-2528.). The trend, however, is not exclusive to the ETP but it is also observed in the Caribbean and the Indo West Pacific (Boschi, 2000Boschi, E.E. 2000. Species of decapods crustaceans and their distribution in the American zoogeographic provinces. Revista de Investigacion y Desarrollo Pesquero, 13: 7-136.). Bauer (2004Bauer, R.T. 2004. Remarkable shrimps-adaptations and natural history of the carideans. Oklahoma, Ed. University of Oklahoma Press, 316p.) and Poupin (2008Poupin, J. 2008. Biogeography of the decapod and stomatopod crustacean of the Tropical Pacific: issues and prospects. Pacific Science, 62: 377-383.) have suggested that Alpheidae and Palaemonidae dominance is related to its ability to inhabit a large number of substrates, its reduced size and the capacity of its individuals to establish symbiotic relations with vertebrates or other invertebrates. In other invertebrate and vertebrate taxa such as ophiuroids (Granja-Fernández et al., 2014Granja-Fernández, R.; Herrero-Pérezrul, D.; López-Pérez, R.A.; Hernández, L.; Rodríguez-Zaragoza, F.A.; Wallace-Jones, R. and Pineda-López, R. 2014. Ophiuroidea (Echinodermata) from coral reefs in the Mexican Pacific. ZooKeys, 406: 101-145.) and gobies (Zepeta-Vilchis et al., 2013Zepeta-Vilchis, R.C.; Ayala-Bocos, A.; Valencia-Méndez, O. and López-Pérez, R.A. 2013. First record and range extension of the jawfish Opistognathus panamaensis (Perciformes: Opistognathidae) from western Mexico. Marine Biodiversity Records, 6: e132.), such attributes have also been signaled as responsible for increasing species richness and dominance of the groups in marine habitats. On the contrary, taxa included on Pasiphaeidae or Pandalidae members of which often constitute an important component of the mesopelagic and bathypelagic planktonic communities (Komai et al., 2012Komai, T.; Lin, C.W. and Chan, T.Y. 2012. Bathypelagic shrimp of the genus Pasiphaea (Decapoda: Caridea: Pasiphaeidae) from waters around Taiwan, with descriptions of four new species. Journal of Crustacean Biology, 32: 295-325.) are relatively larger than Alpheidae and Palaemonidae, and does not establish symbiotic relations with other taxa and hence not as species rich as Alpheidae.

Grid data indicates that species richness concentrates in Galapagos and Panama, but relatively rich spots can also be observed in Colombia, Costa Rica, and the Gulf of California. Boschi (2000Boschi, E.E. 2000. Species of decapods crustaceans and their distribution in the American zoogeographic provinces. Revista de Investigacion y Desarrollo Pesquero, 13: 7-136.) and Wicksten and Hendrickx (2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.) recognized the Panamic Province as the most diverse caridean shrimp area in the entire Eastern Tropical Pacific. However, if we further analyze the number of caridean species against the number of caridean references per grid, we will notice that rich spots correspond with the most heavily studied areas in the ETP; in fact there is a meaningful positive relationship (Spearman rank correlation r = 0.88, p = 0.00) between species richness and the number of caridean references per grid. While the number of references is not an adequate proxy for sampling intensity and a correlation is not a cause-consequence proof, the observed relation suggests to some extent that the observed benthic caridean species richness in the ETP may have not exclusively resulted from habitat heterogeneity or large-scale evolutionary processes currently invoked as one of the several speciation or richness drivers in the marine realm (Tittensor et al., 2010Tittensor, D.P.; Mora, C.; Jetz, W.; Lotze, H.K.; Ricard, D.; Berghe, E.V. and Worm, B. 2010. Global patterns and predictors of marine biodiversity across taxa. Nature, 466: 1098-1103.), instead they may have also arisen from differences in research effort and hence from sampling bias. In the same way, spatial analyses of rare species are not meaningfully related to latitude in the studied area (i.e., no meaningful increase toward Central America). The observed results, rather than plea against the currently recognized caridean spatial richness pattern (Boschi, 2000Boschi, E.E. 2000. Species of decapods crustaceans and their distribution in the American zoogeographic provinces. Revista de Investigacion y Desarrollo Pesquero, 13: 7-136.; Wicksten and Hendrickx, 2003Wicksten, M.K. and Hendrickx, M.E. 2003. An update checklist of benthic, marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the Eastern Tropical Pacific. p. 49-76. In: M.E. Hendrickx (ed), Contributions to the Study of East Pacific Crustaceans, Vol. 2. Mexico, ICMyL, UNAM.), call for a more thoughtful and systematic sampling protocol in order to get information from poorly sampled areas in the Eastern Tropical Pacific.

The function that explains the historic patterns per country and for the entire region suggest that the inventory of the marine benthic caridean shrimp fauna from the ETP is still incomplete. The extent of the gap between the observed and the expected caridean diversity in the ETP is unknown, but several evidence lines may suggest its size. First, if we considered that the benthic caridean records are restricted to an area that occupies 8.8% of the entire ETP, it becomes obvious that most of the region (~91%) is still not inventoried. In this regard if the species-area relationship is invoked, then marine benthic caridean shrimp fauna of the ETP must be nine times larger than the observed and hence is still far from being complete. According to the species-area relationship and the Chao2 species estimator, the marine benthic caridean shrimp fauna of the ETP must be 11% to 900% larger than the current observed. Species-area relationships have been invoked to explain species richness for several biological groups (Connor and McCoy, 2001Connor, E.F. and McCoy, E.D. 2001. Species-area relationships. Encyclopedia of Biodiversity, 5: 297-411.) including marine invertebrates (Alvarado et al., 2010Alvarado, J.J.; Solís-Marín, F.A. and Ahearn, C.G. 2010. Echinoderm (Echinodermata) diversity in the Pacific coast of Central America. Marine Biodiversity, 40: 45-56.; Costello et al., 2010Costello, M.J.; Coll, M.; Danovaro, R.; Halpin, P.; Ojaveer, H. and Miloslavich, P. 2010. A Census of Marine Biodiversity Knowledge, Resources, and Future Challenges. PLoS ONE, 5: e12110.; Granja-Fernández and López-Pérez, 2012López-Pérez, R.A.; Bastida-Zavala, R.; García-Madrigal, M.S.; Barrientos-Luján, N.A.; Torres-Huerta, A.M.; Montoya-Márquez, A.; Benítez-Villalobos, F. and Meráz-Hernando, J.F. 2012. ¿Cuanto sabemos de la diversidad de la fauna marina y costera de Oaxaca? p. 435-449. In: A.J. Sánchez, X. Chiappa-Carrara and R. Brito-Pérez (eds), Recursos Costeros del Sureste: tendencias actuales en investigación y estado del arte, Vol. 1. Mexico, CONCITEY, FOMIX, SIIDETEY, UNAM, CONACYT, ECOSUR, PROMEP, RECORECOS, UNACR, UJAT . ). Nonetheless, this number must be considered with caution since habitat diversity, area per se, passive sampling and edge effects are not mutually exclusive mechanisms and may operate individually and/or in synergy to cause and or modify species-area relationships (Connor and McCoy, 2001Connor, E.F. and McCoy, E.D. 2001. Species-area relationships. Encyclopedia of Biodiversity, 5: 297-411.). On the other hand, the non-parametric Chao2 species index indicates that caridean richness is 11% to 47% larger than recorded; this number, however, is sensibly minor compared to the suggested by species-area relationship, though it is in agreement with estimations regarding the entire crustacean fauna from the Mexican Tropical Pacific using different approaches (González-Oreja et al., 2010González-Oreja, J.A.; de la Fuente-Díaz Ordaz, A.A.; Hernández-Santín, L.; Buzo-Franco, D. and Bonache-Regidor, C. 2010. Evaluación de estimadores no paramétricos de la riqueza de especies. Un ejemplo con aves en áreas verdes de la ciudad de Puebla, México. Animal Biodiversity Conservation, 33: 31-45.; García-Madrigal et al., 2012García-Madrigal, M.S.; Jarquín-González, J. and Morales-Domínguez, E. 2012. Panorama del estado del conocimiento de los crustáceos del Pacifico Sur de México. p. 396-414. In: A.J. Sánchez, X. Chiappa-Carrara and R. Brito-Pérez(eds), Recursos Costeros del Sureste: tendencias actuales en investigación y estado del arte, Vol. 1. Mexico, CONCITEY, FOMIX, SIIDETEY, UNAM, CONACYT, ECOSUR, PROMEP, RECORECOS, UNACR, UJAT. ; López-Pérez et al., 2012López-Pérez, R.A.; Bastida-Zavala, R.; García-Madrigal, M.S.; Barrientos-Luján, N.A.; Torres-Huerta, A.M.; Montoya-Márquez, A.; Benítez-Villalobos, F. and Meráz-Hernando, J.F. 2012. ¿Cuanto sabemos de la diversidad de la fauna marina y costera de Oaxaca? p. 435-449. In: A.J. Sánchez, X. Chiappa-Carrara and R. Brito-Pérez (eds), Recursos Costeros del Sureste: tendencias actuales en investigación y estado del arte, Vol. 1. Mexico, CONCITEY, FOMIX, SIIDETEY, UNAM, CONACYT, ECOSUR, PROMEP, RECORECOS, UNACR, UJAT . ). In the absence of complete inventories, nonparametric estimators (for example, Chao2) have been shown to perform better than most other methods, such as observed species richness, species-area curves or asymptotic estimators (Colwell, 2009Colwell, R.K. 2009. EstimateS, Version 7.5: statistical estimation of species richness and shared species from samples. Available at Available at http://viceroy.eeb.uconn.edu/estimates . Accessed on: 16 December 2017.
http://viceroy.eeb.uconn.edu/estimates...
), but the Chao2 estimator still gives a lower bound to expected species richness, thus producing conservative estimates (Hortal et al., 2006Hortal, J.; Borges, P.A.V. and Gaspar, C. 2006. Evaluating the performance of species richness estimators: sensitivity to sample grain size. Journal of Animal Ecology, 75: 274-287.).

It is revealing that 44 species (24%) inhabit a single grid (as much as ~11,200 km2) but as many as 135 species (~74%) are restricted to less than five grids each. This either indicates that most of the caridean shrimp species in the ETP have a narrow distribution (i.e., provincialism) or that current distribution of narrowly distributed species (74%) resulted from sampling deviations. Previous statements about endemism or narrowness in species distribution in taxonomic groups such as corals, crustaceans, echinoderms or fishes in the ETP indicating that species are spatially restricted to Central America or the Gulf of California, have been contradicted by recent genomic analyses, detailed taxonomic studies or intensive sampling protocols across the area (Combosch et al., 2008Combosch, D.J.; Guzmán, H.M.; Schuhmacher, H. and Vollmer, S.V. 2008. Interspecific hybridization and restricted trans-Pacific gene flow in the Eastern Tropical Pacific Pocillopora. Molecular Ecology, 17: 1304-1312.; Zepeta-Vilchis et al., 2013Zepeta-Vilchis, R.C.; Ayala-Bocos, A.; Valencia-Méndez, O. and López-Pérez, R.A. 2013. First record and range extension of the jawfish Opistognathus panamaensis (Perciformes: Opistognathidae) from western Mexico. Marine Biodiversity Records, 6: e132.; Granja-Fernández et al., 2013Granja-Fernández, M.R.; Antonio-Pérez, V. and López-Pérez, R.A. 2013. Euapta godeffroyi (Semper, 1868) (Holothuroidea: Synaptidae): filling the distribution gap between Mexico and Costa Rica, eastern tropical Pacific. Hidrobiológica, 23: 130-132.; Paz-García et al., 2015Paz-García, D.A.; Hellberg, M.E.; García-de León, F.J. and Balart, E.F. 2015. Switch between morphospecies of Pocillopora corals. American Naturalist, 186: 434-440.; Reyes-Bonilla et al., 2017Reyes-Bonilla, H.; López-Pérez, A.; Paz-García, D.A.; Parra-Madrazo, G.; Medina-Rosas, P. and Balart, E.F. 2017. Distribución del coral Pocillopora inflata (Scleractinia) en el Pacifico Mexicano y comentarios sobre su situación taxonómica. Hidrobiológica, 27: 131-135. ; Valencia-Méndez et al., 2017Valencia-Méndez, O.; López-Pérez A.; Martínez-Guerrero, B.; Ramírez-Chávez, E. and Antonio-Pérez, V. 2017. New record of Hymenocera picta (Decapoda: Caridea) in southern Mexican Pacific reefs. Journal of Threatened Taxa, 9: 10571-10576.). While work on this regard is pending on ETP caridean fauna, the relatively large sampling gaps (~ 91%, see above) regarding the group is the most likely scenario for explaining the narrow distribution of a large number of species in the area, not only for caridean shrimps (this contribution) but across several taxonomic marine groups and areas worldwide (Costello et al., 2010Costello, M.J.; Coll, M.; Danovaro, R.; Halpin, P.; Ojaveer, H. and Miloslavich, P. 2010. A Census of Marine Biodiversity Knowledge, Resources, and Future Challenges. PLoS ONE, 5: e12110.).

Finally, integrative databases (such as the current one) resulting from previously published information to make generalizations in ecology, without any doubt is subjected to errors that distort our view. In particular, bias in sample size, but also in space, time and taxonomy may have altered the stated generalizations about the species richness distribution pattern on ETP caridean fauna; nevertheless the current meta-analysis has an advantage over simple verbal reviews because unbiased estimates of the overall strength of a hypothetical relationship can be obtained, heterogeneity in research findings identified and new research questions developed (Møller and Jennions, 2001Møller, A.P. and Jennions, M.D. 2001. Testing and adjusting for publication bias. Trends in Ecology and Evolution, 16: 580-586 ). In this regard, detailed taxonomy of the Caridea and intensive sampling across environments in the ETP still is the most urgent need in the studied area.

ACKNOWLEDGEMENTS

Authors are in debt to Michel Hendrickx and Mary Wicksten for providing critical references, and to Arthur Anker, Sammy De Grave, Michel Hendrickx, Mary Wicksten and Carolina Tavares for taxonomic advice and critical review of the checklist. Authors are grateful to Socorro García (UMAR) and Luis Hernández (UABCS) for early comments to the manuscript and to anonymous reviewers for suggestions and corrections during revision process. While writing this manuscript BMG received a graduate fellowship (257618) from Consejo Nacional de Ciencia y Tecnología (CONACYT), Mexico.

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Publication Dates

  • Publication in this collection
    06 Dec 2018
  • Date of issue
    2018

History

  • Received
    27 Apr 2018
  • Accepted
    14 Sept 2018
Sociedade Brasileira de Carcinologia Instituto de Biociências, UNESP, Campus Botucatu, Rua Professor Doutor Antônio Celso Wagner Zanin, 250 , Botucatu, SP, 18618-689 - Botucatu - SP - Brazil
E-mail: editor.nauplius@gmail.com