An annotated checklist of marine caridean and stenopodidean shrimps ( Malacostraca : Decapoda ) of the Caribbean coast of Panama

A checklist of caridean and stenopodidean shrimps of the Caribbean coast of Panama is presented, based on material collected during two local workshops (2005, 2008) as well as extensive sampling during 2006–2010. Th is material is augmented by an annotated list of previously recorded species, amounting to a total of 157 species, including 20 new records. Doubtful records are discussed. Th e current checklist is however considered relatively incomplete as older records could not always be verifi ed and more taxa remain to be described. Despite the defi ciencies of the present list, the Caribbean coastline of Panama is clearly one of the most species rich areas in the entire Atlantic Ocean for caridean shrimps. key wordS Caridea, Central America, faunal list, new records, Neotropical region, Stenopodidea. orIgInal artIcle This article is part of the tribute offered by the Brazilian Crustacean Society in memoriam of Michael Türkay for his outstanding contribution to Carcinology CORRESPONDING AUTHOR Sammy De Grave sammy.degrave@oum.ox.ac.uk SUBMITTED 16 November 2016 ACCEPTED 29 December 2016 PUBLISHED 5 June 2017 Guest Editor Célio Magalhães DOI 10.1590/2358-2936e2017015 Nauplius, 25: e2017015


IntroductIon
Although a few early records of shrimps exist for Caribbean Panama (e.g.Kingsley, 1879;Coutière, 1909), large scale decapod faunal recording started with the work of Lawrence G. (Larry) Abele, who commenced working on the local fauna in 1968 and sampled a total of 119 stations along both the Caribbean and Pacific coastline over a 15 month period (see Abele, 1972a).In the same year, a significant oil spillage near the Smithsonian Tropical Research Institute (STRI)'s research centre at Punta Galeta, led to a three-year long study of the local habitats and communities, which started in 1970 and resulted in an extensive faunal list (see Birkeland et al., 1976).These faunal lists enabled some ecological work (e.g.Heck, 1977 on sea grass communities) and culminated in a checklist of the invertebrates of Punta Galeta (Cubit and Williams, 1983).
From this, it is immediately obvious that the history of faunal recording in Panama is closely linked to the history of STRI (detailed in Robertson et al., 2009), which has operated several research stations along the Caribbean coast of Panama.The Punta Galeta laboratory was the first to be opened in 1964 and provided a nucleus for early records.Marine research in the San Blas archipelago followed in 1970, increasing with the opening of a laboratory in 1977, which continued for 20 years.In 2003, a state-of-the-art research laboratory (Bocas del Toro Research Station or BRS) was opened on Isla Colón in the Bocas del Toro Archipelago, with the majority of shrimp records for Caribbean Panama, detailed in the present list originating in that area.
Since its opening, the BRS station has organised several international invertebrate taxonomy workshops, as well as a series of taxonomic training workshops (Collin, 2005), both of which contributed significantly to the present checklist.Given the presence of wellequipped laboratories, numerous ecological and evolutionary studies have been carried out in the region.As a result many faunistic records are hidden in nontaxonomic publications, in which often working names have been used rather than updated nomenclature.In addition, a number of photographs or personal communications have appeared in recent years, based on species not formally recorded from the area.
The present checklist compiles this range of disparate sources using up-to-date nomenclature and classification, as well as reporting on unpublished collections largely obtained during two workshops (2005,2008) at the BRS station, and additional fieldwork conducted in 2015.Nomenclature and classification generally follows De Grave and Fransen (2011), updated by De Grave et al. (2014) for hippolytoid families and De Grave et al. (2015) for palaemonoid families.For literature records, the area from which the species was recorded is given, unless this was not stated in the original record, in which case "Caribbean Panama" is used.Several taxa, especially Alpheidae, were previously recorded under various codes, e.g.Synalpheus bousfieldi "A" in Morrison et al. (2004); these have all herein been linked to formally described taxa to facilitate future studies.Questionable records are indicated by (?).Material examined for the present work is largely deposited in the Zoological Collections of the Oxford University Museum of Natural History, Oxford, U.K. (OUMNH.ZC) with additional samples in Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP) and the Florida Museum of Natural History, University of Florida, Gainesville, USA (FLMNH UF).
Remarks.Sparsely recorded in the tropical western Atlantic, mainly in the Caribbean Sea, with an unconfirmed record from Brazil; in and under coral rubble, less than 10 m.The species may need a thorough taxonomic revision (Anker et al., 2016).
Remarks.Sparsely recorded in the western Atlantic, from Bermuda southwards to Panama and Bahia, Brazil; in coral rubble, in depths of 0-46 m; a species complex being revised at the moment (Anker, in prep.).
Remarks.Currently known from several localities in the Caribbean Sea, Florida and southern Gulf of Mexico (De Grave et al., 2017); typically under rocks and coral rubble on sandy patches between coral stands, in depths of 2-30 m.Chace, 1972 Previous records.Williams et al., 2001: 377 [Caribbean Panama].

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 Cayo Coral (Coral Cay), 09°14.502'N082°08.288'W,under rocks and coral rubble, 1 m, leg. A. Anker, 11.08.2008, OUMNH.ZC.2008-14-122.Remarks.Known with certainty from several localities in the Caribbean Sea, southern Florida and the Gulf of Mexico (Anker, 2012); records from further north (eastern USA) or south (Brazil) belong to either A. angulosus or one of the new species described in Anker (2012); in mixed sand-rubble habitats, 0-10 m.The true identity of older Panamanian records cannot be resolved as they may refer to any of the Caribbean species of the A. armillatus complex (see Anker, 2012).
Remarks.Amphi-Atlantic species, in the western Atlantic recorded from Bermuda southwards to Rio Grande do Sul, Brazil, including much of the Caribbean Sea and southern Gulf of Mexico (Anker et al., 2009;Soledade and Almeida, 2013); usually in rock or dead coral crevices, intertidal to about 5 m.
Remarks.Sparsely recorded from the Florida Keys southwards to the Caribbean Sea and São Paulo, Brazil (Almeida et al., 2015); in a wide range of habitats (rocky shores, mangroves, sea grass beds), typically under rocks in the intertidal or shallow subtidal (less than 5 m).
Remarks.Currently known from Guadeloupe, Cuba, Panama and several Brazilian states (Pará to São Paulo) (Soledade and Almeida, 2013); restricted to estuaries and mangroves, in burrows in soft mud, intertidal to 5 m.
Remarks.Common and conspicuous species widely distributed in the western Atlantic, from North Carolina southwards to São Paulo and Trindade Island, Brazil, including most of the Caribbean Sea and Gulf of Mexico (Anker et al., 2008b;2016); typically in reef and nearby habitats, especially amongst rubble, in depth of 0-40 m.Older records may refer to the relatively recently described Alpheus paraformosus
Remarks.The presence of A. heterochaelis in Panama is highly questionable and all herein listed records are considered as doubtful.Most likely, these records refer to one or both recently described Atlantic species which are morphologically similar to A. heterochaelis: Alpheus buckupi Almeida, Terossi, Araújo-Silva and Mantelatto, 2013 (known from the Lesser Antilles southwards to São Paulo, Brazil, as well as São Tomé in the eastern Atlantic, see Pachelle et al., 2016) and/ or Alpheus petronioi Almeida, Terossi and Mantelatto, 2014 (known from Colombia and Venezuela to São Paulo, Brazil, see Almeida et al., 2014).
Remarks.Recorded from several scattered locations in the Caribbean Sea and the southern Gulf of Mexico (De Grave et al., 2017); always associated with the corkscrew sea-anemone B. annulata at depths of 8-17 m.
Remarks.Sparsely recorded from Bermuda southwards to Venezuela and Panama (Anker, 2012); typically encountered under large rocks or in rubble, close to coral reefs, 0-2 m.
Remarks.Amphi-Atlantic taxon, in the western Atlantic known based on a few records from Florida southwards to São Paulo, Brazil, including the Caribbean Sea and southern Gulf of Mexico (Soledade and Almeida, 2013); lives deep in cavities in exposed rocky and coral reef substrates, 0-5 m.Material examined.Bocas del Toro. 1 spec., Isla Bastimentos, southern shore near Bastimentos village, fine sand and mud near mangroves, under large pieces of rubble, 0-1 m, leg. A. Anker et al., 29.04.2015, MZUSP 34045.Remarks.Sparsely recorded from Florida southwards to Panama (Anker, 2012); typically found under rocks and coral rubble on fine sand or silt, in sheltered locations, 0-2 m
Remarks.The species has been recorded from the Florida Keys, the eastern Gulf of Mexico and several localities in the Caribbean Sea, including Dominican Republic, Colombia and Panama (Felder et al., 2009); lives in self-constructed galleries inside dead corals, usually in exposed areas and in very shallow water (0-2 m).
Remarks.Amphi-Atlantic species, in the western Atlantic recorded from Bermuda southwards to Bahia and Trindade Island, Brazil (Anker et al., 2016); on the Caribbean coast of Panama typically found inside large loggerhead sponges, Spheciospongia vesparia (Lamarck, 1814), in shallow water (less than 5 m), but also recorded from deep-water rubble down to 45-67 m.In older Atlantic literature known under the name A. cylindricus, which is now restricted to the tropical eastern Pacific (see Anker et al., 2008d).
Remarks.Sparsely recorded from the Florida Keys southwards to São Paulo, Brazil, including some localities in the Caribbean Sea (Soledade and Almeida, 2013); typically found under larger rocks or coral rubble near reefs or rocky-sandy shores, 0-4 m. (Armstrong, 1949) Previous records.Abele, 1976: 273 [around Colón]; Heck and Wetstone, 1977: 200 [around Galeta]; Cubit and Williams, 1983: 24 [Galeta  Remarks.Widely distributed in the tropical northwestern Atlantic, from southern Florida southwards to Venezuela, including the entire Caribbean Sea and southern parts of the Gulf of Mexico (Anker, 2012); typically encountered in sea grass beds or close to mangroves, in burrows or under large rocks, 0-3 m.
Material examined.Bocas del Toro. 1 spec., Isla Colón, Boca del Drago, sea grass bed with sand patches, 0.2-1.5 m, leg. A. Anker et al., 01.05.2015, MZUSP 33574.Remarks.This rarely collected species is known only from two localities, southern Florida (near Fort Pierce) and Caribbean Panama (Bocas del Toro) (Anker, 2011a), but is presumably more widespread in the Caribbean Sea; always encountered in burrows in mud or fine sand, however, the host animal remains unknown; probably confined to intertidal and shallow subtidal waters, with one specimen found in the stomach of a sand perch taken at 31 m.
Remarks.Presently known only from the type series from Isla Bastimentos in the Bocas del Toro Archipelago (Anker, 2011a) and two additional specimens from Isla Grande (present material); inhabits burrows of yet unknown hosts, in shallow water (less than 1.5 m).
Remarks.Presently known only from the southern Caribbean Sea: Venezuela, Colombia and Panama (Anker, 2011a); associated with mud shrimps of the genus Upogebia Leach, 1814, in very shallow water (0-1 m).

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 Remarks.The Panamanian specimens are provisionally assigned to L. forceps following Anker (2011a), as they show some minor morphological differences to the populations from North Carolina (type locality) and elsewhere.Recorded from North Carolina southwards to Panama and Ceará, Brazil (Pachelle et al., 2016); associated with several species of ghost and mud shrimps (Callianassidae, Upogebiidae) in very shallow water (0-5 m).
Remarks.Presently known only from the holotype collected on the southern shore of Isla Grande, Panama, from a burrow of an unknown host (Anker, 2008).
Remarks.A pantropical taxon, in the western Atlantic recorded from Puerto Rico southwards to Fernando do Noronha, off northeastern Brazil, including the eastern Gulf of Mexico (Anker et al., 2016); living in crevices of coralline algae and dead corals, 0-12 m.
Remarks.This species remains known only from two specimens, one from Isla Grande and the other from the Bocas del Toro (Anker, 2010a); inhabits small spaces under rocks in water less than 1 m deep.
Remarks.Presently known only from Panama (Isla Grande, San Blas Islands) and Honduras (Bay Islands) (Anker, 2010a); under coral rubble in very shallow water (less than 1.5 m).
Remarks.Only known from Indian River Lagoon (Florida, USA), Turks and Caicos Islands (Anker, 2010a) and now Panama; associated with burrows of unknown hosts on sandy mud flats, sometimes among or proximal to sea grass beds, 0.5-3 m.
Remarks.Currently known only from the southern Caribbean Sea: Panama (both records from Isla Grande), Colombia, Venezuela and Tobago (Anker, 2011a); likely a commensal in burrows of ghost shrimps, Neocallichirus spp., 0-1 m.

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 Remarks.Although described only recently, S. depressus appears to be widely distributed in the western Atlantic, being known from Panama, Venezuela, St. Martin, Barbados and São Paulo, Brazil (Anker et al., 2013); typically found under rocks or coral rubble on sand flats, 0-7 m.
Remarks.The true identity of the western Atlantic material assigned to this taxon in Anker et al. (2012) is unclear.Earlier records listed above likely refer to the same taxon and not S. tenuispina, a much larger species originally described from southern Brazil (see Anker and Pachelle, 2014 for a redescription).
Remarks.Hultgren and Duffy (2011) included "S.androsi (Panama)" as a terminal in their phylogram, but only listed material from Belize under sequenced material.Thus, the presence of S. androsi in Panama requires confirmation.This pair-living species is known from only a few localities in the Caribbean Sea, Bahamas and north-eastern Brazil; lives in various sponges, 14-55 m (Anker and Pachelle, 2014).
Remarks.This recently described pair-living species from the Synalpheus paraneptunus Coutière, 1909 complex is currently known only from Belize and Panama; associates with several sponge species, 0.5-10 m (Hultgren and Brandt, 2015).Remarks.Widely distributed in the western Atlantic, from coral reefs off Texas southwards to São Paulo, Brazil, including the entire Caribbean Sea and some oceanic islands of Brazil (Anker et al., 2012;2016); in crevices of dead corals, coralline algae etc., 1-42 m.The distributional record for Bocas del Toro, Panama, in Anker et al. (2012) refers to the above sample.
Remarks.Widely distributed in the western Atlantic, from Florida southwards to Santa Catarina, Brazil; found in a variety of rocky and coral reef habitats, 0-20 m.The species is morphologically variable and may comprise a species complex (Anker et al., 2012).
Remarks.Rare species, known only from the original description based on material from the Dominican Republic (Armstrong, 1941) and the above record from San Blas Islands (Panama), which requires confirmation.Associated with sponges; depth range unknown.
Remarks.This species is currently known only from the western Caribbean Sea, with records from Panama, Curaçao and Jamaica (Macdonald et al., 2009); associated with sponges of the genus Xestospongia de Laubenfels, 1932, 0.5-15 m.
Remarks.Widely distributed in the western Atlantic, from the northern Gulf of Mexico and the Bahamas southwards to Bahia, Brazil (Macdonald et al., 2009); inhabiting a variety of sponges at depths of 5-20 m.

Synalpheus brevicarpus Herrick, 1891 sensu lato
Previous records.Cubit and Williams, 1983: 24 [Galeta Point, as S. brevidactylus, lap. cal.];Duffy, 1992: 131  Remarks.Widely distributed in the western Atlantic, however, comprised of several genetically, morphologically and chromatically distinct species (Hultgren et al., 2014), of which several are present in Panama.The record of S. brevicarpus from Isla Bartolomé in the Las Perlas Archipelago, on the Pacific side of Panama (Morrison et al., 2004), most likely refers to one of the three cryptic species in the Synalpheus digueti Coutière, 1909complex (see Hultgren et al., 2014).
Remarks.Only known from the original description; pair living in crevice-boring sponges, 1-3 m.The validity of this species needs to be confirmed by a molecular analysis.
Remarks.The presence of S. brevifrons in Panama requires confirmation.Ríos and Duffy (2007) first stated that this species is possibly present in San Blas Islands, but after having provided some details about the single ovigerous female from San Blas, the authors concluded that it may represent "yet another undescribed species".Otherwise S. brevifrons is known only from the Dominican Republic and Belize (Anker et al., 2012), where it associates with the sponge Aiolochroia crassa (Hyatt, 1875) at depths of 1-15 m.

Synalpheus brooksi Coutière, 1909
Previous records.Duffy, 1992: 131 [San Blas Islands]; Duffy, 1993: 461 [San Blas Islands]; Morrison et al., 2004: 574  Remarks.Widely distributed in the western Atlantic, from Florida southwards to Bahia, Brazil, including much of the Caribbean Sea and parts of the Gulf of Mexico (possibly a species complex) (Anker et al., 2012); lives in subsocial groups in a variety of sponges; 1-73 m, most abundant shallower than 10 m.

Synalpheus carpenteri Macdonald and Duffy, 2006
Previous records.Remarks.Known from Belize, Panama, Jamaica, St Martin and the Bahamas (Anker et al., 2012); in malefemale pairs or small subsocial groups in sponges of the genus Agelas, 10-20 m.
Remarks.Only known from a few records from the Dominican Republic, Grenadines, Venezuela and Panama (Anker et al., 2012); ecology poorly known, apparently free-living in crevices of coral rocks in shallow water.
Remarks.Sparsely recorded in the western Atlantic from the Bahamas and southwestern Gulf of Mexico to the Dominican Republic and Panama (Anker et al., 2012); typically in female-male pairs on rock or coral crevices, at depths of 0-14 m.
Remarks.Known with certainty only from Panama and Jamaica, possibly also occurs in the Florida Keys (Anker et al., 2012); lives in eusocial groups composed of a queen, one or several kings and up to 80 colony members, in a variety of sponges, 0-2 m.

Synalpheus elizabethae (Ríos and Duffy, 2007)
Previous records.Duffy, 1996: 309 [San Blas Islands, as S. "rathbunae A"]; Morrison et al., 2004: 575  Remarks.Uncommon in the western Atlantic, with scattered records from the Florida Keys and the southwestern Gulf of Mexico to Panama (Anker et al., 2012); lives in eusocial groups of up to several hundred colony members and one reproductive female (queen); associated with several sponge species, 0-18 m.
Remarks.Known with certainty only from the Dominican Republic, Belize and Panama (Anker et al., 2012); lives in eusocial groups of up to a few hundreds of colony members and one reproductive female; associated with several sponge species, 0-15 m.

Synalpheus fritzmuelleri Coutière, 1909
Previous records.Abele, 1976: 273 [around Colón]; Birkeland et al., 1976: 137 [Galeta reef ]; Heck, 1977: 338 [around Galeta Island]; Heck, 1979: 199 [Caribbean  Remarks.One of the most common and widely distributed species in the western Atlantic, ranging from North Carolina and the northern Gulf of Mexico to Santa Catarina, Brazil, including the entire Caribbean Sea, and some isolated islands in the Central Atlantic

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 (Anker et al., 2012;2016); lives in male-female pairs in a variety of microhabitats on coral reefs and rocky shore habitats, 0-51 m.The record of S. fritzmuelleri from Isla Bartolomé in the Las Perlas Archipelago, on the Pacific side of Panama (Morrison et al., 2004), refers to a different eastern Pacific species, the identity of which remains to be determined.Remarks.Widely distributed in the western Atlantic, from Florida southwards to Tobago, and possibly further south to Brazil (Anker et al., 2012); typically found in male-female pairs in a variety of sponges, 1-20 m.
Remarks.Widely distributed in the western Atlantic, from North Carolina southwards to Bahia, Brazil, including Caribbean Sea and Gulf of Mexico (Anker et al., 2012;2016); in male-female pairs, most often in tube sponges, 1-51 m.

Synalpheus herricki Coutière, 1909
Previous records.Birkeland et al., 1976: 137 [Galeta reef]; Cubit and Williams, 1983: 24 [Galeta  Remarks.Widely distributed but relatively uncommon western Atlantic species, ranging from southern Florida to the Caribbean Sea and Gulf of Mexico, and further south to Ceará, Brazil (Anker et al., 2012); lives in female-male pairs in a variety of sponges, 5-73 m.Remarks.Relatively rarely encountered species, presently known only from Curaçao, Panama and Barbados (Anker et al., 2012); typically found in malefemale pairs in several host sponge species, 1-6 m.

Synalpheus longicarpus (Herrick, 1891)
Previous records.Duffy, 1992: 131 [San Blas Islands]; Morrison et al., 2004: 574  Remarks.Some of the Panamanian records of S. minus may refer to S. antillensis, S. brevicarpus or another taxon from the S. brevicarpus -minus species complex (see Hultgren et al., 2014).The exact distribution range of S. minus is poorly known, but the species is known with certainty from the south-eastern USA, several localities in the Caribbean Sea and Brazil (under Synalpheus minus bahiensis Coutière, 1909) (Anker et al., 2012;2016).

Synalpheus pandionis Coutière, 1909
Previous records.Birkeland et al., 1976: 137 [Galeta reef]; Heck, 1977: 338 [around Galeta Island]; Heck, 1979: 198  Remarks.Widely distributed in the western Atlantic, ranging from Florida southwards to northern Brazil, including numerous localities in the Caribbean Sea and Gulf of Mexico (Anker et al., 2012); earlier records may refer to related taxa; lives in male-female pairs in several species of sponges, 0-80 m.Remarks.Synalpheus paraneptunus is with certainty only known from the holotype from the Gulf of Morrosquillo, Colombia (Anker and Tóth, 2008; under material examined therein, "Panama" should be substituted by Colombia).All the above records from Panama, as well as those listed in Table I in Anker et al. (2012), need confirmation as they most likely refer to other species in the S. paraneptunus complex.

Synalpheus rathbunae Coutière, 1909
Previous records.Duffy, 1992: 131  Remarks.Presently known from Belize, Jamaica and Panama (Anker et al., 2012); eusocial sponge-dwelling species, living in groups of up to several hundreds of individuals, including usually one reproductive female (queen), unknown number of reproductive males (possibly only one), and numerous non-reproductive colony members; typically below 6 m depth.

Synalpheus ruetzleri Macdonald and Duffy, 2006
Previous records.Macdonald and Duffy, 2006: 8  Remarks.Widely distributed in the western Atlantic: common in the Florida Keys, Caribbean Sea and parts of the Gulf of Mexico, extending to São Paulo, Brazil (Anker et al., 2012); typically in male-female pairs in coral rock crevices, 0-30 m.

Synalpheus townsendi Coutière, 1909
Previous records.Abele, 1976: 273 [around  Remarks.One of the most common and widespread species of Synalpheus in the western Atlantic, ranging from North Carolina southwards to Rio de Janeiro and Trindade Island, Brazil, including much of the Caribbean and Gulf of Mexico (Anker et al., 2012;2016); typically found in female-male pairs in a variety of microhabitats, including coral rocks crevices, calcareous algae and sponges, 0-102 m.

Synalpheus ul (Ríos and Duffy, 2007)
Previous records.Ríos and Duffy, 2007: 63  Remarks.Relatively common western Atlantic species recorded from Belize southwards to Bahia, Brazil, including Panama and several other localities in the Caribbean Sea (Almeida et al., 2012); typically found in male-female pairs in a variety of sponge hosts, 1-3 m.Remarks.Relatively common western Atlantic species recorded from western Florida and the southern Gulf of Mexico (Alacranes Reef) southwards to Ceará, Brazil, including Caribbean Sea (Belize, Jamaica, Panama) (Anker et al., 2012;2016); pair-living spongeassociated species, 1-3 m.

Synalpheus williamsi Ríos and Duffy, 1999
Previous records.Ríos and Duffy, 1999: van Soest, 1986, leg. E. Tóth, 04.01.2007, OUMNH.ZC.2007-14-01.Remarks.Widespread in the western Atlantic, from Bermuda southwards to Espírito Santo, Brazil, including the Gulf of Mexico and Caribbean Sea (Felder et al., 2009), with the present material representing new record for Panama.Although A. antiguensis has been recorded in older literature from a variety of habitats from the intertidal down to 118 m (summarised in Williams, 1984); all recent records, including the present ones, are based on specimens found in the oscula of larger sponges, suggesting that the species might be at least a facultative sponge associate.

Remarks. Common and widely distributed western
Atlantic species, ranging from North Carolina southwards to Santa Catarina, Brazil, including the Gulf of Mexico and Caribbean Sea (Felder et al., 2009); on a variety of substrates, but most frequently living on algae and sea grass, 0-10 m.

Latreutes fucorum (Fabricius, 1798)
Previous records.Heck, 1977: 338 [around Galeta Island]; Heck, 1979: 200 [Caribbean Panama]; Heck and Weinstein, 1989: 631  Remarks.Amphi-Atlantic taxon, in the western Atlantic ranging from southern Newfoundland to Bahia, Brazil (Felder et al., 2009); in sea grass beds or on floating algae, 0-2 m.Remarks.Amphi-Atlantic taxon, in the western Atlantic ranging from New Jersey southwards to Argentina (Felder et al., 2009), with the present material being a new record for Panama; most frequently found in sea grass beds, but also occurs in a variety of other microhabitats, 0-44 m
Remarks.Rhyne and Lin (2006) referred part of the material of L. wurdemanni in Abele and Kim (1989) to L. ankeri, however, without any further comments.

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 The material in Abele and Kim (1989) included two specimens (1 male and 1 ovigerous female) collected from the lower western chamber of Gatun Locks.As neither specimen was discussed by Rhyne and Lin (2006), it is unclear which of them is L. ankeri.Based on the species' wide occurrence in the tropical and subtropical western Atlantic, from southern Florida to São Paulo, Brazil (Rhyne and Lin, 2006), its occurrence in Panama is highly likely but does require confirmation.
Remarks.Although the exact distribution of L. intermedia is unclear due to confusion with the recently described Lysmata jundalini Rhyne, Calado and dos Santos, 2012 (see below), the species seems to be relatively widespread in the western Atlantic, from Florida to Brazil (Pachelle et al., 2016); typically on rocky and rubble-algal bottoms, 0-22 m.The herein examined specimens were assigned to L. intermedia on the basis of their colour pattern and some morphological characteristics listed in Rhyne et al. (2012), although the distinction between preserved L. intermedia and L. jundalini specimens is extremely difficult as most diagnostic characters overlap and are ambiguous.All previous records of L. intermedia from Panama cannot be confirmed at this stage as they could refer to either species.

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 station and Playa Bluff, fossil reef terrace, intertidal, leg. A. Baeza, 2007, OUMNH.ZC.2009-06-004.Remarks.Currently known only from Puerto Rico, Mexico (Alacranes Reef in the southern Gulf of Mexico) and Panama (Bocas del Toro) (Rhyne et al., 2012); on mixed rubble-algal substrates near reefs and sea grass, in old conch shells etc., 0-2 m.The unsubstantiated record from the Caribbean Panama in Rhyne et al. (2012) is based on the herein examined specimens.
Remarks.Amphi-Atlantic taxon, in the western Atlantic sparsely recorded from Bermuda southwards to Paraíba, Brazil (Anker and Cox, 2011); rocky shores and other hard bottom and reef habitats, in shallow water (0-5 m).

Ascidonia miserabilis
Remarks.Rarely recorded species, with just a few records in the western Atlantic, ranging from the Flower Garden Banks, Texas, southwards to Atol das Rocas, Brazil (Pachelle et al., 2012); associated with solitary ascidians, 0-30 m.

Holthuisaeus bermudensis
Remarks.Widely distributed but rather uncommon western Atlantic species, ranging from Bermuda southwards to Espírito Santo, Brazil, with a few scattered records from the Caribbean Sea and Gulf of Mexico; formerly known as Periclimenaeus bermudensis (see Anker and De Grave, 2010); associated with a variety of sponge species, 15-78 m.
Remarks.It remains unclear if this species actually occurs in the Caribbean Sea or is replaced here by the related Neopontonides chacei Heard, 1986 (see below).
Remarks.Sparsely recorded in the western Atlantic, from the Dry Tortugas southwards to Pernambuco, Brazil (Vieira et al., 2012); an obligate sponge associate, 0-37 m.

Periclimenes perryae Chace, 1942
Previous record : Felder et al., 2009:  Remarks.Previously recorded from Florida and Colombia (Felder et al., 2009); the unsubstantiated record from the Caribbean coast of Panama in Felder et al. (2009) refers to the above-listed specimen; obligate associate of the basket star A. muricatum, 0-10 m.

Urocaris longicaudata Stimpson, 1860
Previous records.Heck, 1977: 338 [around Galeta Island, as Periclimenes longicaudatus]; Heck, 1979: 199 [Caribbean Panama, as P. longicaudatus Remarks.Widely distributed in the western Atlantic, from North Carolina southwards to São Paulo, Brazil, including much of the Caribbean Sea and Gulf of Mexico (Felder et al., 2009); found in a variety of habitats, typically with sandy bottom, nocturnally planktonic, 1-40 m.The present specimens constitute the first record of the species from Panama.
Remarks.Widely distributed in the western Atlantic, from North Carolina southwards to Bahia, Brazil (Almeida and Bezerra, 2011); occurs in a variety of habitats, most abundant in sea grass beds, 0-49 m.
Remarks.Cubit and Williams (1983)  Remarks.Circum-tropical taxon, in the western Atlantic recorded from Bermuda and the northern Gulf of Mexico southwards to at least Fernando de Noronha, Brazil (Felder et al., 2009).Although recorded from the Pacific coast of Panama by Abele (1976), the herein studied material represents the first record of T. amboinensis for the Caribbean coast of Panama.The species is usually found in close proximity to sea anemones and other cnidarians, at depths of about 0-10 m.
Remarks.Heck (1977;1979) records this species in his seagrass studies, whilst recording the related Thor manningi Chace, 1972 in other work (e.g. Heck andWetstone, 1977).Whilst it seems highly likely that only one species is involved, it cannot be discounted that both species occur in Panama.Thor floridanus is widespread in the western Atlantic, from North Carolina and the northern Gulf of Mexico southwards to Costa Rica (Felder et al., 2009); most often on seagrass and algae, 0-59 m.

dIScuSSIon
A total of 151 species of caridean and 4 stenopodidean shrimp taxa have been recorded from the Caribbean side of Panama, including 20 species herein recorded for the first time.Numerically, the fauna is dominated by the Alpheidae (99 taxa in total) amounting to 63 % of the fauna, with both Alpheus Fabricius, 1798 (34 taxa) and Synalpheus (44 taxa) well represented.This is not surprising, as the area has acted as one of the focal points of western Atlantic alpheid taxonomy in recent decades (e.g.Anker, 2008;2010a;2010b;2011a;2011b;2012;Anker and Tóth,

Shrimp checklist of Caribbean Panama
Nauplius, 25: 2017015 2008; Anker et al., 2007a;2007b;2008a;2008b;Anker and Vera Caripe, 2016), as well as extensive biological studies on sponge dwelling species of Synalpheus (e.g.Duffy, 1992;1993;1996;Hultgren and Duffy, 2011).As is usual in the western Atlantic, the second most species rich family is the Palaemonidae, with 29 species in Caribbean Panama, i.e. 19% of the total fauna, whilst all other families are represented only by a handful of species each.
A number of factors combine to make the current checklist far from complete.Firstly, the identity of a number of previously recorded taxa could not be confirmed during the present study.For example, Alpheus heterochaelis has been recorded a number of times in the past (Heck, 1977;1979;Abele and Kim, 1989), whilst Processa sp.aff.hemphilli was recorded on a single occasion by Cubit and Williams (1983).Without a direct re-examination of the original material recorded under those names, it is not possible to speculate to which species these could refer to and for the time being these records must stand as is, although the presence of A. heterochaelis in Panama is extremely unlikely.Other such records are annotated in the list.Secondly, it is also clear that more as yet undescribed species are already known from Panama, for instance "Synalpheus brevicarpus A" and "S.guerini B" in Hultgren et al., 2014;"S. pandionis RED" in Morrison et al., 2004 (the latter also listed in Hultgren et al., 2010); and at least one unidentified, possibly undescribed species of Leptalpheus Williams, 1965in Anker (2011), as well as a further, two undescribed species in the genera Alpheus and Odontozona Holthuis, 1946 which are currently being described by the second author.Additionally, a number of unresolved species complexes are known to have representatives in the Panamanian Caribbean fauna, such as the Alpheus normanni and Alpheus paracrinitus species complexes, each with several cryptic taxa (Williams et al., 2001;Anker, pers. obs.).
A biogeographical analysis of the western Atlantic shrimp fauna is rather hampered by the fact that the majority of species, especially in the numerically dominant Alpheidae, have only been described in the last two decades.Nevertheless a few relatively recent studies allow for a broad-brush comparison of species richness of the Caribbean Panamanian shrimp fauna.A total of 122 species of shallow water caridean and stenopodidean shrimps have been recorded from Mexico (Felder et al., 2009;Santana-Moreno et al., 2013;Duarte et al., 2014), a much bigger country with the total Atlantic coastline approximately 2805 km long.Cuba, with a coastline of 3735 km, has 106 species of caridean and stenopodidean shrimps (Martínez-Iglesias et al., 1996;Lalana and Ortiz, 2000;De Grave et al., 2017), although this number appears rather low and is likely due to insufficient sampling and taxonomic work.The Caribbean coastline of Panama is about 1288 km in length, which is roughly a third of those of the Atlantic coast of Mexico and Cuba.Yet it harbours proportionally a much higher number of species, with the total estimated number to be somewhat in excess of 154 species.Although species richness on this scale is largely a reflection of habitat heterogeneity, local sampling effort plays a significant role.It is interesting to note that the majority of recent records are from the immediate vicinity of the STRI Research Station in Bocas del Toro, a far smaller area than the total available coastal sampling area in Panama.In an interesting parallel, Cubagua Island (Venezuela) has been intensively sampled over a number of years and despite being only 22 km 2 in area it harbours 46 species of Caridea and Stenopodidea (Hernández-Ávila et al., 2007).Without any doubts, additional sampling on the Caribbean coast of Panama, especially in waters deeper than 30 m, as well as in the extensive San Blas Archipelago, will yield additional species of shrimps.However, it is already clear that Panama, with its heterogeneous Caribbean and Pacific coastlines, represents one of the major hotspots for marine biodiversity in the Americas and worldwide.
acknowledgementS SDG acknowledges Rachel Colin, Director of the Bocas del Toro Research Station (BRS) for the opportunity to participate in the Third Taxonomy Workshop (August 2005), funded by the Smithsonian Tropical Research Institute (STRI).Both authors acknowledge Rachel Colin for organising the August 2008 Caridean shrimp workshop at BRS, as well as the BRS based logistics experts and the collecting efforts of all the international participants.AA thanks the Programa de Capacitação Institucional (PCI) of the Museu Paraense Emílio Goeldi (MPEG) and Universidade Federal de Goiás (UFG) for providing funding to finalise this study.
Remarks.Widespread in the western Atlantic from Bermuda southwards to São Paulo, Brazil, including much of the Caribbean Sea and Gulf of Mexico(Anker  et al., 2013); under rocks and rubble and in crevices of dead corals, 0-22 m.