Caribbean Naturalist

No. 25

Caribbean Naturalist

2015

Freshwater and BrackishWater Fishes Observed on Montserrat, Lesser Antilles, West Indies Robert E. Schmidt and Erin R. McMullin

The Caribbean Naturalist . . .

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Cover Photograph: Juvenile Mountain Mullet, Agonostomus monticola (Bancroft), a catadromous freshwater fish collected on Montserrat.. Photograph © Robert Schmidt.

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CARIBBEAN NATURALIST

James D. Ackerman, Department of Biology, University of Puerto Rico at Río Piedras, USA Alfonso Aguilar-Perera, Department of Marine Biology, Universidad Autónoma de Yucatán, Mexico Wayne J. Arendt, International Institute of Tropical Forestry, Luquillo, Puerto Rico, USA Rüdiger Bieler, Field Museum of Natural History, Chicago, IL, USA Christopher P. Bloch, Department of Biological Sciences, Bridgewater State University, Bridgewater, MA, USA William R. Buck, Institute of Systematic Botany, New York Botanical Garden, Bronx, NY, USA Leo Douglas, Department of Geography/Geology, University of the West Indies, Mona, Jamaica Robert Erdman, Department of Biological Sciences, Florida Gulf Coast University, Fort Myers, FL, USA Keith Goldfarb, Eagle Hill Institute, Steuben, ME, USA ... Editor-in-Chief Grizelle González, International Institute of Tropical Forestry, San Juan, Puerto Rico, USA Gary R. Graves, Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC, USA S. Blair Hedges, Department of Biology, Pennsylvania State University, University Park, PA, USA Julia A. Horrocks, Dept. of Biological and Chemical Sciences, Univ. of the West Indies, Cave Hill Campus, Barbados Scott Jones, Smithsonian Institution, Caribbean Coral Reef Ecosystems, Carrie Bow Cay, Belize Heather Judkins, Department of Biological Sciences, University of South Florida, St. Petersburg, FL, USA Craig A. Layman, Department of Applied Ecology, North Carolina State University, Raleigh, NC, USA John Leavengood, Department of Entomology, University of Kentucky, Lexington, KY, USA Antonio A. Mignucci-Giannoni, Manatee Conservation Center, Inter American University, Bayamón, Puerto Rico, USA Gregg Moore, Department of Biological Sciences, Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH, USA Dawn Phillip, Department of Life Sciences, University of The West Indies, St. Augustine, Trinidad and Tobago James Pitts, Department of Biology, Utah State University, Logan, UT, USA Robert Powell, Department of Biological Sciences, Avila University, Kansas City, MO, USA Chris Rimmer, Vermont Center for Ecostudies, Norwich, VT, USA Armando Rodríguez-Durán, Dean for Research, Inter American University, Bayamón, Puerto Rico, USA Noris Salazar Allen, Smithsonian Tropical Research Institute, Panama Inés Sastre de Jesus, Biology Department, University of Puerto Rico at Mayagüez, USA J. Angel Soto-Centeno, American Museum of Natural History, Division of Mammalogy, New York, NY, USA Christopher Starr, Department of Life Sciences, University of the West Indies, St. Augustine, Trinidad and Tobago David W. Steadman, Florida Museum of Natural History, Gainesville, FL, USA Kathleen Sullivan Sealey, Department of Biology, University of Miami, Coral Gables, FL, USA Jarrod M. Thaxton, Department of Biology, University of Puerto at Mayagüez, USA Jason M. Townsend, Department of Wildlife, Fish and Conservation Biology, University of CaliforniaDavis, USA ... Managing Editor Jill Weber, Eagle Hill Institute, Steuben, ME, USA ... Production Editor Byron Wilson, Department of Life Sciences, University of the West Indies at Mona, Kingston, Jamaica Graham A. J. Worthy, Department of Biology, University of Central Florida, Orlando, FL, USA Joseph M. Wunderle, International Institute of Tropical Forestry, University of Puerto Rico at Río Píedras, USA The Caribbean Naturalist (ISSN # 2326-7119) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680-0009. Phone 207-546-2821, FAX 207-546-3042. E-mail: [email protected]. Webpage: www.eaglehill.us/cana. Copyright © 2015, all rights reserved. Periodical postage paid in Steuben, ME and additional mailing offices. Special issue proposals are welcome. On-line secure subscription ordering: rate per year for Caribbean subscribers - $15 regular, $10 students, $60 organizations; for Non-Caribbean subscribers - $20 regular, $15 students, $80 organizations. Authors: submission guidelines are available at www. eaglehill.us/cana. Co-published journals: The Northeastern Naturalist (ISSN 1092-6194 [print], ISSN 1938-5307 [online]), the Southeastern Naturalist (ISSN 1528-7092 [print], ISSN 1938-5412 [online]), and the Urban Naturalist (ISNN #2328-8965), journals with separate Boards of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899).

2015 2015

Caribbean Naturalist CARIBBEAN NATURALIST R.E. Schmidt and E.R. McMullin

No. 25 No. 25:1–12

Freshwater and Brackish-Water Fishes Observed on Montserrat, Lesser Antilles, West Indies Robert E. Schmidt1,* and Erin R. McMullin1 Abstract - We conducted a short-term survey of fishes in the fresh and brackish waters of the Caribbean island of Montserrat from December 2013 to January 2014. We documented 13 species; most belonged to marine families except the introduced Poecilia reticulata (Guppy) and some catadromous and amphidromous species. We update the taxonomy of several Caribbean species and report a northward range extension for Genytremus luteus (Torroto Grunt). We were unable to identify to species specimens belonging to the amphidromous goby genus Sicydium, and suggest that more specimens and DNA data need to be collected to clarify the taxonomy in this genus.

Introduction Caribbean islands that have semi-permanent bodies of flowing freshwater have an aquatic macrofauna that at least includes Crustacea, Gastropoda, Insecta, Platyhelminthes, and fishes. Many of the aquatic species are regionally endemic or may be endemic to specific islands (Briggs 1984). The freshwater fish fauna has been well documented on some islands, e.g., Cuba (Vergara 1992), Puerto Rico (Neal et al. 2009), Jamaica (Caldwell 1966), Guadeloupe (Monti et al. 2010), Martinique (Lim et al. 2002), Curaçao (Debrot 2003), Trinidad and Tobago (Phillip et al. 2013), and St. Croix (Smith-Vaniz and Jelks 2014), but the freshwater fish fauna of other Caribbean islands remains poorly known. Montserrat is a small (264 km2) volcanic island in the Lesser Antilles. Freshwater surface flows are scarce and often ephemeral; small streams have cut deeply into the surface forming steep-walled canyons locally termed ghauts. The largest permanent water body was the Belham River, but that stream was buried by a pyroclastic flow in the 1995 eruption of the Soufrière Hills volcano. The purpose of this paper is to report on fishes we collected in some of the freshwater and brackish habitats on Montserrat which suffered from major widespread volcanic effects in 1995. Methods We collected fishes from late December 2013 through early January 2014 using small seine nets (3 m or 6 m long, 3/16-in Ace® knotless nylon mesh) or small dip nets. We preserved all fishes in 10% formalin. Prior to preservation, we clipped the right pelvic fins of several individuals of most species and preserved the fin samples in 95% ethanol. We transported the specimens and fin clips to the fish collection at the New York State Museum (NYSM; Troy, NY), where they are now catalogued. Bard College at Simon’s Rock, Great Barrington, MA, USA 01230. *Corresponding author - [email protected].

1

Manuscript Editor: Dawn A.T. Phillip 1

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We did not set a specific distance or time to spend collecting at each site; rather, we modified collecting effort for each site and sampled for as much time or distance as it took to explore the available habitats. For instance, in the upper end of Runaway Ghaut, we spent only a few minutes collecting with an aquarium dip net. In the Collins River, we spent at least 2 h hauling a 20-ft (6.1-m) seine in a brackish basin. In all cases, we ceased collecting when we no longer caught new species at a given site. Collecting was restricted to the northern portion of the island (Fig. 1) because areas around the volcano were no longer accessible. We identified most species by comparing our specimens to those included in regional faunal survey collections, recent taxonomic monographs, and original descriptions where necessary. For some fishes, we compared our specimens with those catalogued in the NYSM fish collection; this approach was particularly helpful for fish in the families Eleotridae, Gobiidae, Mugilidae, and Poeciliidae. The specific works we used are referenced within the following species summaries. We extracted genomic DNA from 2 Sicydium individuals with different color patterns using a standard CTAB/chloroform protocol and PCR-amplified the DNA at 2 mitochondrial loci (COI and ATPase 8 and 6). We amplified the COI fragment using primers provided in the Bio-Rad Fish Barcoding Kit (Bio-Rad product #1665100EDU, Bio-Rad Laboratories, Hercules, CA, USA) resulting in a 700-base-pair fragment. We amplified an 840-base-pair fragment of ATPase 8 and 6 using the primers L8933 and H9795 (Kontula et al. 2003). Both primer pairs contained M13 “tails” which were then employed as binding sites for the M13 forward and reverse primers used for sequencing with Sequetech (http://sequetech.com/). Forward and reverse sequences were aligned and edited in BioEdit (Hall 1999) and MEGA 6 (Tamura et al. 2013). Using MEGA 6, we compared both sequences to sequences published in the NCBI database using a BLAST search and used ClustalW (MEGA) to download and align DNA sequences similar to the unidentified Montserrat fish. We edited the alignments, removed poorly aligning flanking regions and constructed maximum likelihood phylogenetic trees (Kimura 2-parameter model, complete deletion, uniform rates, 1000 bootstrap replications). Published sequences were also grouped by identified species and used to calculate mean within- and betweengroup differences. Results A Preliminary List of Freshwater and Brackish-water Fishes on Montserrat Poeciliidae - Poecilia reticulata Peters (Guppy) We identified this species by comparing the fine structure of the male gonopodium with illustrations in Rosen and Bailey (1963). We collected 10 specimens (19–22 mm SL) on 30 December 2013 from an artificial basin next to Runaway Ghaut on the main road (Fig. 1), a locality also reported by Stevens and Waldman (2001). We also saw Guppies in the pools in Runaway Ghaut downstream of the collection site and in artificial ponds in the arboretum, ~150 m south of Runaway Ghaut. We think this species was introduced to control mosquitoes. 2

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Mugilidae - Agonostomus monticola (Bancroft) (Mountain Mullet) We collected 7 Mountain Mullet specimens (33–39 mm SL) from the mouth of Bottomless Ghaut (Fig. 1) on 9 January 2014. This species was also recorded from the Daly and Bunkum Rivers (Stevens and Waldman 2001). Note that the Daly

Figure 1. Montserrat showing collection sites (solid dots) and the streams mentioned in the text. The southern portion of the island was devastated in the 1995 eruption of the Soufrière Hills volcano, and the Belham River no longer exists on the surface (indicated by a dashed line). The inset shows the relative location of Montserrat in the Caribbean Sea. 3

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River (Fig. 1) no longer flows over the surface into the sea due to the 1995 eruption of the Soufrière Hills volcano. Mugilidae - Mugil curema Valenciennes (White Mullet) We collected 1 White Mullet specimen (85 mm SL) from the brackish mouth of the Collins River (Fig. 1) on 3 January 2014 and 5 specimens (29–32 mm SL) from the mouth of Bunkum Ghaut (Fig. 1) on 2 January 2014. Centropomidae - Centropomus pectinatis Poey (Tarpon Snook) We collected 4 Tarpon Snook specimens (24–106 mm SL) from the brackish mouth of the Collins River (Fig. 1) on 3 January 2014. It is likely that other species of Centropomus recorded from nearby islands (Hostache 1992, Lim et al. 2002) are present in the area. Eleotridae - Eleotris amblyopsis (Cope) (Large-scaled Spinycheek Sleeper) The specimens that we collected in Montserrat best fit the description of E. amblyopsis (Pezold and Cage 2001). We collected 2 specimens (37–61 mm SL) from the brackish mouth of Collins River (Fig. 1) on 3 January 2014. The taxonomy of this species has been confused in the past (Pezold and Cage 2001). Eleotridae - Gobiomorus dormitor Lacepède (Bigmouth Sleeper) We collected 1 Bigmouth Sleeper specimen (49 mm SL) from the brackish mouth of the Collins River (Fig. 1) on 3 January 2014. Gobiidae - Awaous banana (Valenciennes) (River Goby) Watson (1996) described the extensive variability and consequent synonomy in Awaous banana. We collected 1 small (20 mm SL) River Goby from the brackish mouth of Collins River (Fig. 1) on 6 January 2014. Stevens and Waldman (2001) listed this species from the Daly River (incorrectly as A. tajasica). Gobiidae - Ctenogobius boleosoma (Jordan and Gilbert) (Darter Goby) We collected 2 Darter Goby specimens (13–32 mm SL) from the brackish mouth of the Collins River on 6 January 2014. Gobiidae - Evorthodus lyricus (Girard) (Lyre Goby) We collected 7 Lyre Goby specimens (18–32 mm SL) from the brackish mouth of the Collins River on 6 January 2014. The collection included both sexes; the males had elongate dorsal spines and 2 red stripes on the tail. Gobiidae - Sicydium sp. or spp. Gobies in this genus are amphidromous. The adults reside in fresh water, often at a substantial distance from the sea, and they spawn in fresh water. The eggs and early larvae drift to the ocean and develop in marine salinities. The small larvae and juveniles then ascend streams (Watson 2000). The identification of the Sicydium specimens was more challenging than any of the other fishes discussed in this paper, and was complicated by the fact that our specimens were small juveniles. We used morphological and mitochondrial DNA data to attempt to identify our specimens to species. There are 7 described species of Sicydium in the Caribbean. Three of these species are distributed along the eastern coast of Central America—S. adelum 4

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Bussing, S. altum Meek, and S. gymnogaster Ogilvie-Grant (Smoothbelly Goby)—and 4 are known from Caribbean islands—S. buscki Evermann & Clark (Busck’s Stone-biting Goby), S. gilberti Watson (Gilbert’s Algae-eating Goby), S. plumieri (Bloch) (Plumier’s Stone-biting Goby), and S. punctatum Perugia (Spotted Algae-eating Goby). When we compared our specimens with published descriptions of color patterns (Meek 1907, Ogilvie-Grant 1884) and tooth shapes (Bussing 1996, Ogilvie-Grant 1884) of Sicydium spp., we eliminated the 3 Central American species from consideration. The specimens we collected at the Dry Waterfall in Soldiers Ghaut on 22 January 2014 exhibited 2 different color patterns. In one form, fish had dense spotting over most of the body (5 specimens, 28–36 mm SL). The other form had a prominent midlateral stripe (2 specimens, 28–32 mm SL). Our original thought was that these specimens represented 2 different species. Watson (2000) summarized descriptions of the 4 species known from Caribbean islands. Sycidium plumieri is widespread in the Caribbean, but has 2 rows of premaxillary teeth whereas our specimens had only 1 row. The meristic counts of the fin elements on our spotted specimens agreed with those of S. punctatum (Watson 2000): 6 spines on the first dorsal fin, 10 branched rays on the second dorsal fin, and 1 spine and 10 rays on the anal fin. Each scale on the body had a concentration of melanophores at the center, thus producing a basic spotted pattern, very similar to the photograph of S. punctatum in Monti et al. (2010). The elaborate color pattern on the second dorsal and anal fins also matched fin-color patterns on the available photographs of S. punctatum (Monti et al. 2010). This species was originally described from Martinique and has been documented from Puerto Rico through the Antilles, including Guadeloupe (Monti et al. 2010) and St. Croix (Smith-Vaniz and Jelks 2014), to Trinidad and Tobago, and most recently to Bahia, Brazil (de Lucena et al. 2013). The banded individuals had slightly tricuspid teeth, and although spotting was present on the body, it was not as extensive as on the other specimens. Additionally, the color pattern on the fins was dissimilar. Morphologically, these 2 specimens appeared most similar to S. buscki, however Smith-Vaniz and Jelks (2014) cautioned that life colors and color patterns can be highly variable in this genus. Watson (2000) reported this species from Cuba, Puerto Rico, and the Dominican Republic, and it was recently reported from St. Croix (Smith-Vaniz and Jelks 2014). Of the 4 Sicydium species present in the Caribbean islands (S. punctatum, S. plumieri, S. buscki, and S. gilberti), only COI sequences were available for S. punctatum and S. plumieri, and only ATPase 8 and 6 sequences were available for S. punctatum, S. buscki, and Sicydium sp. (Puerto Rico, Cook et al. 2010) in the NCBI database (http://www.ncbi.nlm.nih.gov/genbank/). No sequences were available for S. gilberti. Based on COI sequence similarity, the Montserrat Sicydium specimens nested firmly within the group formed by S. punctatum (mean difference between sequences: 0.7%), and did not cluster with S. plumieri (mean difference between sequences: 9.4%). The COI sequence of the banded and spotted individuals differed at 4 out of 672 sites. The spotted specimens’ sequences differed from all published 5

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S. punctatum by 2 out of 654 single-nucleotide polymorphisms (SNPs), while all polymorphic sites in the banded specimens’ sequences were shared with at least one other published S. punctatum COI sequence. No COI data were available for S. buscki or S. gilberti. Based on ATPase 8 and 6 similarity, the Montserrat Sicydium specimens nested firmly within a group that Cook et al. (2010) called Sycidium sp. from Puerto Rico. The Montserrat Sicydium sequences were on average 0.3% different from Sycidium sp. individuals, 3.2% different from S. buscki, and 6.5% different from S. punctatum. The ATPase sequences of the banded and spotted individuals differed at 8 of 851 sites. The striped form differed from published ATPase sequences for Sycidium sp. by one SNP and the spotted form by 6 of 655 SNPs (Cook et al. 2010). Both differred by 67 SNPs (out of 655) from S. punctatum. There were no ATPase data available for S. plumeiri or for S. gilberti. At present, we cannot definitively place a species name(s) on our specimens. Although they appear to resemble S. punctatum and S. buscki, this resemblance is not corroborated by the somewhat conflicting DNA data (assuming the published DNA sequences have been accurately identified). The ATPase-sequence data suggest that the Montserrat specimens are not S. buscki, while the COI data exclude S. plumieri. The data for S. punctatum are less clear: a grouping with S. punctatum is strongly supported by the COI data, while it is strongly rejected by the ATPase data. It is also unclear whether these individuals are from the same species. While they consistently cluster together using both COI and ATPase sequences, their sequences differ from each other (0.6–0.9%). Additionally, the spotted form is unique from other published Sicydium sequences at 8 SNPs (2 COI and 6 ATPase). Juveniles as small as ours have not been described for the Caribbean species, but color pattern and tooth shape are known to change with size (Watson 2000). The best fit was with an unidentified Sicydium from Puerto Rico (Cook et al. 2010); only 1 species has been documented from Puerto Rico, Sicydium gilberti, and it lacks DNA data (Watson 2000). Positive identification of the fish at this site must wait for collection of adult material and/or increased availability of DNA sequences. Haemulidae - Genyatremus luteus (Bloch) (Torroto Grunt) We identified this species following Lindeman and Richards (2006). We collected 3 juvenile specimens (28–32 mm SL) from the brackish mouth of the Collins River on 6 January 2014. It is possible that our record represents a range extension for Torroto Grunt in the Caribbean. Haemulidae - Pomadasys crocro (Cuvier) (Burro Grunt) We identified this species following Lindeman and Richards (2006). We collected 2 specimens (27–28 mm SL) from the brackish mouth of the Collins River on 6 January 2014. Achiridae - Achirus lineatus (L.) (Lined Sole) This family has been considered a subfamily within Soleidae, but Chapleau (1993) elevated it to family status. We collected 1 juvenile Lined Sole specimen (17 mm SL) from the brackish mouth of the Collins River on 6 January 2014. 6

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Discussion The fishes of the brackish and fresh waters of Montserrat documented here are not primary-division fishes (Myers 1938), but rather are secondary freshwater fishes belonging to families with high salinity tolerances (Briggs 1984). This faunal composition is typical for Caribbean islands distant from continental landmasses (Burgess and Franz 1989). Trinidad and Tobago, which is very close to South America, has substantial numbers of primary-division fishes in its fresh waters (Phillip et al. 2013). In our brief survey, we documented 13 species in 7 families, 1 of which is the Guppy. Guppies are distributed on continental South America from western Venezuela east to coastal streams of Guyana (Rosen and Bailey 1963). In the Caribbean, this species is found on many islands, but its status as native is in dispute. Rosen and Bailey (1963) are widely cited as stating that the Guppy is native to the Antilles including Antigua but also stated, “Records from the Lesser Antilles may be due to introductions” (p. 56). Welcomme (1988) listed Guppies as native in Barbados and Antigua. Lim et al. (2002) and Monti et al. (2010) listed this species as introduced on Martinique and Guadeloupe, respectively. Burgess and Franz (1989) stated that the population on Dominica (and other Antillean islands) is likely a result of introductions. Smith-Vaniz and Jelks (2014) listed Guppies as introduced on St. Croix. Poeser et al. (2005) accepted native status for this taxon in northern South America and Trinidad, but stated that all other Caribbean populations were probably introductions. We take the conservative view that Guppies were introduced into Montserrat. The Mountain Mullet is very widespread in freshwaters from North Carolina to Venezuela through Central America and the Caribbean islands (McMahan et al. 2013). This species had generally been considered catadromous (Fievet et al. 2001), with adults that live in fresh water but migrate to the ocean to spawn; however new research indicates that it is amphidromous, with juveniles ascending streams from the sea but with no evidence of adults entering marine environments (Smith and Kwak 2014). This species can be found very far upstream (Burgess and Franz 1989) and is often found in fast-flowing streams (Ribeiro and Villalobos 2010). Lim et al. (2002) had one record of White Mullet in fresh water on Martinique. Caldwell (1966) recorded White Mullet from fresh water on Jamaica, and SmithVaniz and Jelks (2014) reported this species from St. Croix. Fanta-Fiofiloff et al. (1986) showed that juvenile White Mullet were not physiologically stressed at low salinities but could not tolerate fresh water for long periods. However, Alayo (1973) reported White Mullet at a considerable distance inland from the coast in Cuba. Monti et al. (2010) did not list Centropomidae from the fresh waters of Guadeloupe, but Hostache (1992) recorded Centropomus undecimalis (Bloch) (Common Snook) from there. Lim et al. (2010) listed both C. undecimalis and C. ensiferus Poey (Swordspine Snook) as present though rarely encountered in the fresh waters of Martinique. The Large-scaled Spinycheek Sleeper is known from northern South America, Central America, the Gulf Coast of the US north to North Carolina, and the Antilles (Pezold and Cage 2001). Three species of Eleotris in the Western Atlantic have 7

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been confused for many years (Pezold and Cage 2001). Eleotris pisonis (Gmelin) (Spinycheek Sleeper) is restricted to Brazil through eastern Venezuela (Pezold and Cage 2001; thus Hostache’s 1992 record of this species from Guadeloupe and Starmühlner and Therezien’s 1982 record from Martinique are misidentifications). Eleotris perniger (Cope) (Smallscale Spinycheek Sleeper) and E. amblyopsis have both been recorded from the Antilles, and both often inhabit brackish and fresh waters (Pezold and Cage 2001). Monti et al. (2010) reported both species from Guadeloupe, but Lim et al. (2002) listed only E. perniger from Martinique. The Bigmouth Sleeper has a wide distribution from southern Florida and Texas to eastern Brazil, including Central America and the Caribbean islands (Robins et al. 1986). This species was reported from both Martinique and Guadeloupe (Lim et al. 2002, Monti et al. 2010). Smith-Vaniz and Jelks (2014) reported it from flowing fresh water on St. Croix, and Caldwell (1966) reported a specimen from fresh water on Jamaica. Burgess and Franz (1989) wrote that this species can occur far upstream. There is considerable literature that uses the genus Philypnus for this species (Gillet 1983). Although the genus is still valid, it is restricted to eleotrids in the vicinity of the Phillipines (Eschmeyer 1998). The literature on Bigmouth Sleeper contains information under both genera. The River Goby is distributed on the Atlantic and Pacific coasts of Central and South America from Mexico to Venezuela and Peru (Watson 1996). There are records in the Eastern Caribbean from Florida through the Antilles to Trinidad and Tobago. There are museum records of specimens from Jamaica, Puerto Rico, Haiti, Dominican Republic, Dominica, Martinique, St. Vincent, Barbados, and Trinidad and Tobago (Watson 2006). Debrot (2003) listed A. banana from streams on Curaçao. Starmühlner and Therezien (1983) reported Awaous awaous (not a valid name) and Hostache (1992) mentioned an Awaous from Guadeloupe, reported as A. banana by Monti et al. (2010). Awaous tajasica (Lichtenstein) (Sand Fish) has often been listed from the Eastern Caribbean (sometimes misspelled) but its distribution is restricted to coastal Brazil south of the Amazon River (Watson 1996). The River Goby prefers clear, flowing, well-oxygenated streams and can ascend considerable distances upstream (Watson 1996). The Darter Goby is widely distributed in the tropical and subtropical Atlantic Ocean, from North Carolina and the northern Gulf of Mexico to Brazil (Robins et al. 1986). This species has a broad salinity tolerance (Wyanski and Targett 2000), was recorded from brackish water on Jamaica (Caldwell 1966), and has been recorded from the freshwater mouths of rivers (Neal et al. 2009). The Lyre Goby is commonly found in low-salinity waters (Hackney and de la Cruz 1981, Walls 1975). It ranges from the Chesapeake Bay to Rio de Janeiro, Brazil, and is common on the Gulf of Mexico coast and throughout the Antilles (Ginsburg 1931, Wyanski and Targett 1985). Lyre Gobies prefer estuarine habitats with silty substrates and are often encountered in backwater lagoons (Smith-Vaniz and Jelks 2014). The Torroto Grunt is commonly encountered in estuaries, and Randall (1968) reported this species as abundant in the Caribbean, often in brackish water. It has 8

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been recorded from the southern Lesser Antilles south to South America from Colombia to Brazil. Burro Grunts are known from Florida and the northeastern Gulf of Mexico through the Antilles to Brazil. This species was reported from fresh water on Guadeloupe (Starmühlner and Therezien 1983) and Jamaica (Caldwell 1966), and from brackish water in Puerto Rico (Neal et al. 2009). The Lined Sole is found from Florida and the northern Gulf of Mexico south to Argentina (Zapfe 2006). The juveniles are often seen in brackish water and were recorded from fresh water in Nicaragua (Villa 1982). This brief survey documented 13 species in 7 families, 1 of which (the Guppy) is believed to be introduced. Comparison of our faunal list to those of nearby islands (Hostache 1992, Lim et al. 2002, Monti et al. 2010) suggests that ours probably represents only a portion of the fishes present on Montserrat. For instance, Anguilla rostrata (Lesueur) (American Eel) has been reported on Guadeloupe and Martinique and probably occurs on Montserrrat but was elusive during our study. There are probably more species of fishes, such as members of Centropomidae and Haemulidae whose adults are marine but whose juveniles inhabit brackish water, which could probably be collected in the low salinity waters of Montserrat but may only be seasonally present. This paper documents a probable northward range extension of Torroto Grunt. We were unable to find previous records of this species in the vicinity of Montserrat, and our collection during this study appears to be the first from the region. We also document the presence of at least 1 amphidromous goby (Sicydium) species that cannot be identified at present. More specimens and more DNA data may resolve the identity of these specimens. There is very little published data on the general biology of most of the fishes reported here. Information is especially sparse for the juvenile stages of the larger marine species and for the majority of the small gobies we collected. Thus, there remain several research opportunities to further document the freshwater fish fauna of Montserrat and elucidate the biology of many of the known inhabitants. Acknowledgments We collected fishes under the auspices of a memorandum of understanding between Bard College at Simon’s Rock and the Department of the Environment, Montserrat. We especially thank Mr. Gerard Gray for his help in facilitating this study. We also thank Tom Coote, Sam Yarmis, Molly Zeigler, Ian Hetterich, Nathan Sadowsky, Clara Woolner, Naomi Pitman, and Greig Fields for helping collect fishes. Additionally, Taylor Polster and Nate Shoobs went above and beyond to make this study successful. Kathy Schmidt and Alec Schmidt collaborated on the map. We thank Dawn Phillip, Kevin Conway, and an anonymous reviewer for improving the manuscript. Literature Cited Alayo, P.D. 1973. Lista de Peces fluviatiles de Cuba. Torreia. Nuevo Series No. 29:1– 55. Briggs, J.C. 1984. Freshwater fishes and biogeography of Central America and the Antilles. Systematic Zoology 33:428–435. 9

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