12 2 Abstract—Western Atlantic synodon- A new species of western Atlantic lizardfi sh tid species were studied as part of an ongoing effort to reanalyze Ca- (Teleostei: Synodontidae: Synodus) and ribbean shorefish diversity. A neigh- bor-joining tree constructed from resurrection of Synodus bondi Fowler, 1939, cytochrome c oxidase I (COI) data as a valid species from the Caribbean with revealed 2 highly divergent genetic lineages within both Synodus inter- redescriptions of S. bondi, S. foetens (Linnaeus, medius (Agassiz, 1829) (Sand Diver) 1766), and S. intermedius (Agassiz, 1829) and S. foetens (Linnaeus, 1766) (In- shore Lizardfish). A new species, Synodus macrostigmus, is described Benjamin W. Frable (contact author)1, 2 for one of the S. intermedius lin- Carole C. Baldwin1 eages. Synodus macrostigmus and 1 S. intermedius differ in number of Brendan M. Luther lateral-line scales, caudal pigmen- Lee A. Weigt1 tation, size of the scapular blotch, and shape of the anterior-nostril Email address for contact author: [email protected] flap. Synodus macrostigmus and S. intermedius have overlapping geo- 1 National Museum of Natural History graphic and depth distributions, but Smithsonian Institution S. macrostigmus generally inhabits P.O. Box 37012 deeper water (>28 m) than does S. Washington, D.C. 20013-7012 intermedius and is known only from 2 coastal waters of the southeastern Department of Fisheries and Wildlife United States and the Gulf of Mex- Oregon State University ico, in contrast to those areas and 104 Nash Hall the Caribbean for S. intermedius. Corvallis, Oregon 97331 Synodus bondi Fowler, 1939, is res- urrected from the synonymy of S. foetens for one of the S. foetens ge- netic lineages. The 2 species differ in length and shape of the snout, num- The aulopiform lizardfish family respectively (Jeffers et al., 2008; ber of anal-fin rays, and shape of the Synodontidae is represented in the Manjarrés et al., 2008). Populations anterior-nostril flap. Synodus bondi western Atlantic by 3 genera: Syno- of Synodus foetens (Linnaeus, 1766) and S. foetens co-occur in the central dus (5 species recognized before this (Inshore Lizardfi sh) are estimated to Caribbean, but S. bondi otherwise study), Saurida (4), and Trachino- be at fully exploited levels as bycatch has a more southerly distribution cephalus (1). Lizardfi shes are ben- in Gulf of Mexico shrimp-trawl fi sh- than does S. foetens. Redescriptions thic predators in numerous ecosys- eries (Garcia-Abad et al., 1999; Wells, are provided for S. intermedius, S. tems, including coral reefs, estuaries, 2007; Jeffers et al., 2008). Proper foetens, and S. bondi. Neotypes are and reef structure or sandy bottom management and ecological inves- designated for S. intermedius and areas on continental shelves (Ander- tigation of commercially fi shed spe- S. foetens. A revised key to Synodus species in the western Atlantic is son et al., 1966; Cressey, 1981; Ran- cies require an accurate understand- presented. dall, 2009). Once thought to occupy ing of species diversity. For example, a mid-trophic position and employ Collette et al. (1978) discovered that a sit-and-wait predation strategy, the Caribbean and Brazilian popula- adult lizardfi shes now are known to tions of the commercially important occupy a high trophic position—apex Scomberomorus maculatus (Spanish in some systems—as active hunters Mackerel) constitute a distinct spe- feeding primarily on other predatory cies, which they named S. brasilien- fi shes (Cruz-Escalona et al., 2005). sis (Serra). Without this systematic Despite having no commercial study, populations of S. brasiliensis value as food fi shes, Synodus spp. would still be managed under the Manuscript submitted 11 February 2012. Manuscript accepted 8 February 2013. and Saurida brasiliensis (Largescale same plan as S. maculatus. Fish. Bull. 111:122–146 (2013). Lizardfi sh) are caught as bycatch in Publication date: 28 March 2013. shrimp-trawl fi sheries in the west- for North Carolina. Job 3: character- doi 10.7755/FB.111.2.2 ern Atlantic, accounting for 1.5% and ization of the near-shore commercial shrimp trawl fishery from Carteret 1.8% of total-catch biomass in the County to Brunswick County, North 1 The views and opinions expressed Gulf of Mexico and North Carolina, Carolina. Southeast Fisheries Science or implied in this article are those of the Center, Miami, FL, 29 p. [Available on- author (or authors) and do not necessar- line from http:/ /www.sefsc.noaa.gov/se- ily refl ect the position of the National 1 Brown, K. 2009. Interstate fisheries dar/download/SEDAR20-ASMFC-DW09. Marine Fisheries Service, NOAA. management program implementa tion pdf?id=DOCUMENT] Frable et al.: Description of a new species of Synodus in the western Atlantic Ocean 123 Scientists have periodically investigated the system- to document living color patterns, and subsequently a atics of synodontids since Linnaeus (1758) described small sample of tissue from the trunk musculature for Esox synodus in the mid-18th century. Throughout the genetic analysis. Voucher specimens were fi xed in 10% 19th century, researchers described 17 Synodus species formalin and ultimately preserved in 75% ethanol for in the western Atlantic (Anderson et al., 1966; Meek, archival storage. 1884; Norman, 1935). In the most recent comprehensive Measurements were taken to the nearest 0.1 mm treatments of western Atlantic lizardfi shes (Anderson with Mitutoyo digital calipers (Mitutoyo Corp., Ja- et al., 1966; Russell, 2003), 5 species of Synodus have pan2). Measurements and counts follow Hubbs and La- been recognized. Ongoing research to evaluate diver- gler (1964) and Randall (2009), except as noted below. sity of the Caribbean ichthyofauna with DNA barcod- Length of the anterior-nostril fl ap was measured from ing (Hebert et al., 2003) and traditional morphological the posterior tip of the anterior nostril to the distal investigation have led to the recent discovery of many end of the fl ap when depressed. For the S. intermedius new cryptic fi sh species and the resurrection of several group only, the length of the scapular blotch was mea- formerly synonymized ones in what were thought to be sured on an anterior–posterior axis at its greatest ex- well-studied taxa (Baldwin and Weigt, 2012; Baldwin panse. For the S. foetens group only, the width of the et al., 2009, 2011; Tornabene et al., 2010; Victor, 2007, adipose lid was measured as the maximum distance 2010). In the course of the present work, discrepancies between the bony orbit and distal edge of the lid. were revealed between barcode data and the currently Numbers of vertebrae and dorsal-, anal-, pectoral-, accepted species classifi cation of western Atlantic Syn- and caudal-fi n rays were counted from digital radio- odus lizardfi shes. Specifi cally, each of 2 Synodus spe- graphs or preserved specimens. Scales fl anking the dor- cies, S. intermedius (Agassiz, 1829) (Sand Diver) and sal- and anal-fi n bases are half the size of other trunk S. foetens comprise 2 distinct cytochrome c oxidase I scales and are reported as half-scales. (COI) lineages. Tissue samples were stored in saturated salt buffer The purpose of this study was to reconcile genetic (Seutin et al., 1990). DNA was extracted from up to lineages with the nominal species of western Atlantic approximately 20 mg minced, preserved tissue through Synodus. Through comparative morphological study, an automated phenol: chloroform extraction on an Au- our fi rst goal was to determine if the “extra” genetic togenprep965 DNA extraction system (Autogen, Hol- lineages correspond with morphologically distinct spe- liston, MA) using the mouse tail tissue protocol to a cies and, if so, to assess whether they represent pre- fi nal elution volume of 50 µL. In the polymerase chain viously synonymized or undescribed species. Herein, reaction (PCR), 1 µL of extracted DNA was used in a we resurrect and redescribe Synodus bondi Fowler, 10 µL reaction with 0.5 U BioLine (Bioline USA, Inc., 1939, from the synonymy of S. foetens and describe S. Boston, MA) Taq polymerase, 0.4 µL 50 mM MgCl2, 1 macrostigmus as a new species distinct from S. inter- µL 10× buffer, 0.5 µL 10 mM deoxyribonucleotide tri- medius. We establish neotypes for S. intermedius and phosphate, and 0.3 µL 10 µM each primer FISH-BCL S. foetens and redescribe both species. We discuss pre- (5′-TCAACYAATCAYAAAGATATYGGCAC) and FISH- liminary evidence of population structure within S. foe- BCH (5′-TAAACTTCAGGGTGACCAAAAAATCA). The tens, S. synodus (Red Lizardfi sh), Saurida brasiliensis, PCR theromcycle protocol was: 1 cycle of 5 min at 95°C; and Trachinocephalus myops (Snakefi sh) and species- 35 cycles of 30 s at 95°C, 30 s at 52°C, and 45 s at 72°C; level genetic structure within Synodus poeyi (Offshore 1 cycle of 5 min at 72°C; and a hold at 10°C. PCR prod- Lizardfi sh). Finally, we provide a revised key for the 7 ucts were purifi ed with ExoSAP-IT (Affymetrix, Santa species of Synodus found in the western Atlantic. Clara, CA) with 2 µL 0.2× enzyme and incubated for 30 min at 37°C. The reaction was then inactivated for 20 min at 80°C. Sequencing reactions were performed with Materials and methods 1 µL of this purifi ed PCR product in a 10 µL reaction that contained 0.5 µL primer, 1.75 µL BigDye reaction Specimens for genetic analysis were collected in Tobago buffer (Life Technologies Corp., Carlsbad, CA), and 0.5 (Trinidad and Tobago), Turks and Caicos Islands, the µL BigDye in the thermal cycler for 30 cycles of 30 s Bahamas, Curaçao, Belize, North Carolina, South Caro- at 95°C, 30 s at 50°C, 4 min at 60°C, and then were lina, and Florida. Type material and additional speci- held at 10°C after completion of cycles.