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Alpheus Angulatus, a New Species of Snapping Shrimp from the Gulf of Mexico and Northwestern Atlantic, with a Redescription of A

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Alpheus Angulatus, a New Species of Snapping Shrimp from the Gulf of Mexico and Northwestern Atlantic, with a Redescription of A 24 March 1995 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 108(1):84-97. 1995. Alpheus angulatus, a new species of snapping shrimp from the Gulf of Mexico and northwestern Atlantic, with a redescription of A. heterochaelis Say, 1818 (Decapoda: Caridea: Alpheidae) Matthew R. McClure Business, Science, & Math Division, Lamar University-Orange, 4 10 Front Street, Orange, Texas 77630, U.S.A. Abstract. -Two species of the Edwardsii-group of snapping shrimp, Alpheus heterochaelis Say and A. estuariensis Christoffersen, have been recorded from coastal estuarine habitats of the Gulf of Mexico and Atlantic coasts of the United States. A new species, A. angulatus, has been discovered to inhabit these habitats across this range and is described and illustrated. Thenew species is morphologically similar to A. armillatus, the latter being a tropical species. A redescription with illustrations of A. heterochaelis is also provided herein. The Edwardsii species group of Alpheus Carolina and south Florida, and proposed consists of at least 10 western Atlantic spe- that A. heterochaelis comprised two or more cies (Chace 1972; Christoffersen 1979, 1984; species. From a reevaluation of Alpheus in- Williams 1984; Abele & Kim 1986), of habiting shallow-estuarine environments of which Alpheus heterochaelis Say, 1818, is the western Atlantic, Christoffersen (1 984) the most abundant and widely distributed. suggested the existence of a species com- Alpheus heterochaelis was originally de- plex, and described a new species, A. estu- scribed from Amelia Island, Nassau Coun- ariensis (holotype from the Rio Potengi es- ty, Florida, and has been reported to range tuary, Rio Grande do Norte, Brazil), rang- from the lower Chesapeake Bay to Aransas ing from the east coast of Florida into the County, Texas, Cuba, Curagao, Surinam, Gulf of Mexico from Mississippi to Texas; and possibly to Slo Paulo, Brazil (Chace Cuba; Dominican Republic; Trinidad; Cu- 1972, Williams 1984). ragao; and from Cearh to Paranh, Brazil. A variety of studies have been done con- Christoffersen (1 984) further concluded that cerning the biology of A. heterochaelis (for the Alpheus occurring in the northern Gulf example, Wilson 1903, Nolan & Salmon of Mexico represents A. estuariensis, and 1970, Knowlton 1973, Conover & Miller described the range of A. heterochaelis to be 1978, Mellon &Stephens 1978). Despite the from North Carolina to Paraiba, Brazil. abundance of studies on A. heterochaelis, However, using Christoffersen's (1984) key, the geographic limits and the systematic sta- specimens from Galveston were identified tus of this species remain problematic. Based by the author as A. heterochaelis. Thus, plus on apparent misidentifications of West In- a reexamination of museum specimens of dian and Brazilian specimens, Chace (1 972) A. estuariensis from the northern Gulf of questioned the occurrence of A. hetero- Mexico suggest the existence of taxonomic chaelis south of Surinam. Knowlton (1973) ambiguity concerning the Edwardsii group noted differences in egg size between labo- of Alpheus from the coastal waters of the ratory reared A. heterochaelis from North Gulf of Mexico and northwestern Atlantic. VOLUME 108, NUMBER 1 8 5 An assessment of allozymic variation of armillatus is not available; it is likely that specimens of Alpheus from Texas revealed the specimens have been lost. As the only the existence of two discrete and markedly detailed descriptions of A. heterochaelis are different gene pools (McClure & Green- from the Carolinas (Christoffersen 1984), baum 1994).Additional allozymic data (un- comparisons to this species were from ma- published) indicate that these forms are terial from the type locality and from the sympatric throughout the northern Gulf of Carolinas. For A. arrnillatus, specimens from Mexico and Northwestern Atlantic coasts the type locality (Antilles) were used as a as far north as Beaufort, North Carolina. reference. Type material of the two species The present study was designed to provide described here have been deposited in the morphological descriptions and identifica- USNM. tions of the two electrophoretically identi- fied species. Alpheus angulatus, new species Figs. 1, 2 Materials and Methods Crangon armillatus. -Hay & Shore, 19 18: 386, fig. 9. (not Alpheus arrnillatus Milne- Snapping shrimps were collected at low Edwards, 1837). tide by dip net in intertidal and shallow sub- Alpheus estuariensis. -Christoffersen, 1984: tidal habitats consisting of sand or mud bot- 19 1 (in part, see discussion). toms covered with oyster clumps or rocks. Alpheus arrnillatus.- Chace, 1972:62 (in Shrimps were collected in coastal waters part, see discussion). from south Texas to North Carolina (see Appendix for localities). Shrimps were Ho1otype.-Male, 28 mm TL, on mud stored at - 80°C to fully preserve color pat- under rocks and rubble, South Padre Island, terns. Starch-gel electrophoresis (McClure Texas, Laguna Madre just north of Brazos- & Greenbaum 1994) was used to sort the Santiago Pass, coll. M. K. Wicksten, 4 Jul individuals into discrete electromorphic 1992, USNM 266804. classes. Morphological characters were then Material examined. -See Appendix. assessed and compared with museum ma- Diagnosis. -Rostra-orbital depressions terial and descriptions in the literature. To- abrupt posteriorly. Ventral margin of car- tal Length (TL) of specimens is the com- apace pronounced at an angle ventrally pos- bined measurements of the carapace, ab- terior to second pereopods. Minor claw of dominal, and telson lengths. male not balaeniceps-shaped. Spine present Museum specimens of A. heterochaelis, on merus offirst pereopod. Third and fourth A. estuariensis, A. nuttingi, and A. armil- pereopods with movable spine on ischium, latus were obtained from the following in- lacking on fifth pereopod. stitutions: Marine Research Division, Flor- Description of holotype. -Rostrum reach- ida Department of Natural Resources, St. ing 0.5 length of first antennular segment; Petersburg, Florida (FBSC); Marine Envi- in form of raised crest extending beyond ronmental Sciences Consortium, Tuscaloo- base of eyestalks and widening into flat tri- sa, Alabama (MESC); Texas A&I Univer- angular area (Fig. 1A). Ocular hoods prom- sity, Kingsville, Texas (TAI); National Mu- inent and unarmed, separated from rostrum seum of Natural History, Smithsonian In- by adrostral depressions abrupt posteriorly stitution, Washington, D.C. (USNM); (Fig. 1A). Ocular hood width 0.27 times Museum national D'Histoire naturelle, Par- length of carapace. Carapace as figured (Fig. is, France (MNHN); Lamar University col- lC), 0.35 times TL (range 0.31-0.40 for all lection, Beaumont, Texas (LU). The origi- specimens examined). Carapace smooth, nal type material of A. heterochaelis and A. posterior of carapace with cardiac notch; 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Alphm angulatus, new species. Adult male (holotype) from South Padre Island, Texas (TL = 28 mm). A, anterior carapace and antennae, dorsal; B, sixth abdominal somite, telson and uropods, dorsal; C, carapace and antennae, lateral; D, abdominal pleura, lateral. Bar indicates 5 mm. VOLUME 108, NUMBER 1 ventral margin acute ventrally just posterior odus as figured (Fig. 2A, B), with upper and to articulation of second pereopod. lower notches forming saddle-like depres- Abdomen as figured (Fig. ID), 0.55 times sions (extending about 0.07 and 0.1 1 times total length (range 0.44-0.56). Pleura of ab- length of propodus, respectively) into the dominal somites 1 through 5 with ventral lateral surfaces of propodus; upper notch margins rounded. Sixth abdominal pleura positioned about 0.5 the length of propodus, with ventral margin acute posteroventrally. lower notch positioned about 0.61, having Telson (Fig. 1B) 0.1 1 times total length tuft of setae on proximal end oflower notch; (range 0.09-0.14). Proximal telson width angular area of upper mesial surface of fixed 0.67 of length, distal width about 0.5 of finger rounded, having granular texture and length; 2 pairs of dorsal spines, anterior- tufts of setae; distal end of propodus round- most pair positioned about 0.4 of telson ed, with tufts of setae. Minor chela robust length, posteriormost pair positioned at and setose, with setal tufts increasing dis- 0.67; posterior margin convex, with 2 pairs tally on both propodus and dactyl (Fig. 2C); of lateral spines, space between spines with not sexually dimorphic, lacking balaeniceps double row of setae. setose crest on dactyl; propodus 0.33 times Antennular peduncle with stylocerite total body length (range 0.15-0.37 for spec- dorsally flattened, terminating anteriorly at imens examined); propodus height about sharp point which reaches anterior margin 0.30 times length in both sexes; dactyl length of first antennular segment; second anten- about 0.5 times that of propodus length in nular segment longer than first and about both sexes. 1.5 times as long as third (Fig. lA, C). Second pereopods slender and weakly Antenna1 spine reaching end of antennal chelate (Fig. 2D); carpus subdivided into 5 peduncle, and just overreaching antennal articles decreasing in length as follows scale; concave at middle and faintly convex (numbered from the proximal end); 1, 2, 5, at distal tip. Basal segment of antennal pe- 3 = 4. Third and fourth pereopods with duncle armed with spine ventrolaterally (Fig. ventral movable spine on ischium (Fig. 2E, 1A, C). F); dactyli simple; third pereopod propodus Mandible with 10 teeth. Third maxilliped with 7 stout spines and 3 or 4 smaller al- reaching just beyond end of antennal pe- ternating lateral spines, fourth with 8 stout duncle; terminal article setose. spines and 5 or 6 alternating lateral spines. First pereopods (Fig. 2A, B, C) strongly Fifth pereopod (Fig. 2G) with 8 or 9 spines chelate and unequal; merus armed with spine on the propodus and several lateral bands distoventrally. Major chela thick, setose of comb-like setae extending on distal hale distally; propodus length 0.5 1 times total ischium without spine. Second pleopod of body length (range 0.28-0.56); upper and male with appendix masculina shorter than lower margins deeply notched proximal to appendix interna (Fig.
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