24 March 1995 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 108(1):84-97. 1995. angulatus, a new species of snapping from the Gulf of Mexico and northwestern Atlantic, with a redescription of A. heterochaelis Say, 1818 (: : )

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, 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 , 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 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. 2H). articulation of dactyl, upper notch width Coloration. -Overall dark olive green to about 0.04 of propodus length, lower notch brown, occasionally with blue tones around width about 0.1 1 of propodus length; max- orbits, distal half of abdomen, and telson; imum propodus height about 0.46 times body speckled with tiny brown or red-brown length; dactyl length about 0.36 times prop- spots. Major chela dark olive green through- odus length; hand height with dactyl closed out mesial surface, unspeckled; upper and about 0.35 times propodus length; dactyl lower notches either colored as rest or major with entire upper and distal margins round- chela or lightened to pale yellow, rarely with ed, with setae arranged in tufts; opposable blue; inner surface of major chela white or margin of dactyl with molar process tilted pale yellow, occasionally bluish; fingertips at angle to axis of dactyl; sculpture of prop- pale yellow or white. Minor chela dark olive 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Alpheus angulatus, new species. A, major first pereopod (right), dorsal; B, same, ventral; C, minor first pereopod (left); D, left second pereopod; E, left third pereopod; F, left fourth pereopod; G, left fifth pereopod; H, left second pleopod. Bar indicates 5 mm. VOLUME 108, NUMBER 1 89 green. Second to fifth pereopods translucent Material examined. -See Appendix. to reddish, lightly speckled with reddish Diagnosis. - Rostro-orbital depressions brown spots or mottled with red. Telson not abrupt posteriorly. Ventral margin of colored as abdomen, highly speckled with carapace evenly rounded, not pronounced tiny brown or red-brown spots; uropods of- at an angle ventrally posterior to second per- ten bluish throughout, occasionally blue eopods. Minor claw of male balaeniceps- distally. shaped. Spine absent on merus of first pe- Size. -Total length for males ranging 17 reopod. Third, fourth, and fifth pereopods to 38 mm, females 17 to 35 mm. with movable spine on ischium. Etymology. -Specific name derived from Description of neotype. -Rostrum cari- the shape of the ventral margin of the car- nate, reaching 0.5 length of first antennular apace, which contains a pronounced angle segment, extending about as far as base of posterior to the second pereopods. eyestalks. Ocular hoods prominent and un- Range. -The known range for this spe- armed, separated from rostrum by shallow cies is throughout the northern Gulf of Mex- adrostral depression. Ocular hood width ico and northwestern Atlantic as far north 0.25 times length of carapace (Fig. 3A, C). as Beaufort, North Carolina, and as far south Carapace as figured (Fig. 3C), 0.35 times TL as Quintana Roo, Mexico, and Haiti. (range 0.32-0.40 for all specimens exam- Habitat. -Intertidal and shallow waters ined). Carapace smooth, posterior with car- in bays and other quiet waters consisting of diac notch. mud bottoms with oyster clumps or rocks Abdomen as figured (Fig. 3D), 0.52 times and rubble. total length (range 0.45-0.55). Pleura of ab- dominal somites 1 through 5 with ventral Alpheus heterochaelis Say, 18 18 margins rounded. Sixth abdominal pleura Figs. 3, 4 with ventral margin acutely rounded pos- teroventrally. Telson (Fig. 3B) 0.12 times Alpheus heterochaelis Say, 18 18:243. - total length (range 0.09-0.14). Proximal tel- Milne-Edwards, 1837:356. - Kingsley, son width about 0.67 of length, distal width 1878a: 194, 1878b:329, 1879:417.- about 0.5 of length; 2 pairs of dorsal spines, Brooks & Hemck, 1892:376, pl. 2.- anteriormost pair positioned about 0.5 of Couticre, 19 10:485.-Verrill, 1922:76, pl. telson length, posteriormost pair positioned 22, figs. 1, 2, 4a-c; pl. 24 figs. 7, 7a; pl. almost 0.75; posterior margin convex, with 30, figs. 1-la, It, 2a-2e; pl. 33, figs. 2 pairs of lateral spines, space between spines 1, 2. -Holthuis, 1959: 102.- Williams, with double row of setae (Fig. 3B). 196566, fig. 54, 1984:95, fig. 65.-Chace, Antennular peduncle with stylocerite 1972:67.-Coelho & Ramos, 1972: 148.- dorsally flattened and terminating anteri- Christoffersen, 1984:200, figs. 5-7. orly at a sharp point which reaches anterior Crangon heterochaelis. -Hay & Shore, margin of first antennular segment; second 1918:386, fig. 8.-Schmitt 1935:144. antennular segment longer than first and Holotype. -Not extant, Amelia Island, about 1.4 times as long as third (Fig. 3A, Nassau County, Florida. C). Neotype.- 1 male, 3 1 mm total length, Antenna1 spine reaching end of antennal Fort Saint George Inlet (8 miles south of peduncle, and just overreaching antennal Amelia Island), Duval County, Florida, in- scale; spine faintly convex at distal tip, tertidal habitat consisting of hard mud and straight or slightly concave at middle. Basal clumps of oysters, coll. M. R. McClure and segment of antennal peduncle armed ven- L. S. McClure, 25 May 1992, USNM trolaterally (Fig.' 3A, C). 268646. Mandible with 9 teeth. Third maxilliped PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Alpheus heterochaelis Say, 18 18. A-L, Adult male (neotype) from Fort St. George Inlet, 8 miles south of Amelia Island, Florida (TL = 3 1 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

Fig. 4. Alpheus heterochaelis. A, major first pereopod (right), dorsal; B, same, ventral; C, minor first pereopod (left); D, left second pereopod; E, left third pereopod; F, left fourth pereopod; G, left fifth pereopod; H, left second pleopod; I, Adult female from Beaufort, NC, minor first (left) pereopod. Bar indicates 5 mm. 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON reaching end of antenna1peduncle; terminal Coloration. -Overall olive green to article setose. brown, often with blue tones around orbits, First pereopods strongly chelate and un- distal half of abdomen, and telson; body equal; merus unarmed distoventrally. Ma- speckled with tiny brown or red-brown jor chela (Fig. 4A, B) thick; propodus length spots. Major chela olive green with pale area 0.45 times total body length (range 0.29- at lower center of palm; raised areas of palm 0.61); upper and lower margins deeply and opposable margin of fixed finger very notched proximal to articulation of dactyl, dark green; upper and lower notches pale upper notch width about 0.07 of propodus yellow to pale blue; inner surface of major length, lower notch width about 0.10 of chela white to pale blue; fingertips pale yel- propodus length; maximum propodus height low or white. Minor chela olive green. Sec- about 0.45 times length; dactyl length about ond to fifth pereopods translucent to pale 0.35 times propodus length; hand height blue, lightly speckled with reddish brown with dactyl closed about 0.32 times prop- spots. Telson colored as abdomen; exopod odus length; dactyl with upper distal margin of uropods blue on distal segment; endopod rounded, opposable margin with molar pro- of uropods blue distally. cess tilted at angle to axis of dactyl; sculp- Size. -Neotype total length 3 1 mm. To- ture of propodus as figured, with upper and tal length for males ranging 16 to 45 mm, lower notches forming saddle-like depres- females ranging 18 to 57 mm. sions (extending 0.07 and 0.14 times length Range. -Lower Chesapeake Bay south- of propodus, respectively) into the lateral ward and westward into the Gulf of Mexico; surfaces of propodus; upper notch posi- Cuba; Cura~ao(Williams 1984); Bermuda tioned about 0.5 the length of propodus, (Verrill 1922); Surinam (Chace 1972); lower notch positioned about 0.67; distal southward to Paraiba (Christoffersen 1984). end of propodus rounded. Minor chela sex- Habitat. -Intertidal and shallow waters ually dimorphic (Fig. 4C, I). Male I-'111nor in bays and other quiet waters consisting of chela balaeniceps-shaped, having setose mud bottoms with oyster clumps or rocks crest on dactyl and accessory crest on op- and rubble. posable margin of the propodus; propodus 0.32 times total body length (range 0.22- Discussion 0.40 for specimens examined); propodus height about 0.24 times lengih iil males, 0.19 Although Alpheus angulatus and A. het- times length in females; dactyl length about erochaelis are morphologically similar, they 0.45 times that of propodus length in males, are not closely related. Alpheus heterochae- and about 0.5 in females. lis and A. angulatus were referred to as Al- Second pereopods (Fig. 4D) slender and pheus group A and group B, respectively in weakly chelate; carpus subdivided into 5 ar- McClure & Greenbaum (1994), where the ticles decreasing in length as follows (num- two species are shown to be highly divergent bered from the proximal end): 1, 2, 5, 3 = electrophoretically. Alpheus heterochaelis 4. Third to fifth pereopods (Fig. 4E, F, G) was the most common species that the au- with ventral movable spine on ischium; thor collected from Texas and from North dactyls simple; third pereopod propodus Carolina. This species differs from the other with 9 or 10 stout spines, fourth with 8 stout Edwardsii-groupAlpheus mentioned herein spines; fifth pereopod with 7 spines on prop- in that A. heterochaelis possesses a balaen- odus, with several lateral bands of comb- iceps-type minor claw in males, and a mov- like setae extending on distal half. Second able spine on the ventral surface of the is- pleopod of male with appendix masculina chium of the fifth pereopod. Alpheus het- shorter than appendix interna (Fig. 4H). erochaelis is further distinguished from A. VOLUME 108, NUMBER 1 93 angulatus in that the rostro-orbital area in the color pattern, the two species are diffi- the former lacks posteriorly abrupt adros- cult to distinguish. Both A. armillatus and tral depressions and lacks the broad trian- A. angulatus have the typical A. armillatus- gular area of the carapace from which the type rostrum, consisting of a broad, flat- rostrum terminates posteriorly. tened triangular area posteriorly, and pos- Material of A. estuariensis was examined teriorly abrupt adrostral furrows. Milne-Ed- from Port Isabel and Gilchrist, Texas, and wards' (1837) description of A. armillatus from Mississippi Sound, Alabama, as well was brief and lacking in detail. However, as the Gulf of Mexico material used by the locality was given as the Antilles. The Christoffersen (1 984). Alpheus estuariensis species was named "the banded Alpheus" was not found in this survey, and thus is for its conspicuous banded color pattern; probably of uncommon occurrence in the this pattern is lacking in A. angulatus. Ver- northern Gulf of Mexico. Alpheus estuarien- rill (1922) stated that A. armillatus from sis is distinguishable from A. heterochaelis Bermuda, when recently caught, was iden- in that it lacks the balaeniceps-type minor tifiable by its conspicuous transverse bands claw in males, and lacks the movable spine of white on the body and rings of color on on the ischium of the fifth pereopod. In ad- the legs and antennae, and provided a pho- dition, the minor claw of A. estuariensis is tograph (plate 20, fig. 4b) showing such a very slender relative to that of A. hetero- pattern. Zeiller (1974, p. 76) provided a col- chaelis. or photograph ofA. armillatus revealing the Alpheus angulatus was the most common conspicuous banded pattern on the abdo- species that the author collected from Flor- men, with the body and chelae colored over- ida and from Louisiana, and has apparently all brown speckled with white. In A. an- been mistaken for Alpheus heterochaelis in gulatus, the body is darker and speckled with a number of museum collections. This is dark red-brown spots, with speckling lack- because they are both similarly colored, and ing on the chelae. commonly taken in a single sampling (both Hay & Shore (19 18) listed A. angulatus have been collected simultaneously at most as A. arrnillatus from North Carolina. Al- of the author's collecting localities). though the author was informed that Hay In one case, Alpheus angulatus has been & Shore's (19 18) collection may no longer mistaken for A. estuariensis by Christoffer- exist (R. B. Manning, pers. comm.), the col- sen (1984). One of the specimens referred or description by Hay & Shore for the A. to as A. estuariensis (USNM 98 137) from armillatus from Beaufort, North Carolina, Louisiana represents A. angulatus and not matches that of A. angulatus. In addition, A. estuariensis. Alpheus angulatus differs the figures provided by Hay & Shore of A. from A. estuariensis in that the former has arrnillatus match A. angulatus with respect a more robust minor claw, whereas the spine to the shape of the rostrum and minor chela. on the merus of the first pereopod is lacking They also mentioned the rarity of this spe- in A. estuariensis. In addition, the upper and cies at Beaufort, North Carolina, which also lower notches of the major chela are wider confirms the authors' findings (one A. an- of A. estuariensis than those of A. hetero- gulatus individual was caught with 54 A. chaelis and of A. angulatus, and the distal heterochaelis individuals). The color pat- end of the propodus is sharply truncated in tern difference between the Carolinian A. A. estuariensis. arrnillatus and the tropical A. armillatus also Alpheus angulatus has previously been re- led Knowlton (1970) to speculate that the ferred to as the Carolinian A. armillatus (Hay Carolinian form may represent a separate & Shore 19 18; Knowlton 1970; Chace 1972; species. Williams 1965, 1984). Except by examining The likelihood that A. armillatus, as cur- 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rently known, represents a species complex chium of the third and fourth pereopods, seems more plausible now than ever, since and the "A. armillatus-type" rostrum is ab- differences in color patterns have been dem- sent in A. nuttingi. onstrated to have systematic importance in Alpheus normanni Kingsley, a "Macro- Alpheus (Knowlton & Mills 1992). Hendrix chins-group" species, was examined from (1 97 1) described three color morphs of A. South Padre Island, Texas, Dauphin Island, arrnillatus from Miami, Florida (the normal Alabama, and Crystal River, Florida. This banded pattern, a blue-grey morph, and a species is mentioned here because it too has yellow morph), and suggested that they may occasionally been labeled as A. heterochaelis represent distinct species. The author was in some collections. This is probably due to unable to locate the specimens used by Hen- both A. normanni and A. heterochaelis hav- drix (197 l), so confirmation is lacking if any ing the balaeniceps-type minor claw in males of his color morphs of A. armillatus from or have both been taken simultaneously. Al- the Miami region correspond with A. an- pheus normanni is distinguished from the gulatus. Edwardsii-group species mentioned herein Hendrix (1 97 1) mentioned in his descrip- in that the major chela ofA. normanni lacks tion that A. arrnillatus (all three color a lower notch, being only notched dorsally; morphs included) had a movable ventral the dorsal notch is sharply undercut prox- spine on the ischium of the fifth pereopod, imally. In addition, A. normanni has an- which is lacking in A. angulatus. Specimens gularly produced ocular hoods anteriorly, of A. armillatus from the Smithsonian-Bre- not smooth or evenly rounded as in these din Expedition (Chace 1972) examined from Edwardsii-group species (Hendrix 197 1). It the Leeward Islands (Lesser Antilles) con- is also brighter green in color relative to A. tain such a spine (appearing more conspic- heterochaelis and A. angulatus. uous in smaller specimens), and also differ The geographic range of A. heterochaelis from A. angulatus in the shape ofthe ventral is apparently widespread throughout the margin of the carapace; the latter having a temperate and tropical western Atlantic. pronounced ventral angle posterior to the Despite past taxonomic ambiguity sur- second pereopods (Fig. 1C) lacking in the rounding this species, records confirm the Antillian specimens. Other specimens iden- presence of A. heterochaelis from eastern tified as A. armillatus by Chace (1972) from and southeastern North America, the Ca- the east coast of the Yucatan peninsula, ribbean region, eastern Central America, and Mexico, had characteristics matching A. an- northeastern South America. However, A. gulatus (color pattern information was lack- arrnillatus should now be considered to have ing). In addition, specimens from Haiti a tropical distribution. Previous records of identified as A. armillatus by Coutihe A. armillatus from temperate North Amer- (MNHN-Na 2 17 1) match the characteris- ica correspond with A. angulatus; the latter tics present for A. angulatus. differs from the former in color pattern and Hendrix (1 97 1) provided a description of in some discrete characters. The geographic A. nuttingi (Schmitt, 1924)as the first record range of A. angulatus includes the Gulf of of that species from the continental U.S.A. Mexico and northwestern Atlantic, and ap- His description differs from the original de- pears to extend into the Caribbean region. scription of the holotype in that the latter Additional ranges of A. angulatus are not lacks a spine on the mens of the first per- currently known. The geographic range of eopods; this spine is present both on A. nut- A. estuariensis appears just as Christoffersen tingi and A. angulatus. However, A. angu- (1984) described, although this species does latus differs from A. nuttingi (both of Schmitt not seem to be abundant in the northern 1924, and Hendrix 197 1) in that A. angu- Gulf of Mexico. latus possesses movable spines on the is- Originally, a single Edwardsii-group Al- VOLUME 108, NUMBER 1 95

pheus species, A. heterochaelis, was known was funded in part by Sigma Xi Grants-In- in coastal waters from the northern Gulf of Aid of Research GIAR 99 12 (to M. R. Mc- Mexico and northwestern Atlantic. Prior to Clure) and Advanced Research Program this study two such species, A. heterochaelis ARP grant 160603 (to M. K, Wicksten). and A. estuariensis, had been reported from the northern Gulf of Mexico. In this study, Literature Cited A. angulatus was found to be equally as common as A. heterochaelis and more com- Abele, L. G., & W. Kim. 1986. An illustrated guide mon than A. estuariensis in the northern to the marine decapod of Florida. Gulf of Mexico and northwestern Atlantic. Florida State University, Tallahassee, 326 pp. Brooks, W. K., & F. H. Hemck. 1892. The embry- The new species resembles A. nuttingi, and ology and metamorphosis of the Macroura.- strongly resembles A. armillatus, both of Memoirs, National Academy of Sciences 5:323- which may in fact consist of more than one 576, pl. 1-57. species throughout the western Atlantic. In Chace, F. A. 1972. The shrimps of the Smithsonian- short, the taxonomic and biogeographic re- Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crus- lationships existing for western Atlantic al- tacea: Decapoda: Natantia). - Smithsonian pheids is far from being resolved. Contributions to Zoology 98: 179 pp. Christoffersen, M. L. 1979. Decapod Crustacea: Al- pheoida. Rtsultats Scientifiquesdes Campagnes Acknowledgments de la Calypso, Fascicule 11. Campagne de la Calypso au large des CBtes Atlantiques de The author thanks the following people 1'AmCrique du Sud (1961-1962). I. Number for their help in the collection of snapping 36.-Annales de 1'Institut OcCanografique, new shrimp: L. S. McClure, M. K. Wicksten series 5 5, fascicule supplement:297-377. (Texas A&M University), E. C. Rives, J. S. . 1984. The western Atlantic snapping shrimps Whorff, D. Hockaday and F. Judd (Uni- related to Alpheus heterochaelis Say (Crustacea, Caridea), with a description of a new species.- versity of Texas-Pan American Marine Field PapCis Avulsos de Zoologia 35: 189-208. Station, South Padre Island, Texas), M. Coelho, P. A., & M. de A. Ramos. 1972. A constru@o Dardeau (MESC), M. Demetz, L. Edmiston e a distribuciglo de fauna de decipodos do lit- and G. Bailey (Apalachicola National Es- toral leste do Ami.rica do Sul entre as latitudes tuarine Research Reserve), D. Felder and de 5"N e 39"s.-Trabalhos do Instituto Ocean- ograficos, Universidade Federal, Pernambuco, T. Zimmerman (University of Southwest- Recife 13: 133-326. ern Louisiana), and B. Kirby-Smith and K. Conover, M. R., & D. E. Miller. 1978. The impor- Reinsel (Duke Marine Laboratory, Beau- tance of the large chelae in the temtorial and fort, North Carolina). S. K. Davis, I. F. pairing behaviour of the snapping shrimp Al- Greenbaum, and H. Wilson (Texas A&M pheus heterochaelis. -Marine Behavior and Physiology 5: 185-192. University) provided the cryogenic field Coutikre, H. 1910. The snapping shrimps (Alphei- equipment and storage space for shrimp in dae) of the Dry Tortugas, Florida. -Proceedings their ultracold freezers. The following mu- of the United States National Museum 37:485- seum curators were helpful with the loaning 487. of snapping shrimp specimens: R. B. Man- Hay W. P., & C. A. Shore. 19 18. The decapod crus- taceans of Beaufort, N.C., and the surrounding ning (USNM), D. K. Camp (FBSC), M. Dar- region.-Bulletin of the United States Bureau of deau and T. S. Hopkins (MESC), A. Chaney Fisheries 35:369-475. (TAI), D. Guinot and N. Nguyen (MNHN), Hendrix, G. Y. 197 1. A systematic study of the and R. Harrell (LU). Thanks also to M. K. Alpheus (Crustacea: Decapoda: Alpheidae) in Wicksten, R. E. Knowlton, N. Knowlton, south Florida. Unpublished Ph.D. dissertation, University of Miami, Coral Gables, Florida, 185 and J. Goy for their advice on alpheid sys- PP. tematics. M. K. Wicksten, J. Goy, and I. F. Holthuis, L. B. 1959. The Crustacea Decapoda of Greenbaum provided reviews. This study Suriname (Dutch Guiana).-Zoologische Ver- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

handelingen Uitgegevin Door Het Rijksmu- bados-Antigua expedition from the University seum Van Natuurlijke Historie Te Leiden 44: of Iowa in 19 18.-University of Iowa Studies 1-296. in Natural History 10:65-99. Kingsley, J. S. 1878a. A synopsis of the North Amer- -. 1935. Crustacea Macrura and Anomura of ican species of the genus Alpheus. -Bulletin of Porto Rico and the Virgin Islands.-New York the United States Geological and Geographical Academy of Science 15: 125-227. Survey 4: 189-199. Verrill, A. E. 1922. Decapod crustacea of Bermuda . 1878b. List of decapod Crustacea of the At- Part 11, Macrura.-Transactions of the Con- lantic coast, whose range embraces Fort Ma- necticut Academy of Arts and Sciences 26:l- con.-Proceedings of the Academy of Natural 179. Sciences of Philadelphia 30:3 16-330. Williams, A. B. 1965. Marine decapod crustaceans . 1879. On a collection of Crustacea from Vir- of the Carolinas.-Fishery Bulletin 65: 1-297. ginia, North Carolina, and Florida, with a re- -. 1984. Shrimps, lobsters, and crabs of the vision of the genera of Crangonidae and Palae- Atlantic coast of the Eastern United States, monidae.-Proceedings of the Academy of Nat- Maine to Florida. Smithsonian Institution Press, ural Sciences of Philadelphia 31:383-427. Washington, D.C. Knowlton, N., & D. K. Mills. 1992. The systematic Wilson, E. B. 1903. Notes on the reversal of asym- importance of color and color pattern: evidences metry in the regeneration of the chelae in Al- for complexes of sibling species of snapping phm heterochaelis. -Biological Bulletin 4: 197- shrimp (Caridea: Alpheidae: Alpheus) from the 210. Caribbean and Pacific coasts of Panama.-Pro- Zeiller, W. 1974. Tropical marine of ceedings of the San Diego Society of Natural southern Florida and the Bahama Islands. John History 18: 1-5. Wiley & Sons, New York, 132 pp. Knowlton, R. E. 1970. Effects of environmental fac- tors on the larval development of Alpheus het- Appendix erochaelis Say and Palaemonetes vulgaris (Say) (Crustacea Decapoda Caridea), with ecological Material Examined notes on larval and adult alpheidae and Palae- Alpheus heterochaelis. -Collected Material: Texas: monidae. Unpublished Ph.D. dissertation, Uni- South Padre Island, Laguna Madre, north of Brazos- versity of North Carolina, Chapel Hill, 544 pp. Santiago Pass near old Coast Guard Station, 0.5 m, n . 1973. Larval development of the snapping = 3 1, 17 Apr 1993; Port Aransas, north jetty at Saint shrimp Alpheus heterochaelis Say, reared in the Joseph Island, intertidal, n = 19, 18 Dec 1991; Gal- laboratory.-Journal of Natural History 1973: veston Bay, off Sportsman's Road, 0.5 m, n = 14, 14 273-306. Sep 199 1, South jetty, 0.5 m, n = 1, 17 Jan 1992. Mellon, D., Jr., & P. J. Stephens. 1978. Limb mor- Alabama: Dauphin Island, Mobile Bay, 0.5 m, n = 4. phology and function are transformed by con- Florida: Pensacola, Escambia Bay, 0.5 m, n = 1, 18, tralateral nerve section in snapping shrimps.- 19 Mar, 1993; Apalachicola Bay, St. George Island State Nature 272:246-248. Park, 1.0-1.5 m, n = 3, 25 May, 1992; Panacea, n = McClure, M. R., & I. F. Greenbaum. 1994. Bio- 5, 26 Feb, 1992; Fort St. George Inlet, 8 miles south chemical variation in Alpheus (Decapoda, Car- of Amelia Island, intertidal, n = 1 (neotype, USNM idea, Alpheidae) from the coast of Texas: evi- 268646), 25 May, 1992. NorthCarolina: Beaufort, near dence for cryptic species.-The Southwestern Duke Marine Lab, 0.5 m, n = 54, 25, 26 Jun 1993. Naturalist 39:63-66. Borrowed Material: Texas: USNM 72186, Ranson Is- Milne-Edwards, H. 1837. Histoire naturelle des land, n = 1; USNM 82 1 16, Galveston, Offat's Bayou, CrustacCs, comprenant l'anatomie, la physiol- n = 2; TAI 166, Salt Lake, Indianola, n = 1; TAI 167, ogie et, la classification de ces animaux. 2:l- Port Isabel, n = 1; TAI 169, Bahia Azul, n = 1; TAI 532. 55 1, Bahia Azul, n = 1. Mississippi: USNM 64243, Nolan, B. A., & M. Salmon. 1970. The behavior and Dier Island, n = 2. Alabama: MESC 6 179-367 1, Mo- ecology of snapping shrimp (Crustacea: Alpheus bile Bay, n = 2; MESC 6179-3672, Pt. Pines, n = 2; heterochaelis and Alpheus normanni).-Forma MESC uncat., Dauphin Island, airport marsh, n = 2. et Functio 2:289-335. Florida: USNM 57635, Key West, n = 2; MESC 6179- Say, T. 18 17-1818. An account of the Crustacea of 5143, St. Joseph Bay, n = 1; FBSC I 110, Bush Key, the United States.-Journal of the Academy of Pinellas Co., n = 3; FBSC I 188, Bush Key, n = 2; Natural Sciences of Philadelphia l(18 17):57-63, FBSC I 364, Egmont Key, Pinellas Co., n = 8; FBSC 65-80,97-101,155-169; (1818):235-253,313- I 1496 Boca Ciega Bay, n = 1; FBSC I 2177, Tampa 319, 374-401, 423-444, 445-458, plate 4. Bay, n = 1; FBSC I 2177 Tampa Bay, n = 1; FBSC I Schmitt, W. L. 1924. Report on the Macrura, Ano- 5126, Bush Key, n = 1; FBSC I 6244, Pumpkin Bay, mura and Stomatopoda collected by the Bar- Collier Co., n = 1; FBSC I 6338, Pakalatchee Bay, n VOLUME 108, NUMBER 1 97

= 7; FBSC I 6378 Mullet Key, Pinellas Co., n = 1; 1 7601, "Marine Research Lab", Pinellas Co., n = 2; FBSC I 7628, Sawyer and Sistu Keys, Boca Ciega Bay, FBSC I 7745 (subsample), Crystal River, n = 1; FBSC n = 12; FBSC I 7685, Crystal River, n = 2; FBSC I I, 2 miles south of Hillsburough River, n = 1. South 14419, W. McIllvaine Bay, n = 1; FBSC I 7745, Crystal Carolina: USNM 63549, Jericho Creek, n = 7. North River, n = 12. Georgia: USNM 181864, 181865, Carolina: USNM 12806 1, Beaufort, Lennoxville Point, 181866, 18 1867, Sapelo Island, n = 1 each. n = 1. Haiti: MNHN-Na 2 17 1, n = 2. Mexico: USNM Alpheus angulatus, new species.-Collected Mate- 135891, Quintana Roo, Ascension Bay, n = 12. rial: Texas: South Padre Island, Laguna Madre, north Alpheus estuariensis.-Borrowed Material: Brazil: of Brazos-Santiago Pass near old Coast Guard Station, USNM 144014, Sao Paulo, paratypes, n = 3; USNM 0.5 m, n = 2, 4 Jul 1992 (holotype, USNM 266804 25800, Mamanquape, paratype, n = 1; USNM 222042, and paratype, USNM 266805); n = 2, 17 Apr, 1993; n = 4; MESC 6 179- 1046 1, Raiba, n = 4. CUBA: USNM Port Aransas, north jetty at Saint Joseph Island, in- 96455, Laguna de Paso Malo, n = 3. U.S.A.: Texas: tertidal, n = 3, 18 Dec 199 1; Galveston Bay, off Sports- USNM 63546, Jetty at Galveston Bay (Evennann 189 1 man's Road, 0.5 m, n = 2, 14 Sep 1991, South jetty, coll.), n = 2; TAI uncat., Port Isabel, n = 4; LU uncat., 0.5 m, n = 4, 17 Jan, 1992. Louisiana: Tembonne Bay, East Galveston Bay at Gilchrist, n = 4. Louisiana: Lighthouse island south of LUMCON-Cocodrie, 0.5 USNM 98137, Lake Pontchartrain, n = 1. Alabama: m, n = 34, 19 Jul 1993; Florida: Pensacola, Escambia MESC uncat., Mississippi Sound, n = 1. Florida: USNM Bay, 0.5 m, n = 45, 18, 19 Mar 1993; Apalachicola 90957, Duval Co., n = 4. Bay, St. George Island State Park, 1.0-1.5 m, n = 28, Alpheus armillatus. -Borrowed Material: Lesser An- 25 May 1992; Panacea, n = 1, 26 Feb 1992; Fort St. tilles: USNM 135869, Leeward Islands, Guadaloupe, George Inlet, 8 miles south of Amelia Island, intertidal, n = 14; USNM 135870 (subsample), Leeward Islands, n = 34, 25 May 1992. North Carolina: Beaufort, near Antigua, n = 4. Duke Marine Lab, 0.5 m, n = 1, 25, 26 Jun 1993. Alpheus nuttingi. -Borrowed Material: USNM Borrowed Material: Texas: TAI 168, Mustang Island, 68700, Holotype and 2 additional specimens from Bar- n = 1. Louisiana: USNM 98138, Lake Ponchartrain, bados, W. Schmitt, 19 18. n = 1. Alabama: MESC uncat., west end Dauphin Is- Alpheus normanni. -Collected Material: Texas: land (subsample), n = 3. Florida: USNM 57635, Key South Padre Island, Laguna Madre, north of Brazos- West, n = 1; FBSC I 1293 Tierra Verde, Pinellas Co., Santiago Pass near old Coast Guard Station, 0.5 m, n n = 1; FBSC I 2901, St. George Bay near Apalachicola, = 3,17 Apr 1993. Borrowed Material: Alabama: MESC n = 3; FBSC I 293 1, Magnolia Beach, Andrews Bay, uncat., "Mussels 2-l", n = 3; MESC uncat., west end n = 2; FBSC I 3626, Mullet Key, Pinellas Co., n = 3; Dauphin Island (subsample), n = 1. Florida: FBSC I FBSC I 6297, Gullivan Bay, Collier Co., n = 5; FBSC 7745 (subsample), Crystal River, n = 1.