AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2953, 10 pp., 3 figs., 3 tables October 3, 1989

Cyprinodon nichollsi, a New Pupfish from , and Species Characteristics of C. bondi Myers (Teleostei: Cyprinodontiformes)

MICHAEL LEONARD SMITHI

ABSTRACT The Jaragua pupfish, Cyprinodon nichollsi n. sp., paniola, is redescribed and characterized on the is described from a hypersaline coastal , La- basis of its tapered caudal peduncle, presence of a guna de Oviedo, Jaragua National Park, Domin- toothplate in the position of the first pharyngo- ican Republic. It is distinguished by its large size, branchial bone, and high number of circumpe- deep body, small cleithral scale, and small cleithral duncular scales. process. Cyprinodon bondi Myers, also from His- INTRODUCTION The cyprinodontid genus Cyprinodon La- Recent collecting shows that there are sev- cepede comprises about 30 pupfishes distrib- eral pupfishes in nonmarine habitats of the uted in arid regions of northern Mexico and island of Hispaniola, in addition southwestern United States and in Atlantic to the long-described, but poorly known, C. coastal habitats of North and Middle Amer- bondi Myers. The systematics and variation ica (Turner and Liu, 1977). The diversity of offorms from Lake , southwestern pupfishes in mainland habitats of North Hispaniola, are topics of a separate study. America is well documented (Miller, 1981; The species characteristics of an additional Humphries and Miller, 1981; Humphries, pupfish from Laguna de Oviedo, western Do- 1984; Minckley and Minckley, 1986), but the minican Republic, are reported here, and the number and systematic status offorms in the new species is compared to other Antillean Caribbean region have not been established. species of Cyprinodon.

1 Kalbfleisch Assistant Curator, Department of Herpetology and Ichthyology, American Museum of Natural His- tory.

Copyright © American Museum of Natural History 1989 ISSN 0003-0082 / Price $1.50 2 AMERICAN MUSEUM NOVITATES NO. 2953

Fig. 1. Cyprinodon nichollsi. AMNH 58278, 1 iolotype, male, 75.3 mm standard length. Patricia J. Wynne, del.

Myers (1935) described as new the species METHODS C. bondi on the basis ofthree specimens col- Measurements were taken as described by lected in Etang Saumatre, . An antici- Miller (1948) and Humphries (1984) with ex- pated, fuller description never appeared, and ceptions as follows. Body depth was taken at the species has seldom been mentioned since, the dorsal-fin origin. Depth of the caudal pe- except to note that it appears to stand out duncle is the distance between the first dorsal sharply from the "C. variegatus complex" and ventral procurrent rays of the caudal fin. (Miller, 1962). The collection of additional Mouth width is the greatest distance across material shows that it is indeed one of the the dental arcade of the lower jaw (the lips most distinctive species ofCyprinodon, char- are disregarded). acterized by a number of features that are Every ray in the dorsal and anal fins is inferred to be derived. counted as one (i.e., the last two are not com- bined). Vertebrae are counted from radio- graphs and cleared-and-stained specimens ACKNOWLEDGMENTS and include the hypural plate as one vertebra; I thank C. E. Badgley (University ofMich- the first precaudal vertebra is taken to be that igan), K. J. Lazara (U.S. Merchant Marine which bears the first complete hemal arch, Academy), G. R. Smith (UMMZ), and W. C. detected in radiographs by a white spot where McNiff for assistance in the field and K. W. the hemal processes fuse. Gill rakers were Nicholls for support offieldwork. G. H. Bur- counted on the first gill arch of the right side gess (UF), K. J. Lazara, R. R. Miller (UMMZ), and include all rudiments on both upper and and G. R. Smith reviewed the manuscript lower limbs. All counts and measurements and P. J. Wynne prepared figures 1 and 2. F. are taken from the left side. For meristic char- X. Geraldes (Department of Fisheries Re- acters, the value for the holotype is indicated sources, Secretary of State for Agriculture) in boldface. granted permission to collect fishes in the Do- Referred material is deposited in the Acad- minican Republic, and M. Valverde Podesta emy of Natural Sciences, Philadelphia (National Bureau of Parks) gave permission (ANSP), American Museum of Natural His- to conduct research in national parks. Re- tory (AMNH), Florida Museum of Natural search was supported by the National Science History (UF), and University of Michigan Foundation (BSR 8600683). Museum of Zoology (UMMZ). 1 989 SMITH: HISPANIOLAN PUPFISHES 3

Fig. 2. Cyprinodon bondi. UF 45043, male, 46.8 mm standard length. Patricia J. Wynne, del.

SYSTEMATICS shown in figure 1 and proportional measure- ments are given in table 1. Body deep, strong- Cyprinodon nichollsi, new species ly compressed. Predorsal profile usually in- Jaragua pupfish dented at occiput, but strongly convex from Figures 1, 3; tables 1-3 occiput to dorsal-fin origin. Dorsal-fin origin HOLOTYPE: AMNH 58278, male, 75.3 mm behind midbody, slightly in advance of pel- SL, sandy beach at Laguna de Oviedo, 4 km vic-fin origin. Mouth wide, terminal. east ofOviedo on Hwy. 44, Jaragua National First dorsal-fin ray slender and short, never Park, ; W. C. McNiff, spinelike as in some populations of C. var- K. J. Lazara, and M. L. Smith, 9 Sept. 1987. iegatus. Dorsal-fin rays 11(23 counts), 12(21), PARATYPES: AMNH 58279, 135 juveniles, 13(6). Anal-fin rays 10(11), 11(37), 12(2). males and females, including 10 cleared-and- Pectoral-fin rays 15(6), 16(28), 17(16). Pel- stained, 19.8-69.1 mm SL; USNM 300935, vic-fin rays 5(1), 6(10), 7(38), 8(1). Branched 10 males and females, 24.6-46.0 mm SL; col- caudal-fin rays 11(3), 12(39), 13(6), 15(1). lected with the holotype. AMNH 58280, 10 Branchiostegal rays, from alizarin prepara- males and females, 25.0-55.1 mm SL; tions, 6(10). Gill rakers 19(2), 20(8), 21(20), UMMZ 214865, 107 juveniles, males and 22(12), 23(7), 24(1). Scales between dorsal females, 14.3-66.7 mm SL; UF 79181, 10 and anal fins 11(1), 12(26), 13(21), 14(2). males and females, 22.3-49.5 mm SL; beach Other meristic characters are given in tables at Laguna de Oviedo, 4 km east of Oviedo, 2 and 3. Jaragua National Park, Dominican Republic; Pectoral fin round and short in adults, fail- G. R. Smith and M. L. Smith, 8 Jan. 1988. ing to reach pelvic-fin origin in specimens DIAGNOSIS: A large Cyprinodon (to 75 mm greater than 30 mm in standard length. Ab- SL) with a long head and snout, wide mouth, domen usually fully scaled, but elliptical strongly convex predorsal profile, first pha- scaleless areas sometimes occur on either side ryngobranchial toothplate absent, caudal fin of the ventral midline near bases of pectoral truncate, pectoral fin round, not reaching or- fins. Exposed scale margins entire or slightly igin of pelvic fins, 14-16 circumpeduncular crenate, never laciniate as in C. laciniatus and scales, 23-26 lateral-series scales, scales en- some populations of C. variegatus. Scales be- tire or slightly crenate, never laciniate. Large tween pelvic fins imbricate, not developed as size and small pectoral fin are derived. Com- a lingulate interpelvic process as in C. bondi parisons to particular congeners are given in and Floridichthys carpio. Scapular scale not discussion. notably enlarged relative to first scales in lat- DESCRIPTION: General morphology and eral series; underlying cleithral process broad, pigmentation of preserved specimens are expanded posteriorly beyond the pectoral-fin 1989 SMITH: HISPANIOLAN PUPFISHES 5

TABLE 2 Frequency Distribution of Vertebrae in Antillean Species of Cyprinodon

Species Precaudal Caudal Total Locality 10 11 12 13 13 14 15 16 24 25 26 27 bondia E. Saumatre 1 60 11 2 27 43 20 52 laciniatusb Bahamas 14 1 3 12 2 13 nichollsic L. Oviedo 1 44 11 8 37 11 5 35 16 variegatusd Great Inagua 24 6 7 19 4 3 21 6 a AMNH 37341, 58281; UF 45043, 45045; UMMZ 214866; USNM 100960, 100961. b ANSP 110366. c AMNH 58278, 58279. dAMNH 43282. radials, not greatly enlarged as in C. nazas and contact organs develop as lateral bony and C. variegatus. projections from the first seven anal-fin rays. Acoustico-lateralis system on head con- The fifth and sixth rays of the anal fin are sists ofcombination ofpit organs, canals, and swollen in males. Coloration (below) also dis- pores as follows (data from left side ofhead): tinguishes the sexes. mandibular 2(50); preopercular 6(1), 7(42), COLORATION: Live individuals at the type 8(6), 9(1); preorbital 2(1), 3(2), 4(47). Su- locality are milky white over the entire body praorbital canal uninterrupted, opening to the and fins. Milky coloration, which obscures surface through seven pores (N = 50). the patterns described below, is apparently SEXUAL DIMORPHISM: Cyprinodon nichollsi induced by turbidity which is high at Laguna is strongly dimorphic, a feature that distin- de Oviedo, and it quickly disappears when guishes it from C. bondi. The fins of males, the fish are transferred to clear seawater. The especially the dorsal fin, are longer than in life coloration given below is based on fish females. With increasing size, males become held one hour in seawater; the same color- very compressed and deep-bodied (fig. 1), de- ation is shown by stocks from the type lo- veloping a high, convex predorsal crest, and cality maintained in aquaria for several the caudal peduncle is longer in males than months. in females. In breeding males, scales on the Live adult males are silver on the back and head and sides of the body become ciliate, sides. The breast, periproct, and lower sides

TABLE 3 Frequency Distribution of Circumpeduncular and Lateral Scales in Antillean Species of Cyprinodona Around peduncle In lateral series Species 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 bondi E. Saumatre 5 2 41 1 13 20 11 2 laciniatus Bahamas 6 1 7 1 1 5 7 2 nichollsi L. Oviedo 3 3 46 3 26 18 3 variegatus Great Inagua 1 29 4 21 5 a Material as in table 2. 1989 SMITH: HISPANIOLAN PUPFISHES 5

TABLE 2 Frequency Distribution of Vertebrae in Antillean Species of Cyprinodon

Species Precaudal Caudal Total Locality 10 11 12 13 13 14 15 16 24 25 26 27 bondia E. Saumatre 1 60 11 2 27 43 20 52 laciniatusb Bahamas 14 1 3 12 2 13 nichollsic L. Oviedo 1 44 11 8 37 11 5 35 16 variegatusd Great Inagua 24 6 7 19 4 3 21 6 a AMNH 37341, 58281; UF 45043, 45045; UMMZ 214866; USNM 100960, 100961. b ANSP 110366. c AMNH 58278, 58279. dAMNH 43282. radials, not greatly enlarged as in C. nazas and contact organs develop as lateral bony and C. variegatus. projections from the first seven anal-fin rays. Acoustico-lateralis system on head con- The fifth and sixth rays of the anal fin are sists ofcombination ofpit organs, canals, and swollen in males. Coloration (below) also dis- pores as follows (data from left side ofhead): tinguishes the sexes. mandibular 2(50); preopercular 6(1), 7(42), COLORATION: Live individuals at the type 8(6), 9(1); preorbital 2(1), 3(2), 4(47). Su- locality are milky white over the entire body praorbital canal uninterrupted, opening to the and fins. Milky coloration, which obscures surface through seven pores (N = 50). the patterns described below, is apparently SEXUAL DIMORPHISM: Cyprinodon nichollsi induced by turbidity which is high at Laguna is strongly dimorphic, a feature that distin- de Oviedo, and it quickly disappears when guishes it from C. bondi. The fins of males, the fish are transferred to clear seawater. The especially the dorsal fin, are longer than in life coloration given below is based on fish females. With increasing size, males become held one hour in seawater; the same color- very compressed and deep-bodied (fig. 1), de- ation is shown by stocks from the type lo- veloping a high, convex predorsal crest, and cality maintained in aquaria for several the caudal peduncle is longer in males than months. in females. In breeding males, scales on the Live adult males are silver on the back and head and sides of the body become ciliate, sides. The breast, periproct, and lower sides

TABLE 3 Frequency Distribution of Circumpeduncular and Lateral Scales in Antillean Species of Cyprinodona Around peduncle In lateral series Species 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 bondi E. Saumatre 5 2 41 1 13 20 11 2 laciniatus Bahamas 6 1 7 1 1 5 7 2 nichollsi L. Oviedo 3 3 46 3 26 18 3 variegatus Great Inagua 1 29 4 21 5 a Material as in table 2. 6 AMERICAN MUSEUM NOVITATES NO. 2953 of the head are white, but the pectoral fins SL; to 18.6 g, fixed in ethanol). Only C. maya, sometimes become light yellow in dominant an elongate but less deep-bodied species, sur- males. A terminal black margin is prominent passes it in length (to 85 mm SL; Humphries on all fins; width of the terminal caudal-fin and Miller, 1981). The two species further band is about equal to the diameter of the resemble each other in sharing a large head, pupil. Interradial membranes of the median wide mouth, and heavy lower jaw which are fins are hyaline, but melanophores are scat- characters associated with greater size in pup- tered across the dorsal fin giving it a dusky fishes (Humphries and Miller, 1981; Hum- color. The sides of breeding males are some- phries, 1984). Cyprinodon nichollsi differs times marked by 7-16 indistinct vertical bars, from C. maya in its deeper body (34- 48 vs. although bars are often absent. The nape is 27-39% SL, respectively), greater degree of marked by an iridescent blue sheen that be- lateral compression, and presence oftwo pairs comes intense in dominant males. In ethanol, of mandibular pores in all material exam- silver and blue colors are replaced by buff. ined. Mandibular pores are always absent in Lateral bars and the black borders of fins be- C. maya (Humphries and Miller, 1981). come prominent, and isolated melanophores Cyprinodon variegatus Lacepede, the most produce a pattern of fine speckles over all widespread and variable member of the ge- parts of the body and fins. Pigmentation nus, is usually understood to include six sub- shown in figure 1 is based on a specimen in species distributed along the coast of North ethanol. America from Massachusetts to the Yucatan Live adult females are also silver on the Peninsula and in the Caribbean islands from sides and back. Fins are hyaline except for a the Florida Keys to the Bahamas and Cuba prominent ocellus, about as large as the eye, (Miller, 1962; Humphries and Miller, 1981). in the trailing corner of the dorsal fin. The Cyprinodon nichollsi is distinguished from ocellus consists of a central patch of white this complex of forms by its smaller cleithral surrounded by an irregular black margin. Lat- scale, smaller cleithral process, larger head, eral bars are more conspicuous than in males; and wider mouth. In all populations currently 6-19 irregular vertical bars span the middle classified in C. variegatus, the head is small third ofthe body in a series from the gill cleft and wedge-shaped in lateral view, and the to the caudal base. Dots in lateral series, as mouth is small and narrow, thereby contrast- seen in C. bondi, are absent. Cheeks are ing to those ofboth C. nichollsi and C. bondi marked by a patch oflarge melanophores be- (fig. 3). The Antillean populations of C. var- low the eye. Markings are similar in spirits iegatus are usually characterized by a weak except that silver is reduced and white in the to moderate lappet covering the dorsal mar- dorsal-fin ocellus disappears. gin ofthe iris and by the presence of strongly HABITAT: Cyprinodon nichollsi is reported crenate or laciniate scales on the crests ofthe only from Laguna de Oviedo, a hypersaline caudal peduncle; both features are lacking in coastal lagoon on the southern coast of His- C. nichollsi. paniola. The lagoon is separated from the sea Cyprinodon nichollsi resembles C. bondi in by a narrow barrier which, according to local its large size, large head, and wide mouth. It account, is sometimes breached allowing ma- differs in having fewer circumpeduncular and rine fishes to enter. On 9 Sept. 1987, salinity lateral-series scales (table 3), strongly convex was estimated to be 69 ppt using a YSI con- (rather than straight) predorsal profile, round ductivity meter. Cyprinodon nichollsi is pectoral fin (rather than narrow and elongate, abundant in shallow water (to 1.5 m) over figs. 1 and 2), truncate caudal fin, and no sand, mud, and mats of filamentous algae toothplate on the first pharyngobranchial (N where it was taken by seine. = 10). DISCUSSION: The new species is assigned to Cyprinodon nichollsi differs from C. lacin- the genus Cyprinodon with which it shares a iatus Hubbs and Miller (New Providence Is- terminal black band on the male caudal fin land, Bahamas) in having a round pectoral and thickened rays 5 and 6 in the male anal fin that fails to reach the pelvic-fin origin, fin. Cyprinodon nichollsi is distinguished from fewer vertebrae (table 2), fewer scales around most pupfishes by its greater size (to 75 mm the caudal peduncle and in lateral series (table 1989 SMITH: HISPANIOLAN PUPFISHES 7

C(R A E

D F Fig. 3. Dorsal and lateral views of heads of(A, B) Cyprinodon variegatus, AMNH 76600, male, 34.3 mm SL (Fort Johnson, S.C.); (C, D) C. nichollsi, AMNH 58279, male, 35.1 mm SL (Oviedo, Dominican Republic); (E, F) C. bondi, AMNH 58281, male, 36.2 mm SL (Lago El Fondo, Dominican Republic). All drawn to same scale.

3), larger size (C. laciniatus reaches 50.7 mm ADDITIONAL SPECIMENS: AMNH 37341, 12 SL; ANSP 110366), and in lacking laciniate males and females including one cleared-and- scales. stained, 37.4-62.7 mm SL, Etang Saumatre, ETYMOLOGY: The new species is named for Haiti; W. Beebe and J. Tee-Van, 27 April Kenneth W. Nicholls. 1927. AMNH 58281, 50 juveniles, males, and females, 12.2-38.4 mm SL; UMMZ 214866, 67 juveniles, males, and females, Cyprinodon bondi Myers 13.5-49.5 mm SL; Lago El Fondo (=Etang Hispaniolan pupfish Saumatre) 1 km east of Haitian border at Figures 2, 3; tables 1-3 Jimani, Dominican Republic; G. R. Smith Cyprinodon bondi Myers, 1935 (original descrip- and M. L. Smith, 10 Jan. 1988. UF 45045, tion, Etang Saumatre, Haiti). Hubbs and Miller, 6 males including one cleared-and-stained, 1942 (listed). Miller, 1962 (listed). Turner and 49.0-61.0 mm SL, south shore Etang Sau- Liu, 1977 (distribution in map). Miller, 1981 matre, 3 km W Malpasse, Haiti; J. B. Thorb- (distribution in map). Burgess, 1983 (in part, jarnarson, 18 Aug. 1983. ANSP 2 Etang Saumatre only). Lazara, 1984 (synony- 159201, my). Thorbjarnarson, 1988 (Etang Saumatre). males and 1 female, 46.0-62.5 mm SL, Etang Saumatre, 2.5 km W Malpasse, Haiti; W. C. HOLOTYPE: USNM 100960, female, 62.7 McNiff, K. J. Lazara, and D. Fromm, 11 Jan. mm SL, Etang Saumatre, Haiti; R. M. Bond, 1986. 20 Feb. 1933. DIAGNOSIS: A large Cyprinodon (to 63 mm PARATYPES: USNM 100961, 10 juveniles SL) distinguished by the following characters: and females, 18.7-34.9 mm SL; collected with 18-20 scales around the caudal peduncle; the holotype. caudal peduncle strongly tapered posteriad, 8 AMERICAN MUSEUM NOVITATES NO. 2953 constricted at base ofcaudal fin; median scales gans develop as bony lateral projections of between pelvic fins elongate or lingulate, free fin rays in the male anal fin. As in all Cy- at their distal end; caudal fin slightly emar- prinodon, the fifth and sixth anal-fin rays of ginate; sides marked by a lateral series ofdots males are swollen compared to those that fol- on 3-4 longitudinal rows of scales at mid- low. body; toothplate and well-developed teeth Live adults are olivaceous on the back and present in the position of the first pharyn- silver on the lower sides, belly, and underside gobranchial bone. of the head. The sides are crossed by 7-18 DESCRIPrION: Proportional measurements irregular vertical bars that are narrower than are given in table 1. Body strongly com- the silver interspaces. Scales on the upper pressed and rhomboidal in lateral view (fig. back are outlined by dark melanophores re- 2); predorsal profile straight, no indentation sulting in a reticulated pattern. The species at occiput; caudal peduncle strongly tapered is distinguished from other Cyprinodon by with its narrowest point at insertion of the three or four series of dots (one per scale) in procurrent caudal-fin rays. Dorsal-fin inser- midlateral series. tion at midbody opposite pelvic-fin insertion. The fins ofjuveniles and females are most- Lower jaw oblique to nearly vertical; cleft of ly clear except for a white-and-black ocellus mouth oblique. Dorsal margin of iris entire, in the posterior corner of the dorsal fin. In not bearing a process or lappet. other Cyprinodon, the dorsal-fin ocellus dis- First dorsal-fin ray slender and short, never appears as males reach maturity, but in C. spinelike. Dorsal-fin rays 11(22), 12(26), bondi it is retained in mature males as large 13(2). Anal-fin rays 10(3), 11(43), 12(6). Pec- as 45 mm SL. The terminal black margin of toral-fin rays 15(4), 16(19), 17(25), 18(2). Pel- the male caudal fin is narrower than the pupil. vic-fin rays 6(7), 7(41). Branched caudal-fin HABITAT: Etang Saumatre, the only known rays 10(1), 11(3), 12(34), 13(7), 14(4). Bran- habitat of C. bondi, is one of a series of chiostegal rays 6(10). Gill rakers 18(2), 19(3), in a graben that extends across Hispaniola 20(5), 21(9), 22(4), 23(3). Vertebral counts from Port-au-Prince Bay, Haiti, to Barahona are given in table 2 and scale counts in ta- Bay, Dominican Republic (Woodring et al., ble 3. 1924). The lake lacks a surface outlet and Pectoral fins narrow and long, surpassing fluctuates in volume and salinity (8-10 ppt) pelvic-fin insertion in individuals ofall sizes. in response to rainfall (Woodring et al., 1924; Breast and prepelvic area fully scaled. Ex- Bond, 1935). Aquatic plants are abundant posed scale margins mostly entire, becoming near freshwater springs and in bays with crenate in large specimens, particularly on freshwater input, but vegetation in open water crests of caudal peduncle. Inter-pelvic ap- is limited to a few halophytic algae and vas- pendage composed of several median lingu- cular plants (Thorbjarnarson, 1988). The late scales covering inner margins of pelvic shoreline is bare or fringed with buttonwood fins. Scapular scale not noticeably enlarged mangrove, Conocarpus erectus. compared to first scales in lateral series; Adults ofC. bondi occur in open water over cleithral process among the least developed gravel, sand, or mud; juveniles and young in the genus. adults are abundant in vegetation where they Number ofpores ofcephalic acoustico-lat- are the dominant fish in terms of number. eralis system: mandibular 0(3), 1(1), 2(43); Native fishes collected with C. bondi are preopercular 6(1), 7(44), 8(2); preorbital 2(1), Cichlasoma haitiensis (Cichlidae), Dormita- 3(4), 4(40), 5(1), 6(1). Supraorbital canal un- tor maculatus (Eleotridae), Gobionellus sp. interrupted, opening to surface through seven (Gobiidae), Limia melanotata, L. tridens, and pores. Gambusia hispaniolae (Poeciliidae). A native SEXUAL DIMORPHISM AND COLORATION: belonid, Strongylura timucu, has also been External differences between the sexes are less collected in the lake (ANSP 163387). marked in C. bondi than in most Cyprinodon. DISCUSSION: Several ofthe characters given The dorsal and anal fins tend to be longer in in the diagnosis above appear to be derived. males than in females. Ciliate scales are pres- Myers (1935) referred to the slender or ta- ent on the cheeks of males, and contact or- pered caudal peduncle of the species of Cy- 1989 SMITH: HISPANIOLAN PUPFISHES 9 prinodon as a feature that distinguishes the ever, these pupfishes can be excluded from genus from Floridichthys, in which the pe- C. bondi because they lack the high circum- duncle is deep throughout its length. The peduncular scale count, tapered caudal pe- shape of the peduncle is actually variable in duncle, and lingulate interpelvic scales. Cyprinodon (closely resembling that of Flo- ridichthys in inland pupfishes such as C. al- varezi and C.fontinalis), but it is more strong- REFERENCES ly tapered in C. bondi than in any other Bond, R. M. cyprinodontine and is therefore inferred to 1935. Investigations of some Hispaniolan be autapomorphic. lakes. II. Hydrology and hydrography. The presence of a toothed dermal plate in Arch. Hydrobiol. 28: 137-161. the position of the first pharyngobranchial Burgess, G. H. bone is an unusual feature of C. bondi. A 1983. Cyprinodon bondi Myers. In D. S. Lee, similar toothplate, though much larger, has S. P. Platania, and G. H. Burgess (eds.), been reported in Floridichthys carpio in which Atlas of North American freshwater it was interpreted as an autapomorphy fishes, 1983 Supplement, p. 52. Occas. (Humphries, 1981). A toothed plate in this Pap. North Carolina Biol. Surv. 1983- 6: 67 pp. position also occurs in Garmanella pulchra Hubbs, C. L., and R. R. Miller (ANSP 140554, UMMZ 184668,196534) but 1942. Studies ofthe fishes ofthe order Cyprin- otherwise seems to be unreported among eu- odontes. XVIII. Cyprinodon laciniatus, teleosts. It may therefore be a derived feature new species, from the Bahamas. Occas. indicating close relationships. C. bondi fur- Pap. Mus. Zool. Univ. Michigan 458: ther resembles F. carpio and G. pulchra in 11 pp. lacking both an enlarged scapular scale and Humphries, J. M. an expanded cleithral process, but the lack of 1981. The evolution of a species flock in the these characters is probably primitive among genus Cyprinodon (Pisces: Cyprinodon- cyprinodontines and, if so, would be unin- tidae). Ph.D. thesis, Univ. Michigan, Ann Arbor. 233 pp. formative as an indicator of relationships. 1984. Cyprinodon verecundus, n. sp., a fifth In other cyprinodontines including out- species of pupfish from Laguna Chi- groups such as Cualac, Jordanella, and the chancanab. Copeia 1984(1): 58-68. Old World genus Aphanius, the scales around Humphries, J. M., and R. R. Miller the caudal peduncle number 16 or fewer; C. 1981. A remarkable species flock ofpupfishes, bondi and C. laciniatus have more circum- genus Cyprinodon, from Yucatan, Mex- peduncular scales (modally 20, table 3), a fea- ico. Copeia 1981(1): 52-64. ture that is therefore likely to be apomorphic. Lazara, K. J. These two species also share vertebral num- 1984. The killifish master index. A checklist bers and lateral-scale counts that are high for of oviparous cyprinodontiform fishes. American Killifish Assoc., Cincinnati. Antillean Cyprinodon (tables 2, 3), but these Miller, R. R. characters are more general among cyprino- 1948. The cyprinodont fishes of the Death dontines and, therefore, probably are not evi- Valley system ofeastern California and dence of a close relationship. Cyprinodon southwestern Nevada. Misc. Pap. Mus. bondi is readily distinguished from C. lacin- Zool. Univ. Michigan 68: 155 pp. iatus by its color pattern, rhomboid rather 1962. Taxonomic status of Cyprinodon ba- than elongate shape, lack of laciniate scales, coni, a killifish from Andros Island, Ba- and presence of a first pharyngobranchial hamas. Copeia 1962(4): 836-837. toothplate. 1981. Coevolution of deserts and pupfishes As here construed, C. bondi is restricted to (genus Cyprinodon) in the American a brackish lake on Haiti, Southwest. In R. J. Naiman and D. L. the border between Soltz (eds.), Fishes in North American where the lake is called Etang Saumatre, and deserts, pp. 39-94. New York: Wiley- the Dominican Republic, where it is known Interscience. as Lago El Fondo. Pupfishes from a nearby Minckley, W. L., and C. 0. Minckley Dominican lake, Lago Enriquillo, were re- 1986. Cyprinodon pachycephalus, a new ferred to C. bondi by Burgess (1983). How- species of pupfish (Cyprinodontidae) 10 AMERICAN MUSEUM NOVITATES NO. 2953

from the Chihuahuan Desert of north- in Haiti. Bull. Florida St. Mus., ern Mexico. Copeia 1986(1): 184-192. Biol. Sci. 33(1): 86 pp. Myers, G. S. Turner, B. J., and R. K. Liu 1935. An annotated list of the cyprinodont 1977. Extensive interspecific genetic compat- fishes of Hispaniola, with descriptions ibility in the New World killifish genus of two new species. Zoologica 10(3): Cyprinodon. Copeia 1977(2): 259-269. 301-316. Woodring, W. P., J. S. Brown, and W. S. Burbank Thorbjarnarson, J. B. 1924. Geology ofthe Republic ofHaiti. Dept. 1988. The status and ecology ofthe American Publ. Works, Port-au-Prince. 631 pp.

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