Three New Pupfish Species, Cyprinodon (Teleostei, Cyprinodontidae), from Chihuahua, México, and Arizona, USA Author(S): W

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Three New Pupfish Species, Cyprinodon (Teleostei, Cyprinodontidae), from Chihuahua, México, and Arizona, USA Author(s): W. L. Minckley, Robert Rush Miller and Steven Mark Norris Source: Copeia, Vol. 2002, No. 3 (Aug. 15, 2002), pp. 687-705 Published by: American Society of Ichthyologists and Herpetologists (ASIH) Stable URL: http://www.jstor.org/stable/1448150 . Accessed: 23/07/2014 15:57

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Three New Pupfish Species, Cyprinodon(Teleostei, Cyprinodontidae), from Chihuahua, Mexico, and Arizona, USA

W. L. MINCKLEY,ROBERT RUSH MILLER,AND STEVENMARK NORRIS

Three new species of Cyprinodon(Teleostei, Cyprinodontidae)are described,each long recognized as distinct. Cyprinodonpisteri occupies a varietyof systems and habitats in the Lago de Guzmin complex basin in northern Chihuahua,Mexico. It is distinguishedby its dusky to black dorsal fin and narrowor inconspicuousterminal bar on the caudal fin in mature males. Cyprinodonalbivelis is distributedlargely in relativelyhigh elevation sites in the upper Rio Papigochic (Rio Yaquibasin), with a single occurrence in the Rio Santa Mariabasin. It is distinguishedmost notablyby the strikingwhite dorsal and anal fins displayed by breeding males. Cyprinodonar- cuatusis a distinctive form most similar to Cyprinodonmacularius and Cyprinodon eremusand is restricted to the upper Santa Cruz River basin in southern Arizona and Northern Sonora. It is distinguished in having a sharply convex dorsal body profile form the head to dorsal fin origin and in lacking yellow or orange pigmentation on the part of nuptial males. Extensive and careless human alterationsto this system drove C. arcuatusto extinction, with the last naturalpopulation perishingin Monkey Spring (Santa Cruz County,Arizona), devoured by introduced sport fish. Where appropriate,biogeography and conservationissues are discussedwith regard to these species.

80 while the staff of Miller and n. NEARLY years ago, on Cyprinodonpisteri Minckley, sp. the Field Museum of Natural History, Carl Cachorrito de Guzmin, Guzman Pupfish L. Hubbs recognized as new two of the three Figures 1-2 pupfishes described here, designating as types eximius an abundance of preserved material from the Cyprinodon (misident.): Bean, 1898:168; Evermann and Lago de Guzmin and Rio Yaqui basins of north- Goldsborough, 1902:149; 125 ern Chihuahua, Mexico. A third species from Meek, 1904:xxxii-xxxiii, (in part); Fowler, 1916:430; de Buen, 1947:277 Ever- the Gila River drainage (Colorado River basin) (from mann and 1902); Contreras- of southeastern Arizona, and likely also in Goldsborough, 1969:297. northern Sonora, Mexico, was also known to Balderas, (misident.): Garman, 1895: Hubbs, who referred it to macularius Cyprinodon elegans Cyprinodon 1902:97-98 de Hubbs and The two senior 23-24; Meek, (in part); Buen, (e.g., Miller, 1941). 1940:31 authors have collected and studied these fishes (in part). bovinus (misident.): 1907:83- over their for Cyprinodon Regan, respective geographic ranges 84 (in Fowler, 1916:429-430. and offer their formal part); many years descriptions bovinus bovinus (misident.): de Buen, below. Cyprinodon 1947:277 (in part). Cyprinodonsp. or "sp. nov.": Koster, 1957:83 (in and Arnold, 1969:225; Anas- MATERIALS AND METHODS part); Minckley cavage, 1973 (as "Ascencion [sic)] pupfish"; det. by RRM as UMMZ 211153); Contreras et in Tables Morphometric measures (listed 1- al, 1976:table 3 (Rio del Carmen); Turner meristic counts in Table and de- 2), (listed 3) and Liu, 1977:fig. 1, no. 32 (in part); Echelle scriptions of pupfish scale morphology follow and Echelle, 1978:572; Miller and Chernoff, Miller (1943, 1948). Professor Hubbs' notes and 1980:79-80; Gehlbach, 1981:270 (Palomas); data on these pupfishes (and others) were given Miller, 1981:fig. 2 [17, 19], p. 74; Minckley to RRM in the 1950s and proved invaluable for and Brown, 1983 (reprinted 1994), table 32, their descriptions. Vertebral counts taken from p. 234 (as "Casas Grandes pupfish"); Miller, radiographs, with urostyle counted as one. CS 1986:table 1; Smith and Miller, 1986:table indicates cleared-and-stained specimens. Abbre- 13.1, p. 469 (as "Palomas pupfish"); Williams viations for depositories are from Leviton et al. et al., 1989:10 (as "whitefin pupfish/cachor- (1985). Note variable authorship of species de- rito de Palomas"); Sublette et al., 1990:351; scribed in this report. Minckley et al., 1991:table 15-1, p. 253 (as

2002 the American of and ? by Society Ichthyologists Herpetologists

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TABLE 1. SELECTEDMORPHOMETRIC FEATURES FOR Cyprinodon pisteri (UMMZ 162629, 162630) ANDCyprinodon albivelis(UMMZ 211582, 235040). Presented in 1000s of standard length, which is in millimeters. Mean is followed by 1 SD, range in parentheses; n = 11 for male columns (including holotypes), n = 10 for female columns.

Cyprindonpistei Cyprinodonalbivelis Holotype Males Females Holotype Males Females Standardlength (mm) 41.5 41.6 ? 5.1 39.2 ? 5.8 36.8 40.8 ? 6.6 38,1 ? 1.7 (33.7-49.0) (29.5-49.3) (34.4-53.1) (36.1-40.7) Predorsalfin length 598 592 ? 11.2 595 ? 13.3 587 588 + 21.4 592 ? 11.9 (577-606) (572-609) (562-627) (577-615) Prepelvicfin length 571 557 ? 12.6 555 ? 24.1 560 555 ? 16.6 566 ? 6.2 (538-576) (524-610) (529-580) (551-572) Preanal fin length 684 678 ? 11.9 673 ? 8.4 679 669 ? 16.2 684 ? 13.8 (657-696) (661-686) (641-697) (659-701) Anal origin to caudal fin base 383 391 ? 13.0 363 ? 4.4 397 398 ? 12.4 372 ? 15.6 (373-416) (358-373) (383-429) (344-401) Body depth 402 396 ? 19.4 363 ? 27.6 418 424 ? 37.2 363 ? 14.7 (366-434) (310-398) (370-508) (337-385) Body width 255 234 ? 15.0 229 ? 18.4 242 229 ? 13.2 246 ? 11.3 (204-255) (193-252) (206-245) (232-266) Head length 311 309 ? 8.8 311 ? 11.5 321 333 ? 14.7 318 ? 8.8 (294-318) (295-330) (301-356) (306-332) Caudal-peduncle length 289 285 ? 16.8 269 ? 15.6 280 293 ? 11.5 282 ? 1.30 (248-311) (242-290) (271-313) (261-301) Caudal-peduncle depth 214 206 ? 7.7 191 ? 6.0 215 216 ? 12.5 184 ? 6.1 (190-216) (183-201) (189-237) (174-193) Interorbitalwidth 118 118 ? 10.0 117 ? 6.2 125 123 ? 12.1 114 ? 4.8 (103-134) (106-128) (107-151) (106-121) Snout length 104 103 ? 5.9 101 ? 8.5 106 110 ? 9.8 106 ? 10.2 (94-113) (86-115) (92-126) (89 ? 127) Orbital length 75 81 ? 4.9 81 ? 5.9 87 87 ? 6.0 85 ? 4.6 (75-89) (75-92) (77-98) (80-93) Mouth width 106 98 ? 9.4 97.2 ? 11.4 109 103 ? 7.9 103 ? 6.0 (83-124) (75-111) (88-113) (97-116) Mandiblelength 65 63 ? 5.4 64 ? 8.7 73 64 ? 5.6 69 ? 5.0 (50-71) (47-78) (57-73) (64-78) Dorsal-finlength 258 257 ? 13.9 239 ? 9.1 304 290 ? 21.8 245 ? 12.1 (228-279) (225-249) (251-328) (229-271) Anal fin length 207 221 ? 13.2 187 ? 19.1 245 235 ? 14.7 197 ? 11.1 (202-244) (162-230) (218-256) (117-211) Caudalfin length 222 226 ? 13.1 220 ? 16.2 236 237 ? 13.8 220 ? 10.5 (204-247) (202-244) (215-271) (201-237) Pectoral fin length 214 226 ? 12.9 220 ? 13.2 239 235 ? 14.1 221 ? 12.0 (206-241) (196-234) (205-252) (202-233) Pelvic fin length 108 124 ? 9.4 109 ? 9.7 130 118 ? 6.7 109 ? 7.3 (108-140) (92-121) (108-130) (96-120)

"Guzmin pupfish"), p. 254 (as "Palomas Paratypes.-All Chihuahua, Mexico: UMMZ pupfish"); Propst and Stefferud, 1994:233- 162630, 103 ex., 15 CS, 2 dry skeletons, 19-53 234; Echelle and Dowling, 1992:200-201; mm SL, taken with the holotype; UMMZ Echelle and Echelle, 1993:276. 136126, 50 ex., 12-37 mm SL, Pozo de las De- licias, Rio del Carmen basin 3.2 km W Ahuma- Holotype.-UMMZ 162629, male, 41.5 mm SL; da, S. B. Benson,16 May 1937; UMMZ 181124, Mexico: Chihuahua, Ojo de Palomas Viejo (now 17 ex., 13-44 mm SL, spring near Lago de Guz- dry), 5.2 km S Las Palomas, 31'45'N, 107040'W; man, J. M. Legler, 2 Sept. 1950; UMMZ 162616, R. R. Miller and H. E. Winn (M50-8), 9 March 181 ex., spring-fed pond 0.9 km SE Las Palomas, 1950. R. R. Miller and H. E. Winn (M50-4), 7 March

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1950; UMMZ 182395, 305 ex., 10 CS, 10-46 mm its cleft only slightly oblique when closed. Body SL, Laguna Bustillos, R. R. Miller and family shape and proportions variable among popula- (M64-8), 20 June 1964; UMMZ 182399, 2 ex., tions (Figs. 2-3). Some populations deeper bod- 21, 24 mm SL, Rio del Carmen, R. R. Miller and ied, some with upper body contour more evenly family (M64-9), 21 June 1964; UMMZ 203017, rounded than others, some have larger fins (es- 206 ex., 11-45 mm SL, Rancho la Nariz R. R. pecially dorsal fins of males, as in Rio del Car- Miller et al. (M78-11), 27 May 1978; UMMZ men basin). 203024, 306 ex. (6 CS), 13-41 mm SL, Ojo Cal- First dorsal fin ray broad, spinelike; pectoral iente WNW Janos, R. R. Miller et al. (M78-14), fin long, ovoid, extending beyond pelvic base in 29 May 1978; UMMZ 208225, 2 ex., 35, 42 mm nuptial males; pelvic fin also long, in males of- SL, Laguna Bustillos, B. Chernoff and M. L. ten extending beyond anus; outer caudal rays Smith (BC79-15), 27 May 1978; UMMZ 208230, in males often longer than central ones, result- 7 ex., 16-20 mm SL, Rio Santa Maria at Bachi- ing in weakly trapezoidal outline. niva, B. Chernoff and M. L. Smith (BC79-16), Venter and breast fully scaled; scale margins 29 May 1979; UMMZ 208235, 1 ex., 28 mm SL, entire or irregular, not crenate; scales between Rio Santa Maria at Santa Ana de Bavicora, B. pelvic fins and anus well developed; those be- Chernoff and M. L. Smith (BC79-17), 29 May tween pelvics thick, free distally, and covering 1979; UMMZ 208241, 21 ex., Ignacio Zaragoza, inner pelvic fin rays; body scales lack conspicu- B. Chernoff and M. L. Smith (BC79-20), 30 ous surface reticulations (i.e., circuli lack erect, May 1979; UMMZ 209017, 57 ex., 13-38 mm SL, spinelike projections or lepidonts; similar to trib. Laguna Bustillos, R. R. Miller and M. L. Miller, 1943: pl. 5, fig. 2; pl. 7, fig. 1); scale radii Smith (M80-2), 23 April 1980; UMMZ 209021, numerous, body scales deep; cleithral scale en- 99 ex., 13-41 mm SL, Rio Santa Clara, trib. Rio larged, 1.5-2.0 times as wide as those following; del Carmen, R. R. Miller and M. L. Smith cleithral process extending posteriorly beyond (M80-3), 24 April 1980; UMMZ 211147, 51 ex., pectoral fin base. 18-38 mm SL, Ojo Caliente, E. Zaragoza, R. R. As with congenors, C. pisteri shows noteable and F. H. Miller (M82-81), 3 April 1982; UMMZ sexual dimorphism and dichromatism (Miller 211153, 102 ex., 16-40 mm SL, Ojo de San Juan and Fuiman, 1987; also compare males with fe- ESE Ascencion, R. R. and F. H. Miller (M82- males in Figs. 2-3). Pigmentation differences 84), 5 April 1982; UMMZ 211625, 104 ex., 13- noted below. Adult males tend to develop lon- 35 mm SL, Ojo Varelefio W Nuevas Casas Gran- ger dorsal and anal fins than females. Sexually des, R. R. and F. H. Miller, K. Bowman (M78- mature males more robust in many body mea- 45), 21 June 1978; UMMZ 212314, 45 ex., 10- sures, particularly becoming more compressed 37 mm SL, Lago de Guzmain, C. Lowe and Todd and deep-bodied. Predorsal profile slightly con- (UAZ75-70), 28 April 1975; ASU 9258, 88 ex., vex in females, nearly straight in males, some- 10-44 mm SL; UANL 15048 (ex-UMMZ times with dorsal crest. Adult characters of form 203024); UNAM 12922 (ex-UMMZ 203024). and color often develop in unusually small males (23-25 mm SL), although not to extent Diagnosis.-A species of Cyprinodon distin- as in larger ones. guished from congeners by the following combination of characters: in breeding males dorsal Coloration.-Life colors of nuptial males ob- fin dusky to black, not yellow or orange; band served at various times from early April to July. on caudal fin narrow (narrower than eye-di- Back and upper sides brilliant, deep metallic- ameter) and often diffuse or absent; first dorsal blue, paler (becoming silvery to chalky-blue) on fin ray thickened and spikelike; ocellus usually lower sides, whitish on belly. Opercles, cheeks, well developed in dorsal fin of females, no anal and branchiostegal areas suffused with yellow fin ocellus; breast fully scaled; modally 10 dorsal over blue. Sides with 7-8 dark vertical bars, and anal fin rays; 7 pelvic fin rays, 25-28 (mode sometimes coalescing just before distal part of 26) scales in lateral line, and modally 26 verte- caudal peduncle. In life, caudal peduncle distal brae. third lighter and scarcely (but often obviously) suffused with yellow (but never as bright as in Description.-Morphometric data presented in C. macularius). Dorsal fin dusky to black or Table 1; meristic data presented in Table 3. dusky near base with broad, black margin. Anal Maximum size observed 53 mm SL. General fin slate-gray, overlain by chalky- to turquoise- morphology typical of genus. Dorsal fin origin blue at base, yellow medially, with intensely positioned at midbody or slightly nearer tip of blackened distal border. Pectoral and pelvic fins snout; eye small, lower jaw small, neither robust milky- or chalky-blue to yellow or orange, with nor strongly upturned; mouth nearly terminal, rays and distal margin blackened. Caudal fin

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions TABLE2. SELECTEDMORPHOMETRIC FEATURES FOR Cyprinodon arcuatus [UMMZ 162700, 162701, n = 12 FORMALES (INCLUDING HOLOTYPE) AND 10 FORFEMALES] AND TWO rD = REGIONALCONGENORs Cyprinodon macularius (n 10, EACHCOLUMN) AND Cyprinodoneremus. Mean is followed by 1 SD, range in parentheses (data for C. eremusfrom Miller and Fuiman, 1987, UMMZ162661, 162662, n = 15, each gender, standarddeviation for these values not available).

Cyprinodonarcuatus Cyprinodonmacularius Cyprinodoneremus Holotype Males Females Males Females Males Females Standardlength (mm) 39.3 37.25 t 2.9 31.1 + 3.7 33.7 + 1.9 34.1 ? 2.8 35.5 34.9 (31.8-42) (26.7-39.3) (30.5-36.0) (29.5-37.8) (29.5-41.5) (27.9-39.0) 0 Predorsalfin length 539 564 ? 11.1 557 ? 10.6 559 + 10.5 569 + 9.7 584 583 (547-579) (546-573) (534-571) (553-582) (554-604) (552-602) Prepelvicfin length 519 528 t 8.6 530 + 12.3 550 ? 11.6 565 ? 12.6 573 579 (512-543) (509-544) (532-574) (547-591) (557-588) (545-602) Preanal fin length 628 648 + 12.6 652 + 13.0 660 + 16.4 682 ? 4.0 693 699 (627-666) (633-668) (623-683) (675-687) (676-704) (667-728) Anal origin to caudal fin base 415 420 ? 12.3 400 ? 16.2 432 ? 18.0 376 ? 15.3 386 359 C) (400-437) (375-418) (389-450) (349-400) (366-407) (340-377) Body depth 402 404 + 20.5 358 ? 22.0 420 + 18.1 399 + 8.8 441 396 0 (364-428) (329-394) (383-445) (388-414) (393-449) (369-421) t?1 width 211 207 7.6 209 10.0 221 ? 9.9 244 8.2 262 z Body + + + 263 9;t• (194-217) (192-228) (203-237) (233-258) (235-285) (239-292) Head length 313 316 + 10.3 309 ? 13.1 323 + 13.8 308 ? 8.0 336 325 ?c (294-330) (281-325) (302-354) (299-321) (322-353) (310-339) Caudal peduncle length 300 301 ? 16.7 290 + 18.7 290 ? 14.7 269 + 10.9 257 252 (267-321) (259-417) (269-315) (254-291) (231-275) (235-268) Caudal peduncle depth 206 202 ? 8.5 189 ? 11.1 201 + 6.4 179 t 3.8 205 186 (191-215) (177-211) (190-209) (174-185) (190-217) (172-203) Interorbitalwidth 130 120 ? 2.3 116 t 11.2 119 ? 4.6 112 t 3.6 123 123 (115-123) (91-134) (114-127) (107-117) (112-140) (115-129) Snout length 104 99 + 5.9 86 ? 13.6 104 ? 5.1 98 ? 4.8 100 95 (88-108) (62-104) (97-111) (91-107) (85-111) (86-103) Orbitallength 87 91 ? 3.1 97 ? 4.5 87 + 3.8 82 + 3.5 85 81 (86-94) (89-105) (81-93) (74-86) (78-94) (74-86) Mouth width 107 100 ? 7.7 105 ? 11.8 91 + 3.8 91 t 2.5 122 113 (86-108) (90-135) (82-95) (87-96) (115-131) (101-126) Mandible length 66 65 ? 3.4 57 ? 5.6 65 ? 4.5 59 t 5.3 102 101 (60-71) (47-65) (59-75) (53-68) (81-111) (82-109) Dorsal-finlength 282 288 ? 14.6 257 ? 20.4 307 t 25.4 278 + 9.1 305 252 (265-309) (208-287) (255-344) (260-288) (289-330) (227-279)

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gray washed with yellow, with narrow black border that may be weak to very nearly absent (con- M o C) 0 af) cc, Ite O -;a C\ ) C>- ~-- in males from Bustillos basin; -400 0) spicuous Laguna E C4 II C o cq r--4~-C-0 Other variants include fish from L, 00C) C> C) Fig. 2). major Rio del Carmen at Santa Clara (Fig. 2), where i caudal peduncle of males have bronze to brassy sheen, caudal fins deep-orange in advance of a black terminal border, and Arroyo Nopabechic o Bustillos) where male abdomens or- h3 I-- I- CN4 (Laguna X0 ar) X0 r--4 ange-yellow from pelvic origins onto throat and C114 I\ E C4 00 c, CN 4\ lower sides to above upper base of pectoral fin. In females, dorsal ocellus prominent in some populations, weak in others. Tending to be disrupted or lost in old (large) fish, apparently absent in very large adults (Fig. 2). Live females C C) 6 00C?~.00 olive-brown with bronze to p, C4 a<,) generally dorsally c;a C114 C4 C14 sides that often become reticulate. [" EO +1 +1 +1 +1 I yellowish 24 - -O C 0 400 r--4 Both vertical and paired fins often yellowish. 3 u Most fish from de Las Palomas (d Ojo Viejo (e.g., E UMMZ mm with vertical bars c 162630; 28-42 SL) (as many as 15, Fig. 1) disrupted ventrally, while c h - some with at least one bar continuous across o - C4 r0-4 4 belly. In life, females from Ojo Caliente (UMMZ -;a +1 +1 I E +d +101 1> 211147) with yellow streak along entire dorsal C, t 0c, C~4 00 00 e\ Ci( c C~c - midline. of both sexes similar in 0) t0 C> a'XO Cl - Young pigmentation to females, as usual in pupfishes, with -8 z lateral bars of generally uniform width and usu- Q z ally prominent dorsal fin ocellus. 0 X, C~ a<, X0 00 0) ?c C I C4 o Comparisons.-Cyprinodon pisteri belongs to a H c~J +1 +14+1 +1 E clade of western pupfishes that also includes the Z4o o') 0 -4 nC 0 " C'qC'q - - C" Death Valley species (C. diabolis, C. nevadensis, C. radiosus, C. salinus), Colorado River basin forms [C. macularius, C. eremus,and presumably C>O~~c C> ~ - C. arcuatus (see below) ], and other Guzman and Rio Yaqui basin forms (C. albivelis, C.fontinalis). r-00 XC4I- - 00ow t the southern rnO Relationships among (Guzman/ +1 4 +1I E X0+1I1 +11I-VC4 0 Yaqui basins) forms are yet unresolved (phylo- o cc, - - t- ccg c characterized molecular O e n ~ "- genetics using data, (MC>C O Echelle and Dowling, 1992; Echelle and Echelle, 1993), and additional undescribed species exist in the Guzman Basin (S. Contreras- Balderas, pers. comm.). All three described O Guzman Basin show a O C14 C14 r- pupfishes frequent (ap- 3: proximately 40%) lack of mandibular pores (Ta- ble 3; Smith and Miller, 1980); a trait not observed in any of the three Colorado River forms.

and is re- -\ 0 01 Distribution habitat.--Cyprinodonpisteri stricted to the Lago de Guzmrn basin of northern Chihuahua, Mexico, and possibly southwestern New Mexico, USA and its former con- nective, Laguna Bustillos, Chihuahua (Fig. 3; Smith and Miller, 1986:fig. 13.1, nos. 8-12). In the Pleistocene and earlier, this vast endorheic region (pluvial Lago de Guzmin system of Smith and Miller, 1986:fig. 13.3) supported a

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions TABLE3. FREQUENCYDISTRIBUTIONS OF SELECTEDMERISTIC FEATURES FOR Cyprinodon pisteri, Cyprinodonalbivelis, Cyprinodonarcuatus, COMPAREDWITH Cyprinodon macularius (UMMZ 162680), AND Cyprinodoneremus (FROMMILLER AND FUIMAN,1987); VALUESFOR HOLOTYPE (WHERE KNOWN) INDICATED BY AN ASTERISK(*).

Dorsalfin rays Anal fin rays Pectoralfin rays 7 8 9 10 11 n Mean + 1 SD 6 7 8 9 10 11 n Mean ? 1 SD 12 13 14 15 16 17 n Mean - 1 SD C. pisteri - 1 71* 146 15 233 9.8 ? 0.6 - - - 21 177* 31 229 10.0 ? 0.5 - - 23 132* 66 10 231 15.3 + 0.7 - - C. albivelis 5 12 15* 6 - 38 8.6 ? 0.9 1 - 7 25 7* 40 8.9 + 0.8 2 22 33* 2 - 59 14.6 + 0.6 C. arcuatus - - 5 15* - 20 9.8 ? 0.4 - - - 2 17* 1 20 9.9 ? 0.4 1 - 11* 7 1 - 20 14.3 + 0.8 C. macularius - - 8 11 - 19 9.6 ? 0.5 - - - 1 17 2 20 10.1 0.4 - - 5 14 1 - 20 14.8 ? 0.5 C. eremus - - 4 20 6 30 10.1 ? 0.6 - - - - 25 5 30 10.2 + 0.4 - - - 17 37 6 60 15.8 + 0.6 C, 0

z p

TABLE 3. CONTINUED

Pectoralfin rays Lateralline scales Scalesfrom dorsalfin origin to pelvicfin insertion 4 5 6 7 8 n Mean - 1 SD 24 25 26 27 28 29 n Mean + 1 SD 9 10 11 12 13 n Mean 1 SD _ C. pisteri - - 2 17* 1 20 7.0 ? 0.4 - 4 11 4* 1 - 20 26.1 ? 0.8 5 10 6* - - 21 10.0 0.7 - - C. albivelis 1 2 18 39* 60 6.6 + 0.6 - - 7 25* 5 2 37 27.1 ? 0.7 9 17* 14 1 41 11.2 + 0.8 - C. arcuatus - 1* 4 16 - 21 6.7 -+ 0.6 1* 7 8 5 - - 21 25.9 ? 0.8 16 5 - - 21 10.2 0.4 C. macularius - 12 8 - - 20 6.5 ? 0.5 2 13 4 - 1 - 20 26.3 ? 0.9 - 6 14 - - 20 10.7 ? 0.5 C. eremus - 1 - 58 1 60 7.0 ? 0.3 - 18 12 - - - 30 25.4 ? 0.5 no data

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Circumferentialscales Predorsalscales Mean Mean 24 25 26 27 28 29 30 31 32 33 34 35 36 n I1SD 14 15 16 17 18 19 20 21 22 23 24 n 1SD C.pisteri 2 4 1 6 2 4* 1 ------20 27.0 1.8 - - - 5 5 3 4 - - - - 17 18.4 + 1.2 - - C. albivelis 1 8 10 7* 8 3 1 -- 38 28.7 ? 1.4 1 1 1 2 2 6 4 1 1 19 20.4 + 2.0 C arcuatus 1 5* 4 5 2 3 - - - - 20 26.6 ? 1.5 - - - 1 3 5* 1 1 - - 11 18.6 0.8 C. macularius 1 - 9 3 4 1 2 - - - 20 29.8 ? 1.4 3 8 6 2 ------19 15.4? 0.9 -•- - - =z C. eremus -- - 3 2 12 2 10 1 30 32.6? 1.5 no data

trj

r-? TABLE 3. CONTINUED

Mandibularpores Preorbitalpores Preopercularpores Mean Mean Mean 0 1 2 3 4 5 n 1SD 2 3 4 5 n +1SD 5 6 7 8 9 10 n l1SD ------C. pisteri 84 2 215* 1 1 303 1.4 + 0.9 1 19*" 20 4.0 0.2 21 180* 2 203 6.9 0.4 - - - C. albivelis 99 3 239* 1 1 343 1.4 ? 0.9 4 55* 1 60 4.0 ? 0.3 28 207* 4 2 2 243 6.9 ? 0.5 ------C. arcuatus - 20* 20 2.0 1 2 17* 20 3.8 + 0.5 1 17 2* 20 6.0 0.4 - - - - - Vo C. macularius 1 1 17 1 20 2.0 + 0.9 2 16 2 20 4.0 + 0.5 - 6 13 1 20 7.8 + 0.6 C. eremus 1 - 59 - - - 60 2.0 ? 0.3 1 10 49 - 60 3.8 ? 0.4 - 1 54 5 - - 60 7.1 0.3

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TABLE 3. CONTINUED

Totalvertebrae 25 26 27 28 29 n Meant 1 SD

C. pisteri 8 36 16 - - 60 26.1 ? 0.6 C. albivelis - - 6 42 12 60 28.1 ? 0.5 C. arcuatus 8 20 5 - - 30 25.9 ? 0.7 C. macularius 1 25 24 - - 50 26.5 ? 0.6 C. eremus 1 25 4 - - 30 26.1 ? 0.4

complex of lakes, streams, and springs (Miller, range are at lower elevations, where springs and 1981:fig. 3, nos. 5-7; fig. 4, no. 3). Former trib- other surface waters are disappearing with hu- utaries remain clearly identified within a now- man development (see below). The species disrupted array of desiccated intermontane ba- once occurred in the United States in artificial sins by distributions of at least eleven native fish- ponds near Columbus, Luna County, New Mex- es in remnant aquatic habitats (Smith and Mill- ico (MSB 949, 38 specimens collected in 1951), er, 1986; Propst and Stefferud, 1994). where it was apparently introduced from Mexi- Although certainly less abundant now than in can sources for mosquito suppression (pers. the recent past (Fig. 3), C. pisteri still enjoys a comm., W. J. Koster to RRM, 1978). This pop- wide geographic range including much of the ulation is no longer extant, and Prof. Koster fur- Lago de Guzmain basin. Greatest reductions in ther reported that springs supporting the spe-

Fig. 1. Cyprindonpisteri, holotype (above), UMMZ 162629, 42 mm SL, male; paratype (below), UMMZ 162630, 37 mm female, Mexico: Chihuahua,Palomas Spring (Ojo de PalomasViejo; now dry).

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1130 1090 1050 1010

SM I? I

29'

250

Fig. 3. Distributionsof Cyprinodonpisteri (circles) and Cyprinodonalbivelis (triangles), n. spp., in Chihua- hua, Mexico. Some symbolscover multiple, closelyad- jacent sites; type localities = open figures. B = La- guna Bustillos;C = Rio del Carmen;CG = Rio Casas Grandes;P = Rio Papig6chic;SM = Rio Santa Maria; Y = Rio Yaqui.

dered upslope by sparse Bermuda grass (Cyno- don dactylon) and a few shrubby mesquite (Pro- sopis glandulosa) and acacia (Acacia greggi). Iso- lated stands of cattail (Typha angustifolia) were in the pool. A thick, spongy mat of sedges oc- cupied the shallow, natural outflow that passed east-southeast from the spring, although most outflow was through a newly dug canal leading eastward to agricultural fields. The pupfish was abundant at the type locality Fig. 2. Cyprinodonpisteri, from top to bottom (para- in situations where it occurred in the ab- mm and fe- largely types): UMMZ 209017, male (35.8 SL) sence of other fishes and in male mm Mexico: along pool margins (34.8.9 SL), Chihuahua, Laguna outflow channels and It was rare to Bustillos;UMMZ 209021, male (37.3 mm SL) and fe- cienegas. absent in areas > 1.0 that male (40.9 mm SL), Mexico:Rio Santa Claraat Santa open, deeper (to m) Clara (Rio del Carmen basin). were coinhabited mostly by Pimephalespromelas (native) and Cyprinella formosa (native), and Ameiurus melas (nonnative). Gambusia affinis cies in Mexico immediately south of Pancho Vil- (nonnative) was not yet present but had be- la State Park, New Mexico dried in the 1940s. come common by 1972, when the pupfish still The species is catholic in habitat, occupying remained abundant. The system at the type lo- springs, cienegas, shorelines and cutoff chan- cality was dried by about 1975, presumably be- nels of rivers and creeks, even colonizing cause of groundwater mining in both Mexico ephemeral canals and ditches along roadsides and adjacent USA. (RRM and WLM, field obs.). When first visited by RRM in 1950, the type locality was a partially Status.-The Lago de Guzmain complex of impounded water body perhaps 1.5 ha in sur- northern Chihuahua, including in the past wa- face area, with spring inflows along its north ters of pluvial Lago de Palomas, the now-inde- and west sides and old, shallow, cienega-lined pendent Laguna Bustillos basin, and lesser lakes natural channels draining southward to appar- (Fig. 3), is composed of four major subbasins ently end in a desiccated salina or playa of un- west-to-east: (1) Rio Casas Grandes, originating known extent. Conditions were similar in 1968 in Madrean pine-oak forests along the Sierra (WLM, pers. obs.), but water was muddied by Madre Occidental crest, with a historic terminus livestock and humans, and surroundings were in Lago de Guzmain; (2) Rio Santa Maria, col- severely denuded by grazing cattle, goats, and lecting water from the Sierra Madre and Mesa horses. Shorelines that were sufficiently boggy del Norte west of Ciudad Chihuahua and end- to exclude livestock were vegetated by dense ing in Lago de Santa Maria; (3) Rio del Carmen stands of sedges (Scirpus olnyi, Scirpus american- (also called Rio Santa Clara), beginning just us), with an undergrowth of Eleocharissp. bor- east of the Sierra crest and terminating in Lago

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions 696 COPEIA, 2002, NO. 3 de los Patos north of Villa Ahumada (see Brand, Rio Casas Grandes but likely ended most re- 1937: pl. 8c); and (4) Laguna Bustillos south of cently in Bols6n de los Muertos, another basin the Rio del Carmen basin supporting Cyprinodonfontinalis and Cyprinella In keenly perceptive observations in 1929- bocagrande (Smith and Miller, 1980; Chernoff 1931 and 1935-1936, Brand (1937:72-74) re- and Miller, 1982; Mayden and Hillis, 1990). The corded the onset of major environmental deg- Mimbres River evidently united with other radation in this region. He noted that none of drainages of the Lago de Guzmin complex in its three rivers ". .. has flowed constantly the recent past only during highest stages of throughout a year and continuously from pluvial Lake Palomas. source to terminal lake since 1924." During his The direct or indirect human role in chang- period of observation, none of the streams was ing the hydrography and ecology of aquatic dammed, although several diversions took water habitats in the Chihuahuan Desert may be sum- from them. "The increasing use of spring and marized as follows. First, settlement and agri- river water for irrigation in the haciendas and cultural development led to channel incision colonias of the region has contributed markedly (arroyo cutting) through sills protecting alluvial to the lessened flow of the rivers in their lower plains above them, with resulting reductions in courses. Many of the abundant springs that fed base level of each succeeding basin. Although this system -80 years ago have failed" (Brand, perhaps in part attributable to climate change 1937:21). (Hastings and Turner, 1966), this regional event In the recent past, contributions of water flow (Cooke and Reeves, 1976) must have com- from New Mexico by Arroyo de Palomas and menced shortly after 1640 as a likely result of Mimbres River (see U.S. Geol. Surv. Map, El watershed damage from extensive timbering, Paso, 1:250,000, NH 13-1, AMS Series V502, agriculture, and grazing by innumerable herds to had minor influences. 1953) seems have only of domestic livestock originating with Spanish, Arroyo de Palomas crosses from the USA into Mexican, and Mormon settlers. Since the 1920s, Mexico near Columbus, New Mexico and Palo- several streams, the lower rios Casas Grandes, Chihuahua as subsurface flow that mas, may Santa Maria, and del Carmen, and arroyos near have fed springs near Palomas. One major Ascenci6n, have incised 6 m or more below was near the International spring Boundary their original floodplains. Second, building of (Gehlbach, 1981) and another (type locality for diversions, dams, and other structures to divert C. was about 5.0 km south of the pisteri) present water from natural channels for agricultural village of Palomas Vieja. A small barrial lake lo- and domestic use dried some reaches and called de los Patos" or de cally "Lago "Lago changed patterns of flow in others. Third, Palomas" to be confused with a (not larger, plu- groundwater mining (water extraction in excess vial basin further south in bor- Chihuahua), of natural recharge) with the advent of electric dered inflow channels and on the by cienegas pumps contributed to drying of springs and north and west and low sand dunes on the smaller channels, reducing reliability of base- northeast, was fed the lower directly by (larger) flow in essentially all systems. Last, the transfer floods, overflow from spring(s). During major and naturalization of nonnative fishes (e.g., ic- this depression may have reached the Rio Casas talurids, centrarchids) or interdrainage translo- Grandes, only about 15 km further south. These cations of regionally indigenous forms (e.g., were visited in 1892 Mearns (1907: lagunas by Gambusia forces new and sometimes "Lake Palomas in the Mimbres affinis, etc.) 10; Valley"), fatal and on the who described "Lakes Guzman and Palomas" as predation competition indigenous fauna. "considerable sheets of shallow, alkaline water. The sand wastes surrounding them are covered is a distinct to honor our with a white deposit intolerable to the eye." Etymology.-It pleasure mutual California Edwin Although Mearns (1907: 28) noted surface close, friend, biologist For al- flow of the Mimbres River crossing the Inter- Philip Pister, with the patronym pisteri. Phil Pister has national Boundary in 1892, "flowing south into most four decades, unerringly the Palomas Lakes, toward Lake Guzman in and effectively performed the daunting task of Chihuahua," today's extreme freshets rarely preserving the integrity of natural aquatic hab- reach the Boundary. Only a faint channel can itats and biotas in North American deserts, be traced toward Lago de Tiuldio about 30 km along the way teaching others to do the same. further south of the village of Arena, Chihua- His infectious and tireless persistence, enthusi- hua, approximately 22 km east of Palomas Vieja. asm, optimistic outlook, and unique capability Careful study of maps suggests surface water to redirect conflicting views toward common from the Mimbres River seldom reached the goals have led to significant and enviable suc-

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions MINCKLEYET AL.-NEW PUPFISHES 697 cesses in equating science and a strong environ- Description.-Morphometric data presented in mental ethic with political reality. Table 1, meristic data in Table 3. Maximum observed size 54 mm SL. Body shape and form typical of pupfishes, compressed and deep albivelis and n. Cyprinodon Minckley Miller, sp. (such more pronounced in males); both great- Cachorrito Aletas Whitefin Blancas, Pupfish est body depth and dorsal fin origin are at mid- 4 Figure body; mouth is almost terminal; lower jaw not overly robust; predorsal profile nearly straight or Cyprinodonsp. Cyprinodonsp. nov.-Minckley in males, tending toward convex in females. and Hendrickson et Arnold, 1969:255; al., First dorsal fin ray not sharply distinct from no. 72- 1981:78-79; Miller, 1981:fig. 2, 18, pp. others; pectoral fin ovoid, reaching posteriorly 73 "whitefin and (as pupfish"); Minckley to vertical from pelvic fin midpoint in males; 234 Brown, 1983 (republ. 1994):table 32, p. pelvic fins also long in males, surpassing anus "whitefin et 1991: (as pupfish"); Minckley al., and sometimes anal fin origin. Outer, branched "whitefin aleta 254 (as pupfish/cachorrito caudal rays of males distinctly longer on edges Rinne and blanco"); Minckley, 1991:25-26, than in center, resulting in trapezoidal shape. 44 (as "whitefin pupfish/cachorrito aleta Scale margins entire; breast and venter fully blanco"); Echelle and Dowling, 1992:fig. 1); scaled, with scales well developed between pel- Echelle and Echelle, 1993:fig. 1. vic fins, their posterior margins free and covering inner pelvic rays; body scales lack conspic- Holotype.--UMMZ 235040, adult male 36.8 mm uous surface reticulations. Cleithral scale greatly SL, Mexico: Chihuahua, Rio Papig6chic at Ye- enlarged, 2-4 times as wide as those following; pomera, 28032'W, 107029'N, 2048 m elevation; cleithral process also expanded, extending be- R. R. Miller, F. H. Miller, and R. L. Minckley, 19 yond pectoral fin base. June 1978. Coloration.-Nuptial males with white, third or more of dorsal and anal fins. of males Chihuahua, Mexico: UMMZ Body Paratypes.-All seined from territories 211597, 225 ex., 3-46 mm SL, collected with the directly breeding chalky- blue, darker to irridescent on head and holotype; UMMZ 211628, 237 ex., 8-43 mm SL, sky-blue dorsum, to white below. Faint vertical outlet of Ojo de Arrey (Array; Rio Santa Maria grading bars (3-4) extend from darker basin) 4.8 km SSE Galeana, 30'02'N, 107032'W, may ventrally dorsal to lower sides. Caudal fin, 22 June 1978, R. R. Miller and F. H. Miller pigmentation distinct, black, terminal band, clear (M78-46); ASU 806, 76 ex., 15-51 mm SL, out- excepting or only lightly dusted with melanophores on let of Ojo de Galeana, SE Galeana, 28 May 1964, rays and interradial membranes. Pelvic fins W. L. Minckley et al.. (WLM 934-937); ASU white, as on ex., 10-23 mm SL, Rio at adjacent belly, may develop pearly- 9524,16 Papig6chic white cast as on anal fin. Pectoral fins clear to Rancho de San Pedro near Mifiaca, 28024'N, dusky, often with diffusely darkened, distal mar- 107026'W, 2100 m elevation, 16-17 June 1978, In life, a few males with faint wash of light- D. A. Hendrickson et al. (DAH 76-78); ASU gins. yellow on pelvic and anal fin bases, which quick- 9938, 121 ex., 15-43 mm SL, El Ojo de Yep6m- preservation. era, 29003'N 107051'W, 1900 m elevation, 18 ly disappears upon Dorsal fin ocellus of females usually promi- June 1978, D. A. Hendrickson et al.; UANL nent. Speckled body pattern sometimes orga- 15049, 30 ex. (ex-UMMZ 211597); UNAM nized into diffuse, broken, vertical bars, espe- 12923, 30 ex. (ex-UMMZ 211597). cially on posterior third, which may sometimes consolidate into broken lateral band. Many fe- Diagnosis.--A Cyprinodondistinguished from its males with an evident unpigmented area be- congeners by the combination of a strikingly tween last vertical bar on body and diffuse dark white, one-third or more of the dorsal and anal bar on caudal fin base, separated medially by fin in nuptial males, when viewed in clear, shal- diffuse lateral band. Other than dorsal ocellus low water as a brilliant, milky- to pearly-white and some pigmentation on basal rays of caudal flag with a proximal, blackened blotch on its and dorsal fins, fins of females clear or diffusely posterior margin; terminal caudal bar strongly darkened by scattered melanophores, mostly as- developed in adult males; first dorsal fin ray un- sociated with rays. Juvenile pigments resemble modified, narrower than or only as broad as the those of adult females. second; breast fully scaled; modally 9 dorsal and anal fin rays; 7 pelvic fin rays, 26-29 (mode 27) Comparisons.--See discussion above for C. pisteri. scales in lateral line; modally 28 total vertebrae. Cyprinodon alvivelis is unique among taxa ex-

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Fig. 4. Cyprinodonalbivelis, from top to bottom (holotype and paratypes):UMMZ 235040, male (36.8 mm SL, holotype) and UMMZ211582, female (40.3 mm SL), Mexico:Chihuahua, Rio Papig6chic (Rio Yaquibasin)

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions MINCKLEYET AL.-NEW PUPFISHES 699 amined in having reduced dorsal and anal fin adjacent Rio del Carmen) must have been sym- ray counts, and an elevated vertebral count (Ta- patric in the past. We anticipate that C. albivelis ble 3). and C. pisteri are in contact near Ojo de Arrey and perhaps elsewhere in the Rio Santa Maria Distribution and habitat.-Cyprinodon albivelis is system. Clearly, the assemblage of pupfishes in distributed mostly in the upper Rio Papigochic the Guzmain basin must as yet be considered subbasin of the Pacific slope Rio Yaqui drainage poorly understood (also S. Contreras-Balderas, (Fig. 3), where it is exceptional among pupfish- work in progress). es in its consistent occurrence at relatively high elevations (1800-2300 m) (Hendrickson et al., Status.-In 1978, Hendrickson et al. (.1981) 1981:fig. 19; Blasius, 1996). It also is known found C. albivelis widespread and locally abun- from a single, low-elevation (1430 m) spring dant in the Rio Papig6chic system. Although we complex, Ojo de Arrey, in the Rio Santa Maria expect local stocks to have disappeared because drainage (interior Lago de Guzman Basin). of increased human activities, we have no rea- This latter occurrence is enigmatic, and its na- son to believe the species is as yet in jeopardy. tivity has been supsect. In 1972, a resident hes- The population at Ojo de Arrey was intact in itantly reported to WLM that he thought the 1999 when visited by P. J. Unmack (pers. pupfish was translocated there from springs comm.). near Yepomera, Chihuahua (an unquestioned native population) to control mosquitoes. The Etymology.-The name albivelis, suggested by C. fact that it was known only from one site in the L. Hubbs, is from the Latin albus (white), and Santa Maria basin, essentially surrounded by C. velum (a sail), alluding to pigmentation of the pisteri (Fig. 3), added credence to nonnativity, dorsal and anal fins in nuptial males. as did mtDNA and allozyme data. With regard to these, the Rio Santa Maria populations are Cyprinodonarcuatus Minckley and Miller, n. sp. identical to another of C. al- essentially sample Santa Cruz (Monkey Spring) Pupfish bivelis from the Rio (Echelle assayed Papig6chic Figure 5 and Dowling, 1992; Echelle and Echelle, 1993). It was also determined, however, that mtDNA Holotype.--UMMZ 162700, adult male, 39.3 mm of C. albivelis from both Ojo de Arrey and Rio SL, USA, Arizona: Santa Cruz County, pond fed Papig6chic were essentially the same as C. pisteri by Monkey Spring, trib. to Santa Cruz River (Echelle and Dowling, 1992), results tentatively (Gila River basin), 12 km north-northeast Pata- attributed by Echelle and Echelle (1993) to re- gonia, 27 April 1950, R. R. Miller et al. (M50- placement of C. pisteri mtDNA through intro- 60). gressive hybridization from C. albivelis. Such an event surely cannot be attributed to a modern Paratypes.--All Arizona, USA: UMMZ 162701, 41 fish transfer, since replacement of such magni- ex., 27-42 mm SL, same locality, date, and col- tude from one or a few introduced stocks of a lectors as holotype; AMNH 232395 (ex-UMMZ species through myriad, disjunct populations of 125050), 20 ex.; ASU 729, 82 ex. 24-45 mm SL; another must require far more time. Further, ASU 600, 96 ex., 15-46 mm SL; ASU 4672, 7 although similar in most meristic and morpho- ex., 35-44 mm SL, same locality as holotype; logic features, considerable difference exists in CAS 18561, 2 ex., Sonoita Cr. at Patagonia, San- number of gill rakers in specimens of C. albivelis ta Cruz County, Sept. 1927, J. Mailliard; CAS from Ojo de Arrey (mean 17.2, n = 20) versus 214579 (ex-UMMZ 125050), 20 ex.; UMMZ Rio Yaqui (mean 14.6, n = 80), a degree of dif- 125050, 223 ex., 14-46 mm SL; USNM 130002, ferentiation unlikely to be attained in a short 54 ex.; USNM 21315, 2 ex., "Yuscon" (= Tuc- time unless resulting from a severe bottleneck son, Pima County), undated, 1840s?, A. Schott; (not indicated by molecular data). USNM 44096, 1 ad., "drainage of small lake or To our knowledge, C. albivelis and C. pisteri pond, Santa Cruz River," in Tucson, Pima have not yet been collected in sympatry, but County, 1891, P. L. Jouy; USNM 45441, 2 ex., based on hydrographic history, Miller (1981:46) "Tucson, Santa Cruz River," Pima County, coll. suggested that they and C. fontinalis (from the 1893, H. Brown.

at Yepomera;UMMZ 211628, male (37.9 mm SL), and female (39.3 mm SL), Mexico: Chihuahua, Ojo del Arrey (Array),Rio Santa Mariadrainage (Lago de Guzmin basin).

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Fig. 5. Cyprinodonarcuatus (paratypes), UMMZ 125050, 38.0 mm SL, male (above);and 34.0 mm SL, female (below), USA:Arizona, Santa Cruz County,pond fed by MonkeySpring.

Diagnosis.-A species of Cyprinodon distin- Coloration.-Notes by C. L. Hubbs and ourselves guished from congeners by the following com- (WLM and RRM) concur in life colors of nup- bination of characters: a distinctive, dorsal-body tial males. Dorsal pigmentation olivaceous to surface, which is highly convex before the dor- black, with scintillating surface coating of blue- sal fin but changing abruptly at the dorsal ori- green. Anal fin tended toward bluish-violet, be- gin into a deep, postdorsal concavity most de- coming metallic blue in submarginal band. veloped in breeding males; absence in nuptial Hubbs noted both dorsal and anal fins with males of distinctive yellow or orange pigment black margins and brilliant blue overcast; we on either the caudal fin or peduncle; weak de- also noted the former but observed only a ten- velopment of lepidodonts, and modally six dency toward blue, sometimes chalky-blue, on preopercular pores. fins. Most males with greenish gilt on sides, but otherwise brilliant, robin-egg blue laterally and Description.-Morphometric data presented in anterodorsally, silvery below. No yellow or or- Table 2, meristic data in Table 3. Maximum ob- ange pigmentation on caudal fin or caudal pe- served size 46 mm SL. Predorsal profile of ma- duncle, but bronze reflections on caudal fins of ture animals with distinctive convex shape, oth- a few individuals inspected in full sunlight. Cau- erwise, in general morphology, a typical mem- dal fin brilliant silver-blue on base, olive on rays ber of Cyprinodon.First dorsal fin ray much like and chalky blue-gray on membranes. Pectoral the others; pectoral fin oval, extending to at fins clear or weakly olivaceous, pelvic fins whit- least pelvic fin midlength; pelvic fin reaching to ened with light-blue cast. WLM noted adult anus; caudal fin square. Venter and breast fully males in winter (31 January) with opercles and scaled; body scales with weak surface reticula- cheeks white with apparent wash of blue, bases tions or spines. Cleithral scale enlarged 1.5-2.0 of pectoral and caudal fins with light yellow cast times as wide as those following. Unfortunately, no one recorded life colors of

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions MINCKLEYET AL.-NEW PUPFISHES 701 females in detail, except in winter dorsal ocellus c - f California Ilk Arizona was jet-black, delimited behind and sometimes , surrounded by distinct, clear area; dorsal fin was otherwise darkened. Anal, caudal, pectoral, and pelvic fins clear. Lower two-thirds of sides / G strongly marked with continuous or broken vertical bars, more complete on posterior half of Baja california the body. Tendency for bar-forming pigmenta- c SP 7 o 113 So r/ tion to widen into interrupted lateral band, then coalesce dorsally contributing to diffusely reticulate dorsum. Juveniles resembled females. k Sonora )170 1131 ( Comparisons.-Three pupfishes are native to southern Colorado River basin and closely as- Fig. 6. Distributionsof Cyprinodonarcuatus, n. sp. sociated Rio Sonoyta drainage. Cyprinodonma- (triangles), Cyprinodonmacularius (circles), and Cypri- nodon eremus in Colorado River and Rio cularius was formerly widespread in the Gila Riv- (squares) and Califor- er basin and lower Colorado River (it now per- Sonoyta drainages,Arizona, Sonora, Baja sists in artificial or isolated habi- nia (modified from Miller and Fuiman, 1987: Fig. 2 largely highly and et Some cover with C. eremus C. Echelle al., 2000: Fig. 1). symbols tats), (formerly macularius sites. localities are de- see Echelle et al. confined to multiple closely adjacent Type eremus; 2000) Qui- noted C = Colorado River;G = Gila and the Rio by open figures. tobaquito Springs adjacent Sonoyta River;M = MonkeySpring; Q = S = Rio Sonoyta;SC and C. arcuatus to the Santa Cruz River system. = Santa Cruz River;SP = San Pedro River. These three species are similar enough that they were long confounded under C. macularius. Cyprinodonmacularius and C. eremusare closely related, possibly sister taxa (Echelle et al., tirely across deeply alluviated intermontane ba- 2000). The phylogenetic position of C. arcuatus sins with high floodplain infiltration. For ex- is unknown, but biogeographic considerations ample, in a flood on 12 September 1965, a 9.7 suggest that its affinities lie with C. macularius mS X sec-1 peak was reduced 44% to 5.4 m3 X and C. eremus(or a common progenitor), likely sec-1 on the 13th after flowing about 80 km, having differentiated within its special and iso- and 88% to only 1.2 mS X sec-1 by the 14th 70 lated habitat (see below), once separated from km further downstream (Conde de la Torre, other populations. 1970). More than 1.0% of the water was there- fore lost for each 2.0 km of channel, mostly to Distribution and habitat.-Through most of the infiltration. An additional 40% of the volume 1900s, this species was thought to be restricted was estimated to be lost (reducing the peak to to the immediate vicinity of Monkey Spring. A 0.7 m3 X sec-1, approximately 7.0% of the orig- few records of native Cyprinodonin the Santa inal) over the remaining 46 km of channel lead- Cruz River basin outside this area [two speci- ing into Tucson (Burkham, 1970). mens (CAS 18561) from Sonoita Creek in the Obviously, such high infiltration leads also to town of Patagonia in 1927, 12 km downstream dramatic, up-to-downstream reduction in com- from Monkey Spring, and five others caught in petency of the stream to transport sediment, so and near Tucson in the 1800s (USNM 21315, stream-carried material is deposited in and 44096, 45441) ] were consistently identified as C. along lower reaches. This was amplified region- macularius (Miller, 1961; Minckley, 1969, 1973). ally, through Holocene and perhaps before, by We reidentify all of them as C. arcuatus. Original progressively increasing aridity, resulting in ag- distribution of the species was thus throughout gradation of "a broad, flat alluvial plain with the upper Santa Cruz River basin, Arizona-So- intermittent channels" termed the "Santa Cruz nora (Fig. 6). Flats" (Woods et al., 1999). Flood flows spread Human alteration of the Santa Cruz River sys- over this deltalike area in a broad pattern of tem has been extensive, based both on historic anastomosing distributaries. Except in the great- records (Hendrickson and Minckley, 1985) and est floods that must have connected with the evidence from prehistory (Haynes and Huckell, mainstem Gila River, infiltration, ponding, and 1986). Even in the natural state, the stream was evapotranspiration intercepted surface flows, at most a flood tributary to the Gila River. It has impeding fish dispersal. This may well explain a relatively small watershed (approximately the vicariant event leading to pupfish speciation 22,000 km2 above the lower-most gauging sta- and help explain the historic absence of other tion at Laveen, Arizona), and flows almost en- Gila River stream fishes (Gila robusta, Meda ful-

This content downloaded from 128.123.44.23 on Wed, 23 Jul 2014 15:57:21 PM All use subject to JSTOR Terms and Conditions 702 COPEIA, 2002, NO. 3 gida, Tiaroga cobitis)from the Santa Cruz system tails, history, and broader context, see Minckley (Minckley, 1973). et al., 1991:261-264). Both were ultimately lost Sometime around 1890, human activity in in 1971 when nonnative largemouth bass (Mi- both the upper and lower Santa Cruz basin be- cropterussalmoides) became abundant and de- gan to influence surface discharges even more. voured them. The downstream part of the sys- Water depletion was caused by increasing arid- tem today supports the bass, nonnative bluegill ity, groundwater pumping for irrigation, and (Lepomis macrochirus),and a few native topmin- flow diversion for domestic use. By the early now, which also survives in the headspring, too 1900s (Woods et al., 1999), even the uppermost small to support the predatory exotics and re- reaches had become intermittent, dry in sum- maining isolated by a precipitous outflow. mer and with occasional flows at other times of year associated only with heavy precipitation. Status.-This species is described almost 150 Channel incision ("arroyo cutting"; summa- years following its discovery and more than rized by Cooke and Reeves, 1976 and Hendrick- three decades after the last known population son and Minckley, 1985) lowered water tables in was destroyed. Cyprinodonarcuatus was first col- the upper system, draining floodplain backwa- lected (and misidentified as C. variegatus Lace- ters and cienegas. Channelization downstream pede) from the Santa Cruz River in "Yucson" on the Santa Cruz Flats consolidated flows for (= Tucson, Pima Co., Arizona), almost certainly a time in that area, until groundwater pumping in the late 1840s, during the U.S. and Mexican lowered water tables so infiltration again ab- Boundary Survey (Emory, 1857). Three addi- sorbed any overland discharges that might oc- tional specimens (originally identified as C. ma- cur. cularius) were preserved from that same area in Endemic Santa Cruz basin fishes persisted in the later 1800s. F. W. Chamberlain, a biologist only a single refugium. This special habitat, for the U.S. Bureau of Fisheries (Jennings, Monkey Spring (Santa Cruz County, Arizona), 1987; Minckley, 1999a), collected 54 specimens is perched on a terrace about 10 m above a in April 1904 from a pond fed by Monkey small, ephemeral arroyo tributary to Sonoita Spring, where the species persisted until the late Creek, in turn tributary to Santa Cruz River. 1960s or early 1970s. Carbonate-rich water from the spring originally The fate of habitats in the Santa Cruz River precipitated an extensive shield of tufa about system are intimately interlaced with that of C. 20.5 km downstream, protecting the arroyo wall arcuatus (as noted above). This species is extir- from eroding and ultimately creating a steep, pated in the wild, and unless an unknown rem- vertical to overhanging waterfall. Upstream tufa nant is in culture by a hobbyist, which is con- deposition, about 100 m below the spring ceivable, it is extinct. Unfortunately (or fortu- source, accumulated further to form a natural nately in a case such as this, if such proves true), dam impounding a marshy cienega where fishes theft of a few individuals of an imperiled species were isolated. Then, about a century ago, hu- for personal use is not unknown. Such an event mans altered the system, deepening the cienega was possible, even probable, because of popu- into an open pond and diverting spring outflow larity of pupfish with aquarists and vulnerability into a canal, thereby beheading the system. of C. arcuatus to theft while it was under culture Most fishes persisted downstream in a human- by agency and university individuals attempting made impoundment. to perpetuate the species. If such a population Apparently, isolation of Monkey Spring, in ad- exists, its keeper is urged to return it to the dition to protecting and isolating the pupfish, world ownership from which it came. had been sufficiently long and complete that a chub (Gila sp. cf. intermedia) and topminnow Remarks.-Minckley (1973:192-194) initially rec- (Poeciliopsisoccidentalis) both differentiated from ognized this species as an undescribed taxon he other stocks. The topminnow persisting today is called "Monkey Spring pupfish." Otherwise, it genetically unique, proposed as an evolution- has been consistently referred to either C. ma- arily significant unit for conservation purposes cularius, "Monkey Spring pupfish," or "Cyp- (Parker et al., 1999; Sheffer et al., 1999; Minck- rinodon sp." in systematic, biogeographic, and ley, 1999b), and the chub, now extinct, was mor- conservation-oriented works that have noted its phologically distinct from other known popu- existence and extinction (detailed in Minckley lations (DeMarais, 1986). Pupfish extinction et al., 1991). and chub extirpation occurred during repair of Marsh and Sada (1994) reported a nonnative the irrigation system. Attempts to maintain and macularius-likepupfish of questionable origin, il- repatriate both taxa failed because of erroneous licitly stocked (but no longer present) to Bog- assumptions and unexpected events (for full de- Hole Tank, an artificial pond in the extreme

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Santa Cruz headwaters in San Rafael Valley, Ar- though never published, this name is well izona. A similarly questionable population still known in southwestern ichthyological circles, occupies a cattle tank on the private Research and it can be encountered in correspondence, Ranch near Elgin, Arizona, in the adjacent San unpublished materials, and museum collec- Pedro River basin (Marsh and Sada, 1994). Al- tions. though within or just beyond the presumed of these stocked original range C. arcuatus, pop- LITERATURECITED ulations do not represent C. arcuatus. ANASCAVAGE,J. 1973. The Ascencion pupfish. Adv. Etymology.-The name is derived from the Latin Aquar.Mag. 47:3. arcus meaning arch or shaped or bent like a BEAN,R. H. 1898. Notes on Mexican fishes obtained Dr. Carl Lumholz. Bull. Am. Mus. Nat. Hist. 10: bow. This is in reference to the highly convex by 168. dorsal body profile. BLASIUs,H. B. 1996. Biogeographyof freshwaterfishes of northwestern Mexico. Unpubl. master's thesis, Additional material arcu- examined.--Cyprinodon Arizona State Univ., Tempe. atus: CU 18202, UAZ uncat., UAZ 95-79, UAZ BRAND,D. D. 1937. The natural landscape of north- 95-174; C. albivelis: UMMZ 182391, UMMZ western Chihuahua. Univ. New Mexico Bull. 316, 211607, UMMZ 211623; C. eremus: UMMZ Geol. Ser. 5:1-74. 162661; C. macularius: UAZ 65-79, UMMZ BURKHAM,D. E. 1970. Depletion of streamflowby in- 162680. filtrationin the main channel of the Tucson Basin, southeastern Arizona, p. 827-863. In: Floods of 1965 in the United States. U.S. Geological Survey ACKNOWLEDGMENTS Water-SupplyPap. 1939-B,Reston VA. CHERNOFF,B., ANDR. R. MILLER.1982. Notropisboca- This report was in near final form when Pro- grande,a new cyprinid fish from Chihuahua,Mexi- fessor Minckley passed away unexpectedly on 22 co, with comments on Notropisformosus. Copeia June 2001. Because Professor Miller was no lon- 1982:514-533. DELA A. 1970. in the ger active because of advancing age, SMN (who CONDE TORRE, Streamflow upper Santa and Pima had assisted in this research) was added as third Santa Cruz River Basin, Cruz cos., author to the and Arizona. U.S. Geological SurveyWater-supply Pap. complete manuscript arrange 1939-A,Reston, VA. for its review, revision, and A. A. publication. CoNTRERAS-BALDERAS,S. 1969. Perspectivasde la ictio- Echelle (OSUS), P. J. Unmack (ASU), P. C. fauna de las zonas aridas del norte de Mexico. In- Marsh (ASU), and S. Contreras-Balderas ternat. Cent. Arid-SemiaridLand Stud., Tex. Tech. (UANL) assisted in this. Figure 3 was prepared Univ. Publ. 3:293-304. with the assistance of M. H. Gach, using Fish- , V. LANDA-SALINAS,T. VILLEGAS-GAYrAN,AND G. map by P. A. Buckup. Photographs in Figure 2 RODRIGUEZ-OLMOS.1976. Peces, Piscicultura,Pre- were taken by E. Theriot (UMMZ), other pho- sas, Poluci6n, Planificaci6n Pesquera y Monitoreo en la Danza de las P. Memoriasdel Prim- tographs by unspecified UMMZ staff. Many of Mexico, o the Rio were collected er Simposio de Pesqueriasen Aguas Continentales, Yaqui system specimens 1:315-346.Secretaria de Industria Sub- under Permit 3618 issued de y Comercio, by Departamento secretariade Pesca, Mexico. Mexico. and Miller's Pesca, Minckley original COOKE,R. U., AND R. W. REEVES.1976. and follow unaltered: We are in- Arroyos acknowledgments environmentalchange in the AmericanSouthwest. debted to C. L. Hubbs for sharing his insights Clarendon Press, Oxford. and data on pupfishes, without which their de- DE BUEN,F. 1940. Lista de peces de agua dulce de scriptions would be far less detailed. We dedi- M6xico, en preparacionde su cataloga.Estac. Lim- cate this paper to his memory. M. L. Smith also nol. Patzcuaro,Trab., 2:1-66 (mimeo.). contributed information and data toward com- -- . 1947. Investigaciones sobre ictiologia mexi- cana. de los de la neartica of the manuscript. S. M. Norris provided I. Catalogo peces region pletion en suelo mexicano. Ann. Inst. Biol. 18: measurements, counts, and other (Mexico) comparative 257-292. amenities. We thank T. Hunt of the especially DEMARAIS,B. D., 1986. Morphological variation in Rail-X Ranch, Arizona, where Monkey Spring is Gila (Teleostei: Cyprinidae) and geologic history: located, the long-time manager (through a Lower Colorado RiverBasin. Unpubl. master'sthe- number of owners), for access and other cour- sis, Arizona State Univ., Tempe. tesies over the years. ECHELLE, A. A. 1993. Allozyme perspective on mito- chondrial DNA variation and evolution of the Note added in proof: Cyprinodonpisteri was Death Valley pupfishes (Cyprinodontidae:Cyprino- discussed in an unpublished manuscript by C. don). Copeia 1993:275-287. L. Hubbs (circa 1935), in which Hubbs intend------, ANDT. E. DOWLING.1992. MitochondrialDNA ed to name the species "Cyprinodonartatus." Al- variationand evolution of the Death Valleypupfish-

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es (Cyprinodon,Cyprinodontidae). Evolution 46: MAYDENR. L., ANDD. M. HILUS. 1990. Natural history 191-206. and systematicsof the bigmouth shiner, Cyprinella 1- ANDA. F. ECHELLE.1978. The Pecos pupfish, bocagrande(Teleostei: Cypriniformes), with com- Cyprinodonpecosensis, n. sp. (Cyprinodontidae),with ments on conservation status. Copeia 1990:100- comments on its evolutionaryorigin. Copeia 1978: 111. 569-582. MEARNS,E. A. 1907. Mammalsof the Mexicanbound- -, R. A. VANDEN BUSSCHE, T. P. MALLOYJR.,M. ary of the United States:a descriptivecatalogue of L. HAYNIE,AND C. O. MINCKLEY.2000. Mitochon- the species of mammalsoccurring in that regional; drial DNA variation in pupfishes assigned to the with a general summaryof the natural history,and species Cyprinodonmacularius (Atherinomorpha: Cy- a list of trees. Bull. U.S. Natl. Mus. 56(Pt. 1):i-xv, prinodontidae): Taxonomic implications and con- 15-30. servation genetics. Ibid.2000:353-364. MEEK,S. E. 1902. A contribution to the ichthyology EMORY,W. H. 1857. Chap. 1, Part II, General geolog- of Mexico. Publ. Field Columbia Mus. (Zool. Ser.) ical features of the country, p. 1-24. In: Report on 3:63-128. the United Statesand Mexicanboundary survey. W. . 1904. The fresh-water fishes of Mexico north H. Emory. Vol. 1. Sen. Exec. Doc. No. 108, 34th of the Isthmusof Tehuantepec. Ibid.5:i-lxii, 1-252. Congr., 1st Sess. A.O.P. Nicholson, Printer,Wash- MILLER,R. R. 1943. The status of Cyprinodonmacular- ington, DC. ius and Cyprinodonnevadensis, two desert fishes of EVERMANN,B. W. AND E. L. GOLDSBOROUGH.1902. A western North America.Occ. Pap. Mus. Zool., Univ. report on fishes collected in Mexico and Central Mich. 473:1-25. America, with notes and descriptions of five new 1948. The cyprinodont fishes of the Death species. Bull. U.S. Fish Comm. 21:137-159. Valley system of eastern Californiaand southwest- FOWLER,H. W. 1916. Notes on fishes of the orders erna_--e. Nevada. Misc. Publ. Mus. Zool. Univ. 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vation and management of short-livedfishes: ex- Fishes. C. H. Hocutt and E. O. Wiley (eds.). John amples from the cyprinodontoids,p. 247-282 + lit. Wiley and Sons, New York. cited. In: Battle against extinction: native fish man- SUBLETTE,J. E., M. D. HATCH,AND M. SUBLETTE.1990. agement in the AmericanWest. W. L. Minckleyand Fishes of New Mexico. Univ. of New Mexico Press, J. E. Deacon (eds.). Univ. of ArizonaPress, Tucson. Albuquerque. PARKER,K. M., R.J. SHEFFER,AND P. W. HEDRICK. 1999. TURNER, B. J., AND R. L. LIU.1977. Extensiveinterspe- Molecular variation and evolutionarilysignificant cific genetic compatibilityin the NewWorld killifish units in the endangered Gila topminnow.Conserv. genus Cyprinodon.Copeia 1977:259-269. Biol. 13:108-116. WILLIAMS,J. E., J. E. JOHNSON, D. A. HENDRICKSON,S. D. M. PROPST,D. E., ANDJ. A. STEFFERUD.1994. Distribution CONTRERAS-BALDERAS, J. WILLIAMS, NAVARRO- D. E. AND E. DEACON. 1989. and status of the Chihuahua chub (Teleostei: Cy- MAEDA, MCALLISTER, J. Gila with notes on its Fishes of North America:endangered, threatened, prinidae: nigrescens), ecology or of concern: 1989. Fisheries 14:2-20. and associatedspecies. Southwest.Nat. 39:224-234. special WOODS,M. L., P. K. HOUSE ANDP. A. PEARTHREE.1999. REGAN,C. T. 1907. Pisces. Biologia Centrali-America- na 8:33-160. Historical geomorphology and hydrology of the Santa Cruz River. Arizona Geol. Surv. Open-file RINNE, J. N., AND W. L. MINCKLEY.1991. Native fishes Rept. 99-13:1-92 , maps. of arid lands: a dwindling resource of the desert southwest.U.S. For. Ser. Gen. Tech. RM-206: Rept. (WLM) DEPARTMENT OF BIOLOGY, ARIZONA 1-43 + covers, frontis. STATE UNIVERSITY,TEMPE, ARIZONA; (RRM) SHEFFER, R. P. W. HEDRICK, AND A. L. VELASCO. J., FISH DvISION, MUSEUM OF ZOOLOGY,UNIVER- 1999. Testing for inbreeding and outbreeding de- SITY OF MICHIGAN, ANN ARBOR, MICHIGAN pression in the endangered Gila topminnow.Anim. Conserv.2:121-129. 48109; AND (SMN) DEPARTMENTOF ZOOLOGY, MIAMI OXFORD, OHIO 45056. PRE- SMITH, M. L., ANDR. R. MILLER.1980. Systematics and UNIVERSITY, SENT 1 UNIVERSITY variation of a new cyprinodontid fish, Cyprinodon ADDRESS: (SMN) DRIVE, fontinalis,from Chihuahua,Mexico. Proc. Biol. Soc. CALIFORNIA STATE UNIVERSITY-CHANNEL IS- Wash.93:405-416. LANDS, CAMARILLO, CALIFORNIA93012. Send , AND . 1986. The evolution of the Rio reprint requests to SMN. Submitted: 19 Nov. Grande basin as inferred from its fish fauna, p. 2001. Accepted: 1 March 2002. Section editor: 457-485. In: Zoogeography of North American S. A. Schaefer.

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