Variation in the Rare Lizard Barisia Rudicollis (Wiegmann) (Anguidae) with Description of a New Species from Central Mexico

Herpetologica, 58(3), 2002, 313–326 ᭧ 2002 by The Herpetologists’ League, Inc.

VARIATION IN THE RARE LIZARD BARISIA RUDICOLLIS (WIEGMANN) (ANGUIDAE) WITH DESCRIPTION OF A NEW SPECIES FROM CENTRAL MEXICO

ALEJANDRO ZALDı´VAR-RIVERO´ N1 AND ADRIA´ N NIETO-MONTES DE OCA2 Museo de Zoologı´a, Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico, Me´xico, Distrito Federal 04510, Me´xico

ABSTRACT: We describe a new species of Barisia from the oak-pine and cloud forests on the southern slopes of the central Transvolcanic Belt in the states of Mexico and Morelos, Mexico, and report on variation in a sample of 21 specimens of the poorly known B. rudicollis. The new species and B. rudicollis appear to have allopatric distributions and to be sister taxa. Because of increasing fragmentation and destruction of their habitat, we suggest that these species be regarded as threatened. We discuss synapomorphies for the genus Barisia. Key words: Anguidae; Barisia; Barisia rudicollis; Gerrhonotinae; Mexico; New species

THE MEXICAN, anguid lizard genus Bar- Hacienda La Gavia in western Mexico, was isia contains three recognized species. reported by Martıń del Campo (1939) over Barisia levicollis ranges in the Sierra Tar- 100 yr after the species was described, and ahumara and Sierra del Nido in western was subsequently examined by Tihen and north-central Chihuahua, respectively (1949) in his revision of Barisia. This spec- (Guillette and Smith, 1982; Lemos-Espin- imen also served as the basis for the real et al., 2000); B. imbricata, with four sub- striction of the type-locality of B. rudicollis species, occurs from southern Chihuahua (Smith and Taylor, 1950). A second spec- south and east to central and southern Oa- imen from the same locality was examined xaca (Good, 1988; Guillette and Smith, by Good (1988) in his phylogenetic anal- 1982; Lemos-Espinal et al., 2000); and B. ysis of the subfamily Gerrhonotinae (in rudicollis occurs in western and south- fact, the only specimen examined) and also western regions of the state of Mexico (ϭ used by Gonza´lez-Romero and Lo´pez- ‘‘Mexico’’ for simplicity hereafter) and the Gonza´lez (1990) for natural history obser- northwestern corner of Morelos (McCoy vations. More recently, Schmidt (1991) re- and Fox, 1992; Schmidt, 1991). ported on three specimens from Sultepec, Wiegmann (1828) described Gerrhono- in southwestern Mexico. However, he pro- tus rudicollis on the basis of three syntypes vided only brief information on the mor- from ‘‘Mexico.’’ A female syntype con- phology and color pattern of these speci- tained five embryos, which were examined mens. Finally, McCoy and Fox (1992) re- by Wiegmann in a subsequent, more de- described B. rudicollis on the basis of tailed redescription of the species (Wieg- three specimens: two from Valle de Bravo mann, 1834). A few years later, G. rudi- in western Mexico and one from the collis and other species of this genus were Parque Nacional Lagunas de Zempoala in transferred to the genus Barisia by Gray the northwestern corner of Morelos (ad- (1838). Although B. rudicollis was one of jacent to Mexico). They did not examine the first gerrhonotine lizards described, the specimens previously reported by oth- few specimens have been reported in the er workers. literature since its description by Wieg- Examination of several specimens as- mann. The first of these specimens, from signed to B. rudicollis in three Mexican collections suggested that this taxon rep- 1 PRESENT ADDRESS: Department of Biology, Im- resented a composite species. Subsequent perial College at Silwood Park, Ascot, Berkshire SL5 7PY UK. field work in several localities in the states 2 CORRESPONDENCE: e-mail, [email protected]. of Mexico and Morelos and an analysis of unam.mx allozyme variation between some of the

313 314 HERPETOLOGICA [Vol. 58, No. 3

TABLE 1.—Enzymes, their abbreviations, and buffer system used in electrophoretic analyses.

Abbreviation Enzymes of enzymes Buffer Glutamate-oxaloacetate-transaminate GOT Tris-malate 0.05 M, pH 7.8 Phosphogluconate Dehydrogenase PGDH Tris-malate 0.05 M, pH 7.8 Phosphoglucomutase PGMI-1 Tris-malate 0.05 M, pH 7.8 Phosphoglucomutase PGMI-2 Tris-malate 0.05 M, pH 7.8 Malate Dehydrogenase MDH Tris-malate 0.05 M, pH 7.8

sampled populations confirmed the exis- (HW) and head length (HL) were mea- tence of an undescribed species (Zaldı´var- sured at the level of maximum width on Riveroń, 1998). Several additional speci- the temporal region and from the tip of mens of B. rudicollis also were obtained. the snout to the anterior margin of the Herein, we formally describe the new spe- tympanum, respectively. All measurements cies and report on variation in a sample of are given in millimeters. Notes on color in 21 specimens of the poorly known B. rud- life were taken in the field. icollis. In addition to the morphological data, we used standard techniques of cellullose METHODS acetate-gel electrophoresis (Richardson et We examined a total of 20 specimens al., 1986) to analyze allozyme variation labeled B. rudicollis in herpetological col- among one specimen from eastern Mi- lections of the Universidad Autońoma de choacań (Tuxpan), two from western Mex- Morelos (EBUM) and Universidad Na- ico (Valle de Bravo), and three from Mex- cional Autońoma de Me´xico [Museo de ico contiguous to northwestern Morelos Zoologıá of the Facultad de Ciencias (Ocuilan). The liver was removed from (MZFC) and Instituto de Biologıá (IBH)]. each specimen and maintained at Ϫ76 C In addition, we collected 27 specimens until its use in the laboratory; carcasses representing B. rudicollis or the undescri- were preserved and deposited in the bed species in several field trips to the MZFC (see Appendix). The enzymes, states of Mexico and Morelos. Specimens their abbreviations, and the gel/buffer sys- collected (except for a molt and a skull) tem used are given in Table 1. were fixed in 10% buffer formalin, stored in 70% ethanol, and deposited in the RESULTS MZFC. We compared all of the above Specimens from eastern Michoacań and specimens with all of the species and sub- western and southwestern Mexico differed species previously described in the genus. consistently in scutellation, color, and pat- A list of the specimens examined and their tern from those from northern and north- localities is provided in the Appendix. Ac- eastern Morelos and areas of Mexico con- ronyms for museum collections follow tiguous with northwestern Morelos. Like- Leviton et al. (1985), except for EBUM, wise, for each of the five enzymes that was MZFC, and EBUAP (Escuela de Biologıá, consistently scored in the samples exam- Universidad Autońoma de Puebla). ined (Table 2), a single electromorphic Nomenclature of scales follows Waddick variant was found in all the specimens and Smith (1974) and Campbell and Frost from eastern Michoacań and western Mex- (1993) in the cases of gular and occipital ico, whereas a single, alternate variant was scales, respectively, and Good (1988) in found in all the specimens from areas of the case of the remaining scales. Some Mexico contiguous with northwestern Mo- scale counts were recorded bilaterally; relos (Fig. 1). Thus, geographic variation therefore, the total of counts for such in external morphology was congruent characters is twice the number of speci- with that found in the enzymes assayed, mens. Measurements were taken with cal- and both morphological and electropho- ipers to the nearest 0.1 mm. Head width retic evidence indicate that the examined September 2002] HERPETOLOGICA 315

TABLE 2.—Allozyme variation among populations of Barisia herrerae sp. nov. Barisia examined. Holotype.—An adult male (MZFC Locus 9580, field number AZR 039), collected on Specimen GOT 6PGDH PGMI-1 PGMI-2 MDH 21 September 1996, approximately 4 km Ocuilan E Ocuilan (approximately 15 km W Cuer- MZFC 9580 A A A A A navaca City limits) on dirt road Cuerna- MZFC 9587 A A A A A vaca-Chalma, near Rancherıá Tlaltizapań, MZFC 9588 A A A A A Municipality of Ocuilan, state of Mexico, Valle de Bravo Mexico; 99Њ 23Ј 25Љ N, 18Њ 57Ј 16Љ W; 2350 MZFC 9585 B B B B B m elevation. MZFC 9586 B B B B B Paratypes.—Twenty-three; eight adults Tuxpan from the type-locality (two males, four fe- MZFC 9594 BBBBB males, one molt, and one skull; MZFC 9588 and 5741; MZFC 9584–85, 9587, and 12549; MZFC 12551; and MZFC populations represent two evolutionary 9592, respectively); 12 young born from species. Populations from eastern Michoa- MZFC 9587 and MZFC 12549 (MZFC cań and western and southwestern Mexico 9593 and 12552–562), and three adults represent B. rudicollis (see Discussion), from the state of Morelos: one male each whereas populations from northern and from Cuernavaca City limits (MZFC northeastern Morelos and areas of Mexico 12550) and the Sierra del Chichinautzin adjacent to northwestern Morelos repre- (MZFC 9589), and one female from Tetela sent an undescribed species. We propose del Volcań (EBUM 2485). this species to be known as: Diagnosis.—Barisia herrerae (n ϭ 23–

FIG. 1.—Locality records for Barisia herrerae (squares) and B. rudicollis (circles). The hollow square represents the type-locality of B. herrerae. Hollow symbols represent populations examined for allozyme variation. Shaded areas represent, from palest to darkest, 0–1000 m, 1000–2000 m, 2000–3000 m, 3000–4000 m, and above 4000 m of elevation, respectively. Discontinuous lines represent political boundaries (GRO ϭ Guerrero; D. F. ϭ Distrito Federal). The question mark represents the Lagunas de Zempoala record. 316 HERPETOLOGICA [Vol. 58, No. 3

24, see below) may be distinguished from all other species and subspecies of the genus (Table 3) by having two anterior gulars between the second chinshields (one an- 4 terior gular in Ն92% of the specimens in 2 4 4–6 (6) 3 the other taxa) and usually (in 78.3% of the No specimens) five scales in the first temporal row, three of them between the levels of the dorsal and ventral margins of the orbit * (four scales in the first temporal row, two of them between the levels of the dorsal Barisia. 4 9–10 (10) 1–3 (1) 1 1 and ventral margins of orbit, in Ն95% of No the specimens in the other taxa). Barisia herrerae also differs from all the other taxa in the genus, except B. rudicollis, in having the rostral in contact with the B. i. planifrons B. levicollis B. rudicollis nasal (in 96% of the specimens); cantho- 3–4 (4) 8 1 1 No loreal series divided vertically (two loreal Usually 3 scales, the anterior one smaller than the posterior); usually (in 89.6% of the specimens) four superciliaries; three occipitals; 4–6 nuchals, the lateral ones strongly keeled, projecting laterally; 29–33 (x¯ ϭ 31) 4–5 (4) 8 1–3 (1) 1 No transverse dorsal scale rows; and 14 lon- Uusally 3 gitudinal ventral scale rows. In the other taxa, the rostral is always separated from the nasal; the cantholoreal series is not divided (one loreal scale) or is divided horizontally (one canthal and one loreal scale); 4 1 8–10 (8) 1–3 (1) No there are usually three or less supercili- Usually 3 aries (four superciliaries in Յ9% of the specimens); one occipital (Ն95%); and 8– 10 nuchals, the lateral ones only weakly keeled, not projecting laterally; and there 4–5 (4) 1 are 34 or more transverse dorsal scale rows 8–10 (8) 1–3 (1) No and 12 longitudinal ventral scale rows. Usually 3 In addition to the above characters, B. herrerae may be distinguished from B. rudicollis in having no dark marks on the subocular region, immaculate ventral surface and olive-green dorsum in adults, and B. herrerae B. i. ciliaris B. i. imbricata B. i. jonesi 3–5 (5) 2 4 4–6 (6) 3 3.—Variation (and mode) in selected characters among recognized species and subspecies of each scale on the dorsal surface finely stip- 25% 23–24 (see text); sample sizes for other taxa as in Appendix.

pled with white in adult males. In B. rud- ϭ ABLE icollis, there is a dark, vertical stripe on the T posterior subocular and adjacent supralabial; dark, short irregular spots on the gu-

lar region and sides of chest and abdomen Barisia herrerae

(more abundant in largest males); a pale Character to dark brown to brownish gray dorsum; and much larger, irregular white marks re-

stricted to the dorsal and lateral surfaces Sample size for temporal row contact 96% No No No No No 100% dorsal scale rowsdorsal scale rowsventral scale rows 14 29–33 (31) 14 39–45 (42) 16 36–41 (38) 12 36–41 (38) 12–14 (14) 35–42 (36) 16 12 45–50 (48) 16 28–31 12 (29) 16 12 12 14 14 * Number of anterior gularsNumber of scales on first Presence of 2 postrostral 1–2 (1) 1–2 (1) 1 1 1 1 Nasal and rostral in Number of loreals Number of superciliaries of the head and limbs and posterior mar- Nuchal scalesNumber of nuchals Number of occipitals Number of longitudinal Number of transverse Number of Strongly longitudinal keeled Weakly keeled Weakly keeled Weakly keeled Weakly keeled Weakly keeled Strongly keeled September 2002] HERPETOLOGICA 317 gins of the dark transverse bands in adult males. Description of the holotype.—Medial postrostral present, small, maximum width h length, flanked on either side by one anterior internasal (Fig. 2a); postrostral and anterior internasals subequal in size; posterior internasals approximately thrice as large as anterior ones; frontonasal absent; two prefrontals, slightly larger than posterior internasals; frontal about twice as long as wide, in contact with prefrontals, second and third medial supraoculars, frontoparietals, and interparietal; five medial and three lateral supraoculars; four superciliaries, anterior two about as high as long, subequal in size, posterior two slightly elongate, distinctly shorter and lower than anterior ones; suture between anteriormost superciliary–posterior loreal approximately 1.6/1.3 times as long as suture between anteriormost medial supraocular– posterior loreal; two frontoparietals; interparietal about twice as long as wide, slightly rugose, sunken relative to adjacent scales; two parietals, slightly shorter and broader than interparietal; two small, obliquely-oriented occipitals in medial contact with each other, bordered posteriorly by third, much larger, medial occipital; parietals followed posteriorly by one enlarged scale on either side of medial occipital; medial occipital and flanking scales subequal in size, slightly smaller than parietals and interparietal; nasal moderately elongate (Fig. 2b), narrowly separated from rostral; supranasal in contact with postnasal, separated from anterior loreal by posterior internasal and postnasal; cantholoreal series divided vertically, composed of two loreals; anterior loreal smaller than posterior loreal, in contact with postnasal, posterior internasal, and third supralabial; posterior loreal in contact with FIG. 2.—Barisia herrerae (holotype, MZFC 9580), third and fourth supralabials, preocular, an adult male. (a) Dorsal, (b) lateral, and (c) ventral anteriormost medial supraocular, and an- aspects of head. Head length ϭ 29.3 mm. Horizontal teriormost superciliary; one preocular; two lines ϭ 10 mm. suboculars; posterior one elongate, approximately twice as long as anterior one; four postoculars; four scales in first tem- than remaining ones; infralabials 9/9; two poral row, three between levels of dorsal postmentals (Fig. 2c), each in contact with and ventral margins of orbit; 11 suprala- two infralabials and one sublabial; sublabials, posteriormost one more elongate bials 6/5; chinshields 4/4, first pair in me- 318 HERPETOLOGICA [Vol. 58, No. 3

Variation.—In addition to the holotype, we examined a total of 23 paratypes (see above) to document morphological variation in B. herrerae. We describe below the variation in the entire sample (including the holotype) for each character that showed conditions other than that found in the holotype. Rostral contacting nasal in all specimens except holotype. One medial postrostral present in six specimens; absent in remaining ones. One postnasal on either side in 12 specimens; on one side in five; absent on either side in remaining specimens; nasal and supranasal broadly contacting anterior loreal in specimens with postnasal absent. Superciliaries 3–4, x¯ ϭ 3.9 (3/3, 2; 3/4, 1; 4/4, 21). Supralabials 9– 11, x¯ ϭ 10.5 (9/10, 1; 10/10, 9; 10/11, 4; 11/11, 10). The following characters could not be recorded in MZFC 9592 (skull only); therefore, n ϭ 23. Temporal rows 3/3 in one specimen, 4/4 in remaining ones. Scales in first temporal row 3–5, x¯ ϭ 4.7 (3/3, 1; 4/4, 1; 4/5, 6; 5/5, 15). Scales in FIG. 3.—Barisia herrerae, holotype (MZFC 9580), first temporal row between levels of dorsal an adult male, SVL ϭ 124.0 mm. and ventral margins of orbit 2–3, x¯ ϭ 2.8 (2/2, 2; 2/3 or 3/2, 6; 3/3, 15). Infralabials dial contact, second pair separated by two 7–10, x¯ ϭ 8.2 (7, 5; 8, 11; 9, 4; 10, 3). gulars. Postmental separated from sublabials on Minimum number of scales in trans- either side in one specimen; on one side verse nuchal rows six; lateral nuchals mod- in another one; contacting one sublabial erately to strongly keeled, projecting lat- on either side in remaining specimens. erally; dorsals in 32 transverse and 14 lon- Sublabials 5–6, x¯ ϭ 5.2 (5/5, 17; 5/6, 3; 6/ gitudinal rows; ventrals in 54 transverse 6, 3). Chinshields 4–5, x¯ ϭ 4.1 (4/4, 21; 5/ and 14 longitudinal rows; six preanals. 5, 2). Minimum number of nuchals 4–6, x¯ Measurements.—Snout–vent length ϭ 4.9 (4, 11; 5, 4; 6, 8). Dorsals in 29–33 (SVL) ϭ 124.0, HW ϭ 23.5, HL ϭ 29.3, transverse rows, x¯ ϭ 31.0 (29, 2; 30, 5; 31, tail length ϭ 159.0. 9; 32, 4; 33, 3). Ventrals in 50–57 trans- Color in life.—Dorsal surface of head verse rows, x¯ ϭ 53.2 (50, 1; 51, 1; 52, 8; olive-green (Fig. 3); lateral surfaces above 53, 5; 54, 4; 55, 3; 57, 1). upper lip olive-green, becoming gradually Measurements.—Measurements (SVL, paler ventrally; lateral surfaces below low- HW, and HL) taken from 20 specimens er lip and ventral surface immaculate (12 newborns, 4 adult females, and 4 adult cream; dorsal and lateral surfaces of body males, including the holotype) are given in and tail olive-green; seven faint, dark Table 4. green transverse bands between levels of Color pattern.—There was sexual and axilla and groin; approximately 19 trans- ontogenetic variation in color pattern. verse bands on tail; dorsal and lateral Newborns (n ϭ 12): Dorsal and lateral sur- scales of head, body, limbs, and tail finely faces brownish gray; 7–8 faint, dark brown stippled with white; ventral surface im- transverse bands between levels of axilla maculate cream. and groin, each bordered posteriorly by September 2002] HERPETOLOGICA 319

TABLE 4.—Measurements (mean Ϯ standard error, over range) in specimens examined of Barisia herrerae and B. rudicollis.

B. herrerae B. rudicollis Measure- Adult Adult Subadult Adult Adult ments Newborns males females Newborns male males females n 12 4 4 3 1 8 7 SVL 37.7 Ϯ 0.7 122 Ϯ 1.7 110.5 Ϯ 3.4 37.3 Ϯ 0.3 82 103.2 Ϯ 3.8 111.7 Ϯ 5.3 (34–42) (117–125) (105–119) (37–38) (85–123) (86–127) HW 7 Ϯ 0.2 23.2 Ϯ 0.2 17.2 Ϯ 0.5 6.6 Ϯ 0.3 16 20.4 Ϯ 0.8 18.8 Ϯ 1.1 (6–8) (23–24) (16–18) (6–7) (18–25) (15–24) HL 9.8 Ϯ 0.2 27.7 Ϯ 0.5 22.5 Ϯ 0.9 9.3 Ϯ 0.3 19 24.5 Ϯ 0.8 24.4 Ϯ 0.8 (9–11) (27–29) (21–24) (9–10) (22–29) (22–27)

narrow white line; white stippling absent. ventral surface absent in both males and Dark, narrow vertical stripe extending on females. to posterior subocular and adjacent su- Distribution and ecology.—Barisia her- pralabial; few, faint dark speckles on sides rerae occurs at several localities in north- of throat and body. ern and extreme northeastern Morelos and Adults.—Dorsal and lateral pattern in areas of Mexico contiguous to northwest- males and females similar to that of holo- ern Morelos (Fig. 1). These localities lie type, except white stippling absent in fe- on the southern slopes of the Transvolcan- males (Fig. 4, top). Dark line on subocular ic Belt at about 2350–2500 m elevation. and supralabial scales and dark speckles on Vegetation near the city of Cuernavaca and Tetela del Volca´n is oak-pine forest, whereas that at the type-locality (Ocuilan, Fig. 5) and the Sierra del Chichinautzin is cloud forest (Contreras-McBeath and Ur- bina-Torres, 1995; Luna-Vega et al., 1989). The holotype of B. herrerae was collected on a branch of a fallen tree. The remaining specimens were found on the ground, usually hiding under logs and leaf litter, in relatively open areas in the forest. Two pregnant females of B. herrerae (MZFC 9587 and 12549) were collected

FIG. 5.—Type locality of Barisia herrerae, along FIG. 4.—(Top) Barisia herrerae, an adult female Cuernavaca–Chalma dirt road, approximately 4 km E (MZFC 13929), SVL ϭ 117 mm; (Bottom) B. rudi- Ocuilan, Municipality of Ocuilan, Mexico, Mexico collis, an adult female (MZFC 12541), SVL ϭ 111.5 (99Њ 23Ј 25Љ N, 18Њ 57Ј 16Љ W; 2350 m elevation). mm. Photograph taken on September 21, 1996. 320 HERPETOLOGICA [Vol. 58, No. 3 on 20 May 1997 and maintained in captiv- ternasals; posterior internasals approxi- ity. They gave birth to five and seven mately twice as large as anterior ones; young, respectively, on 15–18 June 1997. frontonasal present in 11 of 21 specimens; In two other adult females, collected in two prefrontals, slightly larger than poste- October (MZFC 9584) and November rior internasals; frontal about twice as long (MZFC 9585), the size of the largest fol- as wide, in contact with prefrontals, sec- licle was 8.92 and 7.23, respectively. This ond and third medial supraoculars, fron- indicates that ovulation occurs in late fall toparietals, and interparietal; five medial and parturition in summer. This reproduc- and three lateral supraoculars; supercili- tive pattern is also exhibited by all of the aries 3/4 in one specimen, 4/4 in others; subspecies of B. imbricata (Guillette and suture between anteriormost superciliary– Casas-Andreu, 1987). posterior loreal longer than suture be- Three specimens from Ocuilan (MZFC tween anteriormost medial supraocular– 9585 and 9587–88) contained numerous posterior loreal; two frontoparietals; inter- nematodes in the lung cavity. These nem- parietal approximately twice as long as atodes (eggs and parthenogenetic females) wide, rugose, sunken relative to adjacent appeared to belong to the genus Entome- scales in adults; two parietals, slightly las (Rhabdiasidae); however, no species of shorter and broader than interparietal; two this genus are known from the Western small, obliquely-oriented occipitals in me- Hemisphere. The four species of Ento- dial contact, bordered posteriorly by third, melas known to date occur in Europe, much larger, medial occipital; parietals fol- Asia, or southern Africa (Baker, 1980, lowed posteriorly by one enlarged scale on 1982). Of these species, only E. entomelas either side of medial occipital; medial oc- is known from European anguid hosts cipital and flanking scales subequal in size, (Ophisaurus apodus and Anguis fragilis; slightly smaller than parietals and inter- Baker, 1980). Thus, the identity of the parietal; nasal elongate, in contact with nematodes found in B. herrerae remains rostral; one supranasal; postnasal absent to be corroborated. (supranasal in contact with anterior loreal) Etymology.—The name herrerae is a on either side in 10 specimens, one post- patronym based on that of Alfonso L. Her- nasal separating supranasal from anterior rera, one of the greatest naturalists in loreal on either side in remaining speci- Mexico in the nineteenth century. mens; cantholoreal series divided vertical- Remarks.—The presence of a postros- ly, composed of two loreals (posterior lo- tral scale in 6 of the 24 specimens exam- real horizontally divided in one specimen); ined of B. herrerae is unique to this spe- anterior loreal smaller than posterior lo- cies within the genus. real; 1 preocular; 2 suboculars in 12 specimens, 3 in 6, 4 in 3; postoculars 2/1 Barisia rudicollis (Wiegmann), 1828 in one specimen, 2/3 in another one, 3/3 Gerrhonotus rudicollis Wiegmann, 1828, in 17, 4/3 in 1, 4/4 in 1; temporal rows 3/ Isis, 21:380. 3 in 20 specimens, 4/3 in 1; four scales in Barisia rudicollis, Gray, 1838, Ann. Mag. first temporal row, two between levels of Nat. Hist., ser. 1, 1:390. dorsal and ventral margins of orbit; supralabials 10–12, x¯ ϭ 10.6 (10/10, 8; 10/11, Variation.—Except for characters that 1; 10/12, 2; 11/11, 9; 12/12, 1). could not be recorded from a skull (see The characters below, except for the below), the following description of varia- number of transverse nuchal scale rows, tion in B. rudicollis is based on a total of could not be recorded in the examined 21 specimens: 17 adults (nine females, six skull; thus, n ϭ 20. Infralabials 7–10, x¯ ϭ males, one skull, and one specimen of un- 8.5 (7/7, 4; 8/8, 3; 9/8, 1; 9/9, 8; 9/10 or known sex), one subadult male, and three 10/9, 3; 10/10, 1); two postmentals; post- newborns. mental contacting one sublabial on one Scutellation.—Medial postrostral ab- side in one specimen, on either side in sev- sent. Two anterior and two posterior in- en, separated from sublabial in remaining September 2002] HERPETOLOGICA 321 specimens; contacting one infralabial on either side in one specimen, contacting two infralabials on either side in remaining ones; sublabials 5/5 in 7 specimens, 5/6 or 6/5 in 4, 6/6 in 9; chinshields 4/4 in 19 specimens, 4/5 in remaining one; one anterior gular. Minimum number of scales in transverse nuchal scale rows 4–6, x¯ ϭ 5.5 (4, 4; 5, 1; 6, 15); lateral nuchals strongly keeled, projecting laterally; dorsals in 14 longitudinal scale rows, in 28–31 transverse scale rows, x¯ ϭ 29.2 (28, 4; 29, 9; 30, 5; 31, 2); ventrals in 14 longitudinal scale rows, in 47–54 transverse scale rows, x¯ ϭ 50.0 (47, 1; 48, 4; 49, 5; 50, 2; 51, 4; 52, 1; 53, 2; 54, 1); six preanals. Measurements.—SVL, HW, and HL from a total of 19 specimens (eight adult and one subadult males, seven adult females, and three newborns) are given in Table 4. Color pattern.—Barisia rudicollis exhib- its moderate sexual dichromatism. Also, adult females retain the color pattern of FIG. 6.—Barisia rudicollis. (Top) a presumably old newborns, whereas males exhibit some on- adult male (MZFC 9590), SVL ϭ 106 mm (see text); togenetic variation. The following descrip- (Bottom) an adult male (MZFC 9586), SVL ϭ 123 tion is based on eight males (seven adults mm. and one subadult), seven adult females, and three newborns. Newborns and adult females (Fig. 4, Males.—Color pattern of MZFC 9590 at bottom). Dorsal and upper lateral surfaces time of collection (already an adult, SVL of head above level of suboculars pale unrecorded) and remaining specimens brown to brownish gray; suboculars and with SVL Յ 102 (n ϭ 5) similar to that of ventral surface of head cream. Short, dark females, except for numerous, small, irreg- vertical line extending on posterior subo- ular white marks unevenly distributed on cular and adjacent supralabial; dark, nar- dorsal and lateral surfaces of head and row, vertical lines extending on anterior limbs and posterior margins of dark trans- and/or posterior margins of anteriormost verse bands on flanks. Pattern in MZFC three or four supralabials. Dorsal and lat- 9590 at time of death (Fig. 6, top), after eral surfaces of body and tail pale brown over 6 yr in captivity (SVL ϭ 106), com- to brownish gray; six or seven 1–3 scale- posed of a grayish brown ground color, wide, vertical to oblique, dark brown pale on dorsum, slightly darker on flanks; transverse bands between levels of axilla dark brown to black, irregular transverse and groin; bands faint on dorsum; evident bands only visible on flanks; large, con- on first 4–5 longitudinal scale rows on spicuous white marks nearly obliterating flanks and granular fold. Dorsal edge of ground color on dorsal and lateral surfaces anteriormost transverse band extending of head, body, limbs, and tail; ventral sur- anteriorly on to first nuchal row. Numer- face with dark marks throughout gular re- ous (18–20, n ϭ 3), dark transverse bands gion, venter, limbs, and tail. Pattern in on dorsal and lateral surfaces of tail. Ven- largest male (MZFC 9586; SVL ϭ 123, age tral surface cream; short, dark irregular unknown; Fig. 6, bottom) similar to that in streaks on gular region and sides of body. smaller males, except dorsal ground color 322 HERPETOLOGICA [Vol. 58, No. 3 dark brown, white marks on dorsal and lat- at the Lagunas de Zempoala. Huitzilac is eral surfaces of head, body, and tail more closer to the type-locality of B. herrerae numerous and evident, dark flecks numer- (approximately 9 km by air), and the veg- ous, present all over ventral surface of etation in these two localities is cloud for- head, body, limbs, tail. est (though drastically modified at Huitzi- Remarks.—McCoy and Fox (1992) not- lac), as opposed to the coniferous forest at ed the presence of a frontonasal scale (ab- Lagunas de Zempoala. In addition, two sent in all other species and subspecies of specimens recently collected at Huitzilac Barisia) in two out of the three specimens or in its vicinity (not included in this work) of B. rudicollis they examined. The speci- undoubtedly represent B. herrerae. Final- mens with a frontonasal were reported as ly, McCoy and Fox’s specimen apparently two females from Valle de Bravo, Mexico looked identical to a photographed speci- (EBUM 1372–73), whereas the remaining men of B. herrerae (McCoy, personal com- specimen was a male from the Lagunas de munication). On the other hand, the main Zempoala, Morelos (EBUM 1356). We ex- problem with the specimen representing amined a total of 21 specimens of B. rud- B. herrerae is the presence of small, dark icollis, including the two from Valle de spots on its ventral surface, although sim- Bravo (which were actually males) exam- ilar marks are found in newborns of this ined by McCoy and Fox (1992), and found species. a frontonasal in 11 (52.4%) of them. The only specimen examined by us pu- The identity of the male from the tatively from the Lagunas de Zempoala ‘‘Parque Nacional Lagunas de Zempoala,’’ (other than specimens of B. imbricata) was Morelos, assigned to B. rudicollis by Mc- an adult female obtained from a zoological Coy and Fox (1992) is problematical. This park in Mexico City, where there was no locality is only about 15 km north of the written record of its provenance. This closest record of B. herrerae (the type-lo- specimen (MZFC 5689) has four scales in cality), whereas it is at least 50 km (by air) the first temporal row (two between the from the closest record of B. rudicollis levels of the dorsal and ventral margins of (Sultepec, Mexico). Unfortunately, this the orbit), a single gular between the sec- specimen was not available in the EBUM ond chinshields, a grayish brown color in collection. However, it was described as preservative, and few, small dark spots on having an uniformly greenish brown dor- the gular region and venter. Again, these sum with no discernable pattern, dark bars characters are present in B. rudicollis and on supralabials reduced to 2–4 dark spots, absent in most or all adult specimens of B. no stripe on the suboculars, and a light herrerae; however, the locality of this spec- gray or tan venter with irregular, small imen is uncertain, and it was not included darker spots, more numerous on the chin in the report of variation for the former and gular region. No white stipples on the species. All other specimens of Barisia ex- dorsal nor lateral surfaces were men- amined by us from Lagunas de Zempoala tioned. Although small dark spots on the and nearby localities represent B. imbri- ventral surface are actually present in the cata. other known populations of B. rudicollis Distribution and ecology.—Barisia rud- (and absent in B. herrerae), the greenish icollis has been recorded from Hacienda dorsum and absence of both dorsal pattern La Gavia (Gonza´lez-Romero and Lo´pez- and subocular stripe suggests that this Gonza´lez, 1990; Martıń del Campo, 1939), specimen might represent B. herrerae. In Valle de Bravo (McCoy and Fox, 1992), a more recent work, McCoy and Fox and 6 km N of Sultepec (Schmidt, 1991), (1996) reported the locality for this spec- Mexico; El Pinal, Municipality of Tuxpan, imen as ‘‘Huitzilac: Parque Nacional La- Michoacań (Zaldı´var-Riveroń and Nieto- gunas de Zempoala,’’ rather than just Montes de Oca, 2001); and the Parque ‘‘Parque Nacional Lagunas de Zempoala.’’ Nacional Lagunas de Zempoala, Morelos Thus, it is possible that the specimen was (McCoy and Fox, 1992). However, we re- actually collected at Huitzilac, rather than gard the last record as doubtful (see September 2002] HERPETOLOGICA 323 above). The newborns examined herein rudicollis (‘‘Mexico’’) is rather vague and from 5 km SE Zitaćuaro, Michoacań, rep- that we did not examine the species’ types, resent the second record of the species for it seems possible that the populations as- this state. Thus, it may be summarized that signed herein to B. herrerae represented B. rudicollis occurs on the southern slopes in reality B. rudicollis, and vice versa. It of the central portion of the Transvolcanic should be remembered that, although Belt in extreme eastern Michoacań, west- Smith and Taylor (1950) restricted the ern Mexico, and possibly extreme north- type-locality of B. rudicollis to Hacienda western Morelos (see above), at 2000– La Gavia in western Mexico, they also did 2550 m of elevation (Fig. 1). Except for not examine the species’ types; thus, they the specimens putatively from the Lagunas also might have attached the name to the de Zempoala, all other specimens of B. wrong species. Furthermore, it seems that rudicollis known to us have been collected Deppe could have collected the types of in cloud or oak-pine forest. Tihen (1949) B. rudicollis within the geographic range and Gonza´lez-Romero and Lo´pez-Gonza´- of any of these species, because he visited lez (1990) suggested that B. rudicollis localities in the proximity of each of them. might be arboreal. However, McCoy and In 1825 and 1826, Deppe travelled several Fox (1996) found some morphological ev- times to Temascaltepec, a locality only idence suggesting that the species is prob- about 15 km SSE of Valle de Bravo in ably terrestrial. In any event, these con- western Mexico, where several specimens trasting hypotheses were based either on of B. rudicollis have been collected. How- inferences drawn from the limb morphol- ever, in 1826 he also visited Tenancingo, ogy of few preserved specimens or the be- only about 18 km West of Ocuilan, type- havior of a single captive specimen, rather locality of B. herrerae, and also made an than on actual field observations. All of the excursion from Temascaltepec to Cuerna- specimens with collecting data examined vaca (Stresseman, 1954). herein were found in strictly terrestrial Although the original descriptions of B. habitats, such as in leaf litter, under pieces rudicollis by Wiegmann (1828, 1834) did of loose bark or fallen trees, or walking on not include any of the scutellation char- the ground, often in forest openings with acters useful to separate it from B. herrer- piled logs and pieces of loose bark used ae, they do state that the dorsum of a fe- for firewood. Lizard remains (a skull and male syntype and her offspring was oliva- a molt) also were found on the ground in ceus greenish gray. This might suggest that forest openings. The available data on the populations in northern Morelos, Mexico reproductive biology of B. rudicollis were contiguous to northwestern Morelos, and reported by Zaldı´var-Riveroń and Nieto- extreme eastern Morelos represent in re- Montes de Oca (2001). ality B. rudicollis. However, color percep- tion is somewhat subjective. Fortunately, DISCUSSION Wiegmann also noted that his specimens Barisia herrerae is morphologically dis- had dark marks on the gular region and tinct from B. levicollis and all of the sub- venter, which are immaculate in the pop- species of B. imbricata. It is also morpho- ulations of B. herrerae. In addition, a logically (if subtly) and electrophoretically drawing of one of the syntypes of B. rud- distinct from B. rudicollis. However, it is icollis (Wiegmann, 1828) shows that it had remarkably similar to the latter species in two scales on the first temporal row be- scutellation, body proportions, and color tween the levels of the dorsal and ventral pattern. The geographic distributions of margins of the orbit, whereas there are these species are likely allopatric, although usually (in 78.3% of the specimens, n ϭ they might be parapatric if the record of 23) three scales in the first temporal row B. rudicollis from the Lagunas de Zem- between the same levels in B. herrerae. poala is valid. At any rate, these geographic Thus, it seems reasonable to assume that ranges are relatively close to each other. populations in extreme western Michoa- Given that the original type-locality of B. cań, western Mexico, and possibly extreme 324 HERPETOLOGICA [Vol. 58, No. 3 northwestern Morelos (see above) actually collis and homoplastically shared with oth- represent B. rudicollis. er genera. In his phylogenetic analysis of gerrhon- Barisia herrerae shares several synapo- otine lizards, Good (1988) proposed eight morphies with B. rudicollis that are not synapomorphies (the first four shared ho- shared with any other species or subspe- moplastically with other gerrhonotine gen- cies in the genus, including the presence era) for the genus Barisia: (1) frontonasal of rostral-nasal contact, usually four super- absent; (2) postoccipitals rugose; (3) 14 ciliaries, three occipitals, four to six nu- longitudinal dorsal scale rows; (4) reduc- chals (the lateral ones strongly keeled, pro- tion in number of transverse dorsal scale jecting laterally), 33 or fewer transverse rows to fewer than 40; (5) postnasal-supra- dorsal scale rows, and 14 longitudinal ven- nasal fusion; (6) canthal-cantholoreal fu- tral scale rows. Thus, these species appear sion; (7) reduction from 5–7 to 1–3 super- to be sister taxa. ciliaries; and (8) characteristically short, The geographic distributions of B. her- deepened snout. However, we found that rerae and B. rudicollis are relatively small, a frontonasal is present in some specimens and it is likely that they are shrinking. The of B. rudicollis; 16 longitudinal dorsal scale main cause for this reduction is the in- rows occur in B. levicollis and B. imbricata creasing fragmentation and destruction of exclusive of B. i. imbricata; there are more their habitat. The forest is quickly disap- than 40 transverse dorsal scale rows in B. pearing as it is improperly exploited for levicollis and most specimens of B. i. cil- lumber and the land is cleared for pasture, iaris; B. rudicollis and B. herrerae have a agriculture, and urbanization. At the type cantholoreal series that is divided vertically locality of B. herrerae, which is in close as in several species of Abronia and Mes- proximity of Chalma and Cuernavaca, ac- aspis; B. rudicollis and B. herrerae usually tive deforestation was occurring during have four superciliaries; and at least one most of the time we were in the area, and species of Abronia (A. frosti) and several the number of human settlements is in- species of Mesaspis appear to have heads creasing. Similarly, the Valle de Bravo re- that are not distinctly longer or more de- gion is rapidly turning into an important pressed than those in Barisia. vacation spot. In addition, most gerrhon- Therefore, three of the characters pro- otine lizards are believed to be poisonous, posed as synapomorphies for Barisia and they are usually killed by local people. (numbers 3, 4, and 6 above) might be ac- All of this, together with their apparently tually spurious, because they are absent in small population sizes, suggests that both some of the taxa in the genus, unless they species should be regarded as threatened. have reversed in these taxa. Also, we re- gard the presence of a short, deepened RESUMEN head as a questionable synapomorphy for Se describe una especie nueva de Ba- the genus, because this character is diffi- risia de los bosques hu´ medos de pino-en- cult to evaluate objectively (partly because cino y de neblina de la vertiente meridio- of sexual and ontogenetic variation) and nal del Eje Neovolcańico Tranversal en los appears to be present in other gerrhono- estados de Me´xico y Morelos, Me´xico, y se tine genera. Similarly, the presence of ru- reporta la variacioń en una muestra de 21 gose postoccipitals has been considered as ejemplares de la poco conocida B. rudi- an ambiguous character in other gerrhon- collis. La nueva especie y B. rudicollis pa- otine genera because it is difficult to eval- recen tener distribuciones alopa´tricas y ser uate objectively (Campbell and Frost, especies hermanas. Debido a la creciente 1993). Thus, only the reduction in the fragmentacioń de su ha´bitat, sugerimos number of superciliaries (to 1–4, not 1–3) que estas especies sean consideradas como and the fusion of supranasal and postnasal amenazadas. Se discuten las sinapomorfıás are uniquely derived, unreversed synapo- para el geńero Barisia. morphies for Barisia, whereas the absence of frontonasal is polymorphic in B. rudi- Acknowledgments.—The following people and in- September 2002] HERPETOLOGICA 325 stitutions loaned specimens: G. Bustos (EBUM), G. H. M. SMITH. 2000. Geographic distribution. Ba- Casas (IBH), H. Eliosa (EBUAP), J. Wiens (CM), D. risia imbricata ciliaris. Herpetological Review 31: Kizirian (LACM), D. Frost and L. Ford (AMNH), D. 112. Rossman (LSUMZ), A. Resetar (FMNH), and R. He- LEVITON, A. E., R. H. GIBBS,JR., E. HEAL, AND C. yer (USNM). M. Feria, A. Gonza´lez, E. Godıńez, and E. DAWSON. 1985. Standards in herpetology and P. Heimes donated valuable specimens. U. Guzmań, ichthyology: part I. Standard symbolic codes for in- W. Schmidt, F. Vargas, and R. Reyes helped with field stitutional resource collections in herpetology and work. K. Oyama and A. Castan˜ eda provided facilities ichthyology. Copeia 1985:802–832. and helped to conduct laboratory work, respectively. LUNA-VEGA, I., L. ALMEIDA-LEN˜ ERO, AND J. LLO- This paper is based in part upon work supported by RENTE-BOUSQUETS. 1989. Florı´stica y aspectos fi- the National Science Foundation under grant no. togeogra´ficos del bosque meso´filo de montanã de DEB-0102383. We also thank the American Museum las can˜ adas de Ocuilan, Estados de Me´xico y Mo- of Natural History (Theodore Roosevelt grant num- relos. Anales del Instituto de Biologıá de la Uni- ber RP66674), CONABIO (grants numbers FB340/ versidad Nacional Autońoma de Me´xico, Serie Bo- H-245/96 and FB284/H-250/96), CONACyT (grant tańica 59:63–87. number 5-28007N), and DGAPA, UNAM (grant MARTıŃDELCAMPO, R. 1939. Contribucioń al co- number IN214198) for financial support for field and nocimiento de los gerrhonoti mexicanos, con la laboratory work. O. Flores and P. Heimes photo- presentacioń de una nueva forma. Anales del Ins- graphed some specimens, M. Villagrań and N. Man- tituto de Biologıá de la Universidad de Me´xico 10: rı´quez helped to collect reproductive data, and D. 353–361. Osorio helped to identify endoparasites. MCCOY,J.K.,AND S. FOX. 1992. Redescription of the Mexican lizard Barisia rudicollis (Wiegmann) LITERATURE CITED (Sauria: Anguidae). Copeia 1992:114–117. . 1996. Is Barisia rudicollis (Sauria: Anguidae) BAKER, M. R. 1980. Revision of Entomelas Travassos, arboreal? A morphological analysis. Southwestern 1930 (Nematoda: Rhabdiasidae) with a review of Naturalist 41:359–364. genera in the family. Systematic Parasitology 1:83– RICHARDSON, B. J., P. R. BAVERSTOCK, AND M. AD- 90. AMS. 1986. Allozyme Electrophoresis. A Handbook . 1982. Systematics and zoogeography of three for Animal Systematic and Population Studies. Ac- new nematode parasites of the frog Breviceps ademic Press, New York, New York, U.S.A. sylvestris sylvestris FitzSimons from South Africa. SCHMIDT Canadian Journal of Zoology 60:3134–3142. , W. M. 1991. Extensioń del a´rea de distri- bucioń sur para el lagarto Barisia rudicollis (Sauria: CAMPBELL,J.A.,AND D. R. FROST. 1993. Anguid lizards of the genus Abronia: revisionary notes, de- Anguidae). Boletıń de la Sociedad Herpetolo´gica scriptions of four new species, a phylogenetic anal- Mexicana 4:41–42. ysis, and key. Bulletin of the American Museum of SMITH, H. M., AND E. H. TAYLOR. 1950. An anno- Natural History 216:1–121. tated checklist and key to the reptiles of Mexico, CONTRERAS-MCBEATH,T.,AND F. U RBINA-TORRES. exclusive of the snakes. Bulletin of the United 1995. Historia natural del a´rea de proteccioń de States Natural Museum 199:1–253. flora y fauna silvestre Corredor Biolo´gico Chichi- STRESSEMAN, E. 1954. Ferdinand Deppe’s travels in nautzin. Centro de Investigaciones Biolo´gicas de la Mexico, 1824–1829. Condor 56:86–92. Universidad Autońoma de Morelos, Cuernavaca, TIHEN, J. A. 1949. A review of the lizard genus Ba- Morelos, Me´xico. risia. University of Kansas Science Bulletin 33:217– GOOD, D. A. 1988. Phylogenetic relationships among 256. gerrhonotine lizards, an analyisis of external mor- WADDICK,J.W.,AND H. M. SMITH. 1974. The sig- phology. University of California Publications in nificance of scale characters in evaluation of the Zoology 121:1–139. lizard genera Gerrhonotus, Elgaria, and Barisia. Gonza´lez-ROMERO, A., AND C. LO´ PEZ-GONZA´ LEZ. Great Basin Naturalist 34:257–266. 1990. Observations on a Barisia rudicollis Wieg- WIEGMANN, A. F. A. 1828. Beitrage zur Amphibien- mann (sic) (Sauria: Anguidae) with notes on its kunde. Isis von Oken 21:364–383. habitat. Bulletin of the Maryland Herpetological WIEGMANN, A. F. A. 1834. Herpetologia mexicana Society 26:159–168. seu descriptio amphibiorum Novae Hispaniae. Pars GRAY, J. E. 1838. Catalogue of the slender-tongued prima. Saurorum species. Lu¨ deritz, Berlin. saurians, with descriptions of many new genera and ZALDı´VAR-RIVERO´ N, A. 1998. Variacioń morfolo´gica y species (part 2). Annals and Magazine of Natural aloenzima´tica en la lagartija Barisia rudicollis History 1:388–394. (Squamata: Anguidae) y contribucioń a su historia GUILLETTE, J. L., JR., AND G. CASAS-ANDREU. 1987. natural. Bachelor’s Thesis, Universidad Nacional The reproductive biology of the high elevation Autońoma de Me´xico, Me´xico, D. F. Mexican lizard Barisia imbricata. Herpetologica ZALDı´VAR-RIVERO´ N, A., AND A. NIETO-MONTES DE 43:29–38. OCA. 2001. Notes on the natural history and dis- GUILLETTE, L. J., JR., AND H. M. SMITH. 1982. A tribution of the lizard Barisia rudicollis (Anguidae). review of the Mexican lizard Barisia imbricata, and Southwestern Naturalist 46:391–396. the description of a new subspecies. Transactions of the Kansas Academy of Science 85:13–33. Accepted: 7 November 2001 LEMOS-ESPINAL, J. A., R. G. WEBB,D.CHISZAR, AND Associate Editor: Joseph Mendelson 326 HERPETOLOGICA [Vol. 58, No. 3

APPENDIX torio Natural Las Joyas (MZFC 4714); 23 km from El Terrero, Manantlań (MZFC 6001); Cerro Grande, Specimens Examined El Terrero, near El Tapeixtle (MZFC 6002, 6019, Barisia herrerae (n ϭ 25): MEXICO: Mexico: Ap- 6773–74); El Tapeixtle, 3 km N El Terrero; El Ta- proximately 4 km E Ocuilan (approximately 15 km W peixtle (MZFC 8138–40); Mexico: Xalatlaco, El Ca- Cuernavaca City limits) on dirt road Cuernavaca- pulıń (MZFC 3242, 3384–85); Hacienda La Gavia Chalma, near Rancherıá Tlaltizapań, Municipality of (LACM 135555–56); Los Reyes Iztacala (MZFC Ocuilan (MZFC 5741, 9580, 9584–85, 9587–88, 3335, two specimens); Parque Nacional Zoquiapan 9592, 12549, 12551); 12 young born from MZFC (MZFC 3207, seven specimens); Parque Nacional 9587 and MZFC 12549 (MZFC 9593 and 12552– Zoquiapan, Can˜ ada del Quesero (MZFC 3352, two 562); Morelos: Cuernavaca City limits (MZFC specimens); NE extreme Llano Grande (MZFC 982, 12550); Huitzilac (MZFC 13929); Sierra del Chichi- two specimens); Popocate´petl (MZFC 3413); Valle de nautzin (MZFC 9589); Tetela del Volcań (EBUM Bravo (LACM 135552, 135554–55, 135559); vicinity 2485). of Valle de Bravo (LACM 136658–59); Morelos: Par- Barisia imbricata ciliaris (n ϭ 50): MEXICO: Chi- que Nacional Lagunas de Zempoala (MZFC 2333, huahua: Chihuahua (AMNH 1945); Coahuila:1miS 1195–96, 12565; IBH 1671, 3159, 3665); Oaxaca: Cedritos (AMNH 77246); 10.7 mi ESE San Antonio Pen˜ a Verde, Can˜ ada de Cuicatlań (MZFC 12545); de las Alazanas (LSUMZ 30154); Las Vigas (IBH Puebla: km 97 on Huauchinango-Apizaco road 4651, two specimens); Guanajuato:20kmNELeoń, (MZFC 12544); Santa Rita Tlahuapan (EBUAP 460); on road to San Felipe (MZFC 12067); Durango:El Colonia El Cerrito, Puebla City (EBUAP 463); Tlax- Salto (MZFC 12547–48); 5 mi ENE El Salto cala: Volcań La Malinche (MZFC 12564); San Tadeo (LSUMZ 30668); 10 mi E El Salto (AMNH 68357); Huexoyucan (MZFC 12563); Veracruz: on road be- 10 mi W El Salto (AMNH 98164); 56 mi W Durango tween Acayucan and Alvarado (MZFC 2309); Cerro (LACM 92674); Navajas (LACM 121928); Navajas, Azul (MZFC 2344, two specimens); 2 mi ESE Las ca. 40 mi W El Salto (LACM 121314); Hacienda Los Vigas (LSUMZ 28687, 31053); 1.5 mi S Puerto del Coyotes, 3 mi S El Salto (LACM 92675); 3 mi NW Aire (LSUMZ 10978); 4 km W Puerto Morelos Los Coyotes (LACM 136860); 10 mi S Rancho Santa (MZFC 727); 1–5 km W Xometla (LACM 131443, Ba´rbara (LACM 133639); 6 mi SE Llano Grande 131445–47); SE slope Pico de Orizaba (LACM (LACM 121927); Navios (AMNH 118078); Rancho 135557); Orizaba, Texmalaquilla (LACM 121929). Las Canoas, Municipality of Suchil, Reserva de la Barisia imbricata jonesi (n ϭ 13): MEXICO: Mi- Bio´sfera (IBH 2503); Rancho Temascal, Municipality choacań: Coalcomań (MZFC 11602); Coalcomań, of Suchil (IBH 7299); Reserva de la Michilıá, Mu- Cerro El Laurel (MZFC 12147, 11724); Coalcomań, nicipality of Suchil (IBH 7300–01); Vicinity of Palo El Polvorıń, 2 km S Dos Aguas (MZFC 11726–27); Gordo (CM 90195); Hidalgo: Molango (MZFC 495); N Dos Aguas (MZFC 11725); Coalcomań, 5 km S Nuevo Leoń: Ojo de Agua, near Galeana (FMNH Dos aguas (MZFC 11608–14). 30703); slope of Cerro Potosı´ (CM 43782); Cerro Po- Barisia imbricata planifrons (n ϭ 20): MEXICO: tosı´, Ojo de Agua (FMNH 30705–07); Pablillo Oaxaca: Sierra de Jua´rez, Yuvila (MZFC 12546); Cer- (MZFC 11237); Pablillo, near Galeana (FMNH ro San Felipe (AMNH 100713–4); near San Felipe 106168); 1.5 km E Mimbres (MZFC 7795–96); Que- (FMNH 112021); 2 mi S El Punto, NE Cerro San re´taro: Vicinity of Pinal de Amoles (MZFC 6875); Felipe, 7800 ft (AMNH 89638); 1.5 mi S El Punto, San Joaquıń (MZFC 7551); Municipality of Caderey- near Cerro San Felipe, 7200 ft (AMNH 89639); 3.5 ta, 1 km NE El Doctor (MZFC 8430); Municipality mi N Cumbre del Estudiante, on Ixtlań de Jua´rez- of Pinal de Amoles, 6 km NW Rancho Los Vela´zquez (MZFC 9252–53); Municipality of Coloń, Pinal de Oaxaca road, 7200 ft (AMNH 90928); SE of Cuaji- Zamorano (MZFC 9401–03, 9752–53); San Luis Po- moloyas, 8575 ft (AMNH 102717); vicinity of Cerro tosı´: Cerro Conejo, between Llano de Conejo and San Felipe [IBH 7429–31, 7431–32, 7433 (two spec- Llano de Garzas (LSUMZ 341); 2 mi E Catorce imens); field numbers JRS 53956–7, 4003 (IBH un- (LSUMZ 2373); 2 km W Alvarez, Municipality of catalogued)]; Sierra de Monteflor, partes altas de San Dıáz Zaragoza (MZFC 11082); 10 mi NE Bledos Juan Bautista Atlahuaca, Canãda de Cuicatlań (CM 41516); Zacatecas: Temascal, 8.2 mi SW Joaquıń (MZFC 8691–92). Amaro (LSUMZ 35070–01); Municipality of Valpa- Barisia levicollis (n ϭ 14): MEXICO: Chihuahua: raı´so, 20 km N (IBH 5066). north Chihuahua (AMNH 592–3); Chihuahua Barisia imbricata imbricata (n ϭ 74): MEXICO: (USNM 26602–3; 26612; 47067; 47413; 47452; Distrito Federal: Ejido San Nicola´s Totolapan, 2 km 46666–7); Samachique (FMNH 15728); Sierra Tara- W of Albergue El Pino, Delegacioń Magdalena Con- humara (FMNH 15729); Sierra del Nido (CM treras (MZFC 11663); Serranıá del Ajusco (MZFC 90194); 25.2 mi W (by road) Tomachic from la Junta, 282–84); Ajusco, Llano Cieneguillas (MZFC 279–80); 2200 m (LACM 132393). Ajusco, W of Valle Monte Alegre (MZFC 281); ap- Barisia rudicollis (n ϭ 21): MEXICO: Mexico: Ha- proximately 200 m E. P.C.V.U. (MZFC 4326); Hidal- cienda de la Gavia (IBH 1834, 2701); 6 km N Sul- go: Pachuca (MZFC 4696); 2 km W turnoff to Tian- tepec (MZFC 5358, 9590–91); Avańdaro, Valle de guistengo (MZFC 275–76); Atotonilco El Grande Bravo [EBUM 1372–73; IBH 6277 (two specimens); (FMNH 106156); Laguna del Tejocotal (MZFC 5091; MZFC 9581 (skull), 9582–3, 9586, 12541–43]; 3097); 2 km E El Tejocotal (MZFC 277); 4 km W El Michoacań: Municipality of Tuxpan, El Pinal (MZFC Tejocotal (MZFC 278); Jalisco: Manantlań, Labora- 9594); 5 km SE Zitaćuaro (MZFC 13435–37).