An Evolutionary Classification and Checklist of Amphibians and Reptiles on the Pacific Islands of Baja California, Mexico.
Resumen.--La herpetofauna de las islas del Pacifico de Baja California esta revisada basada en el concepto evolutivo. Este resulto en la relegacion de 10 subespecies al sinonimo a sus contrapartes peninsulares y in elevacion de tres subespecies al nivel de especie.
The Pacific coast of Baja California is fringed with a number of islands and small archipelagos. Most are young, relatively small, landbridge islands but others are older and continental or oceanic in origin (Grismer 1993). The herpetofauna of these islands as a whole has received little attention except for the publication of a number of checklists (Bostic 1975; Grismer 1993; Savage 1967; Van Denburgh 1905; Van Denburgh and Slevin 1914; Wilcox 1980; Zweifel 1958) and a biogeographical study of the northern islands (Wilcox 1980). This herpetofauna has never been evaluated taxonomically as a faunal unit.
Subsequent to the latest checklist of this herpetofauna (Grismer 1993), a number of additions (Wong 1997), deletions (Grismer and Hollingsworth 1996; Wong 1997), errors (Aguirre et al. 1999) and/or taxonomic changes (Grismer 1994a, 1999b; Grismer and McGuire 1996; Grismer and Hollingsworth in press; McGuire 1996; Smith et al. 1998; Wong and Grismer in prep.) have been published (Table 1). Additionally, this fauna has never been analyzed in the context of an evolutionary species concept (ESC) as has recently been done for the herpetofauna of the islands in the Gulf of California (Grismer 1999a,b). Grismer (1999b) noted that in the Gulf of California, classifications based on a biological species concept (Mayr 1969) had remained unstable for a number of years. This was because the recognition of insular populations as species or subspecies was largely founded on subjective interpretations of degrees of differentiation even though the data sets being interpreted had remained unchanged. Grismer (1999b) proposed to reduce this subjectivity by employing an ESC (Simpson 1961; Wiley 1978; Frost and Hillis 1990), in which only discretely diagnosable (i.e., no overlap in character state variation) lineages were recognized as species and the subspecies category was abandoned (see Grismer, 1999b for discussion). The subspecies category was not used because its operational duality (i.e., usage for both diagnosable insular populations and portions of continuous continental populations) often misrepresents history. The ESC proved most useful when evaluating insular populations because it divorced decisions of taxonomic rank from subjective interpretations of reproductive compatibility based on degree of differentiation. In instances where subspecies merely delimit the geographic distribution of selected character states within a continuous continental population, there is no evidence that the subspecies represent independent lineages and thus, should not be recognized in a formal taxonomy. This is because the organisms manifes ting these character states merely form a co-extensive integrating section of a continuously distributed species. Therefore, it is philosophically illogical and operationally counterproductive to consider arbitrarily defined sections of continuous populations and allopatric diagnosable entities as equivalent phenomenological systems even though both have been traditionally given the rank of subspecies.
It is proposed here that a classification based on the application of the ESC rather than the biological species concept will be more consistent with historical processes because it discourages the recognition of taxa that are not demonstrable evolutionary lineages. Thus, an evolutionary classification is less likely to be misleading to evolutionary biologists working to discover patterns, processes, and/or adaptations (Brooks and McLennan 1991). Such an interpretation of the Pacific island herpetofauna is warranted.
The following are discussions of those taxa whose reevaluation under the ESC resulted in changes in their taxonomy. Also treated are published distributional errors not corrected elsewhere in the literature. A summary of the proposed taxonornic changes presented herein and taxonomic changes published subsequent to Grismer (1993) are listed in Table 1. Table 2 lists the species and the islands on which they occur and Appendix I lists the islands and the species that inhabit them. The location of the islands is illustrated in Figure 1.
Bostic (1975) reported Batrachoseps major from Isla San Martin. The specimen was purported to be collected in the late 1970s by Tom Cozens, then a student at San Diego State University (Cozens, pers. comm., 1980). The disposition of the specimen remains unknown.
Rana pipiens complex
A single specimen of the Rana pipiens complex (Los Angeles County Museum [LACM] 91380) was collected from Isla Sur of Islas de Los Coronados on 20 June 1959. No other individuals have been reported. Because there is no permanent water on this island, this specimen is considered a single introduction. Interestingly, three tadpoles (LACM 91381) of the same complex were collected from Punta Banda south of Ensenada opposite Islas Todos Santos on 29 March 1952. No additional specimens from this locality have been reported. Perhaps, during the early fifties, an attempt was made to establish these frogs for commercial purposes.
In a genetic study of Urosaurus nigricaudus from Baja California, Aguirre et al. (1999) included what they believed to be a specimen U. nigricaudus from Isla de Cedros (voucher number 1861 of R. W. Murphy). This species, however, has never been reported from Isla de Cedros nor are there any museum records listing its presence despite the numerous collections made from this island in the last 45 years. It is also absent from the adjacent Vizcaino Peninsula of Baja California (Grismer et al. 1994) to which Isla de Cedros was most recently connected. This record is therefore considered erroneous. The specimen in question, if indeed from Isla de Cedros, is most likely Uta stansburiana. This may account for the some of the problems it posed within their data set (Aguirre et al. 1999).
Phyllodactylus xanti zweifeli
Dixon (1964) described Phyllodactylus xanti zweifeli from islas Magdalena and Santa Margarita on the basis of having a higher number of scales across the snout between the third supralabials (19-22, n = 18) as compared to peninsular P. xanti (14-21, n = 105) as well as having a lower number of scales across the venter (26-33, n = 18 vs. 31-41, n = 105). However, because the ranges of variation in these counts overlaps with those of peninsular populations, P. x. zweifeli is not discretely diagnosable and is recognized here as P. xanti.
Cnemidophorus tigris occurs on six islands along the Pacific coast of Baja California (Grismer 1993; Grismer and Hollingsworth 1996). Cope (1892) proposed the name C. multiscutatus for the whiptail lizards of Isla de Cedros. Van Denburgh (1894) proposed the name C. stejnegeri for populations ranging from Los Angeles, California, southward to near Laguna San Ignacio (see Smith 1946). Subsequent to their descriptions, these two species went through a multitude of taxonomic rearrangements, terminating with Burger (1950) who, without comment, placed both populations under the name C. t. multiscutatus. Zweifel (1958) noted that lizards from coastal southern California were very different in striping pattern and coloration from lizards of Isla de Cedros. He stated that it probably was not "proper to refer the two populations to the same subspecies." I have observed living individuals from islas de Cedros and Natividad and found them to be less distinctly striped than lizards from southern California populations bu t they do form a cline with populations from the adjacent Vizcaino Peninsula. This is most notable in the dark blotching of the gular and temporal regions where it is prominent in the Vizcaino Peninsula whiptails, less so in lizards from Isla Natividad, and even less so in lizards from Isla de Cedros. Clinal patterns have been observed in other reptiles through these regions as well (Grismer et al. 1994). Therefore, C. t. multiscutatus is not discretely diagnosable and is recognized here as C. tigris.
Walker (1981) proposed the name Cnemidophorus tigris vividus for the whiptail lizards of islas Norte and Sur of Islas de Los Coronados. He compared a series of lizards from Isla Sur to C. tigris of Isla de Cedros, a population from over 500 km to the south, rather than the presumable source population of C. tigris from the adjacent peninsula less than 10 km away. Grismer (1994b) noted that the Islas de Los Coronados populations and those from the adjacent mainland overlap in the number of granules around the body (80-100, n = 11 vs. 92-116, n = 15, respectively), granules between the occiput and rump (167-204, n = 11 vs. 192-231, n = 15), number of femoral pores (32-40, n = 11 vs. 36-43, a = 15), and the number of subdigital lamellae on the fourth toe (27-32, n = 11 vs. 29-32, n = 15). The striping pattern in lizards of the Islas de Los Coronados populations tends to be more vivid but it is not distinct from some individuals of adjacent peninsular populations. Therefore, the Islas de Los Coronados population s are considered here as C. tigris.
Fitch (1934) separated Elgaria multicarinata nana of islas Norte and Sur of Islas de Los Coronados from E. m. webbi of San Diego County, California by its smaller size, weak keeling, having fewer transverse ventral scale rows, and subtle differences in the head and body color pattern. However, only adult body size discretely differentiates these two populations, and according to Fitch (1934), "no overlapping size could be shown among the 49 adult specimens from Los Coronados Islands and 35 from San Diego County." Average SVL for E. m. nana was 93.3 mm with a maximum SVL of 114 mm and for E. m. webbi from San Diego County, average SVL was 135 mm with a maximum of 166 mm (Fitch 1938). Fitch (1938) was aware that body size differences could be the result of an ecological effect imposed by an insular environment rather than a genetically based difference as has been proposed by more recent authors (e.g. Petron and Case 1997 and citations therein). However, Fitch (1938) concluded that the body size differences ha ve a genetic basis, noting ontogenetic changes and sexual dimorphism in E. m. webbi-specifically, that the largest individuals have relatively thicker, more heavily keeled scales, and in males, much wider temporal regions. Fitch (1938) pointed out that the largest individuals of E. m. nana showed these characters which were indicative of their advanced age even though they were smaller than the small adults of E. m. webbi. Additionally, Zweifel (1952) observed a 79 mm SVL female E. m. nana copulating, and Burrage (1965) indicated that the hatchling size range of the Islas de Los Coronados population is 32-35 mm SVL, essentially the same as that of hatchling E. m. webbi (30-36 mm) from San Diego County, California. This strongly suggests that adulthood, and its accompanied morphological changes, is reached at a much smaller SVL in E. m. nana than in E. m. webbi. Based on this, E. m. nana is recognized here as a distinct species, E. nana.
Van Denburgh (1898) described Elgaria multicarinata ignava from Isla San Martin as an endemic subspecies characterized by its heavy keeling and darkened dorsal pattern. Fitch (1938), however, noted that an increase in the degree of keeling was a trend that developed in all of the southernmost populations of E. m. webbi and did not accord it taxonomic status. Its overall dark color pattern is likely a result of substrate matching, being that it inhabits dark, rocky volcanic habitats. A similar color pattern is found in populations from the volcanic areas of the adjacent peninsular coastline west of San Quintin. Therefore, there are no discretely diagnostic characteristics unique to this population and thus, it is recognized here as E. multicarinata.
Van Denburgh and Slevin (1923) proposed the name Diadophis punctatus anthonyi for the population on Isla Sur of Islas Todos Santos. This population was diagnosed as having a more robust body (Blanchard 1942) and an obscure neck ring with poorly defined edges (Van Denburgh and Slevin 1923). It is difficult to interpret from Blanchard (1942) how he determined that D. p. anthonyi was "larger and stouter-bodied" than adjacent populations of D. punctatus because all measurements he gave fall within the range of D. punctatus from the adjacent peninsula. Additionally, many specimens from northern Baja California and southern California have a poorly defined neck ring as well. Therefore, this population lacks discretely diagnostic characters and is considered as D. punctatus.
Zweifel (1958) described Hypsiglena torquata baueri from Isla de Cedros on the basis of a number of subtle character state trends of dorsal blotching, nuchal blotching, and the lateral striping on the head. In a review of these snakes, Tanner (1966), noting that these character states overlapped with those occurring in adjacent peninsular populations of H. Torquata, could not differentiate this subspecies. Based on this, H. t. baueri is recognized here as H. torquata.
Tanner and Banta (1962) described Hypsiglena torquata martinensis on the basis of a single female differentiated from all other Baja California populations in having 54 subcaudals, two loreal scales, and a dorsal scale row formula of 2321-19-17. Moquard (1899) demonstrated that central peninsular populations of H. torquata had 47-58 subcaudal scales and I have observed specimens from the vicinity of Bahia de Los Angeles with two loreal scales. The diagnostic value of the scale row formula is uninterpretable. Tanner and Banta (1962:24) state that "In martinensis, .. .the first fourth of the body has the increase to 23 rows and nearly half have 22 rows. This formula is more nearly the standard pattern for Hypsiglena, that is, the greatest number of rows from the nape to near mid-body and then one or more reductions before the vent." Therefore, these characters do not discretely diagnose H. t. martinensis and it is considered here a synonym of H. torquata.
Van Denburgh and Slevin (1923) described Lampropeltis zonata herrerae from Isla Sur of Islas Todos Santos. In a revision of L. zonata, Zweifel (1952) diagnosed L. z. herrerae from all other L. zonata by having a combination of no red in the color pattern and the posterior margin of the first white band being anterior to the angle of jaw. This population is often considered as lacking red bands (e.g., Rodriguez-Robles et al. 1999: Fig. 4). However, homologous bands do occur in the anterior portion of the body but are unique in that they have become white or sometimes brownish. Hayes (1975) demonstrated that L. z. herrerae had a significantly higher mean number of ventral scales (218.3, 216-220, n = 10) compared to L. zonata from the adjacent peninsula (210.4, 207-216, n = 6) and that their ranges only slightly overlapped. Additionally, he showed that the SVL of L. z. herrerae averaged over 50 mm larger than the SVLs of southern L. zonata. This character state is considered here to be unique because it is the result of an increase in the number and the overall size of the body vertebrae (Hayes 1975). Based on these characteristics, Hayes (1975) recommended that L. z. herrerae be elevated to species status. Using mtDNA, Rodriguez-Robles et al. (1999) demonstrated that L. z. herrerae is an exclusive population most closely related to individuals of L. zonata from the Sierra San Pedro Martir. Based on the above (excluding ventral scale counts), I elect to follow Hayes (1975) and recognize L. z. herrerae as a full species L. herrerae.
Klauber (1946) described Pituophis catenifer coronalis from Isla Sur of Isla de Los Coronados on the basis of four specimens. He diagnosed this population as usually having suboculars, frequent abnormalities in the head plates, and fewer number of dorsal blotches. Additionally, he detailed subtle differences from adjacent continental populations of P. catenifer in the color pattern of the head. Klauber (1946) also noted, however, that none of these characters discretely separated P. c. coronalis from many other subspecies of P. catenifer and therefore, this population is recognized here as P. catenifer.
Klauber (1946) described and differentiated Pituophis catenifer fuliginatus from Isla San Martin from peninsular P. catenifer on the basis of having a darker color pattern, large amounts of black spotting on the head, a lower average number of body blotches (62.6, 55-70, n = 15 vs. 75.5, 57-90, n = 52), usually a single instead of two preoculars, a high frequency of aberrant prefrontals, and paired dark subcaudal streaks. Klauber (1946) noted that the overall darkness of the population was probably related to substrate matching and I have observed similarly colored individuals from the volcanic regions west of San Quintin in coastal areas adjacent to Isla San Martin. None of the scale characters discretely separates P. c. fuliginatus from adjacent peninsular populations and subcaudal streaking also appears in other P. catenifer from Baja California (Klauber 1946). Therefore, this population is recognized here as P. catenifer.
Klauber (1949) described Crotalus viridis caliginis from Isla Sur of Islas de Los Coronados. Although he could find no differences in color pattern or squamation between it and the adjacent peninsular populations of C. viridis, he did note significant differences in body proportions and SVL. Klauber (1949) noted that C. v. caliginis would not grow to the size of an average adult C. v. helleri of the adjacent peninsula. The largest specimen he measured was 674 mm total length and he notes that C. v. helleri regularly reach 1100 mm in length. Also, the rattles of C. v. caliginis reach parallelism at a width of 10-11 mm whereas in C. v. helleri, parallelism is reached at 15-17 mm (parallelism happens when adulthood is reached). Finally, specimens of C. v. caliginis of the same size as C. v. helleri have disproportionally smaller heads (Klauber, 1938). Klauber (1949) noted that a 1100 mm C. v. helleri "would have 5 times the bulk" of a 650 mm C. v. caligin is. Based on these data, C. v. caliginis is considered h ere to be a distinct species, C. caliginis.
For comments on the manuscript and/or discussions of species concepts I thank B. Hollingsworth, H. Wong, and E. Mellink. I wish to thank K. Beaman for assistance with Rana records of the Los Angeles County Museum.
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Accepted for publication 1 March 2000
Summary of proposed taxonomic changes used herein. Referenced changes are those that occurred subsequent to Grismer (1993). Taxonomy of Grismer (1993) Newly proposed taxonomy Caudata Plethodontidae Batrachoseps pacificus major B. major (Wake and Jockusch 2000) Squamata (Lizards) Crotaphytidae Gambelia wislizenii copei G. copei (McGuire 1996) Phrynosomatidae Sceloporus magister monserratensis S. zosteromus (Grismer and McGuire 1996) Uta stellata U. stansburiana (Grismer 1999a; Upton and Murphy 1997) Gekkonidae Phyllodactylus xanti zweifeli P. xanti Teiidae Cnemidophorus tigris multiscutatus C. tigris Cnemidophorus tigiris vividus C. tigris Anguidae Elgaria paucicarinata cedrosensis E. cedrosensis (Grismer and Hollingsworth in prep.) Elgaria multicarinata ignava E. multicarinata Elgaria multicarinata nana E. nana Squamata (Snakes) Colubridae Chilomeniscus cinctus C. stramineus (Wong and Grismer in prep.) Diadophis punctatus anthonyi D. punctatus Hypsiglena torquata baueri H. torquata Hypsiglena torquata martinesis H. torquata Lampropeltis zonata herrerae L. herrerae Masticophis flagellum fuliginosus M. fuliginosus (Grismer 1994a) Pituophis melanoleucus coronalis P. catenifer Pituophis melanoleucus fuliginatus P. catenifer Pituophis vertebralis insulanus P. insulanus (Grismer 1994a) Crotalus viridis caliginis C. caliginis Crotalus exsul exsul C. ruber (Smith et al. 1998) Checklist of the herpetofauna of the Pacific islands of Baja California, Mexico, Asterisks following taxa indicate insular endemics. Taxon Insular locale CAUDATA Plethodontidae Aneides lugubris Los Coronados (Isla Norte) Batrachoseps major Los Coronados (all islands), Isla San Martin, Todos Santos (Isla Sur) ANURA Bufonidae Bufo punctatus Santa Margarita Hylidae Hyla regilla Cedros SQUAMATA (Lizards) Crotaphytidae Gambelia copeii Cedros, Magdalena, Santa Margarita Iguanidae Dipsosaurus dorsalis Magdalena, Santa Margarita Phrynosomatidae Callisaurus draconoides Santa Margarita Phrynosoma coronatum Cedros Sceloporus occidentalis Cedros, Todos Santos (Isla Sur) Sceloporus zosteromus Cedros, Magdalena, Santa Margarita Urosaurus nigricaudus Magdalena, Santa Margarita Uta stansburiana Asuncion, Cedros, Las Brozas, Las Piedras, Los Coronados (all islands), Magdalena, Natividad, Pata, San Benito, San Geronimo, San Martin, San Roque, Santa Margarita, Todos Santos (Isla Sur) Eublepharidae Coleonyx variegatus Cedros, Santa Margarita Gekkonidae Phyllodactylus xanti Magdalena, Santa Margarita Teiidae Cnemidophorus hyperythrus Magdalena, Santa Margarita Cnemidophorus tigris Cedros, Los Coronados (islas Norte and Sur), Magdalena, Natividad, Santa Margarita Scincidae Eumeces skiltonianus Los Coronados (islas Norte and Sur), Todos Santos (islas Norte and Sur) and Sur) Anguidae Anniella geronimensis San Geronimo Anniella pulchra Los Coronados (islas Norte and Sur), Todos Santos (Isla Sur) Elgaria cedrosensis Cedros Elgaria multicarinata San Martin Elgaria [nana.sup.*] Los Coronados (islas Norte and Sur) SQUAMATA (Snakes) Leptotyphlopidae Leptotyphlops humilis Cedros Pythonidae Lichanura trivirgata Cedros, Natividad Colubridae Chilomeniscus stramineus Cedros, Magdalena Diadophis punctatus Todos Santos (Isla Sur), San Martin Eridiphas slevini Santa Margarita Hypsiglena torquata Cedros, Los Coronados (all islands), San Martin Lampropeltis [herrerae.sup.*] Todos Santos (Isla Sur) Masticophis fuliginosus Magdalena, Santa Margarita Pituophis catenifer Los Coronados (Isla Sur), San Martin Pituophis [insulanus.sup.*] Cedros Pituophis vertebralis Magdalena, Santa Margarita Salvadora hexalepis San Geronimo, Todos Santos (islas Norte and Sur) Viperidae Crotalus [caliginis.sup.*] Los Coronados (Isla Sur) Crotalus enyo Magdalena, Santa Margarita Crotalus mitchellii Santa Margarita Crotalus ruber Cedros, Santa Margarita
Checklists of the herpetofauna of the islands along the Pacific coast of Baja California, Mexico. Asterisked taxa are insular endemics.
Isla Asuncion. Uta stansburiana.
Isla de Cedros. Hyla regilla, Gambelia copei, Phrynosoma coronatum, Sceloporus zosteromus, Sceloporus occidentalis, Uta stansburiana, Coleonyx variegatus, Cnemidophorus tigris, Elgaria [cedrosensis.sup.*], Leptotyphlops humilis, Lichanura trivirgata, Chilomeniscus stramenius, Hypsiglena torquata, pituophis [insulanus.sup.*], Crotalus ruber.
Isla Las Brozas. (Within Laguna Ojo de Liebre). Uta stansburiana.
Isla Las Piedras. (Within Laguna Ojo de Liebre). Uta stansburiana.
Isla Magdalcna. Dipsosaurus dorsalis, Gambelia copei, Sceloporus zosteromus, Urosaurus nigricaudus, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus hyperythrus, Cnemidophorus tigris, Chilominescus stramenius, Masticophis fuliginosus, Pituophis vertebralis, Crotalus enyo.
Isla Natividad. Uta stansburiana, Cnemidophorus tigris, Lichanura trivirgata.
Isla Pata. (Within Laguna Ojo de Liebre). Uta stansburiana.
Isla San Geronimo. Uta stansburiana, Anniella geronimensis, Salvadora hexalepis.
Isla San Martin. Batrachoseps major, Uta stansburiana, Elgaria multicarinata, Diadophis punctatus, Hypsiglena torquata, Pituophis catenifer.
Isla San Roque. Uta stansburiana.
Isla Santa Margarita. Bufo punctatus, Callisaurus draconoides, Dipsosaurus dorsalis, Gambelia copei, Sceloporus zosteromus, Urosaurus nigricaudus, Uta stansburiana, Coleonyx variegatus, Phyllodactylus xanti, Cnemidophorus hyperythrus, Cnemidophorus tigris, Eridiphas slevini, Masticophis fuliginosus, Pituophis vertebralis. Crotalus enyo, Crotalus mitchellii, Crotalus ruber.
(1.) Islas de Los Coronados. (Three major islands). Aneides lugubris (Isla Norte), Batrachoseps major, Uta stansburiana, Eumeces skiltonianus (islas Norte and Sur), Cnemidophorus tigris (islas Norte and Sur), Elgaria [nana.sup.*] (islas Norte and Sur), Anniella pulchra (islas Norte and Sur), Hypsiglena torquata, Pituophis catenifer (Isla Sur), Crotalus [caliginis.sup.*] (Isla Sur).
(2.) Islas San Benito. (Three islands). Uta stansburiana.
(3.) Islas Todos Santos. (Two islands). Batrachoseps major, Sceloporus occidentalis, Uta stansburiana (islas Norte and Sur), Eumeces skiltonianus (islas Norte and Sur), Anniella pulchra, Diadophis punctatus, Lampropeltis [herrerae.sup.*], Salvadora hexalepis.
(1.) Includes all three major islands (islas Sur, Norte, Medio) unless noted otherwise. Isla Medio consists of two small islets: a larger southern islet and a smaller northern islet. No records are known from the latter.
(2.) Includes all three islands.
(3.) Presence on Isla Sur only unless indicated
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|Author:||Grismer, L. Lee|
|Publication:||Bulletin (Southern California Academy of Sciences)|
|Date:||Apr 1, 2001|
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