Above: A male Mexican Emerald Spiny Lizard (Sceloporus formosus) from El Pasclar (elev. 2,300 m) in the state Guerrero. Below: A Black- bellied Gartersnake (Thamnophis melanogaster) from Lagunas de Zempoala in the state of Morelos, Mexico. During the mid-1800’s, Auguste Ghiesbreght collected numerous zoological specimens in Mexico, of which many are deposited in the Paris Natural History Museum. In the following article, the authors provide information on the identities of many of these specimens, as well as ' the distribution and relevant taxonomic issues for several of the species. © J. Peter Heimes 8844 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles www.mesoamericanherpetology.com www.eaglemountainpublishing.com Identity, origin, and distribution of Auguste Ghiesbreght’s Mexican amphibians and reptiles Beat Schätti1 and chriStoph KucharzewSKi2 1Apartado postal 9, San Pedro Pochutla, Oaxaca 70900, Mexico. E-mail: [email protected] (Corresponding author) 2Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstr. 43, 10115 Berlin, Germany. E-mail: [email protected] aBStract: We examine and revise the distribution and identities of the herpetofaunal species collected in Mexico by Auguste Ghiesbreght between 1841 and 1853, and discuss the areas he explored during this period. The specimens, deposited in the Paris Natural History Museum (MNHN), were obtained in the mountainous Huatusco–Xalapa–Cerro Cofre de Perote region of central Veracruz (1841–1845); near Tlacolula in the coastal plain of northern Veracruz (1841, Pituophis catenifer); along the periphery of Mexico City (1853); and possibly in the Sierra Madre del Sur in the vicinity of Chilpancingo in central Guerrero (1853, Sceloporus formosus). We review the geographic distribution of these species and include their vertical limits, especially those of Laemanctus serratus (maximum elevations in Mexico), S. formo- sus, and Thamnophis melanogaster. The type material of seven taxa collected by Ghiesbreght contains one specimen of Leptophis mexicanus Duméril, Bibron and Duméril 1854, perhaps the lectotype. We clarify the provenance of the original series of Conophis lineatus (Duméril, Bibron and Duméril 1854), and re- strict the type locality of Coryphodon oaxaca Jan 1863 (Coluber constrictor Linnaeus 1758) to Mata de Indio, Municipio de Totutla, in central Veracruz. Key Words: Central Highlands, elevations, endemism, Guerrero, “Oaxaca”, taxonomy, type specimens, Veracruz reSumen: Se examina y revisa la distribución e identidad de las especies herpetológicas mexicanas re- colectadas por Auguste Ghiesbreght entre 1841 y 1853, así como las áreas recorridas por el explorador durante este período. Los especímenes, depositados en el Museo de Historia Natural de Paris (MNHN), fueron hallados entre Huatusco, Xalapa y la zona del Cerro Cofre de Perote en la región montañosa de Veracruz central (1841 a 1845); en los alrededores de Tlacolula en la franja costera del norte de Veracruz (1841, Pituophis catenifer); en las afueras de la Ciudad de México (1853); y posiblemente en la Sierra Madre del Sur cerca de Chilpancingo en Guerrero central (1853, Sceloporus formosus). Repasamos la distribución de estas especies e incluimos sus límites verticales, especialmente los de Laemanctus serratus (elevación máxima en México), S. formosus y Thamnophis melanogaster. Los especímenes tipo de siete taxones incluyen uno de Leptophis mexicanus Duméril, Bibron and Duméril 1854, quizás el lectotipo. Se esclarece la procedencia del material original de Conophis lineatus (Duméril, Bibron and Duméril 1854) y restringimos la localidad tipo de Coryphodon oaxaca Jan 1863 (Coluber constrictor Linnaeus 1758) a Mata de Indio, Municipio de Totutla, Veracruz central. Palabras Claves: Altiplano Central, elevaciones, endemismo, especímenes tipo, Guerrero, “Oaxaca”, taxonomía, Veracruz Mesoamerican Herpetology 85 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles Citation: Schätti, B., and C. Kucharzewski. 2018. Identity, origin, and distribution of Auguste Ghiesbreght’s Mexican amphibians and reptiles. Mesoamerican Herpetology 5: 85–110. Copyright: Schätti and Kucharzewski, 2018. This work is licensed under a Creative Commons Attribution- NoDerivatives 4.0 International License. Received: 2 December 2017; Accepted: 12 March 2018; Published: 12 April 2018. INTRODUCTION Boniface Augustin Lucien, known as Auguste Ghiesbreght (1812–1893), first came to Mexico from 1838 to 1840 as a member of a Belgian mission, accompanied by Jean Linden and Nicolas Funck. In total he spent more than 50 years in Mexico, and until the mid-1850s collected a large number of natural history specimens. Numerous plants, gastropods, amphibians, reptiles, and bird species were described based on Ghiesbreght’s material, and he “was perhaps the botanist with the greatest knowledge of the flora of northern Mesoamerica during the first half of the nineteenth century” (Ossenbach, 2007: 186). Nonetheless, large periods of his life in Mexico, until his death at San Cristóbal de las Casas (Chiapas) in March of 1893, remain undocumented (Rovirosa, 1889; Silvestre, 2014) and we are unaware of an existing portrait or photograph of this remarkable individual. To earn a living, the “plant hunter” shipped botanical material, especially cacti and orchids, which later were sold by professional horticulturists in the Benelux countries. Passionate European collectors such as the banker Benjamin Delessert and the equally wealthy Hugh Cuming also purchased dry specimens, including snail shells. In time, most of these private collections were incorporated into public institutions, such as the former British Museum and the Geneva Botanical Garden. The bulk of Ghiesbreght’s material, however, is deposited in the Paris Natural History Museum, which during the winter of 1840–41 commissioned the well-prepared Belgian before his depar- ture overseas to procure scientific materials. The exact origin of many specimens received from Ghiesbreght, including type series, often was vague and in many cases became controversial or speculative. For example, McVaugh (1972) found that numerous plants labeled “Oaxaca” actually hailed from localities in other Mexican states, and after examining all of the available evidence Schätti et al. (In Press) concluded that Ghiesbreght did not collect any botanical or zoological material from the state of Oaxaca. In this study we address a series of highly interesting specimens of amphibians and reptiles (“un certain nombre de Reptiles fort intéressants”; Duméril et al., 1854a: 238) acquired from “M. Ghuisbreght” between 1842 and 1854, which constitute nearly all of the specimens that remain of his herpetological legacy. Recently, Schätti et al. (In Press) reported on two shipments from “Oaxaca” (1842, 1845), but no serious efforts have been made to clarify the actual origins of the “Mexique” shipments (1844, 1854), including certain type material. In an attempt to elucidate these matters, we investigated the travels of Ghiesbreght during the corresponding years, and compare our results to the verified distributional ranges of the properly identified species. PRESENTATION AND REFERENCES The information we present in this study primarily involves Ghiesbreght’s itineraries and collections within a well-defined area of central Mexico, located between latitudes ca. 17°N along the Pacific in the vicinity of Acapulco and ca. 21°N near Tlacolula on the coastal plain of northern Veracruz (Fig. 1). We used reliable distributional data, including vertical ranges, for all the identified species. For the sake of brevity, but also for completeness, we provide additional information at the end of this paper to help the reader understand certain complex issues or controversies (Notes 1–14). These remarks include distributional information (e.g., elevations), obscure and dubious literature records, specific extralimital data, as well as explanations relating to institutions or collection, such as “in-house” details and taxonomic intricacies. After carefully analyzing these topics, we compiled an extensive bibliography; herein, however, we only provide the significant references. Where necessary, we consulted the original sources for the regional distribution of species in the Central and Southern Highlands and adjacent areas. We investigated the entire geographic range of the Mexican Black-bellied Gartersnake (Thamnophis melanogaster) from the northern Mesoamerican Herpetology 86 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles portion of the Sierra Madre Occidental to the Transverse Volcanic Belt. In the case of this species and the Mexican Emerald Spiny Lizard (Sceloporus formosus), we incorporate reliable data for preserved specimens in institutional collections accessible at <www.idigbio.org/portal>. To avoid confusion, throughout the text we distinguish references to the original description of taxa by omitting a comma between the author name(s) and the year of publication, and use the comma in standard literature citations. MATERIAL AND TERMINOLOGY Fifty-one specimens of amphibians and reptiles from unspecified localities in “Mexique,” or erroneously from the state of “Oaxaca” (see Introduction), were obtained by Auguste Ghiesbreght and arrived in four shipments; the specimens were recorded in the Catalogue of Acquisitions no 41 of the Lower Vertebrate section of the Muséum National d’Histoire Naturelle (MNHN) between August of 1842 and April of 1854 (p. 47: 1842; p. 71: 1844; p. 91: 1845, 1854). At least 22 valid species, viz., two anurans, three salamanders, eight lizards, and nine snakes were included in these shipments. While searching for the Ghiesbreght material in the MNHN collections, we encountered 26 specimens (Table 1: nine amphibians, nine lizards, and eight snakes; Schätti et al., In Press), including the type material of Leptophis mexicanus and Pituophis mexicanus, which cannot be unambiguously identified (see Types and Taxonomy). Four species of Sceloporus from the 1854 shipment might be present among the MNHN-RA 2520–22 and/or 2525–26 series, which lack collecting data, as well as any reference to Ghiesbreght (Ivan Ineich, in litt., December 2016). We could not locate a specimen of Sceloporus torquatus (MNHN-RA 6045) and 20 specimens lacked verified accession numbers, including eight Ambystoma mexicanum, one unknown bufonid (Bufo sp.), one unknown teiid (Aspidoscelis sp.), two Barisia imbricata, one Ctenosaura acanthura, one Sceloporus sp. incertae sedis (1844 ship- ment), and six unidentified conspecific natricids (Thamnophis sp.). Some missing vouchers might be misplaced, but others probably have been lost or were destroyed. [Note 1] Aquiloeurycea cephalica auct. is a species complex (Parra-Olea et al., 2010; Frost, 2017). We adhere to Zaldivar-Riverón et al. (2005) or Bryson and Riddle (2012) in treating the Barisia imbricata-ciliaris complex (including B. i. ciliaris Smith 1942c) as conspecific, and refer to the nominotypical subspecies auct. (south of latitude ca. 21°N, Table 2, see Note 2) as B. imbricata, excluding B. [i.] jonesi Guillette and Smith 1982 from the Sierra de Coalcomán in southwestern Michoacán. The species boundaries within the Sceloporus grammicus group are obscure (e.g., Auth et al., 2000; Lara-Góngora, 2004; Hammerson et al., 2007). The systematics of certain Mexican representatives of the milksnake genus Lampropeltis Fitzinger 1843 is far from clear, and the allocation of a “Coronella spec. nov.” recorded in 1842 (p. 47, entry no 3) as L. cf. polyzona is explained elsewhere (Schätti and Stutz, 2016: nota 20; Schätti et al., In Press; see, e.g., Woolrich-Piña et al., 2017). We indicate the complete authorship (including the year, see Presentation and References) of the species-group names, except for Barisia imbricata (see above), in Table 1, Notes 2 and 9 (Sceloporus formosus and Thamnophis melanogaster), or under Types and Taxonomy (synonyms and names of nomenclatural relevance). We compiled the distributional data and the pertinent references used to elaborate the ranges and locality records (in Fig. 2) or elevations (in text) in Table 2 (also, see Appendix 1, and Notes 2 and 8–10). [Note 2] The Plateau denotes the Mexican Highlands, a region of vast elevated plains separated by mountain ridges and situated between the coastal cordilleras from the southern border of the United States southward to the Oaxaca Valley. The Central Plateau, or Central Highlands, encompasses the area from the Lower Río Grande de Santiago in Nayarit (Aguamilpa area) eastward through the southern portions of Zacatecas and San Luis Potosí, and southward roughly along latitude 19°N, i.e., the section of the Transverse Volcanic Belt that extends from the Colima volcanoes to the Pico de Orizaba range beyond the southern confines of the Sierra Madre Oriental. For the purposes of this study, the Lower Huasteca is the region with elevations up to approximately 1,500 m within the Río Moctezuma (Pánuco) drainage in northern Hidalgo, northeastern Querétaro, and southeastern San Luis Potosí. In the context of vertical distribution, we regard “moderate” as elevations between 1,000 and 1,500 m, “high” as elevations from 1,500 to 3,500 m, and “alpine” as elevations above 3,500 m. “Gulf” stands for the Gulf of Mexico, and “Isthmus” for the Isthmus of Tehuantepec. We obtained all of our geodetic positions from <www.geonames.org> and indicate them in sexagesimal format. In certain cases, we verified the elevational data by using 1:50,000 topographical maps issued by the Instituto Nacional de Estadística, Geografía e Informática (INEGI). Mesoamerican Herpetology 87 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles ween 1841 and 1853 (recorded as 1842–1854) in Mexico (“Mexique”) including ober (status or verbatim entry in Catalogue of Acquisitions n 41, if appropriate), is CEarly Citation(s) Pertaining to Ghiesbreght Specimens and omments Brocchi (1883: 106): Amblystoma [sic] mexicanum (coll. “Ghussbreght“) Brocchi (1883: 112, plate 20.2): “Mexique par M. Ghuisbreght” Dunn (1926: fig. 60, map): “Oaxaca,” see Note 14 Unidentified toad (Bufo sp.) Duméril et al. (1854c: 93, plate 104): Bolitoglossa Mexicana [sic] Nob. (“province d’Oaxaca au Mexique, par M. Ghuisbreght”), see Plate 2 Brocchi (1881: 30): Hyla Baudini [sic] Duméril and Duméril (1851: 143): “G.[errhonotus] […] Lichenigerus [sic] Wagler” o 1854 shipment: Gerrhonotus imbricatus (Cat. entry n6) Aspidoscelis sp. other than A. sexlineata (Linnaeus 1766), see Note 13 Duméril and Duméril (1851: 64): “C.[yclura] […] Acanthura [sic] Gray” Duméril and Duméril (1851: 55): L.[æmanctus] […] Longipes [sic] Wiegmann 1834 see Note 1 Duméril and Duméril (1851: 77): “Tr.[opidolepis] […] Microlepidotus [sic] Dum. Bib.” o 1854 shipment (Cat. entry n4), see Note 1 Specimen unlocated Duméril and Duméril (1851: 77): “Tr.[opidolepis] […] Variabilis” [sic] “Tropidolepis undulatus” (Bosc and Daudin in Sonnini and Latreille 1801), see Material and Terminology incl. Notes 1 (1854 shipment) and 13 Duméril et al. (1854a: 183): Coryphodon Constrictor [sic] Nobis (“Oaxaca”); Jan (1863: 63): C.[oryphodon] oaxaca (“Messico”); see Types and Taxonomy, Conclusions Duméril et al. (1854b: 936, plate 73): Tomodon lineatum Nobis (“Mexique”); Jan and Sordelli (1866: plate 6.3, “Musée de Paris”); see Types and Taxonomy, Note 4, and Plate 1 Duméril et al. (1854a: 621): Coronella doliata “Holbrook” (Linn. 1766) [nomen suppr.] Duméril et al. (1854a: 536): Leptophis mexicanus Nobis (“Mexique”) see Types and Taxonomy, Note 5 Duméril et al. (1854a: 236, plate 62): Pituophis mexicanus Nobis (“Mexique […] Ghuisbreght”), see Types and Taxonomy incl. Notes 6–7, Conclusions Duméril et al. (1854a: 464, 468): Petalognathus nebulatus (Linnaeus 1758) “Variété D” Bocourt (1883: 584, plate 36.11): Homalocranion Deppii [sic]Lectotype designation as H. “deppei” [sic] (Wilson and Meyer, 1981) Lectotype of Tropidonotus mesomelanus Jan see Note 3 Duméril et al. (1854a: 585): Tropidonotus saurita (Linnaeus 1758)Unidentified gartersnake species (Thamnophis Fitzinger 1843) m ght beton nu Date 1854 1854 1854 1842 1842 1844 1854 1854 1844 1845 1854 1842 1854 1854 1842 1844 1854 1842 1845 1842 1845 1842 1842 1854 1854 1842 MNHN amphibians and reptiles collected by Auguste GhiesbreTable 1. “Oaxaca” (taxon preceded by an asterisk). The scientific name and accessifollowed by the year of registration, and remarks. Identity Ambystoma mexicanum (Shaw and Nodder 1798): MNHN-RA 4774–75 aAquiloeurycea cephalica (Cope 1865): MNHN-RA 6396 ( holotype of tadSpelerpes sulcatum Brocchi 1883)uaC“Bufo americanus” (catalogue name): ? ,aru* Pseudoeurycea gadovii (Dunn 1926): MNHN-RA 4749 (5 ex.)nA * Smilisca baudinii (Duméril and Bibron 1841): MNHN-RA 4799 Barisia imbricata (Wiegmann 1828): ? (2 ex., one per shipment) “Cnemidophorus sex-lineatus” (catalogue name): ? Ctenosaura acanthura (Shaw 1802): ? * Laemanctus serratus Cope 1864: MNHN-RA 2094aSceloporus formosus Wiegmann 1834: ?irua(*) Sceloporus grammicus Wiegmann 1828: MNHN-RA 3152 (6 ex.) and ? S (2 ex.) Sceloporus torquatus Wiegmann 1828: MNHN-RA 6045 * Sceloporus variabilis Wiegmann 1834: MNHN-RA 3056 (2 ex.) Sceloporus sp. incertae sedis: ? (2 ex.) * Coluber constrictor (Linnaeus 1758): MNHN-RA 7378 ( holotype) * Conophis lineatus (Duméril, Bibron and Duméril 1854b): MNHN-RA 3740 (syntype) * Lampropeltis cf. polyzona Cope 1860: MNHN-RA 0419 * Leptophis mexicanus Duméril, Bibron and Duméril 1854a: MNHN-RA 3453 or 3455 ( lectotype or paralectotype)set* Pituophis catenifer (Blainville 1835): ? MNHN-RA 3188 ( syntype of nepP. mexicanus Duméril, Bibron and Duméril 1854a)reS* Sibon dimidiatus (Günther 1872): MNHN-RA 7297 Tantilla deppii (Bocourt 1883): MNHN-RA 0054 ( lectotype) Thamnophis melanogaster (Peters 1864): MNHN-RA 7321 (paralectotype of Tropidonotus mesomelanus Jan 1863) * “Tropidonotus saurita” (catalogue name): ? (6 ex. incl. skull) Mesoamerican Herpetology 88 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles 104°W ZACATECAS 102°W 100°W S 98°W 96°W NAYARIT I Volcán de TeqRuíiola Grande de Santiago GUANAJUATO QUERÉTARO Río TMaocntegzuoMmjaóetztitEláRnRA ZMacuRíao Altltaippexcáon TlacoluHluaitzilac GULF OF MEXICO 20°N JALISCOLago de Chapala C E N T R A L P L A T E A HIUDALGORío AmajacRíoA TulDancRingEo O VERACRUZ 20°N Ciudad Guzmán ArúMmobraelrioa Ciudad Río Ler Tepeji del Río RIE Las Vigas de R. ToxtlPauceubalyoa Viejo VolcánT de FueRgo MICHOACÁN Hidalgo maMexico City TLAXCALANTA C. Cofre de PeroteXalapa COLIMAColima A N ASpatzingVán E R S E VNevaEdOSoT dAeD TOoL DluEc aMÉHCXuICitOzilaAc PoNpocatéIpetl CPueb la B LPaic MoE daeli nOLcrhiLzeabaETl MiradorHSuaant Juosscéo TVenereajacrpuaz Potrero Viejo Coalcomán S MORELOS PUEBLA Acultzingo Ciudad Mendoza I Tehuacán E Iguala R 18°N R 18°N A Cuicatlán GUERRERO M Chilpancingo OAXACA VALLEY Sra A MEXICO D Acahuizotla Tlaxiaco Cerro Zempoaltepec MIXE R Oaxaca de Juárez E Acapulco OAXACA D Cerro Piedra Larga E PACIFIC OCEAN L S U R 16°N 16°N 200 km Puerto Escondido 104°W 102°W 100°W 98°W 96°W Fig. 1. Localities and geomorphological features indicated in the text (also, see Fig. 2). The solid symbols denote places visited by Auguste Ghiesbreght (including potential localities in Guerrero). The gray area specifies the region in central Veracruz, the southern portion of the Sierra Madre Oriental, and the Pico de Orizaba range explored by the collector until 1844 or 1845. Map courtesy of Andrea Stutz. Hereafter, we use “AG” for Auguste Ghiesbreght, and “Cat.” refers to the Catalogues of Acquisitions no 41 (see above) and 42 (Conophis lineatus, see Types and Taxonomy including Note 4) in the former Herpetology and Ichthyology Department of the Paris Museum, which covers the period from 1839 to 1864. In addition to MNHN, other institutional acronyms are as follows: CAS (California Academy of Sciences, San Francisco); ENCB (Escuela Nacional de Ciencias Biologicas, Instituto Politécnico Nacional, Mexico City); IBUNAM (Instituto de Biología, Universidad Autónoma de México [UNAM, Colección Nacional de Anfibios y Reptiles CNAR], Mexico City); LVT (Las Vegas Tissue Collection); MCZ (Museum of Comparative Zoology, Harvard University, Cambridge); MZFC (Museo de Zoología, Facultad de Ciencias, UNAM, Mexico City); NMW (Naturhistorisches Museum, Wien); RMNH (Nationaal Natuurhistorisch Museum, Leiden, former Rijks Museum voor Natuurlijke Historie); UCM (Museum of Natural History, University of Colorado, Boulder); UIMNH (University of Illinois Museum of Natural History, integrated into Illinois Natural History Survey [INHS] collections, Champaign); USNM (National Museum of Natural History [Smithsonian Institution] or United States National Museum, Washington); and UTEP (The University of Texas at El Paso Biodiversity Collections). Mesoamerican Herpetology 89 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles RESULTS Types and Taxonomy We located at least six out of the seven name-bearing types described from the material collected by AG and de- posited in the MNHN collection (Table 1, see Pituophis mexicanus). Four recognized snake species are as follows: Coryphodon [Coluber constrictor Linnaeus] oaxaca Jan 1863 ( holotype), Leptophis mexicanus Duméril, Bibron and Duméril 1854 ( , see below), Homalocranion [Tantilla] deppii Bocourt 1883 ( lectotype), and Tomodon [Conophis] lineatum Duméril, Bibron and Duméril 1854 (syntype). Spelerpes sulcatum Brocchi 1883 ( holotype) is a junior synonym of the plethodontid salamander Aquiloeurycea cephalica (Cope 1865). Tropidonotus mesomela- nus Jan 1863 (paralectotype) actually is a subjective senior synonym of the gartersnake Thamnophis melanogaster (Peters 1864). [Note 3] The holotype (MNHN-RA 7378) of Coryphodon oaxaca Jan, originally reported from the homonymic Mexican state (Table 1), was collected in 1841 or early 1842 and recorded as “Psammophis ?” (Cat. 41, entry no 7, 1842), the only taxon within the shipment that agrees with the general aspects of the Nearctic racer genus Coluber Linnaeus 1758 (type species C. constrictor). Contemporaneous examples for the use of Psammophis auct. for Coluber L. are Holbrook’s (1842) account of P. flagelliformis (Daudin 1803) or P. flavigularis Hallowell 1852 (Coluber [Masticophis auct.] flagellum Shaw 1802). Smith and Taylor (1950: 337) accepted the type locality of “Coluber oaxaca” in the state of Oaxaca (“no definite locality”), as indicated by Duméril et al. (1854a), because of “a reasonable possibility [that] the type […] may have been secured” there (op. cit.: 314; see Table 1, Distribution, Discussion, and Conclusions). Based on Smith and Taylor’s (1945) assumption of the existence of a single original specimen (the “Type”), Wellman (1963) designated MNHN-RA 3738 as the holotype of Conophis lineatus (Duméril, Bibron and Duméril). This species was described from at least two vouchers (precise number unknown), of which Wallach et al. (2014) assigned lectotype status on MNHN-RA 3738. Their provenance (coll. “A. Ghiesbreght & F. Schlumberger”) relies on Bocourt (1876, 1886), who supposedly had three specimens from these collectors at hand. The single “Schlumberger” specimen (MNHN-RA 3738 or 3739), however, was incorporated into the collection in March of 1859 (Cat. 42, p. 92, entry no 8), five years after the species was described (see Plate 1). Wellman’s (1963) taxo- nomic act was implemented without having seen the specimen(s) or crediting the Paris curator (Jean Guibé), who provided the data, but his designation might be incorrect. Consequently, we refer to MNHN-RA 3740 (coll. AG) as a syntype of Tomodon lineatum Dum., B. and D. (Table 1). [Note 4] Smith and Taylor (1950) restricted the type locality of Tomodon lineatum Duméril, Bibron and Duméril (“du Mexique”) to the city of Veracruz, a decision that perhaps should be revisited in view of the probable origin of AG’s “Oaxaca” syntype (MNHN-RA 3740, Cat. 41, entry no 3, 1845), i.e., from the same general area in upland central Veracruz as nearly all of the herpetofaunal specimens gathered by this collector until 1845 (Fig. 1, see Discussion, Conclusions, and Note 8). Leptophis mexicanus Duméril, Bibron and Duméril was described upon MNHN-RA 3453 and 3455 from “Mexique.” Both specimens lack further collecting data and were received through the museum’s “[a]dministra- tion.” AG’s “Dendrophis ?” in the 1845 shipment, originally recorded from “Oaxaca” (Cat. 41, entry no 2), undoubt- edly is one of the former syntypes. Smith and Taylor (1950: 350, as Thalerophis m. mexicanus) restricted the type locality of L. mexicanus Dum., B. and D. to Potrero Viejo (Amatlán de los Reyes Municipality, Fig. 1) in Veracruz, and Mertens (1973) selected MNHN-RA 3455 as the lectotype. His ventral and subcaudal counts for this female (167 and 146, respectively) and the paralectotype (MNHN-RA 3453, : 169, 163) obviously differ from the data reported in Duméril et al. (1854a: 537), i.e., 157–169 and 138–154, respectively. [Note 5] The description of Pituophis mexicanus Duméril, Bibron and Duméril is based on various specimens (pre- cise number unknown) from unspecified localities and received from several unnamed travelers (“envoyée du Mexique par plusieurs voyageurs”) including “Ghuisbreght” (Duméril et al., 1854a). His syntype arrived at the Paris collection in 1842 as “Lycodon ?” (Cat. 41, entry no 4), and Schätti et al. (In Press) associated the specimen with MNHN-RA 3188, but only circumstantial evidence links this “Anasime méxicain” (Duméril et al., 1854a) to AG. [Note 6] Mesoamerican Herpetology 90 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles Plate 1. Conophis lineatus (Duméril, Bibron and Duméril 1854). An illustration of Tomodon lineatum ( ) accompanying the original description (Duméril et al., 1854c: plate 73), which perhaps shows Auguste Ghiesbreght’s specimen (MNHN-RA 3740). Bocourt’s (1886: plate 38.5) reproduction of a Conophis lineatus might rely on a different syntype. A “T.[omodon] lineatus” [sic] (“Mexique. – Musée de Paris”) in Jan and Sordelli (1866: plate 6.3) with eight instead of seven supralabials probably depicts MNHN-RA 3738 or 3739, which was received from Henri D. Schlumberger in 1859. Pituophis mexicanus Duméril, Bibron and Duméril 1854 is a junior synonym of Coluber sayi Schlegel 1837 (Boulenger, 1894; Stull, 1940), i.e., the Nearctic C. [Pituophis] catenifer Blainville 1835 (e.g., Bryson et al., 2011; Wallach et al., 2014; see Note 6). R.[hinechis] mexicanus Duméril 1853 is a nomen nudum, and P. mexicanus Dum., B. and D. has not been used as a valid name after 1899, and thus qualifies for a nomen oblitum. Smith and Taylor (1950) restricted the type locality of this taxon to Sabinas Hidalgo (26°30'12"N, 100°10'54"W) in northern Nuevo León. The type series of Pituophis mexicanus (see above, including Note 6), for example, shows 229–239 ventrals, 9 supralabials (5th in contact with orbit), and usually 4, sometimes (“parfois”) 2, and once 3 prefrontals (Duméril et al., 1854a: 237). This condition (right plate divided) is found in MNHN-RA 3188, which measures slightly over 2 m in total length (snout–vent length = 1,780 mm, tail length = 221 mm), and probably is the largest syntype mentioned Mesoamerican Herpetology 91 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles (“Un sujet qui a 2m,064 de long”) and the only one housed in the Paris Museum today. This male specimen, however, contains a greater number of ventrals (241 plus 2 preventrals) than indicated in the original description of this taxon. [Note 7] DISTRIBUTION Based on their geographic and elevational ranges, 19 verified species of amphibians (three salamanders, one hylid frog) and reptiles (seven lizards, eight snakes; Table 2) attributed to AG can be classified as follows: Conophis linea- tus (vertical range disputed); Leptophis mexicanus (maximum elevation ca. 1,600 m, McCranie, 2011); Sceloporus variabilis (ca. 2,500 m); Sibon dimidiatus (ca. 1,600 m; op. cit.); and the tropical lowland Smilisca baudinii. All of these are wide-ranging species found from near sea level (nsl) to moderate or higher elevations (if data sources not specified, see Table 2). [Note 8] Coluber constrictor is distributed throughout the southern Nearctic Region into Mesoamerica. This species has been recorded at an elevation close to 2,000 m (Sierra del Carmen, Coahuila) on the Northern Mexican Plateau, and it occurs along the western Gulf lowlands (nsl to at least 1,000 m) through Chiapas (few records) to Guatemala (Petén). Lampropeltis cf. polyzona possibly shows a similar southern distributional limit, and might be a largely Mexican or northwestern Mesoamerican taxon. The southwestern Nearctic Pituophis catenifer, found at elevations that approach 2,900 m (Wallach et al., 2014), ranges into Mexico all along the border with the United States (Baja California, the northern Pacific coast, the Northern Plateau southward to Nayarit [fide Heimes, 2016], and the west- ern Gulf area). Populations in the vicinity of the city of Veracruz (nsl) are considered introduced (Pérez-Higareda, 1981). The southernmost reported collecting sites for native populations are in northern Veracruz (Bryson et al., 2011: fig. 2), likely as far south as ca. latitude 21°17'N (Soriano-Arista, 2007: table 1, “P. melanoleucus sayi”; see Conclusions and Note 6). The Mesoamerican Laemanctus serratus (nsl to ca. 1,100 m, see Note 8) shows a disjunct distribution. The Gulf population (southern Tamaulipas to central Veracruz, including Lower Huasteca) is separated from those in the Grijalva drainage (Central Depression of Chiapas) to the border with Guatemala, on the Yucatan Peninsula, includ- ing extreme northern Belize (Corozal District), and in Honduras (Wilson and Johnson, 2010). Three amphibian and seven reptile species (coll. AG) are Mexican endemics. Among these, Ctenosaura acan- thura is the only one that inhabits low elevations. This species is known to occur from sea level to approximately 1,000 m (Johnson et al., 2010; see Note 8) in the western Gulf region, along the Grijalva drainage to the border region and into Guatemala, and through the Isthmus to the Pacific coast (Tehuantepec Plain). Sceloporus formo- sus (elev. ca. 1,100–2,900 m; max. elev. 3,450 m fide Flores-Villela et al., 2010) is recorded from the Xalapa area (Veracruz; re-restricted type locality, Smith and Pérez-Higareda, 1992; formerly Acultzingo, Smith and Taylor, 1950) and the vicinity of Tepeyahualco (Puebla, MZFC 7817; www.idigbio.org) along the southernmost portion of the Sierra Madre Oriental. This pine-oak forest inhabitant also occurs in western Oaxaca (possibly approaching extreme southern Puebla) southeastward to the Cerro Piedra Larga area (Townsend-Peterson et al., 2004), which is located beyond the confines of the Mexican Highlands. The distribution of this species extends to the Pacific versant of the Sierra Madre del Sur, and supposedly includes allopatric populations (see Note 2) in central Guerrero. [Note 9] Sceloporus grammicus (elev. ca. 1,600–4,100 m, see Note 8) occurs on the Central and Southern Plateau and the Sierra Madre del Sur, whereas S. torquatus (elev. ca. 1,500–3,000 m fide Hall, 1973; max. elev. 3,200 m fide Flores-Villela et al., 2010) has not been reported south of the Transverse Volcanic Belt (Fig. 2). Thamnophis melanogaster is known to occur on the northern edge of the Sierra Madre Occidental in Sonora along the northwestern Chihuahua border, on the northern part of the Sierra Tarahumara (elev. ca. 1,000–2,250 m), and is found in disjunct populations on the Plateau from the Durango–Chihuahua border to near the limits of the Central Highlands roughly west of Tlaxcala, at elevations up to about 3,000 m (Fig. 2A). [Note 10] The Imbricate Alligator Lizard, Barisia imbricata (Fig. 2B), usually occurs at elevations from 2,100 m to, approximately 4,000 m, but occasionally is found in habitats below 1,500 m (see Material and Terminology, Note 2). The distribution of this species extends from north of Volcán de Tequila in northwestern Jalisco (B. i. imbricata sensu Bryson and Riddle, 2012; including Cerro La Magdalena area fide Guillette and Smith, 1982: 31, fig. 4) along the Transverse Volcanic Belt and to northern Hidalgo, into adjacent Veracruz (Sierra Madre Oriental), and through the Southern Highlands eastward to the Zempoaltepec range that separates the Oaxaca Valley from the Isthmus. Mesoamerican Herpetology 92 April 2018 | Volume 5 | Number 1 Schätti and Kucharzewski Auguste Ghiesbreght’s Mexican amphibians and reptiles operly identified species of Mexican amphibians and reptiles (coll. AG 1841–1853) deposited in the MNHN, including four lizards (preceded by an pecimens (see Material and Terminology, Discussion, and Notes 2 and 9–10). Figures referenced in this paper are in .bold Distribution and Comments Lake Chalco (Mexico City–México border, disappeared), Xochimilco canals (Mexico City, on brink of extinction, Contreras et al., 2009): Fig. 2B E state of México, México City, border area in Morelos and Tlaxcala, northern Puebla, adjacent Veracruz, Hidalgo (excl. W), N Querétaro, S Tamaulipas (Dixon et al., 1972; Vega-López and Álvarez, 1992; Parra-Olea et al., 1999, 2010; Uribe-Peña et al., 1999; Gillingwater and Patrikeev, 2004; Gutiérrez-Mayén and Salazar-Arenas, 2006; Farr et al., 2009; Dixon and Lemos-Espinal, 2010; Ramírez-Bautista et al., 2009, 2010, 2014; Fernández-Badillo and Goyenechea-Mayer-G, 2010; Huitzil-Mendoza and Goyenechea-Mayer-G., 2011; Badillo-Saldaña et al., 2015; Lemos-Espinal and Dixon, 2016; Woolrich-Piña et al., 2017: table 19): (see Note 2)Fig. 2A Michoacán through S Querétaro to N Hidalgo and montane central Veracruz, southeast to México–Guerrero–Morelos border Colima, W Jalisco and Ntriangle and Puebla (except SW) to Mt. Zempoaltepec (Tihen, 1949; Guillette and Smith, 1982; Auth et al., 2000; Castro-Franco and Zagal, 2003; Navarro-López et al., 2003; Zaldivar-Riverón et al., 2005; Reyes-Velasco et al., 2009; Bryson and Riddle, 2012): (see Note 2)Fig. 2B S Nearctic, Atlantic NW Mesoamerica (S Canada to Guatemala); Coahuila and Durango southeast along Gulf to Chiapas (absent from Oaxaca or Yucatan Peninsula in Mexico) and through Belize into Guatemala (Wilson, 1966, 1978) Mesoamerica (S Mexico to Costa Rica); S Tamaulipas to Yucatan Peninsula, Chiapas, Central Isthmus, and possibly Tehuantepec Plain (Wallach et al., 2014: Jalisco, Querétaro or Hidalgo records in error; W Isthmus fide Mata-Silva et al., 2015: table 4) S Tamaulipas along Gulf (Veracruz, inland to SE San Luís Potosí, NE Hidalgo, extreme SE Puebla) through Chiapas (Central Depression) into Guatemalan border area (Stephen, 2011), Central Isthmus and Tehuantepec Plain (Mata-Silva et al., 2015: table 4) Atlantic Mesoamerica (absent from most of Belize and Guatemala; south to Honduras, Solís et al., 2014); S Tamaulipas to central Veracruz (incl. Lower Huasteca or adjacent Puebla), Chiapas (Central Depression) incl. Guatemalan borderland, Yucatan Peninsula (McCranie and Köhler, 2004) Distribution unclear due to disputed species concepts, taxon may occur in Guatemala, and possibly in contiguous Mesoamerica Mesoamerica (E and S Mexico to Costa Rica); Nuevo León, S Tamaulipas and along Gulf (incl. Lower Huasteca, Puebla) to Yucatan Peninsula, Isthmus, Chiapas (Camarillo-R., 1995; Dixon and Lemos-Espinal, 2010; Lemos-Espinal and Dixon, 2013; Nevárez-de los Reyes et al., 2017) SW Nearctic (incl. SW Canada, W United States); Baja California, south to Sinaloa and Zacatecas, N Veracruz (Bryson et al., 2011: fig. 2; see text) La Malinche, Pico de Orizaba range, and Mount Cofre de Perote area (Solano-Zavaleta et al., 2009; Parra-Olea et al., 2010) Montane central Veracruz and adjacent Puebla; Oaxaca Highlands to Mt. Zempoaltepec area and Pacific versant of Sierra Madre del Sur incl. east- central Guerrero (Smith, 1939, 1984; Davis and Dixon, 1961; Flores-Villela et al., 1991; Canseco-Márquez and Gutiérrez-Mayen, 2005, 2010; Auth et al., 2000; Pérez-Ramos and Saldaña-de la Riva, 2008; Ramírez-Bautista and Pavón, 2009; García-Mares, 2013): (see Note 9)Fig. 2A Jalisco to Hidalgo and Veracruz, through Morelos to central Oaxaca and along Sierra Madre del Sur into Guerrero: (see Note 2)Fig. 2B W Jalisco to E Durango, S Nuevo León and SW Tamaulipas, south to N Morelos and Puebla, N central Veracruz west of Toxtlacuaya (Smith, 1936; Olson, 1990; Vega-López and Álvarez, 1992; Ramírez-Baut. et al., 2010; Lemos-Espinal and Dixon, 2013; Magno-Benítez et al., 2016): Fig. 2A Mesoamerica (S Texas to NW Costa Rica); along Gulf through Isthmus, absent from Yucatan Peninsula (Mather and Sites, 1985) S Mexico and Central America; N Veracruz along Gulf (absent from Tabasco and Yucatan Peninsula in Mexico) and Chiapas (Kofron, 1990) Mexico (incl. extreme S Texas) to Panama; Pacific coast and along Gulf to Yucatan Peninsula (Duellman, 2001: incl. fig. 287) Wilson and Mata-Silva, 2014): Guerrero (Río El Saltre), México (Chalma), N Morelos, W Oaxaca Highlands (Camarillo-Rangel, 1983;Fig. 2B Upper Río Bavispe (NE Sonora) to N Sierra Tarahumara (W Chihuahua) and NW Durango to E state of México: (see Note 10)Fig. 2A Distribution of 19 prTable 2. asterisk) not documented by s Taxon Ambystoma mexicanum Aquiloeurycea cephalica * Barisia imbricata Coluber constrictor Conophis lineatus * Ctenosaura acanthura Laemanctus serratus Lampropeltis cf. polyzona Leptophis mexicanus Pituophis catenifer Pseudoeurycea gadovii * Sceloporus formosus Sceloporus grammicus * Sceloporus torquatus Sceloporus variabilis Sibon dimidiatus Smilisca baudinii Tantilla deppii Thamnophis melanogaster Mesoamerican Herpetology 93 April 2018 | Volume 5 | Number 1