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Home , Hemidactylus, Hemidactylus angulatus, Lepidodactylus, Lepidodactylus lugubris

Check List 8(1): 164-167, 2012 © 2012 Check List and Authors Chec List ISSN 1809-127X (available at www.checklist.org.br) Journal of lists and distribution

n New records of the mourning lugubris (Duméril and Bibron, 1836) (: ) from

istributio D

* raphic Juan D. Daza , Scott L. Travers and Aaron M. Bauer g eo

G Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085, USA. n

o * Corresponding author. E-mail: [email protected]

otes N Abstract: The mourning gecko () is a nocturnal, parthenogenetic species that has been introduced in Colombia. Despite more than 70 years of collecting activity in the country, there has yet to be a thorough evaluation of its distribution in Colombia. Here we review all records from eight museum collections, along with literature reports, to generate a locality map documenting this gecko’s distribution in space and time. Additionally, new sightings are reported and its range expansion in the Cauca rift valley is discussed.

The mourning gecko (Lepidodactylus lugubris, Figure in order to better understand its current distribution and 1) is widely distributed throughout much of the Indo- its pattern of colonization. For example, it is known that

its presence in the Cauca rift valley is based on sporadic this species is more abundant in the Pacific region, while ,Pacific region , (Bauer Costa 1994), Rica, while , in the Colombia, New World , it has records, and it has not been recorded in the Yotoco forest, andbeen Chileintroduced (Smith in andPacific Grant coastal 1961; areas Schauenberg of the mainland 1968; of a locality on the east slope of the western range of the Henderson et al. 1976; Kraus, 2009; Uetz 2011) as well as Andes, despite extensive herpetological survey work over et al. 2009) and Easter a 30-year period (see Castro-Herrera et al. 2007). Island (Garman 1908). More recently, it has also been recordedthe Galapagos in Atlantic (Wright and 1983; Jadin coastal areas from Panama, Suriname and Guadeloupe (Bauer et al. 2007; Lorvelec et al. 2011). In Colombia this species is most thecommon southwestern in the Pacific departments region, of throughout Valle del Cauca, the lowlands Cauca, Nariño,and Pacific and versant on Gorgona of the western Island. Additionally,range of the thereAndes, are in intermittent records documenting it in the city of Santiago de Cali (Ayala 1986; Castro-Herrera and Vargas-Salinas 2008). Despite over 70 years of collecting activity in the region, the distribution of this species in Colombia remains unclear. In a recent herpetofaunal inventory (Castro- Herrera and Vargas-Salinas 2008) it was recorded in the the Andes (Cordillera Occidental), and the Cauca rift valley (ValleValle del interandino Cauca in the del Pacific río Cauca). region, Boththe western L. lugubris range and of “ brookii H. brookii” are actually assignable to H. leightoni or H. haitianus, members of the H. angulatus” (all or most group New that World have “ probably been introduced from mainland [Powell and Maxey 1990; Powell et al. 2007], whereas true H. brookii is restricted to tropical 1996; Rivas 2002; Weiss and Hedges Figure 1. Lepidodactylus lugubris Asia [Bauer et al. 2010; Rösler and Glaw, 2010]) have San Cipriano, Valle del Cauca (photograph by Richard Thomas). been hypothesized to have reach Colombia via cargo ships (UVC 14948). Specimen collected in during the Spanish colonial period in the Americas from

Although today there is a general idea of the dissemination dataWe to reviewedgenerate a the new literature distribution and map collection (Figure data 2) using from of1492 this to species, 1898 a(Castro-Herrera revision of all available and Vargas-Salinas records is required 2008). ArcGISeight zoological 10.0 (ESRI, collections Redlands, (Appendix CA, USA) that1). Werepresents used these the

164 Daza et al. | New records of Lepidodactylus lugubris from Colombia distribution of the species in space and time. Our map L. lugubris is thought to have had a sporadic presence in unpublished records from the Caribbean and Andean Only recently does it appear to have become established regionsincludes 24 from localities, Colombia, including and recently new sightings. published Although records, inCali this since area. the Until time this it point, was firstit may collected have been at thisinhibited locality. by the larger, more dominant “”, which those where this information was not available we have is a more conspicuous species in this area and has been approximatedmany records include by mapping specific the geographic records incoordinates, the smallest for suggested to have niche overlap with L. lugubris (F. Castro geographic unit that is consistent with the collection data. personal communication). Localities on the map are color-coded in 20-year intervals Based on the data gathered, we suspect that colonizing based on the date specimens were collected in order L. lugubris to facilitate visualization of their temporal distribution after the Spanish colonial period (early 20th century). It is within Colombia since the earliest record to the present first arrived in the Colombian Pacific region time. The earliest record of L. lugubris in Colombia dates Canal,almost iscertain responsible that the for voluminous its introduction traffic ofin shipsthe Americas crossing (Smiththe Pacific and , Grant especially 1961) and after not the European opening of cargothe Panama ships C. Murphy in Tumaco (Nariño, AMNH R-63567) during crossing the Atlantic and Caribbean. This introduction in theto 1941, Askoy based Expedition on a single to thespecimen areas collected between by the Robert Gulf the Americas is also likely to be a consequence of repeated of Panamá and Guayaquil (Ecuador). In Colombia this colonization events from different sources at multiple expedition covered the pacific shoreline, Malpelo and localities along the Pacific shoreline. The populations inferredGorgona its(Murphy presence 1941). based In the on hisliterature, review this of specimensspecies is citiesare likely are locatedto have (e.g., first Buenaventura, established in Tumaco). Colombia After along these the fromfirst mentionedPanamá and from Ecuador. Colombia The oldest by Mechler recorded (1968), specimen who initialsouthwestern populations Pacific were coast, established, where several L. important lugubris likely port from the Cauca rift valley was collected in Santiago de Cali by Vernon Lee (USNM 151576) in 1963. Today only four lowland region, including Ecuador (Schauenberg 1968). records are known from Santiago de Cali. According to Somebecame widespread subsequently and dominant moved throughout eastward the to Pacific higher the dates of collection and personal observations (JDD), elevation localities in the Cauca rift valley (between 900

Figure 2. Localities of Lepidodactylus lugubris in Colombia.

165 Daza et al. | New records of Lepidodactylus lugubris from Colombia and 1000 m). Since Santiago de Cali is separated from foraging areas and daytime retreat sites with Hemidactylus geckos, which may prey upon the smaller mourning gecko m), the invasion of this zone was probably facilitated by (Petren et al. 1993; Brown et al. 2002). This behavior may terrestrialthe Pacific transport lowlands of by foreign higher and elevations local freight (above such 2000 as be a limiting factor for the expansion and establishment lumber through two main roads, Buenaventura-Cali and of the Colombian populations. A more complete sampling Buenaventura-Buga. In the Cauca rift valley localized across the country is required in order to provide a better populations went from numerically rare to dominant over understanding of the distribution of this non-native a period of at least 50 years, a process that appears to have species in Colombian territory. Also, continued monitoring of Lepidodactylus lugubris and Hemidactylus aff. angulatus (= H. brookii in literature records) may help to understand L.accelerated lugubris arein the parthenogenetic last five years. and reproduce clonally the interactions between these two invasive species. Like conspecifics in other parts of the world, Colombian identity of all Colombian populations, but specimens (Orozco-Cardona 1996). We have not confirmed the clonal Acknowledgments: from Valle de Cauca (Fig. 1) appear to belong to the chief for providing additional assistance in gathering the locality records of Lepidodactylus lugubris We would like to thank the following persons (1999) and to be rather similar in dorsal pattern to the Bolivar-García (Universidad del Valle, Cali), Colin McCarthy (The Natural EasterCentral Island and South clone American (Ineich and clone Ota as 1993). identified Ineich by (1999) Ineich History Museum, London), for MarthaColombia: Lucía Fernando Calderón Castro-Herrera, (Instituto de Ciencias Wilmar postulated that L. lugubris has had a long presence in the Americas, as supported by its apparently endemic clone NaturalNaturales, History, Bogotá), New AddisonYork). Wynn (United States National Museum and broad distribution, but our data suggest that its spread of Natural History, Washington), David Kizirian (American Museum of has been relatively recent, at least in Colombia. Literature Cited Previous herpetological surveys did not report this Ayala, S.C. 1986. Saurios de Colombia: Lista actualizada, y distribución species in the Chocó Department (Castaño-Mora et al. Caldasia 15(71-75): 557- et al. 1993), but a recent 575. de ejemplares ColombianosDas Tierreich, en los Teilband museos. 109. Gekkonidae (Reptilia, study (García Rentería et al. 2006) recorded it in San José Sauria). Part 1. Australia and Oceania de2004; Purré Dixon near 1979; Atrato Ruiz-Carranza town, this being the northernmost Bauer,Gruyter. A.M. 306 1994. p. Bauer, A.M., T.R. Jackman, E. Greenbaum, and. T.J. Berlin: Papenfuss. Verlag 2007. Walter First de record of Lepidodactylus lugubris in Suriname. Applied Herpetology

(Cordobarecord for andthis Atlántico,species in ICNthe Pacific10399; region UVC 15627of Colombia.–15629), Bauer, A.M., T.R. Jackman, E. Greenbaum, A. de Silva, V.B. Giri and I. Das. whereWe this found species some was records unknown from or the was Caribbean not reported region in 2010.4(1): 84-85. Molecular evidence for the taxonomic status of Hemidactylus brookii group taxa (Squamata: Gekkonidae). The Herpetological et Journal 20(3): 129-138. al. 2007). Aside from a record from Suriname (Bauer et Brown, S.G., R. Lebrun, J. Yamasaki and D. Ishii-Thoene. 2002. Indirect al.a recent 2007), regional these are scientific the only survey reported (Carvajal-Cogollo occurrences of L. competition between a resident unisexual and an invading gecko. Behaviour 139(9): 1161-1173. lugubris from the South American mainland east of the Isthmus of Panama. Although there are only two records interference behavior among sexual and asexual geckos. from this region, this opens several questions: 1) when Bolger,Behavior D.T. and T.J. Case. 1992. Intra-specific and inter-specific was this species introduced in the Colombian Caribbean?; Urbina-Cardona. 44(1): 21-30. 2007. de áreas asociadas a humedales de 2) are these specimens derived from colonizing Carvajal-Cogollo,la planicie del J.E., Departamento O.V. Castaño-Mora, de Córdoba, G. Cárdenas-Arevalo Colombia. Caldasia and 29(2): J.N.

Castaño-Mora, OV, G. Cardenas-A, E.J. Hernandez-R and F. Castro-H. from established Caribbean populations from Central 427-438. America?individuals (e.g., from Panama, the Pacific Bocas region del Toro or didarea); they and disperse 3) are In J.O. Rangel (ed.). Colombia Diversidad Biótica. Volume IV El Chocó Biogeográfico2004. Catálogo. Bogotá: de reptiles Universidad en el Chocó Nacional biogeografico; de Colombia. p. 615-631

departamento del Valle del Cauca, Colombia. Biota Colombiana 9(2): theseAn populations unexpected recordcontinuous was onewith from the Boyacá,Pacific populations in a locality Castro-Herrera,251-277. F and F. Vargas-Salinas. 2008. Anfibios y reptiles en el onor disjunct?the eastern versant of the eastern range of the Andes . Guía de los anfibios y reptiles del Bosque de Yotoco, Valle del Cauca-Colombia. Cali: from an altitude of about 1000 m. Given the geographic Castro-Herrera,Imprenta Departamental F, B.G. Wilmar del and Valle M.I. delHerrera-Montes. Cauca. 70 p. 2007 barriers that separate this locality from the Colombian Dixon, J.L. 1979. Origin and distribution of reptiles in lowland tropical coastal regions, this specimen is very likely to have arrived In The South American herpetofauna: Its origin, evolution, and dispersal: by anthropogenic transportation. rainforestsMonograph ofof South the Museum America; of p.Natural 217-240 History W.E. Number Duellman 7. Lawrence: (ed.). The extent of the invasion of Colombia by L. lugubris and The University of Kansas. its ecological impact on the native biota, especially other Duméril, A.M.C. and G. Bibron. 1836. Érpetologie Générale, ou Histoire Naturelle Compléte des Reptiles. Paris: Libraire Encyclopedique de gekkotans, is still unclear. In Colombia the scarcity of native Roret. 518 p. nocturnal geckos opens up opportunities for the expansion García Rentería, U., J.T. Rengífo Mosquera, F. Moreno Mosquera and A.M. of L. lugubris. On the other hand, and Jiménez Ortega. 2006. Diversidad de lagartos (Squamata: Lacertilia) en el sotobosque de cuatro localidades asociadas a la cuenca del río the presence of more aggressive and larger Hemidactylus Cabí, Chocó-Colombia. Revista Institucional Universidad Tecnológica geckos can reduce its population density considerably del Chocó Garman, S. 1908. The Reptiles of Easter Island. Bulletin of the Museum of Comparative 2006(25): Zoology 47-55. 52(1): 3-13 XI pls. or indirect factors such as stress-induced decrease in reproductiveeither by direct output factors or such competition as predation for of illuminated juveniles, and asexual gecko species (Lepidodactylus) in French . foraging areas or daytime retreat sites (Bolger and Case Hanley,Evolutionary K.A., D.T. Bolger Ecology and T.J. Case. 1994. Comparative ecology of sexual Lepidodactylus lugubris 1992; Petren et al. 1993; Hanley et al. et (Reptilia: Gekkonidae). 8(4): A 438-454. recent addition to the herpetofauna of al. 2002). Lepidodactylus lugubris is reluctant to share Henderson,Nicaragua. R.W, Herpetological J. Villa and Review J.R. Dixon. 1976. 1994; Brown 7(4):173. 166 Daza et al. | New records of Lepidodactylus lugubris from Colombia

Ineich, I. 1999. Spatio-temporal analysis of the unisexual-bisexual Uetz, P. 2011. The database. Electronic database accessible at Lepidodactylus lugubris complex (Reptilia, Gekkonidae); p. 199-228 http://www.reptile-database.org/. Captured on 18 July 2011. In H. Ota (ed.). Tropical Island Herpetofauna: Origin, Current Diversity, and Conservation. Amsterdam: Elsevier. of the Antillean geckos Phyllodactylus wirshingi, Tarentola americana, Ineich, I. and H. Ota. 1993. Morphological variation and distribution of the Weiss,and A.J. Hemidactylusand S.B. Hedges. haitianus 2007. Molecular (Reptilia, phylogeny Squamata). and biogeography Molecular unisexual-bisexual complex of the gecko, Lepidodactylus lugubris, in Phylogenetics and Evolution French Polynesia and Easter Island. Bulletin of the College of Science, Gonatodes collaris in University of the Ryukyus 1993(56): 113-120. Galapagos Archipelago. Herpetological45(1): 409-416. Review Jadin, R.C., M.C. Altamirano, M.H. Yánez-Muñoz and E.N. Smith. 2009. Wright, J.W. 1983. The distribution and status of First record of the (Hemidactylus frenatus) in 14(1): 32. Ecuador. Applied Herpetology 2009(6): 193-195. Kraus, F. 2009. Alien reptiles and amphibians. A scientific compendium and : August 2011 analysis : January 2012 Leviton, A.E., R.H.G. Gibbs Jr, E. Heal and C.E. Dawson. 1985. Standards Received : February 2012 in herpetology. Springer and Verlag. ichthyology: XII, 564 Part p + CD-ROM.I. Standard symbolic codes for Accepted : Philippe J. R. Kok institutional resource collections in herpetology and ichthyology. Published online Copeia 1985(3): 802-832. Editorial responsibility Lorvelec, O., A. Levesque and A.M. Bauer. 2011. First record of the Appendix 1. Locality and collection date (DD/MM/YY) information for Mourning Gecko (Lepidodactylus lugubris) on Guadeloupe, French Colombian specimens of Lepidodactylus lugubris. Collection acronyms Herpetology Notes follow Leviton et al. Mechler, B. 1968. Les Geckonidés de la Colombie. Revue Suisse de Zoologie 75(10):West Indies. 203-371. 4: 291-294. CHOCÓ: : San (1985) José de and Purré Sabaj (March–October Pérez (2010). 2003, García Rentería et al. 2006). VALLE DEL CAUCA: Science ( Atrato Murphy,57-58. R.C. 1941. The Askoy Expedition of the American Museum of Buenaventura: Bajo Anchicayá: Orozco-Cardona,Natural History M. indel the S. 1996.Eastern Ciclo Tropical reproductivo Pacific. de Lepidodactylus 94 2429): BMNH 1980.373-374 (28/V/1976), ICN 3061–3062 (30/05/1976), lugubris (Sauria Gekkonidae). Unpublished B.Sc. thesis. Cali: ICN 3067, ICN 3100 –3104, ICN 3105–3106 (04/06/1976), ICN 3063 (05/06/1976), ICN 3064–3066, ICN 3068–3070 (08/06/1976), MCZ Petren, K., D.T. Bolger and T.J. Case. 1993. Mechanisms in the competitive R154445 (1/9/1977), MVZ 190572–190579 (20/7/77), KU 192645- successUniversidad of an del invading Valle. 46 sexual p. gecko over an asexual native. Science UVC192647, 5085–5086 ICN 3099 (27/9/85),(07/04/1978), UVC UVC 9736 6–7 (23/10/92);(27/5/1980), Cuartel UVC 264, B-IV-5 UVC 5054, UVC 5055, 5057–5067 (2/3/83), UVC 266–269 (12/4/1984), Powell, R. and S.A. Maxey. 1990. Hemidactylus brookii. Catalogue of American259(5093): Amphibians 354-358. and Reptiles Concesión Bajo Calima: UVC 11049-11050; Campamento Pulpa de Powell, R., R. Henderson, K. Adler and H. Dundee. 1996. An annotated Papel: UVC 11142 (10/9/1992); Desembocadura del Río Mayorquín: 1990(493): 1-3. (6/5/1993UVC 14649); (28/07/2001); Isla Palma: UVC 15631 (17/12/2009); Río In Contributions to West Indian Cajambre,(27/12/2006). Punta Bonita: UVC ICN 14647 3087 (5/7/2001); (18/08/1977). Queremal: ICN 7076 Herpetology:checklist of West A Tribute Indian to amphibians Albert Schwartz and reptiles;. Ithaca: p. Society 51-93, forpls 1-8 the Pianguita: UVC 15778; San Cipriano: UVC 14948–14953: StudyR. Powell of Amphibians and R.W. and Henderson Reptiles. (ed.). Calima: Guadalajara de Rivas, G. 2002. Geographic distribution. Hemidactylus leightoni. Buga: (23/12/2010, 3°54’5.00”N, 76°18’6.85”W), Santiago de Cali Herpetological Review USNM 151576 (13/02/1963); Barrio El Lido: ICN 3084 (27/03/1979), Rösler, H. and F. Glaw 2010. Morphologische Variation und Taxonomie Barrio Champagnat: UVC 270 (28/7/1984), CAUCA: UVC 9805 (14/11/92), von Hemidactylus brookii 33(2): 148. (20/7/1965),Barrio Nápoles AMNH (25/12/2010, R-109815-109820 3°23’17.22”N, (2/2/73). 76°32’56.76”W). Bonanza: Tuluá: Spixiana 33(1): (23/12/2010,ICN 3071–3075 4° (15/08/1976); 4’60.00”N, 76°12’0.00”W). ICN 652 FMNH(03/02/1961), 166248 139-160. Gray, 1845 und Guapi, Ruiz-Carranza,Hallowell, 1854 P.M., sowie J.I. Hernández-Cphänotypisch andähnlicher M.C. Taxa. Ardila-R. 1993. La ICN 3089–3098, (__/08/1978), Isla ICN Gorgona: 3087 (18/08/1977), ICN 051970 In (27/03/1979),ICN 3109, ICN 3085–3086ICN 3110 (20/05/1979);(06/04/1977), ICN Campamento 3088 (04/07/1978), del Penal P. Leyva-F. (ed.). Colombia Pacífico. Bogotá: Editorial FEN. herpetofauna de la Standardprovincia symbolic biogeográfica codes delfor institutionalChocó; p 257-269 resource collections in herpetology and ichthyology: an Online Reference. UVC 5088–5090 (1/5/82), UCV 5069-5070 (2/4/83), UVC 5071-: SabajVersion Pérez, 1.5M.H.. Electronic 2010. database accessible at http://www.asih.org/, ICN5081 3108 (21/4/83), (22/05/1979). UVC 5082-5084 NARIÑO: (12/7/83), UVC 265 (4/4/84), UVC (11/7/1965);7324-7325 (7/2/87), UVC 7812 ICN (13/1/89),3107 (16/10/1978). UVC 5078. La EsperanzaAMNH DC. Captured on 18 July 2011. R- AMNH R- 108308, FMNH 16249 Schauenberg,American Society P. 1968. of Ichthyologists Sur la presence and deHerpetologists, Lepidodactylus Washington, lugubris 3081, (25/11/1976),La Guayacana: ICN 3082 (29/11/1976); El Tangarial:Tumaco: ICN 3083 (Dumeril et Bibron, 1836) (Reptilia, Gekkonidae) en Equateur. Revue (08/12/1976).63567 (19/4/1941). BOYACÁ: El Mira: ICN 3076 km 2, (17/11/1976), carretera a Guateque: ICN 3077– ICN Suisse de Zoologie 5826 (26/01/1981). CORDOBA: Bahía Cispatá - Amaya, Smith, H.M. and C. Grant. 1961. The mourning gecko in the Americas. INVEMAR: ICN 10399 (21/09/2000).Santa María, ATLÁNTICO: : UVC Herpetologica 17(1): 75(2): 68. 415-417. 15627–15629 San Antero, Barranquilla

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