18 3 NOTES ON GEOGRAPHIC DISTRIBUTION Check List 18 (3): 483–491 https://doi.org/10.15560/18.3.483 Updating the distribution of Dicrodon guttulatum Duméril & Bibron, 1839 (Reptilia, Teiidae) with a disjunct population in the eastern slope of the Peruvian Andes Antonio García-Bravo1, 2, 3*, Betty K. Guzman1, Jani E. Mendoza1, Cristóbal Torres Guzmán1, Manuel Oliva1, Elgar Barboza1, 4, Jhon Quiñones Rámirez5, J. Luis Zabarburu-Veneros3, Pablo J. Venegas2, 6, 7 1 Instituto de Investigación para el Desarrollo Sustentable de Ceja de Selva), Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas, Chachapoyas, Amazonas, Peru • AGB: willy.garcia@untrm.edu.pe https://orcid.org/0000-0002-1607-0500 • BKG: bguzman@indes-ces.edu. pe https://orcid.org/0000-0002-0097-3289 • JEM: jmendoza@indes-ces.edu.pe https://orcid.org/0000-0001-8324-039X • CTG: cristobal. torres@untrm.edu.pe https://orcid.org/0000-0002-0932-7224 • MO: manuel.oliva@untrm.edu.pe https://orcid.org/0000-0002-9670-0970 • EB: ebarboza@indes-ces.edu.pe https://orcid.org/0000-0002-9628-8138 2 División de Herpetología, Centro de Ornitología y Biodiversidad, Lima, Peru 3 Escuela de Ingeniería Forestal, Facultad de Ingeniería y Ciencias Agrarias, Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas, Chachapoyas, Amazonas, Peru • JLZV: jaimeluiszabarburuveneros@gmail.com https://orcid.org/0000-0003-0679-0827 4 Dirección de Desarrollo Tecnológico Agrario, Instituto Nacional de Innovación Agraria, Lima, Peru 5 Gerencia de Desarrollo Olmos, Proyecto Especial Olmos Tinajones, Gobierno Regional de Lambayeque, Chiclayo, Lambayeque, Peru • JQR: jodaquira79@gmail.com 6 Rainforest Partnership, Austin, TX, USA • PJV: pablo@rainforestpartnership.org https://orcid.org/0000-0002-6501-4492 7 Instituto Peruano de Herpetología, Lima, Peru * Corresponding author Abstract We report a disjunct population of Dicrodon guttulatum Duméril & Bibron, 1839 on the eastern slope of the Cordillera Occidental in the inter-Andean Seasonally Dry Forests of the Marañón River, in the Departments of Cajamarca and Piura in northwestern Peru. We include an updated range distribution map using records from museum specimens, the Global Biodiversity Information Facility, and available photographic records on iNaturalist. In addition, we identify widespread cultivation of rice crops as the main threat to D. guttulatum in the inter-Andean Seasonally Dry Forests of the Marañón. Keywords Dry forest, Huancabamba Depression, Chamaya River, Marañón River, inter-Andean valley Academic editor: Rafael de Fraga | Received 26 January 2022 | Accepted 25 April 2022 | Published 11 May 2022 Citation: García-Bravo A, Guzman BK, Mendoza JE, Torres CG, Oliva M, Barboza E, Quiñones JR, Zabarburu-Veneros JL, Venegas PJ (2022) Updating the distribution of Dicrodon guttulatum Duméril & Bibron, 1839 (Reptilia, Teiidae) with a disjunct population in the eastern slope of the Peruvian Andes. Check List 18 (3): 483–491. https://doi.org/10.15560/18.3.483 © The authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 484 Check List 18 (3) Introduction At each site, we opportunistically searched for D. guttulatum within suitable microhabitats. Some individ- The teiid lizard Dicrodon guttulatum Duméril & Bibron, uals were collected following the protocols by Koch et al. 1839 is distributed along the dry coastal fringe between (2018). At each site we recorded the following data: geo- northern Peru and Ecuador (Peters and Donoso-Bar- graphical coordinates and altitude above sea level using ros 1970). In Peru it inhabits the Sechura desert and the a GPS receiver (Garmin Montana 680 using the WGS84 equatorial dry forests in the departments of Tumbes, geodetic datum), air humidity (RH%), and temperature Piura, Lambayeque, and La Libertad; in Ecuador it is (°C) taken with a digital thermohygrometer (Elitech found mainly on the south-central coast where it inhab- RC-4HC) mounted with an external sensor. its shrublands and deciduous forests in the provinces of We photographed live individuals (uncaptured and in El Oro, Guayas, Santa Elena, and Manabí (Carrillo and their microhabitat), then we attempted to capture them Icochea 1995; van Leeuwen et al. 2011; Havik et al. 2014; by hand or using a slingshot. Once captured, we placed Perez et al. 2017; Torres-Carvajal et al. 2019). lizards in cloth bags. After capture, we photographed Dicrodon guttulatum is herbivorous and feeds on dif- individuals dorsally, ventrally, and laterally. We then euth- ferent types of food; it is not specialized on “algarrobo”, anized lizards by injecting 0.5 ml of Halatal® into the heart Prosopis L. species (Pollack et al. 2007). On the Peru- area, using a tuberculin syringe. We also photographed the vian coast, its diet is based on shoots and tender fruits microhabitat, lizard burrows, and threats at each locality. of Prosopis pallida (Humb. & Bonpl. ex Willd.) Kunth, We used QGIS v. 3.16.6 (https://qgis.org/es/site/index. leaflets of Vachellia aroma var. huarango (Ruíz & J. html) to create the map of D. guttulatum (Fig. 2) in Peru Macbr.) Seigler & Ebinger, and Vachellia macracantha and Ecuador using our new data (UNTRM/INDES-CES (Humb. & Bonpl. ex Willd.) Seigler & Ebinger; seeds of expedition, 21–24.VI.2021), data from specimens in the Encelia canescens Lam., Scutia spicata (Humb. & Bonpl. herpetological collection of Centro de Ornitología y Bio- ex Schult.) Weberb., and Morisonia crotonoides (Kunth) diversidad (CORBIDI, http://www.corbidi.org/), data Christenh. & Byng, which confirms a certain degree of from specimens in the Museo de Zoología de la Pontificia selectivity in its diet (Pollack et al. 2007). Moreover, it Universidad Católica del Ecuador (QCAZ/PUCE), and is considered an efficient seed disperser of some of the data from other specimens obtained through the Global shrubby plants on which it feeds (Pollack et al. 2007). Biodiversity Information Facility (GBIF; https://www. Dicrodon guttulatum is categorized as Least Con- gbif.org/) in the National Museum of Natural History, cern by the International Union for Conservation of Smithsonian Institution (USNM), Museum of Vertebrate Nature (IUCN 2022), despite the severe fragmentation of Zoology, University of California, Berkeley (MVZ), its populations, and the continuous decline in the number Biodiversity Institute & Natural History Museum, the of individuals (Perez et al. 2017). University of Kansas (KU), Natural History Museum During a recent inventory of biodiversity carried out of Los Angeles County (LACM), California Academy by the Proyecto Especial Olmos Tinajones (Gobierno of Sciences (CAS), Museum of Comparative Zoology, Regional de Lambayeque) and Universidad Nacional Harvard University (MCZ), and San Diego Natural Toribio Rodríguez de Mendoza de Amazonas/Instituto History Museum (SDNHM), and photographic records de Investigación para el Desarrollo Sustentable de Ceja from iNaturalist (https://www.inaturalist.org/). A poly- de Selva (UNTRM/INDES-CES), we recorded a dis- gon of the potential geographic range, as a shapefile, was junct population of D. guttulatum in the ecoregion of obtained from the IUCN (https://www.iucnredlist.org/) the inter-Andean Seasonally Dry Forests of the Marañón for inclusion on our map. River (ISDFM) (Linares-Palomino 2006), in the Depart- The GBIF data were filtered according to the follow- ments of Cajamarca and Piura, on the eastern slope of the ing criteria: no coordinates, repeated coordinates, human Cordillera Occidental. observations without any evidence such as photographs, We produce a new map of the geographic distribu- altitude outside the defined range, and finally citizen- sci- tion of D. guttulatum using our new data and data from ence observations with photographic records belonging specimens in museums and online platforms with photo- to iNaturalist. The iNaturalist data were reviewed one graphic or voucher evidence. by one and all misidentified records were deleted [e.g., https://www.inaturalist.org/observations/24982312, Methods which is Ameiva ameiva (Linnaeus, 1758)]. We collected specimens under a research permit issued by the Ministerio de Desarrollo Agrario y Riego Results (Resolución de Dirección General No D000010-2021- On 16 January 2020, during biological monitoring at the MIDAGRI-SERFOR-DGGSPFFS, 12/01/2021). Limón dam, Caserío Huabal, which is part of the area We conducted an expedition on 21–24.VI.2021 with affected by the Olmos Irrigation Diversion, two of us team members from the UNTRM/INDES-CES (Appen- (AGB and JQR) recorded three individuals of D. guttula- dix Table A1). The area assessed is in the lower basin of tum (Fig. 1A), including a photograph of a juvenile indi- the Chamaya River. vidual on the left bank of the Chamaya River. García-Bravo et al. | Distribution of Dicrodon guttulatum 485 During the UNTRM/INDES-CES expedition, we Zabarburu-Veneros obs.; 8 (sex unverified) – Piura recorded 13 individuals of Dicrodon guttulatum (Appen- Department • Huancabamba Province, Huarmaca Dis- dix Table A1), including males (Fig. 1B), females, and trict, Sector el Mango, Caserío el Progreso; −05.7809, juveniles. Four specimens were collected, three of which −079.3942; 1203 m a.s.l.; 22.VI.2021; Antonio García- were adult males, and one was a juvenile of indetermi- Bravo obs.; 1 (sex unverified). nate sex. In Caserío Huabal, we observed the species living in burrows that may be interconnected. The burrows are New record. PERU – Cajamarca Department • Jaén built in sandy soils (Fig. 1C) in which Prosopis juliflora Province, San Felipe District, Caserío Huabal; −05.8787, (Sw.) DC., Vachellia macracantha, and Vallesia glabra −079.3400; 997 m a.s.l.; 22.VI.2021; Antonio García- (Cav.) Link were common plants within the ISDFM eco- Bravo leg.; 3 ♂, CORBIDI 23341–23344, 23345, 1 juv. system (Fig. 1D). We found seeds of V. glabra in the feces CORBIDI 23342 • Jaén Province, San Felipe District, surrounding the burrows of D. guttulatum (Fig. 1E). Caserío Huabal; −05.8787, −079.3400; 997 m a.s.l.; The search for additional records of D. guttula- 22.VI.2021; Antonio García-Bravo, Betty K. Guz- tum yielded 103 localities between Ecuador and Peru: man, Jani E. Mendoza, Jhon Quiñones Rámirez, J. Luis 27 from the CORBIDI Herpetology Division, 54 from Figure 1. Dicrodon guttulatum. A. An adult individual, reported on 16.I.2020, on the bank of the Limón dam, Caserío Huabal, on the left side of the Chamaya River. B. An adult individual, reported on 22.VI.2021, on the banks of the Limón dam, Caserío Huabal, on the left side of the Chamaya River. C. Burrows, Caserío Huabal. D. Microhabitat, Caserío Huabal. E. Feces with seeds of Vallesia glabra. F. Threats to the microhabitat: Rice fields burned. Photographs A–D by Antonio García-Bravo, E by J. Luis Zabarburu-Veneros and F by Jani E. Mendoza. 486 Check List 18 (3) Figure 2. Distribution map of Dicrodon guttulatum in Peru and Ecuador. García-Bravo et al. | Distribution of Dicrodon guttulatum 487 GBIF, six from QCAZ/PUCE, and 16 from iNaturalist remains of the plantation are burned (Fig. 1F). (Fig. 2; Appendix Table A1). Pollack et al. (2007) examined the feces of D. gut- Identification. We identified our collected specimens tulatum in coastal Peru and reported seven plant spe- as D. guttulatum by presence of the following charac- cies as part of its diet: Prosopis juliflora, P. pallida, ters: (1) bicuspid teeth, (2) an entire frontal scale, and Vachellia macracantha, V. aroma var. huarango, Encelia (3) granular dorsal scales (Harvey et al. 2012). These canescens, Morisonia crotonoides, and Vallesia glabra. three characters readily distinguish D. guttulatum from We also identified V. glabra, a common scrub in ISDFM (Fig. 1D), in the feces of D. guttulatum (Fig. 1E). other species of teiids from Peru. The most similar spe- With our new records and the secondary data, we cies to D. guttulatum is Dicrodon heterolepis (Tschudi, confirm that D. guttulatum is distributed on both sides of 1845) which possesses keeled dorsal scales. In addition, the Cordillera Occidental of northern Peru. Other species Ameiva aggerecusans Koch, Venegas, Rödder, Flecks & of lizards that were previously only known from the coast Böhme, 2013, Ameiva nodam Koch, Venegas, Rödder, and Pacific slope of Andes from Peru and Ecuador have Flecks & Böhme, 2013, and Ameiva concolor (Ruthven, likewise been recently reported in the ISDFM: Callo- 1924) also inhabit the IASDFM (Koch et al. 2018). These pistes flavipunctatus (Duméril & Bibron, 1839) and Phyl- Ameiva species can be easily distinguished by having the lodactylus reissii Peters, 1862 (Crespo and Koch 2015; frontal scale transversally divided (Koch et al. 2018). The Koch et al. 2018). We hypothesize that these trans-An- photographic records of D. guttulatum from iNaturalist dean lizards probably used the Huancabamba Depression were identified as having yellow or cream dots on the as a corridor between opposite slopes of the western Cor- back and a blue or reddish head, shown mainly in males, dillera Occidental in northwestern Peru. which differs from the pattern of longitudinal lines of Seven natural protected areas occur in the IASDFM, other coastal species of Ecuador and Peru such as Medo- covering an area of over 45,000 ha and accounting for pheuos edracanthus (Bocourt, 1874) and Holcosus sep- 1.97% of area of this ecoregion (Guzman et al. 2021). temlineatus (Duméril, 1851). However, D. guttulatum is not protected in any of these protected areas in the ISDFM, and we, therefore, suggest Discussion areas where this species occurs should be afforded pro- tection. The main threats to the disjunct population of D. With our new records and other data, we extend the geo- guttulatum in the ISDFM are the widespread cultivation graphical distribution and altitudinal range of Dicrodon of rice crops (Fig. 1F), which is mainly done in the lower guttulatum to include the eastern slope of Cordillera part of the Chamaya River valley, the extraction of aggre- Occidental of northern Peru, extending the altitudi- gates (sand and rocks) for construction, and, as a potential nal range of this species to about 1400 m a.s.l., and the threat, the planned Limón dam project. Importantly, local geographical distribution by approximately 50 km in a people do not consume this species here, as is the case in straight-line northeast of the previously known eastern- some towns along the Peruvian coast (Holmberg 1957). most record, 12 km north of Olmos on the Pan American Highway (MVZ 163081 in Appendix Table A1). The previously known distribution of D. guttula- Acknowledgements tum only included the northern coast and Pacific slope We thank the support of the SNIP Project no. 316114 Cre- of Peru and central Ecuador (Pollack et al. 2007; Van ation of the Services of a Biodiversity Laboratory and Leeuwen et al. 2011; Havik et al. 2014; Perez et al. 2017; Conservation of Genetic Resources of Wild Species of Torres-Carvajal et al. 2019). However, there are unpub- the Toribio Rodríguez de Mendoza National Univer- lished historical records available in GBIF: collections of sity, Amazonas” – BIODIVERSIDAD, financed by the G.K. Noble (1.IX.1916) from Perico, San Ignacio prov- National Public Investment System of the Ministry of ince, Cajamarca Department, MCZ R-18778, R-18779, Economy and Finance of Peru. We would also thank the R-18780, R-18781, R-18782 and R-18783; and collections Olmos Project for logistical support in the expedition and of R.B. Huey (20.VIII.1967) from El Arenal, Chamaya Mr. Jaime Gonzales for his logistical assistance. We are River, province of Jaén, Cajamarca Department, MVZ grateful with Luis Alza and Luis Cueto, and William W. Herp:82426 and 82427. These have gone unnoticed until Lamar for reviewing the manuscript, Gladys Marlo for now despite their biogeographical importance. translation, and Betsy Inches for providing information. We believe that D. guttulatum has not been recorded We are also indebted to Alessandro Catenazzi and one from the IASDFM in the last two decades by Koch et anonymous reviewer for their valuable comments on the al. (2018) probably because some populations have dis- manuscript. appeared (e.g., thoseat recorded by G.K. Noble in 1916). The likely cause for local extirpation is that the flat and gently sloping areas with sandy soils used by this spe- Authors’ Contributions cies for digging burrows are highly suitable for rice cul- Conceptualization: AGB, BKG. Data curation: AGB, tivation. The cultivation of rice crops floods the soils for BKG. Formal analysis: AGB, BKG. Funding acqui- weeks during the planting stage, and after the harvest the sition: AGB, BKG, MO. Investigation: AGB, BKG. 488 Check List 18 (3) Methodology: AGB, BKG, CTG, JEM, JLZV, PJV, JQR. IUCN (2022) The IUCN Red List of Threatened Species. Version Project administration: AGB. Supervision: MO, PJV. 2021-3. Internation Union for the Conservation of Nature. https:// Visualization: AGB, BKG. Writing – original draft: www.iucnredlist.org. Accessed on: 2022-4-22.Koch C, Venegas PJ, Cruz RS, Böhme W (2018) Annotated check- BKG, CTG, EB, JEM, JLZV, JQR, MO. Writing – review list and key to the species of amphibians and reptiles inhabiting and editing: AGB, PJV. the northern Peruvian dry forest along the Andean valley of the Marañón river and its tributaries. Zootaxa 4385: 1–101. https://doi. References org/10.11646/zootaxa.4385.1.1Van Leeuwen JP, Catenazzi A, Holmgren M (2011) Spatial, ontoge- Carrillo N, Icochea J (1995) Lista taxonomica preliminar de los rep- netic, and sexual effects on the diet of a teiid lizard in Arid South tiles vivientes del Perú. Publicaciones del Museo de Historia Nat- America. Journal of Herpetology 45: 472–477. https://doi.org/ ural, Universidad Nacional Mayor de San Marcos 49: 1–27. 10.1670/10-154.1 Linares-Palomino R (2006) Phytogeography and floristics of Sea- Crespo S, Koch C (2015) Notes on natural history and distribution sonally Dry Tropical Forests in Peru. 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García-Bravo et al. | Distribution of Dicrodon guttulatum 489 Appendix Table A1. Records of the locations of Dicrodon guttulatum Duméril & Bibron, 1839 in Peru and Ecuador. Geographic coordinates Voucher Locality District (Parroquia) Province (Cantón) Department Altitude (province) Country Date Institution Latitude Longitude (m) Specimen Photograph Caserío Huabal San Felipe Jaén Cajamarca Perú 22.VI.2021 UNTRM/INDES-CES −05.8787 −079.3400 997 X X Sector el Mango, Caserío el Progreso Huarmaca Huancabamba Piura Perú 22.VI.2021 −05.7809 −079.3942 1203 — X Puerto López Puerto López Puerto López Manabí Ecuador 1.I.2012 iNaturalist −01.5353 −080.7352 74 — X Machalilla Machalilla Puerto López Manabí Ecuador 5.I.2017 −01.4846 −080.7723 19 — X Salinas Salinas Salinas Santa Elena Ecuador 18.X.2020 −02.1942 −081.0021 7 — X ACP Bosque de Cañoncillo San José Pacasmayo La Libertad Perú 21.VIII.2016 −07.3723 −079.4217 122 — X Túcume Túcume Lambayeque Lambayeque Perú 2.III.2016 −06.5155 −079.8507 48 — X Salinas Salinas Salinas Santa Elena Ecuador 29.VIII.2019 −02.1877 −080.9977 71 — X Salinas Salinas Salinas Santa Elena Ecuador 13.XI.2019 −02.2029 −080.9959 7 — X Agua Blanca Machalilla Puerto López Manabí Ecuador 27.XII.2019 −01.5309 −080.7408 68 — X Salinas Salinas Salinas Santa Elena Ecuador 2.I.2020 −02.2027 −080.9955 7 — X Machalilla Machalilla Puerto López Manabí Ecuador 15.III.2019 −01.4949 −080.7902 14 — X Machalilla Machalilla Puerto López Manabí Ecuador 15.III.2019 −01.5172 −080.7715 32 — X Machalilla Machalilla Puerto López Manabí Ecuador 10.VII.2017 −01.4877 −080.7924 30 — X SN Bosque de Pómac Pítipo Ferreñafe Lambayeque Perú 10.X.2015 −06.4920 −079.7950 67 — X SN Bosque de Pómac Pítipo Ferreñafe Lambayeque Perú 13.XI.2019 −06.4919 −079.7950 66 — X Near Salinas Puerto Larrea Tosagua Manabí Ecuador 11.VII.2016 −00.6558 −080.2996 3 — X Salinas Salinas Salinas Santa Elena Ecuador 1.IV.2017 −02.2029 −080.9958 6 — X Manglares de San Pedro de Vice Vice Sechura Piura Perú 25.II.2009 CORBIDI −05.5134 −080.8926 0 X — El Peñal Cristo nos Valga Sechura Piura Perú 29.II.2009 −05.5365 −080.6400 0 X — Vega de Palomino Chulucanas Morropón Piura Perú 9.III.2009 −05.2268 −080.2973 173 X — Cerro Huacrupe Olmos Lambayeque Lambayeque Perú 24.III.2009 −05.8296 −079.9516 161 X — Estuario Virrilá Sechura Sechura Piura Perú 27.II.2009 −05.7809 −080.8577 0 X — Zorritos Zorritos Contralmirante Villar Tumbes Perú 29.V.2009 −03.6890 −080.6958 0 X — Monte de La Virgen Lambayeque Lambayeque Lambayeque Perú 16.X.2004 −06.7177 −079.9041 29 X — Alrrededores de la laguna La Niña Sechura Sechura Piura Perú 4.VI.2012 −05.7295 −080.5867 9 X — Estuario de Virrila Sechura Sechura Piura Perú 8.VI.2012 −05.8717 −080.7472 6 X — Peninsula de Illescas Sechura Sechura Piura Perú 4.VII.2012 −05.8622 −080.9944 75 X — Cerca al canal madre de CHAVIMOCHIC Chao Virú La Libertad Perú 15.III.2014 −08.4846 −078.5882 236 X — San Pedro de Lloc (alrededores de la ciudad) San Pedro de Lloc Pacasmayo La Libertad Perú 4.IX.2014 −07.4267 −079.5039 43 X — Zona reservada de Illescas Sechura Sechura Piura Perú 28.III.2013 −05.8241 −081.0658 167 X — Zona reservada de Illescas Sechura Sechura Piura Perú 20.VII.2013 −05.8329 −081.0628 182 X — Zona Reservada Illescas Sechura Sechura Piura Perú 6.IV.2016 −06.1063 −081.0526 13 X — Zona Reservada Illescas Sechura Sechura Piura Perú 6.IV.2016 −06.1070 −081.0516 10 X — Carretera Mancora-Puntasal Canoas de Punta Sal Contralmirante Villar Tumbes Perú .X.2002 −04.0515 −080.9549 27 X — 490 Check List 18 (3) Locality District (Parroquia) Province (Cantón) Department Geographic coordinates Voucher (province) Country Date Institution Altitude Latitude Longitude (m) Specimen Photograph Montes de La Virgen Lambayeque Chiclayo Lambayeque Perú .I.2002 CORBIDI −06.7164 −079.9005 34 X — La Arenita parte baja Razuri Ascope La Libertad Perú 2.V.2018 −07.6650 −079.3340 81 X — Campamento San Carlos Chao Virú La Libertad Perú 17.IX.2018 −08.5310 −078.6400 129 X — Quebrada de Palo Redondo Chao Virú La Libertad Perú 11.VII.2017 −08.6450 −078.4830 436 X — Campamento San José Virú Virú La Libertad Perú 27.IV.2017 −08.4090 −078.7220 127 X — Los Organos Los Organos Talara Piura Perú 20.X.2018 −04.2172 −081.0090 337 X — Campiña Zaraque Virú Virú La Libertad Perú 14.XI.2018 −08.3920 −078.7210 118 X — dunas Lambayeque Lambayeque Lambayeque Perú 3.III.2020 −06.6510 −080.0030 6 X — Dunas con mucha vegetación Lambayeque Lambayeque Lambayeque Perú 21.IX.2020 −06.6510 −080.0029 16 X — Por carretera a avícola Lambayeque Lambayeque Lambayeque Perú 19.IX.2021 −06.6707 −079.9797 14 X — San Clemente (between Charapote & Bahia), 2 km N of Charapoto Sucre Manabí Ecuador 17.X.1965 GBIF −00.7306 −080.5014 60 X — Playas, 1 KM N General Villamil Playas Guayas Ecuador 3.I.1959 −02.6167 −080.3917 12 X — Colonche, 1 KM SW Colonche Santa Elena Santa Elena Ecuador 8.X.1965 −02.0250 −080.6833 8 X — Playas, 12 KM SE, Villamil General Villamil Playas Guayas Ecuador 9.I.1959 −02.7083 −080.3167 6 X — Manta, 6 km SE of, 5 km NW of Montecristi Manta Manta Manabí Ecuador 15.X.1965 −00.9500 −080.6714 20 X — El Empalme (= Velasco Ibarra), junction of Quito- Velasco Ibarra El Empalme Guayas Ecuador .II.1965 −01.0406 −079.6361 70 X — Manta-Guayaquil Road Valle de Casanga, El Empalme El Empalme Paltas Loja Ecuador .III.1961 −04.1333 −079.8333 699 X — Pariñas Valley Pariñas Talara Piura Perú .XI.1940 −04.5278 −081.2000 30 X — 6 km SSE Bayovar Sechura Sechura Piura Perú 31.VII.1968 −05.9001 −081.0459 352 X — 5 km SE Bayovar Sechura Sechura Piura Perú 10.VIII.1968 −05.8820 −081.0347 355 X — vicinity of Tric Trac, 2 km E Bayovar Sechura Sechura Piura Perú 31.VII.1968 −05.8333 −081.0167 10 X — 5 mi SSE Bayovar Sechura Sechura Piura Perú 13.VIII.1968 −05.9168 −081.0390 366 X — 3 km SE Morrope Morrope Lambayeque Lambayeque Perú 15.VII.1967 −06.5525 −079.9975 30 X — 4 km SE Bayovar Sechura Sechura Piura Perú 22.VII.1967 −05.8589 −081.0245 141 X — 7 km SSE Mocupe Lagunas Chiclayo Lambayeque Perú 6.IX.1967 −07.0462 −079.5515 91 X — 11 km N & 28 km E Barranco, Cabo Blanco Cabo Blanco Talara Piura Perú 7.VIII.1967 −04.2172 −080.9978 234 X — El Arenal, Río Chamaya San Felipe Jaén Cajamarca Perú 20.VIII.1967 −06.1289 −079.0572 944 X — Parinas, 7 km N & 15 km E Talara Pariñas Pariñas Talara Perú 3.VIII.1967 −04.5125 −081.1340 129 X — Playas, 60 mi SW Guayaquil General Villamil Playas Guayas Ecuador 3.III.1964 −02.5507 −080.2814 24 X — Playas General Villamil Playas Guayas Ecuador 3.III.1964 −02.6330 −080.3830 7 X — 3 mi N Oyotun Oyotun Chiclayo Lambayeque Perú 29.VIII.1969 −06.8119 −079.2914 305 X — 2 mi SE Morrope Morrope Lambayeque Lambayeque Perú 16.VIII.1970 −06.5705 −079.9962 30 X — Quebrada, between Talara & Lobitos Lobitos Talara Piura Perú 1.XII.1980 −04.4783 −081.2703 21 X — Quebrada, ca. 5 km NE (by air) Negritos La Brea Talara Piura Perú 2.XII.1980 −04.6230 −081.2704 66 X — 12 km N Olmos on Pan American Hwy. Olmos Lambayeque Lambayeque Perú 30.VIII.1978 −05.8925 −079.7831 152 X — Parinas, 7 km N & 14 km E Talara Pariñas Talara Piura Perú 20.II.1972 −04.5333 −081.1500 30 X — 2.2 km N Playas General Villamil Playas Guayas Ecuador 17.VI.1971 −02.6083 −080.3917 13 X — Playas General Villamil Playas Guayas Ecuador 14.VII.1971 −02.6333 −080.3833 8 X — García-Bravo et al. | Distribution of Dicrodon guttulatum 491 Department Geographic coordinates Altitude VoucherLocality District (Parroquia) Province (Cantón) (province) Country Date Institution Latitude Longitude (m) Specimen Photograph Sierra de la Vieja 7 km S Motupe Motupe Lambayeque Lambayeque Perú 4.VI.1976 GBIF −06.2028 −079.6944 119 X — 5 km E, 4.2 km NNE Lobitos Lobitos Talara Piura Perú 6.XII.1968 −04.4042 −081.2528 19 X — 3 km S Talara La Brea Talara Piura Perú 3.XII.1968 −04.6167 −081.2889 43 X — 2.4 km SW Chulucanas, in Mesquito thicket along Rio Chulucanas Morropón Piura Perú 1.XII.1968 −05.1194 −080.1694 87 X — Piura 3.7 km ENE Los Organos Breach Dunes Los Órganos Talara Piura Perú 4.XII.1968 −04.1458 −081.1083 8 X — 3 km ENE Las Lomas in sandy area Las Lomas Piura Piura Perú 6.XII.1968 −04.6361 −080.2222 274 X — Quebrada Seca, 11.8 km ENE Mancora, under bark in Canoas de Punta Sal Contralmirante Villar Tumbes Perú 4.XII.1968 −04.0500 −080.9125 162 X — misquito grove Puerto Pizarro on estaru beach with mesquite, cacti Tumbes Tumbes Tumbes Perú 5.XII.1968 −03.4833 −080.3833 1 X — 1.2 km S., 1 km E Puerto Pizarro in Cacti, Acacia, thorn Tumbes Tumbes Tumbes Perú 5.XII.1968 −03.4942 −080.3743 3 X — thicket Songora, Quebrada Marcavelica Sullana Piura Perú 22.IX.1940 −04.7000 −081.0694 204 X — Sullana Sullana Sullana Piura Perú .VII.1916 −04.8833 −080.6833 39 X — Grau Tumbes Tumbes Tumbes Tumbes Perú   −03.5654 −080.4570 10 X — Santa Rosa (nr.Salinas),Prov.Guayas Santa Rosa Daule Guayas Ecuador 13.II.1963 −02.2167 −080.9667 6 X — Perico Chirinos San Ignacio Cajamarca Perú 1.IX.1916 −05.3439 −078.7947 441 X — Rocks of Morro,Guayas Prov La Puntilla Samborondón Guayas Ecuador 13.VI.1964 −02.0000 −079.8333 3 X — Buenos Aires,Guayas Prov Chanduy Santa Elena Guayas Ecuador 6.VI.1964 −02.7857 −079.9267 31 X — nr.East data:Guayas Prov Nobol Nobol Guayas Ecuador 8.III.1963 −01.8960 −080.0160 5 X — El Prado, Guayas Prov. Chongón Guayaquil Guayas Ecuador 8.III.1963 −01.7167 −079.8833 9 X — Ecuador: Progresso W. of Guayaquil Progreso Guayaquil Guayas Ecuador 19.VIII.1964 −02.4000 −080.3667 97 X — Ecuador:Santa Rosa Santa Rosa Santa Rosa El Oro Ecuador 26.IV.1963 −03.4488 −079.9595 11 X — Ecuador:Guayaquil,Guayas Prov. Rocafuerte Guayaquil Guayas Ecuador 29.I.1963 −02.1667 −079.9000 4 X — Isla de los Muertos or Isla Santa Clara Jambelí Santa Rosa El Oro Ecuador 3.V.1963 −03.1667 −080.4500 1 X — Peru,Piura Talara Alta Pariñas Talara Piura Perú 1.I.1980 −04.5758 −081.2692 15 X — Posorja, Gulf of Guayaquil Posorja Guayaquil Guayas Ecuador   −02.7000 −080.2500 3 X — Guayas, Buenos Aires Chanduy Santa Elena Guayas Ecuador 6.VII.1964 −01.3829 −080.3624 396 X — Picos Point Zorritos Contralmirante Villar Tumbes Perú 16.II.1938 −03.7500 −080.8000 1 X — El Progreso El Progreso Pasaje El Oro Ecuador 3.I.1986 QCAZ/PUCE −03.2883 −079.75812 92 X — 108 km vía Guayaquil-Salinas La Libertad La Libertad Santa Elena Ecuador 12.XI.1992 −02.2330 −080.83499 57 X — Atahualpa Atahualpa Santa Elena Santa Elena Ecuador 19.X.1993 −02.3092 −080.7709 46 X — El Aromo San Lorenzo Manta Manabí Ecuador 1.V.2010 −01.0467 −080.8328 297 X — La independencia, Dique - La Independencia Sechura Sechura Sechura Perú 26.XI.2009 −05.3799 −080.6774 29 X — Isla Puná, Comuna Bellavista Posorja Guayaquil Guayas Ecuador 14.XI.2015 −02.7772 −080.2233 9 X —