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Distribution of Bothrops asper in Tamaulipas, Mexico and a review of prey items.

Bothrops asper is a wide-ranging Neotropical pit viper. We follow Liner and Casas-Andreu (2008) in using their standardized English and Spanish names for the species, terciopelo and cuatro narices, respectively. Terciopelos occur on the northwestern Pacific and Caribbean versants of South America (from extreme northwestern Peru to northern Venezuela), throughout much of Central America and into eastern Mexico, with vertical distributions from sea level to 1,300 m in Mexico and Central America and up to 2,640 m in South America (Campbell and Lamar, 2004). Townsend et al. (2006) reported a maximum elevation of 1,450 m in Honduras. Bothrops asper has a relatively lengthy synonymy with twenty synonyms and combinations (McDiarmid et al., 1999). An extended discussion on the taxonomy is beyond the scope of this review; however, we note that literature specifically addressing Tamaulipas populations have used Bothrops atrox (e.g., Martin 1955, 1958), Bothrops atrox asper (e.g., Smith and Taylor, 1945), and Bothrops asper (e.g., Auth et al., 2000; Campbell and Lamar, 2004; Lavin-Murcio and Lazcano, 2010; Farr, 2015).

Martin del Campo (1937) included Tamaulipas in the distribution of B. atrox asper; however, it was noted that the record "seems to be based upon probability of occurrence instead of actual specimens" (Smith and Taylor, 1945:180). A specimen from Tamaulipas (CAS-71773) collected in 1920 is notable, although no specific locality is available for the snake, and we are unaware of any references to this specimen in the literature (Appendix 1; institution abbreviations per Sabaj Perez, 2014). Martin (1955) included B. asper (as B. atrox) among 39 amphibian and reptile species representing new state records from the Gomez Farias region of Tamaulipas. Martin (1958) provided further details on specimens collected in the vicinity of Gomez Farias, with explicit localities and the associated vegetation zones. Subsequent to Martin (1955, 1958), the occurrence of B. asper in Tamaulipas has been reflected in the literature; however, like many species, additional localities have rarely been reported despite the accumulation of specimens in museums for over a half a century (Farr et al., 2013). Auth et al. (2000) reported one additional record from Tamaulipas (SMBU-BCBF 68-97; Appendix 2). Campbell and Lamar (2004) included Tamaulipas in the distribution but their map only indicted generalized shading in the state: no dots were included and specific localities or museum vouchers were not identified.

One of us (WLF) queried over 100 institutions for information on their holdings of Tamaulipan herpetofauna. Sixty-one institutions responded providing a total of 28 B. asper records from eight collections. We have examined and verified all of these specimens (Appendix 1). The localities of specimens were plotted on a map along with a literature record from Tamaulipas, literature records from San Luis Potosi, Mexico, and an additional museum specimen (TNHC, verified by personnel from that institution) from Veracruz, Mexico (Fig 1; Appendix 2). Coordinates are expressed in map datum WGS84. Coordinates and elevations for specimens we collected were recorded with a handheld global positioning system unit and are presented in parentheses in Appendix 1. For museum and literature records without coordinates in the original data, we determined the approximate coordinates using Google Earth based on recorded localities, and provide them here in brackets in Appendices 1 and 2. We follow Sabaj Perez (2014) in the abbreviations for symbolic codes of institutional resource collections. We describe the distribution and associated vegetation zones and identify all known municipality records of B. asper in Tamaulipas. A specimen (UANL-6608) with a conspicuous bulge, collected in the municipality of Tula, was dissected and we found a small mammal in the stomach. The skull of this prey item was cleaned and identified. Literature was reviewed for references identifying specific prey items (e.g., family, genus, and species) in the diet of B. asper (Table 1). Literature that only identified generalized prey items (e.g., class, order, frogs, lizards, rodents, small mammals, etc.) is not included here.

Bothrops asper occurs in a relatively limited area of south-central Tamaulipas, where it is generally associated with humid, mid-low elevations on the eastern slopes of the Sierra Madre Oriental and adjacent lowlands (Fig. 1). Martin (1958) reported B. asper from tropical deciduous forest, tropical evergreen forest, and lower sections of cloud forest in Tamaulipas (vegetation zones defined therein). To that we add habitat in the municipality of Tula along the Rio Gallos Grandes, where B. asper follows the river well into the Sierra Madre Oriental. The riparian areas there have elements of temperate Madrean oak woodlands, characterized by conspicuous stands of what appear to be Acer grandidentatum (bigtooth maple), although we were unable to verify the identification of these trees with certainty. The slopes above the riparian areas to the west are slightly more arid with Quercus (oak) and Juniperus (juniper) forest. The slopes above the riparian areas to the east are more humid with Quercus (oak) and Sabal (palm). We confirmed B. asperfrom seven municipalities: Antiguo Morelos, El Mante, Gomez Farias, Llera, Ocampo, Tula, and Xicotencatl. The occurrence of B. asper should be anticipated from the municipality of Nuevo Morelos with near certainty, as every adjacent municipality holds records, including two municipalities (El Naranjo and Ciudad Valles) in the neighboring state of San Luis Potosi, (Taylor, 1949; Auth et al., 2000; Lemos-Espinal and Dixon, 2013). A vertical distribution between 98-1,190 m can be confirmed in Tamaulipas. Bothrops asper is apparently absent from the lower elevations to the east which support semiarid thorn scrub habitat in Tamaulipas. However, literature and museum records are available from lower elevations to the east in adjacent areas in San Luis Potosi and Veracruz (Fig. 1; Appendix 2). An extensive area of marshlands and lagoons associated with the Rio Tamesi on the Veracruz border, which sustain extensive Cyperus and Typha vegetation, might be a factor in limiting the dispersal of the species in southeast Tamaulipas.

Bothrops asper reaches range-wide distributional limits in Tamaulipas. A record (UANL-6608) from the Sierra Madre Oriental in the municipality of Tula represents the westernmost locality, located less than 17 km east of Chihuahua Desert scrub in the Tula Valley. The arid environment of the Mexican Plateau presents a natural barrier, and the species should not be anticipated significantly west of this locality. Three specimens from the municipality of Llera (LACM-20229, 20230, 20231) represent the northernmost locality, approximately 37 km south of the Tropic of Cancer. Significant areas of tropical deciduous forest extend northward to the general vicinity of the Tropic of Cancer and the Rio Guayalejo, and future collecting might produce new records 25-50 km to the north, but this remains to be demonstrated. Martin (1958) noted that B. asper should be anticipated in the tropical deciduous forest of the Sierra de Tamaulipas. We are unaware of any literature or museum records from the Sierra de Tamaulipas. Much of the habitat in the Sierra de Tamaulipas would appear suitable for the species, but this mountain range is isolated on the costal plan and separated from the tropical deciduous forest associated with the Sierra Madre Oriental by [mayor que o igual a]35 km of low-elevation arid thorn scrub habitat.

We found a juvenile B. asper, approximately 325 mm in total length, with a conspicuous bulge in its abdomen (UANL-6608). Although this snake was found dead on the road, the head was pulverized or shredded beyond recognition but the rest of its body was in relatively good condition, suggesting it might not have been run over by a vehicle but deliberately killed by a predator or perhaps a person. The specimen was later dissected and the stomach content was identified from its skull as a Cryptotis parva (North American least shrew or musarana), now in the Universidad Autonoma de Nuevo Leon, Mammal Collection (UANL-4320). Averaging 79 mm and 4-7.5 g, C. parva are notable as one of the smallest mammals of North America (Schmidly, 1994). To our knowledge the genus Cryptotis, the Family Soricidae, or the order Eulipotyphla have not previously been reported in the diet of B. asper.

Martins et al. (2002) reviewed the feeding habits of 22 species of Bothrops and found 32 prey items in the digestive tracts of their B. asper sample, although the prey items were not identified beyond class, order, or suborder. Martins et al. (2002) concluded B. asper is a diet generalist, with an ontogenetic diet shift from a greater percentage of ectotherms in juveniles to a higher percentage of endotherms, particularly small mammals, in adults. Boada et al. (2005) dissected 21 B. asper from Ecuador and found prey items in 14 specimens. Boada et al. (2005) found no clear distinction between juveniles and adults regarding ectothermic and endothermic prey, suggesting B. asper is a diet generalist regardless of size or age, although they noted their sample size was limited. Sasa et al. (2009) reviewed the natural history of the terciopelo in Costa Rica and analyzed the gut contents of 56 adult or subadult terciopelos. They published a remarkable table identifying 33 species and seven additional genera and orders of prey items in the diet of B. asper gleaned from both literature and their samples. We reviewed the literature identifying specific prey items of B. asper and found a number of items reported both prior and subsequent to Sasa et al. (2009) that were not included in their review (Table 1). These data do support a generalized ontogenetic diet shift from a greater percentage of ectothermic prey in juveniles to a higher percentage of endothermic prey in adults. However, clearly B. asper will opportunistically prey on a remarkably wide spectrum of both ectothermic and endothermic species regardless of size or age. Only Aves (birds) and Actinopterygii (ray-finned fishes) are exclusively identified in the diet of adults.

Several authors have reported invertebrate and insect remains in the digestive systems of B. asper including Coleoptera, Diptera, Hemiptera, Hymenoptera, and Orthoptera. Most authors interpreted these as instances of secondary ingestion from anuran and lizard prey. Boada et al. (2005) interpreted insect remains found in specimens along with anuran and lizard remains as secondary ingestion, but in specimens containing only insect remains (Coleoptera and Hemiptera) as primary prey. Boada et al. (2005) found plant matter (leaves) in the digestive tract of one specimen, presumably ingested with insect prey. Sasa et al. (2009) noted that centipedes, squamata, and fishes were an infrequent but important part of the diet, particularly for juveniles, but they did not include any examples of the latter. March (1928) commented that young opossums appear to be the favorite food in the region ofTela, Honduras, although no data were provided. Picado (1931) referred to zorrillos (skunks) in the diet of B. asper; however, no further details were provided and it is not clear if this was a documented prey item or simply an example of a small mammal that might fall prey to B. asper. Cannibalism has been reported for juveniles of both captive (Burger and Smith, 1950; Tryon, 1985; Correa Sanchez et al., 1998) and wild individuals (Buttenhoff and Vogt, 1995). Scavenging of anurans and rodents has been reported (Logan and Montero, 2009; Sasa et al., 2009). Ryan et al. (2010) provided an account of a B. asper succumbing to the toxins of a Leptodactylus savagei while preying on the frog, and Boada et al. (2005) noted an observation in which a Bufo marinus (=Rhinella horribilis) was suspected in contributing to the death of a B. asper found dead on a road. Beebe (1946) identified Leptodactylus pentadactylus, ameivas and other unidentified lizards, a small snake, remains of mice, spiny rats, and fur of a small mammal as prey items of B. atrox from two localities, Kartabo, British Guiana and Caripito, Venezuela. His samples from British Guiana were clearly B. atrox, but specimens from Venezuela were likely B. asper. Unfortunately, Beebe (1946) did not distinguish which specimens or prey items came from which localities.

APPENDIX 1--Specimens Examined--MUNICIPIO DE ANTIGUO MORELOS: AMNH 172091, 12 mi [19.3 km] SW of Mante on Mexico 80 [22.60076[degrees]N, 99.03015[degrees]W; elev. 278 m.]. UANL 4456, Antiguo Morelos- Cd. Valles a 4.6 Km al W del Antiguo Morelos [22.55250[degrees]N, 99.14750[degrees]W; elev. 397 m.]. UANL-6452, Near cave at Al Abra, 8 km NE of Antiguo Morelos (22.60666[degrees]N, 99.02430[degrees]W; elev. 263 m.). MUNICIPIO DE EL MANTE: AMNH 161864, 7 mi. S Ciudad Mante [22.65369[degrees]N, 99.02221[degrees]W; elev. 109 m.]. MUNICIPIO DE GOMEZ FARIAS: AMNH 172092, AMNH 172093 (2), Gomez Farias, between Cd. Victoria and Cd. Mante, in sugar cane fields [23.02214[degrees]N, 99.09088[degrees]W; elev. 98 m.]. TCWC 71142, Gomez Farias, estacion Los Cedros [23.04726[degrees]N, 99.15498[degrees]W; elev. 391 m.]. TU 15681, 7 km. NE. Gomez Farias [23.09676[degrees]N, 99.11260[degrees]W; elev. 134 m.]. UANL-6508 (skin only), In the town of Gomez Farias (23.03388[degrees]N, 99.15000[degrees]W; elev. 364 m.). UANL-7874, 3 km al SE del Ej. San Jose [23.02738[degrees]N, 99.22388[degrees]W; elev. 1173 m.]. UIMNH 51144, 2.5 mi W Gomez Farias [23.05541[degrees]N, 99.19327[degrees]W; elev. 913 m.]. UMMZ 101263 (head only), Rancho del Cielo, 5 mi. NW of Gomez Farias [23.10000[degrees]N, 99.19250[degrees]W; elev. 1190 m.]. UMMZ 101264, Rancho del Cielo, 5 mi. NW of Gomez Farias [23.10000[degrees]N, 99.19250[degrees]W; elev. 1190 m.]. UMMZ 110824, Gomez Farias [23.04726[degrees]N, 99.15498[degrees]W; elev. 391 m.]. UMMZ 110825, Gomez Farias [23.04726[degrees]N, 99.15498[degrees]W; elev. 391 m.]. UMMZ 111348 (head only), UMMZ 111349 (head only), Gomez Farias, near [ca. 23.04726[degrees]N, 99.15498[degrees]W; elev. 391 m.]. UMMZ 112901, Gomez Farias, 3 mi. NNE of at Ejido La Azteca [23.08217[degrees]N, 99.12704[degrees]W; elev. 116 m.]. MUNICIPIO DE LLERA: LACM-20229, LACM-20230, LACM-20231, Rio Sabinas, 2 mi. W Encino [23.14181[degrees]N, 99.14586[degrees]W; elev. 145 m.]. MUNICIPIO DE OCAMPO: UANL-6573, On hwy 66, 18.5 km NW of Ocampo (22.91527[degrees]N, 99.49975[degrees]W; elev. 651 m.). UMMZ 111053, Ocampo, 7.2 airline mi. N of on road to Refugio [ca. 22.97652[degrees]N, 99.35948[degrees]W; elev. 1000 m.]. UMMZ 111350 (head only), Aserradero del Paraiso (near), 9 mi. NNW of Chamal [ca. 22.96306[degrees]N, 99.19276[degrees]W; elev. 500 m.]. MUNICIPIO DE TULA: UANL-6608, 11 km. SE of Gallos Grandes along the Rio Gallos Grandes, 43.5 km SE of Tula (22.643532[degrees]N, 99.543063[degrees]W; elev. 814 m.). MUNICIPIO DE XICOTENCATL OR GOMEZ FARIAS: UMMZ 102911, Pano Ayuctle [=El Azteca], 5 mi. NE of Gomez Farias along the Rio Sabinas [23.11300[degrees]N, 99.14340[degrees]W; elev. 123 m.]. UMMZ 104114, Pano Ayuctle [=El Azteca], 5 mi. NE of Gomez Farias along the Rio Sabinas [23.11300[degrees]N, 99.14340[degrees]W; elev. 123 m.]. MUNICIPIO UNKNOWN: CAS 71773, Tamaulipas.

APPENDIX 2--Literature Records and Other Museum Specimens--San Luis Potosi: MUNICIPIO DE CIUDAD VALLES: 3.2 km E of Ciudad Valles (21.9833[degrees]N, 98.9833[degrees]W; 99 elev. m.) (Lemos-Espinal and Dixon, 2013). 17.2 km W, 14.2 km E of Tamuin (21.9671[degrees]N, 98.9000[degrees]W; 181 elev. m.) (Lemos-Espinal and Dixon, 2013). MUNICIPIO DE EBANO: Ebano. [22.2117[degrees]N, 098.3696[degrees]W; 56 elev. m.] (Lemos-Espinal and Dixon, 2013). KU-24032, Hda. Limon, 10 mi. [16.09 km] W of Ebano [22.1504[degrees]N, 98.5114[degrees]W; 29 elev. m.] (Taylor, 1950). MUNICIPIO DE EL NARANJO: BCB-7546, El Salto [22.5861[degrees]N, 99.3825[degrees]W; 427 elev. m.] (Auth et al., 2000). KU-24033, El Salto [22.5861[degrees]N, 99.3825[degrees]W; 427 elev. m.] (Taylor, 1949). Tamaulipas: MUNICIPIO DE OCAMPO: SMBU BCBF 68-97, Chamal, 6 mi. NW of - [22.90224[degrees]N, 99.26285[degrees]W; elev. 354 m.] (Auth et al., 2000). Veracruz: MUNICIPIO DE TAMPICO ALTA: TNHC-87818, 32 km S of Tampico [21.9350[degrees]N, 97.7607[degrees]W; elev. 68 m].

Submitted 17 June 2016. Accepted 6 February 2017.

Associate Editor was Neil B. Ford.

We thank the following individuals who provided data, allowed us the opportunity to examine voucher specimens held in the respective institutions, or both: D. R. Frost, D. Kizirian, D. Dickey, M. Arnold, A. Schweitzer, R. Pascocello (AMNH); J. Vindom (CAS); R. Brown, D. Blackburn, A. Campbell, L. Trueb (KU); G. Pauly, N. Camacho (LACM); J. Rosado (MCZ); J. R. Dixon, T. J. Hibbitts (TCWC); D. Cannatella, T. J. LaDuc (TNHC); H. Dundee, N. Rios (TU); C. Phillips, D. Wylie, M. Dreslik (UIMNH); G. Schneider, R. A. Nussbaum (UMMZ); R. Heyer, T. Hartsell, S. Gotte, K. Tighe (USNM);J. A. Campbell, C. Franklin (UTA); and C. Lieb, R. Couvillon (UTEP). T. J. LaDuc (TNHC) verified a specimen on our behalf, and we thank A. N. Ruiz Heredia for help identifying Cryptotis parva. We thank the Universidad Autonoma de Nuevo Leon and its research programs (Programa de Apoyo a la Investigacion Cientifica y Tecnologica CN361-15). N. Ford provided some difficult-to-obtain literature. Fieldwork was conducted under Secretaria del Medio Ambiente y Recursos Naturales (now Instituto Nacional de Ecologia y Cambio Climatico) permits 7150/97, FAUT-0045, 1624/05, 0800/06, 1085/07, 1255/08.


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11019 Wainfleet Lane, Houston, TX 77096 (WLF)

Universidad Autonoma de Nuevo Leon, Facultad de Ciencias Biologicas, Laboratorio de Herpetologia, Apartado Postal 513, San

Nicolas de los Garza, Nuevo Leon, C.P. 66450, Mexico (DL)

* Correspondent:

Caption: FIG. 1--The distribution of Bothrops asper in Tamaulipas and adjacent San Luis Potosi and Veracruz, Mexico. Solid dots = verified museum vouchers, open dots = literature records, question mark = area of probable occurrence.
TABLE 1--Specific prey items identified in the diet of Bothrops asper.

Prey                                          Note             Size

Invertebrates                         Dissection             Juvenile
Class Chilopoda
Scolopendra                           Dissection             Adult
Scolopendra                                    --            Juvenile
Scolopendra                           Dissection             Juvenile
Scolopendra angulata                  Dissection             Juvenile
Class Insecta
Coleoptera                            Dissection             Juvenile
Diptera                               Dissection                --
Hemiptera                             Dissection             Juvenile
Hymenoptera                           Dissection                --
Orthoptera                            Dissection                --
Orthoptera                                     --            Juvenile
Orthoptera                            Dissection             Adult
Class Malacostraca
Crayfish                                       --               --
Class Actinopterygii
Synbranchus marmoratus                Dissection             Adult
Class Amphibia
Dermophis parviceps                   In situ observation    Sub adult
Anuran                                In situ                Sub adult
                                        observation (b)
Acris gryllus                         Captivity              Juvenile
Craugastor fitzingeri                 Dissection                --
Dendrobatid frogs                     In situ                Juvenile
                                        observation (c)
Eleutherodactylus                     In situ observation    Juvenile
Eleutherodactylus                              --            Juvenile
Eleutherodactylus                     Dissection             Juvenile
  [Craugastor] rhodopis
Eleutherodactylus                     Dissection             Juvenile
  [Pristimantis] achatinus
Hyla crucifer                         Captivity              Juvenile
Leptodactylus bolivianus              Dissection                --
Leptodactylus savagei                 In situ                Adult
                                        observation (c)
Leptodactylus savagei                 Dissection             Adult
Lithobates forreri                    In situ observation    Adult
Lithobates forreri                    Dissection             Adult
Lithobates vaillanti                  Dissection                --
Lithobates warzewitzchi               Dissection                --
Mannophryne trinitatis                         --            Juvenile
Rhinella marina                       Dissection                --
Bufo [Rhinella] marinus               In situ observation    Subadult
Smilisca baudinii                     Dissection             Juvenile
Smilisca phaeota                      Dissection                --
Smilisca sordida                      Dissection                --
Class Reptilia
Amphisbaena alba                      Dissection             Adult
Ameiva [Holcosus] festiva             Dissection                --
Ameiva [Holcosus] undulata            Dissection             Juvenile
Anolis                                Captivity              Adult
Anolis                                Captivity              Juvenile
Anolis                                         --            Juvenile
Anolis limifrons                      Dissection             Juvenile
Ctenosaura similis                    Dissection             Adult
Gonatodes fuscus [albogularis]                 --               --
Lepidophyma reticulatum               Dissection             Adult
Lepidophyma tuxtlae                   In situ observation    Sub adult
Lygosoma [Sphenomorphus]              Dissection                --
Skinks                                         --            Juvenile
Bothrops asper                        Captivity (c)          Juvenile
Bothrops asper                        Dissection             Juvenile
Bothrops asper                        Dissection             Juvenile
Bothrops asper                        Captivity              Juvenile
Bothrops asper                        Captivity              Juvenile
Crotalus horridus                     Captivity (c)          Juvenile
Erythrolamprus bizona                          --            Adult
Ninia atrata                                   --               --
Ninia sebae                           Dissection                --
Storeria dekayi                       Captivity              Juvenile
Class Aves
Feathers                              Dissection                --
Feathers                              Dissection                --

Unidentified passerines (feathers)    Dissection             Adult
Cantorchilus [Thryothorus]            Dissection             Adult
Eucometis penicllata                  In situ observation    Adult
Troglodytes                           Dissection             Adult
Volantinia jacarina                   Dissection             Adult
Class Mammalia
Hair                                  Dissection             Adult
Mammalian hair                        Dissection             Juvenile
Small marsupials                               --            Adult
Full-grown opossum                             --            Adult
Caluromys derbianus                   Dissection             Adult
Didelphis                                      --            Adult
Didelphis marsupialis                          --            Adult
Didelphis marsupialis                 A. B. Carr story       Adult
Philander                                      --            Adult
Philander opossum                     Dissection             Adult
Rat sized rodent                      Dissection             Adult
Muridae                               Dissection             Subadult
Heteromys                                      --            Adult
Heteromys desmarestianus              Dissection                --
Melanomys caliginosus                 Dissection                --
Mus musculus                          Dissection                --
Mus musculus                          Captivity              Juvenile
Mus musculus                          Captivity              Juvenile
Oligoryzomys fulvescens               Dissection             Adult
Oryzomys                              Dissection             Adult
Proechimys semispinosus               Dissection             Adult
Proechimys semispinosus               Dissection             Adult
Rattus rattus                         Dissection             Juvenile
Rattus rattus                         Dissection             Juvenile
Rattus rattus "probably"              Dissection             Adult
Sigmodon hirsutus                     Dissection                --
Sigmodon peruanus                     Dissection             Subadult
Wood Rat [Neotoma]                             --               --
Coendou rothschildi                   Dissection             Adult
Sylvilagus brasiliensis                        --            Adult
Skunks                                         --            Adult
Cryptotis parva                       Dissection             Juvenile

Prey                                   Country

Invertebrates                         Mexico
Class Chilopoda
Scolopendra                           Ecuador
Scolopendra                           Guatemala
Scolopendra                           Colombia
Scolopendra angulata                  Colombia
Class Insecta
Coleoptera                            Ecuador
Diptera                               Ecuador
Hemiptera                             Ecuador
Hymenoptera                           Ecuador
Orthoptera                            Ecuador
Orthoptera                            Guatemala
Orthoptera                            Costa Rica
Class Malacostraca
Crayfish                              Trinidad
Class Actinopterygii
Synbranchus marmoratus                Venezuela
Class Amphibia
Dermophis parviceps                   Costa Rica
Anuran                                Costa Rica
Acris gryllus                         Mexico
Craugastor fitzingeri                 Costa Rica
Dendrobatid frogs                     Venezuela
Eleutherodactylus                     Costa Rica
Eleutherodactylus                     Guatemala
Eleutherodactylus                     Mexico
  [Craugastor] rhodopis
Eleutherodactylus                     Ecuador
  [Pristimantis] achatinus
Hyla crucifer                         Mexico
Leptodactylus bolivianus              Costa Rica
Leptodactylus savagei                 Costa Rica
Leptodactylus savagei                 Costa Rica
Lithobates forreri                    Costa Rica
Lithobates forreri                    Costa Rica
Lithobates vaillanti                  Costa Rica
Lithobates warzewitzchi               Costa Rica
Mannophryne trinitatis                Trinidad
Rhinella marina                       Costa Rica
Bufo [Rhinella] marinus               Ecuador
Smilisca baudinii                     Mexico
Smilisca phaeota                      Costa Rica
Smilisca sordida                      Costa Rica
Class Reptilia
Amphisbaena alba                      Venezuela
Ameiva [Holcosus] festiva             Costa Rica
Ameiva [Holcosus] undulata            Mexico
Anolis                                Costa Rica
Anolis                                Costa Rica
Anolis                                Guatemala
Anolis limifrons                      Panama
Ctenosaura similis                    Costa Rica
Gonatodes fuscus [albogularis]        Colombia
Lepidophyma reticulatum               Costa Rica
Lepidophyma tuxtlae                   Mexico
Lygosoma [Sphenomorphus]              Guatemala
Skinks                                Guatemala
Bothrops asper                        Mexico
Bothrops asper                        Mexico
Bothrops asper                        Mexico
Bothrops asper                        Mexico
Bothrops asper                        Costa Rica
Crotalus horridus                     Mexico
Erythrolamprus bizona                 Costa Rica
Ninia atrata                          Trinidad
Ninia sebae                           Costa Rica
Storeria dekayi                       Mexico
Class Aves
Feathers                              Costa Rica
Feathers                              Guatemala
Unidentified passerines (feathers)    Costa Rica
Cantorchilus [Thryothorus]            Ecuador
Eucometis penicllata                  Panama
Troglodytes                           Venezuela
Volantinia jacarina                   Costa Rica
Class Mammalia
Hair                                  Panama
Mammalian hair                        Ecuador
Small marsupials                      Costa Rica
Full-grown opossum                    Honduras
Caluromys derbianus                   Costa Rica
Didelphis                             Mexico
Didelphis marsupialis                 Costa Rica
Didelphis marsupialis                 Trinidad
Philander                             Guatemala
Philander opossum                     Costa Rica
Rat sized rodent                      Ecuador
Muridae                               Ecuador
Heteromys                             Guatemala
Heteromys desmarestianus              Costa Rica
Melanomys caliginosus                 Costa Rica
Mus musculus                          Costa Rica
Mus musculus                          Mexico
Mus musculus                          Costa Rica
Oligoryzomys fulvescens               Costa Rica
Oryzomys                              Costa Rica
Proechimys semispinosus               Ecuador
Proechimys semispinosus               Costa Rica
Rattus rattus                         Mexico
Rattus rattus                         Mexico
Rattus rattus "probably"              Costa Rica
Sigmodon hirsutus                     Costa Rica
Sigmodon peruanus                     Ecuador
Wood Rat [Neotoma]                    Trinidad
Coendou rothschildi                   Ecuador
Sylvilagus brasiliensis               Costa Rica
Skunks                                Costa Rica
Cryptotis parva                       Mexico

Prey                                           Reference (a)

Invertebrates                         Buttenhoff and Vogt (1997)
Class Chilopoda
Scolopendra                           Boada et al. (2005)
Scolopendra                           Campbell (1998)
Scolopendra                           Greene (1992)
Scolopendra angulata                  Parker (1926)
Class Insecta
Coleoptera                            Boada et al. (2005)
Diptera                               Boada et al. (2005)
Hemiptera                             Boada et al. (2005)
Hymenoptera                           Boada et al. (2005)
Orthoptera                            Boada et al. (2005)
Orthoptera                            Campbell (1998)
Orthoptera                            Sasa et al. (2009)
Class Malacostraca
Crayfish                              Emsley (1977) (1'2'3)
Class Actinopterygii
Synbranchus marmoratus                Hertz et al. (2009)
Class Amphibia
Dermophis parviceps                   Jones et al. (2014)
Anuran                                Logan and Montero (2009)
Acris gryllus                         Burger and Smith (1950)
Craugastor fitzingeri                 Sasa et al. (2009)
Dendrobatid frogs                     Lemos-Espinal and Dixon (2013)
Eleutherodactylus                     Greene (1997)
Eleutherodactylus                     Campbell (1998)
Eleutherodactylus                     Buttenhoff and Vogt (1997)
  [Craugastor] rhodopis
Eleutherodactylus                     Boada et al. (2005)
  [Pristimantis] achatinus
Hyla crucifer                         Burger and Smith (1950)
Leptodactylus bolivianus              Sasa et al. (2009)
Leptodactylus savagei                 Ryan et al. (2010)
Leptodactylus savagei                 Sasa et al. (2009)
Lithobates forreri                    Campbell and Lamar (2004)
Lithobates forreri                    Sasa et al. (2009)
Lithobates vaillanti                  Sasa et al. (2009)
Lithobates warzewitzchi               Sasa et al. (2009)
Mannophryne trinitatis                Mole (1924) (1)
Rhinella marina                       Sasa et al. (2009)
Bufo [Rhinella] marinus               Boada et al. (2005)
Smilisca baudinii                     Buttenhoff and Vogt (1997)
Smilisca phaeota                      Sasa et al. (2009)
Smilisca sordida                      Sasa et al. (2009)
Class Reptilia
Amphisbaena alba                      Hertz et al. (2009)
Ameiva [Holcosus] festiva             Sasa et al. (2009)
Ameiva [Holcosus] undulata            Buttenhoff and Vogt (1997)
Anolis                                Picado (1931)
Anolis                                Picado (1931)
Anolis                                Campbell (1998)
Anolis limifrons                      Sexton and Heatwole (1965)
Ctenosaura similis                    Sasa et al. (2009)
Gonatodes fuscus [albogularis]        Niceforo-Maria (1930) (2,3)
Lepidophyma reticulatum               Sasa et al. (2009)
Lepidophyma tuxtlae                   Urbina-Cardona (2009)
Lygosoma [Sphenomorphus]              Stuart (1948)
Skinks                                Campbell (1998)
Bothrops asper                        Burger and Smith (1950)
Bothrops asper                        Buttenhoff and Vogt (1995)
Bothrops asper                        Buttenhoff and Vogt (1997)
Bothrops asper                        Correa Sanchez et al (1998)
Bothrops asper                        Tryon (1985)
Crotalus horridus                     Burger and Smith (1950)
Erythrolamprus bizona                 Sofirzano (2004)
Ninia atrata                          Murphy (1997)
Ninia sebae                           Sasa et al. (2009)
Storeria dekayi                       Burger and Smith (1950)
Class Aves
Feathers                              Picado (1931)
Feathers                              Barbour and Loveridge
                                        (1929) (2,3)
Unidentified passerines (feathers)    Sasa et al. (2009)
Cantorchilus [Thryothorus]            Boada et al. (2005)
Eucometis penicllata                  Moody (2015)
Troglodytes                           Hertz et al. (2009)
Volantinia jacarina                   Sasa et al. (2009)
Class Mammalia
Hair                                  Sexton and Heatwole (1965)
Mammalian hair                        Kuch et al. (2004)
Small marsupials                      Picado (1931)
Full-grown opossum                    March (1928)
Caluromys derbianus                   Hirth (1964)
Didelphis                             Buttenhoff and Vogt (1997)
Didelphis marsupialis                 Greene and Hardy (1989)
Didelphis marsupialis                 Mole (1924) (1'2'3)
Philander                             Campbell (1998)
Philander opossum                     Sasa et al. (2009)
Rat sized rodent                      Kuch et al. (2004)
Muridae                               Kuch et al. (2004)
Heteromys                             Campbell (1998)
Heteromys desmarestianus              Sasa et al. (2009)
Melanomys caliginosus                 Sasa et al. (2009)
Mus musculus                          Sasa et al. (2009)
Mus musculus                          Correa Sanchez et al (1998)
Mus musculus                          Tryon (1985)
Oligoryzomys fulvescens               Sasa et al. (2009)
Oryzomys                              Sasa et al. (2009)
Proechimys semispinosus               Segovia-Nunez et al. (2014)
Proechimys semispinosus               Sasa et al. (2009)
Rattus rattus                         Buttenhoff and Vogt (1995)
Rattus rattus                         Buttenhoff and Vogt (1997)
Rattus rattus "probably"              Hirth (1964)
Sigmodon hirsutus                     Sasa et al. (2009)
Sigmodon peruanus                     Kuch et al. (2004)
Wood Rat [Neotoma]                    Mole (1924) (1)
Coendou rothschildi                   Segovia-Nunez et al. (2014)
Sylvilagus brasiliensis               Greene and Hardy (1989)
Skunks                                Picado (1931)
Cryptotis parva                       This study

(a) Reference: fide Murphy (1997); fide Campbell and Lamar (2004);
fide Sasa et al. (2009).

(b) Scavenged.

(c) Attempted.
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Title Annotation:Notes
Author:Farr, William L.; Lazcano, David
Publication:Southwestern Naturalist
Article Type:Report
Geographic Code:1MEX
Date:Mar 1, 2017
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