Siphonops Annulatus (Mikan, 1820) (Amphibia: Gymnophiona: Siphonopidae) in the State Of

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n Siphonops annulatus (Mikan, 1820) (Amphibia: Gymnophiona: Siphonopidae) in the state of

istributio Pará,Rediscovery Brazil, withof an updated geographic distribution map, D and notes on size and variation raphic g

eo Adriano Oliveira Maciel 1*, Henrique Caldeira Costa 2 , Jerriane G n Oliveira Gomes 1 1 3 o , Leandro de Oliveira Drummond

otes and Annelise D’Angiolella N 1 Museu Paraense Emílio Goeldi, Departamento de Zoologia. Avenida Perimetral, 1901 - Terra Firme. CEP 66077-530. Belém, PA, Brazil. * 2 CorrUniversidadeesponding Federal author. de E-mail: Viçosa [email protected] – Campus Florestal, Instituto de Ciências Biológicas. CEP 35690-000. Florestal, MG, Brazil. 3 Autonomous Researcher. Rua Chaves Faria, 534/202, Bairro São Cristovão. CEP CEP 20910-140. Rio de Janeiro RJ, Brazil.

Abstract: Siphonops annulatus has a wide distribution in South America. Here we provide a new geographic distribution map for this species and two new records from the state of Pará, Brazil, from where it has not been reported since 1876. A S. annulatus ever recorded.

specimen collected in the municipality of Senador José Porfírio is the largest specimen of

Caecilian amphibians (Gymnophiona) are distributed annulatus in Pará. All specimens were collected in mainly across the wet tropics, but also occur in subtropical regions (Gower and Wilkinson 2005). The fossorial, semiaquatic or aquatic habit of the known species hamper pitfall traps (collection permit numbers: 01/11; Process the collection of specimens, although recent data shows 02018.001265/2010-41-NUFAP/IBAMA and 1325/2011 that some species are much more abundant than has been SEMA/PA), and deposited in the collection of the Museu assumed (Maciel and Hoogmoed 2011). 220Paraense mm total Emílio length) Goeldi, were Belém. collected Two in specimensthe municipality (male Siphonops annulatus (Mikan, 1820) is the caecilian MPEG 33733, 282 mm total length; male MPEG 33734, with the largest distribution (Wilkinson et al. 2008), occurring throughout most of Cis-Andean South America. of Itaituba (06°19’12” S, 55°47’24” W, elevation 270 m) Although commonly documented in several localities over and a single specimen (Figure 1) in the municipality of its distribution, this species was not found in the state Senador José Porfírio (male, MPEG 32559; 03°35’19” of Pará since 1876 when Spengel received in Hamburg, S, 51°57’00” W, elevation 165 m). TheS. annulatus latter specimen (Taylor Germany, some specimens of S. annulatus from “Pará”, as 1968).measures 539 mm total length, greater than the previous the current municipality of Belém was named at that time reported maximum size of 450 mm for (Spengel 1915). Herein, we report two new localities for Siphonops Both specimens from Itaituba haveS. annulatus 79 annuli (Taylor and the specimen from Senador José Porfírio has 91 annuli. This character ranges from 78–98 in

Figure 1. Specimens of Siphonops annulatus

from Pará, Brazil. Left: MPEG 33734, 220 mm total length, municipality of Itaituba (photo by AOM); Right: MPEG 32559, 539 mm total length, municipality of Senador José Porfírio (Photo by LOD). 106 Maciel et al. | Geographic distribution map of Siphonops annulatus

1968), and presents a low geographic variation, even in a et al. (2006) present a photograph of two caecilians from ca. referred as S. annulatus 4000 km long distribution range; a similar condition Reservato be a Adolpho member Ducke, of the state genus of CaeciliaAmazonas,, not one Siphonops of them. Hoogmoedis observed 2011). in other Neotropical caecilian species with Thus, to the best of our; theknowledge, specimen, the however, occurrence appears of S. wideMaciel distribution and Hoogmoed (Savage (2011)and Wake present 2001; a Maciel geographic and annulatus distribution map for Siphonops annulatus, which lacks The absencein Ceará, of records and in of theSiphonops Reserva annulatus Adolpho from Ducke, the with locality descriptions and coordinates in Appendix largeremains open unconfirmed. vegetation diagonal of Brazil, composed of the 1.some Two literature literature records. records An were updated not considered map is in Figure by us: 2, i) Caatinga, Cerrado, and Pantanal-Chaco morphoclimatic

This distribution pattern can be explained by the lack of Borges-Nojosa and Cascon (2005) presented a record collectionsdomains or inbiomes those (sensu regions Ab’Sáber or, more 1977), likely, is bynoteworthy. historical from Reserva Natural Serra das Almas, in the Brazilian causes, but only a phylogeographic study of the species state of Ceará; however, when visitingS. annulatus the collection. ii) Lima of will elucidate this question. Universidade Federal do Ceará, AOM did not find any specimen that could be identified as

Figure 2. Geographic distribution map of Siphonops annulatus

. Circles = literature records. Triangle 1 = Senador José Porfírio, Pará, Brazil; triangle 2 = AcknowledgmentsItaituba, Pará, Brazil. :The record from Alagoas, although shown on the map (Square), lacks specific localityAnálise data (see das Table Variações 1). da Biodiversidade do who provided valuable suggestions on the manuscript. Bioma Caatinga We thank Ulisses Caramaschii and Ross MacCulloch M.R.V. Barbosa (org.). Herpetofauna dos Literature Cited . Brasília: Ministério do Meio Ambiente. Condez,sudeste T.H., do Brasil. R.J. Sawaya Biota Neotropica and M. Dixo.9(1): 157-185. 2009. Geomorfologia 52: 1-21, 1 map. remanescentes de Mata Atlântica da região de Tapiraí e Piedade, SP, Ab’Sáber, A.N., 1977. Os domínios morfoclimáticos na América do Sul. Bollettino del Museo regionale di Primeira aproximação. De laScienze Riva, I. naturali, 1990. Lista Torino preliminar comentada de los anfibios de Bolivia, Araujo, C.O., T.H. Condez, R.P. Bovo, F.C. Centeno and A.M. Luiz. 2010. con datos sobre su distribución. 2000. Ten years of Brazil.Amphibians Biota Neotropica and reptiles of the Parque Estadual Turístico do Alto research on Bolivian amphibians: 8(1): 261-319. updated checklist, distribution, Ribeira (PETAR), SP: an Atlantic Forest remnant of Southeastern De lataxonomic Riva, I., problems, J. Köhler, S.literature Lötters and and iconography. S. Reichle. Revista Española de 10(4): 257-274. In Herpetologia Barrio-Amorós, C.L. and C.M. Rodríguez. 2010. Herpetofauna del Ramal de Calderas, Andes de Venezuela; p. 74-80RAP A. Bulletin Rial, J.C. ofSeñaris, Biological C.A. Composition 14: of 19-164. the Herpetofaunal Communities of Tambopata AssessmentLasso and A.L. Flores (ed.). Rapid assessment of the biodiversity of Doan, T.M. and W.A.Biotropica Arriaga. 2002. Microgeographic Variation in Species the Ramal Calderas, Venezuelan Andes. Montingelli. 56: 1-183. AmazonianRegion, Peru. Ecuador. The 34(1): University 101-117. of Kansas Museum of Natural Bertoluci,fragmento J., M.A.S. de Mata Canelas, Atlântica C.C. do Eisemberg, estado de Minas C.F.S. Gerais, Palmuti sudeste and G.G. do Duellman,History, W.E. Miscellaneous 1978. The Publications Biology of an Equatorial Herpetofauna in Brasil. Biota Neotropica2009. Herpetofauna da Estação Ambiental de Peti, um Bulletin of the Museum of Comparative Zoology 65: 1-352. 9(1): 147-155.In Dunn, E.R. 1942. The American caecilians. Borges-Nojosa, D.M. and P. Cascon. 2005. Herpetofauna da Área Reserva 91(6): 439-540. da Serra das Almas, Ceará; p. 245-260 F.S Araújo, M.J.N Rodal and Faria, H.A.B. and T. Mott. 2011. Geographic distribution of caecilians 107 Maciel et al. | Geographic distribution map of Siphonops annulatus

(Gymnophiona, Amphibia) in the state of Mato Grosso, Brazil with Herpetology Notes a new state record for Caecilia mertensi Taylor 1973. 4: 053-056. Revista Feio,Ceres R.N. and U. Caramaschi. 1995. Aspectos zoogeográfiicos dos Anfíbios do Médio Rio Jequitinhonha, nordeste de Minas2010. Gerais, Amphibians Brasil. of 42(239): 53-61. Ferreira,Salamandra R.B., T. Silva-Soares and D. Rödder. Vitória, an urban area in south-easternGuia Ilustrado: Brazil: first A herpetofauna approximation. das REFERENCE

Caatingas e Áreas46(4): de 187-196. Altitudes do Nordeste Brasileiro. Pelotas: USEB. et al. 2008 Freitas, M.A. and T.F.S. Silva. 2007. et al. 2006 388 p. Revista Dunn 1942 Taylor 1968 Taylor Dunn 1942 Dunn 1942; De la Riva 1990 Dunn 1942; De la Riva 1990 1968; De la Riva Dunn 1942; Taylor Dunn 1942 Silva Silva Tonini et al. 2010 Tonini Maciel and Hoogmoed 2011 1968 Dunn 1942; Taylor Ferreira et al. 2010 Ferreira Dunn 1942 Zhang and Wake 2009 Zhang and Wake González Fernández 2006; González Fernández Faria and Mott 2011 Faria Freitas and Silva 2007 and Silva Freitas Gayer,Brasileira S.M.P., L. de Krause Zoologia and N. Gomes. 1988. Lista preliminar dos anf(bios Wilkinson da Estação Ecológica do Taim, Rio Grande do Sul, Brasil. 5(3): 419-425. González Fernández, J.E. 2006. Anfibios colectadosGraellsia por62(1): la 111-158. Comisión

Científica del Pacífico (entre 1862 y 1865) conservados en el Museo DMS

amphibians.Nacional de CienciasConservation Naturales Biology de Madrid. LONGITUDE Not specified 55°20’ W 40°22’12” W 66°06’ W 67°13’ W 40°36’00” W Not specified 40°32’07” W 66°04’08” W 64°42’ W 40°20’16” W 69°56’16” W 40°39’32” W 38°40’ W 53°27’50” W 38°30’ W Gower, D.J. and M. Wilkinson. 2005. Conservation biology of caecilian 39°02’56” W 19: 45-55. Hedges, S.B., R.A. Nussbaum and L.R. Maxson. 1993. Cecilian Phylogeny Herpetologicaland biogeography monographs inferred from mithocondrial DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia: Gymnophiona). DMS LATITUDE Not specified 27°15’ S 19°49’55” S 10°59’ S 15°12’ S 19°56’09” S Not specified 20°14’04” S 03°28’51” S 03°30’ S 20°19’08” S 04°15’10” S 20°21’46” S 12°53’ S 10°33’48” S 12°58’ S 7: 64-76. Herpetological Review 14°47’20” S Lehr, E. 2001. New Records for Amphibians and Reptiles from Departamentos Pasco and Ucayali, Peru. 32(2): 130-132. Lima, A.P., W.E. Magnusson, M. Menin, L.K. Erdtmann, D.J. Rodrigues, C. DECIMAL LONGITUDE Not specified Not -55.33 -40.37 -67.22 -66.10 -40.60 Not specified Not -40.54 -64.70 -66.07 -40.34 -69.94 -40.66 -38.67 -53.46 -38.51 Keller and W. Hödl. 2006. Guia de Sapos da Reserva Adolpho Ducke, -39.04 (Amphibia:Amazônia Central. Gymnophiona). Manaus. Átemma Revista Design de la Academia Editorial. Colombiana168 p. de Lynch,Ciencias J.D. 1999. Exactas, Una Físicas approximacion y Naturales a las cuebras ciegas de Colombia

eastern Colombia. Caldasia 23: 317-337. DECIMAL LATITUDE Not specified Not -27.25 -19.83 -15.20 -10.98 -19.94 Not specified Not -20.23 -3.50 -3.48 -20.32 -4.25 -20.36 -12.88 -10.56 -12.97 Lynch,Maciel, J.D. A.O. 2006. and M.S. The Hoogmoed. Amphibian 2011. fauna Taxonomy in the Villavicencio and distribution region of -14.79 caecilian amphibians (Gymnophiona) 28(1): 135-155. of Brazilian Amazonia, with a Zootaxa

key to their identification. 2984: 1-53. In

Malagoli, L.R. 2008. Anfíbios do município de São Além Paulo: do histórico,concreto: COUNTRY Brazil Argentina Brazil Bolivia Bolivia Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil contribuiçõesconhecimento para atual a e proteçãodesafios para da biodiversidade a conservação; paulistana p. 206-233. Brazil L.R. Malagoli, F.B. Bajesteiro and M. Whately (ed.). Savage, J.M. and M.H. Wake. 2001. . São AmericanPaulo: Instituto caecilians Socioambiental. (Amphibia: Gymnophiona), with comments on their origin and evolution. CopeiaReevaluation of the status of Central

2001: 52-64. / STATE Schlüter,ecological A., J. Icocheaand biogeographical and J.M. Perez. notes. 2004. Salamandra Amphibians and reptiles of DEPARTMENT Espírito Santo Espírito Missiones Espírito Santo Espírito La Paz El Beni Espírito Santo Espírito Alagoas Espírito Santo Espírito Amazonas Amazonas Espírito Santo Espírito Amazonas Espírito Santo Espírito Bahia Mato Grosso Mato Bahia the lower Río Llullapichis, Amazonian Peru: updated species list with Bahia 40(2): 141-160. Silva,65-76 S.T., In U.G. Silva, G.A.B. SenaA Mata and Atlântica F.A.C. Nascimento. em Alagoas 2006. A biodiversidade da Mata Atlântica alagoana: anfíbios e répteis; p. F.B.P. Moura (org.). Blätter. Maceió: für EDUFAL.Aquarien 88und p. Terrarienkunde 26: 220. Spengel,Taylor, E.H. F.W. 1968.1915. BrieflicheThe Caecilians Mitteilung of the an World.den Herausgeber. A taxonomic review.

Lawrence: University of Kansas Press. 848 p. Tonini, J.F.R., L.M. Carão, I.S. Pinto, J.L. Gasparini, Y.L.R. Leite and L.P. Costa. 2010. Non-volant tetrapods from Reserva Biológica de Duas Bocas,State of Espírito Santo, Southeastern Brazil. Biota Neotropica / CITY MUNICIPALITY Jatiun10(3): Sacha339-351. in the lowland rainforest of Amazonian Ecuador: a 20- new records. data and two , based on literature Vigle, G.O. 2008. TheBreviora Amphibians and Reptiles of the Estacíon Biológica San Ignacio

Ibiraçu [Pau Gigante] [Pau Ibiraçu Riberalta Santa Teresa Cariacica Juruá Vitória Tabatinga Domingos Martins Itaparica Parque do Xingu Parque Salvador Ilhéus Year Record. 513: 1-30. Von May, R., K. Siu-Ting, J.M. Jacobs, M.M. Müller, G. Gagliardi, HerpetologicalL.O. Rodríguez Conservationand M.A. Donnelly. and Biology 2009 Species Diversity and Conservation status of Amphibians in Madre de Dios, southern Peru. and C. Jared. 2008. One 4(1): hundred 14-29. million years of skin feeding? Wilkinson, M., A. Kupfer, R. Marques-Porto, H. Jeffkins, M.M. Antoniazzi Gymnophiona). Biology Letters Extended parental care in a Neotropical caecilian (Amphibia: phylogeny and biogeography 4: 358-361. of living caecilians (Amphibia: Zhang,Gymnophiona). P. and M.H. Molecular Wake. 2009. Philogenetics A mitogenomic and Evolution perspective on the LOCALITY 53: 479-491. Siphonops annulatus of Siphonops . Locality records : August 2012

Received Upper Rio Beni Reserva Biológica Duas Bocas Reserva Lagoa Japaranão, near Tefé Lagoa Japaranão, Isla Itaparica Accepted: January 2013 Rio Doce Published online: March 2013 1 Appendix Editorial responsibility: Ross MacCulloch 108 Maciel et al. | Geographic distribution map of Siphonops annulatus REFERENCE et al. 2009 et al. 2009 et al. 1993 et al. 2010 et al. 1988 Lynch 1999 Lynch Lynch 1999 Lynch Faria and Mott 2011 Faria Dunn 1942 Maciel and Hoogmoed 2011 Lynch 1999 Lynch Bertoluci Bertoluci Maciel and Hoogmoed 2011 Taylor 1968; Lynch 1999 1968; Lynch Taylor 1999 Lynch Feio and Caramaschi 1995 and Caramaschi Feio Dunn 1942 Hedges Spengel 1915 Dunn 1942 Duellman 1978 2008 Vigle This study Dunn 1942 Dunn 1942 Dunn 1942 This study Malagoli 2008 Dunn 1942 Condez Dunn 1942 Araujo Araujo Dunn 1942; Taylor 1968 Dunn 1942; Taylor Lynch 1999 Lynch Dunn 1942 Lynch 1999 Lynch Taylor 1968 Taylor Lynch 1999 Lynch Dunn 1942 Dunn 1942 Gayer Gayer Dunn 1942; Taylor 1968; Lynch 1999 1968; Lynch Dunn 1942; Taylor Lynch 1999 Lynch Lynch 2006 Lynch DMS LONGITUDE 74°02’ W 57°59’33” W Not found 65°20’20” W 73°55’ W 43°21’ W 61°00’46” W 76°31’ W 68°28’ W 42°36’ W 48°50’45” W 77°46’ W 48°30’14” W 47°24’03” W 76°59’ W 77°37’ W 51°57’00” W 45°33’18” W 77°29’ W 77°38’ W 55°47’24” W 46°38’09” W 42°31’51” W 47°20’ W 41°47’13” W 48°27’ W 42°57’57” W 69°57’ W 43°10’44” W 69°44 ’ W 43°12’28” W 72°33’ W 52°20’34” W 73°04’ W 52°34’37” W 73°37’ W 73°15’ W 73°46’ W 73°51’ W DMS LATITUDE 03°22’ N 13°42’36” S Not found 10°46’58” S 02°45’ N 19°52’ S 11°31’30” S 00°30’ N 05°19’ N 17°12’ S 26°18’14” S 00°59’ S 01°27’21” S 20°32’20” S 00°03’ N 01°04’ S 03°35’19” S 23°01’33” S 01°44’ S 02°01’ S 06°19’12” S 23°32’52” S 22°16’55” S 23°49’ S 22°22’15” S 24°17’ S 22°24’43” S 04°09’ S 22°30’18” S 02°52’ S 22°54’10” S 05°12’ N 31°46’19” S 04°40’ N 32°43’22” S 04°09’ N 04°08’ N 03°59’ N 03°47’ N DECIMAL LONGITUDE -74.04 -57.99 Not found -65.33 -73.92 -43.35 -61.00 -68.47 -76.52 -42.61 -48.83 -77.82 -48.50 -47.40 -77.62 -76.98 -51.95 -45.56 -77.63 -77.48 -55.79 -46.64 -42.52 -47.33 -41.78 -48.45 -42.95 -69.95 -43.17 -69.73 -43.21 -72.55 -52.34 -73.07 -52.58 -73.62 -73.25 -73.77 -73.85 DECIMAL LATITUDE 3.38 -13.71 Not found -10.78 2.75 -19.87 -11.53 5.32 0.50 -17.20 -26.30 -0.99 -1.46 -20.54 -1.07 0.05 -3.59 -23.03 -2.01 -1.73 -6.32 -23.55 -22.28 -23.82 -22.37 -24.28 -22.41 -4.15 -22.51 -2.87 -22.90 5.20 -31.77 4.67 -32.72 4.15 4.13 3.98 3.78 COUNTRY Colombia Brazil Brazil Brazil Colombia Brazil Brazil Colombia Colombia Brazil Brazil Ecuador Brazil Brazil Ecuador Ecuador Brazil Brazil Ecuador Ecuador Brazil Brazil Brazil Brazil Brazil Brazil Brazil Colombia Brazil Colombia Brazil Colombia Brazil Colombia Brazil Colombia Colombia Colombia Colombia STATE / STATE DEPARTMENT Meta Minas Gerais Mato Grosso Mato Rondônia Meta Minas Gerais Rondônia Vichada Putamayo Minas Gerais Santa Catarina Napo Pará São Paulo Napo Napo Pará São Paulo Pastaza Pará São Paulo Rio de Janeiro São Paulo Rio de Janeiro São Paulo Rio de Janeiro Amazonas Rio de Janeiro Amazonas Rio de Janeiro Casanare Rio Grande do Sul Rio Grande Cundinamarca Rio Grande do Sul Rio Grande Meta Meta Meta Meta MUNICIPALITY / CITY MUNICIPALITY Mesetas Campo Novo dos Parecis Campo Novo Guajará-Mirim La Maracena São Gonçalo do Rio Abaixo and Santa Bárbara Santa and Abaixo Rio do Gonçalo São Espigão d’Oeste Espigão Puerto Asís Puerto Minas Novas Joinville Tena Belém Franca Santa Cecilia Senador José Porfírio Taubaté Sarayacu Itaituba São Paulo Nova Friburgo Nova Tapiraí and Piedade Tapiraí Macaé Apiaí and Iopranga Teresópolis Leticia Petrópolis Tarapacá Rio de Janeiro Aguazul Pelotas Paratebueno Rio Grande and Santa Vitória do Palmar do Palmar and Santa Vitória Rio Grande Villavicencio ca. 5 km E. Villavicencio Acacias Cubarral LOCALITY . Continued. Mendez, on Rio Jequitinhonha Estação Ambiental de Peti Parque Nacional Natural El Tuparro Nacional Natural Parque Estacíon Biológica Jatun Sasha Rio Pastaza Serra de Macaé Serra Parque Estadual Turístico do Alto Ribeira (PETAR) Ribeira Alto do Turístico Estadual Parque Guaicaramo Estação Ecológica de Taim Estação Ecológica Río Ocoa Appendix 1 Appendix

109 Maciel et al. | Geographic distribution map of Siphonops annulatus REFERENCE et al. 2000 Dunn 1942 Dunn 1942 Dunn 1942 Dunn 1942 et al. 2004 Schlüter Barrío-Amorós and Rodríguez 2010 and Rodríguez Barrío-Amorós Taylor 1968 Taylor Dunn 1942 Dunn 1942 Dunn 1942 Dunn 1942 Dunn 1942 et al. 2009 May Von De la Riva De la Riva Doan and Arriaga 2002 Doan and Arriaga Lehr 2001 DMS LONGITUDE 76°58’ W 52°19’ W 77°38’ W 78°12’ W 74°57’ W 70°25’ W 76°14’ W 70°12’ W 73°12’ W 73°14’ W 75°18’ W 74°54’ W 70°06’ W 69°11’ W 69°37’02” W 75°32’11” W DMS LATITUDE 06°03’ S 04°55’ N 04°27’ S 04°35’ S 09°37’ S 08°45’ N 09°55’ S 08°38’ N 03°43’ S 03°47’ S 07°10’ S 07°15’ S 12°34’ S 12°36’ S 13°08’43” S 10°07’01” S DECIMAL LONGITUDE -76.97 -77.63 -78.20 -74.95 -52.32 -70.42 -76.23 -70.20 -73.20 -73.23 -75.30 -70.10 -74.90 -69.18 -69.62 -75.53 DECIMAL LATITUDE -4.45 -4.58 -9.62 4.92 8.75 -9.92 8.63 -3.72 -3.78 -7.17 -12.57 -7.25 -12.60 -13.15 -10.11 -6.05 COUNTRY Peru Peru Peru French Guiana French Venezuela Peru Venezuela Peru Peru Peru Peru Peru Peru Peru Peru Peru STATE / STATE DEPARTMENT Amazonas Amazonas Huanuco Barinas Huanuco Barina [Zamora] Loreto Loreto Loreto Madre de Dios Madre Loreto Madre de Dios Madre Madre de Dios Madre Pasco San Martín MUNICIPALITY / CITY MUNICIPALITY Cayenne Barinitas Huanuco Barinas Iquitos San Antonio Ucayali Ucayali Puerto Maldonado Puerto Tambopata Oxapampa Moyobamba [Moyabamba] Moyobamba LOCALITY . Continued. Mouth of Santiago, upper Maranon Rio Cenipa, upper Maranon Río Llullapichis lower Rio Itaya Pampa Pampa Hermosa, middle Cushatabay Ucayali, mouth of East of Contamana [Contamna], Brazil frontier East of Contamana [Contamna], Brazil Los Amigos Research Center Los Amigos Research Tambopata Research Center Research Tambopata Pozuzo Appendix 1 Appendix

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