Assessment of the Rarity and Conservation Status of the Colombian Endemic Brown Hairy Dwarf Porcupine Coendou Vestitus

Assessment of the rarity and conservation status of the Colombian endemic brown hairy dwarf porcupine Coendou vestitus

M ARÍA M. TORRES-MARTÍNEZ,HÉCTOR E. RAMÍREZ-CHAVES E LKIN A. NOGUERA-URBANO and F ERNANDO C. PASSOS

Abstract The brown hairy dwarf porcupine Coendou vesti- Introduction tus is a poorly studied Andean species endemic to Colombia. Its current Red List category is contradictory: globally it is he brown hairy dwarf porcupine Coendou vestitus is categorized as Data Deficient but in Colombia it is cate- Tone of the rarest of the seven species of porcupines gorized as Vulnerable. This contradiction has limited the im- (genus Coendou) that occur in Colombia (Ramírez-Chaves  – – plementation of conservation programmes. We evaluate et al., ). It is a small species (head body length  the level of rarity of the species and provide consolidated mm), characterized by having three types of fur: long  information for a new assessment of its Red List status. We dorsal fur, bicolored defensive fur, and bristles (Voss, ). reviewed literature, photographs, and voucher specimens Since its description more than a century ago (Thomas,  in natural history collections. Using the confirmed records, ), it has been recorded from only six localities. we estimated the extent of occurrence (EOO) based on This species is considered endemic to both sides of the the minimum convex polygon and the area of occupancy Eastern Cordillera in the Colombian Andes, which is a (AOO) summing the area of grid squares occupied by the complex ecosystem with topographical and biological di- species. We found that C. vestitus is very rare, with a small versity and high levels of endemism (Olson & Dinerstein,    range, low estimated population density, occurrence in only ; Armenteras et al., ; Sánchez-Cuervo et al., ). one habitat type and small body size. The species has an Andean ecosystems are a global conservation priority as    EOO of , km and an AOO of  km , based on six only % of their original extent remains (Armenteras  confirmed localities, all on the western slopes of the East- et al., ). ern Cordillera, in the central Andean region of Colombia. Although C. vestitus is considered rare (Ramírez-Chaves  Based on the species’ rarity, restricted distribution, and et al., ), this condition has not been evaluated, and it has threats to its natural habitat, we recommend its categoriza- been suggested based only on the absence of data and the tion as Endangered on the IUCN Red List. This will help paucity of voucher specimens. Information on the ecology, delineate research and conservation efforts for this porcu- genetics, natural history and conservation status of the spe- pine, which has a highly restricted range and inhabits the cies is also scarce, and in the case of the latter, contradic- threatened Andean forest. tory. This porcupine is categorized as Data Deficient on the IUCN Red List (Weksler et al., ), with the assessment Keywords Andes, Coendou vestitus, Colombia, Endan- considering the presence of the species from only two local- gered, porcupine, threat category, voucher specimen ities, although in the same assessment three localities were mentioned. Nationally, the species has been categorized as Vulnerable based on its reduced geographical range as a re-

MARÍA M. TORRES-MARTÍNEZ (Corresponding author, orcid.org/0000-0002- sult of habitat loss and fragmentation (Amori & Gippoliti, 5892-0788) Programa de Pós-graduação em Ecologia e Conservação, ; Alberico & Moreno, ; MADS, ). Currently, Laboratório de Biodiversidade, Conservação e Ecologia de Animais Silvestres, Universidade Federal do Paraná, Setor de Ciências Biológicas, Postal box it is the only porcupine species categorized as threatened 19031, CEP 81531-990, Curitiba, Brazil in Colombia (Alberico & Moreno, ). Here, based on E-mail [email protected] available literature, specimens in natural history museums HÉCTOR E. RAMÍREZ-CHAVES ( orcid.org/0000-0002-2454-9482) Departamento and collections, and data from recent records, we evaluate de Ciencias Biológicas, Facultad de Ciencias Exactas y Naturales, and Centro de Museos, Museo de Historia Natural, Universidad de Caldas, Manizales, the level of rarity of the species and reassess its conserva- Colombia tion status.

ELKIN A. NOGUERA-URBANO ( orcid.org/0000-0002-4391-4852) Instituto de Investigación de Recursos Biológicos, Alexander von Humboldt, Bogotá, Colombia Methods FERNANDO C. PASSOS ( orcid.org/0000-0002-8994-3130) Programa de Pós- graduação em Ecologia e Conservação, Laboratório de Biodiversidade, Conservação e Ecologia de Animais Silvestres, Universidade Federal do Assessing rarity Paraná, Curitiba, Brazil Received  May . Revision requested  June . To evaluate the rarity of the species we followed the crite- Accepted  August . First published online  November . ria of Yu & Dobson (), based on four characteristics:

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloadeddistribution, from https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.35.93 the original work, onis 26 properly Sep 2021 cited. at 11:13:47, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2021, 55(5), 765–770 ©. https://doi.org/10.1017/S0030605319001029The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001029 766 M. M. Torres‐Martínez et al.

() local population density, ()range,() the number of habi- overlap of the AOO and EOO with protected areas, we esti-  tat types in which the species occurs and () body size. In mated the extent (km ) of protected areas inside the EOO addition, we suggest factors that may have determined the polygon using as a limit the elevational range of the species, rarity category of this species. As the species population den- and determined the number of confirmed localities inside sity has not been assessed, we documented the number of protected areas. records per year since the species description. We con- sider the population density to be low if the number of records evaluated is ,  for each -year interval Results (there are c. , records of mammals in databases for Rarity Our findings indicate that C. vestitus matches the Cundinamarca, the department in which C. vestitus has criteria of an extremely rare species (Category H; Yu & been historically recorded). We also estimated the species Dobson, ), based on all four factors evaluated. range and related this to the rarity level using the extent of occurrence (EOO) and area of occupancy (AOO). We calculated the EOO using the minimum convex polygon Population density From the description of C. vestitus to (linking the known points of occurrence for the species), the present, we found only  voucher specimens and photo- and the AOO by summing the area of grid squares in graphs of three living specimens. Three records had no  which the species is known (using grid squares of  km as precise locality information. The records date from the recommended in IUCN, )inGeoCAT (Bachman et al., species description in  to photographic records from ). For this we used data from six confirmed localities. Cundinamarca in  (Plate , Table ). The scarcity of re- Three of these are voucher specimens housed at Colombian cords suggests a low population density, considering that collections and the other three are photographic records several biological expeditions have visited the area in (Table ). The photographs showed characters used to differ- which the species occurs (Cundinamarca is the Depart- entiate C. vestitus from other species in this genus: dorsal ment in which Bogotá, the capital of Colombia and home pelage with long blackish fur that partially or completely of the main Colombian academic institutions, is located). conceals defensive quills, and bicoloured bristle-quills (Voss    & da Silva, ; Voss, ; Plate ). We excluded the record Range It has been suggested that the elevational range of  ICN from our analyses because the specimen was C. vestitus is ,–, m (Barthelmess, ), –, m transported from another locality in the western part of the (Alberico et al., )and–, m (Ramírez-Chaves et al., Eastern Cordillera; the location on the label of the specimen ). However, based on the information from confirmed  is in a market area (Alberico & Moreno, ). localities, and discarding a dubious record from Villavicencio, We also considered the number of habitats occupied by the elevational range appears to be from , m (Cundina- the species. For this, we overlaid the EOO on the ecoregions marca, Quipile) to , m (Cundinamarca, Chicaque). The of Colombia, following the classification of terrestrial eco- estimated area of the species range is based on only six con-  regions (Olson et al., ). We evaluated body size based on firmed localities (Table , Fig. ), with an estimated EOO of   information from the labels of the reviewed voucher speci- , km and an AOO of  km . mens and from the literature (Voss, ; Ramírez-Chaves et al., ), and compared this trait with other species of Coendou. Number of habitat types in which the species occurs Overlaying the EOO on the terrestrial ecoregions map indi- cated that the species only occurs in tropical moist broadleaf Conservation status forest (Olson et al., ). This ecoregion corresponds to the To reassess the species conservation status, we used in- highly threatened sub-Andean and Andean forests. formation on the level of rarity, EOO and AOO (IUCN, ). We included the level of rarity and the ecoregions Body size The adult body size of C. vestitus (total length of the species inhabits to infer the conservation status of the head and body – mm) is within the range observed species because of the absence of information on other for small Coendou species (C. insidiosus – mm; C. factors that can influence the AOO, such as biotic interac- nycthemera – mm; C. pruinosus – mm; C. ru- tions (predation, competition) and landscape (connectivity fescens – mm; C. melanurus – mm; C. quichua and shelter) (Bernard et al., ). We also examined whether – mm; C. speratus – mm; C. spinosus – the EOO or AOO of C. vestitus overlaps with protected areas, mm; C. bicolor – mm; C. prehensilis – mm), by using the protected areas layer for Colombia (October being the third smallest species of the genus after C. ichillus ; SINAP, ), and evaluated the per cent of forest (– mm) and C. roosmalenorum ( mm). Coendou area that remained unchanged during – within the vestitus has, however, a considerably shorter tail than the polygon of the species range (IDEAM, ). To evaluate latter species.

Oryx, 2021, 55(5), 765–770 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001029 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 26 Sep 2021 at 11:13:47, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319001029 Brown hairy dwarf porcupine 767

TABLE 1 Records of Coendou vestitus in Colombia. Records from six confirmed localities were used in the analyses; one record (ICN ) was excluded.

Altitude Latitude, Record1 Locality (m) longitude Sources & comments IAvH 7956; MLS 753 Boyacá, Villa de Leyva 2,100 5°37′59″N, Weksler et al. (2016) 73°31′32″W AMNH 70529 Cundinamarca, Quipile 1,250 4°44′50″N, Voss & da Silva (2001), 74°31′59″W Voss (2011, 2015) AMNH 70596, 71359; BMNH Cundinamarca, San Juan 1,440 4°51′4″N, Voss & da Silva (2001), 24.2.21.2; MNHN 1929.631, de Rio Seco 74°38′1″W Voss (2011, 2015) 1929.632; USNM 240035 Photograph Cundinamarca, Bogotá, 2,625 4°36′19.3″N, 7 Oct. 2012, Pedro Pulido Parque Natural Chicaque 74°18′19.8″W Photograph Cundinamarca, Ubaté, 2,556 5°20′14.72″N, 7 Sep. 2018 by a local environmental vereda Volcán 73°49′57.16″W entity (Corporación Autonóma Regional), La Villa (2018) Photograph Cundinamarca, Tena, 2,080 4°40′49″N, 25 June 2018 by Sergio Chaparro Laguna Pedro Palo 74°23′24.3″W Herrera (Plate 1) ICN 3505 Meta, Villavicencio 610 4°10′0″N, Alberico et al. (1999), 73°39′0″W Ramírez-Chaves et al. (2016); label indicates it was kept captive, illegally MLS 249 Cundinamarca No precise locality MLS 640 Cundinamarca No precise locality BMNH 54.6.26.1 Colombia No precise locality Holotype

 IAvH, Instituto de Investigaciones Biológicas Alexander von Humboldt; MLS, Museo de La Salle; AMNH, American Museum of Natural History; BMNH, British Museum of Natural History; MNHN, Muséum National d’Histoire Naturelle; USNM, National Museum of Natural History; ICN, Instituto de Ciencias Naturales, Universidad Nacional de Colombia.

Nacionales de Colombia). Specifically, these  areas in- clude one Soil Conservation District, five Regional Protect- ed Forest Reserves, eight Regional Integrated Management Districts, and  Natural Civil Society Reserves (UNEP– WCMC, ). The portions of the protected areas within the   EOO polygon have a total area of , km , with , km of the EOO not lying within protected areas (Fig. a). When overlapping the forest coverage with the EOO polygon,  the forest coverage during – was  km (.%). Considering the species’ range and its rarity, we recommend that C. vestitus is recategorized from Data Deficient to En- dangered based on the following criteria: ()Bb(iii,iv,v)  and Bb(iii,iv,v): with an EOO of , , km (B) and continuing decline inferred (b) in extent and/or quality of habitat (iii), number of locations or subpopulations (iv) and the number of mature individuals (v), and similarly with an  AOO ,  km (B); ()Cb: population size estimated to number , , mature individuals with a continuing decline, observed, projected, or inferred (C), and, continu- PLATE 1 Photographic record (June )ofCoendou vestitus from ing decline, observed, projected or inferred, in numbers of Pedro Palo, Tena, Colombia (Table ). Photo: Sergio Chaparro. mature individuals, and extreme fluctuations in number of mature individuals (b). Conservation status We found only two localities (of   the six confirmed) within conservation areas ( . %of Discussion the occurrences), although the EOO polygon intersected with  protected areas, managed by two institutions Our recommendation to categorize C. vestitus as Endangered (Corporación Autónoma Regional and Parques Naturales follows IUCN () recommendations to assess poorly

FIG. 1 Extent of occurrence (EOO) of Coendou vestitus, with (a) protected areas within the EOO, indicating the two records in protected areas, and (b) forest coverage in –.

known taxa based on information on inferred habitat loss distributions in the northern part of South America com- and restricted distribution, to avoid assigning a Data De- pared to the larger species that have only one type of quills ficient category. The current IUCN information (Weksler in adulthood (e.g. C. prehensilis and C. bicolor; Voss, ). et al., ) is incomplete (with AOO and EOO unknown) The rarity of C. vestitus is perhaps associated with homo- because it includes data for only from two localities (Voss, plastic functional traits such as the presence of three types ). The periodic re-evaluation of a species’ Red List status of hairs in adulthood (i.e. with less protection against preda- is an important tool for the planning, monitoring and man- tors than species with a body fully-covered by quills) (Voss agement of biodiversity conservation (Hoffmann et al., ; et al., ) and small body size (Gaston & Blackburn, ; Angulo et al., ). Yu & Dobson, ). Several species of African small mam- Besides being an endemic and rare species, our findings mals have been categorized as rare or Vulnerable (Schlitter, confirm that C. vestitus is restricted to an ecosystem in ; Decher, ) because their size could influence preda- which habitat is being lost and there are continuing threats tor–prey relationships (Brose et al., ; Tsai et al., ). from anthropogenic transformation (Thiollay, ;Armen- In this context, additional morphological characters may teras et al., ;Anguloetal.,). The Eastern Cordillera confer an adaptive advantage to large Coendou species: a comprises % of Andean ecosystems, but only %ofitsori- body mostly covered by quills in adulthood provides a pos- ginal vegetation cover remains. Although this region is charac- sible advantage against predators (Scharf et al., ; Speed terized by high species richness and endemism it is one of the & Ruxton, ), the swollen nasofrontal sinuses protect the least known and least protected ecosystems (Armenteras et al., brain, and a larger tail facilitates arboreal locomotion, as ob- ; Rodríguez-Erazo et al., ). Habitat loss affects the per- served in C. bicolor and C. prehensilis (Voss et al., ; Voss, sistence of small mammals, which play important ecological ). The morphological and/or evolutionary reasons for roles, for example as seed dispersers of pioneer species, and the restricted range of small-sized Coendou are, however, in trophic and predator–prey relationships (Decher, ; as yet unclear, and require further research. Brose et al., ;Limaetal.,;Tsaietal.,). Although  genera,  species and  subspecies of ro- Although C. vestitus has previously been considered a dents are considered threatened and have high endemism rare species because of the scarcity of records (Ramírez- (Ceballos & Brown, ; Amori & Gippoliti, , , Chaves et al., ), there are other reasons for it to be con- IUCN, ) a relatively lower per cent of rodents are sidered rare (Cofré & Marquet, ). Although other por- categorized as threatened compared to other mammalian cupines in Colombia are known from fewer specimens (e.g. groups (Ceballos & Brown, ; IUCN, ; Amori & C. ichillus), they are not endemic to the country, having a Gippoliti, ). No Coendou species are as yet categorized wider distribution (Voss, ). Our confirmation of rarity as threatened (IUCN, ): six are categorized as Data is based on a combination of apparent low local population Deficient, including C. vestitus, and eight as Least Concern. density, a small range, occurrence in only one habitat type, The only locations of C. vestitus within protected areas  and small body size. are in the . km Nature Reserve Parque Natural  In general, small-sized species of Coendou (e.g. C. ichil- Chicaque and the . km Natural Civil Society Reserve lus, C. roosmalenorum and C. vestitus) have more restricted Tenasuca de Pedro Palo, which together correspond to

only .% of the species’ range (Fig. a). In addition to the Author contributions Project design: MMT-M, HER-C; data collec- small EOO, the area of the  protected areas within the tion: MMT-M, HER-C, EAN-U; evaluation of rarity and threatened  EOO is small (a mean of  km per protected area). The status: MMT-M, FCP; writing: MMT-M, HER-C. area surrounding the EOO is severely affected by extensive Conflicts of interest None. commercial plantations and urban settlements, with only c. % of the ecoregion unaffected (Armenteras et al., ; Ethical standards This research abided by the Oryx guidelines on ethical standards. Sánchez-Cuervo et al., ; Angulo et al., ). This limits connectivity (Bright, ), which is important for the persist- ence of a species (Passos et al., ). Maintenance, exten- References sion or connection of protected areas, connecting the relict ALBERICO,M.&MORENO, J.G. () Puerco espín pardo Coendou habitats, could help to protect C. vestitus (Cofré & Marquet, vestitus.InLibro Rojo de los Mamíferos de Colombia. Serie Libros ;Amori&Gippoliti,; Armenteras et al., ). Rojos de Especies Amenazadas de Colombia (eds J.V. Rodríguez- Conservation strategies and financial resources need to Mahecha, M. Alberico, F. Trujillo & J. Jorgenson), pp. –. be established for threatened and endemic species and for Conservación Internacional Colombia, Ministerio de Ambiente, Vivienda y Desarrollo Territorial, Bogotá, Colombia. species with restricted distributions (Cofré & Marquet, ; ALBERICO, M., ROJAS-DÍAZ,V.&MORENO, J.G. () Aporte sobre  Isaac et al., ). However, for mammals most monitor- la taxonomía y distribución de los puercosespines (Rodentia: ing and conservation efforts are directed at large or cha- Erethizontidae) en Colombia. Revista de la Academia Colombiana rismatic species. Less attention has been directed at rodents, de Ciencias Exactas, Físicas y Naturales, , –.  even though the group has a high extinction rate (Amori & AMORI,G.&GIPPOLITI,S.( ) Identifying priority ecoregions for rodent conservation at the genus level. Oryx, , –. Gippoliti, , ;Armenterasetal.,). Prioritizing AMORI,G.&GIPPOLITI,S.() A higher-taxon approach to rodent the conservation of C. vestitus has the potential to contribute conservation priorities for the st century. Animal Biodiversity and to the protection of the ecosystems in which it occurs and of Conservation, , –. co-occurring species. We recommend that national agencies ANGULO, A., VON MAY,R.&ICOCHEA,J.() A reassessment of the prioritize this porcupine species, together with other species extinction risk of the Critically Endangered Oxapampa poison frog  – in urgent need of monitoring. Ameerega planipaleae (Dendrobatidae). Oryx, , . ARMENTERAS, D., GAST,F.&VILLAREAL,H.() Andean forest Porcupines remain a poorly known group, both at nation- fragmentation and the representativeness of protected natural areas al and Neotropical levels (Voss, ). Knowledge of the ecol- in the eastern Andes, Colombia. Biological Conservation, , –. ogy of C. vestitus is mostly based on inference from other BACHMAN, S., MOAT, J., HILL, A.W., DE LA TORRE,J.&SCOTT,B. porcupine species (e.g. Alberico & Moreno, ;Weksler () Supporting Red List threat assessments with GeoCAT:  – et al., ). Our compilation of information on C. vestitus geospatial conservation assessment tool. ZooKeys, , . BARTHELMESS, E.L. () Family Erethizontidae. In Handbook of highlights the need for further fieldwork and data collection. Mammals of the World. Vol. . Lagomorphs and Rodents: Part  Nevertheless, threats to porcupines are evident, in particular (eds D.E. Wilson, T.E. Lacher & R.A. Mittermeier), pp. –. loss of habitat, illegal trade, road-kills, and hunting for con- Lynx, Barcelona, Spain. sumption (de Freitas et al., ; Racero-Casarrubia et al., BERNARD, E., NASCIMENTO, J.L. & AGUIAR, LM.S. () Flagging a ). In Colombia, illegal captivity has also been documen- species as threatened: the case of Eptesicus taddeii, an endemic bat from the Brazilian Atlantic Forest. Biota Neotropical, , –. ted (on a voucher specimen label; Table ) as a threat to the BRIGHT,P.W.() Habitat fragmentation-problems and predictions species. Our compilation of data and our findings form the for British mammals. Mammal Review, , –. basis for further research and for the establishment of con- BROSE, U., JONSSON, T., BERLOW, E.L., WARREN, P., BANASEK- servation strategies and future evaluations of the distribution RICHTER,C.,BERSIER,L.,BLANCHARD,J.L.etal.() Consumer– and conservation status of C. vestitus. resource body-size relationships in natural food webs. Ecology, , –. Acknowledgements We thank Hugo López (Instituto de Ciencias CEBALLOS,G.&BROWN, J.H. () Global patterns of mammalian Naturales, Universidad Nacional de Colombia) and Cristian Cruz diversity, endemism, and endangerment. Conservation Biology, Rodríguez (Museo de La Salle) for allowing us to review specimens , –. under their care; Luiz H. Varzinczak for useful comments on the COFRÉ,H.&MARQUET, P.A. () Conservation status, rarity, and text; Pedro Pulido, Sergio Chaparro and Carlos Aya for sharing records geographic priorities for conservation of Chilean mammals: and photographs; and Mario Ernesto Jijón Palma for help with the an assessment. Biological Conservation, , –. map. MMTM thanks Coordenação de Aperfeiçoamento de Pessoal DECHER,J.() Conservation, small mammals, and the future of de Nível Superior, Brazil (Finance Code 001) for support. HERC sacred Groves in West Africa. Biodiversity and Conservation, thanks Instituto de Investigación de Recursos Biológicos Alexander , –. von Humboldt (Andrés M. Cuervo and María López Rodríguez) for DE FREITAS, M.A., DE FRANÇA, D.P.F. & VERÍSSIMO,D.() First access to voucher specimens, and The Rufford Small Grants (Grants record of the bicoloured-spined porcupine Coendou bicolor 23710-1 and 29491-2), the Universidad de Caldas (projects 0743919 (Tschudi, ) for Brazil. Check List (Luis Felipe Toledo), , –. and 0223418), and the Science and Scholarship Committee of the GASTON, K.J. & BLACKBURN, T.M. () Rarity and body size: some Field Museum of Natural History, Chicago, for support. FCP thanks cautionary remarks. Conservation Biology, , –. the Brazilian National Council for Scientific and Technological HOFFMANN, M., HILTON-TAYLOR, C., ANGULO, A., BÖHM, M., Development (CNPq) (Grant 307303/2017-9). BROOKS, T.M., BUTCHART, S.H.M. et al. () The impact of

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