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

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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
                                                               MARÍA M. TORRES-MARTÍNEZ, HÉCTOR E. RAMÍREZ-CHAVES
                                                                    E L K I N A . N O G U E R A - U R B A N O and F E R N A N D O C . P A S S O S

               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
               categorized as Data Deficient but in Colombia it is cate-                                      T     he brown hairy dwarf porcupine Coendou vestitus is
                                                                                                                    one of the rarest of the seven species of porcupines
                                                                                                              (genus Coendou) that occur in Colombia (Ramírez-Chaves
               gorized as Vulnerable. This contradiction has limited the im-
               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 canasmarianita@gmail.com
                                                                                                              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:

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                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
Assessment of the rarity and conservation status of the Colombian endemic brown hairy dwarf porcupine Coendou vestitus
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, ) in GeoCAT (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
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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 con-
                                                                                                      tinuing decline inferred (b) in extent and/or quality of habi-
                                                                                                      tat (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 ) of Coendou 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

              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
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768         M. M. Torres‐Martínez et al.

                                                                                                                                                   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., ; Angulo et al., ). The Eastern Cordillera                        confer an adaptive advantage to large Coendou species: a
                comprises % of Andean ecosystems, but only % of its ori-                            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., ; Lima et al., ; Tsai et al., ).                                  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

                            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: 46.4.80.155, on 05 Dec 2021 at 01:48:27, 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                   769

              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
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              ; 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. –.
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                                                                                                         Vivienda y Desarrollo Territorial, Bogotá, Colombia.
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              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
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                                                                                                         rodent conservation at the genus level. Oryx, , –.
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                                                                                                      A R M E N T E R A S , D., G A S T , F. & V I L L A R E A L , H. () Andean forest
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              al and Neotropical levels (Voss, ). Knowledge of the ecol-                             in the eastern Andes, Colombia. Biological Conservation, , –.
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              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.,                            B E R N A R D , E., N A S C I M E N T O , J.L. & A G U I A R , 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                           B R I G H T , P.W. () Habitat fragmentation-problems and predictions
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              Acknowledgements We thank Hugo López (Instituto de Ciencias                             C E B A L LO S , G. & B R O W N , 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                         C O F R É , H. & M A R Q U E T , 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                              D E C H E R , 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                           D E F R E I TA S , M.A., D E F R A N Ç A , D.P.F. & V E R Í S S I M O , 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                          G A S T O N , K.J. & B L AC K B U R N , 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                         H O F F M A N N , M., H I LT O N -T AY LO R , C., A N G U LO , A., B Ö H M , M.,
              Development (CNPq) (Grant 307303/2017-9).                                                  B R O O K S , T.M., B U T C H A R T , S.H.M. et al. () The impact of

              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
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                             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
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