A Study of the Giant Armadillo Priodontes Maximus in Central Brazil

Surveying in highly-modified landscapes to document the occurrence of threatened species: a study of the giant armadillo Priodontes maximus in central Brazil

F REDERICO G. LEMOS,ALAN N. COSTA,FERNANDA C. AZEVEDO C ARLOS E. FRAGOSO,MOZART C. FREITAS-JUNIOR and E DNALDO C. ROCHA

Abstract Studies on threatened species in highly modified Supplementary material for this article is available at and unprotected landscapes are necessary for the develop- https://doi.org/./S ment of appropriate conservation policies. This is particu- larly important for species with large home ranges, such as the giant armadillo Priodontes maximus, whose occur- Introduction rence in anthropogenic landscapes is poorly known despite its categorization as Vulnerable on the IUCN Red List. We ata on the demography, ecological interactions and en- searched and surveyed for the giant armadillo within Dvironmental requirements of Neotropical wildlife, and human-modified areas in central Brazil using direct and in- anthropogenic threats, have been used in the management direct methods across a wide region dominated by diverse of several species (e.g. Sanderson et al., ; Medici et al., farming environments and scattered remnants of natural ; ICMBio, b). Although data accuracy and general- vegetation. During a -year period (–), we located ity are paramount for conservation and management, most  records of the species, including three road-kills and two studies are context-dependent, concentrated either in re- instances of poaching. Most of the occurrence points (%) gions dominated by pristine ecosystems or in medium to were in native vegetation, with % in anthropogenic envi- large protected areas (Fazey et al., ). For threatened spe- ronments (pastures and roads). We confirmed the presence cies, however, studies across landscapes modified and man- of the giant armadillo within a wide, intensely human- aged by humans are fundamental for the development of altered region. These findings indicate that Cerrado and appropriate conservation policies (Chazdon et al., ). Atlantic Forest remnants in modified landscapes in central Armadillos (Cingulata: Dasypodidae) are limited to the Brazil play an important role as refuges for this armadillo Neotropics (Wetzel, ) and some species are sensitive species. In addition to habitat loss, road-kills and poaching to environmental change (Abba & Superina, ). Of  persist as threats to the giant armadillo. Conservation ac- species recorded in Brazil (ICMBio, a), the giant arma- tions are necessary to minimize human impacts and facili- dillo Priodontes maximus is categorized as Vulnerable on tate the persistence of the giant armadillo in this region. the IUCN Red List (Anacleto et al., ). Populations of Policies that both deter illegal deforestation and strengthen this armadillo are decreasing as a result of habitat loss, incentives for the protection of natural vegetation remnants poaching, road-kills and the indiscriminate use of fire to re- and restoration of biological corridors such as gallery forests move natural vegetation or induce regrowth of pastures would aid conservation of the giant armadillo in this area. (Abba & Superina, ; Martins et al., ). The lack of Keywords Agro-ecosystems, biodiversity loss, Cingulata, knowledge about the giant armadillo hinders the implemen- extinction debt, fragmented landscapes, Neotropical habi- tation of conservation actions, especially those focused on  tat, Priodontes maximus, wildlife conservation human-dominated landscapes (Meritt, ; Superina & Abba, ). For example, although the predicted range of P. maximus extends from Venezuela to northern Argentina, including a large portion of Brazil (Anacleto FREDERICO G. LEMOS (Corresponding author) and ALAN N. COSTA Programa de   Conservação Mamíferos do Cerrado, Unidade Acadêmica Especial de et al., ; Chiarello et al., ), it encompasses broad re- Biotecnologia, Universidade Federal de Goiás, Av. Lamartine P. Avelar 1120, gions without records, and the species has been considered Catalão, CEP 75704-020, Goiás, Brazil. E-mail [email protected] extinct in areas with high levels of urbanization/agricultural FERNANDA C. AZEVEDO Programa de Conservação Mamíferos do Cerrado, activities or without official records (Chiarello et al., ; Programa de Pós-graduação em Ecologia, Universidade Federal de Viçosa,  – Viçosa, Minas Gerais, Brazil Srbek-Araujo et al., ). Despite its size ( kg; Superina & Abba, ) the giant armadillo is rarely ob- CARLOS E. FRAGOSO and MOZART C. FREITAS-JUNIOR Programa de Conservação Mamíferos do Cerrado, Cumari, Goiás, Brazil served because of its naturally low density and because it  EDNALDO C. ROCHA Universidade Estadual de Goiás, Ipameri, Goiás, Brazil is nocturnal and fossorial (Noss et al., ; Silveira et al.,    Received  March . Revision requested  July . ; Srbek-Araujo et al., ; Aya-Cuero et al., ). Accepted  November . First published online  August . The absence of records of this species in highly modified

Oryx, 2020, 54(1), 133–139 © 2018 Fauna & Flora International doi:10.1017/S0030605317001867 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.19, on 01 Oct 2021 at 17:42:30, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605317001867 134 F. G. Lemos et al.

areas could be a result of low survey effort rather than ab- temperature of – °C and mean annual total precipitation sence per se. of ,–, mm. Central Brazil lies within the range of P. maximus (Anacleto et al., ; Chiarello et al., ) and is dominated by the Cerrado biome, with the Atlantic Forest biome repre- Methods sented by enclaves of seasonal forest along watercourses (e.g. – the Paranaíba river basin; Ribeiro et al., ). As a result, Data on P. maximus occurrence were obtained during  there is a rich mosaic of physiognomies, ranging from open in natural vegetation remnants in farmlands, private grasslands to forest patches, favouring a high regional bio- reserves, and protected areas, from camera trapping    diversity (Lopes et al., ). However, agricultural activities (Plate a), sightings (Plate b), tracks (Plate a) and fresh  and urbanization have resulted in the replacement of . % burrows (Plate b). We did not include old, abandoned bur- of the natural vegetation by anthropogenic environments rows or those where there was evidence they had been used  (Machado et al., ). The landscape now comprises a ma- by other species (Desbiez & Kluyber ). Camera-trap sur- trix of exotic pastures and crops surrounding fragments of veys were in private natural vegetation remnants in the mu- – natural vegetation, the latter located mainly on higher slopes nicipalities of Araguari (Minas Gerais; ), in – or in rugged terrain (Klink & Machado, ; Carvalho pastures with dense bushes in Cumari (Goiás; ), et al., ). The occurrence of P. maximus in central in the Serra de Caldas Novas State Park (Caldas Novas, – Brazil remains poorly known (Martins et al., ). Here, Goiás; ), and in the private reserve Pé do Morro we present records of the giant armadillo in modified land- Farm of the Universidade Federal de Goiás (Catalão, – scapes of the Cerrado biome and ecotone areas with the Goiás; ). Camera-trap records in different months Atlantic Forest biome in the border area of the states of at each location were considered separately. To determine if Goiás and Minas Gerais. We also present data on the spe- camera-trap photographs were taken during the day or cies’ activity period and its habitat use in agroecosystems. night we calculated the time of sunrise and sunset for the location and date of each photograph, using Rv.. (R Development Core Team, ). We also included data  Study area from road-kills (Plate a) and poaching by local people, reported voluntarily during occasional visits to farms Our study area lies in  municipalities of Goiás and Minas (Plate b). We compared our findings with those of previous Gerais States (Supplementary Table ; Fig. ), in a region studies (Martinelli et al., ; Araújo et al., ; Chiarello where –% of the area is occupied by cattle ranches et al., ; Estrela et al., ; Gomes et al., ; Rocha et al., with exotic pastures, and the remainder comprises scattered ; Fig. ). natural patches of savannah (Cerrado) and mesophytic sea- To determine habitat use we classified the vegetation in sonal forest (Atlantic Forest). The regional climate is mark- each location as savannah, forest or pasture. Savannah has a edly seasonal (Alvares et al., ), with a mean annual relative predominance of native grasses and scattered shrubs

FIG. 1. Records of the giant armadillo Priodontes maximus in  municipalities in the states of Goiás and Minas Gerais, Brazil. Municipalities: , Água Limpa; , Caldas Novas; , Ipameri; , Urutaí; , Campo Alegre; , Catalão; , Goiandira; , Cumari; , Araguari; , Uberaba; , Perdizes. Previous records are from Martinelli et al. (), Araújo et al. (), Chiarello et al. (), Estrela et al. (), Gomes et al. (), and Rocha et al. ().

PLATE 3. Carcasses of the giant armadillo recorded in the state of Goiás: (a) killed on the GO- highway, and (b) the carapace of a poached animal.

PLATE 1. Giant armadillos Priodontes maximus recorded by (a) seasonal forests, riverine forests or gallery forests camera trapping and (b) sighting in Pé do Morro Farm reserve  and Serra de Caldas Novas State Park (respectively), in the state (Oliveira-Filho & Ratter, ). To assess differences in of Goiás (Fig. ). proportions of used and available habitat we compared the frequency of records and available habitat for the agro-ecosystems surveyed in Araguari (Minas Gerais; F.C. and small trees, which in the Cerrado includes the vegeta- Azevedo unpubl. data) and Cumari (Goiás; F.G. Lemos un- tion physiognomies campo sujo, campo cerrado and cerrado publ. data), and the reserve Pé do Morro Farn in Catalão .  sensu stricto. Forest consisted of trees m tall and with (Goiás; Cardoso & Moreno, ). canopy formation such as the cerradão physiognomy,

Results

During the -year period we obtained a total of  records of P. maximus:  from  private farmlands,  in a private reserve,  in three protected areas, three animals killed on paved roads and two poached (Fig. ; Supplementary Table ). Forty-six per cent of the records were from camera trapping (), carcasses () and sightings (), and %() were from fresh burrows and tracks, often on trails or at the edge of dirt roads. Camera-trap photographs and sightings were almost ex- clusively during the night (Fig. ), with just one in daylight, shortly after dawn (Supplementary Table ). Most records were in areas with native vegetation cover (%), with  in forest and  in savannah. In Cumari and Catalão six re-

PLATE 2. Typical evidence of the giant armadillo used to record cords were in pastures. The frequency of records in different the species: (a) track and (b) fresh burrow on a leaf-cutter ant habitats generally followed their relative availability in each nest. study site (Table ). The % of records in anthropogenic

southern range of the species (Martinelli et al., ; Fig. ). Previous studies in south-east Goiás state located only six records of P. maximus for – (Araújo et al., ; Chiarello et al., ; Estrela et al., ; Gomes et al., ), but we found  records within the same general area. Based on the home range size of a giant armadillo (–, ha; Silveira et al., ) we assume that at least one animal was living in the vegetation remnants surrounding the record points, and thus our findings, in combination with previous records, potentially indicate the persistence of several populations of this species in natural patches across a  FIG. 2. Activity pattern of the giant armadillo, based on large section of agro-ecosystems in central Brazil. However, camera-trap photographs and three sightings during July – there are many potentially suitable vegetation remnants October  in four municipalities in the states of Goiás and Minas Gerais (Fig. ). within the area that remain unsurveyed. Our findings corroborate previously identified traits of environments, including road-kills, were , . km from giant armadillos: individuals are essentially nocturnal natural vegetation remnants. (Noss et al., ; Silveira et al., ; Srbek-Araujo et al., ; Aya-Cuero et al., ), occur in both open savannah Discussion and closed forest habitats, and their use of habitat tends to reflect habitat availability in the landscape (Silveira et al., The giant armadillo is widely distributed in central Brazil ). It is most likely that giant armadillos select areas (Anacleto et al., ; Chiarello et al., ), and elderly with populations of termites or ants, which are their main rural residents reported to us that formerly it was not un- food items (Anacleto, ), independently of the dominant usual to find this armadillo’s tracks or to sight the species vegetation of an area. on their properties. However, the giant armadillo is now Although we surveyed extensive areas of pasture in the rarely seen, most likely as a result of habitat loss combined matrix between natural vegetation patches, the low fre- with road-kills and poaching (Chiarello et al., ). quency of records (%) in this environment suggests that Intensive mammal surveys have only recently been con- exploration of human-modified habitats is uncommon for ducted in natural vegetation remnants across the study re- giant armadillos even in predominantly altered landscapes; gion (Bruna et al., ; Araújo et al., ; Estrela et al., they mainly explore natural vegetation patches within these ; Gomes et al., ; Rocha et al., ). Our study highly modified areas (Silveira et al., ). Similarly, the  spans the longest period ( years) and covered  km of bush dog Speothos venaticus tends to use native vegetation Cerrado and Atlantic Forest remnants. preferentially (% of records in savannah and forest Protected areas on the border of Minas Gerais and patches), even where this vegetation occupied only Espirito Santo states are considered the last stronghold for one-third of a cultivated landscape in Mato Grosso state P. maximus in the Atlantic Forest (Srbek-Araujo et al., (Lima et al., ). This highlights the importance of the ) but our findings indicate that the species is still pre- protection of private reserves and biological corridors, sent in Atlantic Forest remnants within the basins of the mainly gallery forests, for the conservation of rare and Paranaíba and Araguari rivers. In addition, a record from threatened species in modified landscapes. Vale do Encantado Private Natural Heritage Reserve in Habitat fragmentation often results in vegetation patches the municipality of Uberaba (Minas Gerais) expands the decreasing in size and increasing in number and isolation

TABLE 1 Availability of habitats and number of records (from camera trapping, sightings, carcass recovery, tracks and burrows) of the giant armadillo Priodontes maximus during – in the municipalities of Araguari (Minas Gerais) and Cumari (Goiás), and the reserve Pé do Morro Farm in Catalão (Goiás).

Area, ha (%) No. of records (%) Habitat Araguari Cumari Pé do Morro Farm1 Araguari Cumari Pé do Morro Farm Savannah 396 (0.4) 2,104 (4.4) 38 (42.0) 1 (10.0) 1 (12.5) 4 (33.3) Forest 30,024 (31.9) 10,033 (21.1) 31 (34.0) 9 (90.0) 4 (50.0) 5 (41.7) Pastures & crops 56,065 (59.6) 34,945 (73.6) 21 (24.0) 0 (0) 3 (37.5) 3 (25.0) Others2 7,644 (8.1) 404 (0.9) 0 (0) 0 (0) 0 (0) 0 (0)

 Data from Cardoso & Moreno ().  Urban areas, farm facilities, roads, artificial dams and degraded areas.

(Fahrig, ), negatively affecting species with large spatial connections between natural vegetation remnants and po- requirements such as the giant armadillo (Chiarello, ). tentially result in biodiversity loss (Michalski et al., ; However,  of our records were from fragments smaller Paul et al., ). (– ha) than an armadillo’s home range and therefore Our research suggests that the giant armadillo may still potentially unsuitable to support even a single giant arma- occur in other highly modified areas of central Brazil and dillo. This could be explained by an extinction debt elsewhere, undetected because of low survey efforts. This (Kuussaari et al., ) for the giant armadillo. The species’ may also be the case for other rare and threatened relatively long life expectancy (– years; Nowak, ) in- Brazilian species (e.g. bush dogs were recently recorded in dicates low population turnover and may mask the long- our study area, where the species was expected but had term effects of fragmentation. The irregular topography of not previously been observed; Azevedo et al., ). Our re- the landscape in the border areas between Goiás and sults suggest that road-kills and poaching, in addition to Minas Gerais states results in habitat remnants being close habitat loss, continue to threaten the giant armadillo in cen- to each other, allowing giant armadillos to forage in small tral Brazil. Policies that both deter illegal deforestation and areas and still survive within fragmented landscapes for a strengthen incentives for the protection of natural vegeta- limited time. This extinction delay could present an oppor- tion remnants and restoration of biological corridors such tunity for recovery of this species via habitat restoration and as gallery forests are therefore necessary for the conservation landscape management (Kuussaari et al., ). of the giant armadillo in this area. Road-kills of armadillos and other mammal species have been recorded in central Brazil (Fischer et al., ), and we Acknowledgements We thank Hugo Costa, Frederico Cigano Souza, located three road-kills, but this problem is probably under- Leandro Abade, Guilherme Dias, Lucas Ribeiro and Tiago Borges for reported. Illegal hunting is still a common practice in central their assistance with fieldwork, Daniel Rocha for helping with time Brazil and game meat consumption remains a cultural habit, calculations, Leonardo Gomes for providing coordinates for mapping mainly in rural communities (FGL pers. comm.), although literature records, Stacie Castelda-Bickley, Arnaud Desbiez, Ernane probably insignificant compared to that observed in the Vieira-Neto, Martin Fisher and anonymous reviewers for their cri-  tiques, landowners and protected area managers for their hospitality, Amazon forest (e.g. Baía-Junior et al., ; van Vliet Capim Branco Energy Company for financial support for part of the  et al., ). The low frequency of hunting in agricultural re- study and Minas Gerais Electrical Company for the permit to work at gions (Rushton et al., ) such as central Brazil may be Galheiro Private Natural Heritage Reserve. FCA was supported by a associated with access to domestic sources of animal protein doctoral fellowship from HIDROEX (UNESCO/BID), and ECR by a and stricter environmental laws. We recorded only two grant from Goiás State University (PROBIP/PrP 009/2016). poached giant armadillos, and suspect that hunting of this species in central Brazil is opportunistic. Author contributions Conception and design of the study: FGL, The decline of the giant armadillo (at least % since ANC, FCA, CEF and ECR; data collection: FGL, ANC, FCA, CEF, ; ICMBio a) may influence community diversity MCF and ECR; writing: FGL, ANC, FCA and CEF; revisions: FGL, ANC, FCA, CEF, MCF and ECR. and vegetation structure. The species is an ecosystem engi- neer (Leite-Pitman et al., ; Desbiez & Kluyber, ; Aya-Cuero et al., ), a potential prey for large predators Conflicts of interest None. such as the jaguar Panthera onca and cougar Puma concolor, and a specialized insect-predator (Anacleto, ). With re- Ethical standards This study abided by the Oryx Code of Conduct. spect to the latter, the absence of a top-down effect on insect herbivores, especially leaf-cutter ants (Costa & Vieira-Neto, References ), may affect vegetation structure and dynamics in  modified environments (Terborgh et al., ; Silva et al., ABBA, A.M. & SUPERINA,M.() The / Armadillo Red List ). The border region of the states of Goiás and West Assessment. Edentata, , –. Minas Gerais has experienced intensive landscape modifica- ALVARES, C.A., STAPE, J.L., SENTELHAS, P.C., GONÇALVES, J.L.M. & tion, and , –% of natural vegetation remnants lie within SPAROVEK,G.() Köppen’s climate classification map for Brazil.  – protected areas (Carvalho et al., ). The high degree of Meteorologische Zeitschrift, , . ANACLETO, T.C.S. () Food habits of four armadillo species in the landscape modification and the high cost of land in this Cerrado area, Mato Grosso, Brazil. 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