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(1) An individual of Tolypeutes matacus (arrow) resting inside its burrow at Santa Teresa Ranch, Corumbá, Mato Grosso do Sul. (2-3) Straw-nest built by T. matacus in a pasture area at Duas Lagoas Ranch, Cáceres, Mato Grosso: (2) upper view of the nest (circle) and (3) close-up of the same nest (arrow indicates the entrance of the nest). (4-5) (4) An almost imperceptible individual of T. matacus resting in a depression under leaf litter at Santa Teresa Ranch, Corumbá. The arrow points to the only visible part of the carapace. (5) The same GPS-fitted animal resting in the depression after we manually removed some of the leaf litter to expose part of its carapace (circled).

(1) An individual of Tolypeutes matacus (arrow) resting inside its burrow at Santa Teresa Ranch, Corumbá, Mato Grosso do Sul. (2-3) Straw-nest built by T. matacus in a pasture area at Duas Lagoas Ranch, Cáceres, Mato Grosso: (2) upper view of the nest (circle) and (3) close-up of the same nest (arrow indicates the entrance of the nest). (4-5) (4) An almost imperceptible individual of T. matacus resting in a depression under leaf litter at Santa Teresa Ranch, Corumbá. The arrow points to the only visible part of the carapace. (5) The same GPS-fitted animal resting in the depression after we manually removed some of the leaf litter to expose part of its carapace (circled).

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Article
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It is believed that the two species of Tolypeutes Illiger, 1811are the only armadillos that do not dig their own burrows, and that these species simply re-use burrows dug by other species. Here, we show that Tolypeutes matacus (Desmarest, 1804) and Tolypeutes tricinctus (Linnaeus, 1758) dig their own burrows. We describe the burrows and three other...

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... Smith, 2007;Wetzel, Gardner, Redford, & Eisenberg, 2008). This could make functional sense, as a less fossorial lifestyle requires less force transmission through the vertebral column during high-force digging activities (Hildebrand 1985;Clerici et al 2018;Gaudin and Biewener 1992) and might be associated with less robust TBA, but more recent studies have found that they can and do dig their own burrows (Nina Attias, Miranda, Sena, Tomas, & Mourão, 2016;Galliari et al., 2010). Nevertheless, our results suggest that some aspect of their ecology may lead to unusually light vertebral loading. ...
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Synopsis Trabecular bone is a spongy bone tissue that serves as a scaffolding-like support inside many skeletal elements. Previous research found allometric variation in some aspects of trabecular bone architecture (TBA) and bone microstructure, whereas others scale isometrically. However, most of these studies examined very wide size and phylogenetic ranges or focused exclusively on primates or lab mice. We examined the impact of body size on TBA across a smaller size range in the mammalian clade Xenarthra (sloths, armadillos, and anteaters). We µCT-scanned the last six presacral vertebrae of 23 xenarthran specimens (body mass 120 g–35 kg). We collected ten gross-morphology measurements and seven TBA metrics and analyzed them using phylogenetic and nonphylogenetic methods. Most metrics had similar allometries to previous work. However, because ecology and phylogeny align closely in Xenarthra, the phylogenetic methods likely removed some covariance due to ecology; clarifying the impact of ecology on TBA in xenarthrans requires further work. Regressions for Folivora had high P-values and low R-squared values, indicating that the extant sloth sample either is too limited to determine patterns or that the unique way sloths load their vertebral columns causes unusually high TBA variation. The southern three-banded armadillo sits far below the regression lines, which may be related to its ability to roll into a ball. Body size, phylogeny, and ecology impact xenarthran TBA, but parsing these effects is highly complex.
... Prior to this study, recent records of the Brazilian three-banded armadillo were limited to only 11 localities (Feijó et al., 2015;Attias et al., 2016;Campos et al., 2019;Santos et al., 2019;Magalhães et al., 2021). In contrast, local extinctions have been widely reported across its distribution (Santos et al., 1994;Oliveira, 1995;Feijó et al., 2015;Neto and Alves, 2016). ...
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Conservation of poorly known species is challenging as lack of knowledge on their specific requirements may hamper effective strategies. Here, by integrating biogeographical and landscape analyses, we show that informed actions can be delineated for species with limited presence-only data available. We combine species distribution and connectivity models with temporal land cover changes to define priority areas for conservation of the endemic Brazilian three-banded armadillo, one of the most threatened xenarthrans that was once considered extinct in the wild. We revealed that areas of savanna and grassland are the most suitable habitats for the species and that uplands in the Caatinga ecoregion have a greater likelihood for dispersal. The few remnant armadillo populations are spatially associated with core areas of natural vegetation remnants. Worrisomely, 76% of natural core areas were lost in the past 30 years, mirroring the species’ severe population decline. Preserving the remnant core natural areas should be a high priority to ensure the species’ survival. We highlight key areas for proactive and reactive conservation actions for the three-banded armadillo that will benefit other threatened sympatric species. Our integrative framework provides a set of valuable information for guided conservation management that can be replicated for other poorly known species.
... For instance, only recently the burrowing activity of Tolypeutes spp. and the T. tricinctus predation by native predators have been described in the literature (Attias et al. 2016;Magalhães et al. 2021). Most of the available information about T. tricinctus comes from unpublished works and field observations (e.g., Marinho-Filho et al. 1997), and no published study has systematically assessed its habitat relationships so far. ...
... Tolypeutes tricinctus depends on the soil to forage and dig burrows (Attias et al. 2016). Taber (1945) discusses that Dasypus novemcinctus preferentially digs in soft soils (e.g., sandy soils), which require less energy expenditure. ...
... Taber (1945) discusses that Dasypus novemcinctus preferentially digs in soft soils (e.g., sandy soils), which require less energy expenditure. As Tolypeutes may be the least fossorial of extant armadillos (Attias et al. 2016), they could also prefer to dig in sandy soils, thus preferentially occupying these sites Sand content (g/g)* 0.70 (0.41-0.89) + NE Some camera traps did not work continually during the sampling period. Variation in sampling effort can affect detection probability Sampling effort at each sampling unit and occasion (days)* 4.10 (0-5) NE + Camera trapping and active searches can present distinctive effectiveness in detecting different species (Silveira et al. 2003). ...
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Exploring factors related to species distribution is important to better understand their natural histories and provide their effective conservation. However, the ecology of some threatened species remains poorly understood. Here, we conducted the first quantitative investigation of species-habitat relationships for the Brazilian three-banded armadillo, Tolypeutes tricinctus, a threatened species endemic to Brazil. We combined camera traps and active searches to explore the influence of ecological and methodological factors, including human-related habitat features, on T. tricinctus occupancy and detection probabilities in a human-modified landscape in northeastern Brazil. The T. tricinctus occupancy probability was high throughout the study area, whereas its detection probability was eight times higher by active searches than camera trapping, which should be considered when designing studies on T. tricinctus ecology in the future. Our results suggest that T. tricinctus can be widely distributed in human-modified landscapes under moderate levels of hunting and habitat loss and highlight the importance of the engagement of local people into research and conservation projects for better outcomes, as we found by counting on their local ecological knowledge for the conduction of active searches.
... Also, five physical soil properties at 30 cm depth were included as environmental predictors for glacial and current niches. We chose this soil depth based on the estimation of 35 cm burrow depth for the species in Brazil (Attias et al. 2016). These layers were obtained from the SoilGrids database (Hengl et al. 2017) and rescaled to match WorldClim layers resolution using 'raster' package v. 3.3-13 (Hijmans 2020) in R. Calibration layers were cropped using a mask with an estimation of the hypothetical accessible area (M) for the species (Barve et al. 2011). ...
... Although it can dig to make burrows, T. matacus differs from other armadillos in that its burrows are shallow. In addition, the species also uses depressions in the ground and makes nest-like structures using straw and grass as burrows, without the need to carry out excavations for this (Attias et al. 2016). Considering these characteristics, the soil variables may be of less importance in this species and are mainly relevant in regions where the strategies of making nests and using depressions are not feasible. ...
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Climatic oscillations acted as an essential factor in molding the distribution and composition of Neotropical biota. However, their effects on the South American Chaco are still poorly known. In this work, we present an updated collection of records of one of its flagship species, Tolypeutes matacus. We analyze the species' range dynamics in current and past scenarios to unveil the effects of past changes over the Chacoan biota. We construct three niches, one with current data (current niche), the other using Pleistocenic occurrence data and predictors from the Last Glacial Maximum (glacial niche), and the last one combining current and Pleistocene data to generate a multitemporal niche. Niches were projected to Current, Mid-Holocene, Last Glacial Maximum, and Last Interglacial scenarios. We obtain 398 records for the species, corresponding to current, historical, Holocene, and Pleistocene records. Current distribution occurs mainly in the currently Chaco region. For current niche reconstructions, suitable areas were smaller than those found nowadays, mainly during the Last Interglacial and Last Glacial Maximum. Holocene reconstruction recovers a distribution of suitable areas pretty similar to the present times. In contrast, glacial niche projection shows higher suitability values in the Pampean Region in all periods analyzed. Multitemporal niche projections showed similar results to those obtained from the current niche projections. Climatic oscillations seem to have significantly impacted the distribution of the species, generating retractions during past times and a period of range stability since the Holocene. Differences between niche projections suggest that Pampean populations may have evolved in different conditions than those from the Chacoan region.
... Grassé (1955) e Boily (2002) classificam os tatus como homeotermos imperfeitos, e a construção de estruturas para abrigo, como buracos, pode ser uma consequência da capacidade termorregulatória limitada destes animais (McNab, 1980(McNab, , 1985Abba, 2008). Cada espécie cava sua toca com tamanho e forma característica (Carter & Encarnação, 1983;Emmons, 1990;González et al., 2001;Lima Borges & Tomas, 2004;Attias et al., 2016;Desbiez et al., 2018). Suas tocas são utilizadas para dormir, abrigar os filhotes, escapar de predadores, evitar condições climáticas adversas ou para facilitar o acesso imediato a um reservatório de insetos, pois muitas são escavadas dentro de formigueiros ou cupinzeiros (González et al., 2001;McDonough & Loughry, 2003;Maccarini et al., 2015). ...
... Assim como outros tatus, o tatu-bolinha possui hábito solitário. Com exceção de mães com filhote, os tatus-bola não compartilham abrigos ou reaproveitam estruturas feitas por outros indivíduos ou espécies (Attias et al., 2016). Fêmeas adultas não compartilham sua área de vida com outras fêmeas adultas e adultos em geral não compartilham suas áreas 'core' com outros adultos . ...
... Por muito tempo, acreditou-se que as espécies do gênero Tolypeutes eram as únicas dentre os tatus que não cavavam suas próprias tocas, utilizando tocas feitas por outros animais. No entanto, após extensa observação em campo, Attias et al. (2016) relatam que os tatus-bola, assim como as outras espécies de tatu, cavam suas próprias tocas. Além das tocas, esta espécie também pode se abrigar em pequenas depressões no solo cobertas por folhiço ou em ninhos de palha. ...
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Xenarthra contém duas ordens, seis famílias, 14 gêneros e 38 espécies viventes. A ordem Cingulata (tatus) representa 22 das espécies de xenartros e seis destas são encontradas no Pantanal (Dasypus novemcinctus, Cabassous tatouay, C. squamicaudis, Tolypeutes matacus, Euphractus sexcinctus e Priodontes maximus). Os tatus se diferem de outros mamíferos por terem uma carapaça formada por placas ósseas articuladas que, como um escudo, cobrem a cabeça e o dorso. Neste artigo, baseamo-nos em extensa revisão bibliográfica e décadas de trabalho realizado pelos autores para descrever o estado do conhecimento sobre ecologia, biologia, características morfológicas, saúde, estado de conservação e distribuição dos tatus no Pantanal. Visando dar subsídios a futuros trabalhos com as espécies, também descrevemos melhores práticas para a captura e o manejo (e.g., anestesia e coleta de material biológico) das espécies em campo. Apesar do recente aumento do número de estudos, ainda restam diversas lacunas de informação sobre a ecologia e a biologia da maioria das espécies de tatus do Pantanal. Sendo assim, esperamos que as informações e os métodos descritos aqui sirvam de estímulo e base para o desenvolvimento de novos estudos, que aumentem nosso conhecimento sobre estas espécies na região do Pantanal e permitam o planejamento de estratégias de conservação eficientes.
... Major locomotor adaptations of extant xenarthrans reflect the group's taxonomic diversity. Armadillos are generally characterized by a fully terrestrial locomotor ecology dominated by digging habits (Vizcaíno and Milne 2002;Attias et al. 2016;Amson et al. 2017). Most anteaters are at least semi-arboreal and the three vermilinguan genera represent a cline from fully terrestrial (Myrmecophaga) to fully arboreal (Cyclopes) locomotion, with Tamandua spanning the two ends of the behavioral spectrum (Young et al. 2003;Orr 2005;Amson et al. 2017). ...
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Identifying ecomorphological convergence examples is a central focus in evolutionary biology. In xenarthrans, slow arboreality independently arose at least three times, in the two genera of ‘tree sloths’, Bradypus and Choloepus, and the silky anteater, Cyclopes. This specialized locomotor ecology is expectedly reflected by distinctive morpho-functional convergences. Cyclopes, although sharing several ecological features with ‘tree sloths’, do not fully mirror the latter in their outstandingly similar suspensory slow arboreal locomotion. We hypothesized that the morphology of Cyclopes is closer to ‘tree sloths’ than to anteaters, but yet distinct, entailing that slow arboreal xenarthrans evolved through ‘incomplete’ convergence. In a multivariate trait space, slow arboreal xenarthrans are hence expected to depart from their sister taxa evolving toward the same area, but not showing extensive phenotypical overlap, due to the distinct position of Cyclopes. Conversely, a pattern of ‘complete’ convergence (i.e., widely overlapping morphologies) is hypothesized for ‘tree sloths’. Through phylogenetic comparative methods, we quantified humeral and femoral convergence in slow arboreal xenarthrans, including a sample of extant and extinct non-slow arboreal xenarthrans. Through 3D geometric morphometrics, cross-sectional properties (CSP) and trabecular architecture, we integratively quantified external shape, diaphyseal anatomy and internal epiphyseal structure. Several traits converged in slow arboreal xenarthrans, especially those pertaining to CSP. Phylomorphospaces and quantitative convergence analyses substantiated the expected patterns of ‘incomplete’ and ‘complete’ convergence for slow arboreal xenarthrans and ‘tree sloths’, respectively. This work, highlighting previously unidentified convergence patterns, emphasizes the value of an integrative multi-pronged quantitative approach to cope with complex mechanisms underlying ecomorphological convergence.
... Although armadillos are regularly observed above ground, they dig burrows for shelter (housing, raising offspring), protection (to hide from predators and to buffer against environmental temperatures), and feeding (foraging burrows) (McNab 1980, Eisenberg and Redford 1999, Desbiez et al. 2018. Burrows created by armadillos have specific shapes and sizes (Carter and Encarnaçao 1983, Abba et al. 2005, Trovati 2015, Attias et al. 2016, Desbiez et al. 2018, which are typically influenced by anatomical and morphological differences among species (e.g., Carter andEncarnação 1983, Attias et al. 2016). Several previous studies have used armadillo burrows to estimate population parameters including habitat use and density and to assess activity and behavior (Zimmerman 1990;McDonough et al. 2000;Abba et al. 2005Abba et al. , 2007Desbiez et al. 2018). ...
... Not surprisingly, more records on burrow measurements are available for the species with the widest geographic range, Dasypus novemcinctus (Zimmerman 1990, McDonough et al. 2000, Platt et al. 2004, Sawyer et al. 2012). In addition to D. novemcinctus, several additional species of armadillos were also monitored and had measurements of their burrows reported (Medri 2008, Attias et al. 2016, Desbiez et al. 2018). However, the relationship between the reported burrow measurements and species identification remains tenuous (e.g., Arteaga and Venticinque 2010). ...
... From these works, we extracted and used the mean and standard deviation values to generate synthetic data for 3 measures: width, height, and angle. We chose these measurements because they are the only measurements consistently present on the works related to morphometric measurements of armadillo burrows (Table 1), and they can provide relevant information that may assist in the discrimination of different burrows and the species of armadillo (Medri 2008, Sawyer et al. 2012, Attias et al. 2016, Desbiez et al. 2018. ...
Article
Having accurate information about population parameters of armadillos (Mammalia, Cin-gulata) is essential for the conservation and management of the taxon, most species of which remain poorly studied. We investigated whether we could accurately identify 4 armadillo species (Euphractus sexcinctus, Dasypus novemcinctus, Cabassous tatouay, and Cabassous unicinctus) based on burrow morphometry. We first selected published studies that reported measurements of width, height, and angle of the burrows used by the 4 species of armadillos. Then, using such data we simulated burrow measurements for each of the 4 species of armadillos and we created predictive models through supervised machine learning that were capable of correctly identifying the species of armadillos based on their burrows' morphometry. By using classification algorithms such as Random Forest, K-Nearest Neighbor, Support Vector Machine, Naive Bayes, and Decision Tree C5.0, we achieved the overall accuracy for the classification task by about 71%, including an overall Kappa index by about 61%. Euphractus sexcinctus was the most difficult species to discriminate and classify (approximately 68% of accuracy), whereas C. unicinctus was the easiest to discriminate (approximately 93% of accuracy). We found that it was possible to identify similar-sized arma-dillos based on the measurements of their burrows described in the literature. Finally, we developed an R function (armadilloID) that automatically identified the 4 species of armadillos using burrow morphology. As the data we used represented all studies that reported the morphometry of burrows for the 4 species of armadillos, we can generalize that our function can predict armadillo species beyond our data.
... Os tatus (Dasypodidae: Cingulata) são mamíferos amplamente distribuídos na América Central e do Sul, com apenas uma espécie com alcance na América do Norte (WETZEL, 1985). Este grupo, que só existe no continente americano, possui 20 espécies no mundo, 11 com ocorrência no Brasil (PAGLIA et al., 2012 Esses animais são os únicos mamíferos que possuem uma carapaça, composta por tecido ósseo revestido de epiderme (ENGELMANN, 1985) e todos possuem a habilidade de cavar tocas (EISENBERG; REDFORD, 1999;ATTIAS et al., 2016). Suas características fisiológicas e ecológicas, tais como baixa temperaturas corpórea, frágil sistema imunológico e o hábito de viver literalmente imerso no solo e na matéria orgânica, além de compartilharem suas tocas, contribuem para que se tornem potenciais hospedeiros apropriados de inúmeras doenças (TABER, 1945). ...
... In this instance, the proportions of their hind feet may signify features secondarily modified for terrestrial locomotion. Three banded armadillos are reported as exhibiting the most 'cursorial' habits among armadillos (Vizcaíno and Milne 2002;Costa et al. 2019), and overall, they may be the least reliant on scratch-digging for resource acquisition (Smith 2009) and burrow construction (Attias et al. 2016), especially by use of the hind limbs. Similarly, D. hybridus may scratch-dig shallow burrows while foraging, but this species mainly has shared ecomorphology with To. matacus (Abba and Superina 2016), which is plausibly why these two species are grouped closely in morphospace. ...
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Divergence in forelimb morphology is often associated with functional habits exhibited within the Xenarthra, ranging from terrestrial-digging in armadillos to arboreal-suspension in sloths. We hypothesized that quantitative differences in hind limb form also will be predictive of the diverse lifestyles observed in this small clade. A total of 26 morphofunctional indices were calculated from 42 raw measurements of bone length/width/depth in a sample of N = 76 skeletal specimens (18 species). Index data for each species were categorized by substrate preference and use and then evaluated using a combination of stepwise Discriminant Function Analysis (DFA) and Principal Component Analysis (PCA) to determine significant osteological correlates (traits) among extant taxa. Additionally, character states of the morphometric data were inferred using a recent hypothesis of xenarthran phylogeny. DFA determined 14 distinct morphofunctional indices relating to femur robustness, hip/ankle/limb mechanical advantage, and foot and claw length as the most discriminating features. PCA clearly separated armadillos from sloths in morphospace based on overall robustness versus gracility, as well as proximal versus distal lengths of skeletal elements (including the claws), whereas these characteristics were intermediate in the hind limbs of anteaters and selected armadillos having either a larger greater trochanter or modified foot/claw proportions. Two-toed and three-toed sloths showed further separation from each other in morphospace primarily driven by proportions of their tibia and hind feet despite evidence of convergence for numerous functional traits. Moreover, the majority of the traits measured had significant phylogenetic signal and several of these indicated clear patterns of convergent and divergent evolution in xenarthrans by evaluation of their tip states. Our assessments expand functional interpretations of xenarthran limb form and identify potentially conserved and secondarily modified traits related to fossoriality across taxa, including in three-toed sloths, demonstrate possible morphological trade-offs between digging and climbing habits, and suggest derived traits adapted for arboreal lifestyle and suspensory function.
... Os tatus (Dasypodidae: Cingulata) são mamíferos amplamente distribuídos na América Central e do Sul, com apenas uma espécie com alcance na América do Norte (WETZEL, 1985). Este grupo, que só existe no continente americano, possui 20 espécies no mundo, 11 com ocorrência no Brasil (PAGLIA et al., 2012 Esses animais são os únicos mamíferos que possuem uma carapaça, composta por tecido ósseo revestido de epiderme (ENGELMANN, 1985) e todos possuem a habilidade de cavar tocas (EISENBERG; REDFORD, 1999;ATTIAS et al., 2016). Suas características fi siológicas e ecológicas, tais como baixa temperaturas corpórea, frágil sistema imunológico e o hábito de viver literalmente imerso no solo e na matéria orgânica, além de compartilharem suas tocas, contribuem para que se tornem potenciais hospedeiros apropriados de inúmeras doenças (TABER, 1945). ...