Taxonomic revision of the genus Cabassous McMurtrie, 1831 (Cingulata: Chlamyphoridae), with revalidation of Cabassous squamicaudis (Lund, 1845).

Cabassous comprises armadillos lacking a full osteoderm cover in the tail, justifying its common name naked-tailed armadillos. In the only taxonomic revision of the genus, in 1980, four living species were recognized, including a polytypic taxon with two subspecies. Recent studies have questioned this classification, but a comprehensive taxonomic review is lacking. Here, we revise the taxonomy of the genus Cabassous using complementary morphological approaches and clarify the geographical limits of naked-tailed armadillo species. Based on qualitative and quantitative analyses, we recognize five living species: C. centralis, C. chacoensis, C. squamicaudis, C. unicinctus, and C. tatouay. Most of the species can be easily differentiated using external or cranial traits, except C. centralis and C. unicinctus, which share several morphological features. The scutes pattern on the cephalic shield is an important diagnostic feature in naked-tailed armadillos and can be easily applied in field studies. Cabassous squamicaudis and C. unicinctus were previously treated as subspecies but we show they have conspicuous diagnostic traits, without mixture of characters even in closer contact. Cabassous species can be classified as open-dwellers (C. chacoensis and C. squamicaudis), forest-dwellers (C. centralis and C. unicinctus), or of more generalist habits (C. tatouay). We designate a lectotype for C. unicinctus to preserve its long-term nomenclature use.

Taxonomic revision of the long-nosed armadillos, Genus Dasypus Linnaeus, 1758 (Mammalia, Cingulata).

Dasypus is the most speciose genus of the order Cingulata, including approximately 40% of known living armadillos. Nine species are currently recognized, although comprehensive analyses of the entire genus have never been done. Our aim is to revise the taxonomy of the long-nosed armadillos and properly define the taxa. We examined 2126 specimens of Dasypus preserved in 39 different museum collections, including 17 type specimens. Three complementary methods were applied to explore morphological datasets both qualitatively and quantitatively. Qualitative morphological variation in discrete characters was assessed by direct observations of specimens. Linear morphometric variation was based on external data and cranial measurements of 887 adult skulls. The shape and size of the skull was abstracted through two-dimensional geometric morphometric analyses of dorsal, lateral and ventral views of respectively 421, 211, and 220 adult specimens. Our results converge on the recognition of eight living species (D. beniensis, D. kappleri, D. mazzai, D. novemcinctus, D. pastasae, D. pilosus, D. sabanicola, and D. septemcinctus), and three subspecies of D. septemcinctus (D. s. septemcinctus, D. s. hybridus, and a new subspecies from Cordoba described here). Information on type material, diagnosis, distribution, and taxonomic comments for each taxon are provided. We designate a lectotype for D. novemcinctus; and a neotype for Loricatus hybridus (= D. septemcinctus hybridus).

Taxonomic revision of the Dasypus kappleri complex, with revalidations of Dasypus pastasae (Thomas, 1901) and Dasypus beniensis Lönnberg, 1942 (Cingulata, Dasypodidae).

Dasypus kappleri is the largest species of the genus Dasypus and is restricted to the Amazonian rainforest biome. Over the last century, related taxa have been described and synonymized without comprehensive analyses, and the current classification involving two subspecies, Dasypus k. kappleri and Dasypus k. pastasae, has never been revised. The aim of this work is to clarify the taxonomy of Dasypus kappleri through integrative morphological and morphometric analyses. We examined 70 specimens housed in scientific collections as well as photographs of the type specimens of five nominal taxa. Three methodologies (discrete characters, linear and geometric morphometrics) were employed. All results converged on the recognition of three allopatric groups, each with diagnostic qualitative and quantitative traits, that we recognize as full species: Dasypus kappleri Krauss, 1862, occurs in the Guiana shield; Dasypus pastasae (Thomas, 1901) is distributed from the eastern Andes of Peru, Ecuador, Colombia, and Venezuela south of the Orinoco River into the western Brazilian Amazon; and Dasypus beniensis Lönnberg, 1942, occurs in the lowlands of Amazonian Brazil and Bolivia to the south of the Madre de Dios, Madeira, and lower Amazon rivers. This revision raises to nine the number of living species of Dasypus.

The correct name of the endemic Dasypus (Cingulata: Dasypodidae) from northwestern Argentina.

We show that Dasypus mazzai Yepes 1933 is a senior synonym of Dasypus yepesi Vizcaíno 1995. We present morphological evidence that the holotype of D. mazzai is not a juvenile of Dasypus novemcinctus or any other species of this genus, but a distinct endemic species from northwestern Argentina undistinguishable from D. yepesi. Therefore, the correct name for the long-nosed armadillo of intermediate size occurring in the Argentinean provinces of Jujuy and Salta is Dasypus mazzai Yepes 1933.

When xenarthrans had enamel: insights on the evolution of their hypsodonty and paleontological support for independent evolution in armadillos.

All xenarthrans known to date are characterized by having permanent teeth that are both high crowned and open rooted, i.e., euhypsodont, and with a type of hypsodonty different from that of the rest of Placentalia: dentine hypsodonty. Also, most xenarthrans lack enamel; however, its presence has been reported in the fossil armadillo Utaetus buccatus and in living Dasypus. Considering the divergence of Xenarthra from other eutherians that possessed enameled teeth, the absence of enamel is a derived character. Diverse specializations are known in the dentition of xenarthrans, but the primitive pattern of their teeth and dentitions is still unknown. Here, we describe the mandible and teeth of a fossil armadillo, Astegotherium dichotomus (Astegotheriini, Dasypodidae), from the early Middle Eocene of Argentine Patagonia, with teeth showing both true enamel and closed roots. It is the oldest xenarthran with mandibular remains exhibiting protohypsodonty and is therefore likely representative of ancestral cingulates and xenarthrans generally. Astegotherium supports a recent hypothesis based on molecular data that enamel loss occurred independently not only within xenarthrans but also within dasypodid armadillos.

A new Dasypodini armadillo (Xenarthra: Cingulata) from San Gregorio Formation, Pliocene of Venezuela: affinities and biogeographic interpretations.

We describe Pliodasypus vergelianus gen. et sp. nov., a Dasypodini armadillo from the middle Pliocene of Venezuela (Vergel Member, San Gregorio Formation). Although scarce, the remains are remarkable because of their geochronologic proximity to the main phase of Great American Biotic Interchange (GABI). The cladistic analysis conducted reveals that Pliodasypus groups with Dasypus and both are sister taxa of Propraopus, whereas Anadasypus is at a basal position. With respect to the records of tribe Dasypodini, after its oldest representative (Anadasypus, middle and late Miocene), the chronologically subsequent form is Pl. vergelianus (middle Pliocene), followed by Dasypus bellus in higher northern latitudes (late Pliocene), and then by widespread occurrences in the Pleistocene of North America (D. bellus) and South America (Propraopus, Dasypus punctatus, and Dasypus novemcinctus). Thus, we infer that Dasypus differentiated in the late Pliocene at low latitudes in the northern South America. It leads to two alternative hypotheses of dispersal: (a) some early Dasypus remained cryptically in South America until the Pleistocene, whereas others dispersed to North America between 2.2 and 2.7 Ma, or (b) they dispersed to North America subsequently to the emersion of the Panamanian isthmus and D. bellus differentiated there; later, during the Pleistocene, D. bellus entered South America and experienced speciation. The same process of re-ingression has been proposed to other xenarthrans, breaking with the traditional assumption that the GABI was unidirectional.

New sylvatic hosts of Trypanosoma cruzi and their reservoir competence in the humid Chaco of Argentina: a longitudinal study.

A four-year longitudinal study of the structure of sylvatic transmission cycles of Trypanosoma cruzi, reservoir host competence and parasite discrete typing units was conducted in a disturbed rural area of the humid Chaco in Argentina. Among 190 mammals examined by xenodiagnosis and polymerase chain reaction amplification, the composite prevalence of infection was substantially higher in Dasypus novemcinctus armadillos (57.7%) and Didelphis albiventris opossums (38.1%) than in Euphractus sexcinctus (20.0%), Tolypeutes matacus (12.5%), and Chaetophractus vellerosus (6.3%) armadillos. Trypanosoma cruzi was detected for the first time in Thylamys pusilla small opossums and in two unidentified small rodents. Infection was spatially aggregated only in armadillos. All Didelphis were infected with T. cruzi I and all armadillo species were infected with T. cruzi III, implying two distinct sylvatic cycles with no inputs from the domestic cycle. Dasypus armadillos and Didelphis opossums were much more infectious to vectors than other armadillos, small opossums, or rodents.

Mycobacterium leprae in six-banded (Euphractus sexcinctus) and nine-banded armadillos (Dasypus novemcinctus) in Northeast Brazil

Human beings are the main reservoir of the causative agent of leprosy, Mycobacterium leprae. In the Americas, nine-banded armadillos (Dasypus novemcinctus) also act as a reservoir for the bacillus. In the state of Ceará (CE), which is located in Northeast Brazil and is an endemic area of leprosy, there are several species of armadillos, including D. novemcinctus and Euphractus sexcinctus (six-banded armadillo). Contact between humans and armadillos occur mainly through hunting, cleaning, preparing, cooking and eating. This study identified M. leprae DNA in the two main species of armadillos found in Northeast Brazil. A total of 29 wild armadillos (27 D. novemcinctus and 2 E. sexcinctus) were captured in different environments of CE countryside. Samples from the ear, nose, liver and spleen from each of these animals were tested by a nested M. leprae-specific repetitive element polymerase chain reaction assay. The samples that tested positive were confirmed by DNA sequencing. M. leprae was detected in 21% (6/29) of the animals, including five D. novemcinctus and one E. sexcinctus. This is the first Brazilian study to identify the presence of a biomarker of M. leprae in wild armadillos (D. novemcinctus and E. sexcinctus) in a leprosy hyperendemic area where there is continuous contact between humans and armadillos.

Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos.

Fairy armadillos or pichiciegos (Xenarthra, Dasypodidae) are among the most elusive mammals. Due to their subterranean and nocturnal lifestyle, their basic biology and evolutionary history remain virtually unknown. Two distinct species with allopatric distributions are recognized: Chlamyphorus truncatus is restricted to central Argentina, while Calyptophractus retusus occurs in the Gran Chaco of Argentina, Paraguay, and Bolivia. To test their monophyly and resolve their phylogenetic affinities within armadillos, we obtained sequence data from modern and museum specimens for two mitochondrial genes (12S RNA [MT-RNR1] and NADH dehydrogenase 1 [MT-ND1]) and two nuclear exons (breast cancer 1 early onset exon 11 [BRCA1] and von Willebrand factor exon 28 [VWF]). Phylogenetic analyses provided a reference phylogeny and timescale for living xenarthran genera. Our results reveal monophyletic pichiciegos as members of a major armadillo subfamily (Chlamyphorinae). Their strictly fossorial lifestyle probably evolved as a response to the Oligocene aridification that occurred in South America after their divergence from Tolypeutinae around 32 million years ago (Mya). The ancient divergence date (∼17Mya) for separation between the two species supports their taxonomic classification into distinct genera. The synchronicity with Middle Miocene marine incursions along the Paraná river basin suggests a vicariant origin for pichiciegos by the disruption of their ancestral range. Their phylogenetic distinctiveness and rarity in the wild argue in favor of high conservation priority.

Mycobacterium leprae in six-banded (Euphractus sexcinctus) and nine-banded armadillos (Dasypus novemcinctus) in Northeast Brazil.

Human beings are the main reservoir of the causative agent of leprosy, Mycobacterium leprae. In the Americas, nine-banded armadillos (Dasypus novemcinctus) also act as a reservoir for the bacillus. In the state of Ceará (CE), which is located in Northeast Brazil and is an endemic area of leprosy, there are several species of armadillos, including D. novemcinctus and Euphractus sexcinctus (six-banded armadillo). Contact between humans and armadillos occur mainly through hunting, cleaning, preparing, cooking and eating. This study identified M. leprae DNA in the two main species of armadillos found in Northeast Brazil. A total of 29 wild armadillos (27 D. novemcinctus and 2 E. sexcinctus) were captured in different environments of CE countryside. Samples from the ear, nose, liver and spleen from each of these animals were tested by a nested M. leprae-specific repetitive element polymerase chain reaction assay. The samples that tested positive were confirmed by DNA sequencing. M. leprae was detected in 21% (6/29) of the animals, including five D. novemcinctus and one E. sexcinctus. This is the first Brazilian study to identify the presence of a biomarker of M. leprae in wild armadillos (D. novemcinctus and E. sexcinctus) in a leprosy hyperendemic area where there is continuous contact between humans and armadillos.

Phylogeography of the armadillo Chaetophractus villosus (Dasypodidae Xenarthra): post-glacial range expansion from Pampas to Patagonia (Argentina).

We report a phylogeographic study of Chaetophractus villosus populations in Argentina. Control Region (CR) sequences (484 bp) were obtained for 76 C. villosus from 20 locations across the species whole distribution range. Seventeen new haplotypes were identified. The highest genetic variation and the earliest fossils were found in the Pampean Region, thus appearing as the most probable area of origin of the species. A general pattern of Contiguous Range Expansion (CRE) was revealed by Nested Clade Analysis (NCA) supported by mismatch analysis and Fu's test. The Pampean Region would have been the pre-expansion area, while Patagonia would have been the main dispersal route of contiguous expansion, possibly after the Pleistocenic glaciations.

Sexual dimorphism in the mandible of the armadillo Chaetophractus villosus (Desmarest, 1804) (Dasypodidae) from northern Patagonia, Argentina.

The aim of this study was to study the sexual dimorphism in adult Chaetophractus villosus (Desmarest, 1804), from northern Patagonia, Argentina. Eight mandibular traits were measured in 37 males and 34 females. Univariate and multivariate morphometric analysis were applied to the data set. Results showed that C. villosus was sexually dimorphic, with higher absolute values corresponding to females. The total length of the mandible was the most important variable to discriminate sexes, followed by the height at the level of the last tooth and body length. The percentages of sex discrimination were high, as they were when a new sample (17 males, 13 females) was tested. Females have larger mandibles than males independently of their larger cranial size. They also showed a higher degree of correlation between variables, suggesting a more stable shape for the mandible than in males.

Sexual dimorphism in the mandible of the armadillo Chaetophractus villosus (Desmarest, 1804) (Dasypodidae) from northern Patagonia, Argentina / Dimorfismo sexual da mandíbula do tatu Chaetophractus villosus (Desmarest, 1804) (Dasypodidae) do norte da Patagônia Argentina

The aim of this study was to study the sexual dimorphism in adult Chaetophractus villosus (Desmarest, 1804), from northern Patagonia, Argentina. Eight mandibular traits were measured in 37 males and 34 females. Univariate and multivariate morphometric analysis were applied to the data set. Results showed that C. villosus was sexually dimorphic, with higher absolute values corresponding to females. The total length of the mandible was the most important variable to discriminate sexes, followed by the height at the level of the last tooth and body length. The percentages of sex discrimination were high, as they were when a new sample (17 males, 13 females) was tested. Females have larger mandibles than males independently of their larger cranial size. They also showed a higher degree of correlation between variables, suggesting a more stable shape for the mandible than in males.

O presente estudo examinou o dimorfismo sexual dos adultos de Chaetophractus villosus (Desmarest, 1804) do norte da Patagônia Argentina. Oito parâmetros mandibulares foram medidos em 37 machos e 34 fêmeas. Análises morfométricas univariada e multivariada foram aplicadas à série de dados. Os resultados mostraram que C. villosus apresenta dimorfismo sexual, com todos os valores absolutos mais elevados nas fêmeas. O comprimento total da mandíbula foi a variável mais importante para discriminar os sexos, seguidos pela altura no nível dos últimos dentes e pelo comprimento do corpo da mandíbula. As porcentagens de discriminação dos sexos foram elevadas, quando uma amostra nova (17 machos, 13 fêmeas) foi testada. As fêmeas têm as mandíbulas maiores que os machos, independentemente de seu tamanho cranial maior. Mostraram também um grau mais elevado de correlação entre variáveis, sugerindo uma forma mais estável para a mandíbula que nos machos.

Diversity, seasonality, and context of mammalian roadkills in the southern Great Plains.

Thousands of mammals are killed annually from vehicle collisions, making the issue an important one for conservation biologists and environmental managers. We recorded all readily identifiable kills on or immediately adjacent to roads in the southern Great Plains from March 2004-March 2007. We also recorded distance traveled, whether a road was paved or divided, the number of lanes, and prevailing habitat. Surveys were opportunistic and were conducted by car during conditions of good visibility. Over our 239 surveys and > 16,500 km traveled, we recorded 1412 roadkills from 18 different mammal species (size ranged from Sciurus squirrels to the white-tailed deer, Odocolieus virginianus). The overall kill rate was 8.50/100 km. Four species were prone to collisions: the Virginia opossum (Didelphis virginiana), nine-banded armadillo (Dasypus novemcinctus), striped skunk (Mephitis mephitis), and northern raccoon (Procyon lotor). Together they accounted for approximately 85% (1198) of all roadkills. Mortality rate differed significantly between 2- and 4-lane roads (8.39 versus 7.79/100 km). Kill rates were significantly higher on paved versus unpaved roads (8.60 versus 3.65/100 km), but did not depend on whether a road was divided. Roadkills were higher in spring than in fall (1.5 x), winter (1.4x), or summer (1.3x). The spring peak (in kills/100 km) was driven chiefly by the armadillo (2.76 in spring/summer versus 0.73 in autumn/winter) and opossum (2.65 versus 1.47). By contrast, seasonality was dampened by a late winter/early spring peak in skunk mortalities, for which 41% occurred in the 6-week period of mid-February through March. The raccoon did not exhibit a strong seasonal pattern. Our data are consistent with dispersal patterns of these species. Our results underscore the high rate of highway mortality in the southern plains, as well as differences in seasonality and road type that contribute to mortality. Conservation and management efforts should focus on creating underpasses or using other means to reduce roadkill rates.

Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyses of mitochondrial and nuclear genes.

The 30 living species of armadillos, anteaters, and sloths (Mammalia: Xenarthra) represent one of the three major clades of placentals. Armadillos (Cingulata: Dasypodidae) are the earliest and most speciose xenarthran lineage with 21 described species. The question of their tricky phylogeny was here studied by adding two mitochondrial genes (NADH dehydrogenase subunit 1 [ND1] and 12S ribosomal RNA [12S rRNA]) to the three protein-coding nuclear genes (alpha2B adrenergic receptor [ADRA2B], breast cancer susceptibility exon 11 [BRCA1], and von Willebrand factor exon 28 [VWF]) yielding a total of 6869 aligned nucleotide sites for thirteen xenarthran species. The two mitochondrial genes were characterized by marked excesses of transitions over transversions-with a strong bias toward CT transitions for the 12S rRNA-and exhibited two- to fivefold faster evolutionary rates than the fastest nuclear gene (ADRA2B). Maximum likelihood and Bayesian phylogenetic analyses supported the monophyly of Dasypodinae, Tolypeutinae, and Euphractinae, with the latter two armadillo subfamilies strongly clustering together. Conflicting branching points between individual genes involved relationships within the subfamilies Tolypeutinae and Euphractinae. Owing to a greater number of informative sites, the overall concatenation favored the mitochondrial topology with the classical grouping of Cabassous and Priodontes within Tolypeutinae, and a close relationship between Euphractus and Chaetophractus within Euphractinae. However, low statistical support values associated with almost equal distributions of apomorphies among alternatives suggested that two parallel events of rapid speciation occurred within these two armadillo subfamilies.

Molecular phylogeny of living xenarthrans and the impact of character and taxon sampling on the placental tree rooting.

Extant xenarthrans (armadillos, anteaters and sloths) are among the most derived placental mammals ever evolved. South America was the cradle of their evolutionary history. During the Tertiary, xenarthrans experienced an extraordinary radiation, whereas South America remained isolated from other continents. The 13 living genera are relics of this earlier diversification and represent one of the four major clades of placental mammals. Sequences of the three independent protein-coding nuclear markers alpha2B adrenergic receptor (ADRA2B), breast cancer susceptibility (BRCA1), and von Willebrand Factor (VWF) were determined for 12 of the 13 living xenarthran genera. Comparative evolutionary dynamics of these nuclear exons using a likelihood framework revealed contrasting patterns of molecular evolution. All codon positions of BRCA1 were shown to evolve in a strikingly similar manner, and third codon positions appeared less saturated within placentals than those of ADRA2B and VWF. Maximum likelihood and Bayesian phylogenetic analyses of a 47 placental taxa data set rooted by three marsupial outgroups resolved the phylogeny of Xenarthra with some evidence for two radiation events in armadillos and provided a strongly supported picture of placental interordinal relationships. This topology was fully compatible with recent studies, dividing placentals into the Southern Hemisphere clades Afrotheria and Xenarthra and a monophyletic Northern Hemisphere clade (Boreoeutheria) composed of Laurasiatheria and Euarchontoglires. Partitioned likelihood statistical tests of the position of the root, under different character partition schemes, identified three almost equally likely hypotheses for early placental divergences: a basal Afrotheria, an Afrotheria + Xenarthra clade, or a basal Xenarthra (Epitheria hypothesis). We took advantage of the extensive sampling realized within Xenarthra to assess its impact on the location of the root on the placental tree. By resampling taxa within Xenarthra, the conservative Shimodaira-Hasegawa likelihood-based test of alternative topologies was shown to be sensitive to both character and taxon sampling.

Comparative morphologic placental types in Dasypodidae (Chaetophractus villosus, Cabassous chacoensis, Tolypeutes matacus and Dasypus hybridus).

Information about the morphology of placentas in armadillos is scarce, except for D. novemcinctus. A comparative study of morphologic placental types in armadillos is important in order to have a comprehensive view of the peculiar reproductive physiology in this family. The aim of this paper is to perform a comparative analysis of the morphological features of the placenta in Chaetophractus villosus, Cabassous chacoensis, Tolypeutes matacus and Dasypus hybridus in order to classify them in accordance with Grosser (1909). The placentas were studied macroscopically and histologically (light microscopy in 1 micron thick sections and electron microscopy for fine structure). The macroscopic study in the 4 studied species showed a similar pear-shaped placenta homogeneously villosus in almost all the surface. The histological analysis showed that the 4 studied species had a hemochorial type of placenta. This type of placenta was also found in D. novemcinctus (Dasypodidae), but it is different from those described for other xenarthrans. Hemochorial types of placenta have also been described in more modern mammals. Despite the many primitive features of the armadillos and the different anatomical and physiological features between the genuses of dasypodids, all the studied species share this structural type of placenta.

Comparative morphologic placental types in Dasypodidae (Chaetophractus villosus, Cabassous chacoensis, Tolypeutes matacus and Dasypus hybridus)

Information about the morphology of placentas in armadillos is scarce, except for D. novemcinctus. A comparative study of morphologic placental types in armadillos is important in order to have a comprehensive view of the peculiar reproductive physiology in this family. The aim of this paper is to perform a comparative analysis of the morphological features of the placenta in Chaetophractus villosus, Cabassous chacoensis, Tolypeutes matacus and Dasypus hybridus in order to classify them in accordance with Grosser (1909). The placentas were studied macroscopically and histologically (light microscopy in 1 micron thick sections and electron microscopy for fine structure). The macroscopic study in the 4 studied species showed a similar pear-shaped placenta homogeneously villosus in almost all the surface. The histological analysis showed that the 4 studied species had a hemochorial type of placenta. This type of placenta was also found in D. novemcinctus (Dasypodidae), but it is different from those described for other xenarthrans. Hemochorial types of placenta have also been described in more modern mammals. Despite the many primitive features of the armadillos and the different anatomical and physiological features between the genuses of dasypodids, all the studied species share this structural type of placenta.(AU)