Priodontes maximus (Kerr, 1792) Giant Armadillo Dasypus maximus Kerr, 1792:112. Type locality ���Cayenne,��� French Guiana. Dasypus giganteus ��. Geoffroy Saint-Hilaire, 1803:207. Type locality ���Le Paraguay ��� with reference to ��� Le Grand Tatou d���Azara, t. 2, p. 132;��� but based on specimen ���N˚. CCCCXIV. Individu qui servi de sujet pour la description pr��c��dente. ��� Dasypus gigas G. Cuvier, 1817:221. No type locality given; based on plate ���xlv��� [error for ���xli���], the ��� Autre Kabassou ��� of volume 10 of Buffon (Daubenton in Buffon 1763); Buffon wrote that ��� Le Kabassou ��� was the largest tatou and came from Cayenne, which is the type locality. D [asypus]. gigans Schmid, 1818:164. No type locality given. T [atus]. grandis Olfers, 1818:219. Type locality ��� Paraguay.��� Priodontes giganteus: Lesson, 1827:309. Name combination. D [asypus]. (P [riodontes].) Gigas: Voigt, 1831:261. Name combination. Priodontes gigas: Gray, 1843:120. Name combination. Priodon gigas: Owen, 1845:21. Name combination. Prionodontes gigas: Schinz, 1845:316. Name combination. Prionodos gigas: Gray, 1865:374. Name combination. Prionodon gigas: Gray, 1869:380. Name combination. Cheloniscus gigas: Fitzinger, 1871:227. Name combination. Priodontes maximus: O. Thomas, 1880:402. First use of current name combination. Priodon maximus: J. A. Allen, 1895:187. Name combination. D [asypus]. maximus Larra��aga, 1923:343. Type locality ���nemoribus septentrionalibus paraquarensibus;��� based on Azara���s (1802:110) ��� Maximo;��� a junior homonym and synonym of Dasypus maximus Kerr, 1792. Periodontes maximus: Altrichter, 2006:2729. Name combination. CONTEXT AND CONTENT. Context as for genus. Priodontes maximus is monotypic; synonymy was reformatted from Wetzel et al. (2008). NOMENCLATURAL NOTES. As noted by Wetzel et al. (2008), Kretzoi and Kretzoi (2000:204) designated Dasypus giganteus ��. Geoffroy Saint-Hilaire, 1803, as the type species of Loricatus Desmarest, 1804, which relegated Priodontes F. Cuvier, 1825, to a synonym of Loricatus. Nevertheless, we followed Wetzel et al. (2008:153) who concluded that Loricatus niger Desmarest, 1804 (= Dasypus novemcinctus Linnaeus, 1758) was the valid type species of Loricatus, thus ���retain[ing] Priodontes as the valid generic name for P. maximus.��� Common names of P. maximus include giant armadillo and giant South American armadillo (English); tat�� carreta and tat�� gigante (Spanish); tatu-a���� and tatu canastra (Portuguese); gran tat�� de los bosques, priodonte, priodonte gigante (in Argentina); pejiche and pejichi (Bolivia); ja���ta (Yucuna tribe), jusa trueno and ocarro (Colombia); armadillo gigante, armadillo trueno, cutimbo, and tat�� gigante (Ecuador); carachupa gigante, carachupa maman, kint��ro, and yungunturu (Peru); cachicamo grande, cuspa, cuspa gigante, cuspa grande, and cusp��n (Venezuela); tatou-g��ant (French Guiana); tatu-guaz�� (Guarani); mowoor��mah (Guiana); granman-kapasi (Suriname); and el m��ximo napnalu, tatu-ass��, etopicnic lat��, carreta madre, and carachupa guazu (e.g., Superina and Aguiar 2006; Smith 2007; Trujillo and Superina 2013). DIAGNOSIS Priodontes maximus is by far the largest species of extant armadillos (Emmons and Feer 1997; Nowak 1991; Fig. 1). Its carapace extends only about halfway down its sides���seemingly draped on its back���whereas carapaces of other armadillos appear to wrap around their sides and flanks (Smith 2007). Head of P. maximus is relatively small compared with other species of armadillos and is somewhat domed with a long snout, narrow but blunt on the end, and with widely separated, small, and short ears (Fitzinger 1871; K��hlhorn 1939). Priodontes maximus and 4 species of naked-tailed armadillos (Cabassous) are in the tribe Priodontini (Wetzel et al. 2008), and only a naked-tailed armadillo could be confused for an immature P. maximus (Meritt 1985, 2006; Smith 2007; Trujillo and Superina 2013). Adult size is diagnostic: P. maximus, headbody length> 700 mm, greatest length of skull> 170 mm and Cabassous, head���body length Cabassous implies, their tails lack scutes and ���may or may not have visible scales,��� whereas the tail of P. maximus is ���armored with articulating bony scutes��� (Wetzel 1985a:15). P. maximus has more upper and lower teeth (about 18/19) that are more laterally flattened and broader than those in species of Cabassous (about 9/8���Wetzel 1985a, 1985b). GENERAL CHARACTERS An adult male Priodontes maximus can weigh up to 60 kg (80 kg in captivity), but average adult weight is about 30 kg (Nowak 1991; Redford and Eisenberg 1992; Smith 2007; Wetzel et al. 2008). Sexual dimorphism of P. maximus was reported for 7 of 14 body measurements of 5 males and 2 females from Emas National Park, Brazil: males were larger than females in weight, total length, head���body length, tail length, and circumferences of head, neck, and thorax (Silveira et al. 2009). Carapace of P. maximus is divided into transverse bands of small plates; 3���4 plates on the back of the neck and the central 11���13 plates are quite flexible (Nowak 1991; Wetzel et al. 2008). Pelage of P. maximus is limited to a few beige hairs between the plates. Color is dark brown to black dorsally, with a broad lightcolored band around the lower part of the carapace (Fig. 2). An individual P. maximus can be identified by the scale pattern on its carapace, particularly where dark and light scales meet, and the number of light and dark scales in each band (Noss et al. 2004; Fig. 2). Head shield of P. maximus is oval and does not expand between eyes. Tail is about 500 mm (Wetzel et al. 2008) and covered with small, closely set, rounded plates that are not arranged in rows (Wetzel 1985a, 1985b). Priodontes maximus can balance itself on its large hind feet, with its tail forming a stable tripod while digging (Nowak 1991). Simultaneous use of forefeet permits a P. maximus to excavate a hole quickly to escape (Milne et al. 2009). Unlike smaller armadillos, P. maximus cannot enclose itself into a tight ball within its carapace, so it rarely attempts to do so. Claws on forefeet are thick and powerful; claw on the 3rd forefinger is greatly enlarged, well developed, and sickle-shaped, measuring up to 20.3 cm along the curve���the largest claw of any living mammal (Fitzinger 1871; Smith 2007). DISTRIBUTION Priodontes maximus is found in Brazil, Paraguay, Guyana, Suriname, French Guiana, and east of the Andes in Colombia, Venezuela, Ecuador, Bolivia, and Peru (Meritt 1973; Emmons and Feer 1997; Vaz 2003; Anacleto et al. 2006, 2014; Zapata- R��os et al. 2006; Smith 2007; Meritt 2008; Wetzel et al. 2008; Srbek-Araujo et al. 2009; Porfirio et al. 2012; Zimbres et al. 2012; Fig. 3). In Argentina, P. maximus is probably limited to the northern provinces of Salta, Formosa, Chaco, and Santiago del Estero, as far south as 31��S (Torres and Jayat 2010); further south, temperatures are too cold (McNab 1980). Presence of P. maximus in the Argentinian provinces of C��rdoba, Santa Fe, and Misiones is uncertain (Parera 2002; Massoia et al. 2006), in part because records from C��rdoba are old; new surveys are needed in these provinces (Abba et al. 2012). Although the distribution of P. maximus extends over a large area of South America, it occurs in low-density, discontinuous populations in most areas (Cabrera 1957; Meritt 2006; see ���Conservation��� section). FOSSIL RECORD Ancestral lineages within Cingulata diverged from sloths and anteaters close to the Cretaceous���Paleogene transition about 66 million years ago, and armadillos diversified thereafter in the early to middle Eocene and beyond (Kurt��n 1972; Engelmann 1985; Delsuc et al. 2001, 2002; Croft et al. 2007; Vizca��no et al. 2008; Vizca��no and Bargo 2014; Delsuc et al. 2016). Armored species in the order Cingulata traditionally were placed in 2 extinct families, Pampatheriidae and Glyptodontidae (both were browsers and grazers), and 1 extant family, Dasypodidae (primarily insect-eaters���Fernicola et al. 2008). New phylogenetic evidence concludes that there are 1 extinct family, Pampatheriidae, and 2 extant families, Dasypodidae (dasypodines) and Chlamyphoridae (traditionally including euphractines, chlamyphorines, and tolypeutines) that diverged about 42 million years ago (Gibb et al. 2015; Delsuc et al. 2016). Extinct glyptodonts and pampatheres had been thought to represent a monophyletic clade, sister to dasypodines (Engelmann 1985; Gaudin and Wible 2006); however, new evidence places glyptodonts as a subfamily within Chlamyphoridae (Euphractinae + Glyptodontinae + Chlamyphorinae + Tolypeutinae [including Priodontes]��� Delsuc et al. 2016). Fossil record of Cingulata is rich, with ��� 65 genera of extinct glyptodonts and ��� 35 genera of armadillos and pampatheres (Fernicola et al. 2008). During the Pleistocene, some North American pampatheres evolved in Florida: for example, Holmesina septentrionalis (Simpson 1930) from H. floridanus (Edmund 1985). At least 2 species of giant glyptodont persisted as part of the North American megafauna through the late Pleistocene: Glyptotherium cylindricum as far north as Sonora, Mexico (Ram��rez-Cruz and Montellano-Ballesteros 2014) and G. floridanum in Hunt County, Texas (Gillette and Ray 1981). They may have become extinct because of climate change or human hunting (Cione et al. 2009). Armadillo fossils from the late Paleocene were found in Patagonia, and many species of armadillos, ranging from insecteaters to plant-eaters, were present in South America by the early to middle Eocene (Kurt��n 1972; Delsuc et al. 2016). In the Neogene, some cingulates became large, and by the Pliocene and Pleistocene, species of Pampatherium were the size of rhinoceroses (Kurt��n 1972; Edmund 1985). Glyptodonts arose in the Eocene and diversified greatly in the Miocene and Pliocene; some glyptodonts were gigantic reaching 4 m in length, with their nearly fused plates resulting in their ���inflexible structure��� and common name ���mammalian tortoise��� (Kurt��n 1972:178��� 179). Despite substantial differences in size, comparisons of ���deep and superficial cortexes��� of the osteoderm support close phylogenetic relationships among pampatheres, glyptodonts, and tolypeutine armadillos, including P. maximus (Wolf et al. 2012:388). Priodontes does not have a documented fossil record (de Paula-Couto 1979; McKenna and Bell 1997; Wetzel et al. 2008), but the extinct dasypodid Eutatus had species as large as P. maximus (e.g., E. seguini). They were less fossorial than P. maximus and herbivorous (Vizca��no et al. 2003). E. seguini and the large glyptodont Doedicurus clavicaudatus were common in the late Pliocene to the early Holocene in South America and found at 7,000- to 7,500-year-old archaeological sites in Argentina, confirming early use of cingulates as food by human hunter-gatherers (Fidalgo et al. 1986; Politis and Guti��rrez 1998). FORM AND FUNCTION Skull of an adult Priodontes maximus is ��� 180 mm (Fig. 4). P. maximus can have as many as 100 teeth, the greatest number of teeth of any terrestrial mammal (Ungar 2010). The teeth are hypselodont (shallow-rooted, high-crowned, and ever-growing), have a thin outer layer of acellular cementum (Green and Kalthoff 2015), and lack enamel. All teeth are molariform but not easily differentiated as premolars and molars (Ungar 2010); they are small, with thin blades, twice as long anteroposteriorly as wide, numbering 15���26 per row (Fitzinger 1871; K��hlhorn 1939; Wetzel 1985b). Teeth are not replaced, and toothrows do not articulate and play little role in catching, grasping, or chewing prey (K��hlhorn 1939). Tongue of P. maximus is long and vermiform, which is an adaptation to a diet of small insects; tongue of a recently dead Priodontes was 16 cm (K��hlhorn 1939). Salivary glands of Priodontes extend from under the neck to the sternum; location of submaxillary glands differs from that in other mammals because of the large size and slender head of Priodontes (K��hlhorn 1939). Billet et al. (2015) described the shape of the bony labyrinth of the inner ear of P. maximus; unlike other morphological evidence, bony labyrinth characteristics did not support a Priodontes���Cabassous clade (Billet et al. 2015). Limb measurements related to interspecific differences in digging abilities of armadillos have been assessed; those of P. maximus include (mm �� SD): maximum humeral length, 119.8 �� 4.3; ulnar length, 132 �� 6.4; olecranon length, 64.9 �� 8.8; and rear leg length, 85.5 �� 6.5 (Vizca��no and Milne 2002:tables 2 and 3; Milne et al. 2009)���slightly smaller lengths were provided by Vizca��no et al. (1999). Aside from the small and highly fossorial pink fairy armadillo (Chlamyphorus truncatus), an index of fossorial ability (i.e., length of olecranon process divided by difference between ulnar length and olecranon length) was highest for P. maximus (101.17 �� 34.5) and other species in the tribe Priodontini (92.86 �� 6.04���Vizca��no and Milne 2002). P. maximus walks on the tips of its strongly developed 3rd claws on its forefeet. Worn surfaces of claws suggest that, in contrast to Tolypeutes, P. maximus slides rather than stalks on its forehands (Krieg 1961). Sperm shapes are unique among 4 groups of extant armadillos. P. maximus and the related southern naked-tailed armadillo (Cabassous unicinctus) group together by their sperm shapes: heads are large, long, and frontally wide, with extremely thin profiles and overall paddle shapes (Cetica et al. 1993; Cetica and Merani 2008). Other characteristics of sperm of both species include sperm head lengths of 13.2���18.0 ��m (with acrosomes occupying 74���75% of total sperm head length), sperm head widths of 11.6���16.0 ��m, and total sperm lengths of 79.9��� 98.0 ��m (Cetica and Merani 2008). The pelletized feces of P. maximus are dry and firm, with mean widths of 14.7 mm �� 1.7 SD, mean lengths of 22.7 mm �� 3.0 SD, and mean weights of 2.8 g �� 0.9 SD (Anacleto 2007). Slow and Rapid Eye Movement (REM) sleep patterns and electrical activity of flexor muscles in the neck of P. maximus are typical of mammalian patterns (Affanni et al. 1972). ONTOGENY AND REPRODUCTION A female Priodontes maximus has 2 mammae (Fitzinger 1871); extrudes a watery, bloody fluid from her vulva during estrus (T. S. Carter, in litt.); has 1 (sometimes 2) young per year (Krieg 1929); and lactates for 4���6 months (Neris et al. 2002). Other reports on basic reproductive characteristics of P. maximus seem to be incorrect. According to Merrett (1983), sexual maturity of male and female P. maximus is reached at 9���12 months, gestation is about 4 months, offspring are born with a body mass of 113 g, and weaning occurs at 4���6 weeks. Although these reproductive data have been widely cited, they are likely inaccurate for an armadillo species with the mass of P. maximus, and they cannot be substantiated in other scientific literature or from captive records because P. maximus has never reproduced in captivity (Aya-Cuero et al. 2015). Other armadillo species with similar litter size have birth weights of 6���7% of adult body mass (Superina and Loughry 2012); a neonatal P. maximus is therefore expected to have a body mass of 1.9���3.5 kg (Aya-Cuero et al. 2015). Generation length has been estimated at 6���10 years (Fonseca and Aguiar 2004) and 7 years (Anacleto et al. 2014), but both are very rough estimates given the lack of life-history data for P. maximus. Information on the duration of lactation provided by Neris et al. (2002) is uncertain because it seems to be based on anecdotal reports from indigenous tribes (Aya-Cuero et al. 2015). Nevertheless, it coincides with camera-trap photographs of a juvenile P. maximus estimated at 4���5 months of age that was repeatedly observed with its mother; several photographs taken 3 months later at an estimated age of 7���8 months showed the same individual without its mother, suggesting that weaning had occurred (Aya-Cuero et al. 2015). Parental care behavior was observed in another juvenile estimated at 7���8 months of age that shared the burrow with its mother but sometimes explored the surroundings alone (Aya-Cuero et al. 2015; Fig. 5). The female emerged from the burrow, inspected the environment by sniffing in bipedal posture, introduced its snout into the burrow entrance, and finally allowed its offspring to leave the burrow. The offspring then stood on its hind feet and supported its fore claws on its mother���s back for 2���5 min, a behavior that could be observed several times per day and may be interpreted as play behavior (Aya-Cuero et al. 2015). Adult P. maximus are presumably solitary most of the year, except during breeding and while a female rears her youngof-the-year. Videos of a wild female and her relatively young offspring emerging from a burrow (http://news.mongabay. com/2013/0219-hance-giant-armadillo-baby.html, accessed 27 August 2015) were filmed in February 2013 in Brazil. Next to nothing is known about sexual behavior of P. maximus, but recently a camera-trap image showed 2 adults apparently copulating, with the male mounting the female from the rear (photograph by Rachel Berzins, Office National de la Chasse et de la Faune Sauvage, Guyana; http://www.une-saison-en-guyane. com/article/faune/piegage-photo-de-loncfs-sur-le-centre-spatial-guyanais/, accessed 6 November 2015). Camera-trap videos from Colombia show a female P. maximus digging for 30 min to open the entrance of an existing burrow and an offspring emerging (https://www.youtube.com/watch?v=Q76K-txuWCo, accessed 19 October 2015). Aya-Cuero et al. (2015) hypothesized that the female left its offspring inside the burrow and closed the entrance to impede predator attacks. ECOLOGY Population characteristics. ���Little is known about population characteristics of Priodontes maximus. It has a widespread, discontinuous distribution and has never been numerous (Cabrera 1957; Meritt 2006). There are no longevity records of, Published as part of Carter, Tracy S., Superina, Mariella & Leslie, David M., 2016, Priodontes maximus (Cingulata: Chlamyphoridae), pp. 21-34 in Mammalian Species 48 (932) on pages 22-30, DOI: 10.1093/mspecies/sew002, http://zenodo.org/record/4573372, {"references":["KERR, R. 1792. The animal kingdom or zoological system, of the celebrated Sir Charles Linnaeus. class I. 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