Dasypus kappleri Krauss, 1862

Aya-Cuero, Carlos, Chacón-Pacheco, Julio & Anacleto, Teresa Cristina S, 2019, Dasypus kappleri (Cingulata: Dasypodidae), Mammalian Species 51 (977), pp. 51-60 : 51-55

Dasypus kappleri Krauss, 1862

Greater Long-nosed Armadillo

Dasypus peba: Cabanis, 1848:782 . Not Dasypus peba Desmarest, 1822 .

Das [ypus] [p] eba: Burmeister, 1848:199. Not Dasypus peba Desmarest, 1822 .

Dasypus (Praopus) peba: Burmeister,1854:301 . Not Dasypus peba Desmarest, 1822 .

Dasypus kappleri: Krauss, 1862:20 . Type locality “den Urwäldern des Marowiniflusse in Surinam,” which Husson (1978:261) suggested was probably in the neighborhood of Albina near the mouth of the Marowijne River. First use of current name combination.

Dasypus pentadactylus W. Peters, 1864:179 : Type locality “ Guiana; ” name based on a specimen identified by Cabanis (1848:782) as D. peba from “ am Demerara und auf der Savanne am Berbice,” Guyana.

Tatusia kappleri: Gray, 1865:373 . Name combination.

Praopus kappleri: Gray, 1873:16 . Name combination.

Tatusia kappleri: O. Thomas, 1880:402 . Name combination.

[ Tatusia (Tatusia) ] kappleri: Trouessart, 1898:1139 . Name

[ Tatusia (Tatusia) ] pentadactylus: Trouessart, 1898:1140 . Name combination. T [atu]. kappleri: O. Thomas, 1901:371 . Name combination. Tatu pastasae O. Thomas, 1901:370 . Type locality “Sarayacu,

[ Tatus (Tatus) ] kappleri: Trouessart, 1905:814 . Name combination.

[ Tatus (Tatus) ] pentadactylus: Trouessart, 1905:814 . Name combination.

[ Tatus (Tatus) ] pastasae: Trouessart, 1905:814 . Name combination.

Dasypus kappleri pastasae: Lönnberg, 1928:9 . Name combination.

D [asypus]. k [appleri]. peruvianus Lönnberg, 1928:10. Type locality “Roque in Eastern Peru, S. E of Moyobamba at an altitude of about 1030 m.,” San Martín, Peru.

Dasypus pastasae: Sanborn, 1929:258 . Name combination.

Dasypus kappleri beniensis Lönnberg, 1942:49 . Type locality “near the confluence of Madre de Dios River with Beni River, Victoria, [Pando] Bolivia.”

Dasypus [(Hyperoambon)] kappleri: Wetzel and Mondolfi, 1979:56 . Name combination.

CONTEXT AND CONTENT. Order Cingulata , family Dasypodidae , subfamily Dasypodinae , tribe Dasypodini (Möller-Krull et al. 2007; Wetzel et al. 2008). Two subspecies of D. kappleri are recognized by Cabrera (1958), Wilson and Reeder (2005), and Wetzel et al. (2008):

D. k. kappleri ( Krauss, 1862) . See above; peba ( Cabanis, 1848) , peba (Burmeister, 1848) , pentadactylus (W. Peters, 1864) are synonyms.

D. k. pastasae (O. Thomas, 1880) . See above; peruvianus ( Lönnberg, 1928) and beniensis ( Lönnberg, 1942) are synonyms.

NOMENCLATURAL NOTES. The generic name, Dasypus , meaning hairy or rough-footed, is a combination of two Greek words, dasy meaning hairy and pus (from pous) meaning foot ( Palmer 1904). The species name, kappleri , is in honor of August Kappler (1815–1887), German soldier and naturalist. Common names of D. kappleri include greater long-nosed armadillo, Kappler’s armadillo, great long-nosed armadillo (English); armadillo de Kappler, mulita de Kappler (Spanish); tatuquinze-quilos, tatu-tinga (Portuguese); Gordeldier, Langstaart gordeldier, Stekel-voetgordeldier (Dutch); tatú quinze quilos (in Bolivia); Cayurre (Sikuani tribe), Gouu (Muiname tribe), je’e wani (Yucuna tribe), armadillo espuelón, armadillo arracacho, cachicamo grande, jasuchula ( Colombia); armadillo narizón, tatú-peba grande ( Ecuador); carachupa ( Peru); cachicamo guayanés, cachicamo montañero gigante ( Venezuela); kapasi, kapasie, maka kapasi, platatèrè ( Surinam); grand tatou ( French Guiana — Staffeleu 1975; Husson 1978; Linares 1998; Emmons and Feer 1999; Superina and Aguiar 2006; Trujillo and Superina 2013).

The original designation by Burmeister (1848) of this species as D. peba did not result in a valid name related to D. kappleri because it had been used before by Desmarest (1822) referring to a specimen of Dasypus novemcinctus Linnaeus, 1758 . Cabanis (1848) apparently failed to distinguish this animal from D. novemcinctus , and his specimen was lumped together with examples of D. novemcinctus under Desmarest’s D. peba . Peters (1864), based on a description of a specimen obtained by Cabanis, defines D. pastasae and D. peruvianus as synonyms but they appear to be merely population variations. A specimen of D. kappleri examined in the Museu Paraense Emílio Goeldi (MPEG 041036) showed a haphazard mixture of the diagnostic differences between the types of Krauss and Thomas ( Hamlett 1939). Recently, Feijó and Cordeiro-Estrela (2016) mentioned that over the last century, related taxa had been described and synonymized without comprehensive analyses, and the current classification involved two subspecies (D. k. kappleri and D. k. pastasae ) that were never revised. They proposed recognizing three species with diagnosable autapomorphies, Dasypus kappleri Krauss, 1862 , Dasypus pastasae ( Thomas, 1901) , and Dasypus beniensis Lönnberg, 1942 . However, this hypothesis must be addressed with molecular studies, to analyze the relationships proposed by Feijó and Cordeiro-Estrela (2016).

DIAGNOSIS

Dasypus kappleri is a large armadillo exceeded in size among living armadillos only by the giant armadillo Priodontes maximus . Its mean (± SD) head–body length of 543.2 ± 22.2 mm is considerably larger than that of the nine-banded armadillo D. novemcinctus (460 ± 32 mm —Richard-Hansen et al. 1999) and the northern long-nosed armadillo D. sabanicola (265.7 ± 35.0 mm—Fergusson-Laguna 1984). The hind legs of D. kappleri exhibit prominent spurs that may exceed 17 mm in length ( Wetzel 1985). These spurs are absent in all other armadillos. It usually has eight (sometimes seven) movable bands. This is fewer bands in comparison with D. novemcinctus which typically has nine or the hairy long-nosed armadillo D. pilosus with 9–11 (usually 10) movable bands ( Castro 2015), and more than the southern long-nosed armadillo D. hybridus with usually seven movable bands (Abba and Superina 2016). D. kappleri can be recognized externally by its unique pattern of smooth, flattened, and uniform scales on the pelvic shield, with the central and peripheral scales at the same level (Feijó and Cordeiro-Estrela 2016), as well as the anterior border of the shield scapular sharply angular ( Castro 2015). The color of the carapace varies from uniform grayish or brownish-gray to having a darkened dorsum with a yellowish lateral stripe ( Fig. 1 ; Feijó 2017). D. kappleri is readily distinguished from other Dasypus by the characters of the pterygoid bone; the lateral borders of the palate, posterior to the tooth rows, form definite keels which extend ventrally to the back of the main surface of the posterior palate; the posterior margin of palate is straight ( Fig. 2 ). The rostrum is proportionally longer (62–67% of the length of skull) than in other Dasypus except for D. pilosus ( Tate 1939; Wetzel 1985; Castro 2015; Fig. 3 ). The tail is thick at the base and sharp at the tip, covered with circular rings formed by osteodermic plates ( Linares 1998; Castro 2015), and keeled scales on the proximal tail rings (Feijó and Cordeiro-Estrela 2016).

GENERAL CHARACTERS

Dasypus kappleri is the largest representative of Dasypus . Weight varies between 8.5 and 13 kg ( Wetzel 1985; Emmons and Feer 1999; Richard-Hansen et al. 1999; Castro 2015). Mean external measurements (mm ± SD, with parenthetical n) given by Wetzel and Mondolfi (1979) for adult specimens from Brazil, Colombia, Ecuador, Peru, Surinam, and Venezuela were: head–body length 543.2 ± 22.2 (18); tail length 406.1 ± 45.6 (15); length of hind foot 119.9 ± 8.4 (15); ear length 50.5 ± 3.1 (15); number of scutes in fourth movable band 56.4 ± 2.9 (28). Mean external measurements (mm ± SD) given by Richard-Hansen et al. (1999) for 53 adult specimens from French Guiana were: head–body length 557 ± 25; tail length 400 ± 3; shoulder height 291 ± 19; length of hind foot 120 ± 9; head length 145 ± 7; ear length 54 ± 6. All mean measurements in D. novemcinctus (n = 141) are significantly lower than those in D. kappleri (n = 53) including weight, head–body length, shoulder height, hind foot length, and head length (P <0.0001) and ear length (P <0.05—Richard-Hansen et al. 1999).

The maximum length of the skull 112.1–135.0 mm (Wetzel and Mondolfi 1979), condyle-nasal length is 123.4 ± 4.5 mm (mean ± SD, n = 45), the adjusted rostral length (distance along midline of rostrum from a line through lacrimal foramina to tip of nasal bone) is 79.0 ± 3.7 (n = 47); and the ratio of the adjusted rostral length to condyle-nasal length was 0.64 ± 0.01 (n = 7— Wetzel 1985). Cranially, D. kappleri has a prominent lateral palatine crest raised above the bony surface, and its posterior terminus is conspicuously swollen (Feijó and CordeiroEstrela 2016).

As all Cingulata , D. kappleri bears a carapace consisting of ossified dermal scutes (osteoderms) that are covered by epidermal scales ( Wetzel 1985). There are 7–8 movable bands in the middle of the body, with 51–62 osteoderms on the fourth band ( Castro 2015). Bands also separate the scapular and pelvic shields ( Wetzel 1985), the anterior edge on the shield has an angular form ( Castro 2015). The fixed osteoderms measure 16.9–18.9 mm in length and are 5.1–7.6 mm wide; the movable osteoderms measure 8.1–12.1 mm in length and are 5.5–7.3 mm wide ( Castro 2015). There are 3–8 foramens in the scapular shield and 7–18 in the pelvic shield ( Castro 2015). The mean number of movable bands given by Wetzel (1985) is 7.8 ± 0.4 SD (n = 33), other authors agree that there are 7–8 movable bands in the middle of the body ( Merrett 1983; Linares 1998; Castro 2015), but there may also be nine bands (Trujillo and Superina 2013). Fifty-three individuals from French Guiana revealed that the number of movable bands ranged from seven (25%) to nine (5%), with eight being the majority (70%—Richard-Hansen et al. 1999). The tail (325–483 mm) is very wide at the base, covered with scale-rings, and represents about 75–79% of head– body length ( Wetzel 1985; Linares 1998; Castro 2015). The snout is long and narrow ( Fig. 1 ; Wetzel 1985) and the face is pink. Ears are long (48–55 mm — Wetzel 1985; Emmons and Feer 1999) and without scales, almost touching at the base (Trujillo and Superina 2013). The claws of the front and hind legs are more developed than in D. novemcinctus (Emmons and Feer 1999; Tirira 2007). It is the only species of armadillo with projecting scutes (see “Behavior”) on the hind legs (Wetzel and Mondolfi 1979); scutes are arranged in transverse rows on the posterior surface of the legs ( Fig. 3A ).

Dasypus kappleri retains the fifth finger of the manus complete with metacarpals and phalanges ( Fig. 3B ), while the other species of Dasypus have a vestigial metacarpal ( Castro 2015). However, the use of this character as diagnostic of D. kappleri has been questioned, both Costa and Vizcaíno (2010) and Schulthess (1919) reported the presence of a fifth finger in the manus of some individuals of D. novemcinctus , and Galliari (2014) reported a vestigial fifth finger for D. hybridus .

According to Thomas (1901) and Lönnberg (1928) the subspecies D. k. kappleri and D. k. pastasae are similar in size and general appearance. But overall the scales are larger in D. k. kappleri . Scales in D. k. pastasae on the pelvic shield are of less uniform size and smoothness, the larger scales standing up more prominently above the level of the smaller ones ( Thomas 1901). On the other hand, the tail-scales, especially proximally, are smoother and flatter, the center of each line, although slightly keeled, is not raised into a prominent outwardly directed point, as is the case in D. k. kappleri ( Thomas 1901) . Skull characters are very similar for the two subspecies; however, the following characters differ according to Thomas (1901) and Feijó and Cordeiro-Estrela (2016): the nuchal crest is more strongly developed in D. k. pastasae , surpassing the external occipital crest posteriorly. The malar bones in D. k. kappleri are narrow in lateral view, while in D. k. pastasae malars are broader, projecting further downwards, such that the lower edge between the malars is flush with the palatal edge, rather than shorter than the palatal. The posterior palate in D. k. kappleri is deeply hollowed out, and its edges rise up as sharp bony ridges some 4–5 mm higher than its general level; in D. k. pastasae , on the other hand, the ridges are less developed, barely 2 mm high or less. Finally, at the posterior end of the palatal ridges the lateral walls of the choanae are hollow and considerably inflated, while in D. k. pastasae there is no inflation and the bones appear to be fairly solid.

Thomas (1901) pointed out two additional cranial differences between D. pastasae and D. kappleri . One is that the nuchal crest is more strongly developed in D. pastasae , surpassing the external occipital crest posteriorly. The second is that the jugal bones are broader in lateral view in D. pastasae , projecting further downwards. However, both traits exhibit substantial intraspecific variation and are unreliable for diagnosis.

DISTRIBUTION

The geographic range of Dasypus kappleri is restricted to South America ( Fig. 4 ). The extent of occurrence is about 5,500,000 km 2 (Abba and Superina 2010). It occurs east of the Andes in Colombia, south of the Orinoco in Venezuela to Guyana, Suriname, and French Guiana through the Amazon basin of Brazil, Ecuador, Peru, and northeastern Bolivia ( Wetzel 1982, 1985; Wetzel et al. 2008). D. k. kappleri occurs from southeastern Colombia, through southern Venezuela, in the Guianas, and the lower Amazon basin of Brazil, as far as Pará state and D. k. pastasae occurs in eastern Ecuador and Peru, northeastern Bolivia, and the upper Amazon basin of Brazil ( Wetzel et al. 2008). A significant difference between the distribution maps provided by Wetzel et al. (2008) and Anacleto et al. (2014) is due to a record of the species in the town of Capipi, near the border between the Brazilian states of Pará and Mato Grosso. However, the specimen in question (MZUSP [Museum of Zoology of the São Paulo University, São Paulo, Brazil] 8950) actually came from São Domingos do Capim, north of Pará ( Castro 2015). The easternmost locality is Barra do Garças city, southeast of Mato Grosso, in Cerrado biome (CM [Mammalian Collection at the University of Mato Grosso State] 779). D. kappleri has a potential niche distribution concentrated in the Amazon ( Anacleto et al. 2006).

FOSSIL RECORD

The oldest fossil record for a Dasypus was Dasypus bellus ( Tatus bellus Simpson, 1929 ), a specimen collected from the Blancan age deposits in North America, dating from about 2.4 million BP ( Castro et al. 2014). The fossil record for the genus Dasypus also includes the late Pleistocene and early Eocene of South America ( Vizcaíno et al. 1995; Soibelzon et al. 2013, 2015; Castro et al. 2014); however, this range does not apply to D. kappleri specifically ( Castro 2015). Oliveira and Pereira (2009) recorded an isolated pelvic buckler osteoderm, closely related to Dasypus (Hyperoambon) kappleri in Rio Grande do Sul, southern Brazil.

FORM AND FUNCTION

Form. — The permanent teeth are euhypsodont (evergrowing, high-crowned teeth, with an open pulp cavity), conical when first erupted, and become cylindrical with growth and wear ( Ciancio et al. 2012). Martin (1916) and Ciancio et al. (2010) confirmed the presence of enamel in Dasypus , and dental replacement in D. kappleri ( Castro et al. 2010) . Typical adult dentition of Dasypus consists of 8/8 homogeneous molariform teeth (Mf/mf). None of the upper teeth are placed in the premaxilla ( Ciancio et al. 2012). D. kappleri has Mf 7/mf 7–9 for a total of 28 to 36 teeth ( Linares 1998). The vertebral formula for a single specimen of D. kappleri examined in the collection of Naturalis Biodiversity Center (RMNH.MAM[National Natuurhistorisch Museum].20965) is 7C, 9T, 5L, 8S, and 18+coccygeal; presacral number is 21, for a total of 50 vertebrate (Varela-Lasheras et al. 2011).

Means of selected limb measurements (mm; mean ± SD) for four D. kappleri were: humeral length 78.7 ± 2.2; proximal humeral length 38 ± 1.7; ulna length 92.7 ± 3.6; olecranon length 36.7 ± 2.1; femur length 104.9 ± 2.1; proximal femoral length 58.9 ± 1.2; leg length 89.5 ± 1.2; and midleg width 33.2 ± 0.2 (Vizcaíno and Milne 2002).

Dasypus females have four inguinal mammary glands (Wetzel and Mondolfi 1979); furthermore, D. kappleri males lack penis connector with lateral enlargements, and its smaller terminal portion is blunt in contrast to the trifid appearance and the more pointed terminal portion as in D. novemcinctus , the seven-banded armadillo D. septemcinctus , D. sabanicola , and D. hybridus (Wetzel and Mondolfi 1979; Wetzel 1985).

Function. —Like all Xenarthra , Dasypus kappleri has poor thermoregulatory abilities and it has a low body temperature in comparison to other placental mammals ( McNab 1985), approximately 35°C (Trujillo and Superina 2013) in contrast with the typical mammalian temperature of 37–39°C ( Lopera 2004). Studies in D. novemcinctus reported that the carapace had high thermal conductance, resulting in rapid heat loss into the environment. In contrast, areas such as the tail and ears may have some ability to retain heat (Tettersall and Cadena 2010) and this is probably true for D. kappleri . According to FournierChambrillon et al. (2000a) these two species are biologically very similar.

ONTOGENY AND REPRODUCTION

Dasypus kappleri breeds during the dry season ( Linares 1998), and the litter size is typically one or two (Eisenberg and Redford 1999). It is generally assumed that polyembryony is common to all species of the genus Dasypus , although reproduction of D. kappleri has never been studied ( Superina et al. 2014b). Siblings are monozygotic of the same sex (Wetzel and Mondolfi 1979; Wetzel et al. 2008). Sets of embryo pairs of the same sex and unisexed embryos suggest a similar mode of reproduction but a smaller litter size in D. kappleri , as compared to D. septemcinctus or D. pilosus ( Galbreath 1985) .

Three sets of embryos of D. kappleri preserved in formalin in the Museo de la Estación Biológica de Rancho Grande (MEBRG) were from three females collected on January in Venezuela: Río Grande, El Palmar, and Amanza Guapo. Each set consisted of two fetuses of the same sex. As the fetal membranes and placentae were not preserved, it could not be confirmed that they were monovulvar twins. The Smithsonian Venezuelan Project has deposited in the United States National Museum (USNM) one record of a pregnant D. kappleri with two fetuses collected in January in the state of Amazonas, Cunucunuma River (Wetzel and Mondolfi 1979).

Figures

Fig. 1.—Male Dasypus kappleri inspecting the environment by sniffing in bipedal posture inside burrow in Grand Boeuf Mort, Guyane. Used with permission of the photographer Vincent Rufray/Biotope.

Fig. 2.—Dorsal, ventral, and lateral view of skull and lateral view of the mandible of Dasypus kappleri (EBRG [Estación Biológica Rancho Grande] 1417, female) from Amanza Guapo, Río Grande, Venezuela. Greatest length of skull is 126 mm.

Fig. 3.—Detail of the external diagnostic characteristics: A) spurs on the hind legs; B) fifth finger of the hand complete with metacarpal and phalanges of an adult Dasypus kappleri (EBRG [Estación Biológica Rancho Grande] 1417, female) from Amanza Guapo, Río Grande, Venezuela. Used with permission of the photographer Anderson Feijó.

Fig. 4.—Geographic distribution of Dasypus kappleri (modified from Anacleto et al. 2014).