Taxonomic revision of the Dasypus kappleri complex, with revalidations of Dasypus pastasae (Thomas, 1901) and Dasypus beniensis Lönnberg, 1942 (Cingulata, Dasypodidae) Author Feijó, Anderson Author Cordeiro-Estrela, Pedro text Zootaxa 2016 4170 2 271 297 journal article 10.11646/zootaxa.4170.2.3 5f293e28-4f71-4042-a192-6a23feb15b70 1175-5326 265398 9E3AE604-3656-4B0A-B09A-C970200BF5F6 Dasypus pastasae ( Thomas, 1901 ) Tatu pastasae Thomas, 1901 : 370 (original description) Dasypus kappleri pastasae : Lönnberg, 1928 (name combination) D [ asypus ]. k [ appleri ]. peruvianus Lönnberg, 1928 :10 Holotype . BM 80.5.6.71 collected by Mr. Clarence Buckley ( Figure 9 ). Type locality . “ Sarayacu , upper Pastasa River ”, Pastaza , Ecuador . Etymology. The name pastasae refers to the type locality. Diagnosis. The main diagnostic external features of D. pastasae lying on the scales of the pelvic shield and tail rings. This species has the non-uniform scales of the pelvic shield in size and texture (rough), where the central ones are larger and standing up above the level of the peripherals that are smaller. Further, the posterior scales of the proximal rings on tail are flattened. Cranially, the lateral margin of palatine has a prominent but unswollen crest and the posterior end of palatine is straight (not concave). Distribution. Dasypus pastasae is distributed from the foothills of the eastern Andes in Peru , Ecuador , Colombia , and Venezuela south of the Orinoco River into the western Brazilian Amazon, between the Madeira River and Branco River (Figure 10). Despite the absence of records from Bolivia , it is likely that D. pastasae also occurs in the extreme north of that country, on the left bank of Madre de Dios River. This species is sympatric with D. kappleri in eastern Venezuela . Measures. Dasypus pastasae is the smallest species of the D. kappleri complex, on average smaller than the other species in all 24 cranial measures ( Table 4 ). Remarks. 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. One pregnant female (EBRG 25559) with two fetuses was collected on 4 December 2005 from Ichum River, Salto Ichuno Espuma, Bolivar, Venezuela. TABLE 4. Cranial measurements (in mm) of Dasypus kappleri complex. All measurements are given as the mean ± standard deviation; minimum–maximum (N).

Measurements	Dasypus kappleri	Dasypus pastasae	Dasypus beniensis
GLS	127.86±4.74; 111.11–134.8(24)	121.14±5.18; 114.89–130.03 (16)	131.30±3.77; 122.91–134.63 (8)
CB	115.65±4.53; 100.29–123.94 (24)	110.31±4.85; 103.37–119.89 (16)	116.88±6.99; 100.09–121.33 (8)
APL	26.04±2.01; 21.09–31.84 (28)	23.71±2.8; 20.03–31.94 (20)	25.44±1.21; 22.87–26.9 (8)
PL	87.91±4; 75.77–95.48 (26)	82.99±4.72; 76.04–91.23 (18)	87.59±3.13; 82.24–90.8 (7)
ML	53.48±4.83; 45.77–67.79 (25)	48.04±3.6; 41.12–55.41 (18)	52.33±2.31; 48.93–55.11 (8)
PIL	25.81±2.22; 21.69–31.34 (25)	24.83±2.89; 19.82–30.63 (18)	25.86±1.29; 23.92–27.41 (7)
ICL	7.25±0.98; 5.63–9.3 (26)	7.58±1.09; 5.21–8.93 (22)	11.08±1.46; 9.07–13.05 (8)
MT	29.49±1.89; 24.94–33.51 (25)	29±1.89; 23.83–33.06 (19)	29.66±1.43; 27.47–31.67 (8)
NL	43.11±2.41; 34.94–46.5 (27)	42.94±3.14; 37.25–50.19 (19)	46.51±2.14; 44.32–49.5 (8)
LL	15.57±2.11; 10.29–19.14 (27)	13.15±1.61; 9.86–16.5 (20)	18.8±2.04; 15.30–21.14 (8)
RL	81.66±3.72; 69–88.38 (28)	76.16±4.25; 67.52–84.16 (20)	85.18±3.46; 77.95–89.5 (8)
AB	39.6±1.51; 35.97–41.71 (28)	36.18±1.79; 32.34–38.23 (19)	43.03±1.62; 40.68–45.52 (8)
TL	2.79±0.32; 2.13–3.4 (25)	2.57±0.3; 2.1–3.11 (18)	3.08±0.34; 2.63–3.49 (7)
PB	19.2±2.34; 11.33–21.74 (27)	18.89±1.16; 16.71–20.62 (20)	20.81±0.94; 19.49–21.91 (6)
PIB	21.19±0.88; 19.35–22.91 (27)	19.19±1; 17.35–21.5 (22)	19.39±0.67; 18.55–20.72 (8)
PC	27.47±0.94; 26.15–30.11 (27)	25.31±1.19; 23.01–27.52 (19)	29.75±1.79; 26.53–31.74 (8)
BB	37.93±1.43; 35.33–41.26 (27)	36.02±1.54; 33.06–38.5 (20)	38.08±1.24; 35.58–39.25 (8)
ZB	50.69±2.2; 42.61–53.53 (25)	47.2±2.48; 41.9–51.56 (14)	53.55±3.46; 48.14–58.05 (6)
MB	33.88±1.28; 31.63–37.21 (24)	32.84±1.28; 30.06–35.36 (17)	34.05±1.32; 31.44–35.33 (7)
HJ	9±1.12; 7.27– 12.15 (25)	8.8±0.84; 7.1–10.21 (16)	9.55±1.09; 8.51–11.52 (7)
MAL	101.29±3.73; 86.9–106.44 (25)	93.57±5.77; 80.32–102.08 (19)	102.66±3.64; 95.43–106.07 (7)
AML	24.01±2.13; 19.66–29.27 (27)	21.09±1.84; 16.62–23.62 (19)	23.19±1.51; 20.01–24.41 (7)
LMT	31.6±1.81; 27.59–35.02 (27)	30.05±1.75; 25.61–33.51 (19)	31.88±0.95; 30.78–33.59 (7)
HM	29.4±1.94; 25.1–32.57 (26)	27.14±1.89; 23.37–30.34 (18)	25.63±1.87; 23.55–28.83 (7)

Specimens examined (locality numbers as in Figure 10). BRAZIL : 11 ( MPEG 47762 ). COLOMBIA : ( AMNH 136253 , AMNH 136255 ), 21 ( IAVH 6858 , ICN sn), 22 ( ICN 1621 , ICN 1622 ), 23 (ICN 11625), 24 ( AMNH 136251 ). ECUADOR : 25 ( QCAZ 3370 , QCAZ 3376 ), 26 ( MEPN 7654 ), 27 ( MEPN 8496 , MEPN sn). PERU : 31 ( MUSM 697 ), 32 ( AMNH 98812 ), 33 ( MUSM 23073 ), 34 ( AMNH 98464 ), 35 ( AMNH 76574 , AMNH 76573 ), 36 ( MUSM 11081 ), 37 ( AMNH 268227 , AMNH 268228 ). VENEZUELA : 38 ( EBRG 3305 ), 42 ( AMNH 76933 ), 43 ( MHNLS 3593 ), 44 ( EBRG 17486 ), 47 ( EBRG 2559 ), 48 ( MHNLS 608 ); and photos of the holotype (BM 80.5.6.71).