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Pampatheriidae is a group of South American native cingulates recorded from the Middle Miocene to the Early Holocene. These animals arrived in North America during the Great American Biotic Interchange. During the Quaternary, at least three genera existed: Tonnicinctus Góis, González Ruiz, Scillato-Yané and Soibelzon, Pampatherium Gervais and Ameghino, and Holmesina Simpson. They are differentiated mainly by craniodental and osteodermal characters. In this paper, we describe a new species of Holmesina from Bahia state, Brazil. Two well-preserved specimens possessing osteoderms associated with their skeletons enabled us to determine the genus and species more reliably. Holmesina cryptae sp. nov. differs from the other Quaternary pampatheres by having the simplest ornamental pattern of osteoderms, sharing characteristics between Pampatherium and Holmesina osteoderms and the most robust skull among the Holmesina species, however preserving the main synapomorphies of the genus. Moreover, the integrity of the remains of H. cryptae sp. nov. enabled us to describe elements of the hyoid apparatus, the clavicle, the entire vertebral column and the pelvis. It was also possible to measure its total axial length (2.2 m) more reliably, which is smaller than previously estimated for other pampatheres. Holmesina cryptae sp. nov. constitutes the sixth species of the genus, and it is the fourth in South America.
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by L. Hautier: 27 May 2019; published: 29 Aug. 2019 401
Zootaxa 4661 (3): 401–444
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4661.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:342BAB66-D10B-4E9C-BE36-A2F2B3860A3E
A new and most complete pampathere (Mammalia, Xenarthra, Cingulata)
from the Quaternary of Bahia, Brazil
JORGE FELIPE MOURA1,5, FLÁVIO GÓIS2,
FERNANDO CARLOS GALLIARI3,4 & MARCELO ADORNA FERNANDES1
1Programa de Pós-Graduação em Ecologia e Recursos Naturais, Laboratório de Paleoecologia e Paleoicnologia, Departamento de
Ecologia e Biologia Evolutiva (DEBE), Universidade Federal de São Carlos (UFSCar), Rodovia Washington Luís, KM 235, São Car-
los, 13565-905, SP, Brazil. E-mail: mouradejesus@gmail.com; mafernandes@ufscar.br
2Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción
(CICYTTP–CONICET), Materi y España, 3105 Diamante, Entre Ríos, Argentina. E-mail: flaviogois@gmail.com
3Laboratorio de Morfología Evolutiva y Desarrollo (MORPHOS), Facultad de Ciencias Naturales y Museo, Universidad Nacional de
La Plata, La Plata, Buenos Aires, Argentina. E-mail: fgalliari@fcnym.unlp.edu.ar
4División Paleontología de Vertebrados, Museo de La Plata, Paseo del Bosque S/Nº, La Plata, Buenos Aires, Argentina
5Corresponding author
Table of content
Abstract .................................................................................................401
Introduction ..............................................................................................402
Geological and geographic context ............................................................................403
Material and methods ......................................................................................404
Results .................................................................................................405
Systematic Paleontology ....................................................................................405
Comparative description ....................................................................................406
Armor .................................................................................................406
Skull ................................................................................................. 408
Postcranial skeleton .......................................................................................426
Discussion ...............................................................................................434
Conclusion ..............................................................................................437
Acknowledgements ........................................................................................437
References ...............................................................................................437
Supporting information .....................................................................................440
Abstract
Pampatheriidae is a group of South American native cingulates recorded from the Middle Miocene to the Early Holocene.
These animals arrived in North America during the Great American Biotic Interchange. During the Quaternary, at least
three genera existed: Tonnicinctus Góis, González Ruiz, Scillato-Yané and Soibelzon, Pampatherium Gervais and
Ameghino, and Holmesina Simpson. They are differentiated mainly by craniodental and osteodermal characters. In this
paper, we describe a new species of Holmesina from Bahia state, Brazil. Two well-preserved specimens possessing
osteoderms associated with their skeletons enabled us to determine the genus and species more reliably. Holmesina
cryptae sp. nov. differs from the other Quaternary pampatheres by having the simplest ornamental pattern of osteoderms,
sharing characteristics between Pampatherium and Holmesina osteoderms and the most robust skull among the Holmesina
species, however preserving the main synapomorphies of the genus. Moreover, the integrity of the remains of H. cryptae
sp. nov. enabled us to describe elements of the hyoid apparatus, the clavicle, the entire vertebral column and the pelvis.
It was also possible to measure its total axial length (2.2 m) more reliably, which is smaller than previously estimated
for other pampatheres. Holmesina cryptae sp. nov. constitutes the sixth species of the genus, and it is the fourth in South
America.
Key words: armadillo, osteology, fossil, axial skeleton, cave
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402 · Zootaxa 4661 (3) © 2019 Magnolia Press
Introduction
Xenarthra Cope is one of the major clades of placental mammals comprising 15 genera and 31 species that are wide-
ly distributed in the Americas, however about 200 fossil species were described (McKenna & Bell 1997; Delsuc et
al. 2001; Gaudin & Wible 2006; Möller-Krull et al. 2007). The clade is subdivided into two orders, Cingulata Illiger
(armadillos, pampatheres and glyptodonts) and Pilosa Flower (sloths and anteaters) (Delsuc et al. 2001; Fariña et
al. 2003). The oldest xenarthran recorded is a cingulate from the upper Paleocene (Itaboraian) of Brazil (Scillato-
Yané 1976; Oliveira & Bergqvist 1998; Bergqvist et al. 2004). Until the Early Holocene, Cingulata were much more
diverse than currently observed, and the taxa exhibited a great variety of sizes and habitats (Scillato-Yané 1986;
Vizcaíno 2009; De Esteban-Trivigno 2010; 2011).
Covering their dorsal region, the cingulates have an integumentary armor that consists of hundreds of dermal
bony plates called osteoderms. Externally, the epidermis of the armor consists of epidermal horny scales. The os-
teoderms articulate with one another forming the cephalic shield that covers the head dorsally; the carapace that
covers the trunk dorsolaterally; and the caudal sheath surrounds the tail completely, except in the tail of Cabassous
McMurtrie (naked-tailed armadillos) (Hill 2006; Chen et al. 2011; Krmpotic et al. 2015; Fernicola et al. 2017).
Additionally, isolated osteoderms can be found in the integument of the face, in the ventral region of the body and
in the limbs (Rinderknecht 2000; Tauber & Di Ronco 2000; Soibelzon et al. 2006; Krmpotic et al. 2015). Different
lineages of cingulates show diverse configurations of the carapace, which can comprise fixed osteoderms forming
a buckler; movable (mobile) osteoderms that form belts (bands), or one or two bucklers plus one or more belts (see
Fernicola et al. 2017).
Traditionally, Cingulata included living and extinct forms, which were grouped into two superfamilies: Dasy-
podoidea Gray and Glyptodontoidea Gray (McKenna & Bell 1997). Because pampatheres show morphological
characters similar to dasypodoids and glyptodontoids, their placement in each of these groups has been a matter of
discussion (see Simpson 1930; Hoffstetter 1956, 1958; Robertson 1976; Paula Couto 1979; Cartelle & Bohórquez
1984; Engelmann 1985; Carlini & Scillato-Yané 1993; Abrantes & Bergqvist 2006; Gaudin & Wible 2006; Góis et
al. 2012, 2013; Góis 2013). Recent papers have hypothesized the paraphyly of “Dasypodidae” (when including all
armadillos) and have suggested a new conformation of Cingulata divided into two clades: Chlamyphoridae Bonapar-
te, and Dasypodidae (Gray) (that includes just Dasypodinae) (Gibb et al. 2015; Delsuc et al. 2016; Mitchell et al.
2016). Moreover, molecular phylogenetic studies in glyptodonts have placed them within Chlamyphoridae (Delsuc
et al. 2016; Mitchell et al. 2016). Mitchell et al. 2016 also reanalyzed morphological characters of cingulates includ-
ing the pampathere Vassallia Castellanos and linked the latter to glyptodonts and chlamyphorid armadillos. Thus,
according to these authors, Chlamyphoridae is divided into the subfamilies Glyptodontinae, Pampatheriinae, Eu-
phractinae Winge, Chlamyphorinae and Tolypeutinae Gray (Gaudin & Lyon 2017). Gaudin & Lyon 2017 proposed
the division of Cingulata into two superfamilies, Dasypodoidea and Chlamyphoroidea, the former encompasses the
dasypodines plus their extinct kin, and the latter contains the families Glyptodontidae Gray, Pampatheriidae Paula
Couto, Euphractidae, Chlamyphoridae and Tolypeutidae. These authors argue that previous (mainly traditional)
taxonomic approaches reflect inadequately the family diversity of Cingulata, especially in comparison to Pilosa that
presents younger radiations and shows a more diverse family number.
Pampatheres are herbivorous cingulates (Vizcaíno et al. 1998; De Iuliis et al. 2000) recorded from the middle
Miocene in South America to the early Holocene (Castellanos 1927; Edmund & Theodor 1997; Góis et al. 2012,
2013; Góis 2013). To date, seven genera have been recognized, four of which are recorded from the Neogene:
Scirrotherium Edmund & Theodor, Kraglievichia Castellanos, Vassallia and Plaina Castellanos; three from the
Quaternary: Pampatherium Gervais & Ameghino, Holmesina Simpson, and Tonnicinctus Góis, González-Ruiz,
Scillato-Yané and Soibelzon (Gervais & Ameghino 1880; Castellanos 1927, 1937; Simpson 1930; Scillato-Yané
1982; Edmund 1987; Edmund & Theodor 1997; Scillato-Yané et al. 2005; Góis et al. 2015).
In this paper, two specimens of pampatheres are described. They have well-preserved skeletons and osteoderms,
which enabled us to describe and determine a new species of Holmesina more reliably. In addition, we provide an
osteological description that encompasses most taxa of the family, including first records of hyoid apparatus, com-
plete vertebral column, complete synsacrum and clavicle.
NEW PAMPATHERE FROM THE QUATERNARY OF BAHIA Zootaxa 4661 (3) © 2019 Magnolia Press · 403
Geological and geographic context
Iramaia municipality, Bahia state, Brazil (Fig. 1) has a semiarid climate and a karstic terrain, presenting cave systems
(Vieira et al. 2005). Lapinha Cave (Gruta da Lapinha) (Figs. 1, 2) is located in this municipality (13°19’15.54338’’S
/ 40°58’45.76346’’W), measuring 1,777.47 m in length and it is about 641 m above sea level, and presents bone
accumulation of Quaternary mammals (Moura de Jesus et al. 2015).
Lapinha Cave is formed in carbonate rocks of Gabriel and Nova América undivided Units (Salitre Forma-
tion—Neoproterozoic) (Vieira et al. 2005). In 2012 and 2014, two specimens of Holmesina were found in different
cavities in the second lower level of Lapinha Cave by speleologists from the Grupo Pierre Martin de Espeleologia
(GPME). The skeletons of the specimens were completely articulated, and partially covered by loose sediment of
the cave. The osteoderms were not articulated and scattered around the endoskeletal elements. Additionally, osteo-
derms and bones of juvenile specimens of Glyptodontinae were found at the same site.
FIGURE 1. Map and Location of Lapinha Cave. A–B, location of Iramaia Municipality in Bahia state, Brazil. C, map of Lap-
inha Cave highlighting its second lower level and the points where the fossils were found (adapted map provided by GPME
Speleology Group). Legend: white, ground level of the cave; yellow, first lower level; light blue, second lower level; and dark
green, third lower level of the cave. Abbreviations: G, site where glyptodonts were collected; P1 and P2 (holotype and paratype,
respectively), the sites where these pampatheres were collected.
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FIGURE 2. Exterior and interior of Lapinha Cave. A, External view; in the right below of the picture, notice the cave entrance
closed by a gate. B–C, Specimens of Holmesina cryptae sp. nov. buried in the cave, B, LPP-PV-001; and C, LPP-PV-002 at the
moment of the excavation.
Material and methods
Two specimens of Holmesina sp. nov. were analyzed, LPP-PV-001 and LPP-PV-002. They were compared with
homologous species from the Pliocene and Quaternary: Pampatherium [P. humboldtii (Lund), P. typum (Gervais
& Ameghino), and P. mexicanum Edmund], Holmesina [H. floridana (Robertson), H. septentrionalis (Leidy), H.
occidentalis (Hoffstetter), H. paulacoutoi (Cartelle & Bohórquez), H. rondoniensis Góis, Scillato-Yané, Carlini and
Ubilla], and Tonnicinctus mirus Góis, González-Ruiz, Scillato-Yané and Soibelzon. In addition, other related genera
(Scirrotherium, Vassallia, Kraglievichia and Plaina) were used for comparative purposes (see Appendix 1).
The nomenclature used to describe the osteoderms follows Góis et al. (2013), and the cranial and postcranial char-
acters described by Winge (1915), Castellanos (1927, 1937), Simpson (1930), Bordas (1939), James (1957), Rob-
ertson (1976), Cartelle & Bohórquez (1984), Edmund & Theodor (1997), Gaudin (1999), Góis et al. (2012, 2015),
Góis (2013), Casali & Perini (2017), Gaudin & Lyon (2017) were used.
To measure the osteoderms and bones, a mechanical caliper with an error of 0.5 mm was used. Circumferences
and the total axial size (from the anterior portion of the nasal to the posterior portion of the tail) of LPP-PV-001 were
measured using a tape measure. Moreover, the method proposed by Cartelle & Bohórquez (1984) was followed to
measure the interdental space—the toothrow length minus the sum of the maximum lengths of all molariforms.
The skulls of the specimens LPP-PV-001 and LPP-PV-002 were scanned using a General Electric (GE) Phoe-
nix v|tome|x M Computed Tomography (CT) scanner at the Museum of Zoology from the University of São Paulo,
Brazil (MZUSP), and images were viewed using MyVGL 3.1 software, primarily to analyze some characteristics in
upper molariforms and hard palate.
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The specimens analyzed in this study are housed in the following institutions:
LPP Laboratório de Paleoecologia e Paleoicnologia, Departamento de Ecologia e Biologia Evolutiva, Uni-
versidade Federal de São Carlos, Brazil
ROM Royal Ontario Museum, Toronto, Canada
MCL Museu de Ciências Biológicas da Pontifícia Universidade Católica de Minas Gerais, Belo Horizonte,
Brazil
UF Florida Museum of Natural History, Gainesville, Florida, USA
MUFyCA Museo Universitario Florentino y Carlos Ameghino, Rosario, Santa Fe, Argentina
INAH Instituto Nacional de Antropología e Historia, Ciudad de México, Distrito Federal, Mexico
AMNH American Museum of Natural History, New York, USA
HMNS Houston Museum of Natural Science, Houston, Texas, USA
UZM Universitets Zoologisk Museum, Copenhagen, Denmark
MACN Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina
MHD Museo Histórico Departamental de Artigas, Artigas, Uruguay
MLP División Paleontología Vertebrados, Facultad de Ciencias Naturales y Museo, Universidad Nacional
de La Plata, La Plata, Argentina
Results
Systematic Paleontology
Superorder Xenarthra Cope, 1889
Order Cingulata Illiger, 1811
Superfamily Chlamyphoroidea (Bonaparte, 1850)
Family Pampatheriidae Paula Couto, 1954
Genus Holmesina Simpson, 1930
Type species. H. septentrionalis (Leidy) Simpson
Holmesina cryptae Moura, Góis, Galliari & Fernandes sp. nov.
Figs. 2–23
Type locality. Two specimens (holotype and paratypes) associated to three juvenile individuals of Glyptodontinae
were found in the inferior level of the cave Gruta da Lapinha in Iramaia Municipality, Bahia state, Brazil.
Etimology. Gruta” (cave/grotto in Portuguese) comes from the Latin “crypta” (crypt). At the entrance of Lap-
inha Cave, there is an altar that is used by Catholic believers, and the skeletons of the pampatheres were found in a
chamber below which is analogous to a crypt, reinforcing the choice of this name.
Holotype. LPP-PV-001: Skull. Maxillary teeth (15 – right and left: Mf3, Mf4, Mf5, Mf6, Mf7, Mf8 and Mf9;
and left: Mf2). Mandibles (fragmented). Hyoid bones (right stilohyal and epihyal). Cervical vertebrae (C1–C7).
Thoracic vertebrae (T1–T10). Lumbar vertebra (L1). Synsacral vertebrae (S1–S11). Caudal vertebrae (Cd1–Cd22).
Chevron bones (5). Scapulae (incomplete). Clavicles (right and left). Manubrium. Ribs (10 vertebral portion).
Humeri (right and left). Radii (right and left). Ulnae (right and left). Right trapezoid bone. Hamate bones (right and
left), Triquetral bones (right and left). Lunate bones (right and left). Pisiform bones (right and left). Anterior distal
phalanges (right and left: I, II, III, IV, V). Anterior medial phalanges (II, III and IV). Anterior proximal phalanges
(right: I, II, III, IV and V; left: I, II, III, IV and V). Metacarpi (right and left: I, II, III and IV). Sesamoid bones in right
MOURA ET AL.
406 · Zootaxa 4661 (3) © 2019 Magnolia Press
manus (3), and right and left palmar bones. Pelvis. Femora (right and left). Patellae (right and left). Tibiofibulas
(right one is complete; distal and proximal epiphyses of the left one). Calcanei (right and left). Astragali (right and
left). Naviculars (right and left). Cuboids (right and left). Metatarsi (right and left: II, III, IV and V). Posterior distal
phalanges (right: I, II, IV and V; left: II, III, IV and V). Posterior medial phalanges (right: II, IV and V; left: II, III, IV
and V). Posterior proximal phalanges (right: III; left: I, II, III, IV and V). Right and left plantar bones. Osteoderms
(more than 1000 and fragments).
Paratype. LPP-PV-002: Skull almost complete. Maxilary teeth (right and left M1–M9). Mandible (complete
in right side with nine teeth; the left one is fragmented). Hyoid bones (basithyrohyal). Cervical vertebrae (C1–C7).
Thoracic vertebrae (T1–T10). Lumbar vertebra (L1). Synsacral vertebrae (S1–S11). Caudal vertebrae (Cd1–Cd4).
Chevron bones (4). Manubrium. Ribs (fragments). Incomplete humeri. Radii (right and left). Ulnae (right and left).
Pelvis. Femora (distal and proximal epiphyses). Patellae (right and left). Tibiofibulas (fragments). Calcaneus (left).
Astragalus (left). Cuboids. Anterior distal phalanges (right and left I, II, III, IV and V). Osteoderms (more than 200
and fragments).
Age: SALMA Lujanian (Late Pleistocene).
Diagnosis. Holmesina cryptae sp. nov. differs from other species of pampatheres in having: a less elevated
frontal crests in frontal bones; a less robust, thin and low sagittal crest; a nuchal crest narrower in the lateral borders;
a robust and pointed mastoid process in the petrosal bones; a C-shaped notch anteriorly in lateral view of premaxil-
las; a larger infraorbital foramen in maxillas; very robust postorbital apophyses (postorbital process of jugal). The
shape of the H. cryptae sp. nov. skull is more triangular than the other species and possesses robust zygomatic
arches with an orthogonal process zygomatic of squamosal. The skull also shows the most robust parietal region,
and the widest and more robust snout. The molariforms are farthest from the sagittal line than any of the compared
species. Holmesina cryptae sp. nov. presents osteodermal ornamentations less marked than in Scirrotherium cari-
natum, Kraglievichia, H. paulacoutoi and H. occidentalis, however more defined and punctured than in Pampathe-
rium. Anteriorly to the first lower molariform (mf1) in the dentary, there is a toothless pointed region longer than
in any other species of Holmesina, however it is less developed than in P. typum and P. humboldtii. The first three
anterior upper molariforms (Mf1–Mf3) show an oval morphology as in the rest of the species. The morphology
of the other molariforms is more similar to other Holmesina species (short and wide, eight-shaped), with reduced
interdental spaces, in contrast to Pampatherium. The mandible is much less robust than in P. humboldtii, horizontal
and ascending rami narrower than in P. humboldtii. In the ulna, the lateral and medial fossae are superficial and the
styloid apophysis is rounded. The femur shows a deeper and more delimited trochanteric fossa than other species of
Holmesina. The patella has a longer and straighter apex in comparison to H. floridana.
FIGURE 3. Complete skeleton of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 300 mm.
Comparative description
Armor
Morphological variation among osteoderms of H. cryptae sp. nov. can be seen in Fig. 4 and Appendix 2.
Cephalic shield osteoderms. Their surfaces have diverse shapes. Dorsoventrally, they are concavo-convex and
thinner than the osteoderms of the carapace and tail (Fig. 4A).
Fixed osteoderms. Their lateral margins are narrower than in S. carinatum MLP 69-IX-8-13AC, Kraglievichia,
H. occidentalis ROM 28392, H. paulacoutoi MCL 500/110-126, H. floridana UF 224397, and T. mirus, similar to P.
humboldtii, but more evident than in P. typum (Fig. 4B and 4C). The marginal elevation is lower and less bounded
NEW PAMPATHERE FROM THE QUATERNARY OF BAHIA Zootaxa 4661 (3) © 2019 Magnolia Press · 407
than in Scirrotherium, Kraglievichia or in any other species of Holmesina. In this regard, the osteoderms of H.
cryptae sp. nov. resemble those of Pampatherium. The longitudinal depressions are less deep than in S. carinatum,
Kraglievichia, H. septentrionalis, H. paulacoutoi and H. occidentalis, but deeper than in V. minuta, V. maxima MU-
FyCA 500 and P. typum. The central longitudinal elevation is lower and less sculpted than in S. carinatum, Kragliev-
ichia, H. septentrionalis, H. paulacoutoi and H. occidentalis, and more defined than in P. typum, P. humboldtii and
P. mexicanum, INAH 6201. Nevertheless, its ornamentation is completely distinct to that of T. mirus. Comparative
measurements on fixed osteoderms of pampathere species are given in Table 1 and 2.
TABLE 1. Measurements of greatest scapular fixed osteoderms of pampatheres.
Species Length (mm) Width (mm) Area (mm)
Scirrotherium hondaense 19.75 19.5 385.12
Scirrotherium carinatum 30 26.1 783
Scirrotherium antelucanum 26.9 20.8 559.52
Vassallia minuta 31 23.75 736.25
Vassallia maxima 44 34 1496
Kraglievichia 46 37 1702
Tonnicinctus mirus 39.8 30.7 1221.86
Holmesina septentrionalis 49.6 41 2033.6
Holmesina cryptae sp. nov. 38.8 37.6 1462.2
Pampatherium mexicanum 40 32 1280
Pampatherium humboldtii 45 31 1395
TABLE 2. Measurements of greatest pelvic fixed osteoderms of pampatheres.
Species Length (mm) Width (mm) Area (mm)
Scirrotherium hondaense 22.8 18.66 425.44
Scirrotherium carinatum 36.16 27.71 1001.99
Vassallia minuta 34.8 31.9 1110.12
Tonnicinctus mirus 48.5 34.2 1316.7
Holmesina cryptae sp. nov. 50.2 34.4 1734.1
Holmesina paulacoutoi 63 51.5 3244.5
Pampatherium typum 33.4 30.3 1012.02
Semimovable osteoderms of scapular buckler. Unlike the semimovable osteoderms of the pelvic buckler, the
anterior articular surface is absent in these osteoderms and they have a fixed articulation (Fig. 4F). Some character-
istics such as lateral margins, marginal elevation, longitudinal depression and longitudinal central elevation are as
well marked as in other species of Holmesina, in contrast to those of Pampatherium species.
Movable osteoderms and semimovable osteoderms of pelvic buckler. The movable bands comprise the
middle region of the carapace and consist of successive imbricated osteoderms. In living and extinct cingulates,
the number of movable bands can vary greatly. Nevertheless, pampatheres in which this region is preserved—V.
maxima, H. septentrionalis AMNH 23435, P. humboldtii, and H. cryptae sp. nov.—exhibit three movable bands
(Edmund 1985; Góis et al. 2013). In H. cryptae sp. nov., these osteoderms present a rectangular shape and the
largest ones measure about 110 mm in length (smaller than in H. paulacoutoi) (Fig. 4D and Table 3). Some mov-
able osteoderms show ornamental structures that are more evident than in the fixed osteoderms. Their intermediate
portion is as deep as in other species of Holmesina, in contrast to Pampatherium. Nevertheless, the length of this
portion measures more than the half of the length of the anterior articular surface as in Pampatherium, and different
from other species of Holmesina.
In general, the anterior foramina are small and shallow. The lateral margins are wider than in Scirrotherium,
Kraglievichia, V. minuta and P. typum, but much less marked than in T. mirus and other species of Holmesina. The
marginal elevation is less pronounced than in Scirrotherium, Kraglievichia, Tonnicintus or any other species of Hol-
mesina, but it is more marked than in Vassallia and Pampatherium. The longitudinal depression is deeper than in
Pampatherium, but less deep than in Kraglievichia, Scirrotherium and the other species of Holmesina. The central
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longitudinal elevation is low, long and narrow; it is more detailed than in the species of Pampatherium, but less
marked than in Scirrotherium, Kraglievichia and the other species of Holmesina. The semimovable osteoderms of
the pelvic buckler are very similar to the movable ones (Fig. 4E and Table 4).
TABLE 3. Measurements of greatest movable osteoderms of pampatheres.
Species Length (mm) Width (mm) Area (mm)
Scirrotherium hondaense 60 25 1500
Scirrotherium carinatum 54.5 25 1362
Vassallia minuta 66 28 1848
Vassallia maxima 53 32.5 1722
Kraglievichia paranensis 60.5 26.5 1603
Plaina brocherensis 58 39 2262
Holmesina cryptae sp. nov. 115 37 3108
Holmesina occidentalis 87 35 3045
Holmesina paulacoutoi 127.37 52.19 9065
Pampatherium typum 73 34 2482
TABLE 4. Measurements of greatest pelvic semimovable osteoderms of pampatheres.
Species Length (mm) Width (mm) Area (mm)
Scirrotherium carinatum 42.21 24.31 1026
Vassallia minuta 59.9 24.9 1491
Kraglievichia paranensis 58.25 31.5 1834
Tonnicinctus mirus 76.25 32.7 2493
Holmesina cryptae sp. nov. 68.3 23 1570
Pampatherium typum 44 27 1188
Skull
The skulls of the Pampatheriidae are long due to the prolongation of the rostrum. Comparatively, the pampatheres
resemble more the euphractines (e.g., Euphractus Wagler) among the armadillos, and the Propalaehoplophorinae
Ameghino among the glyptodonts (Winge 1915; Bordas 1939; Vizcaíno et al. 1998).
Frontal. In H. cryptae sp. nov., it is a convex bone as in H. occidentalis, H. floridana, V. maxima and H. oc-
cidentalis, and is more convex than in Kraglievichia, H. rondoniensis, P. typum and P. humboldtii. Anteriorly, it
expands throughout the dorsal view, arranging the lacrimal bones more laterally (Figs. 5A, 6A). The interfrontal
suture is completely fused in both specimens (LPP-PV-001 and LPP-PV-002).
In its posterior portion, the frontal bone presents the temporal lines that form two slightly elevated crests con-
verging posteriorly to the sagittal crest in the parietal bone (Figs. 5A, 6A). These frontal crests are much less marked
than in V. maxima, H. occidentalis, H. rondoniensis and probably in H. paulacoutoi that shows a robust sagittal
crest, but it is damaged along its extension. In H. floridana, these crests are very robust in which these lines form
a ridge.
Dorsally, LPP-PV-001 presents various foramina around the midline of the frontal bone, as it was observed in
H. floridana (UF 191448) by Gaudin & Lyon (2017). In lateral view (Figs. 7A, 8A), the posterolateral region of the
frontal bone shows few small foramina. Anteriorly there are some foramina, including the foramen for the frontal
diploic vein, which is also reported in H. floridana, H. septentrionalis and Vassallia.
Parietal. In H. cryptae sp. nov., the parietal region is robust and rough with several vascular channels and
foramina for rami temporales (from side to side) (Figs. 5A, 6A). This region is longer in H. floridana and Kragliev-
ichia than in late Quartenary species. Longitudinally, the sagittal crest is thin and low, similarly to Kraglievichia,
but much less robust than in H. floridana, V. maxima, H. septentrionalis, H. occidentalis, H. paulacoutoi, P. typum
and P. humboldtii. The nuchal (lambdoid) crest is thick in its medial region, and thinner in the lateral borders. In P.
humboldtii and P. typum, this crest is thicker and rougher, and possesses an anterior convexity that is more marked
than in the species of Holmesina, which implies a stronger muscular insertion. Accordingly, this is a distinctive
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character between the genera Pampatherium and Holmesina (Vizcaíno et al. 1998; De Iuliis et al. 2000; De Iuliis
& Edmund 2002).
Squamosal. The posterodorsal surface shows a variable number of foramina for the rami temporales, as in other
cingulates (Figs. 7A, 8A). Likewise, the suprameatal foramen is present in the base of the zygomatic process of the
squamosal (Figs. 8A, 9). The posteriormost edge that articulates with the occiput forms the lateral parts of the nuchal
crest. In addition, a horizontal crest connects the dorsal edge of the zygomatic process to the nuchal crest, as in other
pampatheres (Gaudin & Lyon 2017). Below the zygomatic process of the squamosal, the postglenoid foramina are
in the glenoid fossa (Fig. 9). The tympanic cavities are small and globular with a semioval base.
FIGURE 4. Osteoderms of Holmesina cryptae sp. nov. A–D, LPP-PV-001, A, osteoderm of cephalic shield. B, fixed osteoderm
of scapular buckler in dorsal and lateral views. C, fixed osteoderm of pelvic buckler. D, movable osteoderm in dorsal and right
lateral views. E, (LPP-PV-002) semimovable osteoderm of pelvic buckler, F, (LPP-PV-001) semimovable osteoderm of scapular
buckler. G, (LPP-PV-002) caudal osteoderm (proximal portion of the tail). H–I, LPP-PV-001, H, caudal osteoderm (median por-
tion), and I, caudal osteoderm (distal portion). Scale bar = 20 mm.
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FIGURE 5. Skull of Holmesina cryptae sp. nov. (holotype, LPP-PV-001), dorsal and ventral views. A, dorsal; and B, ventral
views. Scale bar = 50 mm.
Petrosal. In lateral view, the petrosal bone houses the inner ear and is located posteroventrally to the squamo-
sal bone (Figs. 7A, 8A, 9). In both specimens of H. cryptae sp. nov., it is not fused to the surrounding bones. The
mastoid process is more rosbust and pointed in H. cryptae sp. nov. than in other species of Holmesina. According to
Gaudin & Lyon (2017), this process forms the posterior wall of the porus acusticus. Anteroventrally to the petrosal,
the promontorium is a globose structure, as in H. floridana.
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FIGURE 6. Skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), dorsal and ventral views. A, dorsal; and B, ventral
views. Scale bar = 50 mm.
Sphenoid. In lateral view, the alisphenoid region possesses anteriorly an opening and a fissure with foramina
(Figs. 8A, 9). The ventralmost opening houses the sphenopalatine foramen and the caudal palatine foramen and
continues with the sphenorbital fissure that fuses with the foramen rotundum; this fusion is characteristic in Holm-
esina (Gaudin & Lyon 2017). The optic foramen is observed above the sphenorbital fissure. Posteriorly, behind the
entoglenoid process there is a fossa with a large foramen ovale and a small transverse canal foramen, these foramina
were also described for H. floridana. The basisphenoid and basioccipital are fused in both specimens of H. cryptae
sp. nov., and LPP-PV-002 possesses a crack in the region between the basisphenoid and presphenoid.
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Pterygoid. Posteriorly to the Mf9, the pterygoid apophyses are observed; in H. cryptae sp. nov., they are oval
and very rough, extending out ventrolaterally from the tooth row (Figs. 5B, 6B) as it occurs in H. occidentalis and
H. paulacoutoi; in other species such as H. rondoniensis and H. floridana, these structures are aligned with the
longitudinal line of the molariforms.
In LPP-PV-001, the distinction between pterygoid and palatine is clear, whereas in LPP-PV-002 the palatine
surface forms a continuum with the surface of the pterygoid process. In H. paulacoutoi, the pterygoid forms a trian-
gular pointed “wing” posteriorly, but in H. cryptae sp. nov. it is more rounded.
Occipital. The basioccipital is longer than in H. rondoniensis, H. occidentalis, P. typum, P. humboldtii, similar
to H. paulacoutoi and shorter than in V. maxima and Kraglievichia. In this region (Figs. 5B, 6B), there are two fossae
separated by a crest wherein the rectus capitis ventralis muscle is inserted. These fossae are long and deep, occupy-
ing almost the entire occipital region. This characteristic is absent in the other species.
The occipital condyles are parallelogram-shaped (with round corners) (Figs. 5B, 6B, 7B, 8B, 9), as in all species
of pampatheres. The glenoid (mandibular) fossa is situated dorsally to the condyles, it is deeper in Kraglievichia, H.
floridana and H. rondoniensis, and shallower in V. maxima. In H. cryptae sp. nov., this fossa is long and flat, similar
to P. typum, while in H. rondoniensis and H. occidentalis it is arched. The foramen magnum is suboval in shape, as
in H. floridana, V. maxima, H. rondoniensis, H. occidentalis, H. paulacoutoi, while in P. typum and P. humboldtii it
is subcircular.
In occipital view, the occipital region is semicircular, wider than high, robust, very rough and deep (concave)
(Figs. 7B, 8B); these characteristics are similar to P. humboldtii and P. typum. The lateral borders of the occipital
region are highly robust, thick and rough, due to the lambdoid crest projection. Contrarily, in H. occidentalis, H.
rondoniensis and H. paulacoutoi, the occipital region is high and shows a median occipital crest well defined sepa-
rating two longitudinal depressions. In H. paulacoutoi, this region is rougher and more robust than in H. occidentalis
and H. rondoniensis.
FIGURE 7. Skull of Holmesina cryptae sp. nov. (holotype, LPP-PV-001), lateral and posterior views. A, lateral; and B, and
occipital views. Scale bar = 50 mm.
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Nasal. Long bones measuring about 45% of the greatest skull length (Figs. 5A, 6A, Table 5) in both specimens
of H. cryptae sp. nov. (LPP-PV-001 and LPP-PV-002). According to Gaudin & Lyon (2017), this proportion is a
feature of most pampatheres. Nasal bones occupy centrally the rostrum of H. cryptae sp. nov. in dorsal view; the
maxillary bones are also visible in the approximate proportion which makes the snout of the new species relatively
wider and more robust than other Holmesina species (Figs. 5A, 7A). The nasofrontal suture is V-shaped as in other
Holmesina species.
Anteriorly, the nasal bone extends to the premaxilla. H. cryptae sp. nov. (LPP-PV-001, left nasal bone) shows
the distal portion of the nasal slightly wider than the proximal (in nasofrontal suture), as in other Holmesina spe-
cies (Gaudin & Lyon 2017). Nevertheless, the difference is less marked than in P. typum MACN PV 11543 and P.
humboldtii MHD-P-28.
Premaxilla. The premaxillary bones are damaged in LPP-PV-001 in lateral and ventral views (Figs. 5B, 7A),
but they are more preserved in LPP-PV-002 (Figs. 6B, 8A). The premaxilla-maxillary sutures are completely fused
(Figs. 7A, 8A). Anteriorly in lateral view of the premaxilla, both specimens of H. cryptae sp. nov. show a deep and
tall C-shaped notch. This notch was also observed in H. floridana, H. septentrionalis and Vassallia, but with differ-
ent morphologies (see Gaudin & Lyon 2017).
In dorsal view, anteriorly to the Mf1, there is an edentulous prolongation, more pronounced in P. typum, P. hum-
boldtii and H. floridana. It is shorter in H. cryptae sp. nov. (Fig. 6B) and V. maxima and is absent in H. rondoniensis
and Kraglievichia.
Maxilla. In lateral view, the anterior region of the rostrum shows a maxillary crest more or less marked accord-
ing to the species (Figs. 7A, 8A). In H. cryptae sp. nov., this crest is more robust than in Kraglievichia, V. maxima,
H. floridana and H. rondoniensis, and it is less marked than in H. paulacoutoi, P. typum and P. humboldtii. The latter
two species show a very high maxillary crest.
The antorbital fossa is above the median region of the Mf5 and the infraorbital foramen opens 7.5 mm above
the second third of the Mf6 in LPP-PV-002, and the maxillary foramen is posterior to the maxillary process of the
zygomatic arch. In H. cryptae sp. nov., this foramen is larger than in H. occidentalis, H. paulacoutoi, H. rondoni-
ensis, P. typum and P. humboldtii. In Quaternary pampatheriid species, this foramen is usually almost hidden due
to the robustness of the skull. In H. paulacoutoi, it is completely visible because the maxillary crest is higher. In H.
floridana, Kraglievichia and V. maxima, the foramen opens more anteriorly than in Quaternary species.
In ventral view, the anterior portion of the palate is narrower in Kraglievichia and Holmesina; where the ante-
rior molariforms are imbricated (Góis et al. 2012; Góis 2013). In V. maxima and Pampatherium, although the pal-
ate is anteriorly narrow, the molariforms are not imbricated. The anterior imbrication in H. cryptae sp. nov. occurs
between the Mf1, Mf2 and Mf3 (Figs. 5B, 6B, 10); in H. rondoniensis, between Mf2, Mf3 and Mf4; in H. occiden-
talis, between Mf2 and Mf3; in H. paulacoutoi, between Mf3, Mf4 and Mf5, and in H. septentrionalis HMNS 173
between Mf2 and Mf3. Posteriorly to the Mf5, the palate of H. cryptae sp. nov. is wider than in Kraglievichia, H.
floridana, H. occidentalis and H. rondoniensis; similar to H. paulacoutoi and narrower than in Pampatherium.
The palate presents various foramina as in other xenarthrans (Gaudin & Lyon 2017). In H. cryptae sp. nov.,
the number of foramina is higher in specimen LPP-PV-001 than in specimen LPP-PV-002. The anterior palatine fo-
ramina of H. cryptae sp. nov. are situated around the Mf2 region, whereas in H. septentrionalis and H. paulacoutoi
they are around the Mf3, and in H. occidentalis, H. floridana and Vassallia, these foramina are more posteriorly.
These anterior foramina locate in the palatine grooves, which are discontinuous and extend to the median portion
of the Mf7 in LPP-PV-001, whereas in LPP-PV-002 they are continuous until the Mf5 and become discontinuous
until the Mf7.
Palatine. The palate of H. cryptae sp. nov. presents anteriorly a concavity from the Mf4 to the middle of the
Mf6. From the distal portion of the Mf6 to the first portion of the Mf8 it becomes convex then it finishes concave
again throughout the extension of the palatine (Figs. 5B, 6B).
The palatine is flat in the median region in specimen LPP-PV-002, whereas in LPP-PV-001 it is slightly con-
cave. Both present a median palatine ridge where the median suture fuses, as observed in other Holmesina. The
surface presents a varied number of major and minor palatine foramina, as occurs in other species of xenarthrans
(Gaudin & Lyon 2017). The posterior edge of the palatine is U-shaped, as in other pampatheres. In LPP-PV-001, the
median palatine ridge forms a small beak beyond the posterior edge. Both specimens of H. cryptae sp. nov. present a
transverse crack in the surface of the palatine, as observed by Gaudin & Lyon 2017 in some H. floridana specimens
and have assumed that it is not a suture.
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Zygomatic arch. Anteriorly, the zygomatic arch of H. cryptae sp. nov. is formed by the infraorbital process (zy-
gomatic process of the maxilla) (Figs. 5, 6, 7A), which is an apophysis longer, rougher and more robust than in H.
occidentalis, P. typum and P. humboldtii UZM 2314. This structure is absent in H. rondoniensis (Góis et al. 2012).
Other species, as V. maxima, H. floridana and H. paulacoutoi also show a long, rough and robust infraorbital pro-
cess. Holmesina cryptae sp. nov. shows a very robust postorbital apophysis (= postorbital process of jugal in Gaudin
& Lyon 2017), more robust than any other species. Similarly, H. occidentalis shows this structure well developed.
Pampatheriidae present a robust zygomatic arch. The joint between jugal and zygomatic process of squamosal
in H. cryptae sp. nov. almost reaches the orbital apophysis (= postorbital process of frontal in Gaudin & Lyon 2017).
Holmesina floridana and P. typum also show a robust zygomatic arch, although less marked than H. cryptae sp. nov.
In ventral view, the zygomatic process of the squamosal expands laterally in an orthogonal angle (Fig. 5B) which
makes the H. cryptae sp. nov. skull apparently more triangular than other species.
Lacrimal. This is a parallelogram-shaped bone as in other pampatheres (Figs. 7A, 8A). The lacrimal foramen
is open in the antorbital ridge as in H. floridana, H. paulacoutoi, but differently from H. septentrionalis (UF 889,
243224) and Vassallia (P 14424), which possess the lacrimal foramen anteriorly to the antorbital ridge (Gaudin &
Lyon 2017). Both specimens of H. cryptae sp. nov. have a lacrimal fenestra larger in diameter than the lacrimal
foramen, and located where the lacrimal, frontal and maxillar bones converge. Near the fenestra, a smaller foramen,
the lacrimosinusal foramen, is observed in H. paulacoutoi (Cartelle & Bohórquez 1984).
FIGURE 8. Skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002) , lateral and posterior views. A, lateral; and B, and
occipital views. Scale bar = 50 mm.
Upper molariforms. The anteriormost region of the tooth row is damaged in the holotype (LPP-PV-001), but in
the paratype (LPP-PV-002), the first three molariforms are well preserved, externally they are less imbricated than
other species of Holmesina, but the imbrication is more evident when analyzing CT scanned images (Fig. 10).
Externally, many molariforms (Mf – upper molariform) of the specimens are fragmented or missing; as they are
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euhypsodont teeth, the morphology and size of the crown are inferred according to the shape of the alveoli (Figs.
5B, 6B, 11 and Table 6). CT scanned images were obtained for a better visualization and qualitative description of
H. cryptae sp. nov. molariforms.
In Pampatheriidae, the morphology of the molariforms (upper and lower) shows two well-defined patterns: in
S. hondaensis, Kraglievichia and species of Holmesina, molariforms are short, robust and bilobed (sometimes there
is a third medial pseudolobe), while in V. maxima and Pampatherium they are long, narrow and trilobed (Góis et al.
2012; Góis 2013). Another remarkable difference is the interdental space, i.e., in S. hondaensis, Kraglievichia and
Holmesina the molariforms are closer to each other, while in V. maxima and Pampatherium, they are more separated
(Fig. 11).
FIGURE 9. Ventrolateral view of the skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002) focusing the left auditory
region. aptt, anteroventral process of tegmen tympany; as, alisphenoid; bb, bony bridge between tympanohyal and crista in-
terfenestralis; bcc, basicochlear commissure; bo, basioccipital; bs, basisphenoid; cf, carotid foramen; cp, crista parotica; eam,
external auditory meatus; eo, exoccipital; f, frontal; fo, foramen ovale; fr/sof, fused foramen rotundum and sphenorbital fissure;
fm, foramen magnum; fs, facial sulcus; gf, glenoid fossa; hf, hypoglossal foramen; itc, infratemporal crest; Mf9, ninth upper
molariform; mp, mastoid process (pop, paroccipital process of petrosal of Gaudin, Lyon (2017)); mx, maxilla; nc, nuchal crest;
occ, occipital foramen; of, optic foramen; pcp, paracondylar process of exoccipital (= paroccipital process); pe, petrosal; pf,
piriform fenestra; pgf, postglenoid foramen; pgp, postglenoid process; pr, promontorium of petrosal; pt, pterygoid; pt/pl, fused
pterygoid and palatine; spf/cpf, fused sphenopalatine and caudal palatine foramina; smf, suprameatal foramen; stf, stylomastoid
foramen; sq, squamosal; th, tympanohyal; ttf, tensor tympani fossa on epitympanic wing of petrosal; zp, zygomatic process of
squamosal. Perspective figure, dimensions must be compared to Figs. 6 and 8. Based on Gaudin & Lyon (2017).
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FIGURE 10. Ventral view of frontal plane of skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), computed tomo-
graphic scan image, –15 mm from the palate surface. Abbreviations: ju, jugal bone; ma, maxilla; Mf1, first molariform; Mf2,
second molariform; nc, nasal cavity; pm, premaxilla; pt, pterygoid; zpm, zygomatic process of maxilla. Scale bar = 35 mm
Another character that differentiates the pampatheres is the arrangement of the osteodentine. In S. hondaensis
and Holmesina, the osteodentine is scarce or absent in some molariforms. In V. maxima and Pampatherium, the mo-
lariforms have two layers of osteodentine (central and external) with a certain relief (Vizcaíno et al. 1998; De Iuliis
& Edmund 2002; Góis et al. 2012; Góis 2013).
Mf1. Subcircular or circular, it is the smallest of all the molariforms in the tooth row and inserts in the premax-
illa (Fig. 12). These characters are common to all pampatheres (Catellanos 1927, 1937; De Iuliis & Edmund 2002;
Vizcaíno 2009; Góis et al. 2012; Góis 2013; Gaudin & Lyon 2017). In H. cryptae sp. nov., the Mf1 is subcircular
with a slight lingual inclination.
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FIGURE 11. Schematic representation of the right upper tooth rows of South American Quaternary pampatheres. A, Holmesina
cryptae sp. nov. (paratype, LPP-PV-002). B, H. rondoniensis (holotype, MERO-P-002). C, H. occidentalis (ROM 3881). D, H.
paulacoutoi (holotype, MCL-501/01). E, H. septentrionalis (HMNS 173). F, Pampatherium typum (MACN Pv 11543). G, P.
humboldtii (MHD-P-28). In black, absent molariforms (shape inferred from the alveolus); in some cases, the occlusal surface
is totally or partially destroyed, thus some illustrations were completed from the left tooth row of the same exemplar. Scale bar
= 30 mm.
Castellanos (1927, 1937) states that the first two molariforms of Pampatherium are inserted in the maxilla,
however this observation may be incorrect. As previously mentioned, the CT scan images helped us to verify in
all the specimens analyzed that only the Mf1 inserts in the premaxilla. The same was recorded by Gaudin & Lyon
(2017), who analyzed specimens where the premaxillary-maxillary suture is evident.
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Mf2. Usually larger than the Mf1, with a small distal prolongation in all species, except in P. typum and P. hum-
boldtii. Holmesina cryptae sp. nov. shows a moderate distal prolongation, which is perpendicular to the dental line,
and with a small groove (lingual) resulting in a reniform shape. It is subcircular in S. hondaensis, Kraglievichia, V.
maxima and H. floridana.
Mf3. Obliquely directed to the tooth row with the median groove absent in H. cryptae sp. nov. In the other spe-
cies, this molariform is oriented lingually (e.g., Kraglievichia, H. rondoniensis, H. occidentalis, H. floridana).
Mf4. S-shaped and with slightly defined lobes; the anterior portion is oriented lingually; and the posterior por-
tion labially in H. cryptae sp. nov. The groove is more marked than in H. rondoniensis, H. occidentalis, H. paula-
coutoi and H. septentrionalis, and these species have simpler molariforms than H. cryptae sp. nov. This molariform
is trilobed in P. typum and P. humboldtii. Between Mf5 and Mf9, the molariforms usually have more bilobed or
trilobed morphologies.
FIGURE 12. Sagittal planes of skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), computed tomographic scan im-
ages. A, 1 mm; B, 2 mm; and C, 4 mm from the median sagittal plane. Abbreviations: ec, endocranial cavity; ma, maxilla; Mf1,
first molariform; Mf2, second molariform; nc, nasal cavity; pm, premaxilla. Scale bars = 35 mm.
Mf5. Bilobed, with lobes and grooves well-defined (the labial groove is more pronounced); the anterior lobe
faces lingually while the posterior faces labially in H. cryptae sp. nov. Comparatively, this molariform is more
bilobed than in H. occidentalis but less than in H. septentrionalis. In H. occidentalis, the medial groove is less
pronounced. In H. rondoniensis and H. floridana, the shape is more reniform than bilobed. In H. paulacoutoi, it is
robust as in H. cryptae sp. nov., showing a third lobe (medial). In Kraglievichia, it has the simplest morphology
among the Pampatheriidae. In P. typum and P. humboldtii, it is clearly trilobed, the latter species presents the three
most defined lobes.
Mf6. More robust and nearly trilobed in H. cryptae sp. nov. Nevertheless, this molariform is narrower and com-
pletely different from Kraglievichia and any other species of Holmesina. It differs from P. typum and P. humboldtii
because these species possess a narrower and more marked middle lobe.
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Mf7. Showing three well-defined lobes. It is more robust in H. cryptae sp. nov. than in P. typum and P. humbold-
tii. Holmesina floridana and Kraglievichia possess a rudimentary third lobe, and it is bilobed in H. occidentalis, H.
rondoniensis and H. septentrionalis.
FIGURE 13. Right hemimandible of Holmesina cryptae sp. nov. (paratype, LPP-PV-002). A, lateral; B, medial; and C, occlusal
views. Scale bar = 50 mm.
Mf8. Similar to the Mf7, with the anterior and posterior lobes rounder in H. cryptae sp. nov. In Kraglievichia,
it is bilobed, longer and narrower than the Mf7. In H. floridana it is pseudotrilobed. In H. rondoniensis, H. occi-
dentalis and H. paulacoutoi, they are completely bilobed. In P. typum and P. humboldtii, they are trilobed and much
narrower than in H. cryptae sp. nov.
Mf9. Differently from the anterior molariforms (Mf6-Mf8); it is reniform in H. cryptae sp. nov. It is also larger
in comparison to the other species and possesses a well-marked median groove (labial) and both lobes labially di-
rected in H. cryptae sp. nov.
Mandible. The jaws of pampatheres are isognathous (Vizcaíno et al. 1998). In lateral view, the horizontal ra-
mus is long; its height varies depending on the species (Fig. 13 and Table 7). The maximum height measurement
falls about the seventh and ninth lower molariform (mf7–mf9). In H. cryptae sp. nov., the horizontal ramus is more
robust, and proportionally higher than S. hondaensis, V. minuta, V. maxima, Kraglievichia and H. floridana and H.
occidentalis. It is less robust than in H. septentrionalis, and much less robust than in P. typum and P. humboldtii.
Moreover, the horizontal ramus of H. cryptae sp. nov. decreases considerably from the mf4 to the anterior portion.
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TABLE 5. Skull measurements of pampatheres.
Species Specimen Measurements (mm)
Total length Nasal length Frontal length Parietal length Palatal length MDFM WB
Kraglievichia MACN Pv 2617 194 59 62 73 159 – 69
Holmesina floridana UF 121742 223 75 73 78 121 26 58
Vassallia maxima FMNH P14424 248 117 63 90 161 24 75
Holmesina cryptae sp. nov. LPP-PV-001 hol. 297.7 132.2 93.6 75.1 221.9 34.9 81
LPP-PV-002 par. 296.9 86 82.9 221.6 35.7 86.7
Holmesina rondoniensis MERO–P–002 hol. 325 145 97 83 220 39 88
Holmesina occidentalis ROM 3881 345 162 106 72 230 35 86
Holmesina paulacoutoi MCL-501/01 hol. 282 71 202 32 90
Holmesina septentrionalis HMNS 173 290 220 65 70
Pampatherium typum MACN Pv 11543 350 164 111 75 235 26 70
Pampatherium humboldtii MHD-P-28 360 174.6 113.7 72 237 34 83.5
UZM 2314 315 156 111 75 335 26 70
Hol., holotype; MDFM, maximum diameter of foramen magnum; par., paratype; WB, bicondylar width.
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TABLE 6. Measurements (maximum anteroposterior length) of the upper molariforms of pampatheres.
Species Specimen LTR Molariforms
Mf1 Mf2 Mf3 Mf4 Mf5 Mf6 Mf7 Mf8 Mf9
Kraglievichia MACN Pv 2617 6.5 6.0 6.2 12.6 12.7 14.5 15.5 12 8.5
Holmesina floridana UF 121742 116 6.8 7.1 7.2 12.3 15.4 17.5 16.4 13.1 10.3
UF 248500 - 7.0 7.5 9.0 10.7 15.9 16.8 15.3 13.3 9.8
Vassallia maxima FMNH P 14424 144 6.8 8.0 8.5 14.5 18.5 19.0 17.5 16.7 13.7
Holmesina cryptae sp. nov. LPP-PV-001 hol. 177.8 9.15 9.55 18.1 22.2 23.9 24.4 25.4 17.9
LPP-PV-002 par. 178.6 8.4 9.9 12.4 17.7 24.4 26.6 25.1 22.4 17
Holmesina rondoniensis MERO–P–002 hol. 180 8.1 12 17.5 19 24 27.0 26.5 21.0 17.1
Holmesina occidentalis ROM 3881 190 10 13 16 17.1 25 27.9 26 23.5 14.5
Holmesina paulacoutoi MCL-501/01 hol. 180 12 19 20.4 26.4 26.0 26.2 22.8 15
Holmesina septentrionalis HMNS 173 165 10 12 14.2 17.2 21.2 23.2 23.5 21.2 13.2
UF 234224 - 10 13 15 16 18 22 23 21 20
Pampatherium typum MACN Pv 11543 200 7.8 8.2 9.6 15.8 16.9 19.4 17.5 15.4 17.9
Pampatherium humboldtii MHD-P-28 195 9.2 8 10 14.4 21 23.7 24.2 21.7 20
UZM 2314 200 8.1 11 13 19.1 24.1 26 24.1 22 18
Hol., holotype; Mf, upper (maxillary) molariform; LTR, tooth row length; par., paratype.
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An anterior decrease is more noticeable in H. floridana (UF 224450) and H. occidentalis. Anteriorly to the mf1, the
mandible of H. cryptae sp. nov. (Fig. 13) shows a more pronounced and arched spout, this character is also observed
in H. floridana. In a specimen of P. typum, MACN Pv 11544, this spout is longer but is not arched. The horizontal
rami of P. typum and P. humboldtii are higher throughout all their length and slightly narrower anteriorly. The mental
foramina are located anteriorly in the horizontal ramus and ventrally to mf3, mf4, mf5. In H. cryptae sp. nov., they
are small and very close. In H. floridana, these foramina are associated to grooves. In H. septentrionalis, P. typum
and P. humboldtii, they are aligned, but distant from each other, and they are larger and deeper than in H. cryptae
sp. nov.
TABLE 7. Mandibular measurements of pampatheres.
Species Specimen Measurements (mm)
HAR HHRmf7 Maximum length Tooth row length
Vassallia minuta MLP 29-IV-15-4 hol. 24.5 75
Vassallia maxima FMNH P14424 12 18.5 41 11.6
Kraglievichia MLP 52-IX-30-67 32.5
Holmesina floridana UF 224450 100 19.3 38.5 120
Holmesina cryptae sp. nov. LPP-PV-002 par. 160 224 64 177
Holmesina septentrionalis HMNS 173 160 60 165
Pampatherium typum MACN Pv 11474 170 270 71 183
Pampatherium humboldtii MLP 81-X-30-1 187 290 73 212
HAR, ascending ramus height; HHRmf7, height of the horizontal ramus at the seventh molariform; hol., holotype; par.,
paratype.
In lateral view, the ascending ramus together with the horizontal ramus form an obtuse angle in H. cryptae sp.
nov. Depending on the species, this angle can be more open or closed and this can hide more or less the last molari-
forms in lateral view. This angulation hides the mf9 in S. hondaensis and Kraglievichia. In V. minuta and H. flori-
dana, the distal portions of mf8 and mf9 are hidden. In V. maxima, H. septentrionalis, P. typum and P. humboldtii,
the mf8 is almost totally hidden. In H. cryptae sp. nov., the mf8 is more exposed than the cited species.
The ascending ramus shows the crest of the coronoid process, which is shorter and rougher in V. maxima, lon-
ger and wider in H. cryptae sp. nov. and P. humboldtii, and slightly marked in H. floridana. The masseteric fossa
is more superficial in V. minuta and S. hondaensis, deeper and wider in V. maxima and H. floridana. In Quaternary
pampatheriid species, the masseteric fossa is notably wide and deep in P. typum and P. humboldtii, followed by H.
septentrionalis and lastly, in terms of robustness and deep, H. cryptae sp. nov.
In H. cryptae sp. nov., the mandibular condyle is long, narrow and slightly straight similar to P. humboldtii. In
H. septentrionalis, it is shorter, wider and has a posterior curvature. In H. floridana, it is more rectilinear than the
previously cited species. The angular region is located posteriorly to the mandibular condyle, it is laterally and me-
dially rougher and more robust in P. humboldtii, followed by P. typum and V. maxima. In H. septentrionalis, it is also
deep and striated (less than in Pampatherium), however it is more robust and rougher than in H. cryptae sp. nov.
In medial view, the mandibular symphysis is shorter than in V. maxima and H. floridana, extending to the adja-
cency of the mf4. In S. hondaensis and V. minuta, it extends as far as half of the mf5. This symphysis is longer in H.
floridana and the other Pleistocene species.
Posteriorly to the symphysis, there is the medial mandibular groove, which is short and deep in V. minuta, even
shorter and superficial in V. maxima, and less pronounced in H. floridana. In H. septentrionalis, this structure is
absent. In H. cryptae sp. nov., P. humboldtii and P. typum this groove is very deep.
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TABLE 8. Measurements (maximum anteroposterior length) of the lower molariforms of pampatheres.
Species Specimen LTR Molariforms
mf1 mf2 mf3 mf4 mf5 mf6 mf7 mf8 mf9
Vassallia minuta MLP 29-IV-15-4 hol. 75 *5.1 *7.0 *7.5 *11.0 *11.0 *12.0 *10.5 *8.0
Vassallia maxima FMNH P14424 11.6 7.2 *7.4 *11.5 *18.5 19.5 17.0 15.7 12.8
Kraglievichia MLP 52-IX-30-67 14.5 15.5 12 8.5
Holmesina floridana UF 223813 120~ 8.2 16.0 14.0 10.3
Holmesina cryptae sp. nov. LPP-PV-002 par. 177 7.2 9.7 11.7 17.6 23.2 25.4 25.1 24.1 20.4
Holmesina septentrionalis AMNH 26856 neo. 165 10.2 12.2 13.5 *19.6 22.3 *22.4 19.1 *13.0
Pampatherium typum MACN Pv 11474 183 6.8 8.9 9.2 13.2 23.3 25.1 24.9 23.8 *19.5
Pampatherium humboldtii MLP 81-X-30-1 212 7.1 8.1 10.5 14.1 24.9 27 25.8 24.6 21.2
* alveolar measurement; hol., holotype; LTR, tooth row length; mf., lower molariform; neo., neotype; par., paratype.
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Another structure present is the mylohyoid groove, which varies in length and depth. It is short, narrow and
deep in V. maxima and H. floridana, and equally narrow and deep but longer in S. hondaensis and Kraglievichia. In
H. cryptae sp. nov., it is short, narrow and less deep as in the other Quaternary species. In occlusal view, the lower
molariforms show the same morphology as the upper ones (see above), differing subtly in size (Fig. 13 and Table
8).
Hyoid apparatus. Only the right stylohyal and epihyal of LPP-PV-001 (Figs. 14A, 14B) and the basithyrohyal
(= V-bone) of LPP-PV-002 (Fig. 14C) were preserved. The stylohyal is longer than the epihyal, differently from
most cingulated, but observed in Priodontes Cuvier and Prozaedyus Ameghino and most pilosans. However, the
distal end of the stylohyal is tapered and it is a curved, concave-convex bone in lateral view as in most cingulates
(Casali & Perini 2017). In glyptodonts and Proeutatus Ameghino, the stylohyal and epihyal are fused, but in H.
cryptae sp. nov. and in other cingulates, they are independent bones (Casali & Perini 2017).The ceratohyal of H.
cryptae sp. nov. was not preserved.
As in all Xenarthra, H. cryptae sp. nov. has the basihyal fused to the thyrohyals, forming the V-bone (Flower
1885; Casali & Perini 2017). In H. cryptae sp. nov., this bone shows a basihyal apophysis, a common characteristic
to all cingulates except in Priodontes and Prozaedyus (Casali & Perini 2017). No lingual process in basihyal was
found in H. cryptae sp. nov. This structure is only observed in glyptodonts (Casali & Perini 2017).
FIGURE 14. Hyoid apparatus of Holmesina cryptae sp. nov. A–B, lateral view (holotype, LPP-PV-001); A, stylohyal; B, epi-
hyal; and C, (paratype, LPP-PV-002) ventral view of the basithyrohyal co-ossification (= v-bone). Scale bar = 30 mm.
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FIGURE 15. Vertebral column of Holmesina cryptae sp. nov. A-E, presynsacral vertebrae of the holotype (LPP-PV-001). A,
dorsal view of the articulated presynsacral vertebrae. B–C, mesocervical bone in B, dorsal; and C, lateral views. D–E, postcer-
vical bone in D, dorsal; and E, lateral views. F, xenarthrous vertebrae of the paratype (LPP-PV-002). Scale bars = 50 mm. G,
caudal vertebrae of the holotype (LPP-PV-001); Scale bar = 100 mm. Abbreviations: C1–C7, cervical vertebrae; L1, first lumbar
vertebra; T4–T10, thoracic vertebrae; m, mesocervical bone (C2+C3+C4+C5); p, postcervical bone (T1+T2+T3).
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Postcranial skeleton
Vertebral column. H. cryptae sp. nov. presents seven cervical (C), ten thoracic (T), one lumbar (L), 11 synsacral
(S) and 22 caudal (Cd) vertebrae (vertebral formula: C7T10L1S11Cd22). In detail, the synsacrum is formed by the
fusion of six sacral vertebrae and five caudal vertebrae (Fig. 15; Appendices 3, 4).Other co-ossifications were ob-
served in the vertebral column. The first anterior co-ossification is the mesocervical bone (Scillato-Yané 1982) (Fig.
15B, 15C) which is composed by the axis, the third, fourth and fifth cervical vertebrae (C2+C3+C4+C5). Likewise,
the first three thoracic vertebrae (T1+T2+T3) are fused (Fig. 15D, 15E) forming the second vertebral co-ossifica-
tion. These two co-ossifications also occur in P. humboldtii (MCL 900) and H. paulacoutoi and they were observed
in P. humboldtii (Winge 1915); yet in H. floridana (UF 240947), the mesocervical bone is formed by the synostosis
of C2+C3+C4. The postcervical co-ossifications observed in vertebrae of H. cryptae sp. nov. are characteristic of
Pampatheriidae (Paula Couto 1980). Presacral co-ossifications are also observed in other cingulates and the number
of fused vertebrae varies in the different species even intraspecifically (Scillato-Yané 1982; Cartelle & Bohórquez
1984; Galliari et al. 2010). In H. cryptae sp. nov., the number of lumbar vertebrae is less than most species of living
armadillos (see Galliari et al. 2010).
FIGURE 16. Chevrons of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 30 mm.
Accurately, the xenarthrous vertebrae of H. cryptae sp. nov. are the sixth, seventh, eighth, ninth and tenth tho-
racic and the first lumbar (T6–T10 and L1) (Fig. 15F). Góis et al. (2015) described the xenarthrous vertebrae of T.
mirus (holotype, MLP 54-III-16-1), however just two are preserved. Anteriorly on each side of the vertebral body
of T8–S1, there is the anterior lateral zygapophyseal facet (ALZ) and the anterior xenarthrous facet (AX) forming a
concavity where the anterior vertebral anapophysis fits. The vertebral foramen, which is above the vertebral body,
shows in its anterior upper border the anterior medial zygapophyseal facets (AMZ). Posteriorly on each side of the
foramen vertebral of T7–L1, there is the anapophysis whose surface is the posterior lateral zygapophyseal facet
(PLZ) and the dorsal surface is the posterior xenarthrous facet (PX). In the zygapophyses, there are the posterior
medial zygapophyseal facets (PMZ). PMZ articulates with AMZ of the next xenarthrous vertebra and PX/PLZ ar-
ticulate with AX/ALZ of the next vertebra, and it is the additional articulation of the xenarthrans.
T6 exhibits intermediate morphology; it possesses xenarthry only posteriorly, but resembles other anterior
thoracic vertebrae and differs from other xenarthrous vertebrae. Its anapophysis only has PX that has a very small
area and this articulation plan is anomalously above the PMZ plan. Consequently, T7 metapophysis only bears AX
anteriorly, but its posterior portion is similar to the following thoracolumbar vertebrae. Because its posterior portion
fuses to the sacral vertebrae, S1 presents a xenarthrous process just in its anterior portion.
All thoracic vertebrae (T1–T10) present costal facets for the ribs. In general, there are three articulations, 1) the
posterior costal demifacet for the head of the rib, lying anterolaterally on the vertebral body; 2) the costal facet for
the tubercle of the rib, lying in the metapophysis; and 3) the anterior costal demifacet for the head of the next rib,
lying laterally to the anapophysis continuing with the PLZ (see Fig. 15F). In the postcervical bone, the demifacets
are fused and T1 does not present the costal facet for the tubercle of the rib (see Fig. 15E). The anterior caudal
vertebrae that are not ankylosed to the synsacrum have a transverse process diameter relatively large (Fig. 15G and
Appendix 4).
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Chevrons. Only five chevron bones of LPP-PV-001 were preserved (Fig. 16). The anterior bones are larger
and have two diverging lateral processes, as in Priondontes in which these structures are widely diverging (Flow-
er, 1885). The posterior chevrons are V-shaped with only one medial spine as observed in P. humboldtii (Winge
1915).
FIGURE 17. Vertebral portions of ribs of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Second to tenth right ribs. Scale
bar = 30 mm.
FIGURE 18. Thoracic bones and clavicle of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A, right first anterior rib; B,
manubrium; and C, right clavicle. Abbreviations: ace, acromial end; ssn, suprasternal notch; ste, sternal end; vee, vertebral end.
Scale bar = 30 mm.
Ribs and sternum. Cingulates present ossified vertebral and sternal portion of the ribs. LPP-PV-001 possesses
10 ribs on the right side, but just the vertebral portion of them is preserved. The first left rib is also entire (Fig. 17).
The first anterior pair of ribs are shorter in comparison to the vertebral portion of the other posterior ribs, however
they are broader and flatter (Fig. 18A). Their sternal portion is short and fused to the vertebral portion as in other
armadillos (Flower 1885).
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The manubrium is pentagonal, except in its anterior border where there are the suprasternal (jugular) notch.
This bone is flat, robust and concave-convex; it articulates with the clavicle and with first anterior pair of ribs (Fig.
18B).
Scapular girdle. The scapulae in both specimens are fragmented; however, the right scapula of LPP-PV-001 is
more complete, with the central region of the fossae, medial border and spine broken (Fig. 19). Nevertheless, a well-
developed acromion and coracoid process can be observed. Also there is the second scapular spine, a xenarthran
characteristic (more developed in armadillos and anteaters) shared with some mammals. Propalaehoplophorus
Ameghino also possesses this characteristic but in late diverging glyptodonts it is less developed (Scott 1905).
The clavicle of H. cryptae sp. nov. is more robust and less when compared to armadillos. Its acromial end is
concave and possesses an acute process, yet the sternal end is round (Fig. 18C).
FIGURE 19. Right scapula of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 30 mm.
Pelvic girdle. Pelvic bones form a single co-ossification (Fig. 20) as a result of a fusion of the hip bones (is-
chium, ilium and pubis) with the synsacrum. In dorsal view, anteriorly, the synsacral ala is observed, the lateral
synsacral crests are observed (on the region of the sacral vertebrae); the median crest, as well as ten sinsacral fossae
are observed on the dorsal surface of the pelvis, hence the dorsal plan is formed by the synsacrum. The synsacral
vertebrae of H. cryptae sp. nov. comprises 11 fused vertebrae, in which post-iliac vertebrae are the five posterior-
most. According to Galliari & Carlini (2014, 2018), the number of synsacral vertebrae varies among species, even
among intrageneric species of Xenarthra and also among individuals in the same species, but no difference was
observed between the analyzed specimens of H. cryptae sp. nov.
The lateral borders of the synsacrum are fused to the hip bones, anterolaterally to the ilium (sacral vertebrae)
and posterolaterally to the ischium (synsacral caudal vertebrae). In addition to the obturator foramen, anteriorly
above the acetabulum, there is the synsacroischial foramen, its ventral border is the region where the ilium fuses to
the ischium, and its dorsal border is formed by the borders of the transverse processes of the first synsacral caudal
vertebrae (Table 9).
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TABLE 9. Pelvic measurements of H. cryptae sp. nov.
Specimen Measurements (mm)
Median synsacral crest Posterodorsal border
of the synsacrum
Obturator
foramen
pelvic outlet acetabulum
ML MH MW MD MH MD
LPP-PV-001 hol. 471.2 54.76 302.48 129.75 217.83 52.9
LPP-PV-002 par. 450.07 49.02 308.93 142.41 212.8 52.2
Hol., holotype; MD, maximum diameter; MH, maximum height; ML, maximum length; MW, maximum width; par.,
paratype.
Forelimbs
Humerus. In the proximal epiphysis, H. cryptae sp. nov. presents a round humeral head (Fig. 21A, 21B) similar to
H. paulacoutoi, P. humboldtii and H. floridana. The intertubercular groove is well marked in all species. The greater
tuberosity (laterally to the head and the lesser tuberosity) is less prominent in H. cryptae sp. nov. than in P. hum-
boldtii that is higher and pointed; in H. floridana it is relatively lower, i.e., the humeral head is higher, whereas in
H. paulacoutoi it is similar to H. cryptae sp. nov. The lesser tuberosity is medially projected bellow the neck; in H.
cryptae sp. nov. this tuberosity is small such as in H. floridana and P. humboldtii; in H. paulacoutoi this structure is
well defined. The neck is well marked in all species. Following the neck, the deltoid tuberosity and the epicondylar
crest morphology is typical of most xenarthrans (Rose & Emry 1993). These structures are more robust in armadil-
los than in Pampatheriidae. In H. cryptae sp. nov., the deltoid tuberosity and the epicondylar crest are well marked,
however less marked than in H. paulacoutoi and P. humboldtii (Table 10).
TABLE 10. Humerus measurements of pampatheres.
Species Specimen Measuments (mm)
Maximum
length
Deltoid tuberos-
ity length
Width at proxi-
mal end
Width at distal
end
Holmesina floridana UF 24932 61.5 36.8
H. cryptae sp. nov. LPP-PV-001 228 112.3 60 77
Holmesina septentrionalis HMNS 173 216 118 57 81
Holmesina paulacoutoi MCL-501/02-03 227 124 68.8 92.6
Pampatherium humboldtii MCL-900/01-02 225 122 61.0 83
FIGURE 20. Pelvis of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A, dorsal; and B, lateral views. Abbreviations: ac,
acetabulum; lsc, lateral synsacral crest; of, obturator foramen; sif, synsacroischial foramen; ssa, synsacral ala. Scale bar = 50
mm.
The structures in the distal part of the humerus are different among the compared species. The supracondy-
lar foramen in H. cryptae sp. nov. (both holotype and paratype) is obliterated, and its size is similar to that of H.
paulacoutoi and P. humboldtii. This foramen in pampatheres is longer than wide, in contrast to armadillos where
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the foramen is wider than long. In H. cryptae sp. nov., the lateral supraepicondylar crest is more robust than in P.
humboldtii, however this characteristic is more robust and wider in H. paulacoutoi.
FIGURE 21. Right forelimb bones of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A–B, humerus in A, anterior; and B,
posterior views. C–D, radius in C, anterior; and D, posterior views. E–H, ulna in E, lateral; F, anterior; G, posterior; and H, me-
dial views. Abbreviations: hsp, hook-shaped projection; scf, supracondylar foramen; tn, trochlear notch. Scale bar = 50 mm.
The medial epicondyle in H. cryptae sp. nov. is straight as in H. paulacoutoi, whereas it is round in P. humbold-
tii. The lateral epicondyle is a small tuberosity laterally to the capitulum. In H. cryptae sp. nov., this epicondyle is
round as in P. humboldtii; in H. paulacoutoi it is rounder and shows a lateral projection. In pampatheres, the coro-
noid fossa is well developed. In H. cryptae sp. nov., this fossa is wide and deep, it has a pyramidal shape with a deep
base; the same is observed in P. humboldtii and H. paulacoutoi but the depth is greater in these latter species. The
humeral trochlea (anterior view) is reel-shaped. The capitulum (anterior view) is round as in other species.
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Ulna. This element presents a slight curvature (Fig. 21E–21H). In H. cryptae sp. nov., the olecranon surface is
irregular with a notable lateral hook-shaped projection, similar characteristics are observed in H. paulacoutoi and H.
floridana, however this projection is smaller and less marked in these other species. In P. humboldtii this projection
is absent. Posteriorly to the proximal region, there is a concavity in H. floridana, P. typum and H. paulacoutoi. In
contrast, the same region in H. cryptae sp. nov. and P. humboldtii shows a convexity, more pronounced in P. hum-
boldtii. The olecranon apophysis (lateral view) in H. cryptae sp. nov. is well pronounced and beak-shaped with its
curvature facing medially as in P. humboldtii and H. paulacoutoi; in H. floridana is less marked.
The greater sigmoid cavity (trochlear notch), where the humeral trochlea articulates, is deep and has a transver-
sal disposition. At the distal end, there is also the radial notch which articulates with the radius. In H. cryptae sp.
nov., this notch is well defined as in other pampatheres. The coronoid apophysis is well developed in P. typum. In
lateral and medial view (Fig. 21E, 21H), very deep fossae are observed in P. typum and H. cryptae sp. nov., which
are shallower in H. floridana and P. humboldtii. Distally, there is the ulnar head that is rounder in H. cryptae sp. nov.
than the other species. Comparative measurements of the ulnae of pampathere species are given in Table 11.
TABLE 11. Ulna measurements of pampatheres.
Species Specimen Measurements (mm)
Maximum length Olecranon length
Holmesina floridana UF 24932 129 -
Holmesina cryptae sp. nov. LPP-PV-001 219 73.9
Holmesina septentrionalis HMNS 173 209 -
Pampatherium humboldtii MCL-900/01-02 227 82.4
Radius. The head is separated from the body by a well-marked neck laterally and medially as in H. cryptae
sp. nov. (Fig. 21C, 21D) and in H. paulacoutoi. This characteristic is less defined in P. humboldtii. In the proximal
region, the bicipital tuberosity is located in the anterior surface below the neck. In H. cryptae sp. nov., the tuberos-
ity is ovoid and is well defined, however less than in H. paulacoutoi, but in P. humboldtii it is more superficial; this
tuberosity completes the radius-ulna articulation with the humeral trochlea.
The interosseous crest is located in the most distal portion. In H. cryptae sp. nov. it is less prominent than in H.
paulacoutoi. Its distal border is more robust. Its surface is relatively plane and rough, and a small sigmoid cavity
turned to ulnar head is observed. The styloid apophysis is in the posteromedian part of the sigmoid cavity, and in the
anterolateral part there is a pronounced apophysis wherein its surface articulates with the scaphoid and the lunate.
Manus. Holmesina cryptae sp. nov. has five digits on each hand, as other pampatheres. The distal (ungueal)
phalanges are shorter and less robust than H. paulacoutoi and P. humboldtii and similar to H. septentrionalis. How-
ever, the main morphological characteristics of the structure is preserved. Digits II. III and IV are much longer and
more robust than digits I and V, in which digits II and III are longer than digit IV. H. cryptae sp. nov. has the palmar
bone, also observed in P. humboldtii (Winge 1915) and in other cingulates.
Hind limbs
Femur. In H. cryptae sp. nov. the greater trochanter is slightly higher than the femoral head (Figs. 22A, 22B), yet
in Kraglievichia, P. humboldtii and T. mirus the greater trochanter is much higher than the femoral head. Holmesina
floridana shows an intermediate morphology, and in H. paulacoutoi these structures are at the same level, as in some
glyptodonts (Góis 2013; Góis et al. 2015).
The greater trochanter is robust, laterally compressed, and its maximum diameter is anteroposteriorly oriented
and the femoral head is circular. In H. cryptae sp. nov., the femoral neck is more defined than in P. humboldtii
and H. paulacoutoi, which in turn shows a more defined femoral neck than Kraglievichia. The lesser trochanter
in H. cryptae sp. nov. has a shape slightly suboval and it is distally positioned. In P. humboldtii it is similar to H.
cryptae sp. nov. but it is medially oriented. The trochanteric fossa that is deep in H. cryptae sp. nov., as well as in
Kraglievichia, H. floridana and H. paulacoutoi. In the latter, this fossa is deeper and wider. The third trochanter in
Kraglievichia is located closer to the middle of the bone length than distally, differently to the other species in which
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it is more distal. In H. cryptae sp. nov., this trochanter is less robust than in T. mirus and H. paulacoutoi but is more
robust than in H. floridana and Kraglievichia (Table 12).
TABLE 12. Femur measurements of pampatheres.
Species Specimen Measurements (mm)
Maximum length widht across the
3rd trochanter
width of the distal
end
Kraglievichia MLP 69-IX-8- 13A 164 33.5 38
Holmesina floridana UF 24918 193.5 41.3 51.5
Tonnicinctus mirus MLP 54-III-16-1 holotype 280 62 85
Holmesina cryptae sp. nov. LPP-PV-001 holotype 323 71 70
Holmesina septentrionalis HMNS 173 290 70 86
Holmesina paulacoutoi MCL-501/08 359 83 91
Pampatherium humboldtii MCL-900/05 347 72 92
The lateral epicondyle is greater than the medial one. In H. cryptae sp. nov., both epicondyles are more defined
than in S. antelucanum, H. floridana, P. humboldtii, H. paulacoutoi but less than in T. mirus. Distally, contrary to
the condyles, the patellar facets are asymmetric in H. cryptae sp. nov., as well as in the rest of the species examined.
The condyles (medial and lateral) possess similar sizes, except in T. mirus in which the medial condyle is smaller
than the lateral condyle.
Tibiofibula. Although they differ in size and robustness (Table 13), there are no significant differences among
the main characteristics of both species. In the proximal end of the tiobiofibula, the facets to the condyles of the fe-
mur are asymmetrical, the lateral one is slightly convex and the medial one is concave and proceeded by an apophy-
sis (Fig. 22E, 22F). In H. cryptae sp. nov., the tibiofibula is more robust and compact than in H. floridana.
TABLE 13. Tibiofibula measurements of Holmesina cryptae sp. nov. and H. floridana.
Species Specimen Measurements (mm)
Maximum length Width of proximal
epiphysis
Width of distal
epiphysis
Holmesina floridana UF 294765 121 - 46
Holmesina cryptae sp. nov. LPP-PV-001 holotype 198 85 82
The body of the tibia is more robust than that of the fibula, especially the proximal half that presents a promi-
nent tibial crest, then it finishes distally in the malleolus, this characteristic is common to both species. The body of
the fibula exhibits an anterior crest that terminates in an acute apophyses. The distal end presents a slight torsion in
relation to the proximal end. In the distal epiphysis of the tibia, the articular facet with the astragalus is subdivided
into two, in which the medial one is larger and the lateral one finishes posteriorly in a triangular apophysis facing
downwards. The distal end of the fibula shows an elipsoidal articular facet for the calcaneum, and its malleolus is
lateral.
Patella. It is a triangular bone with rounded angles (Fig. 22C, 22D). In anterior view, H. cryptae sp. nov. pres-
ents a well-defined posterior projection in the base. This characteristic is absent in H. floridana and T. mirus, and in
P. humboldtii is similar but less pronounced. The lateral borders are best defined in H. cryptae sp. nov., followed by
P. humboldtii, T. mirus, and H. floridana. In the latter, the outline is almost imperceptible. A long and narrow apex is
another noticeable characteristic in the patella of H. cryptae sp. nov. In the other species the apex is short and wide.
The articular surface for the femur is similar in all species.
Pes. In H. cryptae sp. nov., the neck of the astragalus is less marked and deep than in H. floridana and P. hum-
boldtii, but it is similar to H. paulacoutoi. The head of the astragalus is spherical in all species. The trochlea of the
astragalus that articulates with the tibiofibula is asymmetrical and the outer part is more robust than the inner one
(Fig. 23A, 23B). In ventral view of the body of the astragalus, the sustentacular facet is larger than the ectal facet in
H. cryptae sp. nov. and P. humboldtii. On the other hand, in H. floridana and H. paulacoutoi; these facets are similar
in size. In H. cryptae sp. nov. and P. humboldtii, both facets are separated by a deep cleft, which is narrower than in
H. paulacoutoi. In H. floridana, both facets are in contact.
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FIGURE 22. Right hind limb bones of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A–B, femur in A, anterior; and B,
posterior views. C–D, patella in C, anterior; and D, posterior views. E–F, tibiofibula in E, anterior; and F, posterior views. Scale
bar = 50 mm.
The calcaneus presents a posterior elongation, that is the calcaneal tuberosity (tuber calcis) for insertion of the
gastrocnemius tendon (Fig. 23C, 23D). Anterodorsally, the calcaneus presents two facets for the astragalus, the ectal
and sustentacuar facets, both separated by a sulcus. The calcaneal tuberosity in H. cryptae sp. nov. is less and less
robust than in H. paulacoutoi and P. humboldtii (Fig12). This structure is less rounded in H. paulacoutoi and in H.
cryptae sp. nov. it presents a crest in its proximal (superior) portion that is more marked than in P. humboldtii, and
that is virtually absent in H. paulacoutoi (Table 14). The size and shape of the facets differ according to the taxa
(Table 14).
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434 · Zootaxa 4661 (3) © 2019 Magnolia Press
TABLE 14. Calcaneus and astragalus measurements of pampatheres.
Species Specimen Measurements (mm)
Calcaneus Astragalus
Maximum length Width across the
articular surface
Maximum length
Holmesina floridana UF 294765 64.1 24.5
Holmesina cryptae sp. nov. LPP-PV-001 holotype 102.2 45.5 49.93
Holmesina paulacoutoi MCL 501/11 and 13 holotype 110 45
Pampatherium humboldtii MCL 900/07 and 08 104.45 43.6 51.08
The other tarsal elements (cuboid and navicular), the metatarsi and the phalanges (five digits in each foot)
are similar to P. humboldtii. Holmesina cryptae sp. nov. presents a plantar bone in each foot, also observed in P.
humboldtii (Winge 1915) and other cingulates. These bones are rounded and dorsally they present two facets that
articulate with tarsal and talar elements.
Discussion
In the present study, new characteristics in Holmesina cryptae sp. nov. different from other species of pampatheres
were identified. Most of these are cranial, facial and dental characteristics. Superficially, this can be seen by the
skull shape that is more triangular than the other species with an orthogonal zygomatic arch, the most robust parietal
region and a robust snout. Regarding the ornamental pattern, the osteoderms of H. cryptae sp. nov. share some char-
acteristics with Holmesina or Pampatherium, whereas some characteristics are intermediate between both genera
(Table 15). Thus, it would be difficult to identify the species of H. cryptae sp. nov. based only on the osteodermal
characters, even because some ornamental patterns in osteoderms of H. cryptae sp. nov. vary depending on the re-
gion in carapace. Some pampatheriid species show distinctive ornamentation in their osteoderms as T. mirus and H.
paulacoutoi however these features are less evident in some regions of the armor as observed in H. cryptae sp. nov.
Therefore, recognizing a species or genus of a cingulate based on patterns of osteoderms from different localities
can be unreliable especially when they are unassociated with endoskeletal elements (Simpson 1948; Hoffstetter
1958; De Iuliis et al. 2000; Perea 2005).
However, most cranial characteristics observed in H. cryptae sp. nov. are also present in other Holmesina spe-
cies (see Table 15). Gaudin & Lyon (2017) identified 13 diagnostic cranial features for the genus Holmesina. In H.
cryptae sp. nov., at least 10 of those 13 morphological characters were recorded (Table 15): a) the nasal becomes
narrower posteriorly; b) premaxillary-maxillary suture M-shaped in ventral view; c) maxillary-palatine suture U-
shaped in ventral view; d) presence of prominent lateral maxillary ridge and deep antorbital fossa; e) reniform Mf4
and bilobate mf5; f) lack of orbital exposure of the palatine; g) ethmoid foramen entirely within the frontal, lacking
orbitosphenoid participation in rim; h) no orbitosphenoid participation in rim of sphenopalatine foramen; i) trian-
gular stylohyal fossa with distally expanded thympanohyal; j) strong medial flange of petrosal marked by pits and
ridges; and k) ventral displaced internal acoustic meatus.
When the contemporary Quaternary genera (De Iuliis et al. 2000; Góis et al. 2015) are compared, it is possible
to confirm certain morphological differences among Holmesina, Pampatherium and Tonnicinctus, for instance:
osteoderms of Holmesina have a more marked and delimited ornamental pattern than Pampatherium, but Ton-
nicinctus has the most complex ornamentation (Góis et al. 2015). The anterior molariforms of Holmesina species
are imbricated, and the posteriormost molariforms of Pampatherium species show a trilobulation whereas they are
less complex in Holmesina (Fig. 11) (Scillato-Yané 1982; Edmund 1996; De Iuliis & Edmund 2002; Góis et al.
2013; Góis et al. 2015). Morphofunctional and paleoecological studies about pampatheres are scarce in the litera-
ture. Vizcaíno et al. (1998) analyzed the masticatory apparatus of V. maxima and H. occidentalis, and verified that
the former applied relative bite forces greater than the latter; consequently, Vassallia was more adapted to feed on
more abrasive vegetation than H. occidentalis; but, in general, all pampatheres are considered grazers. Along the
same lines, De Iuliis et al. (2000) analyzed the masticatory apparatus of Pampatherium species in comparison to
Holmesina and other pampatheres. They suggested that P. typum and P. humboldttii consumed more abrasive plants
than H. occidentalis and H. paulacoutoi. H. cryptae sp. nov. shows characters in masticatory apparatus as gracile as
in other Holmesina species, except for its zygomatic arch that is more robust.
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FIGURE 23. Right astragalus and calcaneus of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A–B, astragalus in A,
dorsal; and B, ventral views. C–D, calcaneus in C, dorsal; and D, ventral views. Scale bar = 50 mm.
Scillato-Yané et al. (2005) inferred the paleobiogeography of the pampatheres in South America, based on di-
etary adaptations and occurrences. De Iuliis et al. (2000) and Scillato-Yané et al. (2005) suggested that even though
some species have been found at the same place, it does not necessarily mean that they co-occurred contemporane-
ously in sympatry; it could represent uncontemporary expansion and contraction of species ranges due to climatic
and environmental changes during the late Pleistocene. Holmesina cryptae sp. nov. is the third pampatheriid species
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436 · Zootaxa 4661 (3) © 2019 Magnolia Press
found in Bahia state together to H. paulacoutoi and P. humboldtii (see Cartelle & Bohórquez 1984; Scillato-Yané et
al. 2005 and Ferreira et al. 2018). They may have co-occurred in sympatry or have lived in different times according
to hypothesis suggested by Scillato-Yané et al (2005).
TABLE 15. Characters of H. cryptae sp. nov. in comparison to Pampatherium and other Holmesina species.
Characters Pampatherium Holmesina H. cryptae
CARAPACE
Fixed osteoderms wide lateral margin 0 2 0
high marginal elevation. 0 2 0
Movable osteoderms and
SMOPB
deep intermediate portion 0 2 2
wide lateral margin 0 2 1
high marginal elevation 0 2 1
high central longitudinal elevation 0 2 1
SKULL
M-shaped premaxillary-maxillary
suture in ventral view
0 2 2
U-shaped maxillary-palatine suture
in ventral view
0 2 2
prominent lateral maxillary ridge
and deep antorbital fossa
0 2 2
reniform Mf4 and bilobate mf5 0 2 2
orbitosphenoid participation in rim
of sphenopalatine foramen
2 0 0
triangular stylohyal fossa with
distally expanded thympanohyal
0 2 2
ventral displaced internal acoustic
meatus
0 2 2
POSTCRANIUM
Humerus straight medial epicondyle 0 2 2
Ulna lateral hook-shaped projection in
olecranon
0 2 2
Femur greater trochanter higher than
femoral head
2 0, 1 1
SMOPB, semimovable osteoderm of pelvic buckler. Characters: 0, absent/opposite; 1, intermediate; 2, present.
Seven genera from the Middle Miocene and the Early Holocene belong to Pampatheriidae family occurring in
Americas with a wide latitudinal distribution: Scirrotherium, Vassallia, Kraglievichia, Plaina, Tonnicinctus, Pam-
patherium and Holmesina (Scillato-Yané et al. 2005; Góis 2013; Góis et al. 2013; Ferreira et al. 2018). During this
time interval they slightly changed, preserving many basic morphological characters, but the terminal taxa show
an enlarged size and robustness (Edmund 1987; De Iuliis et al. 2000), related to an improvement in the capacity
for processing more fibrous food (De Iuliis et al. 2000). Edmund (1987) suggests that the larger pampatheres could
reach 3 m in length. Probably, these estimates were inferred from dimensions of long bones (femur or humerus) of
the largest species. For example, the Quarternary species presents similar sizes of femoral measurements such as H.
cryptae sp. nov. (323 mm), P. humboldtii (347 mm) and H. paulacoutoi (359 mm), except H. septentrionalis (290
mm) (see Table 11). The integral study of the remains of H. cryptae sp. nov. (holotype, LPP-PV-001 and paratype
LPP-PV-002), enabled us to calculate a total axial length of 2.2 m. Quaternary pampatheres could reach around 2.5
m in length. This is larger than the largest living cingulate species, Priodontes maximus (Kerr), which can reach up
to 1.5 m in length (Desbiez & Kluyber 2013).
NEW PAMPATHERE FROM THE QUATERNARY OF BAHIA Zootaxa 4661 (3) © 2019 Magnolia Press · 437
Conclusion
In this paper, a new species of Pampatheriidae is described for Quartenary in South America. Holmesina cryptae sp.
nov. shows different characteristics from other species of pampatheres, however it shares the main cranial synapo-
morphies among the Holmesina species.
The osteoderms of H. cryptae sp. nov. possess some characteristics of Holmesina or Pampatherium, whereas
some characteristics are intermediate between both genera. Although H. cryptae sp. nov. has some characteristics
in the vertebral column also present in other cingulates (except in glyptodonts), such as the meso- and postcervical
co-ossified bones, it shows the most reduced number of thoracolumbar vertebrae.
H. cryptae sp. nov., H. paulacoutoi and P. humboldtii are described in close localities in the Brazilian Intertropi-
cal Region.
Acknowledgements
The authors are very grateful to the speleologist E. Cernawsky Igual (GPME), who collaborated with finding of the
fossils and provided us with all the support during the expedition; L. Joaquim (DEBE - UFSCar) for his support in
the field; A. Hochreiter, C.E. Martins, D.C. Oliveira, E.C. Igual, F. Komuro, G. Tinen, H. Komuro, J. Anjos Neto,
K. Komuro and M. Alzair R. A. Komuro (in memoriam) (GPME), for their tireless research and expeditions that
resulted in new discoveries in Iramaia and the finding of the fossils of this study; A. Ghilardi, B. Peixoto, B. Robbi,
C. Nascimento, L. Fernandes, P. Buck, and W. Dias (LPP-UFSCar) for their support and photographs; A. Alves
Dias, D. Florentino and J.S. Godwin Coury for copyediting our paper; M.E. Bichuette (LES-UFSCar) for having
contacted us about the findings; the reviewers A. K. Oliveira; D. A. Croft; D. Perea; K. Porpino for their construc-
tive suggestions; C. Cartelle (PUC-Minas), P. Auricchio (UFPI); R. Hulbert Jr.; and B.J. Macfadden for their sup-
port that helped us identify the taxon; H.E.D. Zaher and A.B. Carvalho (MZUSP) for allowing and performing the
CT scan of the skulls; and the organizers of I QUATERNE: IX SBPV. This work was supported by CNPq, Capes,
PPGERN-UFSCar.
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Supporting information
APPENDIX 1. Specimens examined for comparative study.
PAMPATHERIIDAE:
Scirrotherium hondaense: UCMP 40056 (paratype), 37979, 38066, 38883.
S. carinatum: MLP 69-IX-8-13AB (holotype), 69-IX-8-13AC (paratype), 69-IX-8-13AD, (paratype), 52-X-1-35 (paratype),
69-IX-8-13AE (paratype), 70-XII-29-1 (paratype).
S. antelucanum: CFM-2867 (holotype) 355, 1639.
Kraglievichia paranensis: MLP 41-XII-13-903, 41-XII-13-911, 41-XII-13-912, 60-VI-18-68, 69-VIII-22-3, 76-VI-12-1.
Kraglievichia cf. paranensis: MLP 69-IX-8-13A.
Vassallia minuta: MLP 29-IV-15-6, 29-X-10-12, 29-X-8-39, 69-IX-5-21, 69-XII-26-17, 95-VIII-1-1.
V. maxima: MLP 29-X-10-65, 52-X-2- 94, FMNH P14424.
Pampatherium humboldtii: MCL 900, 900/05, 900/06, 2308/01–798, MLP 81-X-30-1, MACN Pv 8490, 11905.
P. typum: MLP 34-IV- 12-6, 52-IX-28-20, 69-VIII-22-4, 69-VIII-25-11.
P. mexicanum: INAH 6201 (holotype).
Plaina brocherensis: MUFyCA 769 (holotype).
Holmesina septentrionalis: AMNH 23435, 26856 (neotype), ROM 19787, 19790, UF 16372, HMNS 173.
H. floridana: UF 17476, 24918, 184326.
H. occidentalis: EPN, V. 1068 (paratype), 1176 (paratype), 1086 (paratype), 1103 (paratype), ROM 26121–26170.
H. paulacoutoi: MCL-501/01 (holotype), 501/08 (holotype), 501/86–103 (holotype), MLP 69-VIII-25-13, MACN Pv14400,
CTES-PZ 7495.
H. rondoniensis: MERO-P-002 (holotype).
Tonnicinctus mirus: MLP 54-III-16-1 (holotype).
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APPENDIX 2. Figure of morphological diversity of osteoderms of Holmesina cryptae sp. nov. (holotype, LPP-PV-001).
A–B, appendicular osteoderms; C, cephalic shield osteoderm; D–F scapular buckler osteoderms; G–I, pelvic buckler
osteoderms; J, anterior semimovable osteoderm; K–Q, movable bands odteoderms; K–L, fused osteoderms of the border;
M–N, osteoderms of the border; O–P, medial osteoderms; Q, osteoderms of the border; R–S, osteoderms of the border of
the posterior semimovable band; T, osteoderms of the border of the carapace. Scale bar = 20 mm.
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APPENDIX 3. Table of measurements of precaudal vertebrae including synsacral caudal vertebrae of Holmesina cryptae
sp. nov. (LPP-PV-001 and LPP-PV-002) and Pampatherium humboldtii (MCL 900). C, cervical vertebra; L, lumbar ver-
tebra; S, synsacral vertebra; T, thoracic vertebra. *, also presents xenarthrous articulation.
Spinal region Bone Measurement (mm) LPP-PV-001 LPP-PV-002 MCL 900
Cervical vertebrae C1 Maximum height 51.67 54.23 55.15
Body length 23.19 21.61 20.85
Posterior body height 9.75 9.11 8.25
Posterior body width 73.8875 78.31 78.53
Mesocervical bone
(C2+C3 +C4 +C5)
Maximum height 81.49 71.89 73.5
Body length 85 85.5 84.18
Posterior body height 9.29 8.35 11.15
Posterior body width 29.3 28 30.4
C6 Maximum height
Body length 15.79 17.2
Posterior body height 10.31 9.13
Posterior body width 65.26 70.11
C7 Maximum height 52.6
Body length 17.04 ~13
Posterior body height 9.03 6.55
Posterior body width 29.09
Thoracic vertebrae Trivertebral bone
(T1 +T2 +T3)
Maximum height 96.09 103.6 99
Body length 85.35 78.03 86.2
Posterior body height 17.26 15.95 19.98
Anterior body width 64.78 70.5
Posterior body width 47.54 50.06 50.5
T4 Maximum height 75.14 93.29
Transverse process diameter 70.93 79.06
Body length 33.4 29.08
Anterior body height 17.5 17.16
Anterior body width 39.73 38.7
T5 Maximum height 78.36 88.76
Transverse process diameter 75.7
Body length 32.55 30.65
Anterior body height 20.8 19.43
Anterior body width 38.18 36.99
APPENDIX 4. Table of measurements of the caudal vertebrae of H. cryptae sp. nov. (holotype, LPP-PV-01). Cd,
caudal vertebra.
Bone Measurement (mm) LPP-PV-01
Cd1 Maximum height 77.47
Posterior body height 32.81
Body length 40.76
Transverse process diameter 234.88
Cd2 Maximum height 75.27
Posterior body height 33.36
Body length 39.63
Transverse process diameter 202.2
......continued on the next page
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APPENDIX 4. (Continued)
Bone Measurement (mm) LPP-PV-01
Cd3 Maximum height 73.07
Posterior body height 32.57
Body length 41.81
Transverse process diameter 165.56
Cd4 Maximum height
Posterior body height 32.35
Body length 45.9
Transverse process diameter 155.79
Cd5 Maximum height
Posterior body height 32.35
Body length 45.9
Transverse process diameter 143.16
Cd6 Maximum height
Posterior body height 31.47
Body length 46.96
Transverse process diameter 130.4
Cd7 Maximum height
Posterior body height 31.2
Body length 48.27
Transverse process diameter
Cd8 Maximum height 52.2
Posterior body height 30.66
Body length 48.27
Transverse process diameter
Cd9 Maximum height
Posterior body height 29.2
Body length 48.41
Transverse process diameter 99.41
Cd10 Maximum height
Posterior body height 28.5
Body length 47.9
Transverse process diameter
Cd11 Maximum height
Posterior body height ~27
Body length 48.03
Transverse process diameter
Cd12 Maximum height
Posterior body height 21.6
Body length 41.78
Transverse process diameter
Cd13 Maximum height
Posterior body height 20.23
Body length 40.56
Transverse process diameter
......continued on the next page
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APPENDIX 4. (Continued)
Bone Measurement (mm) LPP-PV-01
Cd14 Maximum height
Posterior body height ~18
Body length 44.78
Transverse process diameter
Cd15 Maximum height
Posterior body height 17.96
Body length 37.91
Transverse process diameter
Cd16 Maximum height
Posterior body height ~14.5
Body length 34.86
Transverse process diameter
Cd17 Maximum height
Posterior body height 13.51
Body length 32.7
Transverse process diameter
Cd 18 Maximum height 13.4
Posterior body height 12.48
Body length 28.9
Transverse process diameter 30.54
Cd19 Posterior body height 10.66
Body length 26.13
Transverse process diameter ~25
Cd20 Posterior body height 9.23
Body length 23.12
Transverse process diameter 20.5
Cd 21 Posterior body height 7.62
Body length 19.53
Transverse process diameter 14.5
Cd22 Posterior body height 5.78
Body length 16.06
Transverse process diameter 11.5
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... Received 10 February 2020; Received in revised form 26 June 2020; Accepted 30 June 2020 Brazil (Fig. 3). Its lithology is part of Gabriel and Nova América Units (Salitre Formation -Neoproterozoic) (Vieira et al., 2005) and presents bone accumulation of Quaternary mammals (Moura et al., 2019). The osteoderms were found associated to three individuals of Holmesina cryptae (Pampatheriidae) that were partially covered by the cave sediment (Moura et al., 2019). ...
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... Was calculated a mean value of humerus and femurs circumferences for several localities in BIR, based on the material found in paleontological collections in Brazil ( (Cartelle and Fonseca, 1982;Cartelle, 1992;Porpino, 1999;Porpino and Bergqvist, 2002;Cartelle et al., 2009;Porpino et al., 2009Porpino et al., , 2010Molena, 2012;Guérin and Faure, 2013;Moura et al., 2019;Silva et al., 2019;Greco et al., 2022; Table S1). ...
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Xenarthra is unique in having striking features in the axial skeleton, usually very different from the rest of the eutherian clades. Some of these features are widely spread among the members of the group (e.g., the presence of xenarthrales in most cingulates and all pilosans, and the synsacrum with sacroischial union to the pelvis in almost all xenarthrans) and others are restricted to some taxa (e.g., deviated number of cervical vertebrae in Bradypus and Choloepus). In this study, we aim to explore the great diversity of vertebral elements composing the xenarthran synsacrum within a phylogenetic framework. Vertebral counts of the adult synsacrum was obtained from almost all extant genera, with the exception of Calyptophractus, and several fossils. The modal number of vertebrae from the adult synsacrum was mapped onto a composite phylogeny of Xenarthra. The ancestral number of synsacrals for Xenarthra was recovered as ambiguous, although one of the optimizations recorded a number of six synsacrals, probably three iliac and three post-iliac vertebrae. The clade Cingulata is characterized by a high number of vertebrae forming the synsacrum (eight synsacrals), which is fused to the ischium through the tip of the transverse processes of the most posterior vertebrae. In pilosans, the ancestral number of synsacral vertebrae seems to be lower, probably formed by five or six vertebrae, and the union with the ischium is achieved through the base of the transverse processes of the most posterior vertebrae. Two exceptions stand out, one involving the extant suspensory sloths and Megalocnus, and the other involving a family of ground sloths, the Mylodontidae. A probable relationship of the synsacral number in the different taxa to the lifestyles is discussed. © 2018 Springer Science+Business Media, LLC, part of Springer Nature
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The genus Neoglyptatelus Carlini, Zurita, Scillato-Yané, Sánchez, and Aguilera has been considered a member of Glyptatelinae, a group encompassing the purportedly basal-most glyptodonts. It is up to now represented by two species from Colombia: Neoglyptatelus originalis Carlini et al., from the middle Miocene (a carapace fragment, isolated osteoderms and postcranial bones), and Neoglyptatelus sincelejanus Villarroel and Clavijo, from the middle or late Miocene (a partial carapace and a caudal armor). More scarce material assigned to this genus was recovered from the late Miocene of Uruguay and Brazil. In this article, we describe a new species, Neoglyptatelus uruguayensis, from the late Miocene Camacho Formation, Uruguay, based on an almost complete carapace and several postcranial bones. We conducted a phylogenetic analysis based on 167 morphological characters (23 new ones and 144 from previous analysis) scored for 19 taxa, encompassing some of the best known glyptodontid genera, one pampathere and four armadillos (including the enigmatic genus Pachyarmatherium Downing and White). In the most parsimonious tree that was obtained, Neoglyptatelus forms a clade with Pachyarmatherium (Pachyarmatheriidae), which is the sister group of the glyptodonts + pampatheres clade; consequently, it is not a glyptodont, as previously believed. This result, together with the known stratigraphic and geographic distribution of Neoglyptatelus and Pachyarmatherium, suggests that this new cingulate clade originated in South America and that Pachyarmatherium reached North America during the Plio–Pleistocene. The carapace of Neoglyptatelus and Pachyarmatherium comprises pelvic and scapular shields overlapping each other without separate intervening transverse mobile bands, an arrangement that differentiates both genera from the remaining cingulates.
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The present study entails descriptions of several well-preserved skulls from the pampathere species Holmesina floridanus, recovered from Pliocene localities in central Florida and housed in the collections of the Florida Museum of Natural History. Bone by bone descriptions have allowed detailed reconstructions of cranial morphology. Cranial foramina are described and illustrated in detail, and their contents inferred. The first ever description of an isolated pampathere petrosal is also included. Cranial osteology of Holmesina floridanus is compared to that of Pleistocene species of Holmesina from both North and South America (Holmesina septentrionalis, Holmesina occidentalis), as well as to the other well-known pampathere genera, to closely related taxa among glyptodonts (Propalaehoplophorus), and to extinct and extant armadillos (Proeutatus, Euphractus). This study identifies a suite of apomorphic cranial features that serve to diagnose a putative, progressive series of more inclusive monophyletic groups, including the species Holmesina floridanus, the genus Holmesina, pampatheres, pampatheres plus glyptodonts, and a clade formed by pampatheres, glyptodonts, and Proeutatus. The study highlights the need for further anatomical investigations of pampathere cranial anatomy, especially those using modern scanning technology, and for analyses of pampathere phylogenetic relationships.
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Glyptodonts were giant (up to ~2,400 kg), heavily armoured relatives of living armadillos, which became extinct during the Late Pleistocene/early Holocene alongside much of the South American megafauna. Although glyptodonts were an important component of Cenozoic South American faunas, their early evolution and phylogenetic affinities within the order Cingulata (armoured New World placental mammals) remain controversial. In the present study we used hybridisation enrichment and high-throughput sequencing to obtain a partial mitochondrial genome from Doedicurus sp., the largest (1.5m tall, and 4m long) and one of the last surviving glyptodonts. Our molecular phylogenetic analyses revealed that glyptodonts fall within the diversity of living armadillos. Reanalysis of morphological data using a molecular "backbone constraint" revealed several morphological characters that supported a close relationship between glyptodonts and the tiny extant fairy armadillos (Chlamyphorinae). This is surprising as these taxa are among the most derived cingulates: glyptodonts were generally large-bodied and heavily armoured, while the fairy armadillos are tiny (~9-17 cm) and adapted for burrowing. Calibration of our phylogeny with the first appearance of glyptodonts in the Eocene resulted in a more precise timeline for xenarthran evolution. The osteological novelties of glyptodonts and their specialisation for grazing appear to have evolved rapidly during the Late Eocene to Early Miocene, coincident with global temperature decreases and a shift from wet closed forest towards drier open woodland and grassland across much of South America. This environmental change may have driven the evolution of glyptodonts, culminating in the bizarre giant forms of the Pleistocene. This article is protected by copyright. All rights reserved.
Thesis
Los Pampatheriidae son una de las familias de mamíferos xenartros nativos de América del Sur. Estos cingulados tuvieron tallas medianas a grandes; los taxones terminales llegaron a pesar más de 250 kg (VIZCAÍNO et al., 1998), cinco veces más que el mayor armadillo actual, Priodontes maximus. Se registran desde el Mustersense (Eoceno tardío) hasta el Holoceno temprano en América del Sur; en América Central solo están representados en el Pleistoceno tardío y en América del Norte desde el Plioceno tardío hasta el Holoceno temprano. Al igual que los gliptodóntidos, son esencialmente herbívoros (WINGE, 1915; VIZCAÍNO et al., 1998). El objetivo de la presente Tesis Doctoral es realizar un estudio sistemático, anatómico y filogenético de los Pampatheriidae, evaluando sus afinidades con otros Cingulata. Tradicionalmente se reconocen seis géneros de Pampatheriidae (sensu SCILLATO-YANÉ, 1980, 1982; EDMUND, 1996; EDMUND & THEODOR, 1997; SCILLATOYANÉ et al., 2005; GÓIS et al., 2012b), considerados válidos por el autor de esta Tesis: Scirrotherium, Kraglievichia, Vassallia, Plaina, Pampatherium y Holmesina. Asimismo, se agregan dos nuevos géneros: Gen. nov. A y Gen. nov. B. Previamente a esta contribución, se reconocían 15 especies, de las cuales 14 resultan válidas. Estas son: 1) Scirrotherium hondaense Edmund, 1997; 2) Kraglievichia paranensis (Ameghino, 1883); 3) Vassallia minuta (Moreno & Mercerat, 1891) 4) Plaina intermedia (Ameghino, 1888); 5) Pl. subintermedia (Rovereto, 1914); 6) Pl. brocherense Castellanos, 1956; 7) Pampatherium humboldtii (Lund, 1839); 8) P. typum Gervais & Ameghino, 1880; 9) P. mexicanum Edmund, 1996; 10) Holmesina major (Lund, 1842); 11) H. septentrionalis (Leidy, 1889); 12) H. occidentalis (Hoffstetter, 1952); 13) H. floridana (Robertson, 1976); y 14) H. paulacoutoi (Cartelle & Bohórquez, 1985). En esta Tesis no se considera válida la especie V. maxima Castellanos, 1946, que pasa ser un sinónimo junior de Pl. intermedia. Además, en el curso del trabajo de Tesis, se reconocieron cuatro nuevas especies (una de las cuales ya fue publicada), a saber: 1) S. carinatum Góis, Scillato-Yané, Carlini & Guilherme (GÓIS et al., en prensa); 2) H. rondoniensis Góis, Scillato-Yané, Carlini & Ubilla (GÓIS et al., 2012b); 3) Gen. nov., sp. nov. A Góis, González Ruiz, Ciancio & Scillato-Yané (en desarrollo); y 4) Gen. nov., sp. nov. B Góis, Scillato-Yané, Ciancio, Gónzalez Ruiz & Soibelzon (en desarrollo). Cabe resaltar que de las cuatro especies nuevas mencionadas anteriormente, dos corresponden a géneros nuevos. De ellos, el Gen. nov., sp. nov. A es hasta el momento el taxón más antiguo conocido para la familia; previamente, los registros más antiguos eran los del Mioceno medio de La Venta, Colombia. El Gen. nov., sp. nov. B es el tercer género que se reconoce para el Pleistoceno de Argentina. Estos dos nuevos géneros y especies, así como las otras dos especies nuevas, incrementan la diversidad de los Pampatheriidae de América del Sur. Probablemente, Argentina fue uno de los centros de diversificación para el citado grupo. Otro resultado obtenido en esta Tesis Doctoral fue la identificación de tres nuevos registros para la Argentina de especies ya conocidas en otros ámbitos, a saber: P. humboldtii, para la provincia de Buenos Aires, y H. majus y H. occidentalis para la provincia de Santa Fe. Dado que la gran mayoría de las especies estaban pobremente descriptas, se procedió a un nuevo tratamiento de ellas: nuevas diagnosis, descripciones e ilustraciones. Como consecuencia, se mejoró notablemente la caracterización morfológica de los géneros y especies que se reconocieron como válidos. Esta situación lleva a cuestionar la validez de algunas especies nominales presentes en la bibliografía. Con los estudios que se están llevando a cabo se ha logrado avanzar en el conocimiento de la anatomía de este grupo. De tal manera, se corrigen, completan y renuevan las descripciones para estos taxones, estudiando la totalidad accesible de los caracteres anatómicos: coraza con sus osteodermos, cráneo, dentición y postcráneo. En síntesis, se ha logrado un conocimiento actualizado de la nomenclatura, anatomía y sistemática de las especies de Pampatheriidae. Uno de los resultados más importantes de esta Tesis, fue el desarrollo de una nueva nomenclatura estándar de los osteodermos para la familia. En tal sentido, los nombres de las estructuras empleados antiguamente resultaban erróneos y confusos desde una perspectiva conceptual. El análisis filogenético realizado evidencia que los Pampatheriidae son un grupo monofilético y están relacionados con los Glyptodontidae, conformando el clado Glyptodontoidea. Además, se confirma la monofilia de Dasypodidae, siendo Peltephilus el grupo hermano de Dasypodidae, ubicándose Peltephilus y Dasypodidae en un clado mayor, los Dasypodoidea. En este análisis se obtiene como otro resultado interesante la exclusión de Proeutatus y Eutatus como el grupo hermano de los GlyptodontoideaLa familia Pampatheriidae queda definida por siete sinapormofías, de las cuales cinco son exclusivas. Vassallia, Plaina y Pampatherium comparten una única sinapomorfía: la elevación central longitudinal de los osteodermos es aplanada. En esta Tesis, se confirman algunas propuestas previas, i.e. la relación Vassallia–Pampatherium (SCILLATO-YANÉ, 1980; SCILLATO-YANÉ et al., 2005), pero con una nueva interpretación: Vassallia es el grupo hermano del clado Plaina–Pampatherium. El clado Pampatherium se encuentra bien soportado, dado que cuenta con seis sinapomorfías. El clado Kraglievichia–Holmesina (excepto H. floridanus) se sostiene a base de tres sinapomorfías. La ubicación de H. floridanus en este clado determina que el género Holmesina sea polifilético.
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The hyoid apparatus reflects aspects of the form and function of feeding in living and extinct organisms and, despite the availability of information about this structure for Xenarthra, it remains little explored from an evolutionary perspective. Here we compare the morphology of the hyoid apparatus in xenarthrans, describing its general morphology and variation in each major clade and score these variations as phylogenetic characters, which were submitted to ancestral states reconstructions. The general hyoid morphology of Xenarthra consists of a v-bone (basihyal fused with the thyrohyals) and three paired bones (stylohyal, epihyal and ceratohyal), which are unfused in the majority of taxa. The clade-specific morphology observed here, allowed us to obtain additional synapomorphies for all major clades of Xenarthra (Cingulata, Pilosa, Folivora and Vermilingua), for Glyptodontididae, and for Nothrotheriidae. The fusion of hyoid elements are convergentelly achieved among the diphyletic extant tree sloths, some extinct ground sloths and glyptodontids. Despite the heavy influence of adaptive evolution related to feeding habits, the morphology of the hyoid apparatus proved to be a valuable source of phylogenetic information.