Anais da Academia Brasileira de Ciências
ISSN: 0001-3765
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Academia Brasileira de Ciências Brasil
Ferreira, José D.; Zamorano, Martín; Ribeiro, Ana Maria
On the fossil Remains of Panochthus Burmeister, 1866 (Xenarthra, Cingulata, Glyptodontidae) from the Pleistocene of southern Brazil
Anais da Academia Brasileira de Ciências, vol. 87, núm. 1, marzo, 2015, pp. 15-27 Academia Brasileira de Ciências
Rio de Janeiro, Brasil
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Panochthus
8
€ •‚ ƒ„ …† †‡ ˆ1 ‰ˆ Š‹Œ •ˆ‰ ˆŒ 2 ˆŒˆ ‰ˆ ‡ˆ ‡ †‡ 3
1 Pro rama de P s- rad a o em eociências Instit to de eociências niversidade ederal do io rande do S l Av. Bento on alves 9500 A ronomia 91501-970 Porto Ale re S Brasil
2Consejo Nacional de Investigaciones Cientí cas y Técnicas/CONICET, División de Paleontología de Vertebrados, Facultad de Ciencias Naturales y Museo, niversidad Nacional de a Plata Paseo del Bos e s/n B1900 A a Plata Ar entina
3Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Rua Dr. Salvador França, 1427, Jardim Botânico, 90690-000 Porto Alegre, RS, Brasil
an scri recei ed n an ar acce ed r ica i n n A ri ˆ •Š ˆ Š
The genus an c s represents the last lineage of “Panochthini” recorded in the Pleistocene. This genus has a wide latit dinal distrib tion in So th America and in Brazil it occ rs in the so thern and northeastern re ions. In this a er we describe new material (isolated osteoderms and ca dal t be fra ments) assi ned to an c s from the state of io rande do S l (so thern Brazil) and disc ss some ta onomic iss es related to an c s erc a s and an c s res e ini based on this material . The occurrence of res e ini is the rst for outside the Brazilian Intertropical Region. In addition, we describe new dia nostic feat res to differentiate the osteoderms of res e ini and erc a s. Unfortunately, it was not possible to identify some osteoderms at the species level. Interestingly, they showed four distinct morphotypes characterized by their external morphology, and thus were attributed to an c s sp. Lastly, we concl de that in addition to erc a s re istered to so thern Brazil there is another s ecies of the en s assi nable to . cf. res e ini. Our analysis reinforce the reliability of caudal tube characters for the classi cation of species of an c s
Ž • : Glyptodontidae, an c s Pleistocene so thern Brazil osteoderms.
Correspondence to: José Darival Ferreira E-mail: darival.fds@gmail.com
‡ŒŠ ƒ• Š‡ Œ
The tribe “Panochthini” is a taxonomically diverse group of glyptodonts restricted to South America. Its fossil record spans the late Miocene to the late Pleistocene (Zamorano 2012). The genera
an c s B rmeister 1866 ac s Ame hino
1888 and r an c s Castellanos 1925 were
traditionally included within this tribe (Castellanos
1942, Hoffstetter 1958, Paula Couto 1979, McKenna and Bell 1997, Zurita et al. 2011, Zamorano 2012); however, in recent cladistic analyses they do not form a natural group (Zamorano and Brandoni 2013, see. Zamorano et al 2013).
This genus has a wide latitudinal distribution in South America (Tonni and Scillato-Yané 1997), incl din both so thern and northeastern re ions ( i . 1) (Por ino and Ber vist 2002 Por ino et al. 200 Ubilla et al. 2004, Zurita et al. 2009a, Zamorano 2012).
JOSÉ D. FERREIRA, MARTÍN ZAMORANO AND ANA MARIA RIBEIRO
The main characteristics of the genus
an c s are the osteoderms of the cara ace
which have a retic lar attern on the e ternal surface, with small polygonal gures that are at and e ivalent in size and the ca dal t be which has a similar ornamentation attern to the cara ace (Castellanos 19 2). Accordin to a recent review of
an c s by Zamorano (2012), six species were
reco nized: s in er edi s Castellanos 1937
in er edi s Lydekker, 1895, erc a s
(Owen, 1845), ren e ian s Ame hino 1889
a ari ensis (Moreira, 1965) and res e ini
Castellanos 19 2.
The rst record of an c s, for the territory
of Brazil was mentioned by Ihering (1891), in corres ondence to lorentino Ame hino from the
coastal lain of the state of io rande do S l Santa Vitória do Palmar Municipality. Subsequently,
an c s was re orted from de osits in northeast
Brazil (tanks) by Branner (1915) and by several a thors (e. Moreira 1971, Bergqvist 1993). The fossil content of these tanks was deposited d rin the late Pleistocene with available electron spin resonance (ESR) ages dating it to between 63,000 to 10,000 years BP (e.g. Kinoshita et al 2005, Oliveira et al. 2009, Silva 2009, Dantas et al. 2011). In the northeastern re ion of Brazil
res e ini and a ari ensis are considered
endemic s ecies (e. . Ber vist 1993 Por ino and Bergqvist 2002, Zamorano 2012, but see Chimento and A nolin 2011). erc a s has only been
recorded in io rande do S l (e. Bombin 1976 ibeiro and Scherer 2009). Another d bio s record of an c s in the Amazon re ion is referred
to by Paula Couto (1956); the material is a single isolated osteoderm collected in the r a iver in the state of Acre.
In this a er we describe new material of the
an c s from the state of io rande do S l
(so thern Brazil) and disc ss some ta onomic as ects of the s ecies re orted.
The material studied here belongs to the paleontological collections of the Museu de Ciências Naturais da Fundação Zoobotânica do Rio Grande do Sul (MCN/FZBRS), Museu de Ciências Tancredo Filho Melo (MCTFM), aborat rio de eolo ia e Paleontolo ia da niversidade ederal de io rande ( P/ FURG) and Museu Nacional do Rio de Janeiro (MNRJ), Brazil. The anatomical nomenclature follows Por ino and Ber vist (2002) while the scheme for the different re ions of the cara ace is based on; systematics follow Zamorano et al (2013) (see. Zamorano and Brandoni 2013). The description and terminology for osteoderms follows ill (2006).
- Map of the geographic distribution of an c s in the Pleistocene.
The material was found in the municipalities of Uruguaiana (Touro Passo Creek), Santa Vitória do Palmar (Balne rio ermene ildo coastal lain and Chui Creek) and Rosário do Sul (Rincão dos
ialho) ( i . 2).
Touro Passo Creek (29°40′S, 56°51′W) is 13 km the north of Uruguaiana Municipality (Da-Rosa 2003). es ite this the bio eo ra hic correlation is disc ssed since it shares fa nal elements with the Sopas Formation (Uruguay), which are not recorded in B enos Aires rovince Ar entina (see Ubilla 1985, Oliveira 1996, Ubilla and Perea 1999). According to Milder (2000), the age obtained by thermol minescence datin encom asses a time s an from 42,600 to 6,400 years BP (Kerber et al 2011).
Balneário Hermenegildo (53°15′S, 33°42′W) com rises the so thern ortion of the coastal lain of Rio Grande do Sul, 20 km from Santa Vitória do Palmar. Over time, it has suffered modi cations to its landscape related to sea level uctuations (trans ressive-re ressive events) which develo ed four lagoon-barrier systems (Villwock and Tomazelli 1995). The fossil remains of the coastal lain are associated with de osits of la oon-barrier system III, with an estimated age of 120,000 years BP (Villwock and Tomazelli 1995). The dates obtained for the coastal lain of io rande do S l show a wide variation from younger than 18,000 to 650,000 years BP (Lopes et al 2008 2010). Accordin to o es et al (2010) the mi t re of fossils from the middle and late Pleistocene is probably the result of reworking of several fossil beds by successive Quaternary transgressive events.
Chuí Creek (33°35′S; 53°20′W) is located in so thernmost art of io rande do S l in Santa Vitória do Palmar Municipality. The material was fo nd in si exposed along the banks of Chuí Creek. The plain through which the creek ows and where the fossil remains of Chuí Creek occur are associated with deposits of lagoon-barrier system
III located between coastal barrier II (westwards) and barrier III (eastwards). The age of the fossils of Chuí Creek was estimated to be at least 120,000 years BP based on their location within the barrier-lagoon system III (Lopes et al. 2005). However, subsequently it was demonstrated that these fossils are more recent ( o es et al. 2010). A sam le from the bank of Chuí Creek dated by ESR suggests an age between 42,000 and 33,000 years BP (Lopes et al. 2010). The mammal fossils collected are assigned to the Lujanian age (Oliveira et al. 2005).
inc o dos ialho is located in os rio do S l Municipality (30°12′S; 55°16′W). It is situated in the southwestern state of Rio Grande do Sul. The material was collected near the ialho farmho se in a stream that cuts through layers of sandy sediments of variable thickness, of Pleistocenic age, that are directly in contact with the Triassic (Ferigolo et al. 1997).
S erorder enarthra Co e 1889 Order Cingulata Illiger, 1811
Suborder Glyptodontia Ameghino, 1889 Superfamily Glyptodontoidea Gray, 1869 Family Glyptodontidae Gray, 1869
en s an c s B rmeister 1866 an c s erc a s (Owen, 1845)
(Fig. 3A; Fig 4C-D)
EFERRED MATERIAL
Ca dal t be ri ht distal ortion P P0212. Isolated osteoderms, MCN-PV 3948; MCN-PV 3953.
EOGRAPHIC PROVENANCE
Balne rio ermene ildo coastal lain of io rande do S l.
ESCRIPTION
The caudal tube is a right distal tip belonging to an c s erc a s. The distal portion
8 JOSÉ D. FERREIRA, MARTÍN ZAMORANO AND ANA MARIA RIBEIRO
- Location map of fossiliferous localities. A. Panoramic view of banks exposing the TouroPasso Formation and stratigraphic sequences (modi ed from Bombin 1976); B. BalneárioHermenegildo and transect of the coastal plain of Rio Grande do Sul, showing its main depositional systems (modi ed from Tomazelli and Villwock 2005); C. Panoramic view of banks exposed at Chui Creek and stratigraphic sequences (modi ed from Lopes 2013); D. Rincão dos Fialho, where the material of an c s s was collected.
erc a s and different from res e ini which
is heavily truncated, and s in er edi s which is ointed and s btrian lar. In dorsal view it is possible to distinguish two subapical gures, as in
ren e ian s a ari ensis and erc a s. In res e ini and s in er edi s there is st one
dorsal gure. In dorsal view, the lateral gure can be seen to be of relatively large size as in ren e ian s and distinct from a ari ensis s in er edi s and erc a s in which it is smaller. The terminal
gure is oriented laterally as in ren e ian s and
erc a s while in a ari ensis and res e ini it is oriented ventrally. The terminal gure
is surrounded by a shallow slope, which differs from
s in er edi s and res e ini. Altho h the
caudal tube is fragmented, it is possible to identify similar gures on the carapace. The morphology of the ca dal t be are the most variable in this s ecies (Zamorano et al. 2012).
The osteoderms of the postero-dorsal region of erc a s are thick and their tubercular
gures are larger than in any other species of
an c s ( i . 3A).
erc a s is resent in the middle Pleistocene
and late Pleistocene of Argentina, Uruguay, southern Brazil, Paraguay and Bolivia (Hoffstetter 1963, 1978, Mones and Francis 1973, Zurita et al. 2009b, Zamorano 2012, Zamorano et al. 2012).
an c s cf. res e ini
(Fig 3B; Fig 4A-B)
EFERRED MATERIAL
Caudal tube, distal right portion, MCN-PV 32182. Isolated osteoderms: MCN-PV 2594; MCN-PV 3296; PV 5324; PV 7131; MCN-PV 8766; MCN-MCN-PV 8804; MNRJ 2106-V; MNRJ 2107-V; MNRJ 2138-V; MNRJ 2152-V; MNRJ 2155-V; MNRJ 3537. EOGRAPHIC PROVENANCE
Balne rio ermene ildo coastal lain of io rande do S l.
ESCRIPTION
The caudal tube, MCN-PV32182, shows an apexian gure, considered an apomorphy of an c s res e ini, visible only on the dorsal surface and
of subcircular outline (Fig. 4A-A’); however, the apexian gure of res e ini (DGM 1M) is less deep than in the specimen MCN-PV 32182. In this specimen, it is possible to identify the gure only in posterior view; in anterior view, it is poorly reserved and it is not ossible to describe it in more acc rate details.
erreira et al. (2013) observed that osteoderms of the lateral re ion of the cara ace resent distinctive feat res to res e ini: they are
thinner and have smaller t bercles which in t rn show a reater distance between the radial s lci (Fig. 3B); the tubercular gures are small and have a at surface, with a diameter of approximately 2-7 mm. Some osteoderms from the ostero-dorsal region show a slight concavity in the center of the plane gures, resembling in this respect
s in er edi s from the early Pleistocene of
Argentina Ensenadan age.
res e ini is recorded in the Pleistocene
of the northeast of Brazil and is considered an endemic s ecies of the Brazilian intretro ical re ion (Ber vist 1993 Por ino and Ber vist 2002).
an c s s .
(Fig. 4E-F; Fig 5; Fig 6)
- A. an c s erc a s. B. an c s cf. res e ini Scale bars: 10 mm.
OS . I A A N A O ANO AN ANA A IA IB I O
- Caudal tubes. A, A′ – B, B′. an c s cf. res e ini A–A′, View ventral; B–B′, View lateral; C, C′ – D, D′. erc a s. C–C′, View ventral; D–D′, View lateral. E, E′ – F, F′. an c s sp. E – E′, View dorsal; F – F′, View lateral. Abbreviations: a. apical gure; ap. apexian gure; m. marginal gure; t. terminal gure; l. lateral gure; v. ventral gure; d. dorsal gure. Scale bars: 100 mm.
EFERRED MATERIAL
Distal left portion of caudal tube (MCN-PV 2960); isolated osteoderms (MCN-PV3977; MCN-PV 3979; MCN-PV 3980; MCN-PV 3981; MCN- PV 3982; MCN-PV 3987; MCN-PV 3958; MCN-PV 3966; MCN-PV 3965; MCN-PV 3964; MCN-PV 3963; MCN-PV 3954; MCN-PV 3960; MCN-PV3971; MCN-PV 3975; MCN-PV 3985; MCN-PV6897; MCN-PV 6321; MCN-PV 4139; MCN-PV 4617; MCN-PV6326; MCN-PV4616; MCN-PV 4136; MCN-PV 4108; MCN-PV 4098; MCN-PV 4099; MCN-PV 4155; MCN-PV 6967; MCN-PV 8772; MCN-PV 8780; MCN-PV 8508; MCN-PV 7260; MCN-PV 7107; MCN-PV 7118; MCN-PV 1150; MCN-PV 3949; MCN-PV 3957; MCN-PV 3976; MCN-PV 6324; MCN-PV 6325; MCN-PV 6345; MCN-PV 6322; MCN-PV 6323; MCN-PV 6319; MCN-PV 6639; MCN-PV 6657; MCN-PV 6842; MCN-PV 6843; MCN-PV 6859; MCN-PV 5350; MCN-PV 5396; MCN-PV 5420; MCN-PV 5345; MCN-PV 5360; MCN-PV 5441; MCN-PV 5442; MCN-PV 5924; MCN-PV 5394; MCN-PV 5325; MCN-PV 5395; MCN-PV 5923; MCN-PV 5443; MCN-PV 5433; MCN-PV 5700; MCN-PV 5421; MCN-PV 5464; MCN-PV 5341; MCN-PV 5327; MCN-PV 5326; MCN-PV 5445; MCN-PV 5589; MCN-PV 3961; MCN-PV 3951; MCN-PV 3955; MCN-PV 3959; MCN-PV 7112; MCN-PV 6320; MCN-PV 5444; MCN-PV 5551; MCN-PV 4988; MCN-PV 3974; MCN-PV 4001; MCN-PV 4002; MCN-PV 4033; MCN-PV 4738; MCN-PV 4739; MCN-PV 4740; MCN-PV 4741; MCN-PV 4742; MCN-PV 4743; MCN-PV 4744; MCN-PV 4745; MCN-PV 4746; MCN-PV 4747; MCN-PV 8800; MCN-PV 8803; MCN-PV 8805; MCN-PV 2960; MCN-PV 448; MCN-PV 461; MCN-PV 145; MCN-PV 210; MCN-PV 134;MCN-PV 1681; MCN-PV 2043; MCTFM-PV 859; MCTFM-MCTFM-PV 850; MCTFM-MCTFM-PV 117); articulated osteoderms of dorsal carapace (MCN-P 5659).
EOGRAPHIC PROVENANCE
Balne rio ermene ildo coastal lain of io Grande do Sul; Rincão dos Fialho and Chuí Creek.
ESCRIPTION
The caudal tube (Fig 4 E-F) has a distal, semi-oval extremity and has a subtriangular shape, endin in a conical ti . In dorsal view the ca dal tube presents only one dorsal gure, shared only by res e ini and s in er edi s, and lacks a secondary dorsal gure, which differs from
ren e ian s a ari ensis and erc a s.
The terminal gure is oriented laterally as in
ren e ian s and erc a s. Therefore, due to
the fragmentary condition of the specimen MCN-P 2960 and the absence of dia nostic feat res its speci c assignment is not possible.
All the carapace osteoderms analyzed are enta onal he a onal rectan lar or s b adran lar form, with thicknesses ranging from17.2 mm (MCN-PV 2016) to 42.8 mm (MCN-(MCN-PV 4139), and show a tendency for merging between the osteoderms, especially those of the lateral side of the carapace. The osteoderms show the general ornamentation attern of the dorsal re ion of the cara ace of
an c s, characterized by the presence of
multiple polygons on the surface; these polygons are unde ned and do not possess the formation of a distinct central gure. In a few osteoderms there is a distinct central gure, typical of the lateral ed es of erc a s and ren e ian s
(Zamorano 2012). In the south of Brazil, most of the records of glyptodonts are isolated osteoderms, which weakens the establishment of species identi cations. Morphological differences that enable us to classify these isolated osteoderms into four main morphotypes were noted (Fig 5).
Morphotype I (MCN-PV 2043, Fig 5A). The contact area between osteoderms has a ro h aspect; in the external view the osteoderm presents several subcircular and concave gures, which are separated from each other by shallow radial sulci
OS . I A A N A O ANO AN ANA A IA IB I O
and limited by shallow radial sulci with foramina in the connection between them. The morphotype I osteoderms are larger than the other morphotypes identi ed and were found only in the Chuí Creek locality (Fig 6).
Morphotype II (MCN-PV 5659, Fig 5B). The contact area between osteoderms does not have a rough aspect as in morphotype I; in external view the osteoderms present several circular gures, small, prominent and trabecular in aspect. The gures are limited by wide and shallow radial sulci with relatively large foramina. The morphotype II osteoderms are smaller than the morphotypes I and I and were fo nd in the os rio do S l inc o dos Fialho locality and Santa Vitória do Palmar, Balneário Hermenegildo locality (Fig. 6).
Morphotype III (MCN-PV 4988, Fig. 5C). This morphotype is distinguished by deep radial sulci and by clearly polygonal gures (pentagonal and hexagonal), at and without any trabecular aspect.
- Morphotypes of the osteoderms of an c s sp. and detail. A–A′, Morphotype I; B–B′, Morphotype II; C–C′, Morphotype III; D–D′, Morphotype IV. Scale bars: 10 mm.
This morphology suggests that these osteoderms were fo nd in a more lateral re ion of the cara ace. The morphotype III osteoderms were found in Balneário Hermenegildo and Touro Passo Creek.
Morphotype IV (MCTFM-PV 117, Fig 5D). This type presents a spongy aspect, making it ossible to differentiate small fi res on the surface of the osteoderm. The morphotype IV osteoderms differ from the other morphotypes by their greater thickness; they are quite similar to e r r s ra ec a s Zurita and Ferrero 2009 ( AP 1510 i 2 ) in e ternal view b t in lateral view ra erc a s resents
layers of compact bone that are much thicker than in an c s with a smaller ma im m
diameter and in internal view fewer foramina are observed (ei ht to ten in ra ec a s and three in the Brazilian material, MCTFM-PV 117). The morphotype IV osteoderms were found in Balneário Hermenegildo and Chui Creek.
The morphotypes present signi cant mor holo ical differences which co ld indicate that the osteoderms are from different areas of the carapace. Chuí Creek and Balneário Hermenegildo are the localities with the most mor holo ical diversity of osteoderms. However, it is interesting to note that the morphotype I osteoderms were found only in Chuí Creek. At present, it is safer to assi n these s ecimens to an c s s .
Thus far, the previous records of an c s erc a s in io rande do S l (Pa la Co to
1943, Bombin 1976) are dubious. Oliveira (1996) in a study on the Xenartha of Rio Grande do Sul, analyzed the material assigned to erc a s
and considered it as an c s s . beca se
the specimens were isolated and insuf ciently reserved osteoderms which were not eno h to differentiate the species. In addition, Kerber and
Oliveira (2008) revised the fauna of mammals from Touro Passo based on new materials, and proposed the resence of an c s sp. previously reported
to the locality by Bombin (1976) as erc a s.
Herein, we identify erc a s and cf. res e ini to the Balne rio ermene ildo while
for the other locations studied (Touro Passo Creek, Chuí Creek and Rincão dos alhos ) assigning
an c s s .
The caudal tubes, although very fragmented, are more informative than osteoderms, and they were cautiously used to identify an c s erc a s and cf. res e ini. Unfortunately,
the ca dal t bes are derived from the continental shelf, so they do not possess a stratigraphic context. Thus, a better identi cation of these species is still dependent on new ndings from the continental Quaternary beds.
All other osteoderms des ite havin different sha es ( adran lar enta onal and he a onal)
OS . I A A N A O ANO AN ANA A IA IB I O
show the same retic lar attern characteristic of the genus. The depth of the radial sulci may vary considerably in an c s accordin to
its localition on the cara ace so it cannot be sed as a diagnostic feature. Most of the osteoderms found could not be assigned to a particular species; however some individ al were ossible to be identi ed at level. An example of this are the osteoderms of ren e ian s located on the ca dal ed e of the cara ace which bear an o enin that narrows down and closes with a ro nded almost cylindrical cross-section (Zamorano 2012). It has also been fo nd that the osteoderms of the ostero-dorsal re ion of erc a s and
lateral osteoderms of res e ini can be sed to
distin ish these s ecies (see i 3).
The species of an c s show the e ternal
s rface of osteoderms i similar mor holo ical pattern comprising small tubercles. Particularly, in in er edi s (antero-dorsal and ostero-dorsal re ions) and a ari ensis (dorsal re ion
following Moreira 1971) the carapace shows the typical rosette pattern (Zurita et al 2011); however, Costa Pereira et al. (201 ) inter ret this fra ment of a ari ensis as havin similarities with the
ce halic shield of other s ecies of an c s.
The pattern differs from that observed on the dorsal re ions of the cara ace of ren e ian s
erc a s and res e ini in which a clear
reticular pattern is observed (Zamorano 2012). Por ino and Ber vist (2002) oint o t that the cara ace of a ari ens is less thick than in res e ini and that this feat re re resents another
distin ishin characteristic amon the ta a. Costa Pereira et al. (201 ) indicate the need detailed revision of a ari ensis.
There are some diagnostic features at least for some s ecies of an c s ( resence of
main gures in osteoderms from the antero and osterodorsal re ions of cara ace in in er edi s and presence of an apexian gure in res e ini for instance). rthermore some a thors (e. . Cr z et
al. 2011) have ar ed that the s ecies of an c s
can be differentiated by unique combinations of cara ace and ca dal t be characteristics.
Moreira (1971) observed a wide variation between s ecimens of res e ini inter reted by him as ontogenetic. Chimento and Agnolin (2011) described a iece fra ment of ca dal t be from Santiago del Estero province, Argentina, this fragment described and gured by them, do not present evidence apexian gure, but a detailed note on the gure provided by these authors in view ventral is observed two ventral gure, which do not corres ond to res e ini. It is noteworthy that that
the fragment described here may actually represent the rst occurrence of res e ini o tside the Brazilian Intertropical Region. The presence of the s ecies s in er edi s (based in MCN-PV 2960) may not be ruled out either, although it is a taxon typical of the early-middle Pleistocene, while res e ini and a ari ensis are from the Pleistocene sens a and considered endemic to the northeast of Brazil (Ber vist 1993 Por ino and Ber vist 2002).
In Brazil the fossil record of an c s is
restricted to the so thern and northeast of Brazil (see Fig 1); the fossils recorded between these regions are mainly in cave of karstic origin (see. Lund 1839, Salles et al. 2006 Castro and an er 2011 hilardi et al. 2011, Silva et al. 2012). The glyptodonts previously reported for this type of depositional environment are d n and r s. The
absence of an c s in these localities may be due
to the lack of further study in these regions, where the Pleistocene deposits are still poorly known.
Zurita et al. (2005) suggested that the strong development of frontonasal sinuses and a strongly pneumatized skull in e sc er ca s and an c s played a major rule in thermoregulation
and wo ld re resent an ada tation for savanna-like environments in a semiarid climate period. Carlini et al. (200 ) s est based on the fa na of Mesopotamia Argentine, southern Brazil
and Uruguay West, would be associated an environmental conditions wetter and warm.
During the Quaternary, there were several glacial cycles, with cold and dry periods interrupted by hot and wetter periods (Haberle and Maslin 1999). Multiple pulses of expansion/contraction of the elds and re ression/trans ression of sea levels were recorded. According to Scillato-Yane et al. (2002) during mainly the last inter lacial eriod there was develo ment of an ecological corridor connecting the Mesopotamia re ion of Ar entine with the intertro ical re ion of Brazil. S nchez et al. (200 ) ro ose a corridor alon the east of So th America and some coastal areas of the Atlantic which was formed d rin re ression of sea level, and was used by mammals adapted to mesic environments. These pulses can justify the concurrent resence of enera from intertro ical and am ean re ions thro h the Pleistocene for e am le:
an c s a a eri ervais and Ame hino 1880 esina Sim son 1930 d n Owen,
1839 and i as d n Cabrera 1929 in so thern
Brazil mainly in the plain coast of Rio Grande do Sul. The interesting thing is the simultaneous presence of purportedly endemic intertropical species and am aean s ecies belon in to those enera in the same area (see Oliveira and Pereira 2009).
The authors thank the Coordenação de A erfei oamento de Pessoal de N vel S erior (CAPES) for nancial support to J.D.F. as a fellowship of the Pro rama de P s- rad a o em eociências/ Universidade Federal do Rio Grande do Sul; FZBRS and LGP-V FURG for the infrastructure provided; D.da Silva, L.Kerber,V.G. Pitana (in e ria )
E.V.Oliveira, K.O.Porpino and A.E.Zurita for reading and commenting on the rst version of the manuscript; we are greatly indebted to D.D.Rego (MNRJ), R.P.Lopes (LGP/FURG), R.R.Machado (DNPM) and J. Pereira (MCTFM) for enabling access to the specimens studied; and to L. Rota for enabling access to com arative material nder his care.
O gênero an c s re resenta a ltima linha em de "Panochthini" registrado no Pleistoceno. Este gênero tem uma ampla distribuição latitudinal na América do Sul, e no Brasil ocorre nas re i es s l e nordeste. No resente trabalho se descreve novos materiais (osteodermos isolados e fra mentos de t bo ca dal) de an c s do estado de io rande do S l (S l do Brasil) e se disc te al mas est es ta on micas relatada ara A ocorrência de res e ini é o primeiro para fora da Região Intertropical Brasileira. Além disso, nós descrevemos novas caracter sticas ara diferenciar osteodermos de res e ini e erc a s. Infelizmente n o foi possível identi car alguns osteodermos em nível de espécie. Curiosamente, eles mostraram quatro morfotipos, caracterizados or s a morfolo ia e terna e atrib dos a an c s s . inalmente n s concl mos e ao lado de erc a s re istrados ara o s l do Brasil h o tras espécies do gênero designada a . cf. res e ini. Nossas analises reforça a con abilidade das características do tubo caudal para a classi cação das espécies de an c s
Glyptodontidae, Panochthus, Pleistoceno, S l do Brasil osteodermos.
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