A FAUNA PLEISTOCENICA DO PIAUi (NORDESTE
DO BRASIL): RELAGOES PALEOECOLOGICAS
E BIOCRONOLOGICAS
THE PLEISTOCENE FAUNA OF PIAUi (NORTHEASTERN
BRAZIL): PALAEOECOLOGICAL AND
BIOCHRONOLOGICAL IMPLICATIONS
Claude Guérin’ , Maria Amélia Curvello”, Martine Faure“, Marguerite Hugueney*
& Cécile Mourer - Chauviré"
‘Centre de Paléontobgie Stratgraphique et Paléoécologie associé au CNRS ( URA 11), Centre des Sciences de la
Terre, Université Claude Bernard - Lyon |, 27-43 Boulevard cu 11 Novembre, 69622, Villeurbanne Cédex, France.
™ Departamento de Geocéncias, Universidade Federal Rural do
Rio de Janero, 23851 Seropédica, Rio de Janeto, Brasil
“Université Lumiere Lyon 2, 7 rue Rauin, 69007 Lyon et URA 11 du CNRS, France, FUMDHAMENTOS
More than fifty species of mammals belonging
to the genera Didelphis, Monodelphis, Marmosa,
Catonyx, Scelidodon, Eremotherium, Dasypus,
Euphractus, _ Tolypeutes,Pampatherium,
Hoplophorus, Panochtus, Glyptodon, Pteronotus,
Tonatia, Phyllostomus, Artibeus, Desmodus,
Myolis. Molossus, Tadarida (Rhizomops),
Holochilus, Pseudoryzomys, Akodon, Calomys,
Oryzomys, Kerodon, Galea, Thricomys, Agout,
Protocyon, Dusicyon, Speothos, Conepatus,
Aretodus, Felis, Panthera, Smilodon,
Macrauchenia, Toxodon, Haplomastodon,
Hippidion , Equus, Dicolyles, Tayassu, Palaeolama,
Mazama, "Cervus" sp.,more than 30 bird species,
Caiman, ‘Dracaena, several turtles, amphibians,
and the fish Plecostomus were collected in the
archaeological area of S40 Raimundo Nonato
(Southeast Piaui, Brazi). This collection was made
in the excavations of three archaeological sites,
Tocada Janola da Barra do Antonidio, Tos de Cima
dos Pilo and Toca do Garrincho; in the first two,
the Paleolthicartifacts are essociated to the feune,
in the Garrincho, human remains were found
associated to the fauna. Test-pits were mage in
the Toca do Serrote do Artur and in various lakes,
specially Sao Vitor and the Lagoa da Pomba.
Finally, some fossil remains were found in the
course of an exploration in the Sumidouro do
Sansdo.
The assemblage of mammal fauna dated from
the Upper Pleistocene is the witness to the
sxistence of a landscape characterized by bush
savanna, crossed by forest zones in aclimate which
was much more humid than the modern one. This
faunas contemporary of one of he oldest currently
known vestiges of the presence of man in the
Americas.
1-Generalitios
Since 1970 the research in Southeast Piaui
(Northeast Brazil, fig. 1) by N. Guidon and her
team has enables the inventory of approximately
400 archaeological sites in the region of Sao
Raimundo Nonato (Amaud et alii, 1984; Guidon,
1986; Guidon ef ali, 1989). Over two thirds of
these sites are known for their rock art (Guidon,
1984a, Possis, 1987); a great number of them
are located in the National Park of the Serra da
Capivara, its boundary being the sendstone
cuesta of the lower Paleozoic; approximately ten
of these sites were excavated, or important test-
pits took place in them. Three of these sites in
the sandstone formation posses Paleolithic
levels: Toca do Caldeirdo do Rodrigues |, dated
from 18,600 600 years BP (Guidon, 1981; 1984
b and c), Toca do Sitio do Meio currently being
excavated , in which the oldest levels were dated
betwoon 12,200 + 600 and 14,300 + 400 BP
56
Mais de cinquenta espécies de mamiferos
pertencentos aos génoros Didelphis, Monodelphis,
Marmosa, Catonyx, Scelidodon, Eremotherium,
Dasypus, Euphractus, | Tolypeutes,
PampatheriumHoplophorus, Panochthus,
Giyptodon, Pteronotus, Tonatla, Phyllostomus,
Artibeus, Desmodus, Myatis, Motossus, Tadarida
(Rhizomops),Holochitus, Pseudoryzomys, Akodon,
Caiomys, Oryzomys, Kerodon, Galea, Thricomys,
Agoutl, Protocyon, Dusicyon, Speothos,
Conopatus, Arctodus, Felis, Panthora, Smilodon,
Macrauchenia, Toxodon, Haplomastedon,
Hippidion, Equus, Dicotyles, Tayossu, Palaoclama,
Mazama, "Cervus" sp., mais de 30 especies de
aves, Caiman, Draceerna, numerosas tartarugas,
anfibios e 0 peixe Plecastmus foram recohidos
nna tea arqueobgica de S20 Raimundo Nonato
(Gudeste do Piaul, Brasil). Esta coleta foi realzada
nas escavacdes de Irés sitios arqueolécicos, Toca
dda Janela da Barra do Antoniao, Toca de Cima dos
Piléo e Toca do Garrincho; nos dois primeiros,
artefates paleolitcos ostao ascociados & fauna, no
Garrincho, restos humanos foram achados
misturedas’com a fauna. Sondagens de foss0
foram realizadas na Toca do Serrote do Arhur &
temvarias lagoas, especialmente Szo Vitor e Lagoa
daPomba. Também, restos fosséisforam achades
numa exploracao realizada no Sumidouro. do
‘Sansao.
© conjunte da fauna de mamiferos, que data
do Pleistoceno superior, é testemunha da
exsténcia de uma paisagem caracterizada pela
savana arbustiva, entrecortada de zonas de
floresta, com um clima muito mais Umido do que
oalual, Esta fauna é contemporanea dos mais
antigos vestigios, atualmente conhecidos, da
presenca do homem nas Américas.
|- Generalidades:
Desde 1970, as pesquisas realizadas no
Sudeste do Piaui (Nordeste do Brasil, fig. 1), por
N. Guidon e sua equipe peimitiram inventariar
aproximadamente 400 sitios arqueolégicos na
regio de Sao Raimundo Nonato (Arnaud ef ali,
1984; Guidon, 1986; Guidon ef ali, 1989). Mais
de dois tergos desses sitios, so conhecidos por
‘sua arte rupestre (Guidon, 1984 a: Pessis, 1987)
um grande numero deles estao localizados no
Parque Nacionel Serra da Capivara, limitado pela
cuesta arenitica do Paleozéico inferior; uma
dezena destes sitios foram escavados ou neles
se realizaram importantes sondagens de fosso.
Tres desses sitios, na formacao arenitica,
apresentam niveis paieolticos: Toca do Caldeirao
do Rodrigues |, datado de 18.600 + 600 anos EP
(Guidon, 1981: 19841b ec), Toca do Sitio do Mei
alualmente em escavacdo, na quel os niveis mis,
artigos se situam entre 12.200 + 600 e 14 300+ 4. Claude Guérin et alii
SBA
Figura 1 -Situacdo geogrica dos sos paleontolbices da regio de Sao Raimundo Nonaio(carontafia
arent)
41: Frente da cuesta
2: Estradas:
3: Rios e riachos temporarios
4: Trihas
6: Serrotes calearios: GA; Garrincho: SA: Serrote ¢o Atur; SBA: Serrote da Barra do Antoniao; SM: Serrote
das Moendas, SS: Sumidoure do Senso
6: Povoados: CJD: Coronel José Diss e SiMe: Sitio do Mocd
188: Toca de Cima dos Pilao
Figure 1 - Geographic locetion of paleontological sites in the Sao Raimundo Nonaio region (cartography F
Parenti)
4: Front of the cuesta
2: Roads
3, Rivers and temporary creeks
4: Traits
5: Limestone outcrops: GA: Garrincho; SA: Serrote do Artur; SBA: Serrots da Bérra do Antonio; SM: Serrote
das Moendas, SS: Sumidouro do Sanséo
6: Hamlets; CJD: Coronel Jose Dias e SiMo: Sitio do Moco
188: Toca de Cima dos Pilg
57 FUMDHAMENTOS
(Guidon & Andreatta, 1980; Guidon, 1984 b and
© Guidon & Delibrias, 1985) and the Toca do
Boqueirto do Sitio da Pedra Furada. This last
site, excavated during ten years, presents ¢ 5
motors thick filling with three Pleistocene and
three Holocene levels. Like in all the sites in
sandstone context, the acidity of the sediment
(pH = 4) has not permitted fossil conservation ,
ut more than 7,000 lithic ertifacts, of which 600
are from the Paleolithic and approximately 160
features whose majority corresponds to hearths
were discovered. Fifty four radiocarson datings,
32 of which from the Pleistocene were produced
from charcoal, the earliest is older than 48,500
BP. Pedra Furada was the subject of several
preliminary papers (Guidon & Delibrias, 1986;
Delibrias ef alii, 1988; Guidon et alii, 1989;
Parenti ef alii, 1990) and most recently a thesis
by F, Parenti (1993 b),
Since 1988, several sites were discovered in a
completely different geological context. Located
southeast of the previous ones (fig. 1) in the
metamorphosed limestone massifs and, thus,
favorable othe conservation of bone materia, hey
are different in that they are able of conserving an
‘e Ursidae
- Arctodus (Paractotherium) brasiliensis
(Lund, 1840)
Acspécie foi definida no Pleistoceno de Minas
Gerais # revisada em 1960 por C. de Paula Couto,
Este pequeno ursideo esia representado na Toca
de Cima dos Pildo por elementos de arcadas
dentarias correspondentes a dois individuos.
7 FUMDHAMENTOS
teeth’s design perfectly correspond to the ones
provided by C. de Paula Couto (1960, fig. 4 and
5; 1979, fig. 360). The species was known only
in Minas Gerais and in Tarija ( Bolivia)
- Aretodus (Arctotherium) cf, bonaerense
(Gervais, 1855)
‘We have attributed to tis very talland short faced
bear, one Galeaneus and thee isolated teeth found in
Garrincho. ‘The species has been determined in the
‘Argentinean Pampas and had never been found with
Certainty in Braz before.
15- Felidae
= The medium sized felines: Falis
yagouaroundi €. Geoffroy, 1803 and Felis
pardalis Linné, 1788
Some mandibles halves, isolated teeth and
entire long bones attributable to the yagouaroundi
‘were found in the Barra do Antoniao. Some parts
were also found in the Toca de Cima dos Piléo
(mandibles fragments, tibia and entire
metapodes, phalanx) belonging to the same
species. This feld had already bean determined
in Brazil's Pleistocene (Paula Couto, 1979)
particularly in the sites of the Rio das Velhas
(Minas Gerais). Other remains in Garrincho and
‘Antonio (particularly one maxillary fragment,
tooth and ono humerus, one radius, one femur,
and one Mtl, all complete) belong to a much
larger feline, probably the ocelot, Felis pardalis,
also known in the Minas Gerais Pleistocene.
The medium and smail felines, that live
currently in the region, are Felis yagouaround,
F trina, F pardalis, F. wiedii (Barbosa Souza
& Olmos, 1991)
- The great felines: Panthera onca (Linné,
1758) and Felis concolor Linné, 1758
A certain number of jeguar remains
particularly one Mtl, one Calcareus anda first
phelanx, very well preserved, are originated from
the Gartincho: other vestiges, particularly one
hand in connection were collected from the Toca
de Cima dos Pildo. The species is known in the
Ple'stocene of South America, specially in Brazil
(Paula Couto, 1979)
Other remains, such as a metapodial anda first
phalanx found at Garrincho and several parts found.
inthe Toca de Cima dos PilAo should be attributed
without any doubt to a large puma, the existence of
which in the Pleistocene was recognized by C. de
Paula Couto (1979). The two species are curently
in existence in the National Park Serra daCapivara,
76
deseno dos dentes corresronde perfeitamente
20s fomecios por C. de Paula Couto (1960. fig. 4
@ 5; 1979, fig. 360). A espécie era connecida
somerte em Minas Gerais e em Tarja (Bolivia).
- Aretodus (Aretotherium) cf. bonaerense
(Gervais, 1855)
Atribuimosa esse grande urso allo e de cara
curta, um calcaneo e tres dentes isolados
encontrados no Garrincho. A espécie foi definida
no Pampa argentino e nunca havia sido,
seguramente, assinalada no Brasil
15- Felidae
= 0s felines de tamanho médio: Felis
yagouaroundi E. Geoffroy, 1803 e Felis
pardalis Linné, 1758
Algumas meias mandibuls, dentes,jsolados ©
‘ossos longos inteiros, alibulvels 20 yagoueround
foram achados na Barra do Antoniao. Algumas pecas
também achadas na Toca de Cima dos Pilao
(fragmentos de mandibulas, tibia e metépodes
Completos, falanges) pertencem a mesma espécie.
Este Felideo ja Navia sido indcado no Pleistoceno
4o Brasil (Paula Couto, 1979), em particular nos sitios
do Rio das Velhas (Winas Gerais). Oultos restos do
Garrincho e do Antondo (em particular um fragmento
cde manila, dentes e um tmero, um rédio, um Emu e
‘um Mil, todos completos) pertencem a um felino bem
‘maior, que muito provavelmente seria um ocelote,
Felis pardolis, conhecido também no Pleistocene
de Minas Gerais
5 fells de tamanho médio e pequeno, que
existem atuaimente na regizo sao Fells
‘yagouaroundi, F.tigrna, F pardalis, F wiedi (Barbosa
Souza & Omos, 1991).
0s grandes felinos: Panthera onca (Linné,
1758) @ Felis concolor Linné, 1758
Um corto ntimero de restos de oneas, em
particular um Mt Il, um calcéneo e uma primeira
felange, muito bam conservados, provim do
Garrincho; outros vestigios, particulamente uma
mao em conexio, foram recolhidos na Toca de
Cima dos Pilo. A espécie ¢ conhecida no
Pieistoceno da América do Sul principalmente no
Brasil (Paula Couto, 1979).
Outros restos, como um metapode e uma
primeira falange, provenientes. do Garrinchoe
diversas pecas achadas na Tosa de Cima dos Pizo,
devem ser alvibuldas, sem duvida, a um grande
puma, cujaexistincia no Peistocen foi admiida por
6.dePaule Couto (1978). As cuas espéciesexistem
‘atualmenie no Parque Nacional Sorra da Capivara. = Smilodon populator Lund, 1840
Represented by only one part in the Barra
do Antoniao, this great machairodont is
abundant in the Toca de Cima dos Pilao where
a crushed skull, two mandible, isolated teeth
and numerous post-cranial skeleton parts,
specially carpal and tarsal bones, metapodials
and phalanx, were collected and form about
16% of the remains or of the assemblages of
determinabie remains. An upper canine tooth
is 265 mm long and an lower one 28,5 mm.
‘This exceptional abundance for a camivorous
has led us to interpret the site as a den of
‘Smilodons, whose favorite preys must have
been the Catonyx, and above all the Pecaris,
Smilodon populator was pointed out by C.
de Paula Couto (1979, 1980) in several sites in
Brazil's Pleistocene, among others in Minas
Gerais and in the Northeast (Curimatas, Paraiba;
ltapipoca, Ceara), The synonymy is confusing
(Hoffstetter, 1952) and some authors only
fecognize one species in the Upper Pleistocene
of the Americas. According to B. Kurten & E.
Anderson. (1980) with whom we agree, the
South American Smiodon is a distinct species
from the North American Smiloden fatalis . the
general size is lerger and the distal segments of
the limbs are very short.
16 - Macraucheniidae
~ Macrauchenia cf. patachonica Owen, 1840
This great Litoptern is not rare in the Barra
do Antoniaio (approximately 40 remains wet
identified - isolated teeth and post -cranial
skeleton - representing 3,7% of the identified
Megafauna). It was also found, but ina smaller
proportion in the other four studied sites,
Recently C. Cartelle & G. Lessa (1988)
proposed the denomination of Xenorhinotherium
bahiense for the Pleistocene Macraucheniidae
living in the tropical regions of Brazil, reserving
M. patachonica for those of the more Southern
zones. Not knowing the boundaries of generic
and speci bility of the latter we will
provisionally utilize the denomination to M. cf
patachonica for the Litoptern from Paul
‘The genus which up to now had not been
found in a prehistoric site (Anderson 1984) has,
been determined in several sites of the
Northeast (Hoffstetter, 1966: Paula Couto,
1960): Lage Grande and Pesqueira
(Pernambuco), Curimatas and Tapero’
3. Claude Guérin et alii
= Smilodon populator Lund, 1840
Representado por uma s6 pega na Barra do
Antonido, esse grande machairodonte é abundanie
na Toca de Cima dos Pilao. onde um cranio
‘esmagado, duas mandibulas e dentes isolados e
numerosos elementos do esqueleio pés-creniano,
sobretuido 0s carpianos, tarsianos, metépodes e
falanges, foram recolhides, constituindo cerca de
16% dos restos e dos conjuntos de restos
determinaveis. Um dente canino superior atinge
um comprimento de 265 mm e um inferior tem.
comprimento de 28,5 mm.. Esta abundancia
excepcional para um carnivoro, nos levou a
interpretar 0 sitio como um esconderijo de
‘Smilodons, cujas présas favoritas deviam ser os
Catonyx e sobretudo os Pecaris.
‘Smilodon populator fol assinalado por C. de
Paula Couto (1979, 1980) em varios sitios do
Pleistoceno do Brasil, entre outros em Minas
Gerais e no Nordeste (Curimatas, Paraiba;
ltapipoca, Ceara). A sinonimia é confusa
(Hoffstetter, 1952) e alguns autores reconhecem
somente uma Unica espécie no Pleistoceno
superior das Américas. Segundo B. Kurten & E.
Anderson (1980), com os quais concordamos, 0
‘Smilocon sulamericano ¢ uma espécie distinta do
‘Smilocon fatalis da América do Norte, o tamanho
320 mm; radius 335 mm
(327 to 343, n= 3); metacarpal cannon 212,7
mm (209,5 to 216, n= 8); tibia 371,7 mm (368
to 375, n=3);, metatarsial cannon 228,7 mm (223
235, n=8),
It_ is classical to consider Hippidion (in
general all of the Hippidiforms) as mammals
adapted to the mountain environments (they
should had hence the distal limb segments
relatively shorter than the ones belonging to the
real horses), but this interpretation cannot be
applied to the region of $40 Raimundo Nonato
where the relief is not high nor very steep. Its
association with Equus is also an argument
derying this hypothesis. R. Hofstetter (1966)
consider Hippidion as 2 hot climate or hot to
temperate animel living on open regions low or
slightly elevated
‘The background of our knowledge on the
genus Equus in South America and particularly
in Brazil is complex (Souza Cunha, 1971; Paula
Couto, 1979). Equus neogaeus which is part
of the sub-genus Amerhippus Hoffstetter is
‘common in the Pleistocene sites of the Central
Planalto of Brazil and appears very frequently
associaled to Hippidion (Paula Couto, 1979,
1980). E. neogaeus is known in the Rio das
Velnas and in Aguas de Arax4. (Minas Gerais),
Pesqueira e Lage Grande (Pernambuco),
Chique-Chique in the valley ofthe Sao Francisco
river (Bahia), Curimatas (Paraiba) and in
Itapipoca (Ceara). It seems to be well
represented in Barra do Antoriao, in the Toca de
Cima dos Pilao and in the Serrote do Artur, but
the exact determination of the species requires
the discovery of more abundant material.
Like every real horse, the Equus neogaeus
inhabited the open plains and consumed a lot
of graminecus herbs.
20 -Tayassuidae
- Dicotyles tajacu (Linné, 1758) and Tayassu
ecari(Link, 1795)
The two peccaries nowadays very
common in South America are present in the
three major faunal sites. particularly in the
Toca de Cima dos Pilao, where they comprise
almost 40% of the material. M.0. Woodburne
(1988) has demonstrated that it is nacessary
to go back to a generic level distinction while
a long time ago, both belonged to the Tayassu
genus. The two species are easily
recognizable from biometric data and they
belong to the modern fauna of the region
(Emperaire, 1989; Barbosa Souza & Oimos,
80
achados no Garrincho so 08 seguintes: mero
320 mm; radio 335 mm (327 a 343, n=3);
canh&o metacerpiano 212,7 mm (209,5 8 216,
n=8); tibia 371.7 mm (368 a 375, n=3); canhéo
metatarsiano 228,7 mm (223 a 235, n=8).
E classico considerarHippidion (numa manera
geral todos 0s Hippidiformes) como mamiferos
adaptedos @ meios montanhosos (eles teriam,
‘nesse caso, os segmontos distais dos membros
telativamente mais curtos que os dos verdadeitos
avalos), mas esta interpretagao no pode ser
aplicaca regiao de Sao Raimundo Nonato onde
© relevo néo ¢ alto nem muito escarpado. Sua
associacdo com Equus & também argumento que
vai contra essa hipétese. R. Hoffstetter (1966)
considera Hippidion como um animal de clima
quente ou quente a temperado, que vive nas
regides abertas, baixas ou pouco clevadas.
© hiisterico de noses conhecimentos sobre
© género Equus na América do Sul, 2
etticularmente no Brasil, € complexo (Souza
Cunha, 1971; Paula Couto, 1979). Equus
Neogaeus que faz parte do sub-género
Amerhippus Hofstetter € comum nos sitios
pleistocénicos do Planelto Central do Brasil ¢
‘aparece, muito frequentemente, associado 20
Hippidion (Paula Couto, 1979, 1980). .
neogaeus € conhecido no Rio das Velhas e em
Aguas de Araxé (Minas Gerais), Pesqueira
Lage Grande (Pernambuco), Chique-Chique no
vale do Rio Sao Francisco (Bahia), Curimatés
(Paraiba) e em Itapipoca (Ceara). Parece bem
Fepresentado na Barra do Antonio, na Toca de
Cima dos Pildo e no Serrote do Artur, mas uma
determinagdo exata da espécie fequer a
descoberta de um material mais abundante.
Como todos os verdadeiros cavalos, Equus
neogaeus frequentava as planicies abertas e
consumia muitas gramineas.
20 - Tayassuidae
= Dicotyles tajacu (Linné, 1758) e Tayassu
pecari (Link, 1795)
Os dois pecaris, hoje muito comuns na
América do Sul, est4o presentes nos trés
principais sitios faunisticos, particularmente na
Toca de Cima dos Pilao, onde constituem quase
40% do material. M. ©. Woodbume (1968)
‘mostrou que é necessario voltar a uma distin¢go
a nivel genérico, ao passo que, faz muito tempo,
todos os dois estavam situados no género
Tayassu. As duas espécies, que sao facilmente
reconheciveis a partir de dados biométricos,
fazemparte da fauna atual da regio (Emperaire,
1989; Barbosa Souza & Olmos, 1991); elas $80 1981); they are known in the Minas Gerais
Ple'stocene (Paula Couto, 1979).
The two forms do not have the same
ecological meaning: the small species (D. tajacu)
is more ubiquitous whilst the large one (T. pscar)
prefers humid forests.
24 -Camelidae
~Paleeolama sp and P. major (Lials, 1872,
‘Winge, 1908)
The genus Palaeolama is present in all of
the sites, its remains (mandible, tooth rows,
complete specimens of all the elements of t
posi-cranial skeleton) respectively constitute
16.1% and 32.7% of the megafaune from Barra
do Antoniao and Garrincho.
This genus has been studied by A. Cabrera
(1932) and was the subject of a delailed revision
by R. Hoffstetter (1952) who considers Auchenia
(pro-parte), Hemiauchenia and Protauchenia
Synonyms and indicates P. major as the only
species of the Brazilian Pleistocene.
The latter one is known in the Rio das Velhas
(Minas Gerais) in the wells and water- holes of
Ceara (Itapipoca) and of Paraiba (Curimatas),
In the caves and in the wells of Bahia, in Lage
Grande (Pemambuco). E. Anderson (1984)
pointed out that Palaeolama never has been
discovered associated to fossil man.
The long bones and metapodials collected
in Antonio and in Garrincho are very similar in
the two sites (fig. 4 and 5). Their average length
are the following: humerus 348,4 (5 of Garrincho,
2 of Antonido: 325 to 400 mm); radius 409,7 (3
from Gartincho, 398 to 416 mm); metacarpal
cannon $05.8 (4 ftom Garnincho, 2 from Antoniao:
300 to 319 mm) : femur 443 (Garrincho): tibia
408.3 (3 from Antoniao: 396 to 420 mm).
metatarsal cannon 330.8 (4 from Garrincho and
1 from Antonio: 328 to 336 mm).
Two long bones originated from the
Serrote do Artur are notably shorter (384 for
one femur, 330 mm for one radius), and
evidently correspond to a different species,
‘A comparison with the values published by
R. Hofstetter in 1952 (P. reissi, from the
Equatorial Andes, major from Lagoa Santa in
Minas Gerais, P paradoxa in the Northwest of
Argentina and also some pieces of P. weddell
from Bolivia, of P aequatorialis rom the Pecific
coast of Equador and P. crassa of the Equalorial
Andes) shows thal the existing form in Garrincho
3. Claude Guérin et alii
conhecidas no Pleistoceno de Minas Gerais
(Pauls Couto, 1979),
As duas formas nao tem a mesma
significagao ecolégica: a poquena espécio (D.
tajacu) & mais ubiquista, enquanto que a grande
(7 pecari) pretere as florestas umidas.
21- Cametida
=Palaeolamasp. e P. major
(Liais, 1872, Winge, 1906)
© género Palaeolama esta presente em
todos os sitios, seus restos (mandibula, arcadas
dentarias, ospécimens completos de todos os
elementos do esqueleto pés-craniano)
constituem respectivamente 16,1% @ 32,7% do
‘material da Barra do Antonia e do Garrincho.
Ogénero foi estudado por A. Cabrera (1932)
e objeto de uma revisdo detalhada por R.
Hoffstetter (1952), que considera sinénimos
Auchenia (pro-parte) Hemiauchenia e
Protauchenia, 0 indica major como a Gnica
espécie do Pleistoceno brasileiro
Esta ultima @ conhecida no Rio das Velhas
(Minas Gerais), nos tanques e cacimbas do
Ceara (Itapipoca) eda Paraiba (Curimatas), nas
grutas e nas cacimbas da Bahia, em Lage
Grande (Pernambuco). _E. Anderson (1984)
precisa que Palaeolama nunca foi descoberta
associada ao homem fossil
‘Os ost08 longos © 08 metapedes recolhidos
no Antonido e no Garrincho sao muito
semelhantes nos dols sitios (fig. 4 € 5). Seus
comprimentos médios sao os seguinies: mero.
348.4 (5 de Garrincno, 2 de Antoniao: 325 a 400
mm); radio 409,7 (3 de Garrincho, 398 a 416
mm); canhao metacar
443 (Garrincho); tibia 408,3 (3 de Antonio: 396
4.420 mm); canhdo metatarsiano 390,8 (4 do
Garrincho, 1 de Antonido: 328 a 336 mm),
cis ossos longos provenientes do Serrote do
‘Artur sao sensivelmente mais curtos (384 mm para
tum férmur, 330 mm para um radio), e correspondem,
evidentemente, a uma espécie diferente,
Uma comparagao com os valores publicadas
por R. Hoffstetter em 1952 (P. reissi, dos Andes
equatorianos, P major de Lagoa Sania em Minas
Gerais, P. paradoxe do Noroeste da Argentina,
e lambém algumas pecas de P. weddelli da
Bolivia, de P. aequetoriais da costa pacitica do
Equador e P crassa dos Andes equatoranos)
‘mostra que a forma presente no Garrincho @ no
ay FUMDHAMENTOS
Figura 4 - Umero direito de Palaeolama sp n* 35150-1, Garrincho, vista antetior (desenho de E. Gomes de
Maios Mede'ros).
Figura § - Radio-cibito direito de Palaeolama sp n® 37534, Garrincho, vista de trés quartos frente (desenho
do E. Gomes de Matos Medeiros).
Figure 4 Right humerus of Palaeolama sp n° 35150-1, Garrincho, anterior view (drawing by E.Gomes de
Malos Mede'ros).
Figure 5 - Right Radius-Cubitus of Palaeolama sp n° 37534, Garrincho, three quarters front view (drawing
by E, Gomes de Matos Medeiros).
82 ‘and in Antonia is clearly larger. Only P paradoxa
approximates itself and this approximation is true
only for the radius.
Much to the contrary, the remaine of the
Serrote do Artur show dimensions closer to the
ones obtained for P. major and P. reissi
Everything happens as if we were in the presence
Of two quite different species, P major in the
Serrote do Artur and P. sp. of a larger size
Garrincho, in Antonio and in the Toca de Cima
dos Pildo.
The relatively hypsodont Palaeolama was
a sprout, leaves, small branches and also 2
graminous eater. The genus was principally
Ubiquitous. To B. Kurten & E, Anderson (1980)
the limbs segments relationships show thal
they were very well adapted to irregular terrain.
22 -Cervidae
- Mazama sp. and "Cervus" sp.
The small Cervine Mazama is rare in the
Barra do Antonido and in the Toca de Cima
dos Pilao where it is represented only by about
20 remains; itis slightly less rare in Garrincho,
L. Emperaire (1989) and M.F. Barbosa & F
‘Olmos (1991) pointed out two recent species
in the region: M. quazoubira (G. Fischer, 1814)
and M. americana (Erxleben, 1777) but the
insufficiency of the available material does not
ensure determination in the specific level. The
genus is known in Brazil's Pleistocene
(Carette, 1922; Paula Couto, 1979, 1980;
‘Souza Cunha & Magalhées, 1986) specially in
the Rio das Velhas (Minas Gerais) where the
two species simplicicornis and americane
were identified, in Curimatas (Paraiba) and in
Ntapipoca (Ceara). Mazama, like almost all
Cervidee, is a genus with forests habits,
‘The remarkable scarcity of South American
cervines of a larger size should be noted, such
as Odocoileus and Blasiocerus, the latter one
seemed to be represented by numerous rock
art figures of the S80 Raimundo Nonato
archaeological area (Pessis, 1991).Two smal
fragments of antlers and a complet second
phalanx from a large deer were however found
in Barra do Antoniao.
B-Birds.
‘The preliminary list of the birds ientiied in the
Jocalty of Barta do Antonio is as follows:
4. Claude Guérin et alii
Antoniao é nitidamente maior. Somente P
peradoxa aproxima-se, e esta aproximacao
Somente & verdadeira para o radio.
‘Ao contrairio, 0s restos do Serrote do Artur
apresentam dimensdes préximas das
levantadas para P. major e P. reissi. Tudo
acontece como se estivéssemos na presenca
de duas especies bem diferentes, P. major no
Serrote do Artur e P. sp. de tamanho muito
grande, no Garrincho, no Antonido @ na Toca
de Cima dos Pildo,
Pélaeclama, relativamente hipsodonte, era
um comedor de brotos, folhas, pequenas ramas
© também gramineas. O génoro ora
principaimente ubiquista. Para B. Kurten & E.
Anderson (1980), es relagdes dos segmentos
dos membros mostra que eles estavam muito
bem adaptados aos terrenos irregulares.
22- Cervidae
-Mazama sp. ¢ “Cervus” sp.
‘O pequeno Cervideo Mazamaé raronaBarra
do Antonido © na Teca de Cima dos Pildo, onde
‘esta representado somente por cerca de 20 pecas;
ele € ligeiremente menos raro (12 vestigios) no
Garrincho. L. Emperaire (1989) e M. F. Barbosa &
F. Olmos (1991) assinaiam na regiao as duas
‘espécies aluais M. quazoubira (G. Fischer, 1814)
eM. americana (Etxieben, 1777), mas 3
insuficiéncia do material disponivel nao autoriza
uma determinagéo ao nivel especifico. O género
@ conhacide no Pleistoceno do Brasil (Carette,
1922; Paula Couto, 1979, 1980; Souza Cunha &
Magalhées, 1986), especialmente no Rio das
Velhas (Minas Gerais), onde as duas espécies
simpliccornis e americana foram reconhecidas, em
Curimatas (Paraiba) e em ltapipoca
(Ceara).Mazama @, como quase todos os
Cervideos, um género de habito florestal
Observar-se-4 a raridade, bastante
surpreendente, dos cervideos sul-americanos de
‘maior tamanho, como Odocoleus e Blastocerus,
sendo que este tiltimo aparece representado em
umerosas figuras rupesties da érea arqueoiégica
de Sao Raimundo Nonato (Pessis, 1991). Dos
pequenos fragmentes de chifies e uma segunda
falange completa de um grande veado foram,
‘entretanto, encontrados na Barra do Antoniao.
B- Aves
Aliista pretiminar das aves identificadas no
lio Barra do Antonio é a saguinte:
83 FUMDHAMENTOS
Tinamifornes
Cryptureltus noctivagus, Yellow - legged
‘Tinamou
Crypturells parvirostris, __Smalt-biled
Tinamou
Ciconifornes
Theristicus caudatus, Buftnecked Ibis
Anseriformes:
‘Amazonetta brasiliensis, Brazilian
Duck
Caiiina moschata, Muscovy Duck
‘Accipitrformes
‘Accipiter bicobor, Bcolored Hawk
Polyborus plancus, Crested Caracara
Faleo rufgulais, Bat Falcon
Faleo sparverius, American Kestrol
Gatiformes:
‘Penelope supercians cr jacucaca,
Rusty-margined or While-browed Guan
Gnuforres
Porzana caroiina, Sora Rail
cf, Porphyrula,
cf, Galle
Columbiiormes,
‘Columba picazuro, Picazure Pigeon,
Zenaida euriculata, Eared Dove
Columbia minuta, Plain-breasted
Ground-Dove
CColumbina picui, Picui Ground-Dove
cf. Columbina falpacot cf. Ruddy
Ground-Dove
Pesitaciformes
‘Arachloroplera, Red-and-Green Macaw
‘Aratinga leucophialmus, White-2yed
Parakeet
cf, Aratinga cactorum, cf. Cactus Parakeet
cf, Amazona aestiva, cf, Turquoise-fronted
Parrot
Strigiformes
‘Tyo alba, Barn Owl
Otus choliba, Tropical Screech-Ow!
Glauciaium minatissimum, Least
Pygmy-Ow!
Gleucicium brasilianum, Ferruginous
Pygmy-Ow!
Clocaba virgata, Mottied Ow!
Rhinoplynx clamator, Striped Owl, or
Ciccaba huhula, Black-banded Ow!
Caprimulgitormes
cf. Hydlopsalis, Nighijar
Tinamiformes
Crypturellus noctivegus, Zabelé
Crypturellus parvirostris, Nnambi
xororo
Ciconiformes
Theristcus caudatus euricaca
Anseriformes
‘Amazonetta brasiliensis, Marreca pé
vermelho
Cairina moschata, pato
Accipitriformes.
Accipiter bicolor, Gaviao bicolor
Polyborus plancus, Caracara
Falco rufigularis, Gavido morcego
Falco sparverius, Quiri-quiri
Galliformes
Penelope supercilieris ou jacucaca,
Jacupemba ou jacucaca
Gruiformes:
Porzana carolina,
cf, Porphynula, frango agua
cf. Gallinule
Columbiformes
Columba picazuro, Asa branca
Zenaida auriculata, Avoanto
Columbine minuta, Rolinha caxexa
Columbina picui, Rolinha branca
ct. Columbina talpacoti, cf. Rolinha
caldo-de-feija0,
Psittaciformes
Ara chloroptera, Arara vermelha
Aratinga loucophtalmus, Periquitao
cf, Aratinga cactorum, cf. Jandaia-gangarra
cf. Amazona aestiva, ef., Papagaio
verdadeiro
‘Stigformes
Tito aba, Coruja-de-igreja, suindara
(Clus choliba, Conyjnha-domato
‘Glaucitium minutissimum, caburé
‘iudinho
Glaucidium brasiianum, Caburé
Caprimulgiformes:
cl. Hydropsalis, Bacurau Apodiformes
Stroptoprocne zonaris, White-collared
Swift
Piciformes
CColaptes melanochioros, Green-barted
Woodpecker
Dryacopus lineatus, Lineated Woedpecker
oF
Campephilus melanoleucos, Crimson-
crested Woodpecker
Passeriformes
Undetermined middle- and small-sized
forms.
The minimal number of individuals (MNI) is
given in Table 2.
The ecological categories have been
established after the Indications provided by Sick
(1993). These categories are:
F: Predominantly Forests, also Groves and
Forest-edges inhabitants,
FSO: Inhabitants of Forests, Scrubs and Open
regions, including Forest borders, Capoeiras,
Trees in the Cerrado and Caatinga, Orchards,
River-edges, Brushy open countries.
©: Live predominantly in Open regions, trom
Savanna with sparse trees, Open countries
with low vegetation and ‘Yields, to Open
countries sometimes almost devoid of
vegetation
MR: Marshes and Rivers.
C: Gifts,
The majority of the species identified are stil
living today inside the Parque Nacional da Serra
da Capivara, where the locality of Antonio is
situated, and in the adjacent areas (Barbosa
Souza & Olmos, 1991). The species which are
no longer found in this National Park are
Theristicus caudatus, Bul-necked Ibis, Falco
rufigularis, Bat Falcon, Porzana carolina, Sora
Rall, Gleucicium minutissimum, Least Pygmy-
Owl, Ciccaba virgata, Mottled Owl, Rhinoptynx
clamator, Striped Owl, or Ciccaba huhula, Black-
banded Owl,
‘Among these species, Theristicus caudatus,
the Buff-necked Ibis, is widespread throughout
Brazil, and its present-day absence in the Park
‘may be fortuitous, The species Falco rufigularis,
Glaucidium minutissimum, Ciccaba virgata and
Ciccaba huhula are forest forms and their
present-day absence may be correlated with a
3. Claude Guerin et alii
Apodiformes
Streptoprocne zanaris, Andorinhao de
coleira
Piciformes
Colaptes melanochioros, Pica-pau
verde-barrado
Diyocopus lineatus, Pica-pau-de-
topete-vermeiho,
ou
Campephilus melanoleucos, Pica-pau
Passeriformes
Formas nao determinadas de tamanho médioe
pequeno.
© numero minimo de individuos (NMI) €
dado na tabela 2.
‘As categorias ecol6gicas foram eslabelecidas
seguindo 5 indicagbes de Sick (1993), Essas
calegorias sao as seguinies:
F: Predominantemente frequentadores ce florestas,
mas também arvoredos e limites de florestas.
FSO: Habitantes de florestas, mata e regides
abertas, incluindo margens de lorestas, capooiras,
rvores no cerrado e caatinga, pomares, beiras de
Tios € regides abertas de mato rasteiro.
©: Frequentam principalmente regioes abertas,
desde a savana com arvores esparsas, regides
abertas com vegetacao baa e campos, als
Areas abertas algumas vezes quase que
inleiramente desprovidas de vegetagao.
MR: Benhatbos e rios,
C: Falésias
A maioria das espécies identificadas ainda
vive hoje no Parque Nacional Serra da
Copivera ¢ areas adjacentes, regio na qual
se situa 0 sitio Barra do Antonido (Barbosa
Souza & Olmos, 1991). As espécies que nao
existem mais no Parque Nacional s&o:
Theristicus caudatus, curicaca, Falco
rutigularis, Gavido morcego, Porzana carolina,
Glaucidium minutissimum, Caburé miudinho,
Ciccaba virgata, Coruja-do-mato, Rhinoptynx
clemator, Corujd-orelhuda ou Ciccaba huhula,
Coruja-pretal.
Entre essas espécies, Theristicus caudatus,
© curicaca, é amplamente distribuido em todo 0
Brasil @ sua auséncia atualmente no Parque,
pode ser fortuita. As especies Falco rufigularis,
Gleucidium minutissimum, Ciccaba virgata &
Ciccaba huhula sao formas florestais ¢ sua
auseénicia hoje pode ser correlacionada com a
85 FUMDHAMENTOS
Tabela 2 : Lista preliminar das espécies de aves da Barra do Antoniao
NMI: Namero minimo de individuos
Categorias ecologicas segundo Sick, 1993: F: Floresta; FSO: Floresta, matae outras regides; O: principamente
regides aberias; MR: Banhados e rios; C: Falésias
Table 2 : Preliminary Ist of the birds species of Barra do Antonio
MNI: Minimal number of individuals
Ecological categories after Sick, 1993: F: Forest, FSO: Forest, scrubs and open regions; O: Predominantly
‘open regions; MR: Mershes and rivers; C: Cifs
Adults Juveniles | Total Ecological |
MNI MNI MN categories
Adultos | Juvenis Total Categorias
NMI NMI NMI ecoléaicas
Crypturellus nootivagus 1 0 1 FSO
‘Crypturellus parvirostris 2 1 3 FSO
|_Theristicus caudatus 1 0 a oO
‘Amazonetta brasiiensis i 0 i MR
Cairina moschata a 0 1 MR
Accipiter bicolor i 0 7 F
| Polyborus plancas i 0 i o
Falco rufigularis 1 0 i i
Falco sparverius a 0 1 0
Penelope supercilaris = - - FSO
or Penelope jacucaca A 0 4 FSO
Porzana carolina 7 0 1 MR
[et Porphiyruta 7 0 7 MR
Columba picazure 1 0 7 FSO
Zenaida auriculata =e 1 a oO
[Columbine minute 5 i 6 o
Columbina picui 7 0 A °
cf. Columbine talpacoti + 0 1 ° i
Ara chioroptera 4 6 F
Aratinga leucophtalmus 3 i a F
i. Arafinga cactorum a 0 2 FSO
ef. Amazona aestiva i 1 z FE
Tyto alba 3 i 4 0
Otus choliba mp 3 | FSO
Glaucidium minutssimum 7 o 7 ad
|" Glaucidium brasiianum i 0 1 FSO
| Ciecaba virgata ale 0 lac FE
Rhinoptyax clamator = = = FSO
‘or Ciccaba huhula 7 0 a F
| cf Hydropsalis 4 0 _ T FSO
‘Streptoprocne zonaris 7 0 7 Cc
Colaptes melanochloros 7 0 7 FSO
| Dryocopus lineatus = = FSO
‘or Campophius molanoleucos_|_ 1 o 1 FSO
Total 52 9 61
86 modification of the environment leading to a
eduction in the forested areas. Porzana carolina
is amigratory bird which breeds in Nortn America
and winters in Central and South America,
southward to Colombia, Venezuela, Guyana,
Ecuador and Peru. At the present time it does
ot reach Brazil (Blake, 1977), but its wintering
‘grounds were perhaps more extended during the
past.
The fossil avifauna from Antoniao differs
from the present- day avifauna of the National
Park by the pronounced scarcity of the aquatic
forms. Indeed, at the present time the aquatic
birds are represented in the park by the following
families:
Podicipedidae, Grebes,
Phalacrocorecidae, Cormorants,
Anhingidae, Darters,
Ardeidae, Herons (9 species),
Anatidae, Swans and Ducks (4 spec
Aramidae, Limpkins,
Rallidae, Rails Gallinules and Coots
(5 species)
Jacanidee, Jacanas,
Charadridae, Lapwings and Plovers,
Scolopacidae, Sandpipers and Snipes,
Recurvirostridae, Stilts,
Alcedinidae, Kinglshers.
8)
These families are absent from the
preliminary list of the fossil material, with the
exception of the Anatidae and Rallidae, which
are each one represented by two species. This
almost total absence can perhaps be explained
by alimentary preferences either of prehistoric
men, or of raptorial birds, who are responsible
for the accumulation of this material.
On the other hand itis interesting to note
thatAra chioroptera, the Red-and-Green Mecaw,
is represented by a large number of remains. It
is possible that this species was selectively
hhunied by the inhabitants of the site. It must have
been rather abundant in the former times, while
Now restricted to a very small population
(Barbosa Souza & Olmos, 1991).
On the whole the majority of the
iduals correspond to forest forms (26 %),
or to forest, scrub, and open region forms (26
%). Next there are forms from predominantly.
‘open regions (39 %), mainly including small
doves. The Eared Dove, Zenaida auriculata,
is very abundant in the fossil material, and it
is known that this species becomes extremely
numerous in the Northeast at intervals of two
or three years (Sick, 1993), Birds of marshes
and rivers are very few (7 %). Finally, only
3. Claude Guérin et alii
mudanga do meio embiente que ocasionou uma
tedugao das areas florestais. Porzana carolina
@ um passaro migratorio que se reproduz na
‘America do Norle © passa os invernos na
‘América Central e do Sul, no sul da Colombia,
Venezuela, Guiana. Equadore Peru. Atualmente,
ele néo atinge 0 Brasil (Blake, 1977), mas seus
territérios invernais podem ter sido mais
extensos no passado,
A avifauna da Barra do Antoniao difere da
avifauna atual do Parque Nacional por uma
pronunciada escassez de formas aquaticas. De
falo, nos dias de hoje, as aves equalicas esto
representadas no Parque pelas seguintes
familias:
Podicipedidae, Merguihdes;
Phalacrocoracidae, Cormordes e biguas;
Anhingidae, Bigud-tinga;
Ardeidae, Gargas © socds (9 espécies);
Anatidge. Cisnes e patos (4 espécies);
‘Aramidae, Carao;
Rallidae, Saracuras (5 ospécio
Jacanidae, Jaganas;
Charadriidae, Macaricos:
Scolopacidae, Batuiras;
Recurvirosttidae, Pernilongos;
Aicedinidae, Martin-pescadores.
Essas familias estéo ausentes da lista
proliminar do material féssil, com excegto de
Anatidae e Rallidae, cada uma representada por
duas especies. Este auséncia quase iotal pode,
talvez, ser explicade pelas preferéncias
alimeniares seja dos homens pré-nistoricos, seja
des aves de repina, que 580 os responsaves
pela acumulacao deste material
Por outro lado, é interessante notar que Ara
chloroptere, a arare Vermetha, € representada
or um grande ndmero de restos. € possivel que
Esta especie era seletivamente cagada pelos
habitantes do sitio. Ela deve ter sido muito
abundante nas épocas passadas, enquanto que
hoje se encontra esiritaa uma populacao muita
pequena (Barbosa Souza & Olmos, 1991).
No conjunto, a maioria dos individuos
corresponde a formas florestals (26%), ou a
formas de areas de floresta, mato e regides
abertas (26%). A seguir, aparecem as formas de
fegides predominantemente abertas (39%),
incluindo principalmente pequenas pombas. A
avoante, Zenaida auriculata, é muito abundanta
No material fossil e é sebido que esta espécie
se tomna extremamente numerosa no Nordeste
a intervalos de dois ou trés anos (Sick, 1993)
Aves de banhados ¢ rios sao raras (7%).
87 FUMDHAMENTOS:
Streptoprocne zoneris, the White-collered
Swift, represents the birds from the cliffs (2
%), which should however have been
numerous in the surroundings of the caves and.
rock-shelters
In conclusion, the preliminary study of the
Antonio avifauna indicates a landscape which is,
not very different from the present - day one, but
where the forests must havebeen more developed.
The main difference is the scarcity of the aquatic
forms among birds, while these aquatic forms are
well represented among macro- (semi-aquatic
Xenarthra) and micromammals (Guérin ef alii,
1993, a and b), and while the fauna also includes
numerous remains of large-sized Amphibians.
C-Reptiles
+ Caiman sp.
Numerous bone and teeth remains were
collected in the Barra do Antonio. Two species
of this genus are present in the recent fauna of
the region, Gaiman crocodilus being the only one
represented in the zone of the Park (Barbosa
Souza & Olmos, 1991).
- Indeterminate Chelonian
Numerous carapace fragments, some
attributable to one large terrestrial turtle specios
belonging to the Geochelone group (F. de Broit
‘verbal communication) were found in Barra do
Antonio and other remains are from Garrincho.
Phrynops geoffroanus and Ph. tuberculatus are
the two species currently known in the region
(Barbose Souza & Olmos, 1981).
- Dracaena sp.
‘Two remains of this Telidae were found in
the Toca de Cima dos Piléo. The genus
currently comprises two species, _D.
paraguaiaensis and D. guianensis; the second
inhabits the Amazon forest; it has amphibious
habits and feeds on fresh water mollusks, fish
D- Amphibians
Numerous remains of different sizes were
collected in the Pleistocene and Holocene levels
in the Barra do Antonio, in the Toca de Cima
dos Piao and in Garrincho.
88
Finalmente, sormente Strepfoprocne zonaiis, 0
andorinhdio de colarinho branco, representa as
aves das falesias (2%), 0s quals, entretanto,
deveriam ter sido numerosos nas vizinhangas das
ferns € abrigos rochosos,
Concluindo, o estudo preliminar da avifauna
da Barra do Antoniao. indica uma paisagem que
‘nao € muito diferente da de hoje, mas na qual as
florestas eram mais conspicuas. A principal
diferenga 6 a raridade das formas equaticas entre
‘8 aves, enquanto que essas formas aquilicas sao
bem representadas entre os macro- (semi-aquatica
Xenarthra) e micro-mamiferos (Guerin etal, 1993,
a e b), sendo que a fauna fossil inclui também
Numerosos restos de grandes Anfibios.
C-Répteis
= Caiman sp.
Numerosos restos ésseos e denies foram
coletados na Barra do Antonio. Duas espécios
desse género figuram alualmente na fauna da
regiao, Gaiman crocodilus @ a nica
representada na zona do Parque (Barbosa
‘Souza & Olmos, 1981).
= Quelénios indeterminados
Numerosos fragmentos de carapaca, alguns
atribuiveis a uma grande espécie de tartaruga
terrestre do grupo das Geochelone (F. de Broin,
‘comunicagéo oral) foram achadas na Bara do
Antonigo e outros restos provém do Garrincho,
Phrynops geoffroanuse Ph, tuberculalus S80 as
duas espécies atualmente conhecidas na regiao,
(Barbosa Souza & Olmos, 1991).
- Dracaena sp.
Dois restos desse Teideo foram achados na
Toca de Cima dos Pilao. © género comporta
atualmente duas especies, D. paraguaiaensis @
D. guianensis, a segunda habita a floresta
amazonica; ela tem costumes anfibios e se
alimenta de moluscos de agua doce, peixes e
anfibios.
D-Anfibios
Numerosos restos de tamanhos variados
foram recolhidos nos niveis pleistocénicos &
holocénicos da Barra do Antoniao, da Toca de
Cima dos Pildo e do Garrincho, E-Fishes
- Plecostomus auroguttatus (Natterer &
Heckel)
A specimen of this small Loricarlidae was
discovered, without the head, mummified in the
layer 2 of Sector A of the Barra do Antonio,
The species is currently frequent in the water
courses of the Northeast (Shuffeldt, 1926;
Fowler, 1954).
IV - Biochronological Comment
The assemblage of mammal fauna
corresponds to approximately fifty species, of
which thirty is part of the recent fauna of South
America. “As for the fossils, the date of their
appearance is unknown. but these species lived
inthe Upper Pleistocene and disappeared at the
‘ond of this period. This disappearance seems
to have taken place about ten thousand years
‘ago. Various genera present in our sites are
‘considered by L.G, Marshallet al(1984) as good
chronological markers (or the final Pleistocene :
there are Eremotherium, Equus (listed in the
corresponding Table 3 as Deing existent since
the Ensenadian) and Haplomastodon that we
will aggregate to the Catonyx. Other species do
not seem known in South America before the
Lujanian ; these are the Dicotyles, Tayassu and
Mazama, possibly Galea. If their presence is
confirmed in the region, Speothos and
Mixotoxodon should be placed in the same
category. Finally for L.G. Marshall et alii (1984)
‘Scelidodon (in @ broad sense) and without any
doubt the Palaeolama had disappeared from the
temperate climate regions for a long time, ike
in Argentina, but survived until the beginning of
the Holocene in the tropical and intertropical
zones of the continent, particularly Brazil. The
fauna of the Sao Raimundo Nonato
‘archasological area cannot therefore be older
than the Lujanian which corresponds to the upper
Pleistocene, and it is highly probable thet it
belongs to the upper Lujanian. On the other
hand, the large number of individuals plus their
great diversity exclude a more recent date: it
Not probable that 2 fauna from the extreme end
of the Pleistocene would be such varied and
abundant. A detailed study of the evolution of
‘each one of the species in the course of time will
certainly permit the obtainment of more accurate
datings,
Several sites, particularly the Sitio do M
have furnished fauna remains dated from the
beginning of the Holocene, Excluding the
micromammals which are currently under study,
this fauna comprises the following mammals:
3. Claude Guérin et alii
E-Poixes,
Plecostomus auroguttatus (Natterer &
Heckel)
Um espécimen deste pequeno peixe
Loricariidae foi descoberto, sem a cabeca,
mumificado, na camada 2 do setor A da Barra
do Antoniao. A espécie é atualmente frequente
nos cursos ¢'Agua do Nordeste (Shuffeldt, 1926.
Fowler, 1954),
IV - Comentario biocronoiégico
© conjunto da fauna de mamiferos
corresponde a cerca de cinquenta espécies, das
quais trinta fazem parte da fauna atual de
América do Sul. Em relagao aos fosseis, a data
de sua aparicao nao conhecida, mas eles
viveram durante o Pleistoceno superior €
desapareceram no final desse periodo, essa
desapari¢ao parece consumada faz cerca de dez
milanos, Varios géneros presentes nes nossos
sitios S80 considerados por L.G. Marshall et ali
(1984) como bons marcadores cronologicos do
Pleistoceno final: trata-se de Eremotherium,
Equus (0 qual, portanto, foi assinalado na
cortespondente tabela 3, como existente desde
© Ensenadiano) e Haplomastodon aos quais
vamos agregar Catonyx. Outras especies nao
parecem conhecidas na América do Sul entes.
do Lujaniano: ¢80 Dicotyles, Tayassu e Mazama,
e possivelmente Galea. Se sua presenga se
confirmar ne regio, Speothos e Mixotoxodon
devem ser colocados na mesma categoria,
Finalmente, para L. , Marshall et all (1984),
Scelidodon (no sentido amplo) e, sem diivida,
Palacolama tinnam desaparecido faz muito
tempo de regides com clima temperado, como 2
Argentina, mas sobreviveram alé o comeco do
Holaceno nas zonas tropicais e intertropicais do
continente, particularmente no Brasil. A fauna
da rea arquoolégica de Sao Raimundo Nenato
‘nao pode, portanto, ser mais antiga que o
Lujaniano, que corresponde ao Pleistoceno
superior, © & allamente provavel que seja do
LLujaniano superior, Por outro lado. a riqueza em
individuos € sua grende diversidade excluem
uma idade mais recente; néo € provavel que
uma fauna do final do Pleistoceno seje tao
abundanle e variavel. Um esludo detalhado da
‘evolucao de cada uma das espécies no decorrer
do tempo, certamenie permitira precisar essa
datagao.
‘Varios sitios, particulermente Sitio do Meio,
forneceram restos da fauna datados do comego
do Holoceno. Excluindo os micromamitsros,
atualmente em curso de estudo, esta fauna
‘compreende os seguintes mamiferos:
89 FUMDHAMENTOS
Euphractus sexcinctus (Linné, 1758)
Dasypus novemenctus Linné, 1758
Priodontes maximus Kerr, 1792
Mymmecophaga tridaciia Linné, 1758
Kerodon rupestris (Wied, 1820)
“Agouti paca (Linné, 1766)
Dasyprocta sp .
Gandae indet.
Felis sp,
Dicotyles tayacu Linne, 1758
Tayassu pecari (Link, 1795)
Mazama sp.
Allare modern forms even ifsome of them
are nowadays absent from the considered
region.
V-Paleoecological Study.
The palececological significance of
Pleistocene mammals (Tab. 3) in the Sao
Raimundo Nonato region is appreciated because
of what is known on the life pattern of each
species present as well as the fauna association
obtained with the help of a modified variant of
the Fleming-Andrews method exposed by C.
Guerin & M. Faure (1987, p. 816-820). It should
be noted that this utilizes a series of 4 histograms
(fig. 6 and 7) representing the composition of
mammal association from the viewpoint of
systemic (a), mass (b), diet adaptations (c) and.
locomotion adaptations (d). The comparison of
the Pleistocene fauna (fig. 7) with the recent one
(fig. 6) was made against a faunal list given by
M.F, Barbosa Souza & F. Olmos (1991). Man
was not considered,
The list of the recent fauna present in the
region and their ecological features ( the
definition of ‘several classes in the legends of
figure 6) is furnished in Table 4
(Our conclusions are the following
~The actual diagrams (fig. 6) highlights the
absence of Insectivora, @ large number of
Rodents end Carnivora, and a very high
proportion for the taxonomic class entitled
‘Other’. The very small mammals are
numerous, specially because of the Chiroptera
and to the Rodents; the very large ones are
absent. The entomophagous prevail, the
omnivorous are determinedly numerous, the
brachiodont herbivores are relatively rare and
the hypsodont herbivores are few. The great
terrestrial runners are absent and the other
large terrestrial mammals are not frequent, the
aquatic ones are absent , the aerial and the
small terrestrial ones are thoroughly
prevailing. The ecological community thus
90
Euphrectus sexcinotus (Linné, 1758)
Desypus novemcinetus Linné, 1758
Prodontes maximus Kerr, 1792
aga tdactyla Linn, 1758
Kerodon rupestris (Wied, 1820)
Agout paca (Linné, 1766)
Dasyprocta sp.
Canidae indet.
Felis so.
Dicotyles tayacu Linné, 1758
Tayassu pecari (Link, 1795)
‘Mazama sp.
‘Todas 20, portanto, formas atuais, mesmo
algumas delas hoje estéo ausentes na regi
considerada.
V - Estudo paleoecolégico
A signficagao palececoldgica dos mamiferos
pleistocénicos (Tab.3) ria regiao de Sao Raimundo
Nonato é apreciada, por ume parte, em fungao do
que se conhece do modo de vida de cada especie
presente, por ouiro lado, como associagao
feunistica, coma ajuda de uma variante modificada
do método de Fleming-Andrews, exposta por C.
‘Guérin & M. Faure (1987, p. 816-820). Lemoramos
que esta uliliza uma séne ded histogramas (fig. 6
€ 7) representando a composicao da associacao
‘do mamiferos dos pontos de vista da sistematica
(@), da massa (o), das adaptacoes alimentares (c)
das adaplagées locomotores (d). A comparagao
dda fauna pleistocénica (ig. 7) com a fauna atval
(ig. 6) 6 estabetecida a partirda uma lista faunistica
fornecida por M. F. Barbosa Souza & F. Olmos
(1991). Nao foi considerado o homem.
A lista dos mamiferos aluais da regigo com
suas caracteristicas ecolégicas (achar-se-8 a
definigdo das diversas classes na legenda da figura
6) 6 forecida na Tabela 4.
Nossas conclusées sto as seguintes:
= Os diagramas atuais (fig. 6) poem em
‘evidéncia uma ausénciade insetivoros, um grande
numero de Roedores © de Camivoros, e uma
‘proporao muito elevada para a classe taxonémica
denominada *Outros’. Os mamiferos muito
equenos s40 muito numerosos, principalmente
‘gragas aos Chiroptera e aos Roedores; 0s muito
grandes estéo ausentes, Os entomdfagos
dominam, os onivoros sao remarcavelmente
numerosce, of herbivoros braquiodontes eo
relativamente raros e os herbivoros hipsodontes
‘Sao pouce representados. Os grandes terresties
corredores estao ausentes, os outros grandes
terrestres sao pouco frequentes, 03 aquatics
esto ausentes, as formas aéreas © 0s pequenos
terrestres dominam amplamente. A comunidade 3. Claude Guérin et alii
Tabela 3: Caracterizacio ecolégica dos mamiteros do Pleistoceno da area arqueolégica de S40 Raimundo
Nonato
Tae Fat ssa___-Regi
Didelphis elbiveniris A ©
Monodelphis domestica A AB
‘Marmosa sp. A AB
Catonye cuviert A c
‘Scelidedon sp. A c
Erometherim lund A H
Yenocnus cearensis A F
Euphractus sexcinctus A c
Dasypus novercinctus A c
Pampatherium humboldt A F
Hoplophorus euphractus A cs
Panochthus gresiebini A Te 1
‘Giyptodon clavipes a G
Pieronotus parneli A AB
Tonatia bidons A AB
Phyjlostomus hactatus A AB
‘Atibeus jamaicensis A AB
‘Myotis sp, A AB
‘Nolossus riolossiis A AB
Tadarda brasiliensis A AB
Dasmodas rotundus A AB
Holoctilus brasivensis R AB
Paeudorhizomys sienplex R AB
‘kode cf. cursor R AB
Calomys callosus R AB
Oryzomys cf. subfavus R AB
Oryzomys sp. R AB
Thricomys apereoides R AB
‘Kerodon rupestrs R c
Galea spixi R AB
‘aout paca R ce
Protocyon irogicdytes c E
Cordosyan thous ce c
Arctodus brasiliensis c F
‘Arctodis cl. bonaerense c 6
Felis yagouaroundi ce c ce orr
Panthera onca c F c ctu
‘Smiloden populator c o ¢ cru
‘Macrauchenia ct patachonica a # HE Gre
Toxodon sp. A 4 HH ctu
Hapiomastedon wariral Pro 4 HB. ctu
Hippidon bonaerensis Por 6 HH Gte
Hippicion sp. Per s HH ore
Equus neogaeus Per G HH Gte
Dicofyies tejacu ar D ° ctu
Tayassu pecari ar D 0 ort
Palaeclama major Ar c HH Gre
Palaoclams ep. Ar 3 HH Gre
‘Mazama sp. Ar D He crf
Table 3 : Ecological features of the Pleisiocene mammals in the archaeological area of S80 Raimundo
Nonato
1 FUMDHAMENTOS:
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