Papéis Avulsos de Zoologia
MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO
ISSN 0031-1049
PAPÉIS AVULSOS DE ZOOL., S. PAULO 42(14):335-350
08.XI.2002
A NEW SPECIES OF LEPOSOMA (SQUAMATA,
GYMNOPHTHALMIDAE) FROM THE REMNANT ATLANTIC
FORESTS OF THE STATE OF BAHIA, BRAZIL
MIGUEL TREFAUT RODRIGUES1
MARIANNA DIXO2
DANTE PAVAN1
VANESSA KRUTH VERDADE1
ABSTRACT
Leposoma puk, a new species of the scincoides group is described from
the Atlantic forests of southern state of Bahia, Brazil. The new species is characterized by a single frontonasal; elongate dorsal and lanceolate ventral scales
arranged in diagonal rows; weakly striated or smooth scales on top of head; a
longer than wide interparietal, with almost straight lateral margins; 17-18
pores; third supraocular the largest one, wider than long and clearly rectangular in shape; a suture between frontal and frontoparietal scales coincident with
the middle of the third supraocular; and, a black pigmentation in the venter or
along venter margins in males. The discovery of this new species, sympatric
with Leposoma annectans, L. nanodactylus, and L. scincoides ranks the Atlantic Rainforest region of Southern Bahia first in species diversity for the genus.
Zoogeography and speciation of Leposoma of the scincoides group are discussed.
KEYWORDS: Leposoma puk, new species, Brazil, Squamata, Gymnophthalmidae.
1
2
Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Caixa Postal
11.461, CEP 05422-970, São Paulo, SP, Brasil.
Universidade de São Paulo, Instituto de Biociências, Departamento de Ecologia, Caixa Postal
11.461, CEP 05422-970, São Paulo, Brasil.
Trabalho recebido para publicação em 18.III.2002 e aceito em 06.VIII.2002.
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RESUMO
Leposoma puk, uma nova espécie do grupo scincoides é descrita para a
região da mata Atlântica do sul da Bahia, Brasil. A nova espécie é caracterizada
por apresentar: frontonasal simples; escamas dorsais alongadas e ventrais
lanceoladas dispostas em fileiras diagonais; escamas do topo da cabeça lisas
ou levemente estriadas; interparietal mais longa que larga, com margens
laterais quase paralelas; 17-18 poros; terceira supraocular mais larga que
longa e a maior da área supraocular; sutura entre a frontal e frontoparietal
coincidente com a região mediana da terceira supraocular; e machos com
pigmentação negra no ventre ou ao longo da região paraventral. A descoberta
desta nova espécie, simpátrica com Leposoma annectans, L. nanodactylus, e L.
scincoides torna a região da mata atlântica do sul da Bahia a área com a
maior diversidade local de espécies do gênero. Discutem-se alguns aspectos
sobre a zoogeografia e a especiação de Leposoma do grupo scincoides.
P ALAVRAS - CHAVE : Leposoma puk, espécie nova, Brazil, Squamata,
Gymnophthalmidae.
INTRODUCTION
The Atlantic rainforests and their highly diverse and endemic biota are
one of the top five threatened hotspots of the world (Mittermeier et al., 1997,
1999; Myers et al., 2000). Originally extending along the eastern coast of Brazil for about 5.500 km from near 6° to 30° S and covering a total area of
approximatelly 1.000.000 km2, less than 6% of the original coverage remains
today. Despite the enormous amount of forests that have been cut down, species surveys are far from complete and several new species have been discovered during the past few years. Under the present regime of forest disturbance,
discovery of new species contributes to phylogenetic understanding of particular groups and consequently the history of these forests and their relationships
with other habitats in South America.
The gymnophthalmid lizards of genus Leposoma are a conspicuous component of the forest litter herpetofauna of Amazonian and Atlantic forests
(Pellegrino et. al., 1999; Rodrigues, 1997; Rodrigues and Borges, 1997; Rodrigues
et al., 2002). Two assemblages of species are presently known: the parietale and
scincoides species groups. Until now, the highest species diversity in Leposoma
was concentrated in the species of parietale group which ranges from Amazonia
to Costa Rica. Nine species are presently recognized in parietale group and a
maximum of two species have been reported to occur sympatrically (Avila-Pires,
1995; Rodrigues, 1997). The scincoides species group of Leposoma is restricted
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to the Atlantic Forest and until recently the only species admitted in the group
was Leposoma scincoides. More recent surveys in the remaining patches of the
Atlantic forests revealed a much higher diversity in this group: Leposoma
baturitensis (Rodrigues and Borges, 1997) and Leposoma nanodactylus
(Rodrigues, 1997) were described and Leposoma annectans resurrected from
synonymy of Leposoma scincoides (Rodrigues et al., 2002).The revalidation of
Leposoma annectans resulted from the study of a large series of specimens obtained during fieldwork at Una, state of Bahia, in the context of a broad project
(RestaUna), designed to study the comparative response of several biological
groups facing forest fragmentation in northern Atlantic forests (Rodrigues et. al.,
2002). Although not describing it, Rodrigues et al. (2002) made reference to an
adult specimen of a still undescribed Leposoma collected more than 10 years ago
in the Atlantic forests of Bahia (see also Rodrigues, 1990). Fortunately, in the last
field campaign of the RestaUna project another specimen of that undescribed
species was obtained and subsequently, an additional specimen was collected on
an adjacent mountain. Therefore, the description presented here is based on three
specimens. The discovery of this new species raises to five the number of species
of Leposoma known from the Atlantic forests and to four the number of species
in the genus that occur sympatrically in the forests of southern Bahia, placing the
area to that of with the highest local diversity for the genus.
SPECIES DESCRIPTION
Leposoma puk sp.nov.
Holotype: MZUSP 87.959, an adult male from Una: state of Bahia: Brazil
(15°10’S, 39°03’W), collected by Marianna Dixo on 25 February 2000; field
number MD 2592.
Paratypes: MZUSP 66.475, an adult male from São José do Macuco, presently
São José da Vitória (Fazenda Unacau): state of Bahia: Brazil (15°09’S,
39°18’W); collected by Miguel Trefaut Rodrigues on 15. x. 1986, field number
86.7231, and MZUSP 89334, an adult male from Serra do Teimoso: Jussari:
state of Bahia: Brazil (15°09’S, 39°31’W); collected by M. T. Rodrigues, D.
Pavan, M. Dixo and V. K. Verdade on 20.iii.2001, field number MRT 6160.
Etymology: puk derives from the vocabulary of the now extinct Botocudo indians
who formerly lived in the Atlantic forests of State of Bahia, Brazil. It means
“the one who cries”, and here is intended to reflect the loss of the Botocudo
culture, as well as the highly threatened and dwindling present condition of the
Atlantic Rainforests, the remaining habitat of these little forest lizards.
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Diagnosis: A Leposoma of the scincoides group characterized by a single
frontonasal, elongate dorsal and lanceolate ventral scales arranged in diagonal
rows and having: 1) weakly striated to smooth scales on top of head; 2) interparietal longer than wide, with almost straight lateral margins; 3) parietals longer
than wide; 4) 17-18 pores; 4) third supraocular the largest one, wider than long
and clearly rectangular in shape; 5) a suture between frontal and frontoparietal
scales that coincides with the middle of the third supraocular, and, 6) a black
pigmentation in the venter or along venter margins in males.
Description of the holotype: (Figs. 1-2). Snout normal, sligthly acuminate.
Rostral broad, wider than high, contacting first supralabial, nasal and
frontonasal. Frontonasal entire, as wide as long; in broad contact with rostral,
nasal, loreal, and prefrontal, always indenting posteriorly the latter; separating or just touching first supraocular. Prefrontals as long as large, in broad
contact at midline and with first supraocular; indented anteriorly by the
frontonasal and posteriorly by the frontal. Frontal hexagonal, with almost
straight lateral margins; approximately twice as long as broad; indenting posteriorly a pair of frontoparietals. Frontoparietals pentagonal, larger than prefrontals, in broad contact; their midline suture approximatelly the same size
as suture between prefrontals. Interparietal wider and longer than frontal,
with almost straight lateral margins, longer than wide; almost straight posteriorly. Parietals irregularly hexagonal, longer than wide; as large as and as
long as interparietal, always reaching the level of its posterior margin.
Supraoculars four; first smallest, triangular, longer than wide and contacting
loreal. Third supraocular the largest, roughly rectangular, wider than second
and fourth supraoculars that are equivalent in size; all supraoculars contacting superciliaries. Nasal above first supralabial, large, longer than wide, with
the nostril in the center. Nasal semidivided superiorly or divided at the level
of posterior margin of nostril. Ventral margin of nasal slightly convex, dorsal
margin straight. Loreal narrow, posterior to nasal, diagonally and ventro-posteriorly oriented; contacting nasal, internasal, first supraocular and first
superciliar, a small preocular granule, and frenocular. Frenocular contacting
nasal and followed by a series of four infraorbital elongate scutes; the first
one quadrangular, smaller than frenocular, the second, third and fourth elongate, the fifth enlarged, quadrangular, and followed by a temporal scute with
about the same size. Six supralabials, first and third the longest, fifth highest,
third and fourth bellow eye. Sixth supralabial followed by a granule much
smaller than last supralabial and separated from the tympanum by two similar
granules. Five superciliaries, first largest, sligthly larger than first supraocular,
expanded both on lateral and dorsal surface of head and in broad contact with
loreal and a small granule placed between the latter and frenocular. Other
superciliaries elongated, the last being the largest one. Eyelid with semitrans-
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339
Figure 1. Dorsal, ventral, and lateral view of head of the holotype of Leposoma puk from Una, state
of Bahia, Brazil (MZUSP 87959).
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Figure 2. Dorsal and ventral view of a paratype of Leposoma puk (MZUSP 66475) from São José
do Macuco.
parent disc formed by two enlarged scales. Temporal region almost all covered with smooth or slightly keeled and juxtaposed scales that are larger around
tympanum, around parietals and surrounding postocular area. Lateral surface
of neck with conical and juxtaposed keeled granules, irregularly arranged in
transversal rows. Ear opening bordered by a series of very small, smooth,
granules; tympanum distinct, subovoid. Scales of the top of head with very
slight, low relief irregular longitudinal striations; almost smooth when compared to most of other Leposoma.
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341
Mental broad, wider than long. Postmental single, contacting first
infralabial. Three pairs of enlarged genials, all contacting infralabials; first pair
the smallest one, second largest, both in broad contact at middline, and in contact with infralabials. Third pair of genials wider than long, separated medially
by two elongate granules. An additional series of relatively large, diagonally
oriented and flat chin-shield like scutes follows the genials. Mental, postmental
and genials smooth. Infralabials five; first largest, followed by second in size.
Scales immediately posterior and lateral to third pair of genials, juxtaposed,
smooth, flat. From there they gradually change to lanceolate, imbricate and
keeled, like posterior gulars. Gulars variable in shape, in nine transverse rows;
anterior rows smooth, rounded and juxtaposed, changing progressively to lanceolate, imbricate, keeled and mucronate. Collar fold indistinct.
Dorsal scales large, strongly keeled, mucronate, lanceolate, and imbricate anteriorly, becoming wider and laterally juxtaposed with almost straight
lateral sides at midbody; 28 regular transverse rows between interparietal and
posterior level of hind limbs. Lateral scales resembling dorsals but more lanceolate; changing progressively to ventrals except for an area with small,
rounded, almost smooth and juxtaposed conical granules around arm level. Groin
scales identical to flank scales except for their smaller size. Twenty seven scales
around midbody. Ventrals leaf-shaped, keeled, mucronate, imbricate, in 21 regular transverse and diagonal rows from interbrachials (included) to preanals.
Figure 3. General view of Serra do Teimoso: Jussari: state of Bahia. The primary forests on the top
are the habitat of Leposoma nanodactylus, L. puk, and L. scincoides.
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Posterior margin of the vent with five scales; central and lateral scales the smallest, paramedials the largest. Total pores 18; 4 of them preanal.
Scales of tail imbricate, keeled, in complete rings that became more regular ventrally. Near the base of tail dorsal and lateral tail scales are more strongly
keeled and wider than ventral tail scales, becoming gradually identical towards
the extremity.
Limb scales keeled and imbricate, except on ventral surface of brachium
and on posterior surface of thigh which are mostly subimbricate or juxtaposed,
sometimes granular. Palmar and plantar surfaces with small, conical granules.
Subdigital lamellae double, 13 on Finger IV and 17 on Toe IV. Fingers and toes
clawed, with the following relative sizes: 1 < 2 = 5 < 3 < 4 and 1 < 2 < 5 < 3 <
4, respectively.
Dorsal surface of body and tail light brown with irregular dark brown or
black spots. Dorsal surface of head lighter. A dorsolateral light stripe one to one
and half scale wide extends from lateral part of parietal scale, converge to occipital region and run to third part of tail. The dorsolateral stripe is more conspicuous anteriorly, fading progressively towards midbody. An irregular longitudinal dark brown to black area one to three scales size marginate inferiorly
the dorsolateral light stripe. The dorsolateral stripes delimit a midorsal light
brown area of about 4 scales wide with the characteristic dark brown reticulate
background. Flanks and lateral surface of tail slightly darker than back with a
few small light spots more conspicuous on neck. Dorsal surface of head lighter
with scattered dark brown reticulation. Lateral surface of head darker, coloured
as flanks. Ventral parts of head and body cream with a very conspicuous reticulate pattern of irregularly disposed dark spots. The dark reticulation is more
concentrated in the ventrolateral area. In the ventral part of tail only the anterior
third remains with some light pigmentation, rest of tail almost entirely black.
Limbs dark brown dorsally, mottled with cream; ventrally yellowish cream,
like in the ventral parts of body and tail.
Left hemipenis everted at preservation; bilobed, with short lobes. Sulcus spermaticus medial, bifurcating centripetally towards the apex. Right and
left lobes symetric with two longitudinal rows of 20-22 enlarged spines on the
apex of naked W-shaped flounces.
Measurements of the holotype: Snout-vent lenght: 38 mm; tail lenght 53 mm.
Comparisons: species of genus Leposoma are separated in two distinctive groups:
parietale and scincoides (Rodrigues, 1997; Rodrigues et al., 2002). Since originally proposed by Ruibal (1952) their content varied due to the description of
new species but their similarity based definition stands until now. Species of
the scincoides group (annectans, scincoides, nanodactylus and baturitensis)
have elongate dorsal and lanceolate ventral scales that are characteristically
Vol. 42(14), 2002
343
arranged in diagonal rather than in longitudinal rows. In the parietale group,
dorsal scales are wider and shorter than those present in species of scincoides
group, and ventral scales are arranged in regular longitudinal rows. Leposoma
puk, is a member of the scincoides group. Leposoma puk, Leposoma
nanodactylus and Leposoma baturitensis are unique in the scincoides group in
presenting an entire and smooth frontonasal and weakly striated or smooth scales
on head (longitudinally divided and highly striated in L. annectans and L.
scincoides). Leposoma puk, Leposoma nanodactylus and Leposoma baturitensis
are also the only species in the scincoides group presenting a characteristically
black pigmentation in the venter or along venter margins in males (venter cream,
immaculate in L. annectans and L. scincoides). Leposoma puk, nanodactylus,
and baturitensis can be separated by the following list of characters.
1) Supraoculars: in Leposoma puk the third supraocular is the largest one, wider
than long and clearly rectangular in shape; in L. baturitensis the second
supraocular is the largest, which is squared. In L. nanodactylus, the scale corresponding to the first supraocular of baturitensis and puk is characteristically
divided in two scales. This implies that the largest supraocular in L. nanodactylus
(the fourth), which is also squared, is actually homologous to the third of
nanodactylus and baturitensis.
2) Frontal: frontoparietal suture: in Leposoma puk the suture between frontal
and frontoparietal scales coincides with the middle of the third supraocular. In
L. baturitensis and L. nanodactylus the suture always reaches the anterior third
of the third supraocular (actually the fourth in L. nanodactylus)
3) Shape of dorsal scales: in Leposoma puk and L. baturitensis midbody dorsal
scales are wider than the anterior ones and are laterally juxtaposed with almost
straight lateral sides. Dorsals in L. nanodactylus are always elongate, lanceolate,
imbricate.
4) In Leposoma puk, L. baturitensis and L. nanodactylus the fingers III and IV
have approximately the same comparative size but in L. puk and baturitensis
the size difference between the referred fingers corresponds to less than the
equivalent size of the distal supradigital lamellae; in nanodactylus to two of
those scales.
5) Leposoma puk has 17-18 total pores; there are 9-11 in baturitensis and 10 in
nanodactylus.
6) Leposoma puk additionally differs from L. baturitensis by slight differences
in the number of dorsal and ventral scales, higher counts of scales around mid-
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body, and of fourth finger infradigital lamellae, and from nanodactylus by a
lower number of dorsal scales and higher counts of IV finger and IV toe
infradigital lamellae (Table 1). Data for L. nanodactylus includes data for two
specimens (one female) recently obtained at Serra do Teimoso. Table 1 also
presents comparative data for other species of the scincoides group.
DISCUSSION
As usual in the Atlantic Forest domain at this latitude, most of the original forest was disturbed for cacao plantations. The paratype and first specimen
of Leposoma puk was collected at São José do Macuco. It was active in the
litter of a cacao grove where only large trees from primary forest (up to 30 m
tall and over approximately 50 cm diameter at breast height) had been preserved to shade the understory cacao plants. At the time, the mountainous area
had been recently disturbed for cacao planting and was adjacent to a hilly region covered with primary forest. The holotype was obtained in a pit-fall trap
located in cacao groves at the Una Reserve, in the same general area but characterized by lower reliefs and located a few kilometers away. The third specimen was also collected in the same area but at Serra do Teimoso, a mountain
reaching approximatelly 1000 meters altitude. Most of the southern face of the
mountain has been preserved as a private reserve (RPPN Serra do Teimoso)
and is still covered with the original forest. At the base of the mountain the
forest is less diverse and humid than at the peak which is dominated by large
trees (up to 30 m tall and approximately 1.0 m DBH), and characterized by
higher abundance of palms, ferns, bromeliads and more leaf litter than the forest situated at lower altitudes. The specimen was obtained in a pit-fall trap
located at about 900 m altitude, near the top of the mountain dominated by
primary rainforest. Maximum strait-line distances among the three localities is
less than 30 kilometers.
Table 1. Condition of frontonasal scale, number of Dorsals, Ventrals, Scales around body (SAB), of
total pores, and IV Finger and IV Toe infradigital lamellae in species of Leposoma of the scincoides
group. Data were taken from specimens examined (Appendix I), and from Rodrigues, Dixo and
Accacio (2002).
Species
annectans
scincoides
baturitensis
nanodactylus
puk
Frontonasal
Dorsals
Ventrals
SAB
Pores
IV Finger
IV Toe
divided
divided
single
single
single
26-30
29-36
25-27
31-34
26-30
16-20
17-23
17-18
16-20
20-22
21-27
24-29
24-26
27-32
27-30
10-13
14-20
9-11
10
17-18
9-12
9-15
11-12
7- 9
13-14
12-16
14-20
16-18
10-12
16-19
Vol. 42(14), 2002
345
The Una area is characterized by low hills dissected by medium to small
size streams. This relatively homogeneous landscape is interrupted only by a
series of isolated, higher mountains like Serra do Teimoso. In contrast to the
lower altitude areas used for growing cacao, these mountains are still partially
covered with the original primary forest (e.g. Serra das Trempes, Serra do Padre, Serra da Lontra or Serra do Teimoso). Additional data on habitat and miscellaneous information on the local herpetofauna was presented previously
(Rodrigues et al., 1989; Rodrigues, 1997; Rodrigues et al., 2002).
Four species of Leposoma occur sympatrically in this region, the highest diversity locality so far reported for the genus. Most of the specimens were
collected in pit-falls traps during field work of the RestaUna project. Table 2
shows capture data for the four species in three field campaigns during the
year in a low relief area (about 100 m altitude) near the Una reserve (Rodrigues
et al., 2002). Leposoma scincoides was the most abundant species, followed
by L. annectans; Leposoma nanodactylus and Leposoma puk were extremely
rare. Table 2 also includes comparative data for species of Leposoma collected at Serra do Teimoso. Leposoma scincoides is the only widespread species, distributed from the state of Bahia to the state of Rio de Janeiro; the other
three species are restricted to the state of Bahia: L. nanodactylus and L. puk
are found in the area around Una, with L. annectans probably extending to
Salvador.
Why is Leposoma scincoides so widespread and locally abundant in
Atlantic Forests, whereas L. annectans is locally common and the other two
sympatric species are so rare? Why do four species of Leposoma occur sympatrically in the general area of Una and not elsewhere? We suggest that the distribution of these species probably reflects historical patterns.
South American tropical forests have fluctuated in time and space, especially during the Quaternary (Haffer, 1979; Vanzolini and Williams, 1970) giving rise to the “refuge hypothesis” of animal and plant distributions in the
Amazon. Biological and geomorphological evidence for Quaternary forest flucTable 2. Number of specimens of L. scincoides, L. annectans and L. nanodactylus collected at
Una, Bahia with pit-fall trapping (sets of four 30 liter buckets connected by 8 meter drift fences)
during three field campaigns. Pit-fall effort was equivalent to 2592/pit-falls/day in the first campaign
and of 5184/pit-falls/day for the other two. Comparative data for Serra do Teimoso are also shown;
pit-fall effort there was 240/pit-falls/day.
Species
L. puk
L. annectans
L. scincoides
L. nanodactylus
January-February
1999
October-November
1999
0
14
46
0
0
23
80
3
January-February
Serra do
2000
Teimoso 2001
1
26
84
2
1
–
6
2
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tuations in the Atlantic forest domain is vast (Ab’Saber, 1977, 1982; Haffer,
1979, 2001; Heyer and Maxon, 1983; Jackson, 1978).
The “brejos”, isolated forest islands located in the semiarid Caatingas of
northeastern Brazil provide textbook examples of refuges (Vanzolini, 1981;
Vanzolini and Williams, 1981; Rodrigues, 1990; 1996). Their relict forest-adapted
fauna persists from eras when forest was continuous between Atlantic forest
and the “brejos” of today. Presently, vestiges of similar forest patches not larger
than 300 km2 occur within the semiarid Caatingas; a region that spans approximately 800,000 km2 (Ab’Saber, 1977). One example is Serra de Baturité, a mountain complex in the State of Ceará, and the type locality of Leposoma baturitensis
the only extralimital species of the scincoides group of Leposoma known. From
the top of the highest Baturité mountain one can see that forest is restricted to the
mountainous area. Other physionomically similar scenarios exist in the Caatingas
of northeastern Brazil as a testimony of times when, under more humid climates,
these areas and Atlantic forest were possibly connected.
If we examined early periods in the Quaternary characterized by higher
humidity than present day and look at the prevalent landscape physiognomy,
the relictual condition dissappears. At those times, Serra de Baturité would
have been connected by forest to the surrounding area. What would have been
the distributional limits of Leposoma baturitensis in those times? Would it have
been restricted to Baturité mountains or widespread around forest areas surrounding them? An affirmative answer to both questions is possible. Restriction of Leposoma baturitensis to the Baturité mountains could be explained by
recent differentiation in isolation and/or by historical contact with another widespread generalist congeneric species that despite the continuous forest coverage prevented expansion of its distribution. As there are no vestiges of a putative relative we could hypothesize that it became extinct when, following climatic deterioration towards present, the forest dwindled and disappeared from
surrounding areas. Under the second possibility, the same arguments could explain disappearance of a more widespread baturitensis from the adjacent areas
today dominated by the semiarid Caatingas.
As in the Baturité example, we may use similar logic to explain the
history of Atlantic forest in the study area. The aridity that progressively affected northeastern Brazil left a formerly continuous forest fauna isolated in
hilly areas surrounding by Caatingas (Oliveira et al., 1999). Forested mountains like Serra das Trempes, Serra dos Padres, Serra do Teimoso, or Serra da
Lontra, may have acted as forest refugia in the recent past. Presently, they are
what we are here calling concealed refuges because they remain as yet unrecognized. This is because under the present humid climate the general green
coverage of these mountains are simply recognized as part of the Atlantic forest
region, a large core area described mainly on geomorphologic grounds, rather
than faunal attributes.
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347
Considering the rarity of Leposoma nanodactylus and Leposoma puk in
the lowlands surrounding Una, we hypothesize that these species may have
differentiated and remained isolated in some of these concealed refuge areas.
Both the widespread and generalist Leposoma scincoides and the relatively
common Leposoma annectans might outcompete these rare species in forest
surrounding the concealed refuge areas.
Presently, even after climatic change and intensive exploration for
wood, agriculture, mining, and cattle ranching, has dramatically altered the
landscape, the contrast between open and formerly forested areas remains
evident in the semiarid Caatingas of northeastern Brazil. Evergreen forested
areas are visually recognized as islands in the grayish-brown dry Caatingas.
This is why putative refugia or centers of differentiation of forest fauna in
the semiarid Caatingas were first surveyed and should receive priority for
conservation. Their insularity and endemisms are so obvious that they are
obvious candidates for conservation priority (Conservation International,
1994). In contrast, several important probable refuges are overlooked because they are part of an otherwise homogeneous habitat. These concealed
refuges and their limits can only be detected with intensive biogeographical
study.
The highly endemic sand dune area of Rio São Francisco in the core
area of semiarid Caatingas of State of Bahia is now recognized as a concealed
refugium (Rodrigues, 1991a,b,c,d; 1993a,b; 1996). We suggest that the relictual
mountains near Una in the Atlantic Forest of State of Bahia provide other examples of concealed refugia. Our suggestion could be tested with additional
field work in the region complemented by molecular phylogeographic studies.
If our hypothesis is confirmed, we hope there will be enough time left to take
adequate conservation measures before the remaining forest habitats and their
associated fauna are destroyed.
ACKNOWLEDGEMENTS
We thank Fundação de Amparo à Pesquisa do Estado de São Paulo
(FAPESP), Conselho Nacional de Desenvolvimento Científico e Tecnológico
(CNPq), and Financiadora de Estudos e Projetos (FINEP) for support. For their
invaluable help in the field we thank A. Langguth, E.X. Freire, C. Bonvicino,
M.N. Stevaux, the staff of Fazenda Unacau and of RestaUna project, especially
R. Pardini and D. Faria, and H. and L.V. Carvalho of the Reserva Particular do
Patrimônio Natural (RPPN) Serra do Teimoso. We also thank L. Seky for valuable help with Botocudo vocabulary, Luciana Lobo for the drawing, and B.
Zimmerman and A. Chek for reading the manuscript and making invaluable
suggestions for its improvement.
348
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ADDITIONAL SPECIMENS EXAMINED
Leposoma nanodactylus – Brazil: Bahia: Una: MZUSP 87955-87957, 8798687987; Serra do Teimoso: Jussari: MZUSP 89332-89333.
Leposoma baturitensis – Brazil: Ceará: Serra de Baturité: Pacoti (Sítio Barbosa)
MZUSP 79378; Pacoti (Sítio São Gonçalo do Freire) MZUSP 79379.
Leposoma annectans – Brazil: Bahia: Una: MZUSP 87792-87804; 8785187874; 87958.
Leposoma scincoides – Brazil: Bahia: Una: MZUSP 87805-87050, 8787587954; Serra do Teimoso: Jussari: MZUSP 89324-89329.
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