Table Of ContentCOMPOSITION OF SEASONALLY DRY TROPICAL FOREST
FLORISTIC
FRAGMENTS CENTRAL NORTHEASTERN
BRAZIL
BAHIA,
IN
Domingos de
Cardoso Luciano Paganucci Queiroz
Benicio
Oliveira Silva
Herbdrio da Universidade Estadual Universidade Estadual de Feira de Santana
da
de de Santana (HUFFS) Depto. de Ciencias Biologicas, Herbdrio
Feira
Km
03-BR Campus. 44031-460 Universidade Estadual de de Santana (HUEF§)
Feira
16,
7
Km
de Santana, BRAZIL 03-BR Campus. 4403 1-460
Feira Bahia,
16,
1
ABSTRACT
Dry F
Tropical
nally
Dry
Seasonally Tropical Forests (SDTF) are one of the v
97%
observed dry remnant;
(2006) that of these forest
ca.
of different anthropogenic threats, such as habitat fragmentation, climatic change, or conversion into
fire,
SDTF
from and
cultivated However, have received attention conservationists ecologists
areas. relatively
little
humid
comparison (Pennington 2006b).
in
to forests et
al.
known SDTF
although
presently about the diversity of neotropical in recent years
Little floristic
is
there has been an increasing interest in this vegetation type (Gentry 1995; Giulietti et 2006; Pennington
al.
SDTF
2006a). However, the growing knowledge of the has stimulated the formulation of various
flora
et
al.
&
hypotheses concerning the historical biogeography of this vegetation (Prado Gibbs 1993; Pennington et
and com-
2000, 2004; Prado 2000; Lavin 2004; Queiroz 2006a) the geographical distribution of
et its
al. al.
SDTF
ponent species, and studies have indicated that are independent phytogeographical units distributed
disjunctly through South and Central America (Pennington al 2000; Prado 2000).
et
SDTF
comprehend
According to the definition elaborated by Pennington et (2000), the neotropical
al.
different physiognomies, and can include both forests as well as more open vegetation types where shrubs
and and Euphorbiaceae
predominate. bromeliads succulent plants belonging Cactaceae are
Terrestrial to
Research of Texas
Journal of the Botanical Institute 2(1)
552
common SDTF component apparent only during the short rainy
an ephemeral herbaceous
well
in areas, as as
SDTF markedly phenology flowering
seasonal of
Another important aspect characterizing their
season. is
between dry
and with many species demonstrating synchronous flowering at the transition the
fruiting,
(Machado
and rainy seasons 1997).
the et
al.
SDTF
The Caatinga domain constitutes the largest and most isolated nucleus of the (Queiroz 2006a).
Km
occupies 800,000 2 in the northeastern section of Brazil (Ab'Saber 1974) with a semi-arid climate
ca.
It
Queiroz
mountain
mainly in the low altitude areas between extensive plateaus and/or ranges (Prado 2003;
domain comprised physi-
2006a). Andrade-Lima (1981) noted that the vegetation of the Caatinga different
These
ognomies, including areas of open vegetation dominated by shrubs or areas with relatively tall forests.
and
commonly dry deciduous
or
seasonally dry forests are referred to as arboreal caatinga, forests, forests,
from
northeastern principally
occur highly fragmented areas throughout the semi-arid region of Brazil,
as
dry
Bahia through northern Minas Gerais In spite of the fact that these tall forest areas are situ-
State State.
domain Andrade-Lima workers have indicated that
different
within Caatinga (Rizzini 1979; 1981),
ated the
domain
they may in be phytogeographically more related to the dry vegetation of the Atlantic Forest
fact
&
may
Amorim
2005, 2006), or represent dis-
Fontes 2000; al 2005; Oliveira-Filho et
(Oliveira-Filho et al.
SDTF humid (Pennington
Coastal Forests
without any the areas of the Atlantic
junct areas of relationship to
and
2000; Prado 2000). The deciduous and semi-deciduous forests of southern southeastern Brazil,
et
al.
with the moister of Atlantic
example, have been demonstrated have strong relationship sites
to a floristic
for
domain
Caatinga
dry included in the
Coastal Forest and therefore should not be treated as a type of tall forest
&
more
have
However, (2006)
Oliveira-Filho
(Oliveira-Filho Fontes 2000; Oliveira-Filho et 2005). et al.
al.
SDTF would same deciduous and semi-deciduous
these
recently proposed wider definition for that treat
a
domain SDTF.
of the Atlantic Forest as another nuclei of
forests
SDTF
As problems be resolved concerning the nature of in eastern
such, clear that there are to
is still
it
SDTF by Penington (2000) and Prado
South America. The delimitation of complicated the fact that et al.
is
on hand,
humid while Oliveira-Filho (2006), the other
(2000) did not include forests in their analyses, et al.
knowledge concerning
the
sampling Caatinga Additionally, the floristic
included only very limited of areas.
a
many SDTF
very
composition areas of in northeastern Brazil incipient.
of is still
fragments seasonal dry in
As present study undertook survey of several of forests
such, the a floristic
dry be
do Orobo Mountains Bahia The relationships of these forests will
the Serra in central State. floristic
we
phy SDTF. intend
togeography
discussed here in terms of the present understanding of the of Specifically,
Orobo Mountains the
demonstrate that the dry forest framents of the Serra do are floristically related to
to
SDTF
SDTF we broader (sensu Oliveira-Filho
To account this goal, will treat the as a definition et al.
floras.
domain from Caatinga
nuclei the
2006) encompasses the dry of the Atlantic Forest as a distinct
that sites
Orobo mountain with former
domain and do range these nuclei.
discuss the relationships of the Serra
Study Area
Ruy and
Barbosa Itaberaba in
The do Orobo mountain range located within the municipalities of
Serra
is
- and 40°19' - 40°30'W) The range occupies an area of
Bahia (12°15' 12°25'S (Fig.
central Brazil
State, 1).
m
maximum (BAHIA The
region has semi-
approximately 7,300 ha and has elevation of 1,014 2002). a
a
arid climate (SEI 1998), and although there are no systematic rainfall records for this area, estimates of
mm
The
800
annual approximately (SEI 1998). aver-
precipitation in the region place the average rainfall at
minimum
maximum
°C
annual temperature 26 with an absolute of 33 °C, and an absolute of 18
age °C,
is
by
sometimes covered
The fragments examined have shallow that are a significant
soils
(SEI 1998). forest
pH
main and moderate high
group present
layer of leaf These soils are classified in the "luvisoil" to
litter.
more
(IBGE/EMBRAPA comparatively than those of
they
and nutrient 2001). In addition, are fertile
levels
nearby
the savannas.
domain
The do Orobo mountain range located within the Caatinga of northeastern Brazil
Serra
is
(Velloso 2002), and several vegetation types can be found there (Fig. 2) according to general altitudi-
et
al.
common
A while occur
shrubby with palm most lower altitudes, forests
nal gradients. caatinga trees is at
Cardoso and de Queiroz, Seasonally dry forest fragments in Bahia, Bi
The localization of the Serra do Orobo Mountains, near the Chapada Diama 1a, central Bahia State, Brazil.
Fig. 1
.
found on
Dry and
450 and 850 fragments riparian forests are
higher (between meters). forest
altitudes
at
humid summit on
more near the
between 450 and 700 There area the
meters. a
the slopes elevations is
at
humid Above
submontane the zone,
700 and 850 harboring forest
southern (between meters) a forest.
slope
with popula-
and surrounding mountain peaks up the 900 meters, a savanna-like vegetation large
the to is
&
Near summit, rock outcrops harbor campo
Ayensu the the
L.B.Sm. (Velloziaceae).
tions of Velloziajurcata
(rocky vegetation.
rupestre
field)
400 European
Orobo anthropogenic in years of
Although do has suffered strong alterations
the Serra
These fragments have extremely high
some fragments well-preserved.
are relatively
colonization, forest still
some fragments of dry forest in northeastern Brazil.
conservation value as they represent of the last intact
minima,
many new belonging groups, such as the fungi (Neojhonstonia
species to different
Additionally,
&
been
Bravo 2006) have described
Gusmao Grandi psychodid (Psychoda serraorobonensis, et
al.
2001), flies
and
were
some new from campo vegetation identified
the rupestre
from species of plants
Besides,
this area.
&
new
comm.; two
Queiroz,
Fabaceae-E.R.Souza L.P.de pers.
are in process of publication (Calliandra,
Journal of the Botanical Research Institute of Texas 2(1)
Fig. 2. Different forest physiognomies found in the Serra do Orobo Mountains, Bahia State, Brazil. A. A fragment of semideciduous forest; B. Riparian
forest extending almost to the top of the mountain, with Eriotheca globosa in bloom; C. A fragment of humid forest covered by fog; D. Interior of
cf.
humid many
the showing
forest, epiphytes.
remnants
In 2002, the
forest as
range were
assigned the status of
si
and
conservation
to biodiversity rese
METHODS
A
floristic survey was carried out from August 2004 to June 2006 in the best preserved sites of semidecidu-
and submontane humid The
ous, riparian, fragments. vegetation used here each
forest classification for for-
est fragment follows Veloso (1991) and stresses the distinct physiognomies and ecological conditions
et
al.
km
observed in each. was studied 8 fragments of semideciduous forests (2.85 2 6 of riparian forests (0.37
It
),
km
km
2 2
and one submontane humid
of forest (0.07
)
).
During
the course of the present study a total of 15 field trips were carried out, each lasting from three
Of humid
to four days. the 15 studied fragments, the submontane one was the unique where our
forest
all
was due and was
collect effort lower to difficult access because discovered only recently. All
its fertile
it
material of vascular plants was collected according the standard methods described by Filgueiras
to et
al.
555
Cardoso and de Queiroz, Seasonally dry forest fragments in Bahia, Brazil
A
some were made species never encountered with flowers or fruits.
In collections for
(1994). cases, sterile
made de Santana University
voucher were and subsequently stored the Feira State
of ,385 collections at
total
1
Herbarium (HUEFS). In some cases, species that were very difficult to collect were included in the checklist
Appendix
they could be positively identified in the field; these are indicated as "Not collected" in 1.
if
HUEFS
specimens were by the authors with the aid of identified collections stored at the
All identified
by Specimens only genus or family, are clearly labeled in
and/or using specific bibliographies. identified to
confirm
the Appendix. Duplicates were sent to specialists in order to provide or to identifications.
APG
and
pteridophytes
the
Angiosperm organized according (2003) in the floristic
families are to list,
II
Brummit and Powell
according
abbreviated to
:
RESULTS
do Orobo mountain range comprises 615 species
The vascular flora of the forest vegetation of the Serra
by 597 365
and (Appendix Angiosperms represented species in
belonging 378 genera 106 families are
to 1).
A
and
genera and 96 The pteridophytes included 18 species belonging to 13 genera 10 families.
families.
some
561 were species rank (91.2%). However, in groups (especially the Myrtaceae),
of taxa identified to
total
rank
many 8.3% or family
taxa were identified only to the genera (51 specimens, or of the total collection)
taxonomic
due
specimens, 0.5%) their difficulty.
to
(3
and 87
while
The semideciduous demonstrated the greatest species richness (465 spp.), 141
forests
humid Considering
submontane
species were sampled in the riparian and in the forests, respectively. all
by herbs
commonest form was followed vines
fragments the trees (182 species), (138),
together,
forest life
The most were Fabaceae spe-
and diverse families (95
shrubs subshrubs epiphytes
(35).
(111), (104), (45),
Bignoniaceae
Rubiaceae Asteraceae
Apocynaceae Orchidaceae
Euphorbiaceae (22),
(24), (24),
cies), (38), (24),
48% number
Myrtaceae Malvaceae and Rutaceae which together comprised of the total
(15),
(19), (18), (17),
by
were only
Orobo mountain represented a
of species surveyed in the Serra do range. Thirty-five families
Senna Eugenia
The most species-rich genera were Croton (11 species), Passiflora (7),
single species. (8), (8),
and
Myrcia Solanum Uiconia Bauhinia Senegalia
(5).
(7), (7), (6), (5),
new
These
included:
by being taxa
were specialists as (Fig.
Several species identified different 3).
&
and
Queiroz 2006b),
Phanera (Fabaceae,
(Moraceae; Castro Rapini 2006), trichosepala
Dorstenia caatingae
comm.), Conchocarpus
(Fabaceae-L.P.de Queiroz, pers.
undescribed species belonging to the genera Cratylia
(Commelinaceae-L.Aona, comm.), Eugenia (Myrt-
comm.), Dichorisandra pers.
(Rutaceae-j.R.Pirani, pers.
comm.), Solanum (Solanaceae-
(Gesneriaceae-A.Chautems,
comm.), pers.
aceae-M.Sobral, Sinningia
pers.
&
and
comm.), Spermacoce
Jardim,
(Rubiaceae-J.G pers.
Queiroz D.Cardoso, comm..), Standleya
L.P.de pers.
number
(Rubiaceae-E.B.Souza and E.L.Cabral, pers. comm.). Additionally, of the total of species collected,
Orobo dry
12 were considered be endemic the dry forest fragments of Serra do or to the adjacent forest
to to
Appendix
marked by an
These species are asterisk in the
areas. 1.
new were time in Bahia including
other taxa collected for the State,
In addition these species, first
to
and
(Oxalidaceae)
Bulbophyllumsanderianum (Orchidaceae), Croton warmingii (Euphorbiaceae), Oxalis roselata
(Solan-
(Boraginaceae), Heteranthia decipiens
Other interesting collections included Cordia blanchetii
(Fig.
3).
known
from
and which were only previously their type material
bahiensis (Passifloraceae),
aceae),
Passiflora
uncommon
and both very sapro-
collected in Bahia; as well as Pogoniopsis schenckii Wullschlaegelia calcarata,
phytic orchids
(Fig.
3).
was canopy
continuous
The were composed plant In general there a
fragments of different strata.
forest
composed (Apocynaceae), Cariniana (Lecythidaceae),
20 meters oiAspidospermaparvijoiium estrellensis
high,
ca.
lis-
(Moraceae),
(Euphorbiaceae), Ficus pulchella
Centrolobium tomentosum (Fabaceae), Cnidoscolus oligandrus
membranacea
Nectandra (Lauraceae),
(Chrysobalanaceae),
blanchetiana (Olacaceae), Hirtellaglandulosa cf.
teria
and
Piptadenia paniculata (Fabaceae),
(Fabaceae),
Myrsine (Myrsinaceae), Peltogyne
ferruginta conjertiflora
30-45 and
up included Eriotheca
Emergent could reach meters,
Vantanea compacta (Humiriaceae). trees to
Journal of the Botanical Research Institute of Texas
2(1)
F
globosa (Malvaceae), Caesalpinia leiostachya (Fabaceae), Cavanillesia arborea (Malvaceae), Ficus cyclophylla,
cf.
and
gomelleira (Moraceae), Goniorrhachis marginata (Fabaceae), Vochysia (Vochysiaceae), Uanilkara
sp. rufula
(Sapotaceae). Several vine species, mostly belonging to the families Apocynaceae, Bignoniaceae, Fabaceae,
and
Passifloraceae, Sapindaceae, occur in the canopy, within treefall gaps, or along the edges of the forest
fragments. The subcanopy was dominated by species of Euphorbiaceae, Fabaceae, Myrtaceae, Melastoma-
and
Rubiaceae, Rutaceae.
taceae,
The submontane humid was
forest fragment the richest area for epiphytes, including Anthurium
pentaphyllum, A. scandens (Araceae), Nematanthus albus (Gesneriaceae), Oncidium cUiatum, O.flexuosum (Or-
Peperomia and
chidaceae), (Piperaceae), Tillandsia (Bromeliaceae), Vriesea simplex (Bromeliaceae).
sp. stricta
humid
submontane had composed
This forest fragment also a dense herbaceous stratum of sciophilous
herbs such as Paepalanthus spatulathus (Eriocaulaceae), Sinningia barbata (Gesneriaceae), Begonia
petasitifolia
(Com-
(Begoniaceae), Dichaea congniauxiana (Orchidaceae), Sobralia (Orchidaceae), Gibasis geniculata
sessilis
and
melinaceae), Ichnanthus grandifolius (Poaceae), Ruellia (Acanthaceae).
affinis
557
Cardoso and de Queiroz, Seasonally dry forest fragments in Bahia, Brazil
commonest and were
The species found at forest edges in disturbed sites the trees Albizia polycephala
and Madura (Moraceae), the shrubs Diospyros inconstant (Ebenaceae), Lantana camara
(Fabaceae)
tinctoria
(Verbenaceae), and Senna (Fabaceae), the vines Arrabidaea cinerea (Bignoniaceae), Chaetocalyx
aversiflora
and
scandens (Fabaceae), Dicella bracteosa (Malpighiaceae), Macroptilium erythroloma Fabaceae), Periandra coc-
cinea (Fabaceae), and the herbs Chaptalia integerrima, Emilia josbergii (Asteraceae), Pavonia martii (Malvaceae),
and
(Lamiaceae).
Hyptis pectinata, H. suaveolens
Some such
tree species present in the best preserved forest fragments, as Galksia integrijolia ("pau-d'alho",
and used medicines. Additionally,
Phytolaccaceae) Caesalpinia leiostachya ("pau-ferro"), are locally as folk
we grown wood,
found evidence of fully trees being harvested for their including Copaifera langsdorffii
and
Fabaceae), Brosimum guianense Moraceae), Ocotea glomerata ("loro", Lauraceae),
("pau-d'oleo", ("oiticica",
Hymenaea
Fabaceae).
courbaril ("jatoba,"
DISCUSSION
Floristic diversity
number
The do Orobo mountain range demonstrated higher of spe-
fragments Serra a relatively
forest of
component
comparison semideciduous even comparing the arboreal only
other Brazilian
cies in to forests,
&
Werneck
Machado Jarenkow
Soares-Filho 2000; 2000;
(Meira-Neto 1989; Oliveira-Filho 1993; et
al.
et
al.
& & &
&
Waechter Moura Sampaio Santin 2002; Paula 2002; Soares
2001; 2001; Cielo-Filho et Silva
al.
& &
&
2002; Nunes 2003; Souza 2003; Andrade Rodal 2004; Toniato Oliveira-Filho 2004; Rodal
et et
al. al.
&
Lemos
Nascimento 2006; Macedo 2007), or shrubby caatinga (Rodal et al 1998; Rodal 2002; Pereira et al.
Andrade Rocha
2002; Alcoforado-Filho et al. 2003; Nascimento et al. 2003; et al. 2004; et al. 2004), riparian
Funch Stradmann 2000; Ribeiro-Filho 2002; Lacerda
1995; 1997; Passos-Junior 1999;
forests (Vilela et
al.
known
humid
2005), or enclaves of forests within the caatinga (locally as "brejo de altitude") (Tavares
et
al.
&
&
2000; Rodal Nascimento 2002; Melo Rodal 2003; Agra et 2004; Barbosa et 2004).
et al. al.
al.
humid was
submontane lower
The encountered surprisingly
species richness in the forest (87 spp.)
than would normally be expected in this forest physiognomy in comparison with other studies (Mori 1989;
&
&
Amorim
Lombardi Goncalves 2000; Tavares 2000; Carvalho-Sobrinho Queiroz 2005; 2005;
et
et al.
al.
Neves 2005; Martini 2007), and lower also that observed in the dry and riparian forest fragments of
et
al.
may
same mountain be due reduced fragment examined, and/or the
the range. This to the size of the to
may
As more presence
low sampling such, the area reveal the of
relatively effort in this area. field trips to
other species not yet recorded.
same
With minor the ten most species-rich families in this survey were the families identi-
variations,
SDTF and
fied as important in other sites, irrespective of the general aspects of the local climate, soil, forest
physiognomy. Similarly, several of these families (usually Fabaceae, Euphorbiaceae, Apocynaceae, Bignoni-
Myrtaceae, and Rubiaceae) were important in sand dune areas of the middle Sao Francisco river in
aceae,
Rocha on sandy
northeastern (Nascimento 2003; 2004), in caatinga areas soils (Araujo
Brazil et al. et al.
Andrade shrub on basement rocks
1998; Rodal 1998; 2004), caatinga areas crystalline
et al. et al. et al.
and
2002; Alcoforado-Filho 2003), semideciduous forests in southeastern southern Brazil
(Pereira et al. et al.
&
& & &
(Jarenkow Waechter 2001; Cielo-Filho Santin 2002; Paula 2002; Silva Soares 2002; Toniato
et
al.
Oliveira-Filho 2004), riparian forests in the Chapada Diamantina of Bahia State (Funch 1997; Passos-Junior
SDTF
1999; Stradmann 2000; Ribeiro-Filho 2002), as well as in several neotropical (Pennington et 2000;
al.
was most
Bridgewater 2003; Linares-Palomino 2003). Fabaceae the species-rich family in the
et
et al.
al.
Serra do Orobo mountain range as well as in the other sites referred to above. Schrire et al. (2005) presented
term between
and biogeographical evidence supporting the hypothesis of a long association le-
historical
numbers
gumes and dry Amongst the vines, the families with the greatest of species in the Serra
forests.
and were encountered
Orobo Sapindaceae)
do (Apocynaceae, Bignoniaceae, Malpighiaceae, Fabaceae, also
&
Udulutsch These
semideciduous fragments (Hora Soares 2002; 2004). too similar
in other forest et al.
physiognomies dry
species richness the family level encountered throughtout different of vegetation in
at
558 Journal of the Botanical Research Institute of Texas 2(1)
may
Brazil as well as in the Central and northern South America also be concerned for justifying that the
Serra do Orobo mountain range should be included into a broad definition of SDTF.
On may
the other hand, more evaluation comparing the most species-rich genera be sugges-
a restrict
SDTF
tive to stress that within the different physiognomies of in eastern Brazil, the dry forest fragments of
the Serra do Orobo Mountains should be related the semideciduous forests of the Atlantic
floristically to
Forest domain rather than the dry shrub-arboreal vegetation of the Caatinga domain. For example, most
to
numbers such
of the genera comprising the largest of species identified in this survey, as Croton, Passiflora,
and semideciduous
Miconia, Myrcia, Eugenia, Solatium, Bauhinia are better represented in several areas of
& &
Nunes Andrade Rodal
Soares-Filho 2000; Paula 2002; Soares 2002; al 2003;
forests et Silva et
(e.g. al.
Amorim
Macedo Rodal
2004; 2005; 2007) than shrub-arboreal caatinga areas (Araujo 1998;
in
et
et al.
al.
&
&
Lemos Cardoso
al 1998; Rodal 2002; Pereira 2002; Alcoforado-Filho al 2003; Queiroz,
et et et
al.
do Orobo Mountains another
unpublished Moreover, the paucity of Cactaceae species in the Serra
data). is
evidence for lack of closer relationship with the Caatinga domain, where this family is usually abundant
&
Gomes
example Taylor Zappi 2002; Rocha al 2004; 2006).
(see for et et
al.
Endemism
Among
Orobo
the 152 arboreal species surveyed in the dry forest fragments in the Serra do Mountains,
&
SDTF
16 (see Appendix are conspicuous representatives of the flora (Prado Gibbs 1993). Besides, the
1)
genera Apuleia (Fabaceae), Brasiliopuntia (Cactaceae), and Pterogyne (Leguminosae) surveyed here are endemic
SDTF
endemic
to the as a whole (Prado 2000). In addition, another 14 species considered to the caatinga
domain occur in the dry forest fragments of the Serra do Orobo Mountains or in other nearby areas, such
Anemopaegma
as laeve (Bignoniaceae), Apodanthera glaziovii (Cucurbitaceae), Averrhoidium gardnerianum
and
Senna
(Sapindaceae), Capparis jacobinae (Brassicaceae), Heteranthia decipiens (Solanaceae), acuruensis
may
do Orobo Mountains be
caatingae (Fabaceae) 2002). This highlights that the Serra
var. (Giulietti et
al.
remmants
indeed related to the other of seasonlly dry forests of the neotropics.
number new
The presence of a considerable of species (ten) in the dry forest fragments in the Serra
may
do Orobo Mountains be explained in term of the processes of metacommunities, a model recently put
SDTF
forward explaining maintenance within (Lavin 2004; Schrire 2005).
for the of diversity et et
al. al.
SDTF)
biome
These authors propose that the biogeography of the succulent (which includes neotropical
can be understood on by metacommunities than by proposing
better a global scale postulating historical
SDTF
processes of speciation due to climatic changes or tectonic movements. According to this model,
much
cur in isolated areas like islands isolated within the continental area), with each area function
(i.e.
random
like a local community that linked to other areas through species exchange based on dispersal
is
SDTF
The more by
events. extreme environmental conditions in restrict colonization species orig:
most immigration
in other vegetation formations (Lavin 2004), limiting the rate of into a give
et
al.
community and resulting in floras that are locally endemic. These processes also result in genera with very
SDTF humid
few species in local communities, in contrast with those observed in forests (Lavin 2006).
SDTF
and
For example, the most diverse genera in Senegalia, Croton, Machaerium, Mimosa, Senna, Zan-
(e.g.
Werneck
more
thoxylum) usually do not comprise than ten species in any given forest area (see et 2000;
al.
&
Nascimento 2003; Rodal Nascimento 2002; Bridgewater 2003; Linares-Palomino 2003;
et al. et al. et al.
& & &
Lemos
Melo Rodal 2003; Rodrigues 2003; Santos Kinoshita 2003; Andrade Rodal 2004; 2004;
et
al.
humid
Rocha and most
et 2004; this study), while the diverse genera in forests (e.g. Eugenia, Inga, Miconia,
al.
same
Myrcia, and Solanum) can have from 15 even 27 species of the genus occurring
Pouteria, Psychotria, to
&
Amorim
in the same locality (see et al. 2005; Neves 2005; Moreno et al. 2003; Lombard! Goncalves 2000;
&
Amaral
Oliveira 2004; Martini 2007).
et
al.
much
The and
dry forests of Brazil are distributed in a fragmented isolated fashion (very like islands),
which has appearance number endemic For example, 12 species en-
led to the of a significant of species.
countered in this study had distributions limited to the forest fragments of the Serra do Orobo Mountains
and adjacent areas. As such, we suggest that the endemism observed in these forests can be explained (on
559
Cardoso and de Queiroz, Seasonally dry forest fragments in Bahia, Brazil
much
a regional by the processes of metacommunities, in the same way as species exchange has been
level)
proposed on a continent scale (Lavin et al. 2004; Lavin 2006; Schrire et al. 2005). In that context, the forest
community
vegetation of the Serra do Orobo Mountains as a whole could be viewed as a local that is isolated
dry
from other dry fragments found in eastern Brazil. Additionally, the recognition that this set of
forest
metacommunity
Orobo supported
forest fragments (together with the Serra do Mountains) is a regional is
—
among been
has observed
by the presence of disjunct species the different areas a pattern that similarly
SDTF
among
(Pennington 2006b; Lavin 2004).
a continental scale the different nuclei of et et
at al. al.
been do Orobo Mountains and another dry
Cordia example, has only observed in the Serra in
blanchetii, for
forest fragment in Jequie, Bahia State; Passiflora bahiensis has been recorded only in a few collections from
an urban fragment in Salvador, Bahia; Croton warmingii demonstrates a disjunct distribution within
forest
and
Dimorphandra Senna
other semi-deciduous forest fragments in southeastern Brazil; jorgei acutisepala
known
Bahia only from few semideciduous fragments in the southern part of that
(Fabaceae) are in a forest
dry from Ceara
and Margaritopsis carrascoana (Rubiaceae) occurs in isolated areas of vegetation State
state;
northeastern Bahia.
to
relationships
Floristic
—
Riparian These forests are closely associated with rivers and are widely distributed throughout
forests.
&
although
principally in the central region of the country (Oliveira-Filho Ratter 1995), local abiotic
Brazil,
may
conditions not always be adequate for the establishment of typical riparian forests (Rodrigues 2000).
The do Orobo Mountains grow and narrow As such,
riverside forests of the Serra in steep relatively valleys.
and
their contact with the neighboring semideciduous forests is very great, frequently results in intercalation
commonly
with found those such Cupania (Sapindaceae), Dictyoloma vandellianum
species in forests, as rigida
(Rutaceae), Machaerium acutijolium (Fabaceae), Pera glabrata (Euphorbiaceae), Piper arboreum (Piperaceae),
(Olacaceae), and Vitex (Lamiaceae). However, a majority of the arboreal species
Shoepfia brasiliensis klugii
aff.
Many
encountered in the riparian forests occur only in this vegetation type. of the species occurring in the
Chapada Diamantina
do Orobo Mountains have been reported from riparian in the in central
Serra forests
Bahia (Funch 1997; Passos-Junior 1999; Stradmann 2000; Ribeiro-Filho 2002) as, for example, Bowdichia vir-
(Fabaceae), Byrsonima sericea (Malpighiaceae), Clusia nemorosa (Clusiaceae), Ficuspulchella (Moraceae),
gilioides
Simarouba
Inga thibaudiana (Fabaceae), Pouteria (Sapotaceae), Sloanea guianensis (Elaeocarpaceae),
ramiflora
and
amara Roupala montana Vantanea com-
(Simaroubaceae), Richeria grandis (Euphorbiaceae), (Proteaceae),
Orobo Mountains
some do widely
However, species observed in the riparian forests of the Serra are
pacta.
such
distributed within the riparian forests of central Brazil, as Copaijera langsdorffiU Hirtella glandulosa, Ta-
&
by
pir guianensis, and the tree-fern Cyathea delgadii (Cyatheaceae) as noted Oliveira-Filho Ratter (1995).
ira
These same authors postulated that these riparian forests could act as ecological corridors linking the two
Amazon
humid and due
South America Coastal Forest the Forest)
principal nuclei of forests in (the Atlantic
Biome
their dendritic distribution throughout of the Cerrado in central Brazil.
to all
—The
Semideciduous and humid fragments. classification of the seasonal dry forests of northeastern
forest
Andrade-Lima being
Brazil a controversial subject. (1981) considered these forests as related to arboreal
is still
and domain. Prado and Pennington based principally
caatinga, thus part of the Caatinga (2000) et (2000),
al.
on adopted circumscription, but considered
the patterns of distribution of the arboreal species, a similar
phy
the caatinga and the seasonal dry forests of eastern Brazil as part of a greater togeographical unit of the
common (SDTF)—
from
neotropics— the so referred seasonally dry tropical forests as distinct the Atlantic
Coastal Forest. This phytogeographical unit would comprise a large variety of vegetation formations, from
in more humid areas cactus scrub in the driest regions. This controversy can be clearly seen
forests to
tall
and
names such dry
physiognomies, subtropical
in the great variety of applied to these different as: tropical
caatinga, mesotrophic, mesophilous or mesophytic forest, semideciduous or deciduous forest, bosque
forests,
and woodland
thorn (Pennington
2000).
caducifolio, et al.
On
the other hand, Oliveira-Filho (2006) performed similarity analyses with areas of seasonal
et
al.
humid and
and
dry shrub-arboreal caatinga, Atlantic Coastal Forest areas, identified greater floristic
forests,
560 Journal of the Botanical Research Institute of Texas 2(1)
between seasonal dry and Atlantic Coastal Forest areas than with shrub-arboreal caatinga
forests
affinity
SDTF
South America
As proposed wider eastern
such, Oliveira-Filho (2006) a definition of of
vegetation.
et
al.
Chaco,
would include the three major nuclei of dry vegetation formations: the Caatinga, the
that floristic
and the seasonal forests of the Atlantic Forest domain, sensu latissimo.
humid
An analysis of arboreal and shrub species distributions from semideciduous and forest frag-
ments in the Serra do Orobo Mountains demonstrated that 106 species (46.5%) also occur in areas of the
gonocarpum
domain including Chrysophyllum (Sapotaceae),
Forest (sensu Oliveira-Filho 2006),
Atlantic et
al.
Myrcia (Myrtaceae), Swartzia apetala
Esenbeckia leiocarpa (Rutaceae), Lonchocarpus cultratus (Fabaceae), rostrata
A common
sampled
(Fabaceae), and TricMia hirta (Meliaceae). total of 56 species (24%) were to areas in the
Mimosa
domain, and included Aspidosperma (Apocynaceae), Poeppigiaprocera,
Caatinga pyrifolium tenuiflora,
Of
(Rhamnaceae) 2006). these species, only
Senna (Fabaceae), andZiziphusjoazeiro (Giulietti et
spectabilis al.
domain
nine of were considered as endemic to the Caatinga (Giulietti et 2002). Additionally, if consider-
al.
non
domain by another
ing the 318 endemic species to the Caatinga cited Giulietti et (2002), eight
al.
all
do Orobo Mountains other nearby
arboreal or shrub species occur in dry forest fragments of the Serra or in
However, make up only 4.5% the endemic of the Caatinga domain,
these 17 species of total flora
areas.
do Orobo Mountains might not be phytogeographically related
dry
suggesting that the forests of the Serra
numbers
the Caatinga domain, but rather are of Atlantic Forest vegetation. Therefore, these reinforce the
to
hypothesis of Oliveira-Filho (2006) that the semideciduous forests of eastern Brazil are floristically
et
al.
domain domain.
more than Caatinga
closely related to the Atlantic Forest to the
made do
between
These suggestions further strengthened by comparisons the flora of the Serra
are
southern Bahia
Orobo Mountains and do Teimoso Reserve fragment of Atlantic Coastal Forest in
the Serra
(a
The Serra do Teimoso Mountains, like the Serra do Orobo Mountains, have dry vegetation at their
State).
more
humid and
more the there closely
lower altitudes but conditions exist at greater elevations, flora is
SDTF (Amorim
domain than sensu Pennington (2000) 2005).
related to the Atlantic Forest to the et al. et al.
Among do Teimoso Mountains, 52 can be found the
the 252 arboreal species that occur in the Serra also in
Orobo (Cactaceae),
Serra do Mountains, including Andira (Fabaceae), Brasiliopuntia brasiliensis
fraxinijolia
Centrolobium tomen-
Brosimum guianense (Moraceae), Cavanillesia arborea (Malvaceae), Caesalpinia pluviosa,
tosum, Goniorrhachis marginata (Fabaceae), Guapira opposita (Nyctaginaceae), Urera caracasana (Urticaceae),
common
and Zanthoxylum (Rutaceae). Examples of shrubs and herbaceous species held in include:
rhoifolium
Faramea
barbata (Gesneriaceae), Coccoloba dedinata (Polygonaceae),
Costus (Costaceae), Sinningia
spiralis
such
and Conchocarpus macrophyllus (Rutaceae). Additionally, species as Croton
hyacinthina (Rubiaceae),
and macrostachya (Malpighiaceae)
warmingii (Euphorbiaceae), Dimorphandra jorgei (Fabaceae), Heteropterys
Orobo Mountains and semideciduous
between do the
demonstrate a disjunct distribution pattern the Serra
domain.
within the Atlantic Forest
forests
between
demonstrating phytogeographical relationship
Thus, while data close
there exists significant a
SDTF
the Serra do Orobo Mountains and the nuclei of seasonal dry forests within the Atlantic Forest do-
domain would
number from Caatinga be useful
main, additional studies involving a larger of areas also the
some
confirming conclusions presented
in of the here.
APPENDIX
1
humid
and submontane do
found semideciduous, forests of the Serra
Species of re plants in riparian,
list
names endemics from
Orobo Mountains, Bahia The before the species highlights the
asterisk
State, Brazil.
Serra do Orobo mountain range and/or adjacent dry forest areas and a circle refers to the representative arbo-
&
SDTF semidecidu-
according Prado Gibbs Abbreviations vegetation types: SF:
real species of to (2002). for
and submontane humid abbreviations collectors LP=L.P.de Queiroz;
ous, RF: riparian, ShF: forests; for are:
^.B.Costa;
