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Lists of Species Check List 12(1): 1842, 16 February 2016 doi: http://dx.doi.org/10.15560/12.1.1842
ISSN 1809-127X © 2016 Check List and Authors
1 Universidade Federal de Pernambuco, Departamento de Botânica, Laboratório de Morfo-taxonomia Vegetal. Av. Prof. Moraes Rego s/n,
Cidade Universitária. CEP 50670-901. Recife, PE, Brazil
2 Jardim Botânico do Recife, km 7,5 da BR 232 s/n, Curado. CEP 50000-230. Recife, PE, Brazil
* Corresponding author. E-mail: aline_vmelo@yahoo.com.br
Abstract: A list of angiosperms from Serra do Urubu, Atlantic Forest domain, and of these, 8,601 (56.3%) are
a montane forest area in the state of Pernambuco, is endemic. The Atlantic Forest is first among Brazilian
here provided. Based on 14 botanical expeditions and domains for threatened species (1,544 species), in
material deposited in herbaria, 832 taxa belonging to addition to those that have insufficient data (265) and
442 genera and 118 families have been recorded in this those which are not endangered but are considered
area, with about 90% of the taxa identified to species of interest for conservation and research (337) (Livro
level. The richest families are Orchidaceae (86 spp.), Vermelho da Flora do Brasil 2013).
Fabaceae (51 spp.) and Rubiaceae (42 spp.). Miconia and According to Stehmann et al. (2009), Orchidaceae,
Solanum (14 spp. each), and Psychotria (13 spp.) are the Fab aceae, Asteraceae, Bromeliaceae, Poaceae, Myrt
richest genera. About 15% of the recorded species are aceae, Melastomataceae and Rubiaceae are the most
endemic to the Atlantic Forest, and another 10% are representative families in the Atlantic Forest. Bromeli
disjunct between this area and the Amazon Rainforest. aceae and Orchidaceae especially noteworthy for their
The results indicate that the Serra do Urubu is one of high richness and endemism. Local floristic surveys
the richest areas in terms of number of species of the along the Atlantic Forest demonstrate that these
Atlantic Forest in the Northeast, and corroborates the families figure among the richest in distinct areas of this
Pernambuco Endemism Center. domain, but in different proportions depending on, for
example, altitude, latitude, and vegetation type (Barros
Key words: Brazilian Northeast; endemic species; et al. 1991; Alves-Araújo et al. 2008; Amorim et al. 2008,
floristics; hotspot; rock outcrops 2009; Scheer and Kinoshita 2009; Coelho and Amorim
2014; Forzza et al. 2014; Landim et al. 2015).
Floristic inventories of the Atlantic Forest in
INTRODUCTION Pernambuco were performed in areas of restingas and
The Atlantic Forest is considered one of the world’s 34 lowland forests (up to 100 m in altitude) by Guedes
biodiversity hotspots, possesses high species richness (1998), Rodal et al. (2005a), Almeida Jr. et al. (2007, 2009),
and endemism levels, a wide latitudinal extension Sacramento et al. (2007), Soares Júnior et al. (2008), Pessoa
(3°S-30°S) and various types of vegetation, yet despite et al. (2009), Cavalcanti (2012) and Alves et al. (2013).
this, only about 7-12% of its original area remains These inventories point out the taxonomic diversity that
(Myers et al. 2000; Mittermeier et al. 2004; Ribeiro et ranges from 101 to 830 species, demonstrating the high
al. 2009). Myers et al. (2000) estimated that there were species richness of some areas (Cavalcanti 2012; Alves et
20,000 species of plants, of which 8,000 (40%) were al. 2013). Inventories that include areas of submontane
endemic to this region and subsequently Stehmann and montane forests (above 100 m altitude) in the state
et al. (2009), in a more detailed compilation, recorded are scarcer (Rodal et al. 2005b; Barreto et al. 2006; Grillo
15,782 species for the Brazilian Atlantic Forest, with et al. 2006; Rodal and Sales 2007; Ferraz and Rodal 2008);
13,708 angiosperms, of which 49% were considered they document the number of angiosperm species as
endemic. Based on the discovery of new species and between 69 and 375.
the general advancement of knowledge of Brazilian The Catende Complex (which includes Serra do
plants, the Lista de Espécies da Flora do Brasil (2015) Urubu) is located in the southern portion of the state
enumerates 15,282 species of flowering plants in the of Pernambuco and is indicated by the MMA (2007) as
of extreme biological importance and is considered a protected area. In addition, Jaqueira was highlighted
priority area for conservation of the flora of the Atlantic by Kasecker et al. (2009) as a Key Biodiversity Area,
Forest domain (Martinelli 2007). The area has been based on the presence of a rare species of Bromeliaceae
the subject of many studies showing great diversity of (Neoregelia pernambucana Leme & J.A. Siqueira-Filho).
birds (Roda 2002; Bencke et al. 2006), frogs (Santos and The area is characterized as the border between
Carnaval 2002), lycophytes and ferns (Barros et al. 2006), montane and submontane seasonal forest (sensu Veloso
Bromeliaceae and Orchidaceae (Leme and Siqueira-Filho et al. 1991), consisting of areas of remaining forest and
2006; Siqueira-Filho and Felix 2006; Pessoa and Alves rock outcrops, and surrounded by areas of sugar-cane
2015b), and even new taxa to science (Carnaval and (Usina Colônia), livestock and subsistence agriculture
Peixoto 2004; Chautems et al. 2005; Leme and Siqueira- on small farms. The average rainfall between 1994 and
Filho 2006; Amorim and Alves 2012a, 2015; Costa-Lima 2013 was 1373.1 mm/year (data from the headquarters of
and Alves 2013; Giacomin et al. 2013; Gregório et al. Usina Colônia). The altitude varies between 600 and 780
2014; Pessoa and Alves 2015a; Pessoa et al. 2014a, 2014b; m, and the area experiences fog at the highest points
Sobral 2013). during the winter months, as well as having ponds and
Therefore, the objectives of this study were the streams.
following: (1) to perform an inventory of angiosperms Samples were collected in the private area of Private
in remnant \montane forests in the southern portion of Reserve of Natural Patrimony (PRNP) Frei Caneca,
the state of Pernambuco, (2) to corroborate the floristic owned by Usina Colônia, and PRNP Pedra D’Anta,
patterns observed in other areas of Atlantic Forest in belonging to Society for Conservation of Birds of Brazil
the region (the richest families, habits and similarity (SAVE Brasil), which are adjacent areas and together
between areas based on representative taxonomic form a continuous fragment of approximately 1,000
groups), (3) to document new occurrences for the ha of Atlantic Forest in the Serra do Urubu (Bencke et
state of Pernambuco and species with some degree of al. 2006). Field expeditions were conducted between
threatened or rare status. August 2010 and December 2014, including both the dry
and rainy seasons, totaling 14 trips and approximately
MATERIALS AND METHODS 50 days of sampling effort. The material collected was
The floristic survey was carried out in the Serra do treated according to the usual botanical techniques
Urubu (08°42ʹ48″ S, 035°50ʹ38″ W), located between (Peixoto and Maia 2013) and deposited at Herbarium
the municipalities of Lagoa dos Gatos, Jaqueira and São UFP with duplicates distributed to the herbaria ASE,
Benedito do Sul, in the Zona da Mata Sul in the state FRA, HRCB, JPB, NY, RB and UFRN (acronyms according
of Pernambuco, about 160 km from Recife (Figure 1). to Thiers 2015).
Besides being considered by MMA (2007) as a priority Identification of the material was performed by
conservation area for the Atlantic Forest domain, the consulting the taxonomic literature, the collections of
municipality of Jaqueira was indicated by Martinelli the UFP and JPB herbaria and specialists as indicated in
(2007) as an area of extreme importance, requiring Table 1. The classification of families, name of the species,
priority action to carry out an inventory and create a abbreviations of authors, geographical distribution and
Figure 1: Map showing the location of the Serra do Urubu, Pernambuco state.
conservation status followed the Lista de Espécies da Flora species were organized into groups based on their source
do Brasil (2015) and the Livro Vermelho da Flora do Brasil of nutrition: (1) Autotrophic, (2) Heterotrophic (parasitic
(2013). The list presented here is mainly the product of or hemiparasitic) or (3) Mycoheterotrophic.
collections made during this work, but other vouchers
collected previously in the study area and deposited in RESULTS
the herbaria HST (Herbarium of Forestry, Federal Rural In the study area 832 taxa were recorded, belonging to
University of Pernambuco - not indexed), JPB and UFP 442 genera and distributed in 118 families. About 90% of
were also used. the morphotypes were identified to the species level (750
The identified species were categorized into structural spp.), 6% only to genus (50), 3% to family level and the
groups as (1) Herbs, (2) Bushes, (3) Trees and related forms remaining 1% undetermined. The most representative
(including treelets and arborescent plants like palms and families were Orchidaceae (86 species), Fabaceae (51
bamboo) and (4) Climbers (woody or herbaceous vines). A spp.), Rubiaceae (42 spp.), Bromeliaceae (41 spp.),
further categorization was also adopted for the substrate: Poaceae (40 spp.), Cyperaceae (34 spp.), Asteraceae (31
(1) Terrestrial, (2) Saxicolous (3) Epiphytic and/or Hemi- spp.), Melastomataceae (30 spp.) , Solanaceae (24 spp.)
epiphytic and (4) Aquatic or Wetland. It is noteworthy and Myrtaceae (23 spp.), comprising about 50% of the
that in this case, some species occupy more than one type, species found in the area. The richest families in number
which is a common condition in some representatives of species also have greater richness in genera such as
of Araceae, Bromeliaceae and Orchidaceae. Finally, the Orchidaceae (51 genera), Fabaceae (30), Poaceae (24),
Table 1. List of the 750 taxa of angiosperms identified to species level of the Serra do Urubu. The species follow in alphabetical order of family and spe-
cies, respectively, with information of habit and voucher specimens of the referenced taxon. Samples are deposited in HST, JPB and UFP. * Alien species.
** Name followed Zanotti et al. (2012).
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Dorstenia bahiensis Klotzsch ex Fisch. & C.A.Mey. Herb A. Melo et al. 702
Ficus cf. calyptroceras (Miq.) Miq. Tree D. Araújo et al. 1783
Helicostylis tomentosa (Poepp. & Endl.) Rusby Tree B.S. Amorim et al. 1847
Myristicaceae (Bruno S. Amorim)
Virola gardneri (A.DC.) Warb Tree A. Melo et al. 755
Myrtaceae (Bruno S. Amorim)
Campomanesia aromatica (Aubl.) Griseb. Tree B.S. Amorim et al. 1567
Campomanesia dichotoma (O. Berg) Mattos Tree E. Pessoa et al. 444
Campomanesia xanthocarpa (Mart.) Berg. Tree B.S. Amorim et al. 1857
Eugenia astringens Cambess. Shrub B.S. Amorim et al. 1988
Eugenia cf. candolleana DC. Tree B.S. Amorim et al. 1280 (sterile)
Eugenia culicina Sobral Tree B.S. Amorim et al. 1782
Eugenia excelsa O. Berg Tree B.S. Amorim et al. 800
Eugenia ligustrina (Sw.) Willd. Tree B.S. Amorim et al. 1840
Eugenia pyriformis Cambess. Tree B.S. Amorim et al. 1851
Eugenia submontana B.S. Amorim & M. Alves Tree B.S Amorim et al. 1080
Eugenia tumescens B.S. Amorim & M. Alves Tree B.S. Amorim et al. 1783
Eugenia umbrosa O. Berg Tree B.S. Amorim et al. 1757
Myrcia amplexicaulis (Vell.) Hook. f. Tree B.S. Amorim et al. 817
Myrcia spectabilis DC. Tree B.S. Amorim et al. 768
Myrcia splendens (Sw.) DC. Tree B.S. Amorim et al. 1781
Myrcia sylvatica (G. Mey) DC. Tree A. Melo et al. 685
Myrcia tenuivenosa Kiaersk. Tree B.S. Amorim et al. 945
Myrcia tomentosa (Aubl.) DC. Tree B.S. Amorim et al. 771
Psidium guajava L. Tree B.S. Amorim et al. 1250
Psidium guineense Sw. Tree B.S. Amorim et al. 770
Syzygium cumini (L.) Skeels* Tree B.S. Amorim et al. 815
Nyctaginaceae (Aline Melo)
Guapira opposita (Vell.) Reitz Tree A. Melo et al. 756
Ochnaceae (Fernanda Silva)
Ouratea crassa Tiegh. Tree B.S. Amorim et al. 793
Sauvagesia erecta L. Subshrub A. Melo et al. 607
Olacaceae (Bruno S. Amorim)
Cathedra rubricaulis Miers Shrub-Tree B.S. Amorim et al. 792
Onagraceae (Bruno S. Amorim)
Ludwigia hyssopifolia (G. Don) Exell Herb E. Pessoa et al. 471
Orchidaceae (Edlley Pessoa)
Acianthera glumacea (Lindl.) Pridgeon & M.W. Chase Herb E. Pessoa et al. 520
Acianthera hygrophila (Barb. Rodr.) Pridgeon & M.W. Chase Herb E. Pessoa et al. 954
Anathallis barbulata (Lindl.) Pridgeon & M.W. Chase Herb E. Pessoa et al. 521
Anathallis sclerophylla (Lindl.) Prodgeon & M.W. Chase Herb E. Pessoa et al. 959
Aspidogyne argentea (Vell.) Garay Herb E. Pessoa et al. 427
Aspidogyne foliosa (Poepp. Endl.) Garay Herb E. Pessoa et al. 416
Brassavola tuberculata Hook. Herb J.A. Siqueira-Filho 985
Camaridium carinatum (Barb. Rodr.) Hoehne Herb E. Pessoa et al. 931
Camaridium micranthum M.A. Blanco Herb E. Pessoa et al. 928
Campylocentrum crassirhizum Hoehne Herb E. Pessoa and A. Melo 470
Campylocentrum micranthum (Lindl.) Rolfe Herb E. Pessoa et al. 1062
Campylocentrum pernambucense Hoehne Herb E. Pessoa and A. Melo 429
Campylocentrum robustum Cogn. Herb M.T. Buril et al. 645
Campylocentrum serranum E. Pessoa & M. Alves Herb E. Pessoa et al. 1051
Cattleya granulosa Lindl. Herb E. Pessoa et al. 941
Continued
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Table 1. Continued.
Rubiaceae (20), Asteraceae (19) and Bromeliaceae (17). (1.5% of total species) found are “hemiparasitic” or
The genera with the highest taxonomic diversity “mycoheterotrophic”; of these, eight species (61.5%) are
were Miconia Ruiz & Pav. and Solanum L. with 14 “hemiparasitic”, belonging to the families Loranthaceae
species each (corresponding to 46.7% of the species of (4 spp.), Orobanchaceae (1 spp.) and Santalaceae (3 spp.),
Melastomataceae and 58.3% of Solanaceae), Psychotria and five species (38.5%) are “mycoheterotrophic”, with
L. with 13 species (30.9% of Rubiaceae) and Epidendrum representatives in Burmanniaceae and Gentianaceae (2
L., Eugenia L., Passiflora L. and Piper L. with 11 species spp. each) and Orchidaceae (1 sp.).
each (12.8% of Orchidaceae, 47.8% of Myrtaceae, 100% About 15% of the total species of Serra do Urubu (122
of Passifloraceae and 61.1% of Piperaceae). spp.) are considered, according to the Lista de Espécies
The most representative structural group in the study da Flora do Brasil (2015), as endemic to the Atlantic
area was “Herbs”, totaling 355 spp. (42.6% of the total Forest. It is worth mentioning that Bromeliaceae
species) being most terrestrial (58.6%) or epiphytes (26 spp.), Orchidaceae (20 spp.), Myrtaceae (8 spp.),
and/or hemiepiphytes (34%). Almost 60% of species Solanaceae (7 spp.) and Marantaceae (5 spp.) together
classified as “Herbs” belong to the following families of account for over 50% of cases. Among this same set of
Monocotyledons: Orchidaceae, Bromeliaceae, Poaceae, species, four are known only from forest remnants of
Cyperaceae and Araceae. the states of Bahia and Pernambuco (Rauvolfia moricandii
The “Bushes” category includes 194 taxa (23.3% of the A. DC., Apocynaceae; Begonia obdeltata Gregório & E.L.
total species), being most represented in terms of number Jaques, Begoniacae; Heteropterys cordifolia Moric. ex A.
of species by Rubiaceae with 34 spp. (representing Juss., Malpighiaceae; and Peperomia estrellensis DC.,
17.5% of the species in the bushes category), followed Piperaceae) and 23 are considered to be restricted to
by Solanaceae with 20 (10.3%), Melastomataceae with the Pernambuco Endemism Center (sensu Prance 1987),
18 (9.3%) and Malvaceae with 15 spp. (7.7%). Followed with 10 known only from the state.
by “Trees and related forms” with 189 spp. (22.7% of all A disjunct distribution between the Amazon and
species) and was represented by Fabaceae with 24 spp. Atlantic Forests has been confirmed for 10.6% of
(12.6% of the Trees and related forms group), Myrtaceae the species in the area. Among the families with this
with 22 (11.6%) and Melastomataceae with 14 spp. distribution pattern are Orchidaceae with 24 spp.
(7.4%). (22.7% of the cases of disjunction), Fabaceae with seven
“Climbers” completes the picture with 116 spp. (14% (8%), Bromeliaceae, Melastomataceae and Piperaceae
of total species in the area), of which 34 are herbaceous with four (4.5% each), Lecythidaceae and Violaceae
(29% of the species of Climbers) and the remainder with three species each (3.4% each). Some examples
woody (71%). The group is well represented by Fabaceae are the following: Lecythidaceae — Eschweilera ovata
and Passifloraceae with 11 spp. each (9.5% each), (Cambess.) Mart. ex Miers, Lecythis lurida (Miers) S.A.
Convolvulaceae with 10 (8.6%), Apocynaceae with nine Mori and L. pisonis Cambess.; Bromeliaceae — Aechmea
(7.7%), Dioscoreaceae with eight (6.9%), Cucurbitaceae mertensii (G.Mey.) Schult. & Schult. f.; Annonaceae —
with seven (6%) and Asteraceae, Malpighiaceae and Cymbopetalum brasiliensis (Vell.) Benth. ex Baill.;
Sapindaceae with six species each (5.2% each). Menispermaceae — Hyperbaena domingensis (DC.) Benth.;
Considering the substrate, most species of the area and Piperaceae — Peperomia glabella (Sw.) Dietr.
are “Terrestrial” with 676 spp. (81.2% of all species)
and comprise several families. The second most repre DISCUSSION
sentative group are “epiphytic and hemiepiphytic” with The Serra do Urubu has the biggest taxonomic richness
124 spp. (about 15% of the total species) and the greatest (832 taxa) when compared with other floristic surveys in
diversity among Orchidaceae (53.2% of epiphytes and the state of Pernambuco (Table 2) (Guedes 1998; Rodal
hemiepiphytes), Bromeliaceae (25.8%) and Araceae et al. 2005a, 2005b; Rodal and Sales 2007; Sacramento
(9.7%) and Piperaceae (5.6%). These are followed by et al. 2007; Ferraz and Rodal 2008; Cavalcanti 2012;
the “Saxicolous” group, represented by 47 spp. (5.6% Alves et al. 2013). The Serra do Urubu slightly exceeds
of all species), and families with the most species were the total number of species only an area of lowland
Bromeliaceae (30% of saxicolous species), Orchidaceae forest in the north of the state, whose inventory has
(19.1%), Begoniaceae and Gesneriaceae (6.4% each). been continuously updated since 2008 and was recently
“Aqua tic and wetland” plants are represented by 21 updated to 830 species of angiosperms (Alves-Araújo et
spp. (2.5% of total species), among which Cyperaceae al. 2008; Melo et al. 2011; Alves et al. 2013).
has seven spp. (33.3% of aquatic and wetland species), The difference is even more striking when the data
Lentibulariaceae and Linderniaceae two species each presented here are compared with floristic inventories
(9.5% each) and the rest of the families have only one of areas of submontane and montane forests of
species. Pernambuco (Rodal et al. 2005b; Rodal and Sales 2007;
Regarding the source of nutrition, only 13 species Ferraz and Rodal 2008), even after considering the
Number of
Study area (state, locale) Vegetation species / families Most representative families Reference
Pernambuco, Serra do Urubu Montane Forest (600–780 832 / 118 Orchidaceae (86 spp), Fabaceae (51 spp.), Rubiaceae (42 This work
m) spp.), Bromeliaceae (41 spp.) and Poaceae (40 spp.)
Pernambuco, Mata do Estado Submontane / Montane 375 / 94 Fabaceae (34), Myrtaceae (21), Rubiaceae (20), Lauraceae Ferraz and Rodal
Forest (200–640 m) and Sapindaceae (14). 2008
Pernambuco, Reserva Municipal Submontane Forest 217 / 65 Fabaceae (26), Asteraceae and Euphorbiaceae (14), Rodal et al. 2005b
de Bonito (450–500 m) Melastomataceae and Rubiaceae (13).
Pernambuco, Parque Ecológico Submontane Forest 332 / 85 Fabaceae (51), Asteraceae (20), Rubiaceae (19), Solanaceae Rodal and Sales 2007
João Vasconcelos-Sobrinho (800–950 m) (14), Myrtaceae (13).
Pernambuco -Usina São José Lowland Forest (60–150 m) 830 / 113 Fabaceae (87), Poaceae (54), Cyperaceae (40), Rubiaceae Alves et al. 2013
(34), Orchidaceae and Melastomataceae (29).
Paraíba, Reserva Ecológica Mata Submontane Forest 309 / 84 Fabaceae (30), Rubiaceae (24), Malvaceae (21), Solanaceae Vasconcelos et al.
do Pau Ferro (400–600 m) (16) and Asteraceae (14). 2004
Ceará, Serra do Baturité Submontane / Montane 419 / 89 Myrtaceae (52), Fabaceae (41), Euphorbiaceae (23), Araújo et al. 2007
Forest (400–800 m) Rubiaceae (21), Lauraceae (13).
Bahia, Serra do Corcovado Submontane / Montane 678 / 100 Orchidaceae (69), Rubiaceae (46), Bromeliaceae (33), Coelho and Amorim
Forest (400–1040 m) Fabaceae (30), Melastomataceae (28). 2014
Bahia, Serra da Pedra Lascada Montane Forest 466 / 88 Orchidaceae (42), Rubiaceae (41), Bromeliaceae (30), Coelho and Amorim
(600–950 m) Melastomataceae (27), Poaceae (23). 2014
Bahia, Serra das Lontras Submontane / Montane 792 / 110 Orchidaceae (108), Bromeliaceae (62), Rubiaceae (57), Amorim et al. 2009;
Forest (400–1,000 m) Melastomataceae (45), Myrtaceae and Asteraceae (33 Leitman et al. 2014
each).
Bahia, Serra Bonita Submontane / Montane 628 / 103 Orchidaceae (52), Rubiaceae (44), Melastomataceae (37), Amorim et al. 2009
Forest (300–1,080 m) Asteraceae (33), Poaceae (28).
Bahia, Serra do Teimoso Montane Forest (850 m) 667 / 100 Fabaceae (51), Myrtaceae (39), Rubiaceae (31), Orchidaceae Amorim et al. 2005
(25), Sapotaceae and Solanaceae (24 each).
Bahia, Reserva Biológica do Una Lowland and Submontane 947 / 108 Fabaceae (71), Rubiaceae (66), Myrtaceae (59), Amorim et al. 2008
Forest (100–350 m) Bromeliaceae (40) and Orchidaceae (38).
size difference of fragments. For example, the region cataloged over 10 years of sampling effort. This region
of Serra do Urubu studied here has more than twice as stands as the most species-rich area in northeastern
many species recorded in an area of the municipality of Brazil (Amorim et al. 2008). However, excluding species
São Vicente Férrer (Ferraz and Rodal 2008), which had that are unique to “tabuleiro” and mangroves, 737
until now held the record for the greatest number of species have been cataloged for the submontane region
species for this type of vegetation in the state. The high of Una, which makes the species richness of the Serra do
floristic diversity of the Serra do Urubu is also reflected Urubu and Rebio UNA comparable.
in other groups of plants such as ferns and lycophytes, The most diverse families in number of species in
which according to Barros et al. (2006), have more than the Serra do Urubu (Orchidaceae, Fabaceae, Rubiaceae,
40% of the species cataloged for Pernambuco occurring Bromeliaceae, Poaceae, Asteraceae, Melastomataceae
in the PRNP Frei Caneca. and Solanaceae) were also the most species-rich in
In relation to similar inventories conducted in floristic studies of montane areas in southern Bahia,
montane forests in the Northeast, the number of such as in the Serra do Corcovado (Coelho and Amorim
recorded species is compatible with values obtained 2014), Serra Bonita and Serra das Lontras (Amorim et al.
by Amorim et al. (2009), Coelho and Amorim (2014) 2009; Leitman et al. 2014). A similar pattern of richness
and Leitman et al. (2014), in areas of southern Bahia was found in Una where Fabaceae, Rubiaceae, Myrtaceae,
(412–792 species) and which are recognized for the high Bromeliaceae, Orchidaceae and Melastomataceae were
diversity and high levels of endemism in their floras. In the most species-rich families, listed in descending order.
addition, the richest fragments in the studies mentioned On the other hand, in studies carried out in Pern
are of sizes comparable with the fragments of the Serra ambuco by Rodal et al. (2005b), Rodal and Sales (2007)
do Urubu (Table 2). Nevertheless, forest fragments in and Ferraz and Rodal (2008), Orchidaceae was poorly
southern Bahia with more than 2000 ha still have lower sampled and represented only 3.2–1.5% of the flora,
species richness than the Serra do Urubu (Amorim et while in the Serra do Urubu Orchidaceae represents
al. 2009; Coelho and Amorim 2014; Leitman et al. 2014). 10.3% of the total number of species recorded. A
The richness of plants in the Serra do Urubu is proportion similar to this has been shown for areas of
comparable to the Una Biological Reserve, located in Bahia, wherein Orchidaceae account for between 8.2%
southern Bahia and home to a mosaic of environments and 10.2% of the flora (Amorim et al. 2009; Coelho and
consisting of submontane forest, “tabuleiro” vegetation, Amorim 2014; Leitman et al. 2014). Furthermore, the
mangrove, flooded and disturbed areas. The Una richness of the family in the Serra do Urubu is among
Biological Reserve has 947 species of angiosperms the greatest ever recorded in floristic surveys in the
Northeastern Brazil. These differences are probably Sapindaceae. However, in the Serra do Urubu this
due to different sampling efforts concentrated on the group also includes Passifloraceae, Apocynaceae,
arboreal layer of the areas of Pernambuco. However, one Convolvulaceae, Asteraceae and Malpighiaceae.
cannot fail to consider the effect of fragmentation and Plants with differentiated forms of nutrition
human pressures on the species, besides the difference (hemiparasitic, parasitic or mycoheterotrophic) are
in the size of the fragments among the studied areas. usually represented by only a few species or absent in
Fabaceae emerges at different proportions in the floristic inventories. No mycoheterotrophic species
various studies performed in Atlantic Forest fragments, were recorded in montane and submontane areas of
despite being one of the most representative families Pernambuco by Rodal et al. (2005b), Rodal and Ferraz
in montane areas of Pernambuco (Rodal et al. 2005b; (2007) and Ferraz and Rodal (2008), who recognized
Rodal and Sales 2007; Ferraz and Rodal 2008) and Bahia only hemiparasitic species of the families Loranthaceae
(Amorim et al. 2005, 2009; Coelho and Amorim 2014). In and Santalaceae. In these works, the hemiparasites were
the Serra do Urubu, the family accounts for about 6.1% represented by one or up to four species, while eight
of total species, while in the Mata do Brejão (Rodal et species have been recorded in the Serra do Urubu. Araújo
al. 2005b) and the Municipal Park Vasconcelos Sobrinho et al. (2007), in the Serra do Baturité (Ceará state), also
(Rodal and Sales 2007), its proportion ranges from recorded hemiparasitic species of Loranthaceae and
12–15.6%. This difference is suggested here to be due to a a species of Balanophoraceae (Langsdorffia hypogaea
small sampling of non-arboreal structural groups in Rodal Mart.). Meanwhile, Barbosa et al. (2004), working in a
et al. (2005b) and Rodal and Sales (2007), as observed for montane forest area in Areia (Paraíba state), cataloged
Orchidaceae and Cyperaceae, among other families. species of hemiparasite and mycoheterotrophy but
It is important to point out that, for various reasons, also cited Cuscuta racemosa Mart. (Convolvulaceae), a
there is a clear bias directed to the effort of collecting parasitic species that was not recorded in other studies.
woody species (including trees, woody climbers In areas of southern Bahia (Amorim et al. 2005, 2008,
and shrubs) in some studies, which usually leads to 2009; Coelho and Amorim 2014), hemiparasites are
low sampling of species from other groups, such as represented with three to nine species in these same
herbaceous plants. This is evidenced in Ferraz and families (Lorantahecae and Santalaceae), in addition
Rodal (2008), in which members of the woody group to parasitic species such as Helosis cayennensis (Sw.)
represented 64.5% of the recorded species. On the other Spreng. (Balanophoraceae) and mycoheterotrophic
hand, when the same sampling effort is performed for species (e.g., Gymnosiphon divaricatus (Benth.) Benth.
other strata of the forest, richness patterns emerge & Hook. f. [Burmanniaceae] and Voyria flavescens
similar to those seen in the Serra do Urubu. Thus, in Griseb. [Gentianaceae]). The mycoheterotrophic species
Rodal et al. (2005b) and Rodal and Sales (2007), trees recorded in this study Campylosiphon purpurascens
represent 25% and 35.7% of the species sampled in the Benth. (Burmanniaceae) and Wullschlaegelia aphylla
areas, respectively, which is close to a value found here. (Sw.) Rchb. f. (Orchidaceae) were previously known only
In addition, Fabaceae, Rubiaceae and Myrtaceae are from single records collected 40 years ago in the state of
among the most representative families in these works Pernambuco from other areas of montane forests in the
(Rodal et al. 2005b; Rodal and Ferraz 2007; Ferraz and state (Melo and Alves 2013).
Rodal 2008). The rock outcrops in the Serra do Urubu are generally
The most representative of structural groupings was at the summit of the mountain, and have been given
“Herbs”, followed by “Bushes” and “Trees and related particular names, such as Pedra do Cruzeiro, which is an
forms”, corroborating the findings of Amorim et al. access point for visitors. Some species recorded in the
(2009) and Coelho and Amorim (2014) in inventories Serra do Urubu are restricted to these environments,
in montane forests in Bahia. In both those studies, among which are Ortophytum disjunctum L.B. Sm.
Orchidaceae is identified as the richest family in number (Bromeliaceae), Paepalanthus myocephalus Mart. ex Körn.
of species, followed by Rubiaceae, Bromeliaceae, (Eriocaulaceae), Euphorbia comosa L. (Euphorbiaceae)
Fabaceae, Myrtaceae, Melastomataceae and Solanaceae, and Sobralia liliastrum Salzm. ex Lindl. (Orchidaceae).
findings which were also found in the Serra do Urubu. These species have also been recorded in other floristic
The similarities also apply to climbers. Rodal et al. surveys of rock outcrops in Pernambuco (Gomes and
(2005b), Rodal and Ferraz (2007), Ferraz and Rodal Alves 2010; Gomes and Sobral-Leite 2013; Pessoa and
(2008), this functional group is 10–15% of the flora in Alves 2014). Other examples are Mandevilla dardanoi
their respective study areas, a pattern into which the M.F. Sales, Kin.-Gouv. & A.O. Simões (Apocynaceae),
Serra do Urubu fits nicely at 14% . The distribution of Paliavana tenuiflora Mansf. and Sinningia nordestina
species richness of climbers into families is also similar, Chautems, Baracho & J.A. Siqueira (Gesneriaceae),
where Fabaceae is among the most representative which are considered restricted to areas of rock outcrops
together with Cucurbitaceae, Dioscoreaceae and of the Borborema Plateau and its extensions (Sales et al.
2006; Gomes and Alves 2009, 2010; Araújo et al. 2015). (Lista de Espécies da Flora do Brasil 2015), an overall
Our results also provide significant advances in the pattern of the Atlantic Forest reflected at the local level
floristic knowledge of the region and on the distribution in the Serra do Urubu.
of species in the northern Atlantic Forest. With this list Ferraz et al. (2004) considered some species
there was a considerable increase in the number of listed recorded here as characteristic of the montane forests
species from Serra do Urubu, as Cardoso et al. (2006) of Pernambuco, among them: Couepia impressa Prance
and Grillo et al. (2006) recorded 241 species of flowering (Chrysobalanaceae), Erythroxylum mucronatum Benth.
plants in the PRNP Frei Caneca. In addition, our results (Erythroxylaceae), Margaritaria nobilis L. f., Senefeldera
improved the initial checklists of these authors with verticillata (Vell.) Croizat. (Euphorbiaceae), Myrsine
the not inclusion of 86 species because we had not been guianensis (Aubl.) Kuntze (Primulaceae), Cinnamomum
located neither the field nor herbaria. Families with triplinerve (Ruiz & Pav.) Kosterm. (Lauraceae), Lecythis
the most corrections were Orchidaceae with 17 species, lurida (Miers.) S.A. Mori (Lecythidaceae), Hyeronima
followed by Fabaceae with 11 and Bromeliaceae with oblonga (Tul.) Müll.Arg. (Phyllanthaceae), Quiina
seven. These improvements work together to enhance cruegeriana Griseb. (Quiinaceae), Rubus brasiliensis Mart.
our knowledge of the biodiversity of the area and for (Rosaceae), Allophyllus racemosus Sw. (Sapindaceae),
future management plans of the protected area. Symplocos neglecta Brand (Symplolacaceae) and Urera
Based on the Lista de Espécies da Flora do Brasil (2015) baccifera (L.) Gaudich. ex Wedd. (Urticaceae). However,
and local floras (Sales et al. 1998; Ferraz and Rodal 2008; Cinnamomum triplinerve, Erythroxylum mucronatum and
Almeida-Júnior et al. 2009; Gomes and Alves 2010; Quiina cruegeriana are also present in lowland areas
Amorim and Alves 2012a, 2012b, 2012c; Amorim et al. (Alves-Araújo et al. 2010; Oliveira and Alves 2013; Costa-
2012; Chagas 2012; Alves et al. 2013; Buril 2013; Oliveira Lima 2014) and Myrsine guianensis is commonly found
and Alves 2013; Sampaio 2013; Viana and Barbosa 2013; in restinga plant formations of coastal Brazil (Freitas
Costa-Lima 2014; Pereira 2014; Pessoa and Alves 2015b), and Carrijo 2008; Almeida Jr. et al. 2009; Alves et al.
it was found that 25 spp. (about 3% of the total cataloged 2013). Furthermore, Hyeronima oblonga is a species
for the Serra do Urubu) are new records for the state found in montane forests (Rebster and Huft 1988),
of Pernambuco. This increase to the flora of the state as is Symplocaceae, which is more species-rich in the
includes families such as Fabaceae (6 spp.), Piperaceae, Brazilian Southeast and South (Fritsch et al. 2008;
Poaceae and Rubiaceae (3 spp. each) and Myrtaceae (2 Aranha-Filho and Pedreira 2009).
spp.). These results reiterate the importance of the study Myrtaceae is usually indicated among the most
of local-regional floras for increasing knowledge of plant species-rich families and as an important component in
diversity in the northern portion of the Atlantic Forest. the tree layer (Oliveira-Filho and Fontes 2000; Amorim
As to the proportion of species endemic to the Atlantic et al. 2008; Coelho and Amorim 2014). The number of
Forest, the present study found that 15% of species have species occurring in submontane and montane forests
this distribution pattern, which is well below that found in relation to lowland forests is similar, but different
by Coelho and Amorim (2014), who verified that more in composition. Eugenia hirta O. Berg and Myrciaria
than 37% of species listed by them are endemic to this ferruginea O. Berg are common in the understory of
domain. Meanwhile, the proportion of species found lowland forests of Pernambuco (Amorim and Alves
with disjunct distributions between the Amazon and 2011) but absent in submontane and montane forests,
Atlantic Forest is 10%, which is similar to that found as is also the case with Marlierea excoriata Mart.,
by Rodal et al. (2005b) and Coelho and Amorim (2014). Myrcia bergiana O. Berg , Myrcia densa (DC.) Sobral
Amorim et al. (2009) indicated that this value is similar and Psidium cattleianum Afzel ex Sabine (Amorim and
to what is found in lowland areas of southern Bahia; Alves 2012c). Although there are species shared by both
however, they also found a decrease in this ratio with environments, some are exclusive to submontane and
increased altitude. montane forest in Pernambuco, such as Eugenia culicina
The families Bromeliaceae, Orchidaceae, Myrtaceae Sobral, E. submontana B.S. Amorim & M. Alves and
and Marantaceae had the greatest number of species Myrcia amplexicaulis (Vell.) Hook.f. (Amorim and Alves
endemic to the Atlantic Forest in our results. These 2012, 2015; Sobral 2013).
families are also among those with the greatest The presence of species occurring in montane and
proportion of endemic species of all families recorded lowland forests reveals the transitional nature of
in this area according to Stehmann et al. (2009) and the the Serra do Urubu, which stands at the altitudinal
Lista de Espécies da Flora do Brasil (2015). More than 88% limit adopted for characterization of submontane
of Bromeliaceae species present in the Atlantic Forest and montane forests. This position may be one of the
are endemic to this domain, followed by Myrtaceae with factors contributing to the high species richness found
77.2%, Marantaceae with 73.15%, Melastomatecae with in this study. Ferraz et al. (2008) found evidence that
71%, Orchidaceae with 68.3% and Rubiaceae with 58% the submontane forests of the Serra da Borborema have
a transitional pattern between lowland and montane According to the Livro Vermelho da Flora do Brasil
forests, following the same trend of altitudinal gradients (2013), 10 species recorded in the area are under some
recorded by Gentry (1988). degree of threat: Phragmipedium sargentianum Rolfe
The high richness of some families in the Serra do (Orchidaceae), Threatened with Extinction; Aechmea
Urubu and other montane and submontane forests gustavoi J.A. Siquerira & Leme (Bromeliaceae), Critically
in the Northeastern Brazil (Amorim et al. 2009; Endangered; Canistrum auratiacum E. Morr., Neoregelia
Coelho and Amorim 2014) is distinct from lowland pernambucana Leme & J.A. Siqueira and Vrisea wawranea
areas. Orchidaceae and Bromeliaceae, well-sampled Antoine (Bromeliaceae), Campylocentrum pernambucense
here and in the cited studies, have less richness in Hoehne and Octomeria alexandri Schltr. (Orchidaceae),
the checklists produced by Guedes (1998), Rodal et al. Endangered; and Canistrum pickelii (A.Lima & L.B.Sm.)
(2005a), Soares-Júnior et al. (2008) and Alves et al. Leme & J.A.Siqueira, Guzmania monostachia (L.) Rusby ex
(2013). In southeastern Brazil, according to Oliveira- Mez. (Bromeliaceae), Cattleya labiata Lindl., C. granulosa
Filho and Fontes (2000), the number of species Lindl. (Orchidaceae) and Euterpe edulis Mart. (Arecaceae),
of Chrysobalanaceae, Sapotaceae, Rutaceae and Vulnerable. Additionally, the Lista de Espécies da Flora
Moraceae tends to decrease with increasing altitude, do Brasil (2015) contains six species considered Near
while for Asteraceae, Melastomataceae, Primulaceae Threatened, they are: Anthurium bromelicola Mayo & L.P.
and Solanaceae the numbers tend to increase. This Félix (Araceae), Bowdichia virgilioides Kunth. (Fabaceae),
assertion is partially corroborated in the Serra do Hortia brasiliana Vand. ex DC. (Rutaceae), Lacistema
Urubu, where Chrysobalanaceae and Sapotaceae are robustum Schnizl. (Lacistemataceae), Ocotea puberula
poorly represented (one and three species, respectively), (Rich.) Nees (Lauraceae) and Zygostates bradei (Schltr.)
while in the Usina São José, a lowland forest area in Garay (Orchidaceae). Aside from Vrisea wawranea, which
northern Pernambuco state (Alves et al. 2013), six and was cataloged here only from herbarium material, the
18 species have been cataloged, respectively. The same other species mentioned above were observed in their
difference of number of species in these areas does natural habitat. Canistrum auratiacum, C. pickelii, Euterpe
not fully apply to Rutaceae and Moraceae (Alves et al. edulis and Lacistema robustum are restricted to the
2013). Primulaceae and Solanaceae are richer in Serra do understory of higher altitude areas of the Serra do Urubu.
Urubu than in Usina São José, partially corroborating Furthermore, Giulietti et al. (2009) considered Canistrum
the findings of Oliveira-Filho and Fontes (2000). auratiacum, Neoregelia pernambucana (Bromeliaceae),
Meanwhile, Asteraceae and Melastomataceae are Dichanthelium assurgens (Poaceae) and Borreria decipiens
represented by the same number of species in the two (Rubiaceae) as rare species.
areas mentioned. Amorim et al. (2009) pointed out that In addition to sheltering a large contingent of
Burseraceae, Combretaceae and Connaraceae are better endangered species, the Serra do Urubu has its relevance
represented in lowland areas than in montane and to conservation highlighted by the fact that it supports
submontane areas of Bahia. However, comparing the significant portions of biodiversity of ecologically
floristic checklists of Usina São José (Alves et al. 2013) important families of the Atlantic Forest and the flora of
and Serra do Urubu, this assertion is not confirmed. It is Pernambuco state. In this sense, the family Bromeliaceae
noteworthy that some families such as Bignoniaceae and has 17 genera and 41 species occurring in the area which
Polygalaceae are more diverse in lowland forests (Alves represents 85% of the genera and 40% of the species
et al. 2013), unlike Gesneriaceae and Begoniaceae that recorded for Pernambuco (Forzza et al. 2015). In turn the
are richer in middle and high elevation areas (Clement Serra do Urubu supports 61.5% of the genera and 39%
et al. 2004; Perret et al. 2013). Piperaceae is another of the species of Orchidaceae occurring in the state of
example of the richness of the flora of Serra do Urubu Pernambuco (Barros et al. 2015; Pessoa and Alves 2015b)
being more pronounced in higher altitude forests than and harbors 26.4% of all species of Myrtaceae recorded
in lowland forests (Guedes 1998; Sacramento et al. 2007; in the state. Other families of less floristic relevance
Soares Jr. et al. 2008; Alves et al. 2013). In addition, also have a significant portions of their biodiversity in
some examples are representative, such as Peperomia Pernambuco preserved in the Serra do Urubu. Such is
glabella (Sw.) Dietr., which in the Northeast is known the case with Gesneriaceae, where all genera and 80% of
only from montane areas in the states of Ceará (Serra the species occurring in Pernambuco, Marantaceae with
do Baturité and Maranguape) and Bahia (Serra Bonita 42.3% of the species occurring in the state, Piperaceae
and Serra das Lontras) (Guimarães and Giordano 2004; with 42.8%, Melastomataceae with about 46%, Violaceae
Amorim et al. 2009), P. pseudoestrellensis C. DC., which with 55.6% and Passifloraceae with 68.75% of its species
in the Northeast has been recorded only in southern in the state are recorded in the Serra do Urubu (Lista de
Bahia (Amorim et al. 2009; Leitman et al. 2014), and P. Espécies da Flora do Brasil 2015).
increscens Miq., which occurs in higher areas in the state Furthermore, in the last 10 years (2004–2014) more
(Gomes and Alves 2010). than 15 new species of angiosperms occurring in the
area have been described, some of which were found in of the herbaria visited; the taxonomists consulted (as
the production of this floristic inventory (Chautems et indicated in the species list); our field assistant, Marcos
al. 2005; Leme and Siqueira Filho 2006; Sales et al. 2006; Chagas; and the management teams of the areas studied
Amorim and Alves 2012a, 2015; Costa-Lima and Alves (PRNP Frei Caneca and PRNP Pedra D’Anta).
2013; Giacomin et al. 2013; Sobral 2013; Gregório et al. 2014;
Pessoa et al. 2014a, 2014b, 2015a). Some taxa are known LITERATURE CITED
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