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Description and phylogenetic position of a new species of Nematanthus (Gesneriaceae) from Bahia, Brazil


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Chautems, A. & M. Perret (2017). Description and phylogenetic position of a new species of Nematanthus (Gesneriaceae) from Bahia, Brazil. Candollea 72: 351–359. In English, English abstract. DOI: Nematanthus exsertus Chautems, a new species of Gesneriaceae from the state of Bahia in Brazil, is described. It is easily distinguished from other Nematanthus Schrad. species by its pendent resupinate flowers with funnel-shaped and laterally compressed corolla combined with striking exserted stamens and style. According to the phylogenetic analyses based on nuclear and plastid DNA sequences, this species belongs to a clade including the morphologically distinct Nematanthus monanthos (Vell.) Chautems and four other species with a similar flower morphology but lacking exserted stamen and style. Field photographs accompany the description. The new species is known from one locality in the municipality of Wenceslau Guimarães in the southern part of Bahia state within the “região cacaueira” [cocoa producing area]. The new species is assigned a preliminary assessment of “Vulnerable” using the IUCN Red List Categories and Criteria. Received: May 31, 2017; Accepted: August 17, 2017; First published online: Septembre 27, 2017
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Addresses of the authors :
AC, MP : Conservatoire et Jardin botaniques de la Ville de Genève, C.P. 60, 1292 Chambésy, Switzerland. E-mail :
Submitted on May 31, 2017. Accepted on August 17, 2017. First published online on Septembre 27, 2017.
ISSN : 0373-2967 – Online ISSN : 2235-3658 – Candollea 72(2) : 353-359 (2017)
Description and phylogenetic position
of a new species of Nematanthus
(Gesneriaceae) from Bahia, Brazil
Alain Chautems & Mathieu Perret
CHAUTEMS, A. & M. PERRET (2017). Description and phylogenetic position of a new species of Nematanthus (Gesneriaceae) from Bahia, Brazil.
Candollea 72 : 351-359. In English, English abstract. DOI:
Nematanthus exsertus Chautems, a new species of Gesneriaceae from the state of Bahia in Brazil, is described. It is easily
distinguished from other Nematanthus Schrad. species by its pendent resupinate owers with funnel-shaped and laterally
compressed corolla combined with striking exserted stamens and style. According to the phylogenetic analyses based on
nuclear and plastid DNA sequences, this species belongs to a clade including the morphologically distinct Nematanthus
monanthos (Vell.) Chautems and four other species with a similar ower morphology but lacking exserted stamen and
style. Field photographs accompany the description. e new species is known from one locality in the municipality of
Wenceslau Guimarães in the southern part of Bahia state within the “região cacaueira” [cocoa producing area]. e new
species is assigned a preliminary assessment of “Vulnerable” using the IUCN Red List Categories and Criteria.
GESNERIACEAENematanthus – Brazil – Atlantic Forest – Endemism – Taxonomy – Phylogeny
352 – A new species of Nematanthus (Gesneriaceae) from Brazil Candollea 72, 2017
The genus Nematanthus Schrader includes 31 species, all
endemic to the Brazilian Atlantic Forest (C et al.,
2005 ; F et al., 2016). Nematanthus species are epi-
phytic perennial herbs or subshrubs, less frequently growing
on rocky substrates in humus pockets. Blossoms display a
large range of morphologies among species, including short
or long pedicelate, brightly colored owers, with resupinate,
or non-resupinate owers. Flowers that are non-resupinate are
pouched. Flowers that are resupinate are funnel-shaped and
laterally compressed, but in some taxa, owers are white and
bell-shaped. Hummingbird pollination has been documented
in the eld for several species and is likely the dominant pol-
lination mode in the genus, according to ower morphology
(S-S et al., 2017). However, owers matching
the bee-pollination have also evolved in a clade of three Nema-
tanthus species (S-S et al., 2015). Despite this
large morphological variation, the monophyly of Nematanthus
has been conrmed by phylogenetic data and traits such as
ower resupination and bee syndrome have been identied
as synapomorphies of specic clades (F et al., 2016 ;
S-S et al., 2015).
The Brazilian Atlantic Forest is recognized as a global
biodiversity hotspot and studies focused on forest remnants
in the State of Bahia in eastern Brazil have revealed a higher
species richness in trees, herbs, vines and epiphytes than in
other areas from southeastern Brazil (A et al., 2005,
2009 ; T, 2008). During fieldwork in the county of
Wenceslau Guimarães in southern Bahia state, plants with
resupinate red owers and exserted anthers and style were
fortuitously discovered by the rst author while walking in a
fragment of well-preserved rain forest. Here we recognize and
describe this new species and used two nuclear loci and seven
plastid DNA regions to investigate its phylogenetic placement
within the genus. Morphology of the new species is discussed
and compared with other closely related and sympatric species.
Field photographs and distribution map are provided as well
as a preliminary risk of extinction assessment using the IUCN
Red List Categories and Criteria (IUCN, 2012).
Material and methods
Specimens of the newly identified species were collected
during eldwork carried out in May 2007 in the county of
Wenceslau Guimarães, southern Bahia state, Brazil. All
pressed material is deposited in the CEPEC and NY herbaria
(acronyms according to I H, 2017). New
collections were later registered from the same area in 2012
and material was deposited in the RB herbarium. e descri-
ption and analysis of the new species were based on fresh and
dried specimens. Data for comparisons with other species were
taken from previous studies (C, 1988, C &
M, 2003) ; directly from herbarium specimens mostly
at G, NY, RB ; from fresh specimens kept in cultivation at the
Conservatoire et Jardin botaniques de la Ville de Genève ; or at
Mauro Peixoto’s private collection in sitio Primavera, Mogi das
Cruzes, São Paulo, Brazil. e map was created using ArcGIS
10 (ESRI Inc.).
To assess the phylogenetic position of the new taxon, we
performed a phylogenetic analysis based on DNA nucleotide
sequence data. A DNA sample of the type collection was ana-
lyzed with sequences data obtained from other representative
species of the genera Nematanthus Schrad. (26 of 31 species),
Codonanthe (Mart.) Hanst. (8 of 8 species), Codonanthopsis
Mansf. (7 of 13 species), Lesia J.L. Clark & J.F. Sm. (1 of
2 species). Two outgroups, Drymonia serrulata (Jacq.) Mart.
and Chrysothemis pulchella (Donn ex Sims) Decne., were
selected according to previous phylogenetic analyses (P
et al., 2013). Phylogenetic relationsips among these taxa were
reconstructed using seven plastid DNA regions (atpB-rbcL,
matK, rps16, rpl16, trnT-trnL, trnL-trnF and trnS-trnG) and
two nuclear regions, the ribosomal internal transcribed spacer
(ITS) and a portion of the plastid-expressed glutamine syn-
thetase gene (ncpGS). New DNA sequences were generated
for trnT-trnL, trnS-trnG and for the new species, N. exsertus,
following the procedure described in P et al. (2003) and
F et al. (2016). All other sequences were obtained
from previous works by S-S et al. (2015) and
F et al. (2016). Newly acquired sequences have been
deposited in GenBank. Voucher information and GenBank
numbers for each sequence used in this study are provided in
Appendix 1.
Newly acquired sequences were manually added to
available alignments in Mesquite 3.03 (M & M-
, 2015). Phylogenetic analyses were performed on the
CIPRES portal in San-Diego, USA [http ://]
using maximum likelihood (ML) and Bayesian Inference
(BI). Maximum likelihood analyses were conducted using the
software RAxML v.8.2.10 (S, 2014) with a rapid
bootstrap analysis followed by the search of the best-scoring
ML tree in one single run. Bootstrap analysis stopped after
456 replicates, which were sucient for getting stable support
values according to the bootstrap convergence test using the
extended majority-rule consensus tree criterion (autoMRE).
e default model, GTRCAT, was used to perform the ML
analyses. Bayesian inference analyses were performed using
MrBayes 3.2.3 (R et al., 2012) following the proce-
dure described in S-S et al. (2015). For each
DNA region, the optimal substitution models were assessed
using jModelTest2 (D et al., 2012) according to the
Akaike information criterion (AIC). e best-t model was
HKY + G for ncpGS, and GTR + G for all other genes. All
the parameters values were unlinked across gene partitions and
estimated during the Markov chain Monte Carlo (MCMC)
Candollea 72, 2017 A new species of Nematanthus (Gesneriaceae) from Brazil – 353
runs. Two independent analyses were run from different
random trees. e chains were run for 20,000,000 generations,
with trees sampled every 1000th generation. We determined
chain convergence and burn-in length (20% of the sampled
generations) by examining trace plots of each parameter in
Tracer v.1.6 (R et al. 2014). A consensus tree was
calculated by removing the burn-in period and combining
the two runs.
Results and discussion
e topology of the phylogenetic trees resulting from the ML
and BI analyses are identical and are summarized in Fig. 1. e
result is also congruent with the maximum clade credibility
tree described in S-S et al. (2015) showing
that Nematanthus species are clustered in five main clades
exhibiting contrasting flower morphologies. Species with
resupinate flowers and hummingbird syndrome belong to
clades I and III, whereas the three species with white non-resu-
pinate owers and bee pollination syndrome belong to clade II
(Fig. 1). e Nematanthus species with non-resupinate owers
Fig. 1. –
Bayesian 50 % majority rule consensus tree of Nematanthus resulting from the combined analysis of plastid loci atpB-rbcL, matK, rps16, rpl16,
trnT-trnL, trnL-trnF, trnS-trnG, and the nuclear regions ncpGS and ITS. Numbers above branches are Bayesian posterior probabilities. Numbers below
branches are maximum likelihood bootstrap when ≥50 %. Asterisks indicate species with funnel-shaped and laterally compressed corollas.
354 – A new species of Nematanthus (Gesneriaceae) from Brazil Candollea 72, 2017
Fig. 2. –
Nematanthus exsertus Chautems. A. Habit with pendent pedicels in native habitat ; B. Flowering shoots ; C. Close-up of shoot showing stem
and petiole indumentum ; D. Close-up of corolla showing exserted stamens and style.
[Jardim et al.
1 cm
3 cm
Candollea 72, 2017 A new species of Nematanthus (Gesneriaceae) from Brazil – 355
and hypocyrtoid (or pouched) corolla are found in clades IV
and V (Fig. 1). Our analysis support that the new species, N.
exsertus, nests in a well-supported clade (bootstrap support, BS
= 90 %, Bayesian posterior probability, PP = 1) with four other
resupinate species (N. brasiliensis, N. crassifolia, N. uminensis
and N. monanthos, Fig. 1). Nematanthus exsertus is sister to
N. monanthos, but interspecic relationships within this clade
are not supported (BS < 50%). Although the new species
can be readily distinguished from N. monanthos, it presents
morphological affinities with N. brasiliensis, N. corticola,
N. crassifolia and N. uminensis. All ve species in this group
are characterized by the presence of pendent flowers with
elongate pedicels with funnel-shaped corolla tubes that
appear laterally compressed in the throat. However, all of the
other species have inserted stamens and pistil. In contrast,
N. exsertus (Fig. 2) is unique by the presence of exserted
stamens and pistil. Scarlet-red corollas are found in N. exsertus,
N. corticola and N. crassifolius, whereas N. brasiliensis displays a
yellow corolla with red stripes and N. uminensis a plain yellow
corolla. However, none of these species have stamens and
pistil as exserted as in N. exsertus (Fig. 2). Scarlet-red corolla
is found in N. exsertus, N. corticola and N. crassifolius, whereas
N. brasiliensis displays a yellow corolla with red stripes and
N. uminensis a plain yellow corolla.
The only closely related species that is sympatric with
N. exsertus is N. corticola (Fig. 3). ese two species are easily
dierentiated when fertile or sterile. When sterile, they can
be dierentiated by the presence of villous indumentum on
the stem in N. exsertus in contrast to the glabrous stems in
N. corticola. In flower, N. exsertus is differentiated by the
presence of exserted stamens and style and shorter corolla tube
with narrower gibbosity (Table 1).
Taxonomic treatment
Nematanthus exsertus Chautems, spec. nova (Fig. 2).
Typus : Brazil. Bahia : Município de Wenceslau Gui-
marães, Estação Ecológica Estadual Nova Esperança,
trilha para o Rio Agua Vermelha, 415-450 m, 13°35’43’’S
39°43’10’’W, 3.V.2007, Jardim et al. 5000 (holo- : CEPEC-
117040! ; iso- : NY!).
Nematanthus exsertus Chautems resembles N. corticola Schrad.
with similar elongate pedicels, short and narrow calyx lobes and
funnel-shaped, laterally compressed red corollas, but diers by
a villous indumentum on stem and petioles, as well as shorter
and more ventricose corollas and presence of exserted stamens
and style.
Subshrub 0.5-1.2 m, epiphytic ; stem ascending, sparingly
branched, 3-6 mm diam., villose, especially in young parts ;
internodes 1.5-5 cm. Leaves strongly anisophyllous ; petioles
0.5-4 cm, dark vinaceous, pubescent-hirsute ; blades 2.5-5 3
0.8-2 cm (smaller ones) to 6–14 3 1.8-3 cm (larger ones),
elliptic, adaxially green, pubescent, abaxially paler green, gla-
brescent, apex acute to acuminate, base cuneate, margin entire,
3-6 lateral veins each side of the midrib, vinaceous, pubescent.
Flowers solitary, 1-3 per stem, pendent-resupinate, in leaf axils,
bracts minute or early caducous ; pedicels 10-18 cm long, vina-
ceous, pubescent. Calyx fused for 3-4 mm at base, sepals 7-8
3 2-3 mm, linear lanceolate, pubescent, vinaceous, apex acute,
greenish, margin entire or remotely toothed. Corolla 3-3.5 cm
long, scarlet-red, obliquely inserted in calyx, funnel-shaped,
laterally compressed on ventral side before mouth, pubescent,
base shortly cylindric, 4-5 mm diam., gradually enlarged to
2-2.2 cm at the opening, lobes revolute at anthesis ; stamens
and style exserted for about 2.5-2.8 cm beyond the corolla
opening, laments 4.5-5 cm, not coiling at the end of anthesis,
N. corticola N. crassifolius N. exsertus
Stem indument glabrous glabrous villose
Petiole indument Sparsely pubescent glabrous villose
Pedicel length [cm] 10-20 3-10(-16) 10-18
Calyx lobes length [mm] 7-12 15-30 8-10
Calyx lobes width [mm] 2-4 4-8 2-3
Corolla length [cm] 3.8-5 4-5 3-3.5
Corolla width at max. gibbosity [cm] 1.4-2 1.8-2.4 2-2.2
Stamens position included included strongly exserted
Distribution BA in most of the southern Montane rain forest in ES, Restricted to one locality in
hylea rain forest RJ, southeastern MG and southern BA
marginally in SP
Table 1. –
Key morphological differences between Nematanthus exsertus Chautems, N. corticola Schrad. and N. crassifolius (Schott) Wiehler ;
abbreviations in capital letters stands for the Brazilian states of Bahia (BA), Espírito Santo (ES),
Minas Gerais (MG), Rio de Janeiro (RJ), and São Paulo (SP).
356 – A new species of Nematanthus (Gesneriaceae) from Brazil Candollea 72, 2017
white, glabrous, pollen cream ; nectary gland bilobed, 3-4 3
2-3 mm, cream, ovary 4-6 mm long, vinaceous, pubescent, style
4.8-5.2 mm, white, glabrous. Fruit unknown.
Etymology. – e epithet “exsertus” refers to the exserted
stamens and style that extend beyond the corolla opening for
at least 2.5 cm.
Distribution, ecology and phenology. – Nematanthus exsertus
is currently known by two collections made in the “Estação
Ecológica Estadual Nova Esperança” in the municipality of
Wenceslau Guimarães in the state of Bahia (Fig. 3). e individuals
were observed growing epiphytically on small trees at 1-2 m above
the ground in the understory of a fragment of wet and dense forest
on at grounds. Specimens were collected with owers in January
and May. According to ower morphology, the species is likely
pollinated by hummingbirds. Pollination by hermit hummingbirds
has been documented for the related species N. crassifolia and
N. brasiliensis presenting similar pendent and resupinate owers
with laterally compressed corolla (SM-G &
V, 2010 ; W et al., 2013).
Conservation status. – Nematanthus exsertus is known only
from a single location of a dozen individuals in an area of
preservation created in 1997 and extended in 2,000, covering
2,418 ha. of Atlantic Rain Forest (“Estação Ecológica Estadual
Nova Esperança”). is area is nevertheless subject to local
preservation conicts, like deforestation, illegal hunting and
illegal occupation [
de-wenceslau-guimaraes]. erefore, with a restricted AOO,
plausible threats that could rapidly push the species to “Criti-
cally Endangered”, N. exsertus is assigned a preliminary assess-
ment of “Vulnerable” [VU D2] using the IUCN Red List
Categories and Criteria (IUCN, 2012).
Notes. – Our first thought, at the time of discovering this
species, was that we had found an individual of Nematanthus
corticola with ill-formed stamens and style, because this taxon
occurs frequently in the southern Bahian rain forest (Fig. 3). After
having observed few individuals, each bearing several owers, it
was readily apparent that we had discovered a new species.
e striking exserted position of stamens and style are a
unique feature in the genus Nematanthus. Nematanthus exsertus
resembles N. corticola with similar pedicel, calyx lobes, corollas,
but diers by a villous indumentum on stem and petioles, as
well as shorter and more ventricose corollas with strikingly
exserted stamens and style (see Table 1).
e total number of known species for the genus Nema-
tanthus is now 32. e new species should be added to the
recent checklist that established a high endemism rate of epi-
phytic plants within the eastern Brazilian Atlantic rain forest
(F et al., 2016). It is also an addition to the number of
Gesneriaceae species that have been registered so far for the
state of Bahia, the total number of registered species reaches
now 29 (C, 1991 ; A et al., 2017).
Paratypus. Brazi l. Bahia : Município de Wenceslau
Guimarães, Reserva Estadual de Wenceslau Guimarães, Trilha
da Petioba, 13°34’49’’S 39°42’17’’W, 411 m, 19.01.2012, Mon-
teiro et al. 545 (RB-536512).
We are grateful to André Amorim and Jomar Jardim for invi-
ting the rst author to join a eld excursion in 2007, Cristiane
I. Aguiar for facilitating access to the collection and providing
an image of the type material at CEPEC herbarium, Mauro
Peixoto for providing access of owering material cultivated in
his collection (sitio Primavera, Mogi das Cruzes, SP, Brazil).
We thank Régine Niba for the lab work and Nicolas Wyler
for the preparation of the distribution map.
Fig. 3. –
Distribution map of Nematanthus exsertus Chautems (star)
showing also the occurrences of morphologically related species
N. corticola Schrad. (circles) and N. crassifolius (Schott) Wiehler
Candollea 72, 2017 A new species of Nematanthus (Gesneriaceae) from Brazil – 357
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358 – A new species of Nematanthus (Gesneriaceae) from Brazil Candollea 72, 2017
Candollea 72, 2017 A new species of Nematanthus (Gesneriaceae) from Brazil – 359
Taxon Voucher ITS ncpGS matK trnL-F rps16 atpB-rbcL rpl16 trnT-L trnS-G
Chrysothemis pulchella Araujo et al. 602 (G) KT958331 KT958276 JX195977 GQ383544 GQ383582 KT958447 KT958388 MF787865 MF787823
Codonanthe carnosa Chautems & Perret 08-001 (G) KT958316 KT958261 JX195956 JX195728 JX195812 KT958433 KT958373 MF787867 MF787825
Codonanthe cordifolia Chautems & Perret 10-103 (G) KT958317 KT958262 JX195958 JX195730 JX195814 KT958434 KT958374 MF787868 MF787826
Codonanthe devosiana Chautems & Perret 99-010 (G) KT958320 KT958265 JX195962 JX195734 JX195817 KT958437 KT958377 MF787871 MF787829
Codonanthe gibbosa Kollmann et al. 4784 (MBML) KT958323 KT958268 JX195965 JX195737 JX195820 KT958439 KT958380 MF787872
Codonanthe gracilis Chautems & Perret 99-003 (G) KT958324 KT958269 JX195968 JX195740 JX195823 KT958440 KT958381 MF787873 MF787830
Codonanthe mattos-silvae Chautems & Perret 99-033 (G) KT958327 KT958272 JX195971 JX195743 JX195826 KT958443 KT958384 MF787875 MF787832
Codonanthe serrulata Chautems & Perret 99-045 (G) KT958328 KT958273 JX195972 AJ439826 GQ383583 KT958444 KT958385 MF787876 MF787833
Codonanthe venosa Chautems & Perret 99-006 (G) KT958330 KT958275 JX195974 JX195745 JX195827 KT958446 KT958387 MF787878 MF787835
Codonanthopsis calcarata Cultivated at CJBG AC-3305 KT958314 KT958259 JX195954 JX195726 JX195810 KT958432 KT958371 MF787866 MF787824
Codonanthopsis corniculata Chautems & Perret 99-043 (G) KT958318 KT958263 JX195959 JX195731 JX195815 KT958435 KT958375 MF787869 MF787827
Codonanthopsis crassifolia Chautems & Perret 06-102 (G) KT958319 KT958264 JX195960 JX195732 – KT958436 KT958376 MF787870 MF787828
Codonanthopsis dissimulata Perret & Chautems 79 (G) KT958335 KT958280 JX195984 JX195753 JX195835 KT958451 MF787879 MF787836
Codonanthopsis macradenia Chautems & Perret 10-107 (G) KT958326 KT958271 JX195970 JX195742 JX195825 KT958442 KT958383 MF787874 MF787831
Codonanthopsis uleana Chautems & Perret 02-105 (G) KT958329 KT958274 JX195973 JX195744 – KT958445 KT958386 MF787877 MF787834
Codonanthopsis ulei Chautems & Perret 07-002 (G) KT958336 KT958281 JX195985 JX195754 JX195836 KT958452 KT958392 MF787880 MF787837
Drymonia serrulata Araujo et al. 601 (G) KT958337 KT958282 JX195992 GQ383548 GQ383587 KT958453 KT958393 MF787881 MF787838
Lesia tepuiensis Ferreira 262 (INPA) KX011577 KX011561 KX011545 KX011539 KX011542 KX011551 KX011569 MF787882
Nematanthus albus Chautems & Perret 99-012 (G) KT958341 KT958286 JX196015 JX195765 JX195848 KT958457 KT958397 MF787883 MF787839
Nematanthus albus Carnauba et al. 20.11.2013 MF787816 MF787931 MF787802 MF787909 MF787924 MF787917 MF787884 MF787840
Nematanthus albus Cultivated at CJBG AC-2705 MF787817 MF787810 MF787932 MF787803 MF787910 MF787925 MF787918 MF787885 MF787841
Nematanthus australis Chautems & Perret 99-011 (G) KT958342 KT958287 JX196016 JX195766 JX195849 KT958458 KT958398 MF787886 MF787842
Nematanthus brasiliensis Chautems & Perret 07-303 (G) KT958344 KT958288 JX196017 JX195767 JX195850 KT958459 KT958399 MF787887 MF787843
Nematanthus corticola Chautems & Perret 99-013 (G) KT958345 KT958289 JX196018 JX195768 JX195851 KT958460 KT958400 MF787888 MF787844
Nematanthus crassifolius Chautems & Perret 07-403 (G) KT958346 KT958290 JX196019 JX195769 JX195852 KT958461 KT958401 MF787889 MF787845
Nematanthus exsertus Jardim et al. 5000 (CEPEC) MF787818 MF787809 MF787936 MF787808 MF787916 MF787926 MF787919 MF787890 MF787846
Nematanthus fissus Ferreira 269 (INPA) MF787819 MF787811 MF787804 MF787911 MF787891
Nematanthus fluminensis Chautems & Perret 99-020 (G) KT958348 JX196020 JX195770 JX195853 KT958462 KT958402 MF787892 MF787847
Nematanthus fornix Chautems & Perret 10-101 (G) KT958349 KT958292 JX196021 JX195771 JX195854 KT958463 KT958403 MF787893 MF787848
Nematanthus fritschii Chautems & Perret 07-404 (G) KT958350 KT958293 JX196022 JX195772 JX195855 KT958464 KT958404 MF787894 MF787849
Nematanthus gregarius Chautems & Perret 99-019 (G) KT958351 KT958294 JX196023 JX195773 JX195856 KT958465 KT958405 MF787895 MF787850
Nematanthus hirtellus Perret, Chautems et al. 61 (G) MF787820 MF787812 MF787933 MF787805 MF787912 MF787927 MF787920 MF787851
Nematanthus hirtellus Cultivated at CJBG AC-3605 MF787821 MF787813 MF787806 MF787913 MF787928 MF787921 MF787896
Nematanthus jolyanus Chautems & Perret 99-028 (G) KT958353 KT958296 JX196025 JX195775 JX195858 KT958467 KT958407 MF787897 MF787852
Nematanthus lanceolatus Chautems & Perret 12-102 (G) KY858387 MF787815 MF787935 KY858634 MF787915 MF787930 MF787923 MF787854
Nematanthus lanceolatus Chautems & Perret 99-023 (G) MF787822 MF787814 MF787934 MF787807 MF787914 MF787929 MF787922 MF787898 MF787853
Nematanthus maculatus Chautems & Perret 07-405 (G) KT958356 JX196027 JX195777 JX195860 KT958469 KT958409 MF787899 MF787855
Nematanthus monanthos Chautems & Perret 08-601 (G) KT958357 KT958299 JX196028 JX195778 JX195861 KT958470 KT958410 MF787900 MF787856
Nematanthus punctatus Chautems & Perret 99-046 (G) KT958358 KT958300 JX196029 JX195779 JX195862 KT958471 KT958411 MF787901 MF787857
Nematanthus pycnophyllus Chautems & Perret 02-010 (G) KT958359 KT958301 JX196030 JX195780 JX195863 KT958472 KT958412 MF787902 MF787858
Nematanthus sericeus Chautems & Perret 99-018 (G) KT958360 KT958302 JX196031 JX195781 JX195864 KT958473 KT958413
Nematanthus strigillosus Chautems & Perret 11-101 (G) KT958362 KT958304 JX196033 JX195783 JX195866 KT958475 KT958415 MF787903 MF787859
Nematanthus teixeiranus Chautems & Perret 07-407 (G) KT958363 KT958305 JX196035 JX195785 JX195868 KT958476 KT958416 MF787904 MF787860
Nematanthus tessmannii Chautems & Perret 07-408 (G) KT958364 KT958306 JX196036 JX195786 JX195869 KT958477 KT958417 MF787905 MF787861
Nematanthus villosus Chautems & Perret 99-041 (G) KT958365 KT958307 JX196037 AJ439825 GQ383608 KT958478 KT958418 MF787906 MF787862
Nematanthus wettsteinii Chautems & Perret 10-102 (G) KT958366 KT958308 JX196038 JX195787 JX195870 KT958479 KT958419 MF787907 MF787863
Nematanthus wiehleri Chautems & Perret 12-101 (G) KT958367 KT958309 JX196039 JX195788 JX195871 KT958480 KT958420 MF787908 MF787864
Appendix 1. – List of sequences used in the phylogenetic analyses. Taxa, voucher information, origin and Genbank accession numbers for
the nine DNA regions (new sequences in bold).
... Thus, the use of data extrapolation can reveal areas with a high number of endemic species recorded in a single densely collected cell, such as the cell located NW of the central-eastern AoE (Fig. 1, highlighted in red), where 26 endemic species contributed to its recognition as an AoE. This cell includes the "Estação Ecológica Estadual Wenceslau Guimarães," a protected forest remnant of about 2.4 ha (Rigueira et al. 2012), with several new endemic species of angiosperms described in recent years (Bacci et al. 2016;Goldenberg et al. 2016;Chautems and Perret 2017). This protected area is only recovered as part of an AoE when data extrapolation is applied (Fig. 2d-f), but due to the likely occurrence of many other endemic species in the region, it should be subjected to more intensive botanical exploration. ...
Given the importance of understanding endemic species distribution for conservation strategies and the status of the Atlantic Forest as a conservation hotspot, we carried out a study to evaluate where are located the areas of endemism (AoE) at Bahia Coastal Forests (BCF), a threatened and species rich area of this hotspot. To achieve this aim, we recovered and filtered the occurrence data of 547 angiosperm taxa endemic to BCF and applied endemicity analyses using different grid cell sizes (5′ × 5′, 10′ × 10′, and 15′ × 15′) and data extrapolation options. A consensus rule was applied to the resulting areas, joining AoE that shared at least 40% of its species. Most of the recovered AoE are located in the central region of BCF, which might be related to a higher humidity compared to northern and southern areas, which had fewer areas recovered. These results could also reflect the higher sampling effort in the central region of BCF, as this area had most intensive botanical studies so far. The use of extrapolation options generated more AoE regardless of cell size; this technique indicated potential AoE that would not have been detected otherwise due to low sampling effort, such as the vicinities of Wenceslau Guimarães at northwestern BCF. We recommend that more botanical exploration should be done at BCF northern and southern areas to overcome sampling bias and define more accurately the number and limits of AoE, highlighting the need for more focused conservation strategies in this hot-point within the Atlantic Forest.
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O estudo das epífitas vasculares foi realizado em uma área de 3.000 m² de floresta na planície litorânea na Ilha do Mel (25°30? S 48°23? W); o levantamento qualitativo foi realizado em toda a área; para o estudo quantitativo, foram sorteados 100 forófitos, divididos em intervalos de 2 m a partir do solo. Em cada intervalo, registraram-se todas as espécies epifíticas ocorrentes, sendo estimado o valor de importância epifítico a partir das freqüências nos estratos, nos forófitos e nas espécies forofíticas. No levantamento total, foram encontradas 77 espécies (16 de Pteridophyta, 53 de Liliopsida e 8 de Magnoliopsida), das quais 70 foram amostradas nos forófitos analisados. As famílias mais ricas foram Orchidaceae, Bromeliaceae e Polypodiaceae e os gêneros foram Vriesea, Encyclia e Maxillaria. A área com maior similaridade florística com este estudo localiza-se no Município de Torres (RS). As espécies mais importantes quantitativamente foram Microgramma vaccinifolia, Codonanthe gracilis, Epidendrum latilabre e E. rigidum. As espécies amostradas foram agrupadas em três categorias quanto à preferência por intervalos de altura: exclusivas, preferenciais e indiferentes. O número de ocorrências de epífitos em um mesmo forófito variou de 1 a 35, enquanto o número de espécies variou de 1 a 21 (médias 14 ± 7,6 e 10 ± 4,6, respectivamente). Os primeiros estratos (0-2 m, 2-4 m e 4-6 m) foram os mais ricos em espécies epifíticas. As espécies forofíticas com maior número de ocorrências foram Andira fraxinifolia e Ternstroemia brasiliensis, e com maior número de espécies Ocotea pulchella e Guapira opposita. Quanto à fidelidade sobre espécies forofíticas, foram encontradas espécies epifíticas exclusivas, preferenciais e generalistas, esta incluindo a maioria das espécies amostradas.
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The study was carried out in a 3,000 m2 area of coastal Atlantic rain forest at Ilha do Mel island (25 degrees 30"S 48 degrees 23'W), on 100 assorted trees separated into 2 meter-high strata starting from the ground. In each stratum all of the occurring epiphytic species were recorded. The sampled species were grouped into three categories: exclusive, preferential, and indifferent, according to their abundance in each strata, and selective, preferential and indifferent, according to abundance on the forophytes. Intermediate strata registered the highest diversity. Six species were considered exclusive to one or two strata, 15 were restricted to some strata and 5 presented a broad distribution. No epiphytic species showed uniform horizontal distribution on the area. The epiphyte richness in a host tree varied from zero to 30. Regarding to fidelity on host tree species, few selective or preferential, and mainly indifferent epiphyte species, were found. A total of 82 epiphyte species were sampled in the surveyed tree, and the Wittaker plot indicate a highly dominant assemblage.
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The effects of specific functional groups of pollinators in the diversification of angiosperms are still to be elucidated. We investigated whether the pollination shifts or the specific association with hummingbirds affected the diversification of a highly diverse angiosperm lineage in the Neotropics. We reconstructed a phylogeny of 583 species from the Gesneriaceae family and detected diversification shifts through time, inferred the timing and amount of transitions among pollinator functional groups, and tested the association between hummingbird pollination and speciation and extinction rates. We identified a high frequency of pollinator transitions, including reversals to insect-pollination. Diversification rates of the group increased through time since 25 Mya, coinciding with the evolution of hummingbird-like flowers and the arrival of hummingbirds in South America. We showed that plants pollinated by hummingbirds have a two-fold higher speciation rate compared to plants pollinated by insects, and that transitions among functional groups of pollinators had a little impact on the diversification process. We demonstrated that floral specialization on hummingbirds for pollination has triggered rapid diversification in the Gesneriaceae family since the early Miocene, and that it represents one of the oldest identified plant-hummingbird associations. Biotic drivers of plant diversification in the Neotropics could be more related to this specific type of pollinator (hummingbirds), than to shifts among different functional groups of pollinators.
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We describe and illustrate Lesia tepuiensis, a new species of subshrub from rock outcrops of the Serra do Aracá tepui in Amazonas, Brazil. Phylogenetic analyses based on 7219-aligned base pairs of the plastid and nuclear DNA sequences recovered the new species as sister to Lesia savannarum, the type species of Lesia, a genus recently described as monotypic. The new species is readily distinguished from L. savannarum by having a red, pouched corolla tube, lanceolate calyx lobes, a hirsute to tomentose leaf indument and a rupicolous habit. Phylogenetic analyses of the relationships of Lesia with other genera in the subtribe Columneinae indicated that Lesia and Codonanthopsis form a monophyletic group sister to a clade endemic to the Brazilian Atlantic forest formed by Nematanthus and Codonanthe. The pouched corolla tube found in the genera Lesia and Nematanthus appears to have evolved independently in these two lineages. Ecological and distributional data, as well as a key to the species of Lesia and related genera, are also provided.
Full-text available
Knowledge of the geographic distribution of plants is essential to underpin the understanding of global biodiversity patterns. Vascular epiphytes are important components of diversity and functionality of Neotropical forests but, unlike their terrestrial counterparts, they are under-represented in large-scale diversity and biogeographic analyses. This is the case for the Atlantic Forest - one of the most diverse and threatened biomes worldwide. We provide the first comprehensive species list of Atlantic Forest vascular epiphytes; their endemism patterns and threatened species occurrence have also been analyzed. A list with 2,256 species of (hemi-)epiphytes - distributed in 240 genera and 33 families - is presented based on the updated Brazilian Flora Checklist. This represents more than 15% of the total vascular plant richness in the Atlantic Forest. Moreover, 256 species are included on the Brazilian Red List. More than 93% of the overall richness is concentrated in ten families, with 73% represented by Orchidaceae and Bromeliaceae species alone. A total of 78% of epiphytic species are endemic to the Atlantic Forest, in contrast to overall vascular plant endemism in this biome estimated at 57%. Among the non-endemics, 13% of epiphytic species also occur either in the Amazon or in the Cerrado - the other two largest biomes of Brazil – and only 8% are found in two or more Brazilian biomes. This pattern of endemism, in addition to available dated phylogenies of some genera, indicate the dominance of recent radiations of epiphytic groups in the Atlantic Forest, showing that the majority of divergences dating from the Pliocene onwards are similar to those that were recently reported for other Neotropical plants.
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Major factors influencing the phenotypic diversity of a lineage can be recognized by characterizing the extent and mode of trait evolution between related species. Here, we compared the evolutionary dynamics of traits associated with floral morphology and climatic preferences in a clade composed of the genera Codonanthopsis, Codonanthe and Nematanthus (Gesneriaceae). To test the mode and specific components that lead to phenotypic diversity in this group, we performed a Bayesian phylogenetic analysis of combined nuclear and plastid DNA sequences and modeled the evolution of quantitative traits related to flower shape and size and to climatic preferences. We propose an alternative approach to display graphically the complex dynamics of trait evolution along a phylogenetic tree using a wide range of evolutionary scenarios. Our results demonstrated heterogeneous trait evolution. Floral shapes displaced into separate regimes selected by the different pollinator types (hummingbirds versus insects), while floral size underwent a clade-specific evolution. Rates of evolution were higher for the clade that is hummingbird pollinated and experienced flower resupination, compared with species pollinated by bees, suggesting a relevant role of plant-pollinator interactions in lowland rainforest. The evolution of temperature preferences is best explained by a model with distinct selective regimes between the Brazilian Atlantic Forest and the other biomes, whereas differentiation along the precipitation axis was characterized by higher rates, compared with temperature, and no regime or clade-specific patterns. Our study shows different selective regimes and clade-specific patterns in the evolution of morphological and climatic components during the diversification of Neotropical species. Our new graphical visualization tool allows the representation of trait trajectories under parameter-rich models, thus contributing to a better understanding of complex evolutionary dynamics.
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The Serra do Teimoso, in southern Bahia, Brazil, is a mountain reaching 850 m at the transition from tropical moist forest to tropical semideciduous forest. A floristic survey of a 200 ha reserve established on the mountain was carried out and a checklist of the vascular flora produced. The survey was conducted by random collecting efforts and the sampling of all specimens ≥5 cm diameter within a one hectare plot. The flora of the Serra do Teimoso Reserve (STR) comprised 727 species in 400 genera and 119 families. The angiosperms comprised 667 species in 363 genera and 100 families and the pteridophytes included 60 species in 37 genera and 19 families. Floristic relationships of the STR flora were discussed with regard to some florest types of eastern Brazil, especially those found at southern Bahia. La Serra do Teimoso, en el sur de Bahia, Brasil, es una montaña alcanzando 850 m en la transición de los bosques tropicales húmedos hacia los bosques tropicales semideciduales. Un inventario floristico de una reserva forestal con ca. 200 ha establecida en la montaña fue hecho y un checklist producido. El levantamiento fue conducido por medio de esfuerzos de coleta aleatorios y el muestro de todos los espécimens ≥5 cm DAP dentro de urn plot de urna hectárea. La flora de plantas vasculares da la Reserva Serra do Teimoso (STR) incluió 727 especies en 400 géneros y 119 familias. Las angiospermas incluiron 667 especies en 363 géneros y 100 familias, y las pteridofitas incluiron 60 especies en 37 géneros y 19 familias. Relacciones floristicas de la flora de STR fueran discutidos con base en algunos tipos de bosques del este de Brasil, en especial aquellos localizados en el sur de Bahia.
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Five new Nematanthus species are described, mapped, and illustrated: N. albus Chautems; N. kautskyi Chautems & J.Rossini; N. punctatus Chautems; N. pycnophyllus Chautems, T.Lopes & M.Peixoto; and N. wiehleri Chautems & M.Peixoto. Their conservation status following IUCN criteria is given. Three species are melittophilous, adding a new pollination syndrome to the genus. The circumscription of Nematanthus therefore is redefined and includes 31 species. A revised key to the whole genus is provided. Cinco novas espécies de Nematanthus são descritas, mapeadas, e ilustradas: N. albus Chautems; N. kautskyi Chautems & J.Rossini; N. punctatus Chautems; N. pycnophyllus Chautems, T.Lopes & M.Peixoto; e N. wiehleri Chautems & M.Peixoto. O estatuto de conservaçao é estabelecido seguindo os critérios da UICN. Tres espécies são melitófilas, adicionando uma nova síndrome de polinizaçâo ao género. Nematanthus é assim redefinido e inclui 31 espécies. Uma chave de identificação revisada para todas as espécies do género é apresentada.
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Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community. I present some of the most notable new features and extensions of RAxML, such as, a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX, and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date, 50 page user manual covering all new RAxML options is available. The code is available under GNU GPL at
Gesneriaceae are represented in the New World (NW) by a major clade (c. 1000 species) currently recognized as subfamily Gesnerioideae. Radiation of this group occurred in all biomes of tropical America and was accompanied by extensive phenotypic and ecological diversification. Here we performed phylogenetic analyses using DNA sequences from three plastid loci to reconstruct the evolutionary history of Gesnerioideae and to investigate its relationship with other lineages of Gesneriaceae and Lamiales. Our molecular data confirm the inclusion of the South Pacific Coronanthereae and the Old World (OW) monotypic genus Titanotrichum in Gesnerioideae and the sister-group relationship of this subfamily to the rest of the OW Gesneriaceae. Calceolariaceae and the NW genera Peltanthera and Sanango appeared successively sister to Gesneriaceae, whereas Cubitanthus, which has been previously assigned to Gesneriaceae, is shown to be related to Linderniaceae. Based on molecular dating and biogeographical reconstruction analyses, we suggest that ancestors of Gesneriaceae originated in South America during the Late Cretaceous. Distribution of Gesneriaceae in the Palaeotropics and Australasia was inferred as resulting from two independent long-distance dispersals during the Eocene and Oligocene, respectively. In a short time span starting at 34Mya, ancestors of Gesnerioideae colonized several Neotropical regions including the tropical Andes, Brazilian Atlantic forest, cerrado, Central America and the West Indies. Subsequent diversification within these areas occurred largely in situ and was particularly extensive in the mountainous systems of the Andes, Central America and the Brazilian Atlantic forest. Only two radiations account for 90% of the diversity of Gesneriaceae in the Brazilian Atlantic forest, whereas half of the species richness in the northern Andes and Central America originated during the last 10 Myr from a single radiation.