Bol. Mus. Biol. Mello Leitão (N. Sér.) 32:5-11. Setembro de 2013
5
Aristolochia bahiensis (Aristolochiaceae) reaches Espírito Santo:
range extension and first description of capsules and seeds
Joelcio Freitas1*, Elton John de Lírio2 & Favio González3
ABSTRACT: Aristolochia bahiensis belongs to Aristolochia ser. Hexandrae
subser. Anthocaulicae, a subseries characterized by the presence of extremely
short and cauliflorous racemes with subtending leaves reduced to small bracts.
Hitherto, this is the only species of this subseries found in the Atlantic Forest.
Here we report a range extension of A. bahiensis, and complement the original
description and illustration, including new morphological traits regarding the
perianth variation during flower development, and description of capsules and
seeds, which had remained unknown until now.
Keywords: Aristolochia ser. Hexandrae, Aristolochia subser. Anthocaulicae,
Brazilian Atlantic forests, perianth variation.
RESUMO: Aristolochia bahiensis (Aristolochiaceae) atinge Espírito Santo:
extensão de alcance e primeira descrição de cápsulas e sementes. Aristolochia bahiensis pertence à Aristolochia ser. Hexandrae subser. Anthocaulicae;
esta subsérie é caracterizada pela presença de racemos caulifloros extremamente
curtos e com folhas reduzidas a pequenas brácteas. Até o momento, esta é a
única espécie desta subsérie encontrada na Mata Atlântica. Neste trabalho são
abordadas a ampliação da distribuição de A. bahiensis, e a complementação da
descrição e ilustração originais, incluindo novidades morfológicas a respeito
da variação do perianto durante o desenvolvimento da flor e a descrição das
cápsulas e sementes, antes desconhecidas.
Palavras-chave: Aristolochia ser. Hexandrae, Aristolochia subser. Anthocaulicae, floresta tropical, variação do perianto, Mata Atlântica.
Museu de Biologia Prof. Mello Leitão. Av. José Ruschi, 04, Centro, Santa Teresa - ES, Brasil. CEP:
29650-000.
2
Programa de Pós-graduação em Botânica, Escola Nacional de Botânica Tropical, Instituto de Pesquisas
Jardim Botânico do Rio de Janeiro. Rua Pacheco Leão, 2040, Solar da Imperatriz, Horto, Rio de Janeiro
– RJ, Brasil. CEP: 22460-030. [email protected].
3
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, AA 7495, Bogotá, Colombia.
[email protected].
* Corresponding author: [email protected]
Recebido: 26 out 2012. Aceito: 15 mar 2013.
1
6
Freitas et al.: Aristolochia bahiensis (Aristolochiaceae) in Espírito Santo
Introduction
The family Aristolochiaceae comprises four genera and about 600 species
(González, 1990, 2012; Capellari Jr., 2001). Aristolochia is the largest genus,
with about 450 species (González, 2011). The genus is highly diversified in
the tropics of the Old and the New World, and it is highly diversified in the
Neotropics, mostly in Hispaniola and Brazil (González, 2000), the latter with
87 species according to Barros & Araújo (2013).
After the monographic work by Hoehne (1927, 1942), devoted mainly to
the Brazilian species, some new taxa have been described from the country
(González, 1998, 2000, 2011). One of these new species, Aristolochia bahiensis F. González was described based on two flowering specimens, both from
the state of Bahia (González, 1998). This species belongs to Aristolochia ser.
Hexandrae subser. Anthocaulicae, which comprises 22 species (González, 1990,
1991, 1997, 1998), all of them characterized by the presence of extremely short
and cauliflorous racemes with leaves reduced to small bracts. Until now, A.
bahiensis is the only species of this subseries found in the Atlantic Forest. The
species of subseries Anthocaulicae are found in western, northern and central
Amazonian forests, in wet forests of the Choco, and Magdalena and Cauca valleys
of Colombia, in stations of Central America and in the Atlantic Forest Brazil.
Here are reported a range extension of A. bahiensis to the state of Espírito
Santo, Brazil, collected by one of us (JF), along with the description and ilustration, for the first time, of the capsules and the seeds of this species, which
had remained unknown until now, and the perianth variation along the flower
development.
Material and Methods
The descriptions, illustrations and the newly reported morphological traits
were made from living specimens of the populations found in the field. Herbarium vouchers were deposited in the herbarium MBML, plus the herbarium
specimens of CEPEC, COL and NY (Mun. Una, Reserva Biológica IBAMA,
J. Jardim et al. 809, CEPEC, NY, the type collection; and Mun. Porto Seguro,
reserva da Brasinheiil Holanda de Industrias S/A, J. G. Jardim et al. 399, COL,
NY). The specimens were examined with the aid of a dissecting microscope
coupled to the light chamber and directly in the field by naked eye. Descriptions
of general morphological characters follow Harris & Harris (2001), except for
those characters unique in Aristolochia flowers, which were described according
to González (1990, 1991, 1994).
Bol. Mus. Biol. Mello Leitão (N. Sér.) 32. 2013
7
Results
Aristolochia bahiensis F. González, Brittonia 50: 8. 1998. Type: Brazil.
Bahía: Município de Una, Reserva Biológica do Mico-leao (IBAMA), entrada
no km 46 da Rod. BA-001 Ilhéus/Una, ramal que 1eva a Faz. Jaqueiral, ca. 8
km da entrada, 15º09’S, 39º05´W, 01 May 1996 (fl), J. Jardim, S. C. Santana
& J. L. Paixão 809 (Holotype: CEPEC; isotype: NY). Figures 1 e 2.
Glabrescent vines. Stem suberose. Twigs cylindrical, internode 6.8-11.1
cm long. Petiole 1.3-5.1 cm long, the proximal half slightly thicker; lamina
ovate-deltoid (narrowly ovate), 10-16 x 5-8.2 cm, base truncate to slightly
cordate (and with a sinus to 4 mm deep), slightly peltate, apex acute to acuminate, chartaceous, glabrous above, sparsely puberulous below, especially along
the veins, basal primary veins 3(5). Pseudostipules absent. Flowers arranged
in short, cauliflorous racemes, 4-13.4 mm long each, with internodes < 1 mm
long, up to 8 flowers, each flower subtended by a deltoid, pubescent bract,
2.0-2.5 x 1.6-2.0 mm. Peduncle plus ovary 1.3-2.3 cm long, slender, shortly
villous, ovary slightly rostrate at its apex. Perianth glabrous, beige with thick
vinaceous grooves on the outside, yellow with whitish indument on the inside,
slightly curved at preanthesis, then the curvature between the utricle and the tube
becomes more pronounced; utricle obovate, 3.7-8 x 3-5.5 mm; syrinx absent;
tube funnel-shape, 1.0-2.3 cm long, 2.0-3.7 mm proximal diameter, 4.5-7.1
mm distal diameter, forming an angle of ca. 110-130o with respect to the utricle;
limb broadly ovate to elliptic, 1.1-2 x 0.6-1.4 cm, forming an angle of ca. 180o
with respect to the tube, vinaceous to yellow and with white trichomes on the
inside, base obtuse, apex obtuse but ending in a very short acumen, margin
revolute at anthesis. Ginostemium stipitate, 1.7-4 mm long; anthers 0.9-1.5
mm long. Capsule cylindrical, 2.9-3.8 x 1.6-1.7 cm, hexagonal in transverse
section, dehiscence acropetal, septa entire; seeds ovate, concave-convex, 4.25.6 x 3.6-4.6 mm, warty, unwinged, prominulous raphe.
Additional specimens examined: Brazil. Bahía: Porto Seguro, reserva
da Brasil Holanda de Industrias S/A, entrada no km 22 da rod. Eunapólis/P.
Seguro, 9.5 km da entrada, mata higrófila sul Bahíana, mata perturbada,
16º27’45’’S, 39º19’31’’ W, 7 abr 1994 (fl), J. G. Jardim et al. 399 (COL, NY).
Espírito Santo: Santa Maria de Jetibá, São José do Rio Claro, prop. Alfredo
Renok, interior da capoeira, próxima à caixa d’água grande, 20º00’08’’S,
40º36’59’’W, 21 abr 2009 (fl), J. Freitas 06 (MBML); 01 jun 2012 (fr), J.
Freitas & L. Tonini 160 (MBML); Santa Teresa, Valsugana Velha, Estação
Biológica de Santa Lúcia, 19º58’4’’S, 40º32’13’’W, 8 fev 2011 (fl) L. Kollmann 12176 et al. (MBML).
8
Freitas et al.: Aristolochia bahiensis (Aristolochiaceae) in Espírito Santo
Figure 1. Aristolochia bahiensis F. González (Freitas et al. 06; Kollmann et al. 12176).
A Twig. B Leaf base with a detail of the indument. C Perianth at anthesis. D Ginostemium during pollen shed. E Dehisced capsule. F Seeds, adaxial (left) and abaxial
(right) views. G Transverse section of capsule before dehiscence.
Bol. Mus. Biol. Mello Leitão (N. Sér.) 32. 2013
9
Discussion
The fruit of Aristolochia bahiensis is a cylindrical capsule, hexagonal
in transverse section (Figure 1G), with dehiscence acropetal and septa entire (Figure 1E). The seeds are ovate, concave-convex, unwinged, warty,
and with a prominulous raphe. This suite of traits is characteristic of most
species of Aristolochia subser. Anthocaulicae (González, 1990). The fact
that the thick, unwinged seeds in capsules located near the ground or in
the understory forest stratum, as those found in A. bahiensis (Figure 1E e
F), could be an adaptation to short-distance dispersal. These traits strongly
contrasts to other new world members of Aristolochia, especially those
from series Thyrsicae (e.g. A. maxima Jacq.) and the group Dipharus (e.g.
A. ringens Vahl), which have capsules that are exposed to the canopy or to
open places, and broad, flat winged seeds, mainly dispersed by wind and/
Figure 2: A-B Young racemes with preanthetic flowers. C Flower at early anthesis,
lateral view. D-E Raceme with preanthetic flowers and a flower at late anthesis.
10
Freitas et al.: Aristolochia bahiensis (Aristolochiaceae) in Espírito Santo
or fresh water streams (González, 1990; González & Stevenson, 2002).
According to the new data reported here, the capsules (Figure 1E) and
seeds (Figure 1F) of A. bahiensis are similar in shape with those of its closely related species, A. guentheri O. C. Schmidt and A. klugii O. C. Schmidt,
which could not be used to separate these three species. Thus, the main
differential traits between these three species are the perianth shape and size
(A. guentheri presented tube 1.0-1.3 cm long, limb ovate to broadly ovate;
A. klugii presented tube 2.0-2.7 cm long, limb bilobed, unequal, lobe adaxial
larger and elliptic to ovate, lobe abaxial smaller and ovate), plus the allopatric
distribution, as the geographic range of A. bahiensis does not overlap with
that of the other two species, which are distributed in the northwestern and
western Amazonian forests (González, 1990, 1994, 1998).
The specimens of A. bahiensis newly collected at Espírito Santo during
the present study show the following variation compared to the type material
of this species: The overall indument is glabrescent (versus glabrous in the
type of A. bahiensis); the shape of perianth is ovate to elliptic (versus oblong
in the type); and the perianth curvature along the tube, which reaches an
angle of ca. 90º during anthesis (versus almost linear in the type). Nevertheless, the variation in all of these traits falls into the expected ranges for
Aristolochia species. The variation of the perianth curvature between the
type collection and the Espírito Santo specimens, is likely due to flower
expansion during anthesis (Figure 2); the young perianth is almost linear
(Figure 2 A), but as the flower approaches anthesis, the tube and limb
gradually curves to ca. 90° with respect to the utricle (Figures. 1C e 2 B)
and remains so until anthesis (Fig. 2 C). At anthesis the tube undergoes
further bending (Figures 2 D-E). These gradual changes on the perianth
curvature have been reported in a number of species (Costa & Hime, 1981;
González & Stevenson, 2000).
Acknowledgements
The authors would like to thank Alfredo Renok for the proper permits to
carry out field observations in situ. Thanks also to the anonymous reviewers for
their thoughtful comments. We acknowledge the staff of the Museu de Biologia
Prof. Mello Leitão, especially, Hélio de Queiroz Boudet Fernandes, curator of
MBML herbarium, for the permission to consult the collection, and Matheus
Poubel for the corrections of the English version.
Bol. Mus. Biol. Mello Leitão (N. Sér.) 32. 2013
11
Literature Cited
Barros, F. & Araújo, A.A.M. 2013. Aristolochiaceae. In: Lista de Espécies da
Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available in: http://
floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB54 (27-02-2013).
Capellari Jr., L. 2001. Aristolochiaceae. In: Rizzo, J.A. (ed.) Flora dos Estados
de Goiás e Tocantins. Coleção Rizzo. Goiânia, Universidade Federal de
Goiás, vol. 27, 34p.
Costa, E. & N. Hime. 1981. Biologia floral de Aristolochia gigantea Mart. &
Zucc. (Aristolochiaceae). Rodriguésia, 33(56): 23-44.
González, F. 1990. Aristolochiaceae. Flora de Colombia. Universidad Nacional
de Colombia, Instituto de Ciencias Naturales, 12: 1-184.
González, F. 1991. Notes on the systematics of Aristolochia subsect. Hexandrae.
Annals of the Missouri Botanical Garden, 78: 497-503.
González, F. 1994. Aristolochiaceae. Flora of Ecuador. In: Harling, G. W. & Andersson, L. (eds.), Flora of Ecuador, p. 1-51. Monograph No. 51. Council
for Nordic Publications in Botany.
González, F. 1997. Hacia una filogenia de Aristolochia y sus congéneres neotropicales. Caldasia, 19(1-2): 115-130.
González, F. 1998. Two new species of Aristolochia (Aristolochiaceae) from
Brazil and Peru. Brittonia, 50: 5-10.
González, F. 2000. A new species of Aristolochia (Aristolochiaceae) from
Bahia, Brazil. Novon, 10: 371-374.
González, F. 2011. A new pseudostipule-bearing species of Aristolochia
(Aristolochiaceae) from Bahía and Espírito Santo, Brazil. Brittonia,
63: 430-435.
González, F. 2012. Florística y sistemática filogenética innecesariamente disyuntas: El caso de Aristolochia, Euglypha y Holostylis (Aristolochiaceae). Revista de la Academia Colombiana de Ciencias Exactas, Físicas
y Naturales, 36 (139): 67-75.
González, F. & Stevenson, D. W. 2000. Perianth development and systematics
of Aristolochia. Flora, 195: 370-391.
González, F. & Stevenson, D. W. 2002. A phylogenetic analysis of the subfamily Aristolochioideae (Aristolochiaceae). Revista de la Academia
Colombiana de Ciencias Exactas, Físicas y Naturales, 26 (98): 25-60.
Harris, J. G. & Harris M. W. 2001. Plant identificacion terminology: An illustrated glossary. 2nd ed. Spring Lake: Spring Lake Publishing. 206p.
Hoehne, F. C. 1927. Monographia illustrada das aristolochiaceas brasileiras.
Memórias do Instituto Oswaldo Cruz, 20(1): 67-175.
Hoehne, F. C. 1942. Aristolochiaceas. Flora Brasílica, 15(2): 3-141.
Download

range extension and first description of capsules and seeds