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A new species of Hirtella (Chrysobalanaceae), and redescription of a closely related taxon, from the Atlantic Forest, Brazil

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Hirtella prancei, a new species from the Brazilian Atlantic Forest, is described, illustrated and its morphological affinities with other species are discussed. The new species differs from the closely related H. hebeclada and H. santosii mainly by its indumentum on young branches and inflorescences, leaves that are usually larger with a bullate surface and greater number of secondary veins, and by its longer stipules and inflorescences. Hirtella prancei is only known from wet forests in southern Bahia State, Brazil. A more detailed morphological description and new data on distribution and habitat are also provided for H. santosii, which was based on an analysis of additional specimens.
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Phytotaxa 265 (3): 259–266
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Ana Giulietti Harley: 6 Jun. 2016; published: 16 Jun. 2016
http://dx.doi.org/10.11646/phytotaxa.265.3.6
259
A new species of Hirtella (Chrysobalanaceae), and redescription of a closely related
taxon, from the Atlantic Forest, Brazil
RENATA CAMARGO ASPRINO1,3 & ANDRÉ MÁRCIO AMORIM1,2,3
¹Programa de Pós-Graduação em Botânica, Universidade Estadual de Feira de Santana, Avenida Transnordestina s/n, Novo Horizonte,
44036-900, Feira de Santana, Bahia, Brazil. E-mail: renata.asprino@hotmail.com
2Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Ilhéus-Itabuna km 16, 45662-900, Ilhéus, Bahia,
Brazil. E-mail: amorim.uesc@gmail.com
3Centro de Pesquisas do Cacau, Herbário CEPEC, Rodovia Ilhéus-Itabuna km 22, 45650-970, Itabuna, Bahia, Brazil.
Abstract
Hirtella prancei, a new species from the Brazilian Atlantic Forest, is described, illustrated and its morphological affinities
with other species are discussed. The new species differs from the closely related H. hebeclada and H. santosii mainly by its
indumentum on young branches and inflorescences, leaves that are usually larger with a bullate surface and greater number
of secondary veins, and by its longer stipules and inflorescences. Hirtella prancei is only known from wet forests in southern
Bahia State, Brazil. A more detailed morphological description and new data on distribution and habitat are also provided for
H. santosii, which was based on an analysis of additional specimens.
Key words: Malpighiales, Northeast Region, taxonomy
Resumo
Hirtella prancei, uma nova espécie da Floresta Atlântica brasileira, é descrita, ilustrada e suas afinidades morfológicas com
outras espécies são discutidas. A nova espécie diferencia-se das proximamente relacionadas H. hebeclada e H. santosii
principalmente pelo indumento dos ramos jovens e inflorescências, pelas folhas comumente maiores, com a face adaxial
fortemente bulada e maior número de nervuras secundárias, além de estípulas e inflorescência maiores. Hirtella prancei é
conhecida até agora de áreas de floresta ombrófila úmida no sul da Bahia, Brasil. É fornecida também uma descrição mais
detalhada e novas informações sobre a distribuição e habitat de H. santosii, possibilitadas pela análise de material adicional
dessa espécie.
Palavras-chave: Malpighiales, Região Nordeste, taxonomia
Introduction
Hirtella Linnaeus (1753:34) comprises 107 species, of which 105 occur in tropical and subtropical America and two
in eastern Africa and Madagascar (Prance & Sothers 2003). Species of Hirtella are trees, treelets or shrubs with lateral
and subpersistent stipules and eglandular petioles. The pedicels are usually longer than the receptacle tube, the flowers
have 3–9 far-exserted stamens arranged unilaterally, and the ovary is unilocular and inserted on the upper part of the
receptacle tube. The fruit is a fleshy drupe with an endocarp that has 4–7 longitudinal lines, which are often visible in
the epicarp of immature fruits (Prance 1972, Prance & Sothers 2003).
Although the monophyly of Hirtella has been suggested (Yakandawala et al. 2010, Bardon et al. 2013), a recent
study proposed the inclusion of a species of Couepia Aublet (1775:519) in the genus (Sothers et al. 2014). Indeed,
these two genera have been considered closely related since the 19th century (Prance 1972, Prance & White 1988). In
spite of Chrysobalanaceae being a predominantly Amazonian group, the Brazilian Atlantic Forest is considered an
important secondary center of diversity, where there are 15 recorded species of Hirtella (Prance 1979, Sothers et al.
2016). Since the major publications for the group in relation to number of species, the Flora Neotropica Monograph
(Prance 1972) and its supplement (Prance 1989), no new species of Hirtella have been published from Brazil.
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260 Phytotaxa 265 (3) © 2016 Magnolia Press
During fieldwork in the Brazilian Atlantic Forest, a new species of Hirtella was discovered that is closely related
to H. hebeclada Moricand ex A.P. De Candolle (1825:529) and H. santosii Prance (1979:34). Hirtella santosii was
described based on a single collection from southern Bahia (Prance 1979), and while studying the taxonomy of the
genus from Bahia many additional specimens of this species were analyzed. The new species is described and illustrated
here, and comments on its morphological affinities with H. hebeclada and H. santosii, geographic distribution and
conservation status are provided. A more detailed morphological description of H. santosii and additional data about
its habitat and distribution are also included.
Taxonomy
Hirtella prancei Asprino & Amorim, sp. nov. Type:—BRAZIL. Bahia: Arataca, Parque Nacional da Serra das Lontras,
acesso à serra pela roça do Sr. Cristóvão, coletada no começo da trilha para o Mirante II, 15º08’23’’S, 39º18’36’’W,
520 m, 16 December 2014, fl., R.C. Asprino & A.M. Amorim 117 (holotype CEPEC!, isotypes HUEFS!, K!, RB!).
(Figs. 1A–K, 2A–I, 3)
Hirtella prancei Asprino & Amorim is distinguished from H. santosii by its densely velutinous indumentum on the young branches and
inflorescences (vs. densely hirsute to hirsute), stipules 6–10 mm long (vs. 2.5–6.5 mm), leaf blades 10.3–18 mm wide (vs. 4.2–9 cm),
subcoriaceous (vs. subchartaceous to chartaceous), and eglandular (vs. with discoid glands at least on the abaxial surface), pendulous
racemes (vs. erect), 18–40 cm long (vs. 4.5–12.5 cm) and bracteoles triangular to deltate and eglandular (vs. ovate to depressed ovate,
bearing numerous glands on the margins).
Trees or treelets, 3–15 m tall; stems ca. 7 cm diam. at breast height; young branches yellowish brown, densely
velutinous, becoming glabrescent with age, lenticels absent. Leaves alternate, distichous; stipules lateral, narrowly
triangular, 6–10 mm long, late deciduous, eglandular; petiole thick, 6–10 mm long, velutinous when young, eglandular;
leaf blade subcoriaceous, discolorous, widely elliptic to ovate, base obtuse to rounded or subcordate, apex acute to
attenuate or acuminate, 17.5–32.5 × 10.3–18 cm, margins irregularly revolute, with 13–16 pairs of secondary veins,
intersecondary veins sometimes present; abaxial surface with velutinous midvein, secondary to quaternary veins
sparsely velutinous, eglandular, primary to quaternary veins prominent; adaxial surface with primary and secondary
veins glabrescent, eglandular, midvein plane to impressed, secondary to quaternary veins deeply impressed, producing
a strongly bullate surface. Inflorescence a single raceme with 30–70 flowers, pendulous, rachis greenish brown in
vivo, yellowish brown in sicco, densely velutinous, 18–40 × 0.25–0.35 cm; bract 1, narrowly triangular to triangular,
sericeous, (1.5–)2–4.5(–6) mm long, late deciduous, eglandular; pedicel thick, velutinous, (5.5–)7–9(–12) mm long,
eglandular; bracteoles 1–2, triangular to deltate, sericeous, 1–2(–3) mm long, late deciduous, eglandular. Flowers 7–10
mm long; receptacle campanulate, slightly swollen on one side, greenish yellow in vivo, yellowish brown in sicco,
exterior velutinous, interior glabrous except for deflexed hairs near throat; sepals 5, elliptic to oblong, 6–7 × 3–3.5
mm, velutinous abaxially, greenish yellow in vivo, yellowish brown in sicco, margin eglandular; petals 5, oblong to
ovate, 5.5–6.5 × 3–3.5 mm, glabrous, pinkish; stamens 6–8, unilaterally inserted on disc, filaments 19.5–22.5 mm
long, glabrous; anthers 0.8–1.0 mm long; staminodes opposite the stamens, filiform, 0.5–0.8 mm long; ovary inserted
at mouth of receptacle, 1–1.5 × 0.5–1 mm, densely villous, unilocular, biovulate; style gynobasic, filiform, 22–24 mm
long, hirsute up to 1/4 of its length, stigma truncate. Drupe fleshy, obovoid, (2.5–)3.5–3.6 × (1.5–)1.9–2 cm, green
when immature, dark purple when mature, calyx persistent, filaments sometimes persistent, epicarp sparsely hirsute,
mesocarp fleshy, endocarp not seen.
Comments:Hirtella prancei is morphologically similar to H. hebeclada and H. santosii (Fig. 4A–G), which
also occur in the Brazilian Atlantic Forest domain. The three species are trees or treelets with dense indumentum
(drying yellowish brown) covering the branches and rachis of the inflorescence, racemose inflorescences, and flowers
6–10 mm long. Hirtella prancei and H. santosii occur in sympatry in forests of southern Bahia, whereas H. hebeclada
is widespread in forests from Minas Gerais to Rio Grande do Sul States. The occurrence of H. hebeclada in the state
of Bahia may have been based on a few misidentified H. santosii specimens.
Hirtella prancei differs from H. hebeclada by its densely velutinous indumentum on the young branches and
inflorescences (vs. tomentose), stipules 6–10 mm long (vs. 3–7 mm), leaves 17.5–32.5 cm long (vs. 5.5–21 cm) and
10.3–18 cm wide (vs. 2.5–9 cm), with 13–16 pairs of secondary veins (vs. 8–11 pairs) and a strongly bullate adaxial
surface (vs. not bullate), and inflorescences 18–40 cm long (vs. 6–20 cm) with 30–70 flowers (vs. 8–22 flowers).
A NEW SPECIES OF HIRTELLA FROM BRAZIL Phytotaxa 265 (3) © 2016 Magnolia Press 261
FIGURE 1. Hirtella prancei. A—Branch with apical leaf and inflorescence; B—Detail of adaxial leaf surface; C—Basal leaf; D—Detail
of abaxial leaf surface; E—Detail of petiole and stipule; F—Bract; G—Bracteole; H—Flower without petals; I—Longitudinal section
of the flower without petals; J—Mature fruit; K—Isolated trichomes from leaf. (A–I, K based on R.C. Asprino 117 and J based on R.C.
Asprino 61).
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262 Phytotaxa 265 (3) © 2016 Magnolia Press
FIGURE 2. Hirtella prancei. A—Detail of stipules; B—Adaxial leaf surface (left) and inflorescence (right); C—Detail of adaxial leaf
surface; D—Branch with leaves and inflorescences; E—Detail of abaxial leaf surface; F—Detail of leaf margin; G—Detail of inflorescence
with old flowers; H—Mature fruit; I—View of the area where the type specimen was collected. Photos A–G, I by A.M. Amorim and H
by R.C. Asprino.
Besides the features stated in the diagnosis, Hirtella prancei can also be distinguished from H. santosii by its usually
longer leaves (17.5–32.5 cm vs. 8.5–21.5 cm), with 13–16 pairs of secondary veins (vs. 8–14 pairs), strongly bullate
adaxial surface (vs. not bullate or slightly bullate) and revolute margin (vs. plane to slightly revolute), inflorescences
with 30–70 flowers (vs. 12–36 flowers), and filaments 19.5–22.5 mm long (vs. 11–14 mm).
Etymology:—The specific epithet “prancei” is an honor to Sir Ghillean T. Prance, because of his relevant work
on Chrysobalanaceae and other neotropical plant families.
A NEW SPECIES OF HIRTELLA FROM BRAZIL Phytotaxa 265 (3) © 2016 Magnolia Press 263
FIGURE 3. Geographic distribution of Hirtella hebeclada (black triangles), H. prancei (black circles) and H. santosii (white squares).
Distribution and habitat:—Hirtella prancei is known only from a few collections from three localities in
southern Bahia State, Brazil, where it grows in the understory of submontane and montane wet forest (Fig. 3).
Conservation status:—The type locality is Serra das Lontras National Park, a protected area that comprises
ca. 11,300 ha where there are at least two populations of Hirtella prancei. Nevertheless, H. prancei is considered
Endangered (EN) according to the following IUCN (2012) criteria, combined with an analysis using GeoCAT (Bachman
et al. 2011): extent of occurrence less than 5,000 km2, area of occupancy less than 500 km2, with distribution in no
more than five locations, and continuing decline in habitat area and quality (B1ab(iii) + B2ab(iii)).
Additional specimens examined (paratypes):—BRAZIL. Bahia: Arataca, Serra das Lontras, trilha para o
Mirante II, 15º08’25’’S, 39º18’37’’W, 500 m, 05 April 2014, fr., R.C. Asprino et al. 61 (CEPEC!). Ibidem, ca. de 7
ASPRINO & AMORIM
264 Phytotaxa 265 (3) © 2016 Magnolia Press
km no ramal que liga o povoado de Itatingui à Serra das Lontras, 15º12’10’’S, 39º24’29’’W, 900 m, 13 September
2009, ste., A.M. Amorim et al. 7910 (CEPEC!). Ibidem, 10 October 2015, fr., A.M. Amorim et al. 9760 (CEPEC! RB!).
Igrapiúna, assentamento Mirante, 13º54’20’’S, 39º20’59’’W, 18 December 2001, fl., D.M. Loureiro et al. 394 (ALCB!,
CEPEC!). Ilhéus, Fazenda Attalea, Distrito de Castelo Novo, próximo ao Ribeirão do Choro, 100 m, 03 March 1995,
fl., L.A. Mattos-Silva 3074 (CEPEC!).
Hirtella santosii Prance, Revista Brasil. Bot. 2: 34. 1979. Type:—BRAZIL. Bahia: Una, Fazenda São Rafael, 10
December 1968, fl., T.S. dos Santos 300 (holotype CEPEC!, isotype NY). (Figs. 3, 4A–G)
FIGURE 4. Hirtella santosii. A—Detail of stipules; B—Detail of abaxial leaf surface; C—Detail of adaxial leaf surface; D—Buds
and flowers at anthesis; E—Detail of leaf margin; F—Inflorescence; G—Branch with leaves and inflorescences. All photos by A.V.
Popovkin.
A NEW SPECIES OF HIRTELLA FROM BRAZIL Phytotaxa 265 (3) © 2016 Magnolia Press 265
Trees or treelets, 4–20 m tall; stems ca. 6 cm diam. at breast height; young branches brownish green in vivo, yellowish
brown in sicco, hirsute to densely hirsute, soon becoming glabrescent, lenticels sometimes present. Leaves alternate,
distichous; stipules lateral, narrowly triangular to triangular, 2.5–6.5 mm long, late deciduous; petiole thick, 2.5–8.5
mm long, densely hirsute when young, eglandular; leaf blade subchartaceous to chartaceous, discolorous, elliptic to
oblong or ovate, base obtuse to rounded or cuneate, apex attenuate to acuminate, 8.5–21.5 × 4.2–9 cm, margins plane
to slightly revolute, with 8–14 pairs of secondary veins, intersecondary veins sometimes present; abaxial surface with
hirsute to densely hirsute midvein, secondary to quaternary veins sparsely hirsute, small discoid glands present at base
and sometimes sparse throughout lamina, primary and secondary veins prominent, tertiary veins slightly prominent;
adaxial surface with primary and secondary veins glabrescent, sometimes with small discoid glands at base, primary
and secondary veins impressed, tertiary veins plane or slightly impressed, sometimes producing a slightly bullate
surface. Inflorescence a single raceme with 12–36 flowers, erect, rachis greenish in vivo, yellowish brown in sicco,
densely hirsute to hirsute, 4.5–12.5 × 0.1–0.15 cm; bract 1, narrowly triangular to triangular, sericeous, 1.5–3.5 mm
long, persistent, sometimes bearing few glands on margin; pedicel slightly thick, densely hirsute, 6–19 mm long,
eglandular; bracteoles 2, ovate to widely ovate or depressed ovate, sericeous, 0.5–2.5 mm long, persistent, bearing
numerous sessile or shortly stipitate glands around margins. Flowers 6–9.5 mm long; receptacle campanulate, greenish
in vivo, yellowish brown in sicco, exterior densely hirsute, interior glabrous except for deflexed hairs near throat;
sepals 5, oblong to ovate, 3.5–6.5 × 2–3 mm, densely hirsute to hirsute abaxially, greenish in vivo, yellowish brown
in sicco, margin sometimes bearing small glands; petals 5, oblong to rounded or ovate, 2.5–4 × 1.5–2.5 mm, glabrous,
white; stamens 5–7, unilaterally inserted on disc, filaments 11–14 mm long, glabrous, purple with white base; anthers
0.8–1.0 mm long; staminodes opposite stamens; ovary inserted at mouth of receptacle, ca. 1.5 × 1 mm, densely villous,
unilocular, biovulate; style gynobasic, filiform, 14–15 mm long, hirsute up to 1/3 of its length, stigma truncate. Fruit
not seen.
Comments:—The original description of Hirtella santosii was based on a single specimen from the Atlantic Forest
of southern Bahia and, subsequently, many additional collections have been made. The analysis of these specimens
made it possible to write a more detailed description, presented here, and make an accurate morphological comparison
to similar species (see comments under Hirtella prancei).
Distribution and habitat:—Hirtella santosii occurs in the Atlantic Forest from Espírito Santo to northern Bahia,
Brazil (Fig. 3). It grows in the understory of submontane and montane forests at low to mid elevations.
Conservation status:—Hirtella santosii was treated as Endangered (EN) in the Livro Vermelho da Flora do
Brasil (Amorim et al. 2013). Even with the numerous additional collections presented here, which led to the extension
of the distribution of this taxon, the status of H. santosii remains the same. In an analysis that included the additional
specimens, based on IUCN (2012) criteria combined with GeoCAT (Bachman et al. 2011), the extent of occurrence is
more than 20,000 km2 but the area of occupancy is less than 500 km2, the number of locations is no more than ten and
the species is suffering from continuing decline in habitat area and quality (B2ab(iii)).
Additional specimens examined:—BRAZIL. Bahia: Almadina, Serra do Corcovado, 14º42’21’’S, 39º36’12’’W,
06 December 2011, ste., M.M. Coelho et al. 538 (CEPEC!). Mun. Almadina, Serra do Corcovado, 03 February 2012,
fl., M.M. Coelho et al. 624 (CEPEC!). Ibidem, 14º42’44’’S, 39º36’10’’W, 463 m, 23 November 2014, fl., R.C. Asprino
et al. 112 (CEPEC!). Arataca, RPPN Caminho das Pedras, Serra do Peito de Moça, 15º10’27’’S, 39º20’22’’W, 400–
900 m, 25 November 2006, fl., A.M. Amorim et al. 6562 (CEPEC!). Barro Preto, Serra da Pedra Lascada, 14º46’13’’S,
39º32’10’’W, 650 m, 10 December 2015, fl., J.G. Jardim et al. 4791 (CEPEC!). Belmonte, Estação Experimental
Gregório Bondar, 29 November 1987, fl., T.S. dos Santos 4343 (CEPEC!). Camacan, RPPN Serra Bonita, 15º23’30’’S,
39º33’55’’W, 800–1000 m, 17 July 2009, ste., A.M. Amorim et al. 7891 (CEPEC!). Entre Rios, Algodões, 12º10’00’’S,
37º58’00’’W, 140 m, 28 October 2014, fl., A.V. Popovkin & J.C. Mendes 1807 (HUEFS!). Itanagra, Fazenda Brejo
Verde, 17 November 1974, fl., E. Gusmão s/nº (ALCB!, CEPEC!, HRB!, HUEFS!). Espírito Santo: Mun. Domingos
Martins, Rio Jucu Braço Sul, 17 January 1995, fl., G. Hatschbach & J.M. Silva 61601 (CEPEC!, MBM!).
Acknowledgments
We thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for awarding a Master’s
Fellowship to RCA (grant # 133493/2014-8) and a Research Productivity Fellowship to AMA (grant # 306992/2012-
4). We also thank Alex Popovkin for kindly permitting us to use the photos of Hirtella santosii, Lucas Marinho for the
graphical support, Klei Sousa for the drawings and Nathan Smith for the English revision. Laboratory and fieldwork
ASPRINO & AMORIM
266 Phytotaxa 265 (3) © 2016 Magnolia Press
was financially supported by CNPq (Reflora Malpighiales grant # 563548/2010-0, Universal grant # 486079/2013–9
and PPBIO Mata Atlântica grant # 457483/2012-1).
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http://dx.doi.org/10.3417/2007175
... Os estudos taxonômicos embora ainda escassos e pontuais, realizados na Bahia (Asprino & Amorim 2016a), na Paraíba (Brito 2010) e em áreas de Pernambuco (Silva & Almeida Jr. 2009;Gomes-Silva et al. 2018), têm contribuído para o conhecimento da família, principalmente no estado da Bahia, onde três novas espécies foram descritas. São elas: Couepia oxossii Amorim & Asprino (Amorim et al. 2018), Hirtella prancei Asprino & Amorim (Asprino & Amorim 2016b) e Licania farinacea Prance (Prance 2017). ...
... Fruto 1,3-1,8 cm compr., elipsoide; epicarpo seríceo-pubescente. Hirtella santosii é endêmica da Mata Atlântica nos estados da Bahia, Espírito Santo (Asprino & Amorim 2016b) e Alagoas (Flora do Brasil 2020, em construção). Considerada como "Em Perigo" (EN) de extinção , é encontrada na Mata Atlântica em Floresta Submontanas, com flores de outubro a dezembro. ...
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The taxonomic treatment of Hirtella (Chrysobalanaceae) from Bahia State, Brazil, is presented. We recognize 14 species; H. bicornis is recorded for the first time in the state and H. prancei was originally described from this study. The treatment includes an identification key to species, descriptions and comments on taxa, as well as illustrations and geographic distribution maps of species in Bahia. An identification key to the genera of Chrysobalanaceae in the state is also provided.
... In a revision of Chrysobalanaceae in the Brazilian Atlantic Forest, Prance (1979) recognized 47 species, of which five were newly described. Since then, eleven new species of Chrysobalanaceae from this same biome have been published (Prance, 1989(Prance, , 1995(Prance, , 1999(Prance, , 2017Asprino & Amorim, 2016;Amorim et al., 2018). Currently, there are 67 species of Chrysobalanaceae recorded from the Atlantic Forest, of which 47 are endemic (Sothers et al., 2023). ...
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... The most comprehensive taxonomic study of the neotropical species is that of Prance (1972), which provides a key, descriptions, and typifications for 16 species, three of them occurring in the Brazilian Atlantic Forest. The Atlantic Forest has been the focus of other taxonomic studies, which has increased the richness of the region (Prance 1979(Prance , 1995Asprino and Amorim 2016; Gomes-Silva and Alves 2020; Barros et al. 2022). ...
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Recent molecular phylogenetic studies in Chrysobalanaceae as well as new analyses presented in this study cast doubt on the monophyly of the three largest genera in the family, Couepia, Hirtella and Licania. Couepia, a Neotropical genus, had species appearing in four separate clades, the majority of species sequenced, however, form a highly supported clade, referred to here as core Couepia (including the type species). These results lend support to a revised taxonomy of the genus, and to resolve Couepia as monophyletic the following taxonomic changes are here proposed: Couepia recurva should be transferred to Hirtella, C. platycalyx transferred to Licania, C. longipendula and C. dolichopoda transferred to Acioa, and a new genus, Gaulettia, is proposed to accommodate species of the Gaulettia clade and allies.
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Monograph of the genera of Chrysobalanaceae: Acioa, Couepia, Maranthes, Dactyladenia, Hirtella, Kostermanthus, and Magnistipula with descriptions, keys and distribution maps
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Some plant families show a striking imbalance in species diversity between the Neotropics and the Palaeotropics. The woody plant family Chrysobalanaceae is a typical example of this pattern, with 80% of the 531 species in the Neotropics. In order to test alternative interpretations for this pattern, we generated a dated phylogenetic hypothesis for Chrysobalanaceae, using DNA sequence data from one nuclear and six plastid markers. Using a maximum likelihood approach, we jointly inferred ancestral areas and diversification rates in the Neotropics and Palaeotropics. We found that Chrysobalanaceae most probably originated in the Palaeotropics about 80 Mya. The family dispersed into the Neotropics at least four times beginning 40–60 Mya, with at least one back‐dispersal to the Palaeotropics. Members of Chrysobalanaceae have experienced higher extinction, speciation and net diversification rates in the Neotropics. Hence, the high species diversity of Chrysobalanaceae in the Neotropics appears to be primarily caused by a higher speciation rate in this region. Several recent studies have shown high diversification rates in Neotropical plant families, but have focused on Andean‐centred taxa. Ours is the first study to find a similar pattern in a family for which the centre of diversity is in eastern and central Amazonia. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••.
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This account is a generic monograph of a medium-sized (494 species) pantropical plant family. Its rank, circumscription and relationships to other families are examined. An attempt is made to provide an objective and repeatable taxonomy and to formulate evolutionary hypotheses consistent with the facts presented. The monograph is also used to test the general application of certain relatively new and still controversial theoretical approaches to descriptive and evolutionary biology, notably numerical taxonomy, cladistics and vicariance biogeography. The practical value of some potentially useful sources of taxonomic information, especially electron microscopy and phytochemical analysis, is also examined. This is possibly the first time that non-traditional approaches to taxonomy have been so comprehensively evaluated within the framework of a worldwide monograph. Based mainly on original investigations, the morphology, anatomy, reproductive biology, distribution and ecology of each of the 17 genera are described and illustrated. Published works by others on chemistry, embryology and chromosome cytology are summarized. The delimitation and arrangement of the genera included here depart somewhat from those of earlier systems. Most of the new taxa have been formally published elsewhere but the following are proposed here for the first time: tribus Chrysobalaneae, tribus Parinarieae Prance & F. White, tribus Couepieae Prance & F. White, tribus Hirtelleae Prance & F. White, Licania subgenus Angelesia (Korth.) Prance & F. White and Atuna latifrons (Kosterm.) Prance & F. White. All Chrysobalanaceae are woody. They vary greatly in size but are uniform in their vegetative architecture. By contrast, the structure of the inflorescence, flower and fruit is much more diverse; detailed examination has revealed a multitude of previously overlooked characters of classificatory importance. Individually these characters are small, but they are well correlated and belong to character complexes of high diagnostic value which are also functional syndromes related to pollination, dispersal and germination. Each genus is unique in the precise way in which its stigmatic and polliniferous surfaces are presented to pollen vectors and in the structure of its fruit, although within a genus there may be more than one principal pollinator or disperser. Except for a few anomalous species, each genus is distinct in its general appearance. Membership of a genus can be detected by intuitive perception, based on rapid inspection `by eye' of herbarium specimens with flowers. This is possible because of the underlying structural uniformity of the inflorescence and the similar proportions of the flower in all congeneric species. For two genera, Couepia and Hirtella, their objectivity and ease of identification by rapid visual means was demonstrated by simple experiments involving colleagues with little experience of taxonomy and no previous knowledge of the group. The characteristics these `subjects' had subconsciously used were then explicitly formulated in taxonomic terms and their individual diagnostic values calculated. Some methods of numerical taxonomy were found to be useful, although less so than their advocates frequently claim. This is because of problems associated with the detection, selection and weighting of characters. Attempts to apply cladistic analysis to the Chrysobalanaceae were largely unsuccessful because of parallelism and the difficulty of identifying sister groups. The occurrence and extent of parallelism was shown by using a `principle of discordant characters', which is defined and discussed. The ideas underlying it are widely known but the principle itself does not seem to have been previously used as an analytical tool in taxonomic research. In the Chrysobalanaceae, parallelism is so rife that convincing cladograms cannot be constructed; for this reason, among others, the methods of vicariance biogeography are inappropriate. Aspects of evolution other than branching patterns, however, are more relevant to general biology, and monographic studies should help to identify them. Examples from Chrysobalanaceae include: (a) the response in growth-form to geomorphological and climatic change; (b) the biological consequences of dispersal across faunistic barriers; and (c) the evolution of ocean-dispersed, obligate strand plants. It is concluded that monographers cannot afford to ignore fine structure and chemistry, though there is no evidence that they are more fundamental in practical taxonomy than other sources of information. The Chrysobalanaceae and other families show that parallelism is as likely to occur in fine structure and chemistry (at least of secondary compounds) as in gross morphology, and it is much more expensive to detect. At present, for tropical plants, some currently fashionable approaches to taxonomy are of limited relevance. It is much more important to develop objective methods of visual analysis for dealing with the copious material that monographic work entails. It is equally necessary to understand the central role of ecology in biology. Without ecological understanding taxonomy is lifeless and dull, and taxonomically important information and interesting evolutionary situations go undetected.