Phytotaxa 172 (2): 176–200
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Copyright © 2014 Magnolia Press
ISSN 1179-3155 (print edition)
Article
PHYTOTAXA
ISSN 1179-3163 (online edition)
http://dx.doi.org/10.11646/phytotaxa.172.3.2
Taxonomic novelties in Neotropical Chrysobalanaceae: towards a monophyletic
Couepia
CYNTHIA SOTHERS¹, GHILLEAN T. PRANCE², SVEN BUERKI³, ROGIER DE KOK4 & MARK W. CHASE5
¹Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England, UK. Email: c.sothers@kew.org
²Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England, UK. Email: siriain01@yahoo.co.uk
³Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England, UK. Email: s.buerki@kew.org
4
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England, UK. Email: r.dekok@kew.org
5
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England, UK. Email: m.chase@kew.org
Abstract
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.
Key words: Acioa, Hirtella, Licania, phylogenetics, rbcL, matK, ndhF, ITS, Xdh
Introduction
Couepia Aublet (1775: 519) is the third largest genus of the pantropical family Chrysobalanaceae, with all 71 species
endemic to the Neotropics (Prance & Sothers 2003b); its distribution is from Mexico to Paraguay, with the greatest
number of species found in the Guianas and the Brazilian Amazon region. The genus occurs mainly in wet lowland
tropical primary rainforests, although it is also present in a variety of habitats such as flooded forests, cerrado, savanna,
campina and wallaba forest and at higher elevations up to 1000 m, with the exception of C. recurva Spruce ex Prance
(1972a: 272), C. platycalyx Cuatrecasas (1950: 66) and C. steyermarkii Maguire (1952: 252), which may occur at
elevations of 2000 m and higher.
Couepia is characterised by a hollow, usually cylindrical receptacle, free stamens inserted in a complete circle, in
a semi-circle or unilaterally around the mouth of the receptacle, a tendency towards numerous stamens, varying from
ca. (10–)15 to 300 (but most species range from 30–40), a unilocular ovary positioned at the mouth of the receptacle
and fruit with a hard, granular endocarp and a single seed.
Couepia was based on the type C. guianensis Aublet (1775: 519). At the same time Aublet also described four
other Neotropical genera of Chrysobalanaceae: Licania Aublet (1775: 119), Acioa Aublet (1775: 698), Parinari Aublet
(1775: 514) and Moquilea Aublet (1775: 521; currently considered a synonym of Licania). He distinguished Couepia
from these by the numerous free stamens, their arrangement around the receptacle, and the position of the ovary at the
mouth of the receptacle. Acioa was defined by the fused and unilateral stamens, a reduced number of stamens and ovary
at the mouth of the receptacle. Licania (and Moquilea) was described with having a variable number of free stamens,
from few to numerous and the ovary positioned at the base of the receptacle, which clearly distinguishes it from
Couepia. Hirtella Linnaeus (1753: 34) was easily separated from Couepia by its fewer stamens situated unilaterally on
the receptacle.
Although Couepia has been considered a well-defined genus, several relationships remain unclear. Phylogenetic
studies (Bardon et al. 2013, Yakandawala et al. 2010 and Sothers unpublished results) have cast doubts on the monophyly
of some genera in Chrysobalanaceae, including Couepia, Licania and Hirtella, the three largest genera in the family.
The molecular results presented here corroborate previous ideas regarding the peculiar species of Couepia, namely C.
176
Accepted by Hans-Joachim Esser: 6 Apr. 2014; published: 18 Jun. 2014
recurva, C. platycalyx, C. dolichopoda Prance (1974a: 302) and C. longipendula Pilger (1914: 141) (Prance 1972a,
1974a, Cuatrecasas 1950, Pilger 1914), and refutes the view of Couepia as a well-delimited genus. Phylogenetic data
supports revision of the generic limits of Couepia sensu lato.
Couepia recurva, C. dolichopoda and C. platycalyx were all described from single specimens. All three species
have been re-collected several times in recent years, and this has enabled us to better understand the morphology of
these species and to carry out molecular work on them.
The aim of this paper is to recircumscribe Couepia based on a combination of morphological characters and
phylogenetic evidence. We propose the following taxonomic changes: 1) transfer of C. recurva to Hirtella; 2) transfer
of C. platycalyx to Licania; 3) transfer of C. longipendula and C. dolichopoda to Acioa; and 4) propose Gaulettia, a
new genus to accommodate species of the Gaulettia clade, namely C. amaraliae Prance (1989: 72), C. canomensis
(Martius 1827: 80) Benth. ex Hooker (1867: 42), C. elata Ducke (1935: 35) and C. parillo Candolle (1825: 526), as
well as allied species (C. racemosa Benth. ex Hooker (1867: 43), C. foveolata Prance (1972a: 221), C. steyermarkii,
C. canescens (Gleason 1931: 370) Prance (1974b: 119) and C. cognata (Steudel 1843: 761) Fritsch (1889: 60).
Material & Methods
The sampling used to infer phylogenetic relationships includes a total of 54 accessions: 29 of Couepia, 24 species in
other genera of Chrysobalanaceae and one outgroup taxon, Euphronia guianensis (Schomburgk 1847: 757) Hallier
(1918: 13); see Appendix 1. GenBank accession numbers will be available in an upcoming publication on the complete
generic phylogeny of Chrysobalanaceae. The matrix presented here is available from the first author on request.
DNA extraction. Total genomic DNA was extracted from 0.3 g of silica-gel dried leaf tissue from field collections
(except for C. platycalyx, C. recurva, C. parillo and M. panamensis (Standley 1935: 22) Prance & White (1985:
76) which were taken from herbarium specimens) and isolated following a modified 2×cetyl trimethylammonium
bromide (CTAB) method (Doyle & Doyle 1987). Leaf tissue was ground to a paste and incubated in 1 ml of preheated
2×CTAB in 100 mM pH 8.0 Tris-HCl buffer with 1.4 M NaCl, 2% PVP (polyvinylpyrrolidone) and 20 mM EDTA
(Doyle & Doyle 1987). Organic compounds were removed with SEVAG (24:1 chloroform: isoamyl alcohol), after
centrifugation, followed by DNA precipitation in 2.5 volumes of 100% ethanol (for herbarium specimens 2/3 volume
of isopropanol was used instead) at -20ºC for at least 24 hrs. Before precipitation, an aliquot of 150 ml was purified
using the NucleoSpin Extract II PCR purification kit (Machery-Nagel, GmbH & Co. KG, Düren, Germany) following
the manufacturer’s protocols; this provided a small amount of DNA that was used the same day for amplification).
The precipitate was then resuspended in 1.55 gmL-1 caesium chloride/ethidium bromide. Samples were then purified
using a density-dependent gradient, followed by removal of the ethidium bromide with butanol and caesium chloride
by dialysis. All DNA samples are stored in 10 mM Tris-EDTA (TE) (pH 8) -80°C and deposited in the DNA Bank at
the Royal Botanic Gardens, Kew (http:// data.kew.org/dnabank/homepage.html).
PCR amplification and sequence production. For this analysis five regions were selected, three plastid, rbcL,
matK and ndhF, and two nuclear, ribosomal ITS and the low-copy Xdh (primers and protocols in Appendix 2). The
PCR protocol for a 20 µl volume reaction was as follows: 4 µl reaction buffer, 2 µl bovine serum albumin (bsa) (0.4%),
1 µl of each primer, 1.2 µl MgCl2 (25 mM), 0.4 µl dNTPs, 0.4 µl Taq, 11.6 µl H2O and 1–1.5 µl total DNA. Amplified
products were purified using NucleoSpin Extract II PCR purification kit (Machery-Nagel, GmbH & Co. KG, Düren,
Germany) following the manufacturer’s protocol.
The cycle sequencing reactions were performed using the BigDye Terminator Kit ver. 3.1 (Applied Biosystems Inc.,
Warrington, UK). For the plastid regions, the PCR protocol was performed in a 10 µl volume reaction as follows: 0.5 µl
BDT 1.0 µl primer (0.01 µg/µl), 1.75 µl 5 × sequencing buffer, 50 ng of PCR product and up to 10 µl double distilled
H2O. For the nuclear regions ITS and Xdh 0.4 µl of DMSO (2% dimethylsulfoxide) was added to reduce secondary
structure. The thermo cycler reaction included 26 cycles of 10 sec denaturation (96°C), 5 sec annealing (50°C) and 4
min elongation (60°C). The primers used for sequencing were the same as those used for amplifications.
All products were cleaned using the ethanol-EDTA precipitation method (Applied Biosystems Inc.). Cleaned samples
were then sequenced on an ABI 3730 capillary DNA sequencer, using the manufacturer’s protocols. DNA sequences
were edited and contigs assembled using Sequencher ver. 4.1 (Gene Codes Inc., Ann Arbor, MI, USA). The sequences
were aligned by eye, following the guidelines of Kelchner (2000).
Phylogenetic inferences. To investigate the phylogenetics of genus Couepia, single-partition and combined
Bayesian and maximum parsimony (MP) analyses were performed as below. If not mentioned, the same settings were
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Phytotaxa 172 (3) © 2014 Magnolia Press • 177
used for the single-partition and combined inferences. Bayesian analyses were performed using a Markov-Chain MonteCarlo (MCMC) approach, as implemented in MrBayes v3.2.1 (Ronquist et al. 2012), following recommendations by
Nylander et al. (2004). Best-fit models for each region were selected using MrModeltest v.2.0 (Nylander 2004) based
on the Akaike information criterion. The best-fit model for ndhF and Xdh was GTR+G, whereas for the other regions it
was GTR+G+I. For the combined analysis, all the parameters were unlinked (except the topology), allowing partitions
to vary independently and fully contribute to the analysis. Two metropolis-coupled MCMCs with incremental heating
temperature of 0.2 were run for 20 million generations, with the parameters and resulting phylogenetic trees being sampled
every 1000 generations. The analysis was repeated twice, starting with random trees. Convergence was accepted when
standard deviations attained values below 0.01 and when the effective sampling size (ESS) was higher than 200. After
a burn-in period corresponding to 2,500 trees, a half-compatible consensus tree (and its associated Bayesian posterior
probabilities; PP) was reconstructed using MrBayes v3.2.1 (Ronquist et al. 2012). In the case of the MP analyses,
heuristic analyses were performed in PAUP* ver. 4.0b10 (Swofford 2002) under equal weights (Fitch parsimony;
Fitch 1971), using 1000 random taxon-addition replicates and tree bisection-reconnection (TBR) branch swapping
saving multiple trees (MULTREES on), but saving only ten trees per replicate. Shortest trees from all replicates were
then used as starting trees for a final round of swapping with a tree limit of 10,000. Internal support for clades was
assessed using bootstrap analysis (BP; Felsenstein 1985) using 1000 replicates with TBR swapping and a limit of 10
trees retained per replicate.
Results
Phylogenetic inferences. The Bayesian and MP single-partition analyses revealed no moderately to well supported
topological incongruence (BS > 75% and BPP>0.8; data not shown), therefore allowing us to combine the partitions.
In addition, the combined Bayesian and MP phylogenetic trees yielded the same clades; to avoid including many highly
similar trees here, only the combined Bayesian tree is presented and discussed below (Fig. 1).
Phylogenetic analyses conducted in this study strongly supported (Fig. 1) the polyphyly of Couepia, with species in
four clades (Fig. 1): Acioa (BPP: 1.0), Hirtella (posterior probability, PP, of 1.0), Gaulettia (PP 1.0) clades, and core
Couepia (PP 1.0); The latter clade including the type species of the genus, C. guianensis. Monophyly of Maranthes
Blume (1825: 89) and Parinari is highly supported (PP 1.0). Chrysobalanus icaco Linnaeus (1753: 513) is sister to
the Acioa clade (Acioa + C. longipendula; PP 1.0). Licania is polyphyletic, with one clade, L. adolphoduckei Prance
(2001a: 325) + L. heteromorpha Bentham (1840: 221) sister to the Gaulettia clade, a highly supported clade composed
of species of Couepia (PP 1.0) plus L. oblongifolia Standley (1937: 257) sister to the Hirtella clade (Hirtella + C.
recurva; PP 1.0), and Licania egleri Prance (1972a: 56) and L. minutiflora (Sagot 1883: 308) Fritsch (1889: 54) are
sister to core Couepia. Couepia platycalyx is in a polytomy in a clade with the Hirtella clade and species of Licania [L.
niloi Prance (1972a: 129) + L. parviflora Bentham (1840: 221) + L. harlingii Prance (1979: 9) + L. oblongifolia].
Taxonomy & Discussion
The four clades indicated on the phylogenetic tree (Fig. 1), core Couepia, Acioa, Gaulettia, Hirtella, as well as the genus
Licania, are discussed in more detail here based on phylogenetic and morphological evidence.
1. Acioa clade
Phylogenetic evidence strongly supports inclusion of C. longipendula in Acioa. Although C. dolichopoda was not
included in this study, the morphological similarities between C. longipendula and C. dolichopoda are extensive and
in themselves sufficient to warrant also including C. dolichopoda in Acioa.
Acioa longipendula (Pilg.) Sothers & Prance, comb. nov.
Basionym: Couepia longipendula Pilger (1914: 141). Lectotype (designated by Prance 1972a: 248):—BRAZIL. Amazonas: Manaus,
Bosossa, September 1908, Ule 8854 (K!; isolectotypes G! L!, LA, MG!; holotype B destroyed).
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SOTHERS ET AL.
Distribution and habitat:—Brazil (Amazonas and Pará). In terra firme forest from 50–400 m.
Conservation status:—This species is assessed here as least concern, LC (IUCN 2001). It is widely distributed
throughout the Brazilian Amazon region, and there are a substantial number of known individuals; it regenerates well
and is also extensively cultivated.
Selected specimens examined:—BRAZIL. Amazonas: foothills of Serra Curicuriari, 5 November 1971, Prance
et al. 16143 (FHO!, INPA!, NY!, US!); Distrito Agropecuário, Reserva 1501, N of Manaus, 21 October 1990, Freitas
et al. 327 (INPA!, K!, NY!); Santo Antônio de Abonari, Manaus–Caracaraí km 220, 24 November 1976, Prance et al.
24269 (INPA!, NY!, U!, US!). Pará: Rio Trombetas, Planalto Saracá, 23 May 1978, Silva & Santos 4612 (MG!, NY!);
Mun. de Oriximiná, Rio Trombetas near Cachoeira Porteira, 18 June 1980, Cid Ferreira & Ramos 1059 (INPA!, MG!,
NY!, US!).
fIguRe 1. Combined Bayesian half-compatible phylogenetic tree of Couepia and its sister lineages. Bayesian posterior probabilities are
shown above branches. The outgroup taxon, Euphronia guianensis, is not displayed due to its long branch lengths.
Acioa dolichopoda (Prance) Sothers & Prance, comb. nov.
Basionym: Couepia dolichopoda Prance (1974a: 302). Type:—PERU. Loreto: Varadura de Mazán, from Río Amazonas to Río Napo, 21
August 1972, Croat 19382 (holotype NY!, isotypes G!, INPA!, MO!, RB!).
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Phytotaxa 172 (3) © 2014 Magnolia Press • 179
Illustrations:—Prance (1974a: 303, 2001b: 26).
Distribution and habitat:—Western Amazonia in Colombia, Peru and Brazil. In terra firme forests from 100–300 m.
Conservation status:—This species is here assessed as near threatened, NT (IUCN 2001). Although Acioa
dolichopoda occurs over an extensive range and is locally abundant in Colombia (Rozo et al. 2009), it is less widespread
than A. longipendula, and occurs in only a few localities in Colombia, Peru and in western Amazonian Brazil. This
species was assessed as near threatened, NT (VU) for Colombia (Calderón et al. 2002).
Selected specimens examined:—COLOMBIA. Amazonas: Correg. de Puerto Santander, La Chorrera, 7 km SW
of Araracuara, 25 November 1993, Cárdenas et al. 4220 (COAH, K!, NY!). Caquetá: Río Messay, 16 March 1976,
Roa 365 (COL!, INPA!). PERU. Loreto: Maynas Prov., Caserío Mishana, Río Nanay, 18 November 1981, Ruíz &
Murphy 178 (K!, MO!, NY!); Estac. Biol. Callicebus, Río Nanay–Mishana, 31 December 1981, Vásquez et al. 2788
(MO!, NY!); near Brillo Nuevo, Río Yaguayaca, 8 November 1977, Gentry & Revilla 20385 (MO!, NY!). BRAZIL.
Amazonas: Mun. de Amaturá, São Domingos, Rio Solimões, 21 November 1986, Daly et al. 4387 (INPA!, NY!); Rio
Javari, behind Estirão do Equador, 10 August 1973, Lleras et al. P17294 (FHO!, INPA!, NY!); Carauari, Rio Juruá, 30
June 1980, Silva et al. 452 (INPA!, MG!, NY!, RB!).
Notes:—Acioa longipendula and A. dolichopoda are known in the Amazon region for their fruits, the seeds
of which are rich in oils and used locally for cooking. These two species were unique in Couepia s.l. as the only
flagelliflorous, red-flowered and bat-pollinated species in the genus; they were also the only two species in the genus
with fleshy leaves, like Acioa but unlike those of Couepia s.l. Both species have similar floral morphologies but
display distinct characters, such as receptacle shape and stamens.
Acioa is a small South American genus previously comprising four species. Leaf, floral and fruit morphological
characters easily distinguish Acioa from Couepia. The morphological synapomorphies shared by A. longipendula
and A. dolichopoda and other members of Acioa include the fleshy, glabrous leaves with widely spaced secondary
venation, the unequal calyx lobes (actinomorphic in Couepia), a distinct pair of sessile glands on each of the calyx
lobes (absent in Couepia), stamens unilaterally arranged around the receptacle, the number of stamens varing from
10–20(–32) (32 in A. longipendula, in two semi-circles), and unilocular ovary positioned at the mouth of the receptacle
(as in Couepia).
The most notable feature of Acioa, stamens fused into a ligule, which was Aublet’s defining character for the
genus, is now found in only three of the species, A. guianensis Aublet (1775: 698), A. somnolens Maguire (1951: 272)
and A. schultesii Maguire (1951: 272); A. longipendula, A. dolichopoda and A. edulis Prance (1972b: 12) all have free
stamens. Molecular results presented here support placement of A. edulis in Acioa and refute the idea of fused stamens
as the main character in defining this genus.
When Prance (1972b) first described Acioa edulis from fruiting material alone, he oscillated between placing it in
Acioa or Couepia, but described it under Acioa. Once flowering collections of A. edulis were made, he transferred it to
Couepia (Prance 1975), since the flowers displayed free stamens, but later transferred it back to Acioa despite lacking
the main feature of Acioa, the fused stamens (Prance & White 1988). The fruit of A. edulis is also rich in oils and used
regionally for cooking, as are those of A. longipendula and A. dolichopoda.
The newly delimited Acioa is less disjunct in distribution with the inclusion of Acioa longipendula and A.
dolichopoda. Previously each species of Acioa occurred in restricted localities. Acioa schultesii is restricted to the
upper Caquetá River region in Venezuela, A. edulis is confined to the region of Tefé and the Purus River of Brazil,
A. longipendula is widely distributed throughout the Amazon basin and A. dolichopoda occurs in isolated localities
in Colombia, Peru and in the western portion of the Brazilian Amazon region. Acioa guianensis and A. somnolens
are known from only a few specimens; A. guianensis is found in French Guiana, Amapá State (northeastern part of
the Brazilian Amazon) and in the Humaitá River region of Brazil, whereas A. somnolens is known only from French
Guiana.
2. Gaulettia clade
This study strongly supports the close relationship between C. amaraliae, C. canomensis, C. elata and C. parillo,
which belong to the Gaulettia clade. This result together with morphological features unique to this clade support
recognition of a distinct genus. The other five species that have not been sequenced (C. cognata, C. canescens, C.
foveolata, C. racemosa and C. steyermarkii) share several morphological characters that support placing them in this
clade. Further analyses will be required to infer phylogenetic relationships within the genus by expanding the taxon
sampling and sequencing additional and more polymorphic DNA regions.
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SOTHERS ET AL.
Gaulettia Sothers & Prance, gen. nov.
Diagnosis: Gaulettia shares affinities with Couepia and Hirtella but the pronounced reticulate venation of Gaulettia distinguishes it from
both Couepia in the restricted sense used here and Hirtella. The receptacle is often narrow-elongate as in some species of Couepia;
but in Hirtella most species have shallow open receptacles. It shares with Couepia s.s. and Hirtella the ovary at the mouth of the
receptacle. The fruits of Gaulettia are hard and of two types, verrucose or puberulous and differs from Hirtella, which are fleshy with
a smooth, glabrous epicarp. Gaulettia is easily distinguished from Licania by the shape of the receptacle, which is shorter and more
open in Licania, and by the ovary, positioned at the base of the receptacle in Licania.
Type: Gaulettia parillo (DC.) Sothers & Prance
Trees or shrubs; leaf venation reticulate on the abaxial surface and with hair-filled stomatal cavities, leaves often
with a whitish bloom on abaxial surface; prominent parallel secondaries at 90° to primaries; pair of glands at base of
leaf; inflorescence and axis densely ferrugineous (except for G. elata); inflorescence panicles or racemes; calyx lobes
acuminate (rounded to acute in G. racemosa and rounded in some G. cognata); receptacle 3(–4)–10(–22) mm long,
receptacle interior glabrous except for the deflexed hairs at the throat; petals 5, white; stamens 12–65, in a complete
circle, semi-circle or unilaterally placed around the disk; style mostly glabrous, pubescent at base only; ovary villous to
densely villous, unilocular, inserted at mouth of receptacle; fruit hard, epicarp pubescent (G. canomensis, G. parillo) or
verrucose (G. amaraliae, G. elata, G. foveolata, G. racemosa, G. cognata, G. canescens). The fruit of G. steyermarkii
is known only from an immature specimen, which is sparsely pubescent.
Distribution and habitat:—Throughout the Amazonian region of Peru, Ecuador, Colombia, Venezuela, Brazil
and the Guianas.
etymology:—The epithet refers to the word ‘gaulette’, a vernacular name often used in French Guiana for G.
parillo and other species of Chrysobalanaceae.
Notes:—In his revision of Neotropical Chrysobalanaceae, Prance (1972a) assigned four species to the Couepia
parillo superspecies: C. canomensis, C. parillo, C. steyermarkii and C. foveolata. He defined the group by the
conspicuously reticulate venation with silver-grey pubescence in the space between the veins on the underside of
the leaves, prominent parallel tertiary venation connecting secondary veins, a racemose inflorescence with a dense
ferrugineous-brown hirtellous pubescence, a slender cylindrical receptacle tube (usually with villous hairs on the
exterior), and fruits with a tomentose epicarp. He cited two species as also having this distinct leaf reticulation, C. elata
and C. racemosa, but did not place them in this group. Couepia amaraliae was unknown at the time, and C. canescens
was considered a synonym of C. cognata (as Parinari canescens), which he also did not place in the superspecies
group. However, in subsequent publications he assigned C. racemosa, C. amaraliae, C. cognata and C. canescens to
this group (Prance 1974b, 1989).
The nine species of Gaulettia occur mostly in habitats distinct from those of Couepia. Although G. parillo,
G. canomensis and G. racemosa occur in terra firme forest, they are more commonly found in open areas such as
secondary forests and sandy soil habitats. Gaulettia elata is the only true lowland wet rainforest species of Gaulettia,
found in terra firme forests with clay soils, and occasionally in campinarana forests. Gaulettia foveolata occurs in
riverine forests, G. amaraliae, G. steyermarkii, G. cognata and G. canescens all occur in drier, sandy soil habitats
such as savanna, campina, campinarana or high elevation habitats. Two species, G. steyermarkii and G. foveolata, are
endemic to the Guayana Highland, a region known for its high plant endemicity.
Key to species of Gaulettia:
1.
-
2.
3.
Leaves with prominent parallel intercostal (tertiary) venation; inflorescences racemose; flowers usually subsessile or pedicels ca.
1–2 mm long; stamens in almost complete circle, > 40; mature fruit with persistent yellow-brown pubescence; Amazon region in
rainforests and secondary vegetation..................................................................................................................................................2
Leaf without prominent parallel intercostal (tertiary) venation or obscured by dense pubescence; inflorescences panicles or
racemes; flowers with pedicels 1–3.5 mm long; stamens 12–40, unilateral or in 2/3 circle; mature fruit verrucose (unknown in G.
steyermarkii, young fruit pilose); savannas or white sand habitats in northern South America and Amazon region ........................3
Leaf size 5.0–15.5 x 1.7–5.8 cm; receptacle 11–22 mm long; stamens 45–62 ...................................................................G. parillo
Leaf size 8.5–18.0 x 3.0–8.0 cm; receptacle 5.5–10.0 mm long; stamens 40–51 ........................................................ G. canomensis
Leaf reticulate, with indistinct stomatal cavities on abaxial surface; inflorescences racemose; flowers distinctly pedicellate, pedicel
2.5–3.5 mm long, slender; receptacle with sparse silver-grey pubescence; stamens 35–40; canopy tree of (mostly) terra firme
forest ........................................................................................................................................................................................G. elata
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Phytotaxa 172 (3) © 2014 Magnolia Press • 181
-
4.
5.
6.
7.
8.
-
Leaf strongly reticulate, often with distinct stomatal cavities on abaxial surface; inflorescences racemose or paniculate (G.
racemosa, G. amaraliae, G. cognata); flowers with pedicels 1–3 mm long; receptacle with dense pubescence (reddish-brown
villous); stamens 12–40; small tree or shrub of savanna, tepui, campina (except for G. racemosa) .................................................4
Leaf on abaxial surface always covered in lanate pubescence, obscuring tertiaries; inflorescences densely tomentose; stamens
12–25, unilateral .................................................................................................................................................................................5
Leaf on abaxial surface not covered in lanate pubescence, not obscuring tertiaries; inflorescences brown-ferrugineous tomentellous;
stamens 20–40, unilateral or in 2/3 of a circle....................................................................................................................................6
Inflorescences (usually) paniculate; receptacle 5–8 mm long; pedicels ca. 3 mm long; stamens 12–25; shrub or small tree of
savannas..............................................................................................................................................................................G. cognata
Inflorescences racemose; receptacle 3–4 mm long; pedicels to 1 mm long; stamens 12–16; small tree of high elevation forest and
tepuis................................................................................................................................................................................G. canescens
Leaf coriaceous, deeply reticulate, parallel secondaries not prominent .............................................................................................7
Leaf chartaceous, reticulate, parallel secondaries prominent .............................................................................................................8
Receptacle length 3–4 mm; stamens 35–40; campinas and campinaranas, Brazil......................................................... G. amaraliae
Receptacle length 8–10 mm; stamen number unknown; tepui and montane forest, Venezuela..................................G. steyermarkii
Receptacle 5.0–7.5 mm long; stamens 24–30; tree or shrub of secondary, terra firme and riverine forest in central and western
Amazon (not Guianas)..................................................................................................................................................... G. racemosa
Receptacle 8.0–10.0 mm long; stamens ca. 20; tree of riverine forest in western Amazon/upper Rio Negro region......G. foveolata
Gaulettia amaraliae (Prance) Sothers & Prance, comb. nov.
Basionym: Couepia amaraliae Prance (1989: 72), as amaralae. Type:—BRAZIL. Amazonas: base of Serra Aracá, 3 km E of Rio Jauari,
27 February 1984, Prance et al. 29261 (holotype INPA!, isotypes K!, NY!).
Shrub 1–4 m tall. Leaves with an oblong to oblong-elliptic, coriaceous lamina, 2.5–6.0 × 1.5–3.2 cm, prominently
reticulate beneath with conspicuous deep stomatal cavities filled with short grey pubescence. Inflorescences are little
branched panicles or racemes. Receptacle campanulate-turbinate, 3–4 mm long. Stamens 35–40, inserted around ½
of circle. Fruit globose, 5 cm in diameter with a glabrous, lenticellate epicarp; pericarp 2.0–2.5 mm thick, hard and
woody, tomentose within.
Illustration:—Prance (1989: 73).
Distribution and habitat:—Brazil (Amazonas). Sandy soil forests (campina and campinarana). Fig. 2.
fIguRe 2. Distribution of G. amaraliae.
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Conservation status:—This species is here assessed as endangered, EN B1ab(i) + 2ab(i) (IUCN 2001). Gaulettia
amaraliae is restricted to one locality in northern Brazil, in the foothills and surroundings of Serra do Aracá, a sandstone
mountain of the Guayana Highlands.
Selected specimens examined:—BRAZIL. Amazonas: 3 km S of central part of Serra do Aracá, 13 March 1984,
Silva et al. 7121 (INPA!, NY!); Rio Aracá near mouth of Rio Jauari, 2 July 1985, Cordeiro 126 (INPA!, K!, NY!,
SP!); Rio Aracá, 4 November 1952, Fróes & Addison 29277 (IAN!, NY!); Rio Jauari near Ipsilon junction, 9 July
1985, Prance et al. 29420 (INPA!, NY!); Serra Aracá, 1–1.5 hours (8 km) along seringueiro trail, 220° S towards Rio
Jauari from base camp, 4 March 1984, Pipoly & Cress 6788 (INPA!, NY!); near Serra Aracá, 01 March 1977, Rosa &
Cordeiro 1713 (IAN!, NY!).
Gaulettia canescens (Gleason) Sothers & Prance, comb. nov.
Basionym: Parinari canescens Gleason (1931: 370), as Parinarium. Couepia canescens (Gleason) Prance (1974b: 119). Type:—
VENEZUELA. Amazonas: Cerro Duida, Aguita Slope, August 1928–April 1929, Tate 870 (holotype NY!, isotype K!).
Tree to 10 m tall. Leaves oblong to oblong-lanceolate, coriaceous, 2–6.5 × 1.5–3.2 cm, densely brown-lanate pubescent
beneath, the pubescence obscuring prominent venation with parallel secondary veins. Inflorescences racemes 1.5–3
cm long. Receptacle campanulate, 3–4 mm long. Stamens 12–16, unilateral with a few short staminodes opposite
them. Fruit globose with a glabrous and verrucose epicarp.
Illustration:—Prance (1998: 209).
Distribution and habitat:—Endemic to the Guayana Highland of Venezuela and Guyana. In lower montane to
montane slope forests from (800–)1000–1700 m elevation. Fig. 3.
fIguRe 3. Distribution of G. canescens.
Conservation status:—Gaulettia canescens is here assessed as vulnerable, VU D1 (IUCN 2001), since it occurs
in higher-elevation habitats with restricted distribution.
Selected specimens examined:—GUYANA. Demerara-Mahaica: track to St. Cuthbert’s Mission, 22 June
1990, McDowell et al. 1741 (NY!). VENEZUELA. Bolívar: SE facing bluffs of NW Chimantá-tepui, 21 May 1953,
Steyermark 75526 (NY!). Amazonas: Sierra Parima, 18–23 May 1972, Steyermark 105936 (NY!, VEN!).
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Gaulettia canomensis (Mart.) Sothers & Prance, comb. nov.
Basionym:—Moquilea canomensis Martius (1827: 80).—Hirtella canomensis (Mart.) Sprengel (1827: 341).—Couepia canomensis
(Mart.) Benth. Ex Hooker (1867: 42). Type:—BRAZIL. Amazonas: Canuma, without date, Martius s.n. (holotype M!, isotypes BR!,
G!, K!).
Parinari pilosa Standley (1937: 257). Type:—BRAZIL. Amazonas: Mun. de Humaitá, plateau between Rios Livramento and Ipixuna,
7–18 November 1934, Krukoff 7109 (holotype F!, isotypes A!, BM!, BR!, G!, IAN!, K!, LE, MICH, MO!, NY!, RB!, U!).
Shrub or small tree to 10 m tall. Leaves ovate to oblong elliptic, 8.5–18.0 × 3.0–8.0 cm, prominently reticulate beneath
with short grey-brown pubescence between reticulations. Inflorescences densely flowered axillary racemes. Receptacle
subcampanulate-cylindrical, 5.5–10.0 mm long, dense ferrugineous-brown pubescent on exterior. Stamens 40–51, inserted
in a nearly complete circle, with a short row of toothed staminodes opposite them. Fruit round to oblong/pyriform, 3.5–5.0
× 2.5–3.5 cm; epicarp short-velutinous; endocarp fragile, granular, sparsely-pubescent within. Fig. 6J.
Distribution and habitat:—Western and central Amazonia and Guyana, Colombia, Peru, Brazil (Amazonas,
Pará, Rondônia) and Venezuela. Primary and secondary forest and clearings on non-flooded ground. Fig. 4.
fIguRe 4. Distribution of G. canomensis.
Conservation status:—The conservation status of this species is here evaluated as least concern, LC (IUCN
2001). This is a widespread species occurring both in primary and secondary forests throughout the Amazon region.
This species was previously assessed as LC for Colombia (Calderón et al. 2002).
Selected specimens examined:—GUYANA. Rupununi District: Kanuku Mts., Two-Head Mt., 28 January 1994,
Jansen-Jacobs et al. 3434 (K!, NY!, P!, U!). VENEZUELA. Amazonas: Savannas of Santo Antonio, Río Orinoco, date
unknown, Williams 15055 (F!, US!, VEN!). COLOMBIA. Vaupés: Mitú, lower Río Kubiyú, 17 April 1975, Zarucchi
1221 (ECON!, NY!). Caquetá: Araracuara, 28 January 1989, Gentry et al. 65297 (K!, MO!). BRAZIL. Amazonas:
Mun. de São Paulo de Olivença, Estrada Bonfim, 23 November 1986, Daly et al. 4408 (INPA!, K!, NY!); Manaus,
Reserva Florestal Ducke, 03 December 1997, Souza 467 (INPA!, K!, NY!). Pará: Alto Rio Tapajós, São Raimundo, 23
May 1977, Rosa & Santos 1926 (FHO!, MG!, NY!). Rondônia: Mun. de Porto Velho, Samuel Dam area, 13 June 1986,
Cid Ferreira 7427 (INPA!, NY!).
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2d. Gaulettia cognata (Steud.) Sothers & Prance, comb. nov.
Basionym: Hirtella cognata Steudel (1843: 761).—Couepia steudeliana Miquel (1851: 28), nom. illeg. superfl.—Moquilea steudeliana
(Miq.) Walpers (1852: 463), nom. illeg. superfl.—Couepia cognata (Steud.) Fritsch (1889: 60).—Parinari hostmannii Fritsch (1890:
13), as ‘Parinarium hostmanni’, nom. illeg. superfl. Type:—SURINAME. 1843, Hostmann 795 (holotype P!, isotypes BM!, CGE!,
G!, GH!, K!, LE, NY!, OxF!, TCD!, U!, W!).
Shrub or small tree, 1–10 m tall (rarely large trees to 35 m). Leaves oblong to oblong-lanceolate, 2.5–6.0(–15.0) ×
1.0–2.0(–4.5) cm, with grey- to brown-lanate pubescence beneath that usually obscures the prominently reticulate
venation. Inflorescences terminal panicles, the axis and branches brown-villous. Receptacle cylindrical, 5–8 mm long,
reddish-brown and often villous on exterior; petals white or pale pink. Stamens 12–25, unilateral, with a few teeth-like
staminodes opposite them. Fruit globose; epicarp pubescent when young, verrucose and glabrous when mature.
Distribution and habitat:—Gaulettia cognata is distributed in Guyana, Suriname, Brazil (Amazonas) and
Venezuela. It is found in savanna, savanna forest, scrubland, wallaba forest, primary and secondary forest, and tepuis
(1100 m), as well as lower montane forests (200–300 m). Fig. 5.
fIguRe 5. Distribution of G. cognata.
Conservation status:—Gaulettia cognata has a wide distribution in northern Brazil, Venezuela and the Guianas
but often occurs in restricted habitat types, from ca. 50–1000 m. It is therefore here assessed as NT (IUCN 2001).
Selected specimens examined:—GUYANA. Upper Demerara-Berbice District: E of Linden, junction of road to
Mabura Hill, 25 March 1987, Boom 7145 (NY!, P!). Demerara-Mahaica District: Linden Highway, 18 October 1997,
Jansen-Jacobs & ter Welle 5616 (K!, U!). SURINAME. Jodensavanna, without date, BW 5229 (A!, K!, MO!, NY!).
VENEZUELA. Bolívar: Mun. Raúl Leoni, 5 km from base of Cerro Camarón, 42.5 km SE of Entre Rios, 30 October–2
November 1988, Aymard & Fernández 7173 (K!, MO!, PORT). BRAZIL. Amazonas: Rio Içana, near Tunui, 28 March
1952, Fróes 28080 (IAN!, NY!, UB!).
Notes:—Three varieties of Gaulettia [Couepia] cognata have been recognized in the past (Prance 1972a, 1974b,
Prance & Sothers 2003b), but currently their taxonomy is uncertain and should be evaluated in light of more recently
available material. The three varieties previously recognized are: var. cognata, var. major Prance (1972a: 249) and var.
membranacea Prance (1972a: 249).
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2e. Gaulettia elata (Ducke) Sothers & Prance, comb. nov.
Basionym: Couepia elata Ducke (1935: 35). Type:—BRAZIL. Amazonas: Manaus, 09 May 1932, Ducke RB 25001 (holotype RB!,
isotypes K!, P!, U!).
fIguRe 6. Gaulettia elata (Ducke) Sothers & Prance. A. Flowering habit × 1(2/3); B. Underside of leaf × 1.5(1); C. Detail of underside
of leaf venation × 9(6); D. Bud × 4½(3); E. Flower × 6(4); F. Flower L.S. × 6(4); G. Stamen, two views × 40(30); H. Fruit × 1(2/3); I. Detail
of fruit surface × 9(6); J. Fruit of Gaulettia canomensis × 1(2/3). A–G: Freitas et al. 760; H, I: Souza et al. 449; J: Nascimento & Silva 710.
Scale bar: A, H, J = 3 cm; B = 2cm; C, I = 3 mm; D = 7 mm; E, F = 5 mm; G = 0.8 mm (drawn by Lucy T. Smith).
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Tree to 40 m in height. Leaves ovate 4.0–7.0 × 2.5–3.0 cm, prominently reticulate beneath with silver-grey pubescence
between reticulations. Inflorescences few-flowered lax racemes, with grey pubescence. Receptacle obconical-turbinate,
4–5 mm long, sparsely pubescent on exterior. Stamens 35–40, inserted in ½ a circle, with a row of short-toothed
staminodes opposite them. Fruit globose, 5–6 cm in diameter, with a densely verrucose epicarp; endocarp fragile,
granular. Fig. 6A–I.
Distribution and habitat:—French Guiana, Guyana, western and central Amazonian Brazil (Acre, Amazonas)
and Colombia. Found in non-flooded forests. Fig. 7.
Conservation status:—This species is primarily a terra firme forest canopy tree distributed over an extensive part of
the Amazon region. Despite being infrequently collected, it is assessed here as least concern, LC, according to the IUCN
criteria (IUCN 2001). This species was assessed as NT (VU) for Colombia (Calderón et al. 2002).
Selected specimens examined:—FRENCH GUIANA. Sinnamary R., above Petit Saut, 07 September 1993, Mori
et al. 23651 (K!, NY!). GUYANA. Cuyuni-Mazaruni Region: Pakaraima Mts., Karowrieng R., 1 km SE of Maipuri
Falls, 15 October 1992, Hoffman 3036 (K!, NY!, US!). COLOMBIA. Amazonas: Río Cahuina, 07 September 1988,
Sánchez et al. 980 (K!). BRAZIL. Amazonas: Manaus, Reserva Florestal Ducke, 28 May 1997, Assunção 506 (INPA!,
K!, NY!, RB!); Rio Camanaú, Mun. de Novo Airão, 24 July 1989, Miller 173 (INPA!, K!, NY!). Acre: Cruzeiro do Sul,
Estrada Alemanha, 15 April 1971, Prance et al. 11943 (FHO!, INPA!, M!, NY!, RB!).
Notes:—Gaulettia elata is a canopy terra firme forest tree and also occurs infrequently in sandy campinarana
vegetation. It has less distinct reticulate venation than all other species of Gaulettia. The leaves and flowers also lack
the dense pilosity of all other species,which may be due to differences in adaptation to forest versus open habitats.
fIguRe 7. Distribution of G. elata.
2f. Gaulettia foveolata (Prance) Sothers & Prance, comb. nov.
Basionym: Couepia foveolata Prance (1972a: 221). Type:—VENEZUELA. Bolívar: Río Caroní, below Urimán, 11 January 1955,
Steyermark & Wurdack 62 (holotype NY!, isotypes BM!, F!, FHO!).
Tree to 30 m tall, lamina oblong, 3.5–9.5 × 1.5–3.7 cm, prominently reticulate with hair-filled stomatal cavities beneath.
Inflorescences of axillary racemes, few-flowered, densely ferrugineous-brown-pubescent. Bracts and bracteoles
oblong, to 7 mm long, persistent. Receptacle subcylindrical, 8–10 mm long, hirtellous-pubescent on exterior. Stamens
c. 20, unilateral, with tooth-like staminodes opposite. Fruit ovoid, 2.6 × 3.2 cm; epicarp verrucose and hard; endocarp
granular, sparsely hirsute within.
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Illustration:—Prance (1972a: 223).
Distribution and habitat:—Venezuela. Riverine forests of the Guayana Highland, 100–600 m. This species
occurs in the same region as G. steyermarkii but is ecologically distinct. Fig. 8.
Conservation status:—Gaulettia foveolata occurs in a restricted habitat type in the Guayana Highland of
Venezuela, and is thus here assessed as NT (IUCN 2001).
Selected specimens examined:—VENEZUELA. Bolívar: Salto Erepuchi, Alto Río Caroní, January 1949,
Cardona 2555 (NY!, VEN!); Mun. Raul Leoni, S of Urimán, March 1986, Fernández 2288 (K!); Distr. Piar, Río
Aparamán, rapids of Yuray-merú, SW base of Amaruay-tepui, 21 April 1986, Holst & Liesner 2669 (MO!, NY!); Río
Paragua between mouth and Quebrada del Trueno, 13 May–13 June 1987, Stergios 10186 (MO!, NY!, PORT).
Notes:—In Prance & Sothers (2003b) two specimens were erroneously cited as C. foveolata. Rosa & Cordeiro
1713 is now considered to be G. amaraliae, and the correct identification of Stevenson et al. 1011 is G. racemosa.
fIguRe 8. Distribution of G. foveolata.
2g. Gaulettia parillo (DC.) Sothers & Prance, comb. nov.
Basionym: Couepia parillo Candolle (1825: 526).—Moquilea parillo (DC.) Steudel (1841: 159) as ‘parilla’. Type:—FRENCH GUIANA.
Without date, Herb. Ventenat s.n. (holotype G!).
Couepia pauciflora Huber (1909: 372). Type:—BRAZIL. Pará, Lago de Faro, Igarapé do Dedal, 4 September 1907, Ducke MG8630
(holotype MG!, isotypes BM!, RB!).
Couepia villosa Fanshawe & Maguire (in Maguire et al. 1948: 378). Type:—SURINAME. Tafelberg, 17 September 1944, Maguire 24782
(holotype NY!, isotypes BR!, F!, K!, M!, MO!).
Tree to 20 m tall or shrubs. Leaves oblong or oblong-elliptic, 5.0–15.5 × 1.7–5.8 cm, prominently reticulate beneath,
with silver-grey pubescence between the reticulations. Inflorescences racemose, 5–21 flowers, or rarely with solitary
flowers, the axis with ferrugineous-brown pubescence. Receptacle tubular, 11–22 mm long, with dense hirtellous
pubescence and some longer hairs on exterior, glabrous within or with hairs nearly to base on proximal side beneath
ovary only; stamens 45–62, inserted almost in a complete circle, with a portion of toothed staminodes. Fruit globose,
c. 2.5 cm diameter; epicarp yellow-brown-velutinous. Fig. 9.
Illustration:—Prance (2002: 204).
Distribution and habitat:—Guyana, French Guiana, Suriname, Venezuela, Amazonian Brazil (Acre, Amazonas,
Amapá, Roraima), Peru, Ecuador and Colombia. Savannas, primary and secondary forests. Fig. 10.
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fIguRe 9. Fruit of Gaulettia parillo (DC.) Sothers & Prance (Souza et al. 211), from Brazil. © Projeto Flora da Reserva Ducke.
Conservation status:—This species meets the criteria of least concern, LC (IUCN 2001). It is widespread,
occurring throughout the Amazon region, in a number of habitats including secondary forests. Gaulettia parillo (as C.
parillo) was also assessed as least concern, LC for Colombia (Calderón et al. 2002).
Selected specimens examined:—GUYANA. Potaro-Siparuni Region: Iwokrama Reserve, 26 September 1996,
Clarke 2598 (K!, NY!, US!). SURINAME. Sipaliwini: Blanche Marie Falls, Nickerie R., 25 March 1997, Evans et al.
2671 (K!, MO!). VENEZUELA. Amazonas: Dept. Río Negro, Río Mawarinuma, Neblina Massif, 09–14 July 1984,
Davidse & Miller 27217 (MO!, NY!, US!, VEN!). COLOMBIA. Vaupés: near Mitú, 02 October 1991, Betancur et al.
2865 (COL!, K!, NY!, US!). ECUADOR. Napo: Aguarico Cantón, Res. Faunística Cuyabeno, Laguna Zancudo Cocha,
28 September 1991, Palacios 7751 (K!, MO!). PERU. Loreto: Prov. Loreto, Nauta, Río Marañón, 10 November 1982,
Vásquez & Jaramillo 3463 (MO!, NY!). BRAZIL. Acre: Mun. de Cruzeiro do Sul, 44 km E of Cruzeiro do Sul, 18
October 1984, Cid Ferreira et al. 5336 (INPA!, K!, NY!, RB!). Amazonas: Mun. de Barcelos, 3 km S of Central Serra
of Serra Aracá, 04 March 1984, Amaral 1698 (INPA!, NY!). Amapá: Mun. de Oiapoque, 109 km SSE of Oiapoque
road to Calçoene, 05 December 1984, Mori et al. 17186 (K!, MG!, NY!). Roraima: Rio Catrimani, Igarapé Camogi,
20 April 1974, Pires et al. 14059 (IAN!, NY!).
Notes:—Gaulettia parillo and G. canomensis have similar leaves and are sympatric, widely distributed species.
They are more easily distinguished by the size and pilosity of their flowers and inflorescences. The two are also the
only species of Gaulettia with villous mature fruit.
2h. Gaulettia racemosa (Benth. ex Hook.f) Sothers & Prance, comb. nov.
Basionym: Couepia racemosa Benth. ex Hooker (1867: 43). Lectotype (selected by Prance 1972a):—BRAZIL. Amazonas: Falls of São
Gabriel, March 1951, Spruce 1776 (K!, isolectotypes OxF!, P!).
Couepia racemosa var. reticulata Pilger (1906: 150). Lectotype (designated here):—BRAZIL. Amazonas: Manaus, 17 May 1902, Ule
6149 (K!; isolectotypes G!, HBG, MG!; holotype B destroyed).
Couepia calophlebia Standley (1937: 248). Type:—BRAZIL. Amazonas: Rio Embira, 19 June 1933, Krukoff 4913 (holotype F!, isotypes
A!, BM!, K!, M!, MO!, NY!, RB!, U!, US!).
Tree to 30 m tall. Leaves elliptic to oblong, 5.0–18.5 × 2.0–7.5 cm, prominently reticulate beneath, or with stomatal
cavities, with appressed grey pubescence. Inflorescences little branched, densely flowered panicles, the axis and
branches with brown tomentellous pubescence. Receptacle cylindrical-turbinate, 5.0–7.5 mm long, with brown
pubescence on exterior. Stamens 24–30, unilaterally placed, with a smooth, glabrous, untoothed portion of the circle
opposite them. Fruit elliptic, pointed at apex, 3.0–5.0 × 2.0–3.5 cm; epicarp verrucose and pericarp undifferentiated,
fragile, sparsely pubescent within.
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fIguRe 10. Distribution of G. parillo.
Distribution and habitat:— Amazon region of Venezuela, Colombia, Peru and Brazil (Amazonas, Pará). Occurs
in non-flooded forest or on periodically flooded ground, riverine forests, riverbanks and secondary forests up to 100
m. Fig. 11.
Conservation status:—The conservation status of this species is here evaluated as least concern, LC (IUCN
2001). As with G. parillo and G. canomensis, this is a widespread species occurring throughout the Amazon region, in
a number of habitat types, including secondary forests. This species was also assessed as LC for Colombia (Calderón
et al. 2002).
Selected specimens examined:—VENEZUELA. Amazonas: Dept. Atures, Río Cataniapo, 2 km from San Pedro
de Cataniapo, 11 November 1980, Guanchez 396 (MYF). COLOMBIA. Amazonas: Resguardo Aduche, Correg. de Pto.
Santander, 01 December 1993, Cárdenas & Andoque 4282 (COAH, K!). PERU. Loreto: Maynas, Llachapa, Río Napo, 19
January 1983, Vásquez & Jaramillo 3762 (MO!, NY!). BRAZIL. Amazonas: Mun. de Tefé, Lago Tefé, Vila Nogueira, 13
October 1982, Amaral et al. 56 (INPA!, NY!, US!); Manaus, Estrada Mauá, 26 March 1971, Prance et al. 11631 (FHO!,
INPA!, K!, NY!, RB!); Rio Negro at confluence with Rio Uaupés, Serra Camaleão, 22 November 1987, Stevenson et al.
1011 (INPA!, K!, NY!). Pará: 25 km up Rio Mapuera, 07 June 1974, Campbell et al. P22563 (INPA!, NY!, US!).
2i. Gaulettia steyermarkii (Maguire) Sothers & Prance, comb. nov.
Basionym: Couepia steyermarkii Maguire (1952: 252). Type:—VENEZUELA. Bolívar: SE slopes of Ptari-tepui, 10–11 November 1944,
Steyermark 60025 (holotype F!, isotypes K image!, NY!, VEN!).
Tree or shrub with elliptic to oblong-ovate leaves, 4.0–6.0 × 2.7–4.0 cm, prominently reticulate beneath, grey-lanate
between the reticulations. Inflorescences few-flowered axillary racemes. Receptacle cylindrical, 8–10 mm long, with
hirtellous pubescence on the exterior. Stamens inserted around 2/3 of the circle; number of stamens unknown. Young
fruit to 21 × 10 mm, epicarp appressed velutinous (lanate), with long yellow hirtellous hairs, apex pointed.
Distribution and habitat:—Venezuela (highlands of Bolívar state). Forests from 1000–1600 m and on tepuis
(rarely at 500 m); one specimen from secondary forest (1112 m). Fig. 12.
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fIguRe 11. Distribution of G. racemosa.
fIguRe 12. Distribution of G. steyermarkii.
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Conservation status:—Gaulettia steyermarkii has a restricted distribution and is known from only a few
collections. It is here assessed as endangered, EN D, according to the IUCN criteria (IUCN 2001).
Specimens examined:—VENEZUELA. Bolívar: Sifontes Dept., Mun. de Urdaneta, 30 January 1985, Hernández
& Dezzeo 112 (MYF, NY!); Distr. Roscio, 15 km ESE of San Ignacio de Yuruaní, 01 March 1984, Huber et al. 9152
(MYF).
Notes:—When Maguire (1952) described this species, he noted the similarity of the pubescence of the inflorescences
with C. pauciflora, a synomym of G. parillo, and that it was “suggestive” of G. canomensis. He referred to the venation
of G. steyermarkii as “reticulate veinwork”. This species has a deeply reticulate venation and distinct stomatal cavities
on the abaxial surface of the leaves, and the fruits are tomentose on the exterior; however, only immature fruits have
been collected.
3. Hirtella clade
Phylogenetic analyses (Fig. 1) place C. recurva in a clade with other members of American Hirtella. These results,
together with fruit morphological characters, strongly support the inclusion of C. recurva in the genus Hirtella. Hirtella
recurva shares with Couepia the greater number of stamens and the usually narrow, elongate receptacle, although this
character occurs in a few Hirtella species, e.g. H. tubiflora Cuatrecasas (1950: 59).
Hirtella recurva (Spruce ex Prance) Sothers & Prance, comb. nov.
Basionym: Couepia recurva Spruce ex Prance (1972a: 242). Type:—ECUADOR. ‘In Andibus Ecuadoriensis’, October 1857, Spruce 5067
(holotype NY!, isotypes BM!, CGE!, G!, K!, LE, OxF!, W!).
Fruit fleshy, 2.5 × 1.2 mm, with a rugose and glabrescent surface. Mature fruit black. Fig. 13.
Illustrations:—Prance (1972a: 243, 1979: 13); Prance & Sothers (2003b: 38).
Distribution and habitat:—Ecuador and Peru. Hirtella recurva is an Andean primary tropical forest species of
higher elevations, found from 1300 to 1930 m.
Conservation status:—This species is here assessed as VU D1+2 (IUCN 2001); Hirtella recurva is restricted to
high altitudes and is only known from a few localities. It is also currently cited as vulnerable, VU D2 (Romero-Saltos &
Pitman 2004), and although cited as endemic to Ecuador it has now been collected in Peru (Campos & Nuñez 4605).
Selected specimens examined:—ECUADOR. Tungurahua: La Victoria, valley of Río Pastaza, 1 December
1939, Asplund 10055 (NY!, S!). Napo: El Charo Cantón, Río Quijos, at Coca Hydroelectric Project, 3–5 October
1990, Palacios 5956 (K!, MO!, NY!, QCNE). Prov. Zamora-Chinchipe: Zamora, P. N. Podocarpus, Guardería Río
Bombuscaro, January 1995, Palacios & Tirado 13264 (K!, MO!, QCNE), January 1995, Palacios & Tirado 13304
(K!, MO!, NY!, QCNE). Prov. Zamora-Chinchipe: Res. San Francisco, road Loja-Zamora, ca. 35 km from Loja, 2
September 2003, Homeier 1245 (GOET!, K!, LOJA, MO, QCNE) [first record of fruiting collection]; 17 March 2006,
Homeier & Hertel 2190 (GOET!, K!, LOJA, MO, QCNE), 19 September 2008, Homeier 4083 (GOET!, K!, LOJA,
QCNE). Zamora-Chinchipe: Zamora, region de la Cordillera de Cóndor, Parroquia San Carlos de las Minas, Nambija,
San Carlos, 30 January 2005, Quizhpe et al. 867 (K!, LOJA, MO!). PERU. Cajamarca: San Ignacio Prov., Huarango,
Nuevo Mundo, 12 November 1997, Campos & Nuñez 4605 (K!, MO!).
Notes:—Couepia recurva was described from a specimen of flowering material made by Spruce in 1857 (Prance
1972a). Prance noted that the collection was distinct from all other Couepia by the leaves and hirsute filaments.
Moreover, he stated that it resembled many Hirtella in floral characters, but placed the new species in Couepia rather
than Hirtella based solely on the number of stamens (15–20, unlike all other Hirtella with 3–9 stamens), a character
long used to separate Hirtella from Couepia (Prance & White 1988). Fruit morphological characters are also used to
distinguish Couepia from Hirtella (Martius 1827, Zuccarini 1832, Prance & White 1988); fruits of Hirtella have a thin
and fragile endocarp, whereas fruits of Couepia have a hard and woody endocarp. When C. recurva was described
its fruits were unknown but now recent collections from Ecuador (Homeier 1245) confirm that the fruits share more
features with Hirtella than with Couepia. The fruits of Hirtella are generally dispersed by birds, and Couepia are
mammal-dispersed.
192 • Phytotaxa 172 (3) © 2014 Magnolia Press
SOTHERS ET AL.
4. Core Couepia
A highly supported clade of core Couepia, including the type species, C. guianensis, was recovered in the Bayesian
analysis presented here (PP: 1.0; Fig. 1). The Acioa clade is sister to the large clade composed of core Couepia,
Gaulettia, American Hirtella and American Licania. Phylogenetic analyses support the new circumscription of
species in Couepia sensu lato presented here, with high support for core Couepia and the Gaulettia and Hirtella
clades. Morphologically these three genera are distinct from each other as well as from Licania, which further justifies
maintaining them as separate genera.
fIguRe 13. Fruit of Hirtella recurva (Prance) Sothers & Prance (Homeier et al. 1245), from Ecuador. © J.Homeier.
5. The polyphyletic genus Licania
Licania is a large and polyphyletic genus (Bardon et al. 2013, Yakandawala et al. 2010 and Sothers unpublished
results) of ca. 220 species (Prance & Sothers 2003a) and results here highlight the need for further studies in this genus
to re-evaluate the previously held view of it as a good genus, circumscribed mainly by the ovary at the base, or near
the base, of the receptacle and a short, actinomorphic receptacle.
Species of Licania appear in four separate clades within a large clade composed of exclusively Neotropical taxa
(Fig. 1). All clades of Licania receive high node support (PP: 1.0; Fig. 1).
In the analyses presented here L.[C.] platycalyx appears in a polytomy with species of Licania and the Hirtella
clade (Fig. 1). Affinities of L.[C.] platycalyx vary between the Bayesian inference and MP analyses, but in neither does
it appear closely related to core Couepia. Although the results presented here do not fully resolve the relationships of
L.[C.] platycalyx with Licania, they clearly refute the inclusion of this species in core Couepia. Therefore until more
sampling of Licania is available to clarify the placement of this species and understand its relationships to Licania and
other genera of Chrysobalanaceae, we tentatively place it in Licania.
Licania platycalyx (Cuatrec.) Sothers & Prance, comb. nov.
Basionym: Couepia platycalyx Cuatrecasas (1950: 66). Type:—COLOMBIA. Cundinamarca, 04 June 1939, Cuatrecasas & PérezArbeláez 15336 (holotype F!, isotypes COL!, US!).
TAxONOMIC NOVELTIES IN NEOTROPICAL CHRYSOBALANACEAE
Phytotaxa 172 (3) © 2014 Magnolia Press • 193
Illustration:—Prance & Sothers (2003b: 50).
Distribution and habitat:—Costa Rica, Venezuela, Colombia and Ecuador. Occurs in higher altitude Andean
and Central American forests, from 1000 m to 2700 m, but has been recorded from 250 m.
Conservation status:—Licania platycalyx occurs over an extensive geographic range encompassing four
countries, however, it is restricted to higher altitudes and therefore is here assessed as near threatened, NT (IUCN
2001). This species was assessed as EN for Colombia alone (Calderón et al. 2002).
Selected specimens examined:—COSTA RICA. Limón: Cantón de Talamanca, Bratsi, Alto Lari, Kivut, 15 March
1992, Aguilar & Schmidt 1133 (INB!, K!, MO!). Heredia: Cantón de Sarapiqui, Rara Avis, 15 km SW of Horquetas, 2
October 1989, Vargas 175 (CR!, K!, MO!). VENEZUELA. Lara: Distr. Jiménez, camino entre San Carlos y el camino
a Sanare, 15 km S of Sanare, 9 August 1970, Steyermark et al. 103618 (M!, NY!, S!, VEN!). COLOMBIA. Antióquia:
Mun. de El Retiro, 4 km WNW of Cabecera of Quebrada La Agudelo, 12 April 1980, Bernal & Galeano 142 (HUA).
Mun. de Yarumal: 1–5 km on road to San Fermin de Briceño, 4–5 June 1989, Luteyn & Escobar 13275 (NY!).
Valle: Res. Nat. El Refugio Torremdinos, Mun. de Dagua, December 1988, Calderón-Sáenz 143C (K!). ECUADOR.
Esmeraldas: Quinindé, Bilsa Biological Station, Reserva Ecológica Mache-Chindul, 11 March 1998, Clark et al. 4639
(K!, MO!, QCNE).
Notes:—Licania [as Couepia] platycalyx was described by Cuatrecasas (1950), who noted that the species was
“an extraordinary member of this genus and of the Chrysobalanoideae”, and placed it in Couepia “on account of the
strongly lateral insertion of the ovary and the numerous stamens”. Prance (1972a) saw only the type and one other
specimen so was reluctant to make a change, although uncertain about its affinity to Couepia, since the fruits and the
position of the ovary differ from all other Couepia.
Licania platycalyx shares a similar receptacle and leaf morphology with Licania, but differs in the position the
ovary, which is laterally inserted on the wall of the receptacle rather than at the base, as in most other Licania. Licania
licaniiflora (Sagot 1883: 308) Blake (1917: 66), also has a laterally inserted ovary, but differs significantly from L.
platycalyx in other characters.
Acknowledgements
We thank Steve Bachman for producing the maps of Gaulettia and reviewing the conservation assessments, Lucy
Smith for the beautiful illustration, Jürgen Homeier for providing plant material and photographs of Hirtella recurva,
Kyle Dexter and Juliette Chamagne for providing DNA samples of G. parillo, Mike Hopkins for logistic support
during field expeditions; and Edith Kapinos, Jim Clarkson, Dion Devey and Lola Lledó for their support in the Jodrell
Laboratory. The curators of the cited herbaria are gratefully acknowledged, in particular of NY and K for providing
loan material and access to specimens.
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APPeNDIX 1. List of taxa used in the molecular analyses with an indication of markers included for each taxon. All from
silica-gel dried leaf material, except those marked with an *, which were extracted from herbarium specimens at the Royal
Botanic Gardens, Kew (K).
Species
Collector/ number
DNA Nº
rbcL
matK
ndhF
ITS
Xdh
30797
√
√
√
√
√
20138
√
√
√
√
√
MWC
Acioa edulis
Sothers 1512; Brazil, Amazonas, BR-174, Reserva de
Fruticultura do INPA
Acioa guianensis
Mansano 04/279; Brazil, R.J., Rio de Janeiro, Jardim
Botânico. Cultivated
*Couepia amaraliae
Stevenson et al. 1011; Brazil, Amazonas, rio Negro
42732
√
√
√
√
√
Couepia belemii
Sothers 1518; Brazil, Bahia, Mun. de Itacaré
30800
√
√
√
√
√
Ribeiro 1550; Brazil, Amazonas, Manaus, Reserva Florestal
28042
√
√
√
√
√
28043
√
√
√
√
√
Couepia bracteosa
Ducke
Couepia canomensis1
Hopkins 1438; Brazil, Amazonas, Manaus, Reserva Florestal
Ducke
Couepia canomensis2
Sothers 1456; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30890
√
√
√
√
√
Couepia caryophylloides
Sothers 1455; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30851
√
√
√
√
√
Couepia coarctata
Sothers 1519; Brazil, Bahia, Mun. de Itacaré
30850
√
√
√
√
√
Couepia elata1
Assunção 502; Brazil, Amazonas, Manaus, Reserva Florestal
28044
√
√
√
√
√
35090
√
√
√
√
√
Ducke
Couepia elata2
Sothers 1450; Brazil, Amazonas, Manaus, Reserva Florestal
Ducke
Couepia excelsa
Sothers 1453; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30852
√
√
√
√
√
Couepia grandiflora
Sothers 1507; Brazil, Brasília, Distrito Federal, Jardim
28052
√
√
√
√
√
28046
√
√
√
√
√
28045
√
√
√
√
√
Botânico
Couepia guianensis
Vicentini 578; Brazil, Amazonas, Manaus, Reserva Florestal
glandulosa
Ducke
Couepia guianensis
Sothers 1451; Brazil, Amazonas, Manaus, Reserva Florestal
guianensis
Ducke
Couepia habrantha
Sothers 1464; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30804
√
√
√
√
√
Couepia impressa
Sothers 1529; Brazil, Pernambuco, Restinga de Ariquindá
34570
√
√
√
√
√
Couepia longipendula1
Sothers 263; Brazil, Amazonas, Manaus, Reserva Florestal
28047
√
√
√
√
√
Ducke
Couepia longipendula2
Sothers 1490; Brazil, Amazonas, Manaus, grounds of INPA
35106
√
√
√
√
√
Couepia magnoliifolia
Sothers 1538; Brazil, Amazonas, Manaus, Reserva Florestal
35097
√
√
√
√
√
Ducke
Couepia morii
Sothers 1470; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30853
√
√
√
√
√
Couepia ovalifolia
Sothers 1509; Brazil, Rio de Janeiro, RJ, Barra da Tijuca,
28054
√
√
√
√
√
M171-
√
√
√
√
x
Lote 2
Couepia parillo
Without collector and number; French Guiana, Montagne
Tortue
16625
Couepia parvifolia
Oliveira 1713; Brazil, R.J., Rio de Janeiro, Estrada da Vista
28053
√
√
√
√
√
Aguilar 3643; Costa Rica, Limón, Cantón de Pococi
42734
partial
partial
√
√
x
Homeier et al. 2190; Ecuador, Zamora-Chinchipe, Reserva
35410
partial
√
√
√
√
Chinesa, Km 1
* Couepia platycalyx
* Couepia recurva
Biológica San Francisco
Couepia rufa
Sothers 1668; Brazil, Pernambuco, Recife, Campus UFRPE
34569
√
√
√
√
√
Couepia sandwithii
Sothers 1465; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30855
√
√
√
√
√
Couepia schottii
Sothers 1501; Brazil, Rio de Janeiro, Restinga da Marambaia
21383
√
√
√
√
√
......continued on the next page
198 • Phytotaxa 172 (3) © 2014 Magnolia Press
SOTHERS ET AL.
APPeNDIX 1. (Continued)
Species
Collector/ number
DNA Nº
rbcL
matK
ndhF
ITS
Xdh
MWC
Couepia spicata
Sothers 1471; Brazil, Amazonas, ZF-2, Reservas do PDBFF
30856
√
√
√
√
√
Couepia ulei
Sothers 992; Brazil, Amazonas, Manaus, Reserva Florestal
28051
√
√
√
√
√
Chrysobalanus icaco
Sothers 1524; Brazil, Bahia, Mun. de Olivença
30798
√
√
√
√
√
Hirtella myrmecophila
Sothers 1503; Brazil, Amazonas, Manaus, Reserva Florestal
28912
√
√
√
√
√
35098
√
√
√
√
√
35093
√
√
√
√
√
35094
√
√
√
√
√
35104
√
√
√
√
√
40407
√
√
√
√
√
35102
√
√
√
√
√
37996
√
√
√
√
x
Ducke
Ducke
Hirtella physophora
Sothers 1539; Brazil, Amazonas, Manaus, Reserva Florestal
Ducke
Hirtella rodriguesii
Sothers 1475; Brazil, Amazonas, Manaus, Reserva Florestal
Ducke
Hirtella sprucei
Sothers 1530; Brazil, Espírito Santo, Linhares, Res. Natural
da CVRD
Hirtella mutisii
Homeier et al. 3476; Ecuador, Napo, Parque Nacional
Sumaco-Galeras
Licania adolphoduckei
Tree number 435-02; Brazil, Amazonas, Manaus, Reserva
Florestal Ducke
Licania celiae
Homeier et al. 3567; Ecuador, Napo, Parque Nacional
Sumaco-Galeras
Licania egleri
Sothers 1545; Brazil, Amazonas, rio Urucu, Área de
Exploração da Petrobrás
Licania harlingii
Homeier et al. 3755; Ecuador, Napo, Cordillera Galeras
35105
√
√
√
√
√
Licania heteromorpha
Sothers 1474; Brazil, Amazonas, Manaus, Reserva Florestal
35092
√
√
√
√
√
Licania macrocarpa
Homeier 3481; Ecuador, Napo, Cocodrilos
38000
√
√
√
√
√
Licania minutiflora
Sothers 1555; Brazil, Amazonas, rio Urucu, Área de
37999
√
√
√
√
√
42725
√
√
√
√
√
37995
√
√
√
√
√
37997
√
√
√
√
√
28905
√
√
√
√
√
35089
√
√
√
√
√
Ducke
Exploração da Petrobrás
Licania niloi
Tree number 219-02; Brazil, Amazonas, Manaus, Reserva
Florestal Ducke
Licania oblongifolia
Sothers 1544; Brazil, Amazonas, rio Urucu, Área de
Exploração da Petrobrás
Licania parviflora
Sothers 1548; Brazil, Amazonas, rio Urucu, Área de
Exploração da Petrobrás
Maranthes chrysophylla
van der Burgt 773; Cameroon, Korup National Park, NW
plot near P transect
Maranthes corymbosa
Prance 30902; Malaysia, Selangor, grounds of Forest
Research Institute
*Maranthes panamensis
Thomsen 1000; Costa Rica, Aguabuena, Osa Peninsula
42735
partial
partial
√
√
x
Parinari occidentalis
Homeier et al. 3709; Ecuador, Napo, Parque Nacional
35103
√
√
√
√
√
Parinari montana
Sothers 1661; Brazil, Amazonas, rio Urucu, Área de
35101
√
√
√
√
√
Parinari parva
Prance 30899; Malaysia, Johore, Endau-Rompin N.P.
35411
√
√
√
√
√
Euphronia guianensis
Mori 23699; Venezuela, Bolívar, Gran Sabana, Salto Kama
1057
√
√
√
√
√
Sumaco-Galeras
Exploração da Petrobrás
TAxONOMIC NOVELTIES IN NEOTROPICAL CHRYSOBALANACEAE
Phytotaxa 172 (3) © 2014 Magnolia Press • 199
Appendix 2: Primers used and thermocycler conditions.
DNA
Primers
Reference
390F-trn2KR
Johnson & Soltis 1995
Profile
markers
matK
1 min denaturation (94°C) and 30 sec at 94°C, 35
cycles of 40 sec annealing (50°C), 1 min (72°C),
5 min (72°C)
matK
ChrysF*-ChrysR*
ChrysF: TTT GTT GAA AAA TGT AGG TTA TGG
ChrysR: TTC AAT CAC AGG AAT AAT TGG AA
1 min denaturation (94°C) and 30 sec at 94°C, 35
cycles of 40 sec annealing (53°C), 1 min (72°C),
5 min (72°C)
ndhF
5.5mF-10.5R
Davis et al. 2001
2 min denaturation (94°C) and 1 min at 94°C, 30
cycles of 1 min annealing (50°C), 1.5 min (72°C),
4 min (72°C)
rbcL
1F-724R
Chase et al. 1993; Lledó et al. 1998
3 min denaturation at 94oC and 1 min at 94oC, 28-
674F-1360R
Chase et al. 1993
30 cycles annealing at 48oC for 1 min, elongation
at 72oC for 1 min, 72oC for 7 min, 4oC hold
ITS
17SE-26SE
Sun et al. 1994
94oC for 2 min, denaturation at 94oC for 1
min, annealing at 50oC for 1 min for 28 cycles,
elongation at 72oC for 1 min, 72oC for 7 min, 4oC
hold
Xdh
471F-1365R
Morton 2011
Touchdown reaction: 94oC for 3 min, 0:45 sec
denaturation at 94oC, annealing at 55oC–1oC for
0:45 sec for 6 cycles (temperature lowered by
1o for each cycle and cycle repeated to 49oC),
elongation at 72oC for 1:30 min, then 27 cycles of
94oC for 0:45 sec, 49oC for 0:45 sec, 72oC for 1:30
min, hold at 72oC for 7 min
* matK ChrysF and ChrysR were designed for this study.
200 • Phytotaxa 172 (3) © 2014 Magnolia Press
SOTHERS ET AL.
Rogier R P de Kok