A new species of Siparuna (Siparunaceae),

S. ficoides, is described, illustrated, and

placed in a phylogenetic context based on morphological and DNA sequence data. The species, which is a monoecious subcanopy tree, is known from three collections made near Manaus, Brazil (two from the same tree), and one in the state of Bolivar, Venezuela.Se describe y se ilustra una nueva especie de Siparuna (Siparunaceae), S. ficoides, ademas se la ubica en un contexto filogenetico basado en datos morfologicos y en sequencias de DNA. Esta nueva especie es un arbol monoico de subdosel, de la cual se han registrado tres colecciones cerca de Manaos, Brasil (dos del mismo arbol) y una coleccion en el Estadod e Bolivar, Venezuela

Hausner, Gerlinde

Renner, Susanne S.

English

Open Access LMU

New Species of siparuna (Siparunaceae) IV. A New Subeanopy Tree from White-Sand Areas in Brazil and Venezuela Susanne S. Renner Fakultat fur Biologie, Ludwig-Maximilians Universitat Munchen, Munich, Germany, renner@lrz.uni-muenchen.de; Missouri Botanical Garden, St. Louis, Missouri 63166, U.S.A. Gerlinde Hausner Werner-Hilpert-Str. 67, D-65197 Wiesbaden, Germany ABSTRACT. A new species of Siparuna (Siparun- aceae), S. ):coides, is described, illustrated, and placed in a phylogenetic ontext based on morpho- logical and DNA sequence data. The species, which is a monoecious subeanopy tree, is known from three collections made near Manaus, Brazil (two from the same tree), and one in the state of Bolivar, Venezuela. RESUMEN. Se describe y se ilustra una nueva es- pecie de Siparuna (Siparunaceae), S. ficoides, ade- mas se la ubica en un contexto filogenetico basado en datos morfologicos y en sequencias de DNA. Esta nueva especie es un arbol monoico de sub- dosel, de la cual se han registrado tres colecciones cerca de Manaos, Brasil (dos del mismo arbol) y una coleccion en el Estado de Bolivar, Venezuela. Key words: Brazil, Central Amazon basin, forest on sandy soils, Siparuna, Siparunaceae, Venezuela. The Ducke Forest Reserve comprises some 10,000 ha of non-flooded forest on the outskirts of Manaus, capital of the state of Amazonas, Brazil. Between 1955 and 1978, about 7000 collections of vascular plants were made there, and another 7354 were added between 1992 and 1998, as part of a British-Brazilian technical cooperation that result- ed in a "Field Guide to the Vascular Plants" (Ri- beiro et al., 1999; M. Hopkins, pers. comm., May 2004). This collection density makes the Ducke forest the best-documented site in the Amazon ba- sin, at least with regard to vascular plants. It is all the more remarkable that new species of trees con- tinue to come to light among collections made in the reserve during the 1990s that are now available for study, mainly in the herbaria INPA, K, MG, NY, RB, and SP. One such species is described here, based on collections from the Ducke reserve and from the Venezuelan state of Bolivar. This distri- butional range makes sense phytogeographically because the Ducke area harbors many species NOVON 15: 202-206. 2005. adapted to sandy soils, such as are typical of the Rio Negro basin and the Venezuelan Gran Savanna. Siparunaceae consist of just two genera, Sipa- runa Aublet (including Bracteanthus Ducke) and its sister group Glossocalyx Bentham, a monotypic West African genus very similar to Siparuna (Ren- ner, 2004; Renner & Hausner, 2005). Most of the 53 species of Siparuna are dioecious, but the phy- logenetically basal species (in a molecular phylog- eny; Renner & Won, 2001) are monoecious and include the large tree species as well as most low- land species found in the genus. Besides monoecy, they share entire-margined leaves, fruitlets that lack stylar arils, and little-developed tepals. ("Sty- lar aril" is a term introduced by Corner [1976] for the fleshy appendages present on some Siparuna- ceae fruitlets; others, for example, Endress [1980] and Barroso [1999], have referred to these append- ages as superarils because they do not derive from the seed or funiculus like a true aril [van der Pijl, 1982: 139, fig. 25]. Instead, the appendages in Si- paruna develop as asymmetric outgrowths at the style base. The stylar arils in Siparuna are orange or red in color and rich in oil, but contain little or no starch. As far as known, monoecious species lack stylar arils, and their drupelets' exo- and me- socarp instead often becomes slimy.) The new spe- cies described here has been sequenced for one chloroplast locus and one nuclear locus (Renner & Won, 2001, under the name S. aff. monogyna), and based on these data it belongs in a clade that also includes S. cristata (Poeppig & Endlicher) A. DC., the species to which it is most similar morpholog- ically (see below). Siparuna ficoides Renner & Hausner, sp. nov. TYPE: Brazil. Amazonas: Res. Flor. Ducke, Manaus-Itacoatiara, km 26, 02°53 ' S, 59°58'W, permanently tagged tree 3325-74, 29 Sep. 1995, C. A. Sothers, E. da C. Pereira & C. F: da Silva 591 (holotype, INPA; iso- types, K, MG, M0, NY, R, SP, U). Figures 1, 2. This content downloaded  on Fri, 22 Feb 2013 05:11:50 AMAll use subject to JSTOR Terms and ConditionsVolume 15, Number 1 2005 Renner & Hausner New Species of Siparuna 203 5 cm A Figure 1. A. Habit of Siparuna ficoides Renner & Hausner (Sothers et al. 591, MO, isotype). B. Distribution of S. JiRcoides. This content downloaded  on Fri, 22 Feb 2013 05:11:50 AMAll use subject to JSTOR Terms and Conditions204 Novon Figure 2. Siparuna ticoides Renner & Hausner. A. Inflorescences with male (left) and female flowers (right, some broken off); (Sothers et al. 591, MO, isotype). B. Lower leaf surface, showing the loose tertiary venation (Briceno 427 NY, paratype). C. Fruiting receptacle (Assunocao et al. 808, NY, paratype). D. Branchlet with young fruiting recep- tacles (Reserva Florestal Ducke, tree 3133-74; vouchers from this tree are Assun,cao et al. 808 and Vicentini et al. 1010). E. Inflorescence with male flowers (Reserva Florestal Ducke, tree 3325-74; a voucher from this tree includes the isotype Sothers et al. 591, MO). Photos D and E: M. J. G. Hopkins. This content downloaded  on Fri, 22 Feb 2013 05:11:50 AMAll use subject to JSTOR Terms and Conditions205 Volume 15, Number 1 2005 Renner & Hausner New Species of Siparuna A Siparuna cristata foliis brevioribus et angustioribus (9-15 x 4-8.8 cm) cum venis tertiaris irregulariter an- astomosantibus laxe dispersis fructibus subglobosus (ca. 2 x 1.1 cm) pilosibusque differt. Monoecious ubeanopy tree, 13-20 m tall, with a DBH of 10-11 cm, the trunk straight, the slash wound light tan or yellowish and strongly lemon- scented; young branchlets terete or flattened at the nodes, glabrous. Leaves opposite; petioles 1-2.2 cm long; lamina drying light green, yellowish or grayish green, shiny, stiff-chartaceous to coriaceous, ob- ovate or elliptic, 9-15 x 4.44.8 cm, the base ob- tuse and shortly attenuate, the apex obtuse, acu- minate, truncate, or emarginate, mature leaves glabrous except for very few peltate or stellate-lep- idote hairs on the lower leaf surface or on the mid- rib; with 9 to 12 pairs of secondary veins, these flat above, slightly raised below, anastomosing near the leaf margin, the tertiary veins irregularly reticulate and loosely spaced (Fig. 2B), the margin entire. Cymes in the axils of extant leaves or on leafless nodes, about 1.5 cm long, with 6 to 12 flowers, the peduncle 44 mm long, the pedicels of male flowers gradually elongating during anthesis and becoming up to 8 mm long, those of the female flowers and fruiting receptacles remaining short (Figs. 1, 2); cymes persistently velvety. Fresh fowers green with a cream-colored center, dried golden-brownish in color; maleflowers cup-shaped, 3-3.5 mm high, I 4.5 mm diam., the indumentum as on the cymes, the tepals reduced to a circular rim surrounding the flower center with its barely developed tomentose floral roof (Fig. 2A, E), stamens 27 to 30, fleshy and with small anther flaps, dorsally with a few stellate hairs; female flowers subglobose, 2.5-3.5 mm high, 4-5 mm diam., the tepals reduced to a circular rim around the flower center (Fig. 2A); the floral roof conical and velvety; carpels ca. 8, the styles basally fused to a short column and hardly protruding from the floral roof, the free parts of the styles thin and easily detached. Fruiting receptacle subglobose, ca. 1.1 cm in height and 2 cm diam. (Fig. 2C), when fresh greenish yellow and densely pubescent (Fig. 2D), dried material persistently vel- vety brown-pubescent, the drupelets protruding (Fig. 2C), drupelets about 6, apparently lacking a stylar aril. Distribution (Fig. 1B), habitat, and phenolo- gy. Siparuna ficoides is known from the Central Amazon basin and the state of Bolivar in Venezue- la, where it grows on sandy soils at altitudes up to about 410 m. Flowers and fruits have been col- lected in August and September. Etymology. The species is named for its leaves resembling certain species of Ficus L. (Moraceae). Common name. The Venezuelan Yekuana (or Yekuna) Indians refer to this species as Medebadi and apply the leaves against snakebites (Briceno 427). Siparuna ficoides resembles S. cristatan which ranges throughout the Amazon basin and into northern Venezuela, adjacent northernmost Colom- bia, and Panama, and which also occurs in the Ducke reserve. The two species have similar gla- brous leathery leaves, but S. cristata has glabrous fruits that are elongate in shape, reaching 3.54 cm in length, with a diameter of (1-)2-3.5 cm, whereas the fruits of S. ficoides are densely velvety pubes- cent and subglobose in shape. In addition, S. cris- tata usually has 10 to 18 stamens, while S. fi;coides has 27 to 30 stamens, the leaves of S. cristata dry dark brown and their blades measure 1>27(-38) x (7-)9-14 cm, while leaves of S. ficoides dry light green or light brown and have blades measuring 9- 15 x 4.4 8.8 cm. Yet another difference lies in the leaf venation: in S. cristata, the tertiary veins cross between adjacent secondaries in parallel paths without branching, while in S. ficoides they are ir- regularly anastomosing and loosely spaced (Fig. 2B) compared to other species of Siparuna. A few duplicates of S. ficoides were distributed as S. mon- ogyna Jangoux, a name that we briefly thought might pertain to the new entity, but a study of the type of S. monogyna revealed that this name does not apply. Another monoecious pecies with velvety fruits is S. gentryana S. S. Renner, occurring in Ecuador and Colombia. However, S. gentryana has tuberculate fruits, rather than smooth, subglobose ones, and dark brown leaves with a venation more like S. cristata. Paratpes. VENEZUELA. Bolivar: Foraneo Aripao, Botadero Norte, 8 May 1994, E. Briceno 427 (NY). BRA- ZIL. Amazonas: Res. Flor. Ducke, Manaus-Itacoatiara, km 26, near observation tower, permanently tagged tree 3133-74, 5 Sep. 1995, A. Vicentini, E A. C. L. Assunfao, C. F: da Silva & E. da C. Pereira 1010 (INPA, K, MG, MO, NY, R, RB, SP, U), same tree collected again, 6 Mar. 1998, P. A. C. L. Assunfao, E. da C. Pereira & C. E da Silva 808 (INPA, K, MG, MO, NY, RB, SP, U, UB). Acknowledgments. We thank Mike Hopkins (MG) and Cynthia Sothers (K) for photos of the new species, and the reviewers David Lorence and Juan Martinez-Laborde for their comments. Literature Cited Barroso, G. M. 1999. Frutos e Sementes: Morfologia Apli- cada a Sistematica de Dicotiledoneas. ViSosa, Editora UFV, Universidade Federal de Vicosa, Brasil. Corner, E. J. H. 1976. The Seeds of Dicotyledons. Cam- bridge Univ. Press, Cambridge. Endress, P. K. 1980. Ontogeny, function and evolution of This content downloaded  on Fri, 22 Feb 2013 05:11:50 AMAll use subject to JSTOR Terms and Conditions206 Novon & H. Won. 2001. Repeated evolution of dioecy from monoecy in Siparunaceae (Laurales). Syst. Biol. 50: 700-712. Ribeiro, J. E. L. da S., M. J. G. Hopkins, A. Vicentini, C. A. Sothers, M. A. da S. Costa, J. M. de Brito, M. A. D. de Souza, L. H. P. Martins, L. G. Lohmann, P. A. C. L. Assunc,ao, E. da C. Pereira, C. F. da Silva, M. R. Mes- quita & L. C. Procopio. 1999. Flora of the Ducke Re- serve: Field Guide to the Vascular Plants of a Terra Firma Forest in Central Amazonia. INPA/DFID, Brazil. extreme floral construction in Monimiaceae. P1. Syst. Evol. 134: 79-120. Pijl, L. van der. 1982. Principles of Dispersal in Higher Plants, 3rd ed. Springer, Berlin. Renner, S. S. 2004. Siparunaceae. Pp. 353-355 in N. P. Smith, S. A. Mori, A. Henderson, D. W. Stevenson & S. V. Heald (editors), Flowering Plants of the Neotrop- ics. Princeton Univ. Press, Princeton. & G. Hausner. 2005. Siparunaceae. Flora Neotro- pica 95. This content downloaded  on Fri, 22 Feb 2013 05:11:50 AMAll use subject to JSTOR Terms and Conditions

New Species of Siparuna (Siparunaceae) IV

Universität München: Elektronischen Publikationen

New Species of siparuna (Siparunaceae) IV. A New Subeanopy 
Tree from White-Sand Areas in Brazil and Venezuela 
Susanne S. Renner 
Fakultat fur Biologie, Ludwig-Maximilians Universitat Munchen, Munich, Germany, 
renner@lrz.uni-muenchen.de; Missouri Botanical Garden, St. Louis, Missouri 63166, U.S.A. 
Gerlinde Hausner 
Werner-Hilpert-Str. 67, D-65197 Wiesbaden, Germany 
ABSTRACT. A new species of Siparuna (Siparun- 
aceae), S. ):coides, is described, illustrated, and 
placed in a phylogenetic ontext based on morpho- 
logical and DNA sequence data. The species, 
which is a monoecious subeanopy tree, is known 
from three collections made near Manaus, Brazil 
(two from the same tree), and one in the state of 
Bolivar, Venezuela. 
RESUMEN. Se describe y se ilustra una nueva es- 
pecie de Siparuna (Siparunaceae), S. ficoides, ade- 
mas se la ubica en un contexto filogenetico basado 
en datos morfologicos y en sequencias de DNA. 
Esta nueva especie es un arbol monoico de sub- 
dosel, de la cual se han registrado tres colecciones 
cerca de Manaos, Brasil (dos del mismo arbol) y 
una coleccion en el Estado de Bolivar, Venezuela. 
Key words: Brazil, Central Amazon basin, forest 
on sandy soils, Siparuna, Siparunaceae, Venezuela. 
The Ducke Forest Reserve comprises some 
10,000 ha of non-flooded forest on the outskirts of 
Manaus, capital of the state of Amazonas, Brazil. 
Between 1955 and 1978, about 7000 collections of 
vascular plants were made there, and another 7354 
were added between 1992 and 1998, as part of a 
British-Brazilian technical cooperation that result- 
ed in a "Field Guide to the Vascular Plants" (Ri- 
beiro et al., 1999; M. Hopkins, pers. comm., May 
2004). This collection density makes the Ducke 
forest the best-documented site in the Amazon ba- 
sin, at least with regard to vascular plants. It is all 
the more remarkable that new species of trees con- 
tinue to come to light among collections made in 
the reserve during the 1990s that are now available 
for study, mainly in the herbaria INPA, K, MG, NY, 
RB, and SP. One such species is described here, 
based on collections from the Ducke reserve and 
from the Venezuelan state of Bolivar. This distri- 
butional range makes sense phytogeographically 
because the Ducke area harbors many species 
NOVON 15: 202-206. 2005. 
adapted to sandy soils, such as are typical of the 
Rio Negro basin and the Venezuelan Gran Savanna. 
Siparunaceae consist of just two genera, Sipa- 
runa Aublet (including Bracteanthus Ducke) and 
its sister group Glossocalyx Bentham, a monotypic 
West African genus very similar to Siparuna (Ren- 
ner, 2004; Renner & Hausner, 2005). Most of the 
53 species of Siparuna are dioecious, but the phy- 
logenetically basal species (in a molecular phylog- 
eny; Renner & Won, 2001) are monoecious and 
include the large tree species as well as most low- 
land species found in the genus. Besides monoecy, 
they share entire-margined leaves, fruitlets that 
lack stylar arils, and little-developed tepals. ("Sty- 
lar aril" is a term introduced by Corner [1976] for 
the fleshy appendages present on some Siparuna- 
ceae fruitlets; others, for example, Endress [1980] 
and Barroso [1999], have referred to these append- 
ages as superarils because they do not derive from 
the seed or funiculus like a true aril [van der Pijl, 
1982: 139, fig. 25]. Instead, the appendages in Si- 
paruna develop as asymmetric outgrowths at the 
style base. The stylar arils in Siparuna are orange 
or red in color and rich in oil, but contain little or 
no starch. As far as known, monoecious species 
lack stylar arils, and their drupelets' exo- and me- 
socarp instead often becomes slimy.) The new spe- 
cies described here has been sequenced for one 
chloroplast locus and one nuclear locus (Renner & 
Won, 2001, under the name S. aff. monogyna), and 
based on these data it belongs in a clade that also 
includes S. cristata (Poeppig & Endlicher) A. DC., 
the species to which it is most similar morpholog- 
ically (see below). 
Siparuna ficoides Renner & Hausner, sp. nov. 
TYPE: Brazil. Amazonas: Res. Flor. Ducke, 
Manaus-Itacoatiara, km 26, 02°53 ' S, 
59°58'W, permanently tagged tree 3325-74, 
29 Sep. 1995, C. A. Sothers, E. da C. Pereira 
& C. F: da Silva 591 (holotype, INPA; iso- 
types, K, MG, M0, NY, R, SP, U). Figures 1, 2. 
This content downloaded  on Fri, 22 Feb 2013 05:11:50 AM
All use subject to JSTOR Terms and Conditions
Volume 15, Number 1 
2005 
Renner & Hausner 
New Species of Siparuna 
203 
5 cm 
A 
Figure 1. A. Habit of Siparuna ficoides Renner & Hausner (Sothers et al. 591, MO, isotype). B. Distribution of 
S. JiRcoides. 
This content downloaded  on Fri, 22 Feb 2013 05:11:50 AM
All use subject to JSTOR Terms and Conditions
204 Novon 
Figure 2. Siparuna ticoides Renner & Hausner. A. Inflorescences with male (left) and female flowers (right, some 
broken off); (Sothers et al. 591, MO, isotype). B. Lower leaf surface, showing the loose tertiary venation (Briceno 427 
NY, paratype). C. Fruiting receptacle (Assunocao et al. 808, NY, paratype). D. Branchlet with young fruiting recep- 
tacles (Reserva Florestal Ducke, tree 3133-74; vouchers from this tree are Assun,cao et al. 808 and Vicentini et al. 
1010). E. Inflorescence with male flowers (Reserva Florestal Ducke, tree 3325-74; a voucher from this tree includes 
the isotype Sothers et al. 591, MO). Photos D and E: M. J. G. Hopkins. 
This content downloaded  on Fri, 22 Feb 2013 05:11:50 AM
All use subject to JSTOR Terms and Conditions
205 Volume 15, Number 1 
2005 
Renner & Hausner 
New Species of Siparuna 
A Siparuna cristata foliis brevioribus et angustioribus 
(9-15 x 4-8.8 cm) cum venis tertiaris irregulariter an- 
astomosantibus laxe dispersis fructibus subglobosus (ca. 
2 x 1.1 cm) pilosibusque differt. 
Monoecious ubeanopy tree, 13-20 m tall, with a 
DBH of 10-11 cm, the trunk straight, the slash 
wound light tan or yellowish and strongly lemon- 
scented; young branchlets terete or flattened at the 
nodes, glabrous. Leaves opposite; petioles 1-2.2 cm 
long; lamina drying light green, yellowish or grayish 
green, shiny, stiff-chartaceous to coriaceous, ob- 
ovate or elliptic, 9-15 x 4.44.8 cm, the base ob- 
tuse and shortly attenuate, the apex obtuse, acu- 
minate, truncate, or emarginate, mature leaves 
glabrous except for very few peltate or stellate-lep- 
idote hairs on the lower leaf surface or on the mid- 
rib; with 9 to 12 pairs of secondary veins, these flat 
above, slightly raised below, anastomosing near the 
leaf margin, the tertiary veins irregularly reticulate 
and loosely spaced (Fig. 2B), the margin entire. 
Cymes in the axils of extant leaves or on leafless 
nodes, about 1.5 cm long, with 6 to 12 flowers, the 
peduncle 44 mm long, the pedicels of male flowers 
gradually elongating during anthesis and becoming 
up to 8 mm long, those of the female flowers and 
fruiting receptacles remaining short (Figs. 1, 2); 
cymes persistently velvety. Fresh fowers green with 
a cream-colored center, dried golden-brownish in 
color; maleflowers cup-shaped, 3-3.5 mm high, I 
4.5 mm diam., the indumentum as on the cymes, 
the tepals reduced to a circular rim surrounding the 
flower center with its barely developed tomentose 
floral roof (Fig. 2A, E), stamens 27 to 30, fleshy 
and with small anther flaps, dorsally with a few 
stellate hairs; female flowers subglobose, 2.5-3.5 
mm high, 4-5 mm diam., the tepals reduced to a 
circular rim around the flower center (Fig. 2A); the 
floral roof conical and velvety; carpels ca. 8, the 
styles basally fused to a short column and hardly 
protruding from the floral roof, the free parts of the 
styles thin and easily detached. Fruiting receptacle 
subglobose, ca. 1.1 cm in height and 2 cm diam. 
(Fig. 2C), when fresh greenish yellow and densely 
pubescent (Fig. 2D), dried material persistently vel- 
vety brown-pubescent, the drupelets protruding 
(Fig. 2C), drupelets about 6, apparently lacking a 
stylar aril. 
Distribution (Fig. 1B), habitat, and phenolo- 
gy. Siparuna ficoides is known from the Central 
Amazon basin and the state of Bolivar in Venezue- 
la, where it grows on sandy soils at altitudes up to 
about 410 m. Flowers and fruits have been col- 
lected in August and September. 
Etymology. The species is named for its leaves 
resembling certain species of Ficus L. (Moraceae). 
Common name. The Venezuelan Yekuana (or 
Yekuna) Indians refer to this species as Medebadi 
and apply the leaves against snakebites (Briceno 
427). 
Siparuna ficoides resembles S. cristatan which 
ranges throughout the Amazon basin and into 
northern Venezuela, adjacent northernmost Colom- 
bia, and Panama, and which also occurs in the 
Ducke reserve. The two species have similar gla- 
brous leathery leaves, but S. cristata has glabrous 
fruits that are elongate in shape, reaching 3.54 cm 
in length, with a diameter of (1-)2-3.5 cm, whereas 
the fruits of S. ficoides are densely velvety pubes- 
cent and subglobose in shape. In addition, S. cris- 
tata usually has 10 to 18 stamens, while S. fi;coides 
has 27 to 30 stamens, the leaves of S. cristata dry 
dark brown and their blades measure 1>27(-38) 
x (7-)9-14 cm, while leaves of S. ficoides dry light 
green or light brown and have blades measuring 9- 
15 x 4.4 8.8 cm. Yet another difference lies in the 
leaf venation: in S. cristata, the tertiary veins cross 
between adjacent secondaries in parallel paths 
without branching, while in S. ficoides they are ir- 
regularly anastomosing and loosely spaced (Fig. 
2B) compared to other species of Siparuna. A few 
duplicates of S. ficoides were distributed as S. mon- 
ogyna Jangoux, a name that we briefly thought 
might pertain to the new entity, but a study of the 
type of S. monogyna revealed that this name does 
not apply. Another monoecious pecies with velvety 
fruits is S. gentryana S. S. Renner, occurring in 
Ecuador and Colombia. However, S. gentryana has 
tuberculate fruits, rather than smooth, subglobose 
ones, and dark brown leaves with a venation more 
like S. cristata. 
Paratpes. VENEZUELA. Bolivar: Foraneo Aripao, 
Botadero Norte, 8 May 1994, E. Briceno 427 (NY). BRA- 
ZIL. Amazonas: Res. Flor. Ducke, Manaus-Itacoatiara, 
km 26, near observation tower, permanently tagged tree 
3133-74, 5 Sep. 1995, A. Vicentini, E A. C. L. Assunfao, 
C. F: da Silva & E. da C. Pereira 1010 (INPA, K, MG, 
MO, NY, R, RB, SP, U), same tree collected again, 6 Mar. 
1998, P. A. C. L. Assunfao, E. da C. Pereira & C. E da 
Silva 808 (INPA, K, MG, MO, NY, RB, SP, U, UB). 
Acknowledgments. We thank Mike Hopkins 
(MG) and Cynthia Sothers (K) for photos of the new 
species, and the reviewers David Lorence and Juan 
Martinez-Laborde for their comments. 
Literature Cited 
Barroso, G. M. 1999. Frutos e Sementes: Morfologia Apli- 
cada a Sistematica de Dicotiledoneas. ViSosa, Editora 
UFV, Universidade Federal de Vicosa, Brasil. 
Corner, E. J. H. 1976. The Seeds of Dicotyledons. Cam- 
bridge Univ. Press, Cambridge. 
Endress, P. K. 1980. Ontogeny, function and evolution of 
This content downloaded  on Fri, 22 Feb 2013 05:11:50 AM
All use subject to JSTOR Terms and Conditions
206 Novon 
& H. Won. 2001. Repeated evolution of dioecy 
from monoecy in Siparunaceae (Laurales). Syst. Biol. 
50: 700-712. 
Ribeiro, J. E. L. da S., M. J. G. Hopkins, A. Vicentini, C. 
A. Sothers, M. A. da S. Costa, J. M. de Brito, M. A. D. 
de Souza, L. H. P. Martins, L. G. Lohmann, P. A. C. L. 
Assunc,ao, E. da C. Pereira, C. F. da Silva, M. R. Mes- 
quita & L. C. Procopio. 1999. Flora of the Ducke Re- 
serve: Field Guide to the Vascular Plants of a Terra 
Firma Forest in Central Amazonia. INPA/DFID, Brazil. 
extreme floral construction in Monimiaceae. P1. Syst. 
Evol. 134: 79-120. 
Pijl, L. van der. 1982. Principles of Dispersal in Higher 
Plants, 3rd ed. Springer, Berlin. 
Renner, S. S. 2004. Siparunaceae. Pp. 353-355 in N. P. 
Smith, S. A. Mori, A. Henderson, D. W. Stevenson & 
S. V. Heald (editors), Flowering Plants of the Neotrop- 
ics. Princeton Univ. Press, Princeton. 
& G. Hausner. 2005. Siparunaceae. Flora Neotro- 
pica 95. 
This content downloaded  on Fri, 22 Feb 2013 05:11:50 AM
All use subject to JSTOR Terms and Conditions


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