American Society of Plant Taxonomists Is Collaborating With JSTOR To Digitize, Preserve and Extend Access To Systematic Botany Monographs

Revision of Neosprucea (Salicaceae)
Author(s): Mac H. Alford

Source: Systematic Botany Monographs, Vol. 85, Revision of Neosprucea (Salicaceae) (Jun. 2, 2008
), pp. 1-62
Published by: American Society of Plant Taxonomists
Stable URL: http://www.jstor.org/stable/25027975
Accessed: 03-10-2015 13:02 UTC
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/
info/about/policies/terms.jsp
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content
in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship.
For more information about JSTOR, please contact support@jstor.org.
American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic
Botany Monographs.
http://www.jstor.org
This content downloaded from 130.102.42.98 on Sat, 03 Oct 2015 13:02:34 UTC
All use subject to JSTOR Terms and Conditions
REVISION OF NEOSPRUCEA (SALICACEAE)
Mac H. Alford
Department of Biological Sciences
University of Southern Mississippi
118 College Drive #5018
Hattiesburg, Mississippi 39406
Abstract. The neotropical genus Neosprucea (Salicaceae), formerly included in Flacourtiaceae, com
nine of trees. Its range extends from Panama to Guyana and to southern Peru. Data on morphol
prises species
ogy, wood anatomy, pollen, and geographical distribution are presented. Morphological and molecular analyses
were used to assess relationships among the species and of the genus within the family. Neosprucea includes five
previously recognized species and four here newly described (N. melastomatoides, N. paterna, N tenuisepala,
and N. wilburiana). The name N. sucumbiensis is considered a synonym of N. grandiflora. The earlier transfer
of N. kuhlmannii to the genus Macrothumia is maintained. For each species a full description and range map is
provided; all species, except N. sararensis, are illustrated.
Resumen. El g?nero neotropical Neosprucea (Salicaceae), incluido anteriormente en Flacourtiaceae,
comprende nueve especies de ?rboles. Se extiende de Panam? a Guyana y al sur del Per?. Se presentan datos
sobre morfolog?a, anatom?a de lamadera, polen y distribuci?n geogr?fica. Se utilizaron an?lisis morfol?gicos y
moleculares para establecer las relaciones filogen?ticas entre las especies (del g?nero) y del g?nero en la familia.
Neosprucea incluye cinco especies reconocidas anteriormente y cuatro especies nuevas aqu? descritas (N. melas
tomatoides, N. paterna, N. tenuisepala y N. wilburiana). El nombre N. sucumbiensis se considera sin?nimo de
N. grandiflora. Se mantiene la transferencia anterior de N. kuhlmannii al g?nero Macrothumia. Se incluyen
descripciones detalladas y mapas de distribuci?n para cada especie; se ilustran todas las especies excepto
N. sararensis.
INTRODUCTION
Neosprucea (Salicaceae) is a neotropical genus of nine species of small to medium

sized trees. It is one of the most distinctive genera of Salicaceae, a family which was re
cently circumscribed to include a large part of the polyphyletic family Flacourtiaceae
(Chase et al. 2002; Alford 2005), in which Neosprucea was originally placed (Sleumer
1936). Unlike other genera in Salicaceae (or former Flacourtiaceae), which usually have
small, non-showy flowers and longitudinal anther dehiscence, Neosprucea has large,
showy flowers, poricidal anther dehiscence, and usually acrodromous leaf venation remi
niscent of Melastomataceae (Alford et al. 2005). The flowers are remarkably similar to
those of Elaeocarpaceae (see illustration in Cronquist, 1981, or photos in Mori et al.,
2002), and some authors have considered the tribe to which Neosprucea belongs, the
Prockieae, as intermediate between Flacourtiaceae and Elaeocarpaceae (Cronquist 1981;
Keating 1973; Miller 1975) or even as part of a broadly defined Tiliaceae including Elaeo
carpaceae (Hutchinson 1967). Evidence based on karyotypes (Morawetz 1981) and analy
ses of DNA sequence data (Chase et al. 2002), however, indicates that allies of
Neosprucea in the tribe Prockieae are closely related to other members of Salicaceae,
which are now placed in the order Malpighiales (APG 2003). Analyses of DNA sequence
data (see Relationships, below) affirm that Neosprucea, too, belongs with Salicaceae.
Elaeocarpaceae are distantly related and are now placed in the order Oxalidales (APG
2003).
Despite the unusual and striking morphological features of Neosprucea, the genus is
poorly known. Neosprucea was not sampled in the systematic treatments of wood
2 SYSTEMATICBOTANYMONOGRAPHS VOLUME 85
anatomy (Miller 1975), pollen (Keating 1973), or chromosome numbers (Grill 1990) of
Flacourtiaceae, and the generic treatment in the monograph of neotropical Flacourtiaceae
(Sleumer 1980) was based on only 32 specimens and described the fruits of only a single
species. The cause for this lack of study was clearly the paucity of material available. Only
six collections of Neosprucea were obtained before 1930 and only sixteen before 1960.
Recent, intensive collecting in Ecuador and Guyana have greatly augmented the material
on which to base a study the genus. The importance of Neosprucea has recently been
highlighted in a report of medicinal properties (Friedman et al. 1993) and in the listing of
one species (N. sararensis) as endangered (Calder?n 1997).
TAXONOMICHISTORY
The first known collections of Neosprucea were gathered in Colombia ("New

Granada") by the Spanish botanist Jos? Celestino Mutis. Unfortunately, little geographic
information is associated with his specimens because most of his labels were lost, and his
specimen numbers, like many of his time, reflected a filing system and not a chronologi
cal series (Blanco y Fern?ndez de Caleya & del Valle Stervinou 1991). According to his
notes and diaries (Blanco y Fern?ndez de Caleya & del Valle Stervinou 1991; Mutis
1957-1958), Mutis obtained the majority of his collections in the highlands near Bogot?
and in the lowlands near Mariquita between 1760 and 1808. He made three collections
of Neosprucea, including both a lowland species (N. wilburiana) and a montane species
(N. sararensis), but these remained unidentified until Sleumer annotated them in the 1930s
and assigned them to the single species of Neosprucea known at that time. Two other
Spanish explorers, Ruiz and Pav?n, collected a specimen of Neosprucea in 1827 from an
unknown locality in Peru. It was annotated as genus novum but also did not receive fur
ther attention.
The first collection of Neosprucea to receive detailed

study was obtained by English
explorer Richard Spruce. He collected the material during a stay in Tarapoto, Peru, be
tween 1855 and 1856, and sent it to George Bentham (Spruce 1908). Bentham described
the new entity in 1861 as Ba?ara grandiflora Spruce ex Bentham. Like Ba?ara Aubl., as
circumscribed today (e.g., Sleumer 1980), Neosprucea has alternate, stipulate, gland
toothed leaves, valvate sepal aestivation, numerous stamens, and highly intruded parietal
placentation. Ba?ara was the obvious genus inwhich to place the new species; at that time
Bentham had a broader circumscription of Ba?ara than modern authors (e.g., Sleumer
1980), which included about half of the currently recognized genera of the tribe Prockieae
(cf. Lemke 1988; Chase et al. 2002). Furthermore, some of Spruce's collection of
Neosprucea from Tarapoto did not include adequate flowering material, and Bentham
(1861) was probably unaware of the linear anthers and poricidal anther dehiscence, char
acters in which Ba?ara and Neosprucea differ. He did not mention these characters in his
description and thus had no reason to propose a new genus.
Additional collections of Neosprucea were gathered in the early twentieth century by
Ule in Peru and P?rez-Arbel?ez in Colombia. Sleumer used this new material to prepare
a revision of the subtribe Prockiinae (in the classification of Gilg, 1925; now
recognized
as tribe Prockieae, cf. Lemke 1988; Chase et al. 2002). Using the flowers collected
by Ule,
Sleumer (1936) recognized that the linear elongate anthers, in combination with the race
mose inflorescences and large flowers and fruit, warranted the creation of a new genus,
which he named Spruceanthus in honor of Richard Spruce (1817-1893) (cf. Seaward &
2008 NEOSPRUCEA 3
FitzGerald 1996). This name is a later homonym of Spruceanthus Verd., and Sleumer pro
posed the name Neosprucea in the revision that he published later (Sleumer 1938).
Elbert Little, Jr., described a second species of Neosprucea in 1948 and extended the
range of the genus to the Pacific basin of Ecuador. Jos? Cuatrecasas collected several
specimens of Neosprucea in Colombia in the 1940s and then published a short treatment
of the genus in 1955 in which he described three new species. Cuatrecasas (1955) pro
vided comments on the species and a key based principally on flowering material to dis
tinguish them. Unfortunately, he had few specimens at his disposal, and the characters
used in the keys often do not work even with the few specimens that he did have. Robyns
(1968) later recorded Neosprucea from Panama but was unable to assign the material to
species, because he had only fruiting material, which prohibited use of Cuatrecasas's di
agnoses and key.
Sleumer (1980) revised Neosprucea in his monograph of all neotropical Flacour
tiaceae. He recognized no new taxa and essentially maintained the circumscriptions of
previous authors, although he noted that Cuatrecasas's Neosprucea sucumbiensis was
"very close toN grandiflora" and assigned Robyns's "Neosprucea aff. sararensis" toN
grandiflora.
This study aims to assess species delimitation inNeosprucea, augment the descrip
tion of the genus with new data (e.g., pollen, wood anatomy, DNA sequence), discern re
lationships of the species within the genus and of the genus within the family using both
morphological and molecular data, and provide useful keys, illustrations, and distribution
maps to facilitate identification.
MATERIALSAND METHODS
Specimens of Neosprucea were borrowed from 21 herbaria (A, BM, BRIT, CAS,
COL, DUKE, F, G, GB, GH, HUA, L, LL, MA, MICH, MO, NY, S, TEX, U, US) and ex
amined at 16 additional herbaria (AAU, AMAZ, BH, HAO, HUT, IBE, K, MOL, OS, P,
QCA, QCNE, USM, VEN, W, WU) (acronyms from Holmgren et al. 1990). No specimens
were available at C, DS, FDG, INPA, GOET, JBSD, LY, M, MG, MSC, or USMS.
Spec
imens have been reported from INPA and GOET (Sleumer 1980); possibly they have been
misfiled or filed in another family, as elements of former Flacourtiaceae often are (e.g., in
Tiliaceae in some systems). Online or e-mailed images of specimens at COL (types), KW,
and BR were also available. In total, 218 collections were examined. Individuals and pop
ulations of Neosprucea were also observed in the field, and the herbarium specimens and
material in ethanol or FAA obtained during field work were studied. Study in the field in
cluded one month in Ecuador (October 2002), two weeks in Guyana (May 2003), two
months in Peru (November and December 2002), and three weeks in Venezuela (Decem
ber 2000 and January 2001).
Species Circumscription. Specimens were sorted into morphologically similar enti

ties, and variability was assessed by comparisons of duplicate collections and field notes.
Correlations of altitude and geography to the sorted entities were considered in order to
differentiate between genetic and environmental variation. The amplified phylogenetic
species concept of Nixon and Wheeler (1990) was employed, by which species are diag
nosable populations that have a unique combination of character states. This species con
cept and later development of its methods (Davis & Nixon 1992) are essentially precise
versions of that used by many museum taxonomists, exemplified by Cronquist (1988),

who defined species as the "smallest groups that are consistently and persistently distinct,
and distinguishable by ordinary means." False resolution of extra species was avoided by
using correlated, presumably unlinked characters and by minimizing reliance on charac
ters known to be variable in other closely related and better collected species (e.g., leaf
pubescence inHasseltia floribunda H. B. K. and Ba?ara guianensis Aubl.). Distribution
maps of the species were then generated using Online Map Creation (M. Weinelt;
www.aquarius.ifm-geomar.de/), based on the Generic Mapping Tools (GMT) software
(Wessel & Smith 1995).
Anatomy and Morphology. Wood anatomy of Neosprucea was investigated by cutting

26 urn sections of 1 cm diameter wood on a sliding microtome and by staining with 0.05%
toluidine blue O in 0.1 M sodium acetate at pH 4.4 for 20 s. Sections were destained with
water, mounted with water, and allowed to air dry. Sections were then soaked in 100%
ethanol, transferred to Hemo-DE (=d-limonene; Scientific Safety Solvents, Keller, TX),
and mounted in Permount (Fisher Scientific Company, Pittsburgh, PA). Five species, one
sample each, were studied (Table 1).
Description of the gross morphology follows the general terminology of Radford et
al. (1974), enhanced by details from Harris and Harris (1994) for vestiture and from
S team (1992) for seed sculpturing. Counts of teeth/cm were made in the middle third of
the lamina.
Pollen was examined using a, LEO DSM 960
LaB6 themoionic scanning electron
microscope (SEM) after coating naked pollen on double stick tape with 90 s of gold
palladium using a Balzar sputter coater (model SCD 050). Pollen of six species, one sam
ple each, was studied. Measurements of size and shape were made from 10 grains per
sample and are summarized with voucher information in Table 2.
DNA. DNA was extracted from approximately 0.02 g of field-collected, silica-dried

material or of leaf material taken from herbarium specimens using the Qiagen DNeasy kit
(Qiagen, Inc., Valencia, CA). The manufacturer's protocol was used, except that tissue
was crushed at ambient temperature (ca. 22?C) in extraction buffer with amortar and pes
tle. The plastid regions trnL-F and the 3' end of ndhF were chosen for amplification due
to their reported variation at the generic or sub-generic level (Soltis & Soltis 1998) and
presumably simple mode of inheritance (but see Shore et al., 1994, and Hansen et al.,
2007, who demonstrated maternal, paternal, and biparental inheritance in the related fam
ilies Turneraceae and Passifloraceae). The non-coding regions trnL and trnL-F are an in
tron and an intragenic spacer, respectively, and show high frequency of mutation. The
gene ndhF is a coding region for a subunit of nicotinamide dehydrogenase, which shows
approximately twice the mutation rate of rbcL (Olmstead & Sweere 1994).
DNA was amplified using a PTC-100 Peltier Thermal Cycler (MJ Research,
Waltham, MA) with a heated lid. The polymerase chain reaction (PCR) consisted of a
regime of two minutes initial denaturation at 94?C followed by 35 cycles of 94?C for 20 s,
55?C for 35 s, and 72?C for 25 s followed by three minutes of final extension at 72?C.
Primers c, d, e, and f of Taberlet et al. (1991) were used to amplify the trnL-F region, and
primers 5.5F and 10.2R of R. Nyffeler (in Davis et al. 2001) were used to amplify the 3'
end of ndhF. DNA was amplified in a volume of 50 uL, consisting of 48 uL of a master
mix and 2 uL of ca. 1-20 ng/uL template. Template often needed to be diluted 1:10 or
1:100. The master mix was formulated according to P??bo (1990), using concentrations
2008 NEOSPRUCEA 5
of reagents appropriate for the amplification of little or deteriorated DNA. This master
mix was composed of 37.75 |uLwater, 5 juLbuffer (1 mL buffer = 670 joL 1.0M Tris, pH
8.0-8.8, 20 |oL 1.0M MgCl2, 25 joLeach dNTP at 100 mM, 2 (iL 10mg/mL bovine serum
albumin, 208 |iL water), 2.5 |oL each primer at 10 ?M, and 0.25 |uL 5 U/|uL Taq poly
merase. Double-stranded PCR products were cleaned using QIAquick PCR Purification
Kits (Qiagen, Inc., Valencia, CA), and were sequenced by the Cornell University BioRe
source Center using an Applied Biosystems Automated 3730 DNA Analyzer with Big
Dye Terminator chemistry and AmpliTaq FS DNA polymerase. Because of large A-T ho
mopolymer regions, some amplifications and almost all sequencing of the trnL-F region
were done using all four primers. Thus, the trnL-F region is listed in the Appendix as two
separate regions, the ca. 600 bp trnL intron (primers c and d) and the ca. 350 bp trnL-F
intragenic spacer (primers e and f). Contiguous sequences were assembled from chro
matograms and checked for accuracy using Sequencher 4.1.4 (Gene Codes Corp., Ann
Arbor, MI). Sequences were obtained for 40 species (see Appendix).
Phylogenetic Matrices. Two phylogenetic matrices were generated: (1) amorpholog

ical matrix, focusing on character states that are variable within Neosprucea, and (2) a
molecular matrix {trnL + trnL-F + 3' end of ndhF), using an expanded sample of species
from tribe Prockieae. Because the sampling of species is quite different, the two matrices
were not combined.
species of Neosprucea were included in the matrix of morphological

All characters.
Hasseltia floribunda served as the global outgroup, and likely sister genera from tribe
Prockieae {Ba?ara, Hasseltiopsis Sleumer, Pineda Ruiz & Pav., and Prockia L.) were in
cluded as additional outgroups. Morphological characters and character states were cho
sen based on consistent or mostly consistent discrete differences (cf. Stevens 1991). In
general, primary homology assessments were made using the criteria of Patterson (1982):
characters were considered putatively homologous, if they passed the tests of similarity
and conjunction. The morphological characters were all unordered, except for leaf vena
tion. In that case, evolution of acrodromous venation (3-veined from base to apex) is hy
pothesized to proceed through actinodromous venation (3-veined from the base, lateral
secondaries not reaching the apex). With the one meristic character (perianth merosity),
the principal character state was coded. All species of Neosprucea are regularly 4-merous.
Some species rarely or infrequently produce 5-merous flowers, but they are coded as 4
merous. Only one species commonly produces 5-merous flowers, and it is coded as 4-mer
ous or 5-merous. Likewise, for the one color character (petal color), the principal charac
ter state was coded. The primary states of color are white to creamy white or yellow.
These are non-overlapping states, but species with petals of either color sometimes have
those petals also infused with pink. This pink color is considered a separate character. The
matrix of morphological characters was generated in and edited using WinClada (Nixon
2002). Inapplicable data were given the score "-", while missing or inconclusive data
were "?".
given
The matrix of DNA sequence data includes only five of the nine species of
Neosprucea but includes a much larger sampling of outgroup taxa (35 species). Homal
ium racemosum Jacq. served as the global outgroup, and numerous species of putatively
related taxa (cf. Chase et al. 2002; Alford 2005) were included. DNA sequences were ini
tially aligned using the default conditions (gap opening cost=10; gap extension cost=0.10)
in ClustalX (Thompson et al. 1997) and were then transferred toWinClada (Nixon 2002)
for alignment by eye. Gaps were coded for phylogenetic analysis using the simple indel
coding method of Simmons and Ochoterena (2000).
Phylogenetic Analyses. Parsimony analyses were conducted using the computer pro
gram NONA (Goloboff 1993) initiated through WinClada (Nixon 2002) in order to search
for most parsimonious trees (MPTs). Because of the small size of the morphological ma
trix, a heuristic analysis consisting of 1000 TBR replications holding 10 trees per replica
tion was performed. Character distributions on the phylogenies were studied using Win
Clada (Nixon 2002). Parsimony analyses of the molecular data were also conducted using
NONA (Goloboff 1993) initiated through WinClada (Nixon 2002) in order to search for
most parsimonious trees. First, uninformative characters were removed from the matrix
(mop command). Then, 10 sequential r?chet operations (Nixon 1999) were performed,
each with 200 iterations, holding two trees per iteration (h/2) at a 10% random constraint
level. These trees were then saved and swapped to completion in NONA (max*)
(Goloboff 1993). Up to 10000 additional trees were saved.
To test the strength of the signal for individual clades given these data sets, jackknife
values (Farris et al. 1996) were generated. Each matrix was sampled 1000 times, where
each replicate consisted of five random addition sequences with 10 trees held during TBR
swapping of each sequence followed by the calculation of the strict consensus of each
replicate (h/10; mult*5).
WOOD ANATOMY
The wood anatomy of five species was studied. Each species was represented by a
single wood collection. See Table 1 for vouchers and for characteristics that differed
among the species.
The wood of Neosprucea is diffuse porous, consisting of vessels in radial multiples
of 1-9 elements and septate fibers (Fig. 1A-C). The pith has numerous crystals. The
Table 1. Variable anatomical features of the wood of Neosprucea.
Length of Length of #-seriate

Radial upright ray procumbent ray procumbent
Species and Vessel multiples parenchyma parenchyma ray
voucher perforations of vessels (number of cells) (number of cells) parenchyma
N. grandiflora simple or 1-6 1-11 1-120 1-3
(Alford3103) scalariform
N. melastomatoides
(Alford3091) simple 1-5 (-7) 1-28 1-21 1-2
N. montana
(Alford3129) simple 1-5 1-15 (+?) 1-5 (+?) 1-2
N. paterna
(Alford3149) simple 1-8 1-12 1-22 (-30) 1-2
N. pedicellata
(Alford3076) simple 1-4 (-9) 1-34 1-13 (-19) 1-2
2008 NEOSPRUCEA 7
FIG. 1.Wood of Neosprucea.

anatomy A. Cross section, showing lack of axial parenchyma (N. paterna).
B. Cross section, indistinct growth rings and short multiples
showing of vessels (N. melastomatoides). C. Lon
gitudinal section, showing homocellular (above arrow) and heterocellular (below arrow) rays and septate fibers
(septum at arrow) (N. grandiflora). D. Longitudinal section, showing simple perforation at arrow (N. pedicel
lata). E. Polarized radial section, showing chambered upright ray parenchyma with prismatic crystals (one crys
tal at arrow) (N paterna). F. Radial section, showing upright and procumbent ray parenchyma (N melastoma
toides). Scale bar: A, C, E, 100 urn; B, 500 urn; D, F, 300 urn. (Based on: A, E, Alford 3149; B, F, Alford 3091;
C, Alford 3103; D, Alford 3076.)
vessel perforations are simple (Fig. ID) or infrequently (N. grandiflora) scalariform with
up to 9 bars. As in most members of Salicaceae (Metcalfe & Chalk 1950) and members
of Flacourtiaceae (Miller 1975) now placed in Salicaceae, axial parenchyma ismostly ab
sent (or scanty). Growth rings are usually present but diffuse and more evident in younger
growth. Intervascular pitting is alternate. Homogeneous and heterogeneous rays are pre
sent. If a ray is homogeneous, it consists of only upright ray parenchyma, 1-34 cells long.
If a ray is heterogeneous, it consists of short to long series of upright ray parenchyma,

1-28 cells long, terminal or alternating with procumbent ray parenchyma (Fig. ID, F).
The series of procumbent ray parenchyma is short to very long, 2-120+ cells long, and
1-2-seriate (sometimes 3-seriate inN. grandiflora). Prismatic crystals are present in both
the upright and procumbent ray parenchyma, occasionally in two or four chambers within
upright ray parenchyma cells (Fig. IE).
MORPHOLOGY
Habit. Species of Neosprucea are small treelets to medium-sized trees, occasionally

reaching 30 m tall with trunks of 60 cm dbh. Some species known only as treelets or small
trees probably grow taller, as most large-sized individuals of species known to attain such
heights and diameters (e.g., N grandiflora, N paterna) were collected following logging
or pipeline operations where individuals of such size were easily accessible. Furthermore,
individuals of some species of Neosprucea flower when only as tall as 1.5m, making them
easy to collect and probably skewing the number of collections to smaller individuals. The
labels of two collections of Neosprucea reported that the individuals were lianas or scan
dent shrubs {Harling & Andersson 12871, Stahl et al. 4338), but I suspect that these indi
vidual plants were lanky shrubs; the liana or scandent habit is unknown in other Sali
caceae and was not observed in the field work for this study.
Vestiture. Here and in the following descriptions the term "pubescent" is used tomean
presence of hairs that are straight, short, and soft.
Species of Neosprucea have an indumentum, although individual parts may be
glabrous or glabrescent. The indumentum consists of simple trichomes, which vary in
length, density, and disposition relative to the surface, resulting in parts that range from
sparsely pubescent, hirtellous, or strigillose to densely pubescent, hirsute, or sericeous.
Trichomes of Neosprucea also differ in color, ranging from white and silvery to golden
brown. The larger trichomes, especially on the abaxial surface of the sepals, are often eas
ily rubbed off.
Leaves. Leaves of Neosprucea are alternate and simple (Fig. 2A, B). They vary
greatly in size, even on a single individual. The margin of the lamina is toothed, and a
gland can be found at or near the apex of each tooth. Early in development, these glands
are typically marginal and crateriform (Fig. 12B) but usually become flat, abaxial, and
torus- or doughnut-shaped upon maturity (e.g., Fig. 20A, B). The teeth and glands are very
evident early in development, as is common in many species of wet-soil environments
(Burnham et al. 2001; Kowalski & Dilcher 2003). Perhaps the teeth make use of transient
light or build up energy reserves for use by the plant before leaves are fully mature (Lar
son & Gordon 1969; Larson et al. 1972; Larson & Dickson 1973; Baker-Brosh & Peet
1997; Roy er & Wilf 2006). The function of the glands inNeosprucea is unknown, but in
related members of Salicaceae they serve as sources of resin or nectar for defense against
insects and are often associated with hydathodes (Trelease 1881; Curtis & Lersten 1974,
1978, 1980; Wilkinson 2007).
Leaf venation inNeosprucea is actinodromous or acrodromous (Fig. 2A, B). In the
two species with actinodromous venation {N. paterna, N. rimachii), three principal veins
arise from the base but only the midvein extends to the apex. The lateral two veins
2008 NEOSPRUCEA 9
FIG. 2. Vegetative and floral morphology of Neosprucea. A. Acrodromous venation of N. melastoma

toides. B. Actinodromous venation of N. paterna. C. Minute stipule of AT.grandiflora. D. Well-developed pul
vini at the base and apex of the petiole inN. montana. E. Inflorescence of N. melastomatoides. F. Rower at late
anthesis and opening flower bud of N. grandiflora. G. Flowers o?N. paterna. H. Young infructescence o?N. pa
terna, showing perianth enclosing the developing fruit. (Based on: A, Alford 3092; B, Alford 3133; G, H, Alford
3149; C, F, Alford 3103; D, Alford 3129; E, Alford 3091.)
(secondaries) extend about 2/3 the distance to the apex. The secondary venation is semi
craspedodromous, meaning that the secondaries form loops near the margin with the pro
duction of another vein, which extends directly to a tooth. The other species of
Neosprucea have acrodromous basal venation. In this case, the three principal veins arise
from the base and all extend to (or near) the apex. Even though additional major veins
may branch from the midvein, I have chosen to call the two principal lateral veins "sec
ondaries" as well, which suggests homology with the basal secondaries of the species with
actinodromous venation. The tertiary veins are perpendicular to the primary and sec
veins. Venation is raised on the abaxial surface. In N. montana and N. sararensis,
ondary
the raised venation is often winged between the midvein and two secondary veins at the
base of the leaf, forming small domatia. Stomata are abaxial, numerous, and paracytic (ru
biaceous) (N=3: N. grandiflora [Alford 3103], N melastomatoides [Alford 3091], N pa
terna [Alford 3149]).
Adaxially, the leaves are usually glabrous, except near the lamina-petiole junction,
but they are infrequently covered in trichomes on the major veins inN paterna and N
pedicellata, and even sparsely to densely covered in trichomes throughout inN rimachii.
Abaxially, the leaves are glabrous, sparsely pubescent or strigillose along the major veins,
or pubescent, hirtellous, or hirsute throughout.
Three species of Neosprucea (N. montana, N sararensis, N. wilburiana) have well
developed pulvini at the base and apex of the petiole (Fig. 2D). This is an unusual feature
for Salicaceae and is unfortunately another character thatmay lead to confusion with Mal
valean families. The pulvini are solid, but their function and behavior are unknown. Less
conspicuous pulvini are sometimes present in the other species, and N. pedicellata occa
sionally develops only the apical pulvinus.
One to five small projecting glands are found at or near the apex of the petiole inN
rimachii and N. paterna. In the other species, glands associated with the leaf teeth are in
frequently located close to the petiole, almost appearing to be petiolar glands.
The leaves of Neosprucea are stipulate, but the stipules are caducous. The stipules are
small, often less than 1mm long (N. grandiflora, N. pedicellata, N tenuisepala', Fig. 2C),
or quite long, up to 1 cm (N paterna). The petioles are pubescent or glabrescent, and the
stipules are always strigillose or densely pubescent.
Leaves of Neosprucea lack cyanogenic glycosides (N montana [Steyermark 118458],
N. pedicellata [Dodson & Gentry 6318, Dodson 10066], and N wilburiana [Robyns 7997
(sic), probably Dwyer & Correa 7997]) (Spencer & Seigler 1985).
Inflorescence. The inflorescence of Neosprucea is a determinate thyrse (=thyrsoid),

which is a branched inflorescence with a terminal flower and cymose partial inflores
cences (Weberling 1989; Fig. 2E-H). In other words, the inflorescence consists of a sin
gle axis bearing several triads, or 3-flowered cymes, and resembles a panicle. This type of
inflorescence is common in closely related families, as reviewed for Passifloraceae by de
Wilde (1971a) and for Violaceae by Hekking (1988). In some cases the determinate thyrse
is reduced or depauperate, consisting of a single axis bearing several 1-flowered cymes,
which forms a raceme-like inflorescence referred to as a pseudoraceme by C. Anderson
(1997), W. Anderson (1981), and Hekking (1988). Herein, this reduced inflorescence is
simply called a "raceme," and the branched inflorescence is called a "panicle."
InNeosprucea, the ultimate part of the inflorescence consists of an axis with a bract,
two bracteoles, an articulation, and a flower. The bracteoles (a kind of "prophyll" sensu
Weberling, 1989) are usually located below the articulation that is below the flower. This
2008 NEOSPRUCEA 11
arrangement has resulted in different interpretations of the pedicel. In Flacourtiaceae (both

those genera now placed in Salicaceae and those placed in other families) and the closely
related Passifloraceae, the unit above the bracteoles has been considered the pedicel, re
sulting in usage such as "pedicel articulated above the base" (Sleumer 1980; Alford et al.
2005; Alford 2006a, b) and floral "stipe" (deWilde 1971b; MacDougal 1994). Others con
sider the articulation the defining point of the pedicel (e.g., C. Anderson 1997). In this
viewpoint, the bracteoles have moved from the articulation to some point lower (some
times much lower) on the peduncle. The latter view is taken here, principally because
pedicels so defined are easier to recognize and measure, especially if bracteoles are ca
ducous. Thus, as used here, one finds bract, proximal part of peduncle, pair of bracteoles,
distal part of peduncle, articulation, pedicel, and flower.
The inflorescence axes are sparsely to densely covered in trichomes, rarely glabres
cent near the base, sometimes with numerous appressed hairs (strigillose) and scattered
longer hairs (hirsute). When dry, the axes appear golden or silvery due to the trichomes,
but when trichomes are scarce, the axis appears black. Bracts and bracteoles are usually
covered in trichomes but are sometimes glabrous on the adaxial surface. The hairs are usu
ally longer and denser at the junction of peduncle and pedicel (the articulation).
Flowers and Perianth. The flowers of Neosprucea are quite large and showy relative
to those of most members of Salicaceae (Fig. 2E-H). The flowers can extend to almost 5
cm in diameter. The perianth is 4-5-merous and consists of both calyx and corolla (Fig.
2E-G). The number of sepals and petals is usually four each, and the few 5-merous flow
ers are apparently randomly arranged, not arranged according to their terminal or lateral
position, as is the case in the related family Ochnaceae (Weberling 1989). Both calyx and
corolla exhibit valvate aestivation. The calyx is green, yellowish green, or brownish
green. The corolla is bright white to creamy white or yellow. Occasionally, the corolla (ei
ther white or yellow) is infused with pink.
The has an indumentum. In some {N. montana, N. tenuise
perianth always species
pala), the abaxial surface of the calyx is sparsely strigillose, and the sepals appear black
when dried. In other species {N melastomatoides, sometimes N sararensis and N. wil
buriana), the trichomes are dense but easily rubbed off in dried specimens, revealing the
dark epidermis below. The corolla usually has more trichomes than the calyx and is some
times sericeous. Both the calyx and corolla may have lines of denser pubescence alter
nating with less dense pubescence, or lines of pubescence alternating with glabrous areas
(e.g., N. melastomatoides, N. montana, N. tenuisepala). The receptacle is moderately to
densely hirsute, with hairs commonly reaching 1mm in length.
Androecium.The androecium consists of numerous stamens ([30-] 60-400), in some

species surrounded by a whorl of filamentous staminodes. The stamens are often posi
tioned tightly against the ovary (Fig. 10D, E) but may also be rather loosely arranged
(Fig. 14C, D). The filaments are short (0.5-6 mm) and inconspicuous. The anthers are
large and linear (2-11 mm), and dehisce by oblique extrorse pores at the apex, features
found nowhere else in Salicaceae. The staminodes may be abruptly differentiated from the
stamens and obvious (Fig. 10D), or they may be obscure and the endpoint of a transition
where the outer stamens have smaller anthers and longer filaments (Fig. 8G, 14C-F).
Filaments are glabrous or hirsute, and the anthers are glabrous to densely hirsute. Hairs on
the anthers, if present, usually arise from the adaxial surface of the connective. The sta
minodes are usually glabrous, rarely hirsute at the base.
Table 2. Pollen of Neosprucea: variation in shape and size.
Species and Voucher Shape Polar Length (urn) Equatorial Width (um)
N. grandiflora spherical 22-24 20-22
(Alford3103)
N. melastomatoides subprolate to prolate 24-30 14-22
(Alford3091)
N. montana subprolate 32-34 26-30
(Campos et al. 3028)
N. paterna subprolate 36-40 28-34
(Alford3149)
N. pedicellata spherical to subprolate 18-22 16-18
(Gentry12001)
N. wilburiana spherical to subprolate 24-26 20-24
(Dressler 4339)
Pollen. The pollen of six species was sampled (Table 2). Each species was repre
sented by a single pollen collection. The pollen grains of Neosprucea are single, isopolar,
and tricolporate (Fig. 3A-C). The pollen of Neosprucea is spherical to prolate, with polar
lengths of 18-40 |im and equatorial widths of 14-34 |im. The exine is tectate, and exine
sculpturing is reticulate (Fig. 3D). These are the same shapes and average sizes as the
fairly monotonous pollen of most Flacourtiaceae now placed in Salicaceae (Keating
1973).
Gynoecium. The gynoecium of Neosprucea consists of one superior paracarpous

ovary of 4-9 carpels capped by a single, persistent style with a minutely or obscurely
lobed stigma. Each slightly to highly intruded parietal placenta bears numerous anat
ropous ovules, except inN. montana, which appears to have four highly intruded fertile
placentae and four less intruded sterile placentae (N=l).
Fruits. Fruits of Neosprucea are poorly collected and thus poorly known, perhaps be
cause the fruits are readily consumed. Only immature fruits have been seen for two
species {N. montana, N. paterna). The remaining species, except M rimachii, have juicy
berries about 1 cm in diameter, which are presented for frugivores when the persistent pe
rianth is open. Neosprucea rimachii, on the other hand, has a large and sericeous fruit that
is probably a "dry berry" (non-fleshy, ind?hiscent, multi-seeded, thin-walled fruit derived
from amulti-carpellate pistil, common in genera of the former Flacourtiaceae now placed
in Salicaceae). Ovaries and fruits inNeosprucea are usually glabrous but can be hirsute
{N. melastomatoides, infrequently N. grandiflora) or even sericeous {N. rimachii).
Seeds. Neosprucea fruits contain ca. 15-28 exarillate seeds, which are irregularly
ovoid or ellipsoid, sometimes several-angled by compression against the fruit wall and
other seeds (Fig. 3E, F). The seeds are glossy chestnut-brown and minutely reticulate
areolate with a brittle testa, a large crateriform hilum, an apiculum formed from the
integuments at the opposite end, and a light brown raphe, which extends the length of the
seed. The few embryos seen {N. grandiflora) were erect and spathulate. No data on seed
anatomy are available.
2008 NEOSPRUCEA 13
FIG. 3. Pollen and seeds of Neosprucea. A. Pollen of N. paterna. B. Pollen oiN. melastomatoides. C. Polar
view of pollen of N melastomatoides. D. Fine detail of the exine sculpturing of N melastomatoides. E. Seed of
N melastomatoides, showing crateriform hilum at apex and raphe. F. Detail of seed sculpturing inN grandi
flora. Scale bar: A, B, 20 urn; C, 10 urn; D, 2 urn; E, 1 mm; F, 200 urn. (Based on: A, Alford 3149; B, C,
D, Alford 3091; E, Palacios 2980; F, Zaruma 583.)
POLLINATIONBIOLOGY
The anther dehiscence in Neosprucea is poricidal, and the perianth of some species
has lines of pubescence. Rowers with these features are often buzz-pollinated (Endress
1994). Large bees were observed visiting the flowers of N. melastomatoides (pers. obs.),
but adequate pollination studies have not been conducted.
In most herbarium specimens of Neosprucea the flowers are closed. The persistent
perianth is also closed around any developing or mature fruit (Fig. 2H). In the field, daily
observations for two weeksin Ecuador indicated that flowers were open only during
sunny days. (No observations were made at night.) This behavior of opening during sunny
periods may be correlated to pollen viability and pollinator visitation. Many pollinators do
not forage in the rain, and pollen viability is sometimes reduced after precipitation (Ed
wards & Jordan, 1992; Huang et al., 2002; but see Garcia & Hoc, 1997, for a case in the
closely related Passifloraceae where pollen viability was not reduced by rain).
RELATIONSHIPS
Thirteen morphological characters (Tables 3,4) were coded for a phylogenetic analy
sis of Neosprucea. A larger matrix of 40 species and 222 parsimony informative charac
ters was generated based on plastid DNA sequence data (trnL-F and the 3' end of ndhF).
DNA was extracted from two collections of Neosprucea grandiflora (Alford 3103, Ro
moleroux et al. 2259), but the resulting sequences were identical. Voucher information and
GenBank accession numbers are listed in the Appendix.
In both the morphological and molecular analyses, Neosprucea is recovered as mono
phyletic (Figs. 4-6). The morphological analysis resulted in seven most parsimonious
trees (MPTs) of length 25 (CI=0.64, RI=0.78). A strict consensus of these trees (Fig. 4)
resulted in an unresolved polytomy for the sister taxon (taxa) of Neosprucea but provided
good resolution within the genus. The molecular results, too, fail to point out confidently
the sister taxon (taxa) o? Neosprucea. The analysis of molecular data resulted in 98 MPTs
of length 279 (0=0.68, RI=0.86). A strict consensus (Fig. 6) yielded a polytomy among
the genera Ba?ara, Pineda, Prockia, and a poorly supported (13% jackknife) Hasseltiop
sis + Neosprucea. Given these molecular data, Ba?ara and Prockia as circumscribed by
Sleumer (1980) are not monophyletic, regardless of how the polytomy is resolved.
TABLE 3. Characters and character states used in the morphological analysis o? Neosprucea and relatives.
1. Leaf venation: pinnate = 0; actinodromous =

1; acrodromous perfect
= 2.
[additive].
2. Leaf base: attenuate to rounded = 0; abruptly cordate or auriculate = 1.
3. Glands at the leaf teeth: papillate/spherulate = 0; = 1.
torus-shaped
4. Ind?ment of glands at leaf teeth: glabrous = 0; pilose = 1.
5. Petiolar glands: absent = 0; present = 1.
6. Pulvini at petiole base and apex: absent = 0; present = 1.
7. Merosity of perianth: 3-merous = 0; 4-merous = 1; 5-merous = 2.
8. Petal color at anthesis: white to creamy white = 0; yellow = 1.
9. Receptacular disk glands: absent = 0; present = 1.
10. Filaments: = 0; villous = 1; hirsute = 2.
glabrous
=
11. Anther connectives: glabrous 0; hirsute = 1.
12. Filamentous staminodes: absent = 0; present = 1.
13. Anther dehiscence: longitudinal slits = 0; apical slits/pores = 1.
2008 NEOSPRUCEA 15
Table 4. Matrix of morphological characters used in the phylogenetic analysis of Neosprucea and rela
tives. See Table 3 for a list of characters; A=0, 1; B=l, 2.
Character number 1 10 11 12 13
Hasseltia floribunda 1 0
Ba?ara guianensis 0 0
Pineda incana 0 0
Prockia crucis ? 1
Hasseltiopsis dioica 1 0
Neosprucea rimachii 1 1
N. paterna 1 1
N. wilburiana 0 2
N. montana 2 0
N. sararensis 2 0
N. pedicellata 2 0
N. tenuisepala 2 0
N. grandiflora 0 2
N. melastomatoides 2 0
Hasseltia floribunda
?
Ba?ara guianensis
-Pineda incana
-
Prockia crucis
100 dioica
-Hasseltiopsis
Neosprucea rimachii
Neosprucea paterna
Neosprucea pedicellata
Neosprucea wilburiana
53 Neosprucea montana
Neosprucea sararensis
Neosprucea tenuisepala
Neosprucea grandiflora
Neosprucea melastomatoides
FIG. 4. Strict consensus of the most parsimonious trees recovered in a phylogenetic analysis of morpho
logical data of Neosprucea and close relatives. Jackknife values >50% are shown above branches. MPTs=7,
L=25, CI=0.64, RI=0.78.
5 10
1 7
Pineda incana
9 10
Hasseltiopsis dioica
8 12
5 12 Ba?ara guianensis
Prockia crucis
3 9
2 4 5
Neosprucea rimachii
Neosprucea paterna
11 13
I? Neosprucea pedicellata
Neosprucea wilburiana
1 8
Neosprucea montana
Neosprucea sararensis
?
Neosprucea tenuisepala
10
1? melastomatoides
Neosprucea
FIG. 5. One of the seven most parsimonious trees recovered in a phylogenetic analysis of morphological
data of Neosprucea and relatives, showing character support and transformations. Numbers above the branches
refer to the characters in the data matrix (Tables 3, 4), black squares represent unique gains, and white squares
represent losses or homoplasious gains. L=25, 0=0.64, RI=0.78, fast optimization (ACCTRAN).
Character support is shown in Fig. 5. Characters are mapped onto one of the MPTs
and are optimized using fast optimization (=ACCTRAN, in equally parsimonious opti
mizations, it chooses evolution and loss instead of parallelism). Neosprucea is supported
as a monophyletic group by having principally 4-merous perianth, hirsute anther connec
tives (later reversed inN. montana and N sararensis), and anther dehiscence by oblique
apical pores. Jackknife values are low for this clade in both the morphological and mole
cular analyses.
Within Neosprucea, two major clades are recovered (Figs. 4-6): one of N rimachii
and N. paterna and one of the remaining species of Neosprucea. Neosprucea rimachii and
N. paterna have the plesiomorphic character states of actinodromous venation and yellow
corolla, and have the synapomorphic character states of abruptly cordate to auriculate leaf
bases and pilose leaf teeth. They also have projecting glands at the apex of the petiole, a
character state that apparently has evolved multiple times in Salicaceae (e.g., in Ba?ara,
Pineda, Pleuranthodendron L. O. Williams). Synapomorphies of the other species of
Neosprucea are white to creamy white corolla and the distinctive acrodromous (melas
tome-like) venation. Within this group, the placement ofN. pedicellata is unresolved, and
the other species form two species groups: (1) a N. grandiflora group {N. grandiflora,
2008 NEOSPRUCEA 17
Homalium racemosum
Xylosma bahamense
Scolopia spinosa
Ahernia glandulosa
Macrothumia kuhlmannii (=Banara kuhlmannii)
Pleuranthodendron lindenii
EHasseltia allenii
Hasseltia guatemalensis
Abatia canescens
Abatia spicata
Abatia mexicana
-
Abatia stellata
Abatia parviflora
Abatia angeliana (=Aphaerema spicata)
Azara alpina
Azara celastrina
Azara lanceolata
Azara salicifolia
Prockia pentamera
Ba?ara domingensis
Ba?ara vanderbiltii
Pineda incana
Pineda ovata
Hasseltiopsis dioica
Neosprucea paterna
Neosprucea montana
63 i-Neosprucea pedicellata
Neosprucea melastomatoides
Ba?ara axilliflora
Prockia costaricensis
Prockia crucis
Prockia flava
Ba?ara minutiflora
Ba?ara tomentosa
Ba?ara umbraticola
Ba?ara regia
Ba?ara guianensis
Ba?ara n?tida
FTG. 6. Strict consensus of the 98 most parsimonious trees recovered in a phylogenetic analysis of molec
ular data (plastid trnL-F and the 3' end of ndhF) of Neosprucea and relatives. Jackknife values >50% are given
above the branches. L=279, CI=68, RI=86.
N. melastomatoides, N. tenuisepala), characterized by hirsute filaments, well-developed

filamentous staminodes, and lack of petiolar pulvini, and (2) aN. montana group {N.mon
tana, N. sararensis, N. wilburiana), characterized by glabrous filaments, well-developed
petiolar pulvini, and lack of filamentous staminodes. Two species of the N. montana
group, N. montana and N. sararensis, form a clade within this group; both have glabrous
anthers. They also have overlapping distributions and grow at high elevations.
Analyses of the molecular data are not fully congruent with analyses of the morpho
logical data. Neosprucea paterna is resolved as sister to the rest of Neosprucea as ex
pected {N. rimachii was not included in the analysis), but N. pedicellata and N. melas
tomatoides are resolved in a clade without N. grandiflora. This anomaly could be the
result of limited variation in the molecules; however, the morphological analysis results
in an unresolved placement for N. pedicellata and a weakly resolved clade of N grandi
flora, N. melastomatoides, and N. tenuisepala. Clearly, more morphological and molecu
lar variation is needed to resolve the fine-scale relationships.
HABITATS,DISTRIBUTION,AND BIOGEOGRAPHY
Neosprucea ranges from Panama to Guyana and to southern Peru (Fig. 7). All species
of Neosprucea grow in areas with no rainfall deficit, meaning that there is adequate water
available for growth throughout the year. Most species grow in low altitude tropical rain
forest, but N. montana and N. sararensis grow in higher altitude cloud forest up to 2300
m.
Like many members of Salicaceae, the species of Neosprucea observed in the field
take advantage of disturbance for reproduction. Saplings of Neosprucea are commonly
found along streams, in small light gaps, and along trails or small roads. Apparently, they
do not prefer or tolerate as much disturbance as other species of Salicaceae, such as Ba
?ara guianensis and Hasseltia spp., which are commonly found along larger streams,
rivers, and even highways or fencerows. In fact, only one label indicates thatNeosprucea
was found in a highly disturbed locality ("potreros"). Few labels indicate disturbed forest,
and most labels indicate primary, undisturbed forest ("bosque primario"). Mature treelets
or trees are found in forest that seems undisturbed, but seedlings or saplings are rarely
seen there, hinting that establishment coincides with creation of small gaps in generally
mature forest.
Most species occur on alluvial or weathered soil, but N. paterna and N. rimachii
occur on sandy soils. The species that grow at higher elevations are often found in rocky
soil along small streams.
Because all close relatives of Neosprucea are neotropical {Abatia Ruiz & Pav., Azara
Ruiz & Pav., Ba?ara, Hasseltiopsis, Pineda, Prockia', Figs. 4-6), one can confidently infer
that the ancestors of Neosprucea were neotropical. The majority of these close relatives,
as well as themajority of the species of Neosprucea itself, presently occur in South Amer
ica, which suggests a South American origin in particular. The sister taxon (taxa) of
Neosprucea is (are) not resolved (see Relationships), but because its relatives are pre
dominantly South American and the uplift of the northern Andes is a fairly recent event,
occurring during Pliocene and Pleistocene times (van der Hammen 1974; Flenley 1979),
the immediate ancestors of Neosprucea probably occupied lowland tropical forest
in South America and diversified by expanding into a new niche {N. montana and
N. sararensis into cloud forest), taking advantage of new geographical opportunities
2008 NEOSPRUCEA 19
85? W 80? 75? 70? 65? 55?

60?
FIG. 7. Distribution of Neosprucea.
(N. wilburiana into the land bridge of Panama), and by allopatric speciation with the up
lift of the Andes (N. pedicellata on the western side of the Andes). In this scenario, the
distribution of the two species sister to the rest of the genus (N. rimachii, N. paterna) re
flect something of the ancestral distribution pattern. Both grow in lowland tropical forest,
one in Amazonian Peru and Colombia and the other in Guyana.
TAXONOMY
Note. Measurements are based on all specimens studied that exhibit the given char
acter. When the number of specimens exhibiting the character is unusually small (e.g.,
specimens with mature fruit) or sampling is artificially small, the measurement is accom
panied by a number indicating sample size (N=#).
Neosprucea Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 14: 47. 1938. Spruceanthus
Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 13: 362. 1936, non Spruceanthus
Verdoorn, Ann. Bryol. 4 (Suppl.): 151. 1934.?TYPE: Neosprucea grandiflora
(Spruce ex Bentham) Sleumer.
Treelets or trees to 30 m tall, sometimes reproductively mature when as small as

1.5 m tall. Trunk to 60 cm dbh, infrequently forming small buttresses (N. montana,
N. pedicellata); bark whitish, gray, or light brown, general aspect smooth and tight, de
tailed aspect finely fissured-scaly, slash orange-brown. Branches light green to coppery
green to dark gray when young, drying chestnut brown to black, but becoming gray with
growth, obscurely or distinctly 3^-sided, becoming terete with growth, sparsely to
densely covered in trichomes, sometimes glabrescent; lenticels rare to numerous, some
times obscure, white to maroon-brown, circular to linear, the long axis parallel to the
branch axis, more conspicuous and elongated with growth. Leaves alternate, simple; leaf
blades elliptic in most species, less often ovate to obovate, infrequently slightly asym
metrical with respect to the midvein, both very large and very small leaves commonly
found on the same branch; apex obtuse to abruptly acuminate or caudate; base attenuate
to truncate or abruptly cordate to almost auriculate, usually eglandular, but glands associ
ated with leaf teeth sometimes located near the lamina-petiole junction; lamina margin
serrate-crenate, rarely obscurely so, or almost entire (N paterna, adult leaves), teeth ob
tuse (convex-convex to convex-retroflexed) or merely like small half-ellipses along the
leaf margin, with a glabrous or pubescent (N. paterna, N rimachii) gland at each tooth,
gland commonly abaxial, abutting the leaf margin or slightly inset at maturity, or less fre
quently marginal; thinly chartaceous to coriaceous; adaxial surface often glossy, dark
green to light green to grayish green (TV.wilburiana), infrequently dull, usually glabrous
except near lamina-petiole junction, infrequently pubescent to glabrescent on major veins
(N. paterna, N pedicellata), rarely sparsely to densely pubescent throughout (N. ri
machii); abaxial surface dull green or light green, glabrous, sparsely pubescent or strig
illose along the major veins, soft pubescent, hirtellous, or hirsute throughout; venation
actinodromous, 3-7-veined from the base, or acrodromous perfect, 3-veined from base to
apex, an upper 0-5 secondary pairs (or secondary veins per side of the lamina) arising
above the 3 basal veins at an acute angle from the midvein near apex, prominent tertiaries
perpendicular to the midvein (and more or less perpendicular to the pair of secondaries,
when venation acrodromous) and alternating with less prominent tertiaries, the first and
last several tertiaries more closely spaced and less prominent than those in the middle,
semicraspedodromous, veins raised abaxially, sometimes winged between the midvein
and two secondary veins producing small domatia (N. montana, N. sararensis). Petioles
sparsely to densely covered in trichomes or glabrescent, sometimes reddish in saplings or
adults (N montana, N paterna), canaliculate or flat on adaxial surface; solid pulvini at the
base and apex absent, small and obscure, or sometimes prominent and well-developed (N.
montana, N. sararensis, N. wilburiana, sometimes N. pedicellata at apex); 1-3 (-5) pro
jecting glands sometimes present (N. paterna, N rimachii), usually on the distal third.
Stipules lanceolate to deltate, entire or with small glandular teeth, strigillose or densely
pubescent, caducous. Inflorescences subterminal, or terminal racemes, raceme
axillary,
like panicles,or panicles of 1-21 (-31) flowers, sometimes reduced to fewer flowers due
to abscission, sometimes congested into an axillary fascicle-like raceme (N. wilburiana),
the "racemes" or "panicles" complete or depauperate determinate thyrses consisting of a
single axis bearing several full or depauperate triads, or 1-3-flowered cymes, the axis
sparsely to densely pubescent or glabrescent near the base, sometimes quite thick (N.
grandiflora, N. melastomatoides, N. sararensis), sometimes flared or expanded at the
nodes of the triads (the ultimate cymose node beneath the flowers); bracts lanceolate to
triangular to ovate, abaxially moderately to densely pubescent, adaxially glabrous to
densely pubescent, caducous; peduncle sparsely to densely pubescent with or without in
termixed erect hairs, sometimes ridged, when dry appearing as the color of the pubescence
or black if pubescence sparse, the articulation between the peduncle and pedicel
2008 NEOSPRUCEA 21
sometimes more densely pubescent; bracteoles lanceolate to triangular to ovate, some

times inconspicuous (even the scars), located just below to well below the articulation,
abaxially densely pubescent, adaxially glabrous to densely pubescent, caducous; pedicels
very short (flowers subsessile) or up to 20.6 mm long, pubescent like the peduncle. Flower
buds oblate to ovoid to pyramidal to squarish with a pyramidal top, commonly capped by
ridges where the sepals abut against each other, green to yellow to shades of pink, red, or
brown, sparsely to densely pubescent; perianth aestivation valvate. Flowers hermaphro
ditic and complete. Sepals 4-5, widely deltate to narrowly elliptic to lanceolate to oblong,
abaxially brownish green to green to yellowish green, drying darker than the petals,
sparsely to densely covered in trichomes, usually drying black when sparsely strigillose
{N. montana, N. tenuisepala), several to numerous veins sometimes evident, adaxially
pale green, densely short-pubescent or tomentose except for a central glabrous strip {N ri
machii, N. wilburiana, some N. montana), the pubescence sometimes easily rubbed off.
Petals 4-5, widely deltate to lanceolate to oblong, not thickened, yellow {N. paterna, N.
rimachii) or creamy white to white, sometimes infused with pink, abaxially moderately,
densely, or unevenly pubescent with hairs longer than those of the calyx, or sericeous,
adaxially short-tomentose or moderately, densely, or unevenly pubescent in lines with
hairs longer than those of the calyx. Receptacular disk absent. Receptacle moderately to
densely hirsute or pubescent, sometimes inconspicuously so {N. rimachii). Androecium of
numerous stamens, ca. (30-) 60-400; filaments glabrous to densely hirsute, flattened; an
thers linear, glabrous to hirsute, especially on the adaxial surface of the connective, de
hiscent by oblique extrorse pores at the apex, the pores sometimes splitting longitudinally
for up to 2 mm below the pores. Staminodes absent, transitional (i.e., with subtle outer
rows of stamens with reduced anthers gradually merging into filamentous staminodes
lacking anther remnants), or filamentous, numerous, white to yellow, flattened, infre
quently hirsute near the base, rarely distally, 1/3 as long as the stamens to longer than the
stamens. Gynoecium a single, superior, paracarpous ovary, spherical to ovoid, sometimes
weakly lobed, light green, usually glabrous or sparsely but thoroughly hirsute {N melas
tomatoides), rarely densely tomentose or sericeous {N. rimachii); placentation parietal,
usually highly intruded, placentae 4-9; ovules numerous (>50); style one, glabrous to hir
sute; stigma minutely lobed or with several small terete projections. Fruit a fleshy berry
or rarely a dry berry {N rimachii), spherical to widely ovoid, yellowish green, green, or
green with white speckles, glabrous to densely tomentose or sericeous {N rimachii),
perianth and style persistent. Seeds ca. 15-28 per fruit, irregularly ovoid or ellipsoid, often
several-angled by compression against the fruit wall and other seeds, glossy, light to dark
chestnut brown, minutely reticulate-areolate, testa brittle; raphe extending the length of
seed, light brown; hilum crateriform; micropyle becoming an apiculum; embryo erect,
spatulate. Chromosome number unknown.
Neosprucea is distinctive in Salicaceae (and among all former Flacourtiaceae sensu

Lemke, 1988) by its combination of actinodromous or acrodromous leaf venation
(3-veined leaves), crateriform or torus-shaped glands at the leaf teeth, and poricidally de
hiscent anthers. The two species with actinodromous venation may be confused with sev
eral genera in Salicaceae, most Ba?ara, Hasseltia H. B. K., Pleuranthoden
commonly
dron, or Prockia, but these genera have globose, longitudinally dehiscent anthers. Ba?ara
and Prockia can also be distinguished by their 3-merous perianth, and Hasseltia and Pleu
ranthodendron can be distinguished by their paired glands at the intersection of petiole
and leaf blade. Neosprucea is infrequently mistaken for Lunania Hook., which also has
+/- acrodromous venation, but the leaves in Lunania are pellucid-punctate, exstipulate,
and commonly entire. Other ways to differentiate Neosprucea from species in the former
Flacourtiaceae may be found inAlford (2003). Species of Neosprucea are more often con
fused with members of other families. The species of Neosprucea with actinodromous ve
nation resemble many taxa of Tiliaceae and Euphorbiaceae, but they can be distinguished
from those families vegetatively by the presence of the torus- or doughnut-shaped glands
associated with the leaf teeth. The species with acrodromous venation are infrequently
confused with members of Melastomataceae or Loganiaceae (Strychnos L.), but those
taxa almost always have opposite leaves and lack the characteristic glands at the leaf teeth.
Species of Neosprucea with acrodromous venation may also be confused with taxa of
Menispermaceae, but menisperms are mostly lianas and have long petioles with flexed
apices. One of the hardest genera to distinguish vegetatively from Neosprucea is Am
pelozizyphus Ducke (Rhamnaceae). It differs from Neosprucea in its liana habit and in
having an entire, hyaline leaf margin and an indumentum of reddish hairs.
Key to the Species of Neosprucea
1. Leaf base cordate or almost auriculate; leaf venation actinodromous (3-7-veined from the lamina
base, secondary veins extending ca. 2/3 or less of lamina length); petals usually yellow at anthesis.
2. Leaves of mature plants elliptic or obovate, toothed at base or essentially entire; ovary and fruit
glabrous. 4. N. paterna
2. Leaves of mature plants ovate, serrate-crenate throughout; ovary and fruit densely tomentose to
sericeous. 6. N rimachii
1. Leaf base rounded, obtuse, acute, or attenuate; leaf venation acrodromous perfect (3-veined from
lamina base to apex; melastome-like); petals usually white to creamy white at anthesis.
3. Anthers glabrous (or rarely with a few scattered hairs); staminodes absent; middle to higher ele
vations, 950-2300 m.
4. Filaments 1.3-1.7 mm
long, shorter than the anthers (2.2-3.3 mm); abaxial epidermis of
sepals usuallydrying black, lightly pubescent throughout, epidermis visible through any
pubescence; inflorescence usually a raceme; leaves 5-16.5 (-20.1) cm long. 3. N montana
4. Filaments 2.2-2.5 mm long, longer than the anthers (2 mm); abaxial epidermis of sepals dry
ing black, but usually concealed by dense pubescence, all or most of the epidermis not visi
ble through the pubescence; inflorescence usually a panicle; leaves 11.5-28.1 cm long.
7. N. sararensis
3. Anthers hirsute; staminodes present (except for N. wilburiana), although sometimes obscure;
lower to middle elevations, up to 1300 (-1850) m.
5. Staminodes absent; inflorescences <4.5 cm long, usually congested and <2.0 cm long; Panama
and northern Colombia. 9. N. wilburiana
5. Staminodes present; inflorescences >3.0 cm long (rarely shorter in N. pedicellata and N.
tenuisepala); southern Colombia, Ecuador, and Peru.

6. Abaxial epidermis of sepals drying black, lightly pubescent throughout, epidermis visible
through any pubescence. 8. N tenuisepala

6. Abaxial epidermis of sepals drying black but concealed by dense golden or silvery
pubescence, all or most of the epidermis not visible through the pubescence.
7. Pedicels 14-27 mm long, pubescence drying white or silvery white; inflorescence
<6.5 cm long, 1-5-flowered. 5. N. pedicellata
1. Pedicels very short (flowers
subsessile) to 9.5 mm long; pubescence usually drying
golden-brown; 4-21 cm long, 4-31-flowered.
inflorescence
8. Staminodes usually 1/3 to 3/4 as long as the stamens; sepals and petals 8-23 mm
long, lanceolate to triangular (uncommonly ovate); leaves coriaceous or subcoria
ceous, glossy and smooth adaxially, rarely pubescent throughout abaxially.
1.N. grandiflora
2008 NEOSPRUCEA 23
8. Staminodes about as long as or longer than the stamens; sepals and petals 3-10
mm long, ovate to widely deltate; leaves subcoriaceous or chartaceous, glossy or
dull and often corrugated adaxially, usually pubescent throughout abaxially.
2. N. melastomatoides
Alternative Key to the Species of Neosprucea
1. Leaf base cordate or almost auriculate; leaf venation actinodromous (3-7-veined from the lamina
base, secondary veins extending ca. 2/3 or less of lamina length).
2. Leaves of mature plants elliptic to obovate, toothed at base to essentially entire; ovary and fruit
glabrous. 4. N. paterna
2. Leaves of mature plants ovate, serrate-crenate throughout; ovary and fruit densely tomentose to
sericeous. 6. N rimachii
1. Leaf base rounded, obtuse, acute, or attenuate; leaf venation acrodromous perfect (3-veined from
lamina base to apex; melastome-like).
3. Abaxial epidermis of sepals drying black, lightly pubescent throughout, epidermis visible through
any pubescence.
4. All tertiary veins usually perpendicular to the midvein; pedicels (5.5-) 8-21 mm long; higher
elevations, 1440-2300 m. 3. N. montana
4. to five upper pairs of tertiary veins acutely angled to the midvein;
Two pedicels 2-9 mm long;
lower elevations, 180-1300 (-1750) m. 8. N. tenuisepala
3. Abaxial epidermis of sepals drying black, but concealed by dense golden or silvery pubescence,
all or most of the epidermis not visible through the pubescence.
5. Pedicels >10 mm long.
6. Sepals and petals 10-15 mm long; stipules 0.6-0.8 mm long; leaves chartaceous to thinly
chartaceous; pulvinus at base of petiole absent or obscure; lower elevations, 80-780 m.
5. N. pedicellata
6. Sepals and petals 8-10 mm long; stipules 1.0-2.0 mm long; leaves chartaceous to coria
ceous; pulvinus usually present at both base and apex of petiole; higher elevations,
950-2300 m.
7. Filaments 1.3-1.7 mm long, shorter than the anthers (2.2-3.3 mm); inflorescence
usually a raceme; leaves 5-16.5 (-20.1) cm long. 3. N. montana (uncommon variant)

7. Filaments 2.2-2.5 mm long, longer than the anthers (2 mm); inflorescence usually a
panicle;leaves 11.5-28.1 cm long. 7. N. sararensis
5. Pedicels very short (flowers subsessile) to 10 mm long.
8. Inflorescences <4.5 cm long, usually congested and <2.0 cm long; stipules 1.5-2.5 mm
long; pulvini at base and apex of petiole usually present. 9. N. wilburiana

8. Inflorescences 4-21 cm long; stipules 0.7-1.1 mm long; pulvini at base and apex of peti
ole absent or obscure.
9. Staminodes 1/3 to 3/4 as long as the stamens; sepals and petals 8-23 mm long, lance
olate-triangular (uncommonly ovate); leaves coriaceous or subcoriaceous, glossy and

smooth adaxially, rarely pubescent throughout abaxially. 1. N grandiflora
9. Staminodes about as long as or longer than the stamens; sepals and petals 3-10 mm
long, ovate to widely deltate; leaves subcoriaceous or chartaceous, glossy or dull and
often corrugated adaxially, usually pubescent throughout abaxially. 2. N. melastomatoides
1. Neosprucea grandiflora (Spruce ex Bentham) Sleumer, Notizbl. Bot. Gart. Berlin

Dahlem 14:47.1938. Ba?ara grandiflora Spruce ex Bentham, J. Linn. Soc., Bot.
5 (Suppl. 2): 93. 1861. Hasseltia grandiflora (Spruce ex Bentham) Sleumer, No
tizbl. Bot. Gart. Berlin-Dahlem 11: 960. 1934. Spruceanthus grandiflorus
(Spruce ex Bentham) Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 13: 363.
1936.?Type: Peru. San Martin: among rocks along streams near Tarapoto [this
detail is given in the protologue by Bentham; labels indicate only "Prope Tara
poto; Peruviae orientalis"], [ca. 06?30'S, 76?22,W], 1855-1856 (fl, fr), R. Spruce
4897 (holotype: K!; isotypes: B-destroyed, BM! BR-photo! G! KW-photo! P!

W!).
Neosprucea sucumbiensis Cuatrecasas, Trop. Woods 101: 23. 1955.?TYPE: COLOM
BIA. Putumayo: selva higr?fila del R?o San Miguel o Sucumb?os, en el afluente
izquierda Quebrada de la Hormiga, [ca. 00?20'N, 76?50'W], 290 m, Dec 1940,
/. Cuatrecasas 11147 (holotype: US!; isotypes: COL! L-fragment!).
Trees, small to large, to 30 m tall. Trunk to 60 cm dbh; bark whitish, smooth.

Branches green when young, drying maroon-brown to black, but becoming gray with
growth, obscurely 3^1-sided, becoming terete with growth, sparsely (to moderately)
strigillose; lenticels rare to infrequent on youngest growth, light brown, elliptic to linear.
Leaf blades 5.3-34.5 cm long, 2.3-13.9 cm wide, narrowly elliptic, elliptic, or oblong
elliptic, infrequently obovate, infrequently slightly asymmetrical with respect to the mid
vein; apex acuminate to abruptly acuminate for up to 3.0 cm; base acute to rounded,
eglandular; margin serrate-crenate, rarely obscurely so, 0.3-1 tooth/cm, teeth obtuse (con
vex-convex to convex-retroflexed) with a glabrous gland at the apex, gland abaxial and
abutting the leaf margin or marginal; subcoriaceous to coriaceous; adaxial surface glossy,
glabrous except near lamina-petiole junction; abaxial surface dull green, glabrous or
sparsely pubescent along the major veins, rarely soft pubescent throughout; venation acro
dromous perfect, 3-veined from base to apex, a distal 1-2 (-3) secondary pairs (or sec
ondary veins per side of the lamina) arising at an acute angle from midvein near apex,
prominent tertiaries 8-41, usually perpendicular to the midvein and the pair of secon
daries, semicraspedodromous, veins raised abaxially. Petioles 0.8-2.9 cm long, 0.6-2.0
mm in diameter, sparsely strigillose or infrequently densely pubescent, canaliculate on
adaxial surface with a small or obscure pulvinus at apex. Stipules 0.7-1.1 mm long,
0.3-0.4 mm wide, entire, caducous. Inflorescences termi
triangular, densely pubescent,
nal, infrequently subterminal or axillary, racemes or raceme-like panicles with 4-16 (-31)
flowers, sometimes reduced to fewer flowers due to abscission, 5.2-12.7 (-21.0) cm long
at anthesis or in fruit, axis 0.8-3.8 mm in diameter, sometimes flared or expanded at the
nodes of the triads (the ultimate cymose node beneath the flowers), densely pubescent;
bracts 3 mm long, 1.8 mm wide, triangular, densely pubescent abaxially, moderately to
densely pubescent adaxially, caducous; bracteoles 3.6 mm long, 1.3 mm wide, lanceolate,
located <2.5 mm below the articulation, densely pubescent, caducous; pedicels very short
(flowers subsessile) or up to 9.5 mm long, densely pubescent, golden-brown when dry,
and sometimes ridged. Flower buds ellipsoidal to ovoid, capped by ridges where the
sepals abut against each other, light green, silvery green, creamy green, yellowish green,
brownish green, pinkish green, or brown, densely pubescent. Sepals 4 (-5), 7.8-20.2 mm
long, 3.4-8.3 mm wide, up to 0.8 mm thick medially, up to 1.2 mm thick at the apex,
lanceolate, narrowly triangular, triangular, or ovate, abaxially brownish green to green,
drying darker than the petals, venation of ca. 10 veins sometimes evident, densely short
pubescent with golden-brown hairs, adaxially lighter green, densely short-pubescent, the
pubescence sometimes easily rubbed off. Petals 4 (-5), 7.8-22.7 mm long, 3.4-8.3 mm
wide, lanceolate, narrowly triangular, triangular, or uncommonly ovate, not thickened,
creamy white to white, occasionally infused with pink, moderately, densely, or unevenly
pubescent with hairs longer than those of the calyx, sericeous. Receptacle densely pubes
cent. Androecium of numerous stamens, ca. (60-) 140-400; filaments 0.9-1.9 mm long,
densely hirsute; anthers 3.3-7.6 mm long, hirsute, especially on the adaxial surface of the
connective. Staminodes filamentous, numerous, flattened, sometimes hirsute near the
2008 NEOSPRUCEA 25
base, infrequently distally, usually 1/3 to 3/4 as long as the stamens, rarely as long as the
stamens. Ovary ca. 2 mm in diameter, spherical, light green, glabrous to infrequently
sparsely but entirely hirsute; placentation highly intruded parietal, placentae 4-6; style
3.8-11.2 mm long, glabrous; stigma minutely lobed or with several small terete projec
tions. Fruit a fleshy berry, ca. 1 cm in diameter, widely ovoid, yellowish green or green.
Seeds ca. 15-28, 3.5 mm long, 2.2 mm wide, ellipsoidal. Fig. 8.
Common Names. Huaorani: "ague" {Espinoza & Coba 394), "damatahue" {Gudi?o
et al. 1035). Quichua (Quechua): "guataracu muyo" {Korning & Thomsen 47468), "pu
raqui paju" {Gudi?o et al. 1853), "sapote" {Gudi?o et al. 1825), "sara muyo," "sara
muyo," or "sara muyu" {Gudi?o 1683, Tipaz et al. 434, Neill & Maries 7010), "uraqui
paju yura" {Gudi?o et al. 1754), "yaeu payatzig" {Alvarado 271).
Phenology. Collected in flower and fruit throughout the year but more frequently
from July to January, approximately corresponding to the drier months of the year (Pour
rut et al. 1995; Neill & J0rgensen 1999).
Distribution (Fig. 9). Western Amazon basin in Colombia, Ecuador, and Peru; low
land rainforest (bosque h?medo tropical, bosque muy h?medo tropical) and lower mon
tane rainforest(bosque h?medo premontano, bosque muy h?medo premontano, bosque
pluvial premontano); 100-1200 (-1850) m.
Additional Specimens Examined. Colombia. Caqueta: Florencia, [01?37'N, 75?37'W], P?rez-Ar

bel?ez 752 (COL, US); Florencia, carretera Florencia-Suaza, Km 20, Vereda "Villar?s," 01?36'N, 75?37'W,
Ram?rez et al. 5613 (MO); Morelia, [01?29.3'N, 75?43.5'W], von Sneidern A1350 (S). Ecuador. Morona
Santiago: Parroquia El Copal, San Jos? Grande, cuenca del R?o Paute entre el R?o Cardenillo Grande y Car
denilloChico, 02?36'S, 78?26'W, Cer?n & Benavides 14114 (MO); Cumand?, southern side of R?o Pastaza, ca.
4 km W of Mera, [01?25'S, 78?10'W], Lugo 343 (AAU, MO, NY); Morona, Cordillera del Cutuc?, Asociaci?n
Shuar Sevilla, Comunidad Angel Ruby, pie del Cerro Muchin, 02?21'39"S, 78?04'08"W, Suin et al. 1973 (BH,
MO); Pozo petrolero "Garza" de Tenneco, 35 km (aprox.) al NE de Montalvo, 01?49'S, 76?42'W, Zak & Es
pinoza 4463 (AAU, MO, NY, QCNE), Zak & Espinoza 4688 (AAU, K, MO, NY, QCNE, TEX), Zak & Espinoza
4747 (AAU, MO, NY, QCNE); Bomboiza, Misi?n Salesiana-Shuar y alrededores, 03?25'S, 78?35'W, Zaruma &
Arguello 511 (AAU, MO, NY, QCA, QCNE).??APO: Carretera Holl?n-Loreto, Km 40-50, alrededores de la
comunidad Huaman? [=Guaman?] y del R?o Pucuno, al margen W del R?o Holl?n, 00?43'S, 77?36/W, Hurtado
653 (AAU, MO, NY, QCNE), Hurtado 683
(AAU, G, MO, NY, QCNE), Hurtado 853 (AAU, G, MO, NY,
QCNE); carretera Holl?n-Loreto, 5 km alW
de Guaman?, faldas del Volc?n Sumaco, 00?43'S, 77?38'W, Neill et
al. 8588 (AAU, K, MO, NY, QCNE); R?o ?apo, 2 km r?o abajo de Campana Cocha, boca del R?o Huambuno,
OrOl'S, 77?30'W, Neill & Maries 7010 (AAU, MO, NY, QCNE); R?o Arajuno, Sola Cocha, 01?07'S, 77?36'W,
Palacios et al. 871 (AAU, IBE, K, MO, NY, QCA, QCNE, US); Archidona, faldas al sur del Volc?n Sumaco,
carretera Holl?n-Loreto, Km 50, Comuna Huahua [=Guagua] Sumaco, 00?44'S, 77?34'W, Alvarado 271 (AAU,
G, MO, QCNE); Archidona, carretera Holl?n-Loreto, Km 50, Comunidad Guagua Sumaco, faldas al sur del Vol
c?n Sumaco, 00?38'S, 77?27'W, Cer?n & Hurtado 6677 (MO); Archidona, Reserva Ecol?gica Antisana, comu
nidad Shamato, entrada por Km 21-Shamato, Plot #1, 00?43,S, 77?49'W, Clark et al. 5193 (BH, MO); Archi
dona, Parque Nacional Sumaco Napo-Galeras, faldas de Galeras, Bloque 19, l?nea s?smica 30, Compa??a Triton,
00?53'S, 77?33'W, Freir? & Cerda 373 (MO); Archidona, southern slopes of Volc?n Sumaco, carretera Hol
l?n-Loreto, centro
Challuayacu, permanent plot number 06, on the property of Pedro Alvarado, 00?43,S,
77?40'W, Pitman & Alvarado 95 (MO); Tena, Estaci?n Biol?gica Jatun Sacha, ca. 8 km E of Misahuall? on the
5 side of the R?o ?apo, 01?03.92,S, 77?36.95'W, Alford 3103 (AAU, BH, BM, DUKE, K, MO, QCA, QCNE,
US), Gentry et al. 60179 (MO), Miller et al. 2459 (MO, QCNE), Palacios 1834 (MO), Palacios 3274 (AAU, K,
MO, NY, QCNE, TEX, US), Palacios 4792 (AAU, F, G, MO, NY, QCNE).?Orellana: Archidona, carretera
Holl?n-Loreto, entre Avila y Loreto, Huiruno, comunidad Quichua, 00?43'S, 77?19'W, Cer?n 7693 (AAU, MO,
QCNE), Cer?n 7770 (MO, QCNE); Loreto, faldas del Volc?n Sumaco, al oeste de Avila Viejo, Bloque 19, l?nea
s?smica 8, Compa??a Triton, 00?38'12"S, 77?27'12"W, Freir? & Cerda 1 (MO, QCNE); Loreto, al pie del Vol
c?n Sumaco, al lado del R?o Suno, 00?40'S, 77?19,W, Hurtado et al. 2753 (AAU, MO, QCNE); Loreto, a 2.5
km al N de la carretera a Campo Alegre, carretera Holl?n-Loreto-Coca, 19, l?nea s?smica, Helipuerto 3,
Bloque
Compa??a Triton, 00?34'S, 77?20'W, Vargas & Cerda 648 (MO, QCNE); Loreto, 2 km al oeste del R?o
FIG. 8. Neosprucea grandiflora. A. Flowering branch, showing variability of leaf size on a single branch,
detail of a torus-shaped gland at the leaf tooth (upper left), and detail of a stipule (lower right). B. Abaxial sur
face of leaf. C. Flower bud, showing valvate sepal aestivation and thick sepals. D. Flower at anthesis, showing
thickness of the sepals and the subtle outer whorl of staminodes. E. Flower at early anthesis, longitudinal sec
tion, with detail of the stigma. F. Staminodes. G. Stamens, showing variability in size from those of the inner
whorls (left) to those of the outer whorls (right), with detail of apical pores. (Based on Alford 3103.)
2008 NEOSPRUCEA 27
W 80? 75? 70?
FIG. 9. Distribution of Neosprucea grandiflora.
Tutapishco, Bloque 19, l?nea s?smica 4, Helipuerto 3, Compa??a Triton, 00?36'S, 77?22'W, Vargas & Cerda 681
(MO, QCNE); Loreto,sector del R?o Suno, Bloque 19, l?nea s?smica 2, cruce con l?nea 1, Helipuerto 1, Com
pa??a Triton, 00?33'S, 77?24/W, Vargas & Cerda 708 (MO, QCNE); Loreto, 500 m al S de la carretera a Campo
Alegre, carretera Holl?n-Loreto-Coca, Bloque 19, l?nea s?smica 4, Helipuerto 4, Compa??a Triton, 00?36'S,
77?20/W, Vargas et al. 588 (MO); Parque Nacional Yasun?, carretera y oleoducto de Maxus en construcci?n, Km
46-52, 00?47'S, 76?30/W, Aulestia et al. 460 (MO, QCNE), Aulestia et al. 454 (MO, QCNE); Parque Nacional
Yasun?, carretera y oleoducto de Maxus en construcci?n, Km 54-58, 00?48'S, 76?30'W, Aulestia & Andi 795
(MO, QCNE); Parque Nacional Yasun?, carretera y oleoducto de Maxus en construcci?n, Km 42, 00?45'S,
76?30'W, Aulestia & Grefa 162 (MO, QCA, QCNE); Parque Nacional Yasun?, carretera y oleoducto de Maxus
en construcci?n, Km 46 al Pozo Capiron, 00?41'S, 76?29'W, Dik 300 (MO, QCNE); Parque Nacional Yasun?,
pozo petrolero Daimi I, Conoco, 01?02'S, 76?10'W, Hurtado et al. 5 (AAU, G, MO, NY, QCA, QCNE); Parque
Nacional Yasun?, plano inundable Km 84.7 carretera Pompeya-Iro, parcela 21, 00?51.859'S, 76?16.723'W,
Macla et al. 1263 (MA); Parque Nacional Yasun?, tierra firme Km 34.6 carretera Pompeya-Iro, parcela 5,
00?37.567,S, 76?27.657/W, Mac?a et al. 3828 (MA); Parque Nacional Yasun?, carretera de Maxus en construc
ci?n, Km 8-10, 00?29'S, 76?34'W, Zuleta 145 (MO, NY); Zona de amortiguamiento del Parque Nacional
Sumaco, l?neas s?smicas de la compa??a Amoco, R?o Paushiyacu, 00?21'S, 77?19'W, Tirado 1914 (MO); La Joya
de los Sachas, Parroquia Pompeya, campamento de Maxus, carretera Maxus Km 1-4, 00?25'S, 76?36/W, Gri
jalva et al. 623 (AAU-2, MO, NY, QCNE); La Joya de los Sachas, Pompeya, carretera Maxus Km 3.9-5.2,
00?25'S, 76?37'W, Gudi?o et al. 1853 (AAU, MO, NY-2, QCA, QCNE), Gudi?o et al. 2194 (AAU, MO, NY,
QCNE), 1683 (MO, QCNE), Gudi?o et al. '1754 (MO, QCNE), Gudi?o et al. 1825 (AAU [3], MO, NY,
Gudi?o
QCNE); La Joya de los Sachas, Pompeya, carretera Maxus Km 3-3.5, 00?25'S, 76?37'W, Gudi?o & Andi 2090
(MO, QCA, QCNE); Parque Nacional Yasun?, a 6 km E del Helipuerto, pasando cuatro veces el R?o Shiripuno,
01?01'S, 77?05'W, Hurtado et al. 535 (AAU, K, MO, NY, QCNE); A?angu, NW c?rner of the Parque Nacional
Yasun?, undisturbed forest in the surroundings of the SEF terra firme line and KTH hectare plot, at SEF point
196, 00?33,S, 76?22'W, Korning & Thomsen 47468 (AAU); 20 km alW de Coca, al sur del R?o ?apo, por V?a
de los Zorros, 00?35'S, 77*03^, Neill et al. 6384 (AAU, F, MO, NY, QCA); A?angu, NW c?rner of the Parque
Nacional Yasun?, along SEF line in terra firme forest, near Tiputini trail, at SEF point 57,00?32'S, 76?22'W, 011
gaard et al. 57040 (AAU, MO, NY, QCA, U); Coca (Puerto Francisco de Orellana), trail along Rio Payamino,
[00?27'S, 77?00'W], Harling & Andersson 11911 (GB); Estaci?n Experimental INIAP-Payamino, 5 km al N de
Coca, Reserva El Chuncho, Inventario Forestal JICA-MAG (Misi?n Japonesa), 00?25'S, 77?00/W, Palacios
1089 (MO); SW0077, ca. 30 km NNW of Coca, R?o Huashito, site of Proyecto Huashito, [00?20'S, 77?04'W],
Pennington 10620 (QCA, QCNE, U, US), Pennington 10626 (NY, QCA, QCNE, U); Aguarico, Reserva ?tnica
Huaorani, Maxus road and pipeline construction project, Km 99-100, 00?56'S, 76?13'W, Pitman 481 (MO,
QCNE); Estaci?n Cient?fca Yasun?, R?o Tiputini, al noroeste de la confluencia con el R. Tivacuno, 6 km este de
la carretera Maxus, Km 44, desvio hacia el pozo Tivacuno, parcela de 50 ha, 00?38'S, 76?30'W, Romoleroux et
al. 2259 (AAU, F [photo], MO); Estaci?n Experimental INIAP-San Carlos, Reserva Flor?stica El Ahuano,
00?19'S, 76?50'W, Zaruma 583 (AAU, K, MO, NY, QCA, QCNE).?PASTAZA: Madre Tierra, ca. 12 km S of
Puyo, [01?33/S, 78?02'W], Lugo 328 (F, MO, QCA, US); 0.4 km E on new road to R?o Bobonaza, leaving
Puyo-Macas road at Km 15, 01?35'S, 77?53'W, Stein 3013 (K, MO, NY, QCNE); Arajuno, campamentos tem
porales 9, 22, y 25, l?nea propuesta del oleoducto Villano-CPF por Arco, Km 25 noroeste del pozo Villano 2,
01?27'S, 77?36'W, Freir? & Santi 3288 (MO, QCNE); Mera, town of Shell, small patch of forest on northern
edge of Shell, 01?29'14"S, 78?03'39"W, Clark & Mailloux 7852 (US); Pastaza, Villano, Pandanuque, encima de
colina al sur del pozo petrolero Villano 2 de Arco, 01?28'S, 77?27'W, Alvarez et al. 2379 (MO, NY, QCNE);
Pastaza, pozo petrolero "Danta 2" de Unocal, 50 km al SSE de Curaray, 01?47'S, 76?48'W, Espinoza & Coba
394 (MO, NY, QCA, QCNE); Pastaza, carretera Puyo-Canelos, 01?30'S, 77?55'W, Gudi?o 451 (AAU, MO, NY,
QCNE); Pastaza,
pozo petrolero Moretecocha de Arco, 75 km al E de Puyo, 01?34/S, 77?25/W, Gudi?o et al.
1035 (MO, QCA, QCNE); Pastaza, pozo petrolero Moretecocha de Arco, R?o Landayacu, 75 km al E de Puyo,
01?34'S, 77?25'W, Gudi?o 1194 (AAU [2], MO-2, QCA, QCNE); Pastaza, pozo Villano 2 de Arco, 2 km del
pueblo de Villano, 01?25'S, 77?20'W, Tipaz et al. 434 (MO); Pastaza, pozo petrolero Ram?rez, 20 km al sur de
la poblaci?n de Curaray, 01?32'S, 76?51'W, Zak & Espinoza 5024 (MO, NY, QCNE), Zak & Espinoza 4963
(AAU-2, MO, QCNE).?Sucumb?os: San Pablo de los Secoyas, cleared primary forest with cattle grazing just
opposite the village, 00?15'S, 76?21'W, Brandbyge et al. 33440 (AAU, MO, NY, U, US); Lago Agrio, Parroquia
Dureno, bosque primario en la Reserva Ind?gena Cof?n-Dureno, 00?02'S, 76?42/W, Cer?n & Cer?n 3151 (AAU,
MO); Dureno, Reserva Forestal de Communidad Cofan de Dureno, R?o Aguarico, transect 4, [00?03/N,
76?40'W], Gentry et al. 64365 (AAU, GH, MO); Limoncocha, 00?24'S, 76?38'W, Holm-Nielsen 39836 (AAU);
El Reventador, colecciones en ambas m?rgenes del R?o Reventador, [0.15?S, 77.38?W], Jaramillo & Grijalva
12903 (MO, NY); Lago Agrio, Reserva Cuyabeno, orilla del R?o Aguarico, comunidad ind?gena cof?n del
Z?balo, 75?45'W, Palacios
00?22'S, et al. 9504 (AAU, MO, NY, QCNE); Shushufindi, Parroquia Pa?acocha,
Sector Pucape?a, pozo exploratorio Tongoy 2, petrolera Occidental, 00?20'S, 76?23'W, Freir? et al. 2775 (MO,
QCNE); Aguarico, Chiro Isla community, north bank of R?o ?apo ca. 195 km downstream from Puerto Fran
cisco de Orellana (Coca), 00?36'S, 75?52'W, Bensman 308 (MO, QCNE); Rio Aguarico, ca. 5 km S of Lago
Agrio, [00?05'N, 76?50'W], Lugo S. 3209 (GB); Aguarico, Reserva Faun?stica Cuyabeno, R?o Aguarico, Zan
cudo, detr?s del campamento militar, 00?29'S, 75?32'W, Palacios et al. 7669 (F, MO, QCNE).?ZAMORA
Chinchipe: Area of the Estaci?n
Cient?fica San Francisco, road Loja-Zamora, ca. 35 km from Loja, 03?58'S,
79?04'W, Homeier et al. 52 (MO), Homeier 257 (MO). Peru. Without locality, 1827, Ruiz & Pav?n s.n. (P).?
Amazonas: Bagua: Soldado Oliva, carretera entre Bagua-Imaza, [05?18'14"S, 78?23,17"W], D?az et al. 10605
(BH).?Loreto: Pampas de Ponasa [=Panaza, Ponaza; near border between Loreto and San Mart?n], [07?00'S,
76?03'W], Ule 6712 (G, K); Maynas, Quebrada Yanomono, Explorama tourist camp, above mouth of R?o ?apo
2008 NEOSPRUCEA 29
on R?o Amazonas, [03?27'S, 72?54'W], Gentry et al. 27848 (MO); Maynas, Yanamono (Explorama lodge), 50
mi NE de Iquitos, 03?30'S, 72?50'W, V?squez & Jaramillo 2742 (MO); Maynas, Santa Mar?a de Nanay, Que
brada Yarina, 03?55'S, 73?40'W, V?squez et al. 12176 (AMAZ, MO).
Neosprucea grandiflora is the type species of the genus and is part of a three
species complex that includes N. melastomatoides and M tenuisepala. Each of these
species is distributed principally in the lowlands of Ecuador and northern Peru, and they
share the characteristics of acrodromous (melastome-like) venation, filamentous sta
minodes, small stipules, short pedicels, hirsute anthers, and lack of (or obscure) petio
lar pulvini. These species differ from each other in leaf shape, leaf texture, sepal pu
bescence, sepal thickness, petal length, staminode length, and thickness of the
inflorescence axis.
Neosprucea grandiflora has coriaceous to subcoriaceous elliptic to narrowly elliptic

leaves, densely pubescent and thick sepals (up to 1.2 mm thick), long petals (8-23 mm
long), small tomedium-sized staminodes (usually 1/3 to 3/4 length of the stamens, some
times obscure in the pubescence of the receptacle), and thick (0.8-3.8 mm in diameter),
long (5.2-21.0 cm), and usually many-(4-31)-flowered inflorescences (Fig. 2F).
Neosprucea melastomatoides differs from N. grandiflora by having thick but chartaceous
or subcoriaceous, often corrugated elliptic leaves (Fig. 2A), thinner sepals (up to 0.5 mm,
but usually less than 0.2 mm thick), smaller petals (3-10 mm long), and longer stamin
odes (about as long as the stamens) (Fig. 2E). Neosprucea tenuisepala has thinly charta
ceous (ovate to) elliptic, narrowly elliptic, or obovate leaves, fewer flowers (1-7), and
sepals which are thin and scarcely pubescent. When dry, the epidermis of the sepals is
black and is not obscured by the pubescence.
The type specimen of Neosprucea grandiflora is from the geographical and altitudi
nal periphery of the species. It has flowers that are smaller, leaves that are less coriaceous,
and pedicels that are thinner and longer than what one finds most commonly for the
species. Cuatrecasas recognized the more common morphotype as a different species,
N. sucumbiensis, but there is a continuum between the type collection of Spruce and the
type collection of Cuatrecasas. Unfortunately, the type of N. sucumbiensis also has leaves
that are softly pubescent throughout abaxially, and many collectors have utilized the
salient feature of leaf pubescence from Cuatrecasas's diagnosis to identify every specimen
of Neosprucea with pubescent leaves as N. sucumbiensis. Neosprucea grandiflora rarely
has thoroughly pubescent leaves abaxially, but the type of N. sucumbiensis happens to be
one of those exceptions; other cases include Gentry et al. 64365 and Cer?n & Cer?n 3151.
Most individuals with thoroughly pubescent leaves belong to another species, N. melas
tomatoides, which was overlooked as a separate taxon, because Cuatrecasas's name was
misapplied to it.
One might argue that this overlap in leaf pubescence and variation in the sizes of
flower and fruit parts is evidence thatN. grandiflora and N. melastomatoides are conspe
cific; however, where they co-occur, there are no intermediates (e.g., at Jatun Sacha Bio
logical Station near Tena, Ecuador). Furthermore, based on one sample from each species,
the wood anatomy differs
(Fig. 1).Neosprucea grandiflora has a long and wide series of
procumbent ray parenchyma (1-120 cells long, 1-3-seriate), while N. melastomatoides
has shorter and narrower series of procumbent ray parenchyma (1-21 cells long, 1-2
seriate).
The flowers ofN. grandiflora were reported once as fragrant (Palacios 4792).
2. Neosprucea melastomatoides M. H. Alford, sp. nov.?Type: Ecuador. Napo:

Estaci?n Biol?gica Jatun Sacha, R?o Napo, 8 km r?o abajo de Misahuall?, margen
derecha, bosque primario, suelos rojos de colinas, 450 m, 01?08'S, 77?30,W, 2
Oct 1986 (fl, fr), W. Palacios 1390 (holotype: US!; isotypes: AAU! K! MO-2!
NY-2! QCA-2! QCNE!).
Arbor parva. Folia elliptica, subtus molliter pubescentia, breviter acuminata, ad

basim subrotundata, margine inconspicue glanduloso-serrato, venatione acrodroma. Inflo
rescentia axillaris vel terminalis, racemus spiciformis vel panicula racemiformis, pedicel
lis <0.9 cm longis sub anthesi; s?pala 4-5, viridi-albida, crassa, ovata, utrinque dense
tomentoso-sericea; p?tala 4-5, albida, forma et indumento sepalis simillima; stamina cre
berrima filamentis capillaribus hirtis 0.5-1.2 mm longis, antheris anguste linearibus 2-3.4
mm longis graciliter hirtis; staminodia angustissime capillaria; receptaculum intra stamina
valde hirsuto-setosum. Ex affinitate Neospruceae grandiflorae Sleumer et specierum
affinium staminodiis longioribus, foliis subtus ubique molliter pubescentibus, et floribus
minoribus distinguenda.
Trees to 10m tall. Trunk to at least 6 cm dbh; bark gray, smooth. Branches light green
when young, drying maroon-brown, becoming gray with growth, obscurely 3^1-sided, be
coming terete with growth, moderately pubescent to hirsute; lenticels few to numerous,
maroon-brown, elliptic. Leaf blades (7.0-) 11.0-32.5 cm long, (3.5-) 5.1-14.8 cm wide,
(ovate-) elliptic, infrequently somewhat asymmetrical with respect to the midvein; apex
abruptly acuminate for ca. 0.8-4.0 cm; base rounded to obtuse, eglandular; margin serrate
crenate, sometimes obscurely so, 0.3-1 tooth/cm, teeth obtuse (convex-convex to convex
retroflexed) with a glabrous gland at or near the apex, gland abaxial and abutting the leaf
margin or marginal; chartaceous or subcoriaceous; adaxial surface dull to glossy, often
corrugated in appearance (with lamina between the major tertiary veins forming ridges
above the grooves of the veins), glabrous except near lamina-petiole junction; abaxial sur
face softly pubescent throughout or rarely only on the primary, secondary, and tertiary
veins; venation acrodromous, 3-veined from base to apex, occasionally a single upper sec
ondary pair (or one secondary vein per side of the lamina) arising at an acute angle from
themidvein near apex, prominent tertiaries (13-) 20-34, the first and last several tertiaries
more closely spaced and less prominent than those in the middle, perpendicular to the
midvein and secondaries, veins raised abaxially. Petioles 0.5-2.0 cm long, 1.7-3.6 mm in
diameter, (moderately-) densely and softly pubescent, flat adaxially, infrequently canalic
ulate near apex; pulvinus absent or obscure, as entire petiole is quite thick. Stipules 1.1
mm 0.4-0.7 mm wide, entire, caducous. Inflores
long, triangular, densely pubescent,
cences or sub-terminal racemes or raceme-like with 5-21 flowers, some
axillary panicles
times fewer due to abortion or abscission, 4.0-14 cm long, axis 1.0-2.4 mm in diameter,
moderately to densely pubescent; bracts 2.4-3.1 mm long, 0.6-0.9 mm wide, lanceolate,
abaxially moderately to densely pubescent, adaxially glabrous or sparsely pubescent, ca
ducous; bracteoles 1.4-1.5 mm long, 0.4 mm wide, narrowly triangular, located just below
(<1 mm) the articulation, abaxially densely pubescent, adaxially glabrous or pubescent
along themargins, caducous; pedicels 2.4-9.0 mm long, densely pubescent, golden-brown
when dry, sometimes ridged. Flower buds spheroidal to ovoid, pale green, densely pubes
cent. Sepals 4-5, 3.1-9.8 mm long, 3.1-6.2 mm wide, up to 0.5 mm thick, usually less
than 0.2 mm thick, ovate to widely deltate, abaxially green, tomentose, adaxially pale
green, tomentose with hairs. Petals 4-5, 3.2-7.4 mm 2.1-3.7 mm
golden-brown long,
wide, ovate to widely deltate, white to creamy white, abaxially short-tomentose, adaxially
2008 NEOSPRUCEA 31
FIG. 10. Neosprucea melastomatoides. A. Leafy branch, with details showing the pubescent abaxial sur
face of the leaves and a torus-shaped gland at the leaf tooth. B. Flowering branch. C. Rower bud, showing val
vate aestivation and thin sepals. D. Flower, view from above and from the side, showing the stamens appressed
to the gynoecium and the staminodes reflexed with the perianth. E. Flower, longitudinal section. F. Staminodes.
G. Stamens. (Based on: A, Alford 3092; B-G, Alford 3091.)
short-tomentose, trichomes sometimes in lines. Receptacle densely pubescent. Androe

cium of numerous stamens, ca. 120-180; filaments 0.5-1.2 mm long, hirsute; anthers
2.0-3.4 mm long, hirsute on the adaxial side of the connective. Staminodes filamentous,
numerous, flattened, about as long as the stamens or slightly longer. Ovary spherical or
widely ovoid; placentation highly intruded parietal, placentae ca. 6; style 2.0-3.5 mm
long, pubescent to hirsute, especially near the base; stigma minutely lobed. Fruit a juicy
berry, ca. 1-1.2 cm in diameter, spherical to widely ovoid, green, moderately hirsute, es
pecially near the apex. Seeds ca. 20, 1.9-2.5 mm long, 1.0-1.5 mm wide, ellipsoid, but
angled by compression of the fruit wall and other seeds. Fig. 10.
Common Name. Quichua (Quechua): "tuclla caspi" {Delinks et al. 284).
Phenology. Collected in flower and fruit principally between mid-August and mid
February, approximately (Pourrut et
corresponding to the months with least precipitation
al. 1995; Neill & J0rgensen 1999). Buds, however, have been collected in the intervening
months of March and April, possibly indicating only a seasonal reduction in flowering.
Distribution (Fig. 11). Western Amazon basin of Napo, Orellana, Pastaza, and Su
cumbios provinces of Ecuador; lowland tropical rainforest (bosque muy h?medo tropical,
bosque h?medo tropical, bosque pluvial premontano); 250-850 m.
82? 76?W
PACIFIC
OCEAN
Neosprucea
melastomatoides
FIG. 11. Distribution o? Neosprucea melastomatoides.
2008 NEOSPRUCEA 33
Additional Specimens Examined. Ecuador. ?apo: Campana Cocha, Comunidad Quichua, margen
izquierda del R?o ?apo, aguas abajo, 01?05'S, 77?30'W, Jaramillo & Coello 3743 (AAU, MO, QCA); R?o ?apo
"Hotel Jaguar" [ca. 2 hours downstream from Misahuall?], [00?57'S, 77?25'W], Jaramillo 4052 (AAU); R?o
Sumino, tributary of the R?o ?apo, ca. 5 km northeast of Santa Rosa, [00?55'S, 77?25'W], Lugo S. 2081 (GB);
Rio Arajuno, Sola Cocha, 01?07'S, 77?36'W, Palacios et al. 844 (AAU, K, MO, NY, QCA, QCNE, US); Tena,
Estaci?n Biol?gica Jatun Sacha, ca. 8 km E of Misahuall? on the south side of the Rio Napo, 01?03.92'S,
77?36.95'W, Alford et al. 3091 (AAU, BH, DUKE, K, MO, QCA, QCNE), Alford 3092 (BH, DUKE, MO, QCA,
QCNE, US), Cer?n & Cer?n 4650 (MO, QCNE), Cer?n & Cer?n 4552 (AAU, MO, QCNE), Cer?n 587 (K,
MO, NY, QCA, QCNE, US), Cer?n & Iguago 5347 (MO, QCNE, US), Cer?n 3590 (K, MO, NY, QCA, QCNE),
Cer?n & Montesdeoca 8008 (MO-2, QCNE), Cer?n & Iguago 5384 (MO, QCNE), Cer?n 642 (K, MO, NY,
QCA, QCNE), Palacios 4163 (AAU, MO, QCNE), Palacios 2980 (AAU, K, MO, NY, QCNE, US); Tena,
Cabanas Chuva Urcu, 01?08'32"S, 77?35'29" W, Delinks et al. 284 (MO).?Orellana: v?a Payamino-Loreto,
4-6 km del r?o, 00?26'S, 77?02/W, Zaruma 730 (MO, NY, QCNE); Loreto, faldas del Volc?n Sumaco, al oeste
de Avila Viejo, Bloque 19, l?nea s?smica 8, Compa??a Triton, 00?37'53"S, 77?27/46,/W, Freir? & Cerda 131
(MO, QCA, QCNE); Loreto, Parroquia San Vicente de Huaticocha, Comunidad Santa Rosa de Arapino, Bloque
#19 Triton, Pozo Santa Rosa, 00?58'S, 77?4(W [more accurately as 00?52'S, 77?28'W?], Freir? et al. 2312
(MO).?Napo-Pastaza border: between Tena and Napo, [01?01'S, 77?48'W], Asplund 9054 (LL, NY, QCA,
S).?Pastaza: about 4 km NW of Canelos, 01?33'S, 77?45'W, St?hl et al. 4338 (AAU).?SUCUMB?OS: 2 km
downstream from Punto Aguarico, 00?05'N, 76?59'W, Brandbyge et al. 30477 (AAU).
Neosprucea melastomatoides is closely related toN. grandiflora (see that species, no.
1) but differs by having large, chartaceous to subcoriaceous, often corrugated leaves, thin
ner sepals (< 0.5 mm thick and usually < 0.2 mm), shorter petals (3-8 mm), and
longer
staminodes (as long as or longer than the stamens). It is the species most commonly found
with softly pubescent leaves; rarely that feature may be found also inN. grandiflora or N.
tenuisepala. The name N. sucumbiensis has been misapplied toN. melastomatoides.
3. Neosprucea montana Cuatrecasas, Trop. Woods 101: 20. 1955.?Type: Colombia.

Norte de Santander: Sarare, Quebrada del Sararito, 1930-1950 m, 22 Oct 1941
(bud, fl, young fr), J. Cuatrecasas et al. 12581 (holotype: F!; isotypes: COL-2!
GH?L-fragment!US?).
Trees to 16m tall. Trunk to 27 cm dbh, forming small buttresses with age; bark ma
roon-brown when young, tight, older bark light brown, tight and smooth. Branches cop
pery green when young, drying to chestnut- or maroon-brown, distinctly 4-sided, strig
illose, glabrescent with age, especially pubescent at petiole base and on buds; lenticels
common, light pinkish brown, circular to narrowly elliptic, elongated by the splitting of
the bark. Leaf blades (5.0-) 7.0-16.5 (-20.1) cm long, (2.5-) 3.7-9.0 cm wide, (ovate to)
elliptic (to narrowly elliptic), symmetrical; apex acute to abruptly acuminate for 0.5-2 cm;
base acute or obtuse, serrate-crenate, 0.4-1.2 teeth ob
eglandular; margin (0.3-) teeth/cm,
tuse or rounded with a glabrous gland near the apex, gland abaxial, slightly inset from the
margin at maturity; chartaceous; adaxial surface dark green, glabrous except near petiole
junction; abaxial surface light green, strigillose to glabrescent on the veins and glabrous
elsewhere to softly but moderately pubescent throughout; venation acrodromous, 3
veined from the base to the apex, rarely a single upper secondary pair (or a single sec
ondary vein per side of the lamina) arising at an acute angle from midvein near apex,
prominent tertiaries 12-23, perpendicular to the midvein and the two secondaries, veins
raised abaxially, sometimes winged between the midvein and two secondaries producing
small domatia. Petioles 0.55-3.2 cm long, 0.6-1.6 mm wide, densely strigillose to
glabrescent, canaliculate adaxially; solid pulvini at the apex and base prominent and
well developed. Stipules 1.0-2.0 mm long, 0.4 mm wide, lanceolate, entire, strigillose,
caducous. Inflorescences axillary or subterminal, few-flowered racemes (or rarely pani

cles) with 3-8 flowers, sometimes fewer due to abortion or abscission, 3.2-11.0 cm long
at anthesis or in fruit, axis 0.7-1.4 mm thick, usually sparsely tomoderately pubescent or
strigillose (rarely densely pubescent); bracts 1mm long, 0.2 mm wide, lanceolate, densely
strigose, caducous; bracteoles (and bracteole scars) inconspicuous, ca. 1mm long, lance
olate to ovate, located 1-3 mm below the articulation, densely strigose, caducous; pedicels
(5.5-) 8.0-20.6 mm long, sparsely to moderately strigillose, drying black. Flower buds
spheroidal to pyramidal, light to silvery green, sparsely to densely pubescent. Sepals 4
(-5), 5.8-9.5 (-10.5) mm long, 2.A-A.9 mm wide, up to 0.4 mm thick, triangular to ovate,
light green, lightly strigillose abaxially (or rarely densely pubescent), the sepal clearly vis
ible through the pubescence (or rarely not), drying black (or rarely golden), adaxially
densely pubescent or pubescent in lines or glabrous near base. Petals 4 (-5), 4.8-9.3
(-11.0) mm 2.2-3.8 mm wide, lanceolate to ovate, white to creamy white, some
long,
times infused with
pink, abaxially densely pubescent, adaxially pubescent in lines or
glabrous except along the margins. Receptacle densely pubescent with hairs up to 1mm
long. Androecium of numerous stamens, ca. 100; filaments 1.3-1.7 mm long, glabrous;
anthers 2.2-3.3 mm long, glabrous or very sparingly hirsute. Staminodes absent. Ovary
ca. 3.3 mm in diameter, ovoid, glabrous or with a few long hairs (or rarely moderately pu
bescent); placentation parietal, apparently (N=l) with 4 highly intruded placentae and 4
non-intruded placentae, the highly intruded placentae bearing ovules, the others not bear
ing ovules; style 3.0-4.5 mm long, glabrous (rarely pubescent); stigma obscurely lobed.
Fruit a berry, green when immature, mature fruit not seen. Seeds not seen. Fig. 12.
Phenology. Neosprucea montana has been recorded to have buds, flowers, or fruits in
every month, except April, May, and September. Reproduction is likely reduced during
the wettest months (April-May and August-December for Colombia/Venezuela, Febru
ary-April and October-December for Peru) and increased during the short drier periods
(January-March and July for Colombia/Venezuela, mid-July to August for Peru) (Snow
1976).
Distribution (Fig. 13). From the Venezuelan-Colombian border along the eastern
slopes of the Andes to south-central Peru but not yet recorded from Ecuador; montane or
submontane rainforest inmossy, streamside gaps and openings; 1440-2300 m.
Additional Specimens Examined.

Colombia. Antioquia: San Luis, Vereda Manizales, 12 km de San
Luis en la v?a San Luis-San bosques a lo largo del R?o Dormil?n,
Carlos, 06?05/N, 75?00'W, Callejas et al. 4283
(MO, NY, US).?Cundinamarca: Estaci?n Santana, arriba de Sasaima, [05?00'N, 74.4?W], Dugand &
Jaramillo 3873 (COL, US).?VALLE: Buga, Finca El Diluvio, Carretera Buga-El Placer a 33 km al E de Hotel
Guadalajara (Buga), Cordillera Central, certiente occidental, bosque nublado perturbado (parcialmente pri
mario), [03?50'N, 76?00/W],
Silverstone-Sopkin & Giraldo 6427 (NY). Peru. AMAZONAS: Mendoza,
[06?18'S, 77?25'W],
Woytkowski 8286 (MO); Bagua, Aramango, Cerros de Nueva Esperanza, 05?28'02"S,
78?23'11"W, V?squez et al. 27535 (BH); Bongara, Shillac, 05?52'S, 77?57'W, Leo Luna 326 (MO); Bongara,
near border with San Mart?n, along road Pedro Ruiz-Rioja, 05?40'S, 77?46'W, van der Werff et al. 16803 (BH);
Luya, Distrito Camporredondo, Anexo Tullanya, Tullanya, [06?07'S, 78?21'W], Campos et al. 3028 (MO).?
CAJAMARCA: San Ignacio, Picorana, 05?01'40"S, 78?54'30"W, Campos ?tal. 5474 (BH, MO); San Ignacio, San
Jos? de Lourdes, base del Cerro Picorana, 04?59'25"S, 78?54'05"W, D?az et al. 10356 (BH); San Ignacio, San
Jos? de Lourdes, base del Cerro El Parco, 05?04'38"S, 78?53'W, V?squez et al. 26725 (BH).?Jun?n: Tarma
Chanchamayo border, Rio Tulumayo drainage, Rondayacu, tributary valley and spur road N of Monobamba, 45
km S of San Ram?n, 11?18'S, 75?2(rW, Foster & Smith 9165 (BH, CAS, MO, USM).?PASCO: Oxapampa,
headwater of Rio Tunqui, trail to Chuchurras-Palcazu, 10?14'S, 75?28'W, Foster et al. 7738 (MO, USM).?SAN
Mart?n: Venceremos, near Amazonas border, Km 291 on Rioja-Pomacocha road, 05?45'S, 77?40'W, Gentry &
Smith 45255 (AAU-2, AMAZ, F, MO, NY, USM); Rioja, ca. 100 m to the E of Km 388 on road between Florida
(=Pomacocha) and Naranjos, just beyond the Amazonas-San Martin border along a hillside stream, 05?39.932'S,
2008 NEOSPRUCEA 35
FIG. 12. Neosprucea montana. A. Leafy branch with young inflorescences. B. Young leaf, showing torus
shaped glands at the teeth. C. Young inflorescence, showing evident pulvini on the petiole of the subtending leaf.
D. Mature inflorescence, showing dark-drying sepals. E. Flower at early anthesis, side view. F. Flower, top view,
showing lack of staminodes. G. Flower, longitudinal section. H. Stamens, showing apical pores. (Based on: A-C,
Alford 3129; D-H, Campos et al. 3028.)
85? W
80? 75? 65?
70?
FIG. 13. Distribution o? Neosprucea montana and N sararensis.
77?45.275'W, Alford 3129 (BH, DUKE, HUT, MO, MOL, USM); Rioja, Pedro Ruiz-Moyobamba road, Km
390, Venceremos, 05?50'S, 77?45'W, Smith 4429 (F,MO, NY), Smith & V?squez 4760 (AAU, AMAZ, F, NY,
USM), Smith & V?squez 4545 (AMAZ, IBE, MO, USM). Venezuela. T?CHIRA: wet forested slopes along
Quebrada Agua Azul, over slate-shale substrate, south of El Reposo, 14 km SE of Delicias, 07?31'N, 72?24'W,
Steyermark & Liesner 118458 (L, MO, NY, S).
Neosprucea montana is a middle- to high-elevation species with glabrous stamens,

long pedicels, and (usually) sparsely pubescent sepals that dry black. It is similar to the
poorly known N. sararensis and differs clearly only in filament/anther length (filaments
shorter than the anthers vs. longer inN. sararensis). It usually differs from N. sararensis
in its sparsely pubescent sepals that dry black, its racemose inflorescence, and its smaller
and generally broader leaves (elliptic, 5-16.5 [-20.1] cm long vs. elliptic to narrowly
elliptic, 11.5-28.1 cm long inN. sararensis). Neosprucea montana may be confused with
N. wilburiana, but the latter has hirsute anthers and highly congested, fascicle-like
2008 NEOSPRUCEA 37
inflorescences, and occurs at lower altitudes. Itmay also be confused with N. tenuisepala
on the basis of its sparsely pubescent and black-drying sepals, but N. tenuisepala has hir
sute anthers, filamentous staminodes, usually shorter pedicels (2.7-8.6 mm), and grows at
lower elevations. Sleumer (1980) and Cuatrecasas (1955) indicated in their keys and de
scriptions that N. montana has hirsute anthers, but I have found this rarely true.
Two collections of N montana from the southern end of its range have densely pu
bescent sepals and inflorescence axes. Although they lack the characteristic black-drying
sepals of the typical N. montana, they are identical in all other characters. No collections
are known from Ecuador, but one can expect to find it there.
4. Neosprucea paterna M. H. Alford, sp. nov.?Type: Guyana. Potaro-Siparuni: forest

just SE of the junction of Kurupukari-Annai road and Surama road, ca. 80 km
[S] by road from the Essequibo River, 60 m, 04?05.487'N, 59?02.491'W, 14May
2003 (fl), M. Alford et al. 3149 (holotype: BRG!; isotypes: AAU! B! BH! BM!
DUKE! F! FDG! K! MICH! MO! OS! NY! P! U! US! VEN!WIS!).
Arbor mediocris. Folia elliptica vel obovata, ad basim cordata, margine glanduloso
serrato vel integro praeter dent?s paucos ad basim, dentibus pubescentibus, venatione
actinodroma. Inflorescentia racemosa; s?pala 4, oblonga, 1.2-1.5 cm longa, utrinque
dense tomentoso-sericea; p?tala 4 (-5), ?urea, oblonga 1.3-1.65 cm longa, utrinque dense
tomentoso-sericea; stamina filamentis capillaribus glabris, 3-4 mm longis, antheris an
guste linearibus (5-) 6-9 mm longis graciliter hirtis; staminodia pauca angustissime cap
illaria, aliqua staminodia antherodiis parvis instructa. Ovarium parvum, glabrum. A
Neosprucea rimachii foliis ellipticis vel obovatis et ovariis parvis glabris differt.
Trees to 25 m tall. Trunk to 15 cm dbh; bark pale gray to whitish, general aspect
smooth, fine aspect finely fissured-scaly. Branches green to dark gray when young, drying
black or brownish gray, obscurely 3-4-sided to terete, lightly pubescent, glabrescent with
growth; lenticels numerous, white to light brown, elliptic to narrowly oblong. Leaf blades
(6-) 8-24 cm long, (3.5-) 5.5-12.5 cm wide, elliptic to obovate, sometimes slightly asym
metrical with respect to themidvein; apex (obtuse to) acute to acuminate, up to ca. 2.5 cm;
base abruptly cordate to almost auriculate; margin serrate-crenate, especially in very
young leaves or leaves of saplings, 0.5-1 tooth/cm, in older leaves essentially entire or
with a few scattered teeth or few teeth near the base, teeth obtuse with a pubescent gland
at or near the apex, gland abaxial or marginal; chartaceous; adaxial surface somewhat
glossy, mostly glabrous but pubescent to glabrescent on major veins; abaxial surface
softly hirtellous; venation actinodromous, 3 (-5)-veined from the base, secondaries
clearly semicraspedodromous in leaves of saplings, appearing brochidodromous or fes
tooned brochidodromous in leaves of mature plants, 1-4 pairs of secondaries above the
basal pair, acutely angled to the midvein, conspicuously raised abaxially, reddish in
saplings, tertiaries more or less perpendicular to the midvein. Petioles 0.8-3 cm long,
1.0-1.6 mm wide, lightly to densely pubescent, reddish in saplings, not canaliculate or
only canaliculate near petiole apex and base; occasionally with 1 (-5) gland(s) near the
apex; pulvinus absent. Stipules 7.0-10.5 mm long, 0.5-0.7 mm wide, linear-lanceolate,
often with small glandular teeth along the margin, densely pubescent, caducous. Inflores
cences terminal racemes or few-branched panicles with 2-6 flowers, 3.5-9.5 cm long, axis
to 2.3 mm thick, densely pubescent; bracts not seen, caducous; bracteoles 5-10.5 mm
long, 1-2.5 mm wide, narrowly ovate to lanceolate, located 1.6^4.9 mm below the
articulation, densely pubescent, caducous; pedicels 5.0-12.6 mm long, densely yellow
FIG. 14. Neosprucea paterna. A. Leaf from sapling, showing serrate-crenate margin and pubescent glands
on the abaxial surface at the leaf teeth. B. Flowering branch, showing the almost entire leaves typical of mature
individuals. C. Flower. D. Flower, longitudinal section, with detail of the stigma. E. Staminode and transitional
stamen. F. Stamens, showing elongated apical pores. G. Pistil and cross section of the ovary, showing highly in
truded parietal placentation and numerous ovules. (Based on: A, Alford 3133; B-G, Alford 3149).
2008 NEOSPRUCEA 39
58? W
FIG. 15. Distribution o? Neosprucea paterna.
pubescent. Flower buds ovoid to squarish with a pyramidal top, densely yellow-pubescent.
Sepals 4,1.2-15.0 mm long, 5.5-8.5 mm wide, quite thick, up to 1.3mm thick at the apex,
ovate to oblong, green, densely pubescent on both sides, conspicuously nerved on the
abaxial side. Petals 4 (-5), 13.0-16.5 mm long, 4.5-5.5 mm wide, narrowly ovate to ob
long, light yellow at anthesis becoming greener or with a tinge of pink after anthesis,
densely pubescent, thinner than the sepals. Receptacle densely pubescent. Androecium of
numerous stamens, ca. 120; filaments 3.0-4.0 mm long, becoming reddish orange after
anthesis, glabrous; anthers (5-) 6-9 mm long, hirsute. Staminodes consisting of subtle
outer rows of stamens with reduced anthers and filamentous staminodes lacking anther
remnants, slightly shorter than the fertile stamens. Ovary ca. 3.5 mm in diameter, spher
oidal or weakly lobed, glabrous; placentation parietal, placentae ca. 7-8; style 15 mm
long, glabrous; stigma minutely expanded and lobed. Fruit ca. 6.7 mm in diameter when
young, spheroidal, green but white-speckled, basically glabrous; mature fruit not seen.
Seeds not seen. Fig. 14.
Phenology. Known to flower and fruit inmid-May and to flower at the beginning of
March, approximately corresponding to the February-April dry season (Bernard 1999;
Clarke et al. 2001).
Distribution (Fig. 15). West-central Guyana in dry evergreen forest near the savan
nah-forest ecotone; forest over white sand dominated by Eperua ("wallaba," Legumi
nosae); ca. 100 m.
Additional Specimens Examined. Guyana. Potaro-Siparuni: forest just SE of the junction of Ku
rupukari-Annai road and Surama road, ca. 80 km by road from the Essequibo River, 04?05.487/N, 59?02.491'W,
Alford et al. 3148 (BRG, US); roadside 7.2 km from Surama village, near intersection with Kurupukari-Annai
road, 04?05.718'N, 59?02.610,W, Alford et al. 3133 (BRG, FDG, US); newly cleared right-of-way, 15-20 km
NE of Surama Village Road, along sides of cleared bulldozer track, in dense forest, 04?15'N, 58?56'W,
McDowell et al. 2103 (BM, G, NY, TEX, U, US).
Neosprucea paterna is the only species of the genus known from the Guayana Shield
region. Its nearest congeneric members are found ca. 1600 km away in Venezuela and
Colombia {N. montana/N. sararensis and TV.rimachii, respectively). Neosprucea paterna
is easily diagnosed by its elliptic to obovate leaves with actinodromous venation and
abruptly cordate to auriculate bases. Like N rimachii, it has actinodromous venation, pro
jecting petiolar glands, leaf teeth with pilose glands, and a yellow perianth. It differs from
that species in having the leaves of mature plants essentially entire or toothed only near
the base and in having small and glabrous fruits.
The specific epithet "paternus" (Latin, "of the father") is chosen to honor my father,
Emmette Earl Alford (b. 1943), a forester who traveled to the tropics for the first time to
help me relocate and collect this species.
5. Neosprucea pedicellata Little, J. Wash. Acad. Sei. 38(3): 104. 1948.?Type:

Ecuador. Pichincha: Santo Domingo de los Colorados, [00?10'S, 79?10'W],
560 m, 2 Apr 1943 (fl, young fr), E. Little 6152 (holotype: US!; isotypes: F! GH!
K! NY-2!).
Treesto 15 m tall. Trunk to 27 cm dbh, sometimes buttressed to 50 cm; bark pale

gray, smooth to slightly scaly, slash orange-brown. Branches dark brown when young, ob
scurely 3-4-sided, lightly to moderately short-pubescent, pubescent, or strigillose;
lenticels infrequent to numerous, sometimes obscure, light brown, (widely elliptic to)
2008 NEOSPRUCEA 41
narrowly elliptic to linear. Leaf blades 8.0-21.0 (-24.0) cm long, 3.0-9.0 (-10.5) cm
wide, elliptic, infrequently asymmetrical with respect to the midvein; apex acuminate;
base rounded to acute (to attenuate), eglandular, but glands associated with leaf teeth
sometimes closely approaching the lamina-petiole junction; margin serrate-crenate,
0.7-2 teeth/cm, teeth obtuse with a glabrescent gland at the apex, gland completely abax
ial but abutting the leaf margin; chartaceous; sparsely strigillose on veins of both sur
faces, rarely short-pubescent, hirtellous, or hirsute abaxially, often glabrescent above; ve
nation acrodromous, 3-veined from the base to the apex, occasionally 1-2 upper
secondary pairs (or secondary veins per side of the lamina) arising at an acute angle from
the midvein near apex, prominent tertiaries ca. 8-28, perpendicular to the midvein and
secondaries, raised abaxially. Petioles 0.7-2.8 (-3.3) cm long, 1mm wide, moderately to
densely short-pubescent throughout, canaliculate or flattened adaxially; pulvinus weakly
or moderately developed at apex, absent at base. Stipules 0.6-0.8 mm long, triangular,
entire, densely pubescent, caducous. Inflorescences axillary racemes with 1-5 flowers,
up to 6.5 cm long, axis 0.5-1.8 (-2.0) mm thick, densely pubescent with short and long
(0.5 mm) hairs; bracts 0.9 mm long, triangular, densely pubescent, caducous; bracteoles
1.8 mm long, lanceolate, located <0.8 mm below the articulation, densely pubescent, ca
ducous; pedicels 1.4-2.7 cm long, articulation more densely pubescent than the pedicel.
Flower buds oblate to ovoid, capped by ridges where the sepals abut against each other.
Sepals 4, 1.0-1.3 cm long, 0.35-0.80 cm wide, up to 0.4 mm thick medially and 0.9 mm
at the apex, ovate, light green, densely pubescent with silvery hairs on both sides. Petals
4, 1.3-1.5 cm long, 0.45-0.70 cm wide, ovate to lanceolate, white to creamy white,
densely pubescent on both sides. Receptacle densely hirsute. Androecium of numerous
stamens, ca. 80-160; filaments 2.5-4.3 mm long, glabrous; anthers 5.0 mm long, bright
yellow, hirsute, especially along the adaxial surface of the connective. Staminodes fila
mentous, flattened, glabrous, about the same size as the stamens or a little longer, white
(to yellowish). Ovary ca. 2.5 mm in diameter, spheroidal, glabrous or with a few scat
tered hairs; placentation highly intruded parietal, placentae ca. 6-7; style 7.0 mm long,
glabrous; stigma more or less terete, slightly expanded at apex. Fruit a berry, ca. 8.5 mm
in diameter or more, spheroidal, glabrous or almost so. Seeds few, 2.5 mm long, 2.0 mm
wide, ellipsoid. Fig. 16.
Phenology. Collected in flower and fruit throughout the year, but with fewer collec
tions during the wettest months of January to June (Pourrut et al. 1995; Neill & J0rgensen
1999).
Distribution (Fig. 17). Pacific basin of Ecuador and Colombia; moist and wet lowland
tropical rainforest and premontane rainforest; 80-780 m.
Additional Specimens Examined. Colombia. Nari?o: Tumaco, vereda de Santa Mar?a, R?o Rosario,
[01 ?46'N, 78?37'W], Romero-Casta?eda 5175 (COL). Ecuador. Carchi: property of Humberto Rosero, on N
side of R?o Mira, just above cable crossing, upstream from Lita and downstream from Cachaco, 00?52'N,
78?26'W, Boyle et al. 3688 (MO), 3689 (MO).?Esmeraldas: R?o Onzole, primary tall forest on Estero Chon
taduro, [00?45'N, 79?10'W], Jativa & Epling 1102 (L-fragment, MO, NY, S, U, US-2); Rio Pambil, primary for
est near junction with Estero Capuli, [00?42'N, 79?05'W], Jativa & Epling 1062 (MO, NY, S, U, US-2); San
Lorenzo, Finca La Chiquita, 01?13'N, 78?49'W, Palacios 2600 (AAU, K, MO, NY, QCNE); Anchayacu, Eloy
Alfaro, Mayronga, 00?53.45'N, 79?13.03AW, Pennington et al. 14300 (K-2, MO, U), Pennington et al. 14684
(K), Pennington et al. 14285 (K); San Lorenzo, Reserva ?tnica Aw?, Centro Ricaurte, colectado a orilla de un
estero, 01?10'N, 78?32'W, Tipaz et al. 2184 (MO, NY, QCNE).?Los R?OS: Buena F?, 2 km S of Patricia Pilar
at the R?o Palenque Science Center, 00?36'S, 79?22'W, Alford et al. 3076 (BH, DUKE, MO,
QCA, QCNE, US),
Dodson & Gentry 6318 (AAU, MO, QCA), Dodson & Gentry 10066 (MO), Dodson & Tan 5340 (AAU, MO,
QCA, US), Gentry 12001 (L, MO-2, NY, QCA), Vodicka 859 (BH, QCA), Watson 172 (NY).?Pichincha: 20
FIG. 16. Neosprucea pedicellata. A. Flowering branch, showing detail of torus-shaped glands at the leaf
teeth. B. Petiole and abaxial surface of leaf. C. Flower at late anthesis, top view and side view, showing the outer
whorl of staminodes. D. Flower, longitudinal section. E. Staminodes. F. Stamens. G. Fruiting branch, showing
persistent perianth. (Based on Alford 3076.)
2008 NEOSPRUCEA 43
82?W 80 78? 76
PACIFIC OCEAN
Neosprucea j
pedicellata \
FIG. 17. Distribution of Neosprucea pedicellata.
km W of Santo Domingo de los Colorados, [00?15'S, 79?22'W], Cazalet & Pennington 5183 (K, L-fragment,
NY, US), Cazalet & Pennington 5046 (K, NY, US); Santo Domingo de los Colorados, 00?16'S, 79?14'W, Cer?n
& Benavides 6172 (MO, NY, QCNE-2); Finlandia, 16 km E of Santo Domingo de los Colorados, [00?20'S,
79?00'W], Gentry et al. 12167 (IBE, L, MO); Reserva Forestal ENDESA, R?o Silanche, "Corporaci?n Forestal
Juan Manuel Durini," Km 113 de la carretera Quito-Pto. Quito, faldas occidentales, a 10 km al N de la carretera
principal, OO^X 79?02'W, Jaramillo 7002 (NY), Jaramillo 7003 (AAU, MO), Jaramillo 7558 (AAU, MO,
NY-2, QCA-2).
Neosprucea pedicellata is the only species of the genus known from the Pacific basin
of South America. It is diagnosed by the combination of large flowers (petals 1.3-1.5 cm
long) and long pedicels (>1.4 cm). Sleumer (1980) reported the sepals as 12-20 mm long,
but I have not found this to be true.
6. Neosprucea rimachii McDaniel ex M. H. Alford, Gr?ndez & V?squez, Arnaldoa 11:

38. 2005 ["2004"].?Type: Peru. Loreto: Maynas, Dtto. Iquitos, Rio Nanay trail
from Astoria to R?o Mazan, upland mature forest over sandy clay, 160-200 m, 9
Jan 1976 (fir), S. McDaniel & M. Rimachi Y. 20402 (holotype: AMAZ; isotypes:
F-photo! IBE-2! MO! NY! US! USM, WIS).
Trees to 12 m tall. Trunk to 15.1 cm dbh; bark unknown. Branches drying coppery
brown when young due to the pubescence, maroon-brown beneath, obscurely 3-4-sided,
moderately to densely short-pubescent; lenticels common, light brown, elliptic to linear.
Leaf blades (6-) 11-20 cm long, (4-) 7-11 cm wide, ovate, symmetrical; apex acuminate
or abruptly acuminate for 0.5-3 cm; base cordate, eglandular (but see petiole); margin ser
rate-crenate, 1-2 teeth/cm, teeth obtuse with a pilose gland at or near the apex, gland
abaxial, abutting the leaf margin or infrequently marginal; chartaceous; scarcely to
densely pubescent on both faces, densely pubescent especially on the veins; venation
actinodromous, 3-7-veined from the base, 1-2 pairs of prominent secondaries (or sec
ondary veins per side of lamina) above the basal ones, acutely angled, semicraspedodro
mous, veins raised abaxially. Petioles 1-3 cm long, 0.5 mm in diameter, densely pubes
cent to hirtellous, not canaliculate or only weakly so at apex; pulvinus absent; 1-3
projecting glands on the distal third. Stipules not seen, caducous. Inflorescences subter
minal, few-flowered racemes with 1-2 (-5+?) flowers, up to 3 cm long, axis up to 1.7 mm
thick, moderately to densely pubescent to hirtellous; bracts and bracteoles caducous, not
seen; pedicels 1.5-2.0 cm long, densely short-pubescent or hirtellous with golden hairs.
Flower buds ovoid, covered in dense yellow pubescence. Sepals 4 (-5), 1-1.7 cm long,
0.5-0.8 cm wide, up to 1.3 mm thick, oblong to narrowly elliptic, yellowish green,
densely pubescent on both sides except for a central glabrous stripe on the adaxial surface.
Petals like sepals but 1.6-1.9 cm long, 0.5-1.1 cm wide, yellow. Receptacle moderately
hirsute but inconspicuously so. Androecium of numerous
stamens, ca. 120; filaments ca.
6 mm long, glabrous; anthers ca. 5-11 mm long, hirsute. Staminodes not seen, but elon
gated filaments with reduced anthers found at the outer edge of the androecium. Ovary
ovoid, densely tomentose to sericeous; placentation parietal, number of placentae and
ovules unknown; style 1.0-1.2 cm long, glabrous; stigma minutely expanded and lobed.
Fruit a dry berry, ca. 1.1-1.4 cm in diameter, spheroidal or widely ovoid, densely tomen
tose to sericeous.Seeds ca. 19, 3.0-3.5 mm long, 2.0 mm wide, ellipsoidal. Fig. 18.
Phenology. Collected in flower and fruit in December and fruit in early January.
Flowering possibly coincides with the somewhat drier months of the year, June to De
cember (Tosi, Jr. 1960).
Distribution (Fig. 19). Amazon basin near Iquitos, Peru, downstream in neighboring
Colombia, and in the Parque Nacional Manu in southern Peru; upland mature forest
over sandy clay (Iquitos and Colombia) or high terraces, slopes, and ravines (Manu);
160-350 m.
Additional Specimens Examined. Colombia. Amazonas: Mpio. Leticia, Parque Nacional Natural
Amacayacu, Quebrada de Agua Pudre, ca. 1.5 km NE de desemboca dura sobre el Rio Amacayacu, Muestreo
Permanente Estrat?gica del MO, ?rbol #517, 03?47'S, 70?15'W, Pipoly 16411 (AMAZ, COL, MO). Peru.
Madre de Dios: Parque Nacional Manu, R?o Manu, Pakitsa Station, Tachigali Trail to 8 km N of the camp,
11?56'S, 71?16'W, Foster &. Balde?n 12812 (US).
2008 NEOSPRUCEA 45
3 cm.
FIG. 18. Neosprucea rimachii. A. Fruiting branch, showing detail of the leaf teeth. B. Petiole and leaf base.
C. Fruit, showing persistent perianth. D. Outer stamens, showing apical pores. (Based on McDaniel & Rimachi
20402.)
75?W
80? 70?
65?
FIG. 19. Distribution o? Neosprucea rimachii.
Neosprucea rimachii is easily recognized by its coarsely toothed, ovate leaves with a
cordate base and its large (ca. 1.1-1.4 cm in diameter) sericeous fruit. It and N. paterna
are the only species with actinodromous venation, projecting petiolar glands, leaf teeth
with pilose glands, and a yellow perianth.
7. Neosprucea sararensis Cuatrecasas, Trop. Woods 101: 21. 1955.?Type: Colombia.

Norte de Santander: Sarare, Cordillera Oriental, hoya del R?o Margua, Quebrada
del Sararito entre Miranda y Alto de laAurora, [07?11TS?, 72?20.6'W], 950-1087
m, 22 Nov 1941 (fl), J. Cuatrecasas 13400 (holotype: US!; isotype: COL! L
fragment!).
Trees, probably small to medium-sized (only one report: 3 m). Trunk size unknown;
bark unknown. Branches maroon-brown when young, 4-sided, moderately strigillose,
glabrescent; lenticels common, light brown, elliptical to oblong. Leaf blades 11.5-28.1 cm
long, 3.8-9.4 cm wide, elliptic to narrowly elliptic, sometimes asymmetrical with respect
to the midvein; apex acuminate, 0.9-2.0 cm; base truncate to acute to abruptly acute,
eglandular; margin serrate-crenate, 0.4-0.8 teeth/cm, teeth obtuse with a glabrous gland
near the apex, gland abaxial, abutting the leaf margin or slightly inset; chartaceous;
2008 NEOSPRUCEA 47
adaxial surface glabrous or lightly strigillose near base; abaxial surface glabrous except
for scattered hairs or sparsely to moderately strigillose veins; venation acrodromous, 3
veined from base to apex, occasionally one upper secondary pair (or a single secondary
vein per side of the lamina) arising at an acute angle from the midvein near apex, promi
nent tertiaries ca. 15-37, perpendicular to the midvein and secondaries, veins raised abax
ially. Petioles 9.0-28.5 mm long, 0.8-1.5 mm in diameter, lightly strigillose to glabres
cent, occasionally canaliculate on the adaxial surface; solid pulvini at the base and apex
present. Stipules 1.3 mm long, 0.4 mm wide, narrowly triangular, entire, densely strig
illose, caducous. Inflorescences axillary racemes or panicles with 2-16+ flowers, 4-16 cm
long, axis (0.5-) 0.8-2.5 mm thick, sparsely to densely strigillose, with a few longer, erect
hairs, or pubescent and strigillose, glabrescent near the base; bracts 2 mm long, 1mm
wide, ovate, densely strigillose abaxially, glabrous adaxially except for a line of pubes
cence near themiddle; bracteoles not seen, presumably caducous; pedicels 7-13 mm long,
moderately to densely strigillose with intermixed erect hairs or pubescent. Flower buds el
lipsoidal, color unknown, thoroughly but thinly strigillose, the sepal epidermis sometimes
visible through the indumentum. Sepals 4 (-5), 9.0-10.5 mm long, 5.0 mm wide, to 0.4
mm thick, ovate, green, thinly but thoroughly strigillose. Petals 4 (-5), like sepals except
densely pubescent on both surfaces, white. Receptacle moderately to densely pubescent.
Androecium of numerous stamens, ca. 60 (N=l); filaments 2.2-2.5 mm long, glabrous;
anthers 2 mm long, glabrous. Staminodes absent. Ovary ovoid, glabrous; placentation
highly intruded parietal, placentae ca. 8-9; style 4 mm long, glabrous; stigma minutely
lobed. Fruit a berry, ca. 7 mm in diameter, spherical, glabrous. Seeds several, 2.8 mm
long, 1.5 mm wide, ellipsoidal to narrowly ellipsoidal.
Phenology. According to the two collections with adequate label information, N.
sararensis flowers in late November and fruits in late July.
Distribution (Fig. 13). Andes of northern Colombia; montane cloud forest; 950
1700 m.
Additional Specimens Examined. Colombia. [Locality and date unknown, probably near Bogot? or in
Cundinamarca, 1760-1808, see Mutis (1957-1958) and Blanco y Fern?ndez de Caleya & del Valle Stervinou
(1991)] (fr), Mutis 1025 (US), Mutis 2211 (US).?BOYAC?: carretera de Chiquinquir? a Pauna, arriba de La Es
peranza, [05?35'N, 73?55'W], Jaramillo-Mej?a et al. 3572 (COL).
Neosprucea sararensis is a poorly known, high-elevation species endemic to Colom

bia. It is one of only three species of the genus that lack staminodes (the other two are N.
montana and N. wilburiana) and can be readily distinguished from N. wilburiana by hav
ing glabrous anthers and occurring at higher elevation. Neosprucea sararensis can be re
liably distinguished from N. montana only by comparison of filament/anther lengths, al
though N. sararensis has longer, narrower leaves, longer and more branched
inflorescences, and sepals with enough pubescence to hide the dark, dried epidermis.
Neosprucea sararensis combines the vegetative and floral features of N montana?
petiolar pulvini, elliptic leaves, glabrous anthers, no staminodes?and the inflorescence
features of N. grandiflora?a long, thick, and branched axis. The Mutis specimens differ
from the type in that they have thinner, less branched inflorescence axes, which makes
them hardly distinguishable from N montana.
The IUCN Red List (Calder?n 1997) reports that N. sararensis is endangered (EN
Bl+2c,ver. 2.3, 1994).
8. Neosprucea tenuisepala M. H. Alford, sp. nov.?Type: Peru. Amazonas: Prov. Con

dorcanqui, Distr. El Cenepa, Comunidad de Mamayaque, R?o Cenepa, Cerro
Sakee gaig, bosque primario, 04?34'49"S, 78?14'01"W, 900-1000 m, 12 Feb
1997 (fl), R. V?squez et al. 22494 (holotype: MO!; isotypes: BH! HUT!).
Arbor parva. Folia elliptica vel obovata, ad basim cuneata vel acuta, margine glan
duloso-serrato. Inflorescentiae axillares, 2.3-7.2 cm longae, pedicellis 2.7-8.6 mm longis
sub anthesi; s?pala 4 (-5), abaxialiter sparsim pubescentia, in sicco nigra; p?tala 4 (-5),
utrinque tomentoso-sericea. Stamina creberrima filamentis capillaribus hirtis 0.9-1.3 mm
longis, antheris anguste linearibus 3.7-4.9 mm longis graciliter hirtis; staminodia an
gustissime capillaria; receptaculum intra stamina valde hirsuto-setosum. A Neosprucea
montana pedicellis brevioribus, antheris longioribus, et habitatione ad altitudinem inferi
orem differt. A N. grandiflora foliis tenuiter chartaceis et sepalis abaxialiter modice pu
bescentibus differt.
Trees to 10 m tall. Trunk to at least 6 cm dbh; bark grayish brown, texture unknown.
Branches light brown to maroon-brown when young, initially 4-sided, becoming terete
with growth, densely strigillose to glabrescent; lenticels obscure, common, reddish brown,
circular to oblong. Leaf blades 6.5-26.5 cm long, 3.0-11.3 cm wide, elliptic, narrowly el
liptic, or obovate, rarely ovate, occasionally slightly asymmetrical with respect to themid
vein; apex acuminate to abruptly long-acuminate, 1-4 cm long; base attenuate to obtuse,
rarely truncate, occasionally with 1-several glands associated with basal leaf teeth; mar
gin serrate-crenate, 0.5-2 teeth/cm, teeth obtuse or rounded, rarely obscure, with a
glabrous gland at or near tooth apex, abaxial and abutting the leaf margin, the margin
sometimes there; thinly chartaceous; adaxial surface glabrous except for pu
thickened
berulent midvein and strigillose lamina base; abaxial surface strigillose on major veins,
glabrescent elsewhere, rarely pubescent throughout (Jaramillo 5607A); venation acrodro
mous, 3-veined from the base to the apex, almost always 2-5 distal pairs of secondary
veins (or 2-5 secondary veins per side of the lamina) arising at an acute angle from the
midvein near apex, prominent tertiaries 12-26, perpendicular to the midvein and lateral
secondaries, veins raised abaxially. Petioles 5.7-15.8 mm long, 0.6-2.1 mm in diameter,
moderately strigillose, canaliculate adaxially; pulvinus absent or obscure. Stipules 0.6-1.0
mm long, 0.2-0.3 mm wide, deltate to narrowly triangular, entire, densely strigillose,
sometimes located as much as 1mm from petiole base. Inflorescences axillary (or sub-ter
minal) racemes (or panicles) of 1-7 flowers, 2.3-7.2 cm long; bracts and bracteoles ca. 0.5
mm long, triangular to lanceolate, densely strigillose, caducous; pedicels 2.7-8.6 mm
long, densely strigillose. Flower buds ovoid, green to yellowish red, moderately strigillose
but epidermis of the sepals visible beneath. Sepals 4 (-5), 6.7-11.2 mm long, 2.8-5.3 mm
wide, thin, to 0.2 mm thick, ovate to lanceolate, light green, green, or yellowish red (?),
lightly to moderately strigillose abaxially, sepal epidermis visible, drying black, densely
pubescent adaxially. Petals 4 (-5), 6.5-11.4 mm long, 2.4-3.2 mm wide, ovate, narrowly
elliptic, or lanceolate, white,
possibly light green or creamy white, densely pubescent
throughout abaxially, densely pubescent or pubescent in lines adaxially. Receptacle
densely pubescent. Androecium of numerous stamens, ca. 60; filaments 0.9-1.3 mm long,
hirsute; anthers 3.7-4.9 mm long, hirsute, hairs mostly on the adaxial surface of the con
nective. Staminodes filamentous, about half the length of the stamens, rarely absent (?).
Ovary ca. 2 mm in diameter at anthesis, ovoid, glabrous or with a few scattered hairs; pla
centation parietal, placentae ca. 4-5; style 4.5-5.0 mm long, glabrous; stigma minutely
lobed. Fruit a berry, ca. 1.2 cm in diameter, spheroidal, green when immature. Seeds
2008 NEOSPRUCEA 49
FIG. 20. Neosprucea tenuisepala. A. Flowering branch, showing the dark abaxial epidermis of the sepals
when dry and detail of a torus-shaped gland at a leaf tooth. B. Petiole and abaxial surface of leaf. C. Leaf, adax
ial surface and venation. D. Flower bud, showing valvate aestivation. E. Flowers, side/top view (early anthesis)
and longitudinal section. F. Staminodes. G. Stamens, with detail of apical pores. (Based on: A, B, V?squez et al.
22494; C-F, Huashikat 1338.)
FIG. 21. Distribution o? Neosprucea tenuisepala.
numerous, ca. 24, 1.9-2.7 mm 1.1-1.7 mm wide, ovoid or

long, irregularly ellipsoid.
Fig. 20.
Common Names. Language source unknown: "namakia betsagnum" (Ancuash 1156),
"shigk?tnum" (Huashicat 376), "tseas" (Kayap 844), "uci betsagnum" (Kayap 1478).
Humabisa: "kantsa" (Tunqui 1101), "uyukmants" (Tunqui 951), or "ujukm?nts"
(Huashicat 586, 848, 1166, 1338, 1572, 2304).
Phenology. Flowers and fruits throughout the year.
Distribution (Fig. 21). Amazonas province of Peru, and Morona-Santiago and
Zamora-Chinchipe provinces of Ecuador; wet and moist tropical rainforest and montane
rainforest; 180-1300 (-1750) m.
Additional Specimens Examined. Ecuador. Morona-Santiago: Centro Shuar Yukutais, W of Pedro

Kunkumas's house and S of school, 02?30'S, 78?08/W, Bennett et al. 4091
(NY); Pumpuentza, primary forest
WNW of the village, 02?25'S, 77?20,W, Brandbyge & Asanza 32322 (AAU, MO, US); San Jos? Grande, cuenca
del R?o Paute entre el R?o Cardenillo Grande y Cardenillo Chico, 02?36'S, 78?26'W, Cer?n & Benavides 14126
(BH); Loma San Jos? Grande-Sendero San Jos?-R?o Bombioza, 02?38'S, 78?27/W, Cer?n 14288 (BH); road
Lim?n (General Plaza) to Macas, ca. Km 20 from Lim?n, [02?55'S, 78?15'W], Harling & Andersson 12871
2008 NEOSPRUCEA 51
(GB).?Zamora-Chinchipe: Shaime, sendero hacia el Hito, [04?20'S, 78?40'W], Jaramillo 14453 (NY, QCA);
Miasi, sendero hacia el Hito, [04?19'S, 78?39'W], Jaramillo 14252 (NY, QCA); Nangaritza Cant?n Shaimi,
frente a destacamento Militar, margen derecha del R?o Nangaritza, 04?18'S, 78?43/W, Palacios et al. 8702
(MO). Peru. Amazonas: R?o Cenepa, vicinity of Huampami, ca. 5 km E of Ch?vez Valdivia, 04?30'S,
78?30'W, Ancuash 1156 (F, IBE, L, MO); Rio Santiago, 800 m atr?s de la comunidad de Caterpiza, banda este
de Caterpiza, [03?55'S, 77?42'W], Huashicat 376 (MO); Rio Santiago, monte virgen, 1 km a la banda de la Co
munidad de Caterpiza, trocha de mitayar, banda este de la Quebrada Caterpiza, [03?55'S, 77?42'W], Huashikat
586 (MO); Rio Santiago, 3 km atr?s de comunidad Caterpiza, banda este de Caterpiza, [03?50'S, 77?40'W],
Huashikat 1166 (MO); Rio Santiago, monte virgen, 2 km atr?s de comunidad Caterpiza, banda este, en trocha,
[03?50'S, 77?40'W], Huashikat 1338 (L, MO); valle del Rio Santiago, approx. 65 km N de Pinglo, Quebrada
Caterpiza, 2-3 km atr?s de la comunidad de Caterpiza, [03?50'S, 77?40'W], Huashikat 1572 (MO), Huashikat
2304 (MO), Tunqui 951 (BH, L, MO), Tunqui 1101 (MO); Rio Santiago, monte virgen, 2 km atr?s de la comu
nidad Caterpiza, [03?50,S, Huashikat
77?40'W], 848 (L, MO); Quebrada Etseketai, Rio Cenepa, [04.3?S,
78.07?W], Kayap 844 (GH, MO); al lado de Huampami, monte, [04?27'S, 78?10'W], Kayap 1142 (MO), 1478
(GH, MO); Quebrada Cunup, monte, cerca a la chacra, Kayap 1291 (MO); Condorcanqui, Distrito El Cenepa,
Comunidad de Mamayaque, bosque primario, arriba de la quebrada Sa?sa, 04?37'08"S, 78?13'46"W, Rodr?guez
et al. 1420 (BH, MO, HUT); Condorcanqui, Distrito El Cenepa, Comunidad de San Antonio, de ribera,
bosque
Rio Cenepa, 04?37'08"S, et al. 180 (MO); Condorcanqui, Distrito
78?13'46"W, Rojas Cenepa, bosque ribere?o
de la quebrada Tayo, 04?31'35"S, 78?10'34"W, V?squez et al. 22129 (HUT, MO); Distrito El
Condorcanqui,
Cenepa, Comunidad de Mamayaque, cabecera de la quebrada Saas?, 04?37'03"S, 78?19'57"W, V?squez &
Quiaco 24554 (HUT, MO); Condorcanqui, Distrito El Cenepa, Comunidad de Tutino, 04?33'05"S, 78?12'54"W,
V?squez et al. 22469 (MO).
Neosprucea tenuisepala is similar to N. grandiflora and N. melastomatoides

but differs in its thinly chartaceous leaves, thin, short, and sparsely-flowered inflores
cences, and abaxially lightly pubescent sepals, which dry black and do not have the epi
dermis obscured by pubescence. Neosprucea tenuisepala commonly has obovate leaves
with an abruptly acuminate apex up to 4 cm long, features rare in either N. grandiflora
or N. melastomatoides. See the discussion under N. grandiflora (no. 1) for additional
comparisons. Neosprucea tenuisepala may be superficially confused with M montana
or N. wilburiana. Neosprucea tenuisepala resembles N. montana due to the lightly
pubescent sepals that dry black; however, N. montana lacks staminodes, has longer
pedicels at maturity ([5.5-]8.0-20.6 mm vs. 2.7-8.6 mm), has pulvini at both the apex
and base of the petiole, and grows at higher elevations in cloud forest. Neosprucea
tenuisepala resembles N. wilburiana in its small inflorescences and lightly pubescent
sepals; N. wilburiana, though, has coriaceous to subcoriaceous leaves, often with
pulvini at both the base and apex of the petiole, has glabrous filaments, and lacks
staminodes.
A record Azuay fromprovince of Ecuador is suspect (Azuay: carretera

Paute-Guarumales, Sector Amaluisa, Parroquia Palmas, cant?n Paute, colecciones en
Pica Guarumales-M?ndez hasta finca del Sr. Absal?n Bravo, ?rea lluviosa y nublada,
[02?47'S, 78?45'W?], Jaramillo & Winnerskjold 5607A [AAU, MO, NY, QCA]). The
label, with an original identification of Fuchsia (Onagraceae), was clearly placed with
the wrong specimen. The description of the collection on the label (flowers red and 1m
tall) better fits Fuchsia, and the given locality has an elevation of 2500 m or more. Such
a high elevation record for this species of Neosprucea is not expected. The labeling
error is unfortunate, because this collection is unusual for the species in its ovate and
pubescent leaves. Additional collections may prove it to represent an undescribed
species.
The specific epithet refers to the thin (Latin tenuis) sepals.
9. Neosprucea wilburiana M. H. Alford, sp. nov.?Type: Panama. Col?n: Lower Rio

Guanche, 18 Apr 1973 (fl), R. L. Dressier 4341 (holotype: MO-3604291!; iso
types: MO-3604290! US-2!).
Arbor parva. Folia cori?cea, oblonga vel anguste elliptica, ad basim acuta vel rotun
data, margine glanduloso-serrato, venatione acrodroma. Inflorescentia axillaris congesta,
<4.5 cm longa sub anthesi vel postea; s?pala et p?tala dense tomentoso-sericea in pagina
abaxiali. A N. grandiflora absentia staminodiorum, inflorescentiis congestis et floribus mi
noribus differt.
Treelets or trees 1.5-10 m tall. Trunk to unknown dbh; bark unknown. Branches ini
tially brown, becoming gray with growth, obscurely to evidently 3-4-sided, puberulent to
strigillose to glabrescent; lenticels common, reddish brown, circular to elliptic. Leaf
blades 6.4-22.7 cm long, 2.3-8.7 cm wide, narrowly elliptic, oblong, or narrowly obovate,
rarely asymmetrical with respect to the midvein; apex abruptly acuminate to caudate for
up to 3.5 cm (rarely obtuse, N=l, Bristan 501); base acute to rounded, eglandular; margin
serrate-crenate, 0.5-2 teeth/cm, teeth obtuse to rounded to inconspicuous, with a glabrous
gland at or near the apex, gland abaxial and abutting the leaf margin; subcoriaceous to co
riaceous; adaxial surface glabrous, sometimes with a few hairs at lamina base, drying
gray-green; abaxial surface mostly glabrous, sparsely strigillose along major veins, dry
ing dull green; venation acrodromous, 3-veined from base to apex, 0-3 (-4) distal pairs of
secondary veins (or 0-3 [-4] secondary veins per side of the lamina) arising at an acute
angle from the midvein near apex, prominent tertiaries (16-) 25-52, mostly perpendicu
lar to the midvein and the secondaries and parallel with each other, veins raised abaxially.
Petiole 0.8-2.3 cm long, moderately strigillose, puberulent, or pubescent (in Colombian
specimens), sometimes adaxially canaliculate; solid pulvini at base and apex present. Stip
ules 1.5-2.5 mm long, 0.2-0.6 mm wide, narrowly triangular to lanceolate, entire, densely
strigillose, caducous. Inflorescences axillary racemes, sometimes so congested as to ap
pear like fascicles, with 1-5 (-13) flowers, up to 4.5 cm long, often less than 2 cm long,
axis up to 1.6 mm thick, (moderately to) densely pubescent; bracts 2.1-2.5 mm long,
0.5-1.7 mm wide, narrowly triangular to ovate, densely golden-pubescent, caducous;
bracteoles 1mm long, 0.5 mm wide, triangular, located ca. 0.5 mm below the articulation,
densely pubescent abaxially, scarcely so adaxially, caducous; pedicels 2-8.5 mm long,
densely pubescent. Flower buds ovoid to pyramidal, light green, sparsely to densely
golden-pubescent. Sepals 4, 3.5-11 mm long, 1.5-7.0 mm wide, up to 0.6 mm thick,
lanceolate, ovate, or triangular, pale green, abaxially sparsely to densely pubescent, adax
ially densely pubescent to pubescent in lines or pubescent only along the margins, dis
tinctly 3 (-5)-veined in less densely pubescent forms. Petals 4, 3.5-11.5 mm long, 1.5-4.7
mm wide, lanceolate to ovate, creamy white to white, densely pubescent abaxially, pu
bescent in lines or pubescent only along margin or apex adaxially. Receptacle moderately
to densely pubescent. Androecium of numerous stamens, ca. 30; filaments 1.0-1.4 mm
long, glabrous; anthers 2.5-4.3 mm long, densely hirsute on the adaxial surface of the con
nective. Staminodes absent. Ovary ca. 2 mm in diameter, spheroid to ovoid, glabrous to
pubescent, especially near apex; placentation highly intruded parietal, placentae 4; style
3.2-4.1 mm long, glabrous or pubescent; stigma minutely lobed. Fruit a berry, 0.85-1 cm
in diameter, spheroidal, green, glabrous or moderately pubescent, especially near apex.
Seeds several, 3.6-4.3 mm long, 1.7-2.1 mm wide, irregularly ellipsoidal. Fig. 22.
Phenology. Flowers and fruits from February until mid-August.
2008 NEOSPRUCEA 53
FIG. 22. Neosprucea wilburiana. A. Branch with flowers and buds, showing the small, congested inflo
rescences in the axils of the leaves. B. Petiole and abaxial surface of leaf, showing pulvini on the petiole and a
detail of a torus-shaped gland at a leaf tooth; an inflorescence axis with aborted flowers is shown in the leaf axil.
C. Flower bud. D. Flower, side view, showing lack of staminodes. E. Stamens, with detail of apical pores. F. Pis
til. G. Fruit, subtended by persistent perianth. (Based on: A-F, Correa 4341; G, Seibert 465.)
FIG. 23. Distribution of Neosprucea wilburiana.
Distribution (Fig. 23). Panama and Colombia; lowland tropical rainforest; 380-870
m. These regions experience a short dry season from December to April (Portig 1976).
Additional Specimens Examined. Colombia. [Locality and date unknown, probably near Mariquita or
Bogot?, 1760-1808, see Mutis (1957-1958) and Blanco y Fern?ndez de Caleya & del Valle Stervinou (1991)],
Mutis 2232 (US).?Antioquia: Puerto Berr?o, Vereda Alicante, en la v?a San Juan de Bedout-La Cabana, a lo
largo de quebrada Penjamo, finca Penjamo, 06?39'N, 74?32'W, Callejas et al. 9259 (MO, NY); Puerto Berr?o,
carretera Cisneros-Puerto Berr?o, sitio La Carlota, Quebrada El Vapor, 06o37TS?, 74?27'W, Cardona et al. 1053
(MO); Remedios, corregimiento de Santa Isabel, 20-30 km de Remedios, en la v?a a Puerto Berr?o, 06?58/N,
74?40'W, Callejas et al. 8135 (HUA); Remedios, corregimiento Santa Isabel, 6 km de Santa Isabel, en la v?a a
Remedios, O?^STS?, 74?50'W, Callejas et al. 8041 (HUA, MO, NY). Panama. Cocl?: El Valle de Ant?n at
the foot of Cerro Pil?n, [08?37.5'N, 80?07/W], Dwyer & Correa 7997 (BR [photo], MO); near sawmill above
El Cop?, Atlantic drainage east of sawmill, [08?40'N, SO^'W], Hammel 3519 (L, MO); El Valle de Ant?n and
vicinity, [08?36'N, 80*06^], Seibert 465 (F,MO).?Col?n: small tributary of R?o Boquer?n, above Peluca, 26
km from Transisthmian Hwy, [09?25'N, 79?33'W], Dressier 4339 (MO, US); forest along lower part of Rio
Guanche, [09o3(rN, 79?40'W], Foster 2781 (F); Santa Rita ridge, 4 hour walk from end of road, [(t?^'N,
79?47'W ?], Hammel 6320 (MO); Upper R?o Piedras headwaters, along trail from end of Santa Rita Ridge Road,
ca. 11 km SW of Cerro Braja, 09?25lSi, 79?35'W, Sytsma et al. 4262 (MO).?Dari?n: Cerro Pirre, [07?56'N,
77?43'W], Bristan 501 (MO).
Neosprucea wilburiana can be easily recognized by the combination of hirsute an

thers and lack of staminodes. Its inflorescences are the smallest in the genus, up to 4.5 cm
long but often much shorter. The adaxial surface of the leaves dries gray, the number of
prominent tertiary veins is often many ([16-] 25-52; no more than 41 recorded for the
other species), and many individuals have well-developed pulvini at both the base and
apex of the petiole. Smaller inflorescences and more evident pulvini are characteristic of
the Panamanian collections, whereas the Colombian specimens have less congested inflo
rescences and obscure pulvini. Perhaps they represent two different species, but I interpret
them as representing continuous variation within a single species. Further collections from
Colombia are needed to elaborate on the variation.
2008 NEOSPRUCEA 55
Neosprucea wilburiana is the species referred to as Neosprucea aff. sararensis Cua

trec. in Robyns's treatment (1968) of Flacourtiaceae for Flora of Panama.
The specific epithet honors the botanist Robert L. Wilbur (b. 1925), professor of
botany and curator of the herbarium at Duke University, and prolific collector of Central
American plants.
Excluded Name
Neosprucea kuhlmannii Sleumer, Lilloa 23: 248. 1950.?Type: Brazil. Espirito Santo:
Colatina, Rio Doce, 30 Nov 1943, J. G. Kuhlmann 6559 (holotype: RB [photo!];
= Macrothumia kuhlmannii (Sleumer) M. H. Alford, No von 16:
isotype: LIL)
296. 2006.?This species was transferred toMacrothumia M. H. Alford, a genus
closely allied to Ahernia Merr., Pleuranthodendron, and Hasseltia (Alford
2006a, b). It differs from Neosprucea in having papillate (violoid) leaf teeth, de
ciduous perianth, globose anthers, and receptacular disk glands.
ACKNOWLEDGMENTS
The author thanks Melissa Luckow, Sue Zmarzty, Jerrold Davis, and Jeff Doyle for
helpful discussions and thoughtful comments on the project or manuscript; Sandra
Borgardt, Mark Chase, Carol Daugherty, Jane Doyle, Elizabeth Hermsen, Kathie
Hodge, Amanda Ingram, Richard Jobson, Damon Little, Miao Liu, Marcela Mart?nez
Mill?n, Dominick Paolillo, and Jennifer Svitko for laboratory assistance; E. Earl Alford,
David Barrington, Cesar Gr?ndez, Carl Lewis, Sumedha Mahadeo, Am?rico
Manuyama, Victor Medina Ib??ez, Fabi?n Michelangeli, Nelson Miranda, Sigrun Moe,
David Neill, Eric Rodr?guez Rodriguez, and Jackeline Salazar for field assistance; Ed
Cope, Robert Dirig, William Dress, Peter Fraissinet, Alejandra Gandolfo, Robert Kiger,
and Tod Stuessy for herbarium, bibliographic, or translation assistance; and Bobbi An
gel? for the excellent illustrations. I also thank the herbarium curators of A, AAU,
AMAZ, BH, BM, BR, BRIT, C, CAS, COL, DS, DUKE, F, FDG, G, GB, GH, GOET,
HAO, HUA, HUT,
IBE, INPA, JBSD, K, KW, L, LL, LY, M, MA, MG, MICH, MO,
MOL, MSC, NY, OS, P, QCA, QCNE, S, TEX, U, US, USM, USMS, VEN, W, andWU
for loans, digital images, or hospitality during visits. Field work was greatly facilitated
by the Jatun Sacha Biological Station (Fundaci?n Jatun Sacha), Bosque Protector Cerro
Blanco, R?o Palenque Science Center (Fundaci?n Wong), and the Iwokrama Centre for
Rain Forest Conservation & Development. The Ministerio del Ambiente del Ecuador
(N? 007-IC-FLO-DBAP/MA), the Environmental Projection Agency of Guyana (N?
080503 BR 001), and the Instituto Nacional de Recursos Naturales del Per? (N? 14 C/C
2002-INRENA-DGANP and N? 050-2002-INRENA-DGFFS-DCB) are thanked for
collection permits. The author especially thanks the Harvey Fellows Program of the
Mustard Seed Foundation, the Andrew Mellon Foundation, the Graduate School of Cor
nell University, and the Harold F. Moore,
Jr., Fund of the L. H. Bailey Hortorium, which
provided funding for this study. This work was part of a dissertation submitted in partial
fulfillment of the requirements for the Ph.D. degree in plant biology at Cornell
University.
LITERATURECITED
Alford, M. H. 2003. Claves para los g?neros de Flacourtiaceae de Per? y del Nuevo Mundo. Arnaldoa 10(2):
19-38.
_2005. Systematic studies in Flacourtiaceae. Cornell University, Ithaca, NY. Ph.D. Dissertation. 290 pp.
_2006a. Nomenclatural innovations in neotropical Salicaceae. No von 16: 293-298.
_2006b. A new species o? Hasseltia (Salicaceae) from Costa Rica and Panama. Brittonia 58: 277-284.
Alford, M. H., C. Gr?ndez R., and R. V?squez M. 2005 ["2004"]. Una nueva especie de Neosprucea (Sali
caceae/Flacourtiaceae) de Per? y Colombia. Arnaldoa 11: 37-42.
Anderson, C. 1997. Monograph of Stigmaphyllon (Malpighiaceae). Syst. Bot. Monogr. 51: 1-313.
Anderson, W. R. 1981. The botany of the Guayana Highland-Part XL Malpighiaceae. Mem. New York Bot.
Gard. 32: 21-305.
Angiosperm Phylogeny Group (APG). 2003. An update of the Angiosperm Phylogeny Group classification for
the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436.
Baker-Brosh, K. F., and R. K. Peet. 1997. The ecological significance of lobed and toothed leaves in temperate
forest trees. Ecology 78: 1250-1255.
Bentham, G. 1861. Notes on Bixaceae
and Samydaceae. J. Linn. Soc, Bot. 5 (Suppl. 2): 75-94.
Bernard, 1999. A new geography
D. M. of Guyana. London: Macmillan.
Blanco y Fern?ndez de Caleya, P., and A. del Valle Stervinou. 1991. Herbarium Mutisianum seu catalogus plan
tarum a Novagranatensi Regia Legatione, sub Iosephi Caelestini Mutisii auctoritate lectarum. Adduntur
aliae ab ipso lectae. Fontqueria 32: 1-173.
Burnham, R. J., N. C. Pitman, K. Johnson, and P. Wilf. 2001. Habitat-related error in estimating temperatures
from leaf margins in a humid tropical forest. Amer. J. Bot. 88: 1096-1102.
Calder?n, E. 1997. Neosprucea sararensis. In 2003 IUCN Red List of threatened species (IUCN Species Sur
vival Commission, ed.). International Union for Conservationof Nature and Natural Resources, Cam
bridge, U. K.
Chase, M. W., S. Zmarzty, M. D. Lled?, K. J.Wurdack, S. M. Swensen, and M. F. Fay. 2002. When in doubt,
put it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew
Bull. 57: 141-181.
Clarke, H. D., V. A. Funk, and T. Hollowell. 2001. Plant diversity of the Iwokrama Forest, Guyana. Sida, Bot.
Misc. 21: 1-86.
Cronquist, A. 1981. An integrated system of classification of flowering plants. New York: Columbia University
Press.
_1988. The evolution and classification of flowering plants, 2d ed. Bronx, New York: New York Botani
cal Garden.
Cuatrecasas, J. 1955. Taxonomic notes on neotropical trees. Trop. Woods 101: 10-28.
Curtis, J. D., and N. R. Lersten. 1974. Morphology, seasonal variation, and function of resin glands on buds and
leaves of Populus deltoides (Salicaceae). Amer. J. Bot. 61: 835-845.
_1978. Heterophylly in Populus grandidentata (Salicaceae) with emphasis on resin glands and extraflo
ral nectaries. Amer. J. Bot. 65: 1003-1010.
_1980. Morphology and anatomy of resin glands in Salix lucida (Salicaceae). Amer. J. Bot. 67:
1289-1296.
Davis, C.C.,W,R. Anderson, and M. J. Donoghue. 2001. Phylogeny of Malpighiaceae: evidence from chloro
plast ndhF and trnL-F nucleotide sequences. Amer. J. Bot. 88: 1830-1846.
Davis, J. L, and K. C. Nixon. 1992. Populations, genetic variation, and the delimitation of phylogenetic species.
Syst. Biol. 41:421-435.
de Wilde, W. J. J. O. 1971a. The systematic position of tribe Paropsieae, in particular the genus Ancistrothyrsus,
and a key to the genera of Passifloraceae. Blumea 19: 99-104.
-1971b. A monograph of the genus Adenia Forsk. (Passifloraceae). Meded. Wa
Landbouwhoogeschool
geningen 71-18: 1-281.
Edwards, J., and J. R. Jordan. 1992. Reversible anther opening in Lilium philadelphicum (Liliaceae): a possible
means of enhancing male fitness. Amer. J. Bot. 79: 144-148.
Endress, P. K. 1994. Diversity and evolutionary biology of tropical flowers. Cambridge: Cambridge University
Press.
Farris, J. S., V. A. Albert, M. K?llersj?, D. Lipscomb, and A. G. Kluge. 1996. Parsimony jackknifing outper
forms neighbor-joining. Cladistics 12: 99-124.
Flenley, J. R. 1979. The equatorial rainforest: a geological history. London: Butterworth.
2008 NEOSPRUCEA 57
Friedman, J., D. Bolotin, M. Rios, P. Mendosa, Y. Cohen, and M. J. Balick. 1993. A novel method for identifi
cation and domestication of indigenous useful plants in Amazonian Ecuador. In New Crops, ed. J. Janick,
and J. E. Simon, 167-174. New York: Wiley.
Garcia, M. T. A., and P. S. Hoc. 1997. Floral biology and reproductive system of Passiflora caerulea (Passiflo
raceae). Beitr. Biol. Pflanzen 70: 1-20.
Gilg, E. 1925. Flacourtiaceae. In Die nat?rlichen Pflanzenfamilien, 2d ed., ed. A. Engler, 21: 377-457. Berlin:
Duncker & Humblot.
Goloboff, P. 1993. NONA. Tucum?n, Argentina, www.cladistics.com.
Grill, I. 1990. Beitr?ge zur Karyosystematik der Flacourtiaceae. Universit?t Wien. Dissertation. 55 pp.
Hansen, A. K., L. K. Escobar, L. E. Gilbert, and R. K. Jansen. 2007. Paternal, maternal, and biparental inheri
tance of the chloroplast genome in Passiflora (Passifloraceae): implications for phylogenetic studies.
Amer. J. Bot. 94: 42-46.
Harris, J. G., and M. W. Harris. 1994. Plant identification terminology: an illustrated glossary. Spring Lake,
Utah: Spring Lake Publishing.
Hekking, W. H. A. 1988. Violaceae:
Part l-Rinorea and Rinoreocarpus. Fl. Neotrop. 46: 1-208.
Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index herbariorum. Part I: The herbaria of the world,
8th ed. Regnum Veg. 120: 1-693.
Huang, S.-Q., Y Takahashi, and A. Dafni. 2002. Why does the flower stalk of Pulsatilla cernua (Ranunculaceae)
bend during anthesis? Amer. J. Bot. 89: 1599-1603.
Hutchinson, J. 1967. The genera of flowering plants. Oxford: Clarendon Press.
Keating, R. C. 1973. Pollen morphology and relationships of the Flacourtiaceae. Ann. Missouri Bot. Gard. 60:
273-305.
Kowalski, E. A., and D. L. Dilcher. 2003. Warmer paleotemperatures for terrestrial ecosystems. Proc. Nati.
Acad. Sei. U.S.A. 100: 167-170.
Larson, P. R., and R. E. Dickson. 1973. Distribution of imported 14C in developing leaves of eastern cottonwood
according to phyllotaxy. Planta 111: 95-112.

Larson, P. R., and J. C. Gordon. 1969. Leaf development, photosynthesis and 14C distribution in Populus del
toides seedlings. Amer. J. Bot. 56: 1058-1066.
Larson, P. R., J. G. Isebrands, and R. E. Dickson. 1972. Fixation patterns of 14Cwithin developing leaves of east
ern cottonwood. Planta 107: 301-314.
Lemke, D. E. 1988. A synopsis of Flacourtiaceae. Aliso 12: 29^3.
Little, Jr., E. L. 1948. New species of trees from western Ecuador. J.Wash. Acad. Sei. 38(3): 87-105.
MacDougal, J. M. 1994. Revision of Passiflora subgenus Decaloba section Pseudodysosmia

(Passifloraceae).
Syst. Bot. Monogr. 41: 1-146.
Metcalfe, C. R., and L. Chalk. 1950. Anatomy of the dicotyledons, vol. 2. Oxford: Clarendon Press.
Miller, R. B. 1975. Systematic anatomy of the xylem and comments on the relationships of Flacourtiaceae. J.
Arnold Arbor. 56: 20-102.
Morawetz, W. 1981. Zur systematischen Stellung der Gattung Prockia: Karyologie und Epidermisskulptur im
Vergleich zu Flacourtia (Flacourtiaceae), Grewia (Tiliaceae) und verwandten Gattungen. PL Syst. Evol.
139: 57-76.
Mori, S. A., G. Cremers, C. A. Gracie, J.-J. de Granville, S. V Heald, M. Hoff, and J. D. Mitchell. 2002. Guide
to the vascular plants of central French Guiana. Part 2. Dicotyledons. New York: New York Botanical
Garden Press.
Mutis, J. C. 1957-1958. Diario de observaciones de Jos? Celestino Mutis (1760-1790). Bogot?: Editorial Min
erva.
Neill, D. A., and P.M. J0rgensen. 1999. Climates. In Catalogue of the vascular plants of Ecuador, ed. P.M. J0r
gensen and S. Le?n-Y?nez, 8-13. St. Louis: Missouri Botanical Garden Press.
Nixon, K. C. 1999. The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics 15: 407-414.
_2002. WinClada, version 1.00.08. Ithaca, New York, www.cladistics.com.
Nixon, K. C, and Q. D. Wheeler. 1990. An amplification of the phylogenetic species concept. Cladistics 6:
211-223.
Olmstead, R. G., and J. A. Sweere. 1994. Combining data in phylogenetic systematics: an empirical approach
using three molecular data sets in the Solanaceae. Syst. Biol. 43: 467^181.
P??bo, S. 1990. Amplifying ancient DNA. In PCR protocols: a guide to methods and applications, ed. M. A.
Innis, D. H. Gelfand, J. J. Sninsky, and T. J.White, 159-166. San Diego: Academic Press.
Patterson, C. 1982. Morphological characters and homology. In Problems of phylogenetic reconstruction, ed. K.
A. Joysey and A. E. Friday, 21-74. London: Academic Press.
Portig, W. H. 1976. The climate of Central America. In World survey of climatology: climates of Central and
South America, ed. W. Schwerdtfeger, 405-478. Amsterdam: Elsevier.
Pourrut, P., O. R?vere, I. Romo, and H. Villacr?s. 1995. Clima del Ecuador. In El agua en el Ecuador: clima,
precipitaciones, escorrent?a, ed. P. Pourrut, 13-26. Quito.
Radford, A. E., W. C. Dickison, J. R. Massey, and C. R. Bell. 1974. Vascular plant systematics. New York:
Harper & Row.

Robyns, A. 1968. Hora of Panama: Family 128. Flacourtiaceae. Ann. Missouri Bot. Gard. 55: 93-144.
Roy er, D. L., and P.Wilf. 2006. Why do toothed leaves correlate with cold climates? Gas exchange at leaf mar
gins provides new insights into a classic paleotemperature proxy. Int. J. Plant Sei. 167: 11-18.
Seaward, M. R. D., and S. M. D. FitzGerald, eds. 1996. Richard Spruce (1817-1893): botanist and explorer.
Kew: Royal Botanic Gardens.
Shore, J. S., K. L. McQueen, and S. H. Little. 1994. Inheritance of plastid DNA in the Turnera ulmifolia com
plex (Turneraceae). Amer. J. Bot. 81: 1636-1639.

Simmons, M. P., and H. Ochoterena. 2000. Gaps as characters in sequence-based phylogenetic analyses. Syst.
Biol. 49: 369-381.
Sleumer, H. O. 1936. ?ber die neue Flacourtiaceengattung Spruceanthus. Notizbl. Bot. Gart. Berlin-Dahlem 13:
362-263.
_1938. Die Gliederung der Flacourtiaceae-Prockiinae Gilg. Notizbl. Bot. Gart. Berlin-Dahlem 14: 45-52.
_1980. Flacourtiaceae. Fl. Neotrop. 22: 1-499.
Snow, J.W. 1976. The climate of northern
South America. In World survey of climates: climates of Central and
South America, ed. W. Schwerdtfeger, 295-403. Amsterdam: Elsevier.
Soltis, D. E., and P. S. Soltis. 1998. Choosing an approach and an appropriate gene for phylogenetic analysis. In
Molecular systematics of plants II: DNA sequencing, ed. D. E. Soltis, P. S. Soltis, and J. J. Doyle, 1-42.
Boston: Kluwer Academic Publishers.
Spencer, K. C, and D. S. Seigler. 1985. Cyanogenic glycosides and the systematics of the Flacourtiaceae.
Biochem. Syst. Ecol. 13: 421-431.
Spruce, R. 1908. Notes of a botanist on the Amazon and Andes. London: Macmillan.
Stearn, W. T. 1992. Botanical Latin: history, grammar, syntax, terminology and vocabulary, 4th ed. Portland,
Oregon: Timber Press.

Stevens, P. F. 1991. Character states, morphological variation, and phylogenetic analysis: a review. Syst. Bot. 16:
553-583.
Taberlet, P., L. Gielly, G. Pautou, and J. Bouvet. 1991. Universal primers for amplification of three non-coding
regions of chloroplast DNA. PL Molec. Biol. 17: 1105-1109.
Thompson, J. D., T. J. Gibson, R. Plewniak, F. Jeanmougin, and D. G. Higgins. 1997. The ClustalX windows
interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucl. Acids
Res. 25: 4876-4882.
Tosi Jr., J. A. 1960. Zonas de vida natural en el Per?: memoria explicativa sobre el mapa ecol?gico del Per?.
Lima: Instituto Interamericano de Ciencias Agr?colas de la OEA.
Trelease, W. 1881. The foliar nectar glands of Populus. Bot. Gaz. 6: 284-290.
van der Hammen, T. 1974. The Pleistocene changes of vegetation and climate in tropical South America. J. Bio
geogr. 1: 3-26.
Weberling, F. 1989. Morphology of flowers and inflorescences. Cambridge: Cambridge University Press.
Wessel, P., and W. H. F. Smith. 1995. The Generic Mapping Tools (GMT), version 3.0. SOEST/NOAA.
Wilkinson, H. P. 2007. Leaf teeth in certain Salicaceae and 'Flacourtiaceae.' Bot. J. Linn. Soc. 155: 241-256.
APPENDIX
GenBank accession numbers and herbarium vouchers for DNA used in the phylogenetic analysis of mol
ecular data for Neosprucea and relatives. Information is presented in the following order: species, herbarium
voucher, GenBank accession numbers for ndhF, trnL, and trnL-F. Herbarium acronyms follow Index herbario
rum (Holmgren et al. 1990). Sequences deposited at GenBank may be found at www.ncbi.nlm.nih.gov.
Abatia M. H. Alford Hatschbach 23260 -

angeliana (=Aphaerema spicata Miers): (BH) AY757093, ?,
AY756985
Abatia canescens Sleumer: Alford & Miranda 3082 -
(BH) AY757087, AY756974, AY756980
Abatia mexicana Standl.: Panero & Calzada 5543A -
(NY) AY757088, AY756975, AY756981
2008 NEOSPRUCEA 59
Abatia parviflora Ruiz & Pav.: Barrington 2104 (BH) - AY757089, AY756976, AY756982
-
Abatia spicata (Turcz.) Sleumer: Alford 3125 (BH) AY757090, AY756977, AY756983
-
Abatia stellata Lillo: Grifo 688 (BH) AY757091, AY756978, AY756984
Ahernia glandulosa Merr.: Liede 3289 (K) - AY757092, ?, ?
- AY756986
Azara alpina Poepp. & Endl.: Seijo 1729 (NY) AY757094, AY756931,
-
Azara celastrina D. Don: Chase 3889 (K) AY757095, AY756932, AY756987
- ?
Azara lanceolata Hook, f.: Alford 3171 (BH) AY757096, AY756933,
-
Azara salicifolia Griseb.: Villamil 3474 (NY) AY757097, AY756934, AY756988
Ba?ara axilliflora Sleumer: Alford & Manuyama 3131 (BH) - AY757098, AY756935, AY756989
-
Ba?ara domingensis Benth.: Solazar 2370 (BH) AY757099, AY756936, AY756990/AY756991
Ba?ara guianensis Aubl.: Michelangeli & Alford 704 (BH) - AY757100, AY756937, AY756992
?anara minutiflora -
(A. Rieh.) Sleumer: A//ow/ & Lewis 3030 (BH) AY757102, AY756938, AY756994
Ba?ara nitida Spruce ex Benth.: Alford 3105 (BH) - AY757103, AY756939, AY756995
Ba?ara regia Sandwith: Alford & Miranda 3088 (BH) - AY757104, ?, AY756997
Ba?ara tomentosa Clos: Alford 3175 (BH) - AY757105, AY756940, AY756998
Ba?ara umbraticola Arechav.: Vodicka 858 (BH) - AY757106, AY756941, AY756999
-
Ba?ara vanderbiltii Urb.: Alford & Lewis 3025 (BH) AY757107, AY756942, AY757000
Hasseltia allenii M. H. Alford: A//ow/ 3023 (BH) - AY757132, AY756914, AY757022
Hasseltia floribunda H.B.K.: A//o?/ 2990 (BH) - AY757133, AY756915, AY757023
-
Hasseltia guatemalensis Warb.: A//wrf 3006 (BH) AY757134, AY756916, AY757024
Hasseltiopsis dioica (Benth.) Sleumer: Alford 3010 (BH) - AY757136, AY756947, AY757026
racemosum -
Homalium Jacq.: Sa/azar 2410 (BH) AY757139, AY756949, AY757029
et al. 6689 -
Macrothumia kuhlmannii (Sleumer) M. H. Alford: Carvalho (MO) AY757101, AY756912,
AY756993
ex Benth.) Sleumer: Romoleroux 2259 -
Neosprucea grandiflora (Spruce (MO) AY757156, AY756953,
AY757041
montana -
Neosprucea Cuatrec: Alford 3129 (BH) AY757158, AY756954, AY757042
Neosprucea pedicellata Little: Alford & Miranda 3076 (BH) - AY757160, AY756956, AY757044
Neosprucea melastomatoides M. H. Alford: Alford et al. 3091 (BH) - AY757161, AY756957, AY757045
-
Neosprucea paterna M. H. Alford: Alford et al. 3149 (BH) AY757159, AY756955, AY757043
Pineda incana Ruiz & Pav.: Alford 3124 (BH) - AY757167, AY756959, AY757049
Pineda ovata M. H. Alford & Zmarzty: Wood 18684 (K) - AY757168, ?, AY757050
Pleuranthodendron lindenii (Turcz.) Sleumer: Alford 2989 (BH) - AY757169, AY756919, AY757051
Prockia costaricensis Standl.: Alford 3018 (BH) - AY757174, AY756960, AY757056
Prockia crucis L.: Alford & Manuyama 3132 (BH) - AY757175, AY756961, AY757057
Prockia flava H. Karst.: Michelangeli & Alford 677 (BH) - AY757177, AY756962, AY757058
Prockia pentamera A. H. Gentry: Alford et al. 3130 (BH) - AY757178, AY756963, AY757059
-
Scolopia spinosa (Roxb.) Warb.: Chase 1288 (K) AY757190, AY756967, AY757072
-
Xylosma bahamense (Britton) Standl.: Alford & Lewis 3031 (BH) AY757198, AY756968, AY757078.
NUMERICALLIST OF SPECIES
1.N. grandiflora 6. N. rimachii

2. N. melastomatoides 1. N. sararensis
3. N. montana 8. N. tenuisepala
4. N. paterna 9. N. wilburiana
5. N. pedicellata
INDEXTO NUMBERED COLLECTIONSEXAMINED
The numbers in parentheses refer to the corresponding species in the text and in the Numerical List of
Species above.
Alford,M. H. 3092 (2), 3103 (1), 3129 (3). Freir?, E., et al. 2312 (2), 2775 (1).
Alford, M. H., et al. 3076 (5), 3091 (2), 3133 (4), 3148 Gentry, A. 12001 (5).
(4), 3149 (4). Gentry, A.; & Smith, D. N. 45255 (3).
Alvarado, A. 271 (1). Gentry, A., et al. 12167 (5), 27848 (1), 60179 (1),
Alvarez, A., ?tal. 2379(1). 64365 (1).
Ancuash,E. 1156(8). Grijalva, A., ?tal. 623(1).
Asplund, E. 9054 (2). Gudi?o, E. 451 (1), 1194 (1), 1683 (1).
Aulestia, M., & Andi, N. 795 (1). Gudi?o, E., & Andi, N. 2090 (1).
Aulestia, M., & Grefa, G. 162 (1). Gudi?o, E., et al. 1035 (1), 1754 (1), 1825 (1), 1853
Aulestia, M., et al. 454 (1), 460 (1). (1), 2194(1).
Bennett, B. C, et al. 4091 (8). Hammel, B. 3519 (9), 6320 (9).
Bensman, R. 308
(1). Harling, G. & Andersson, L. 11911 (1), 12871 (8).
Boyle, B., et al. 3688 (5), 3689 (5). Holm-Nielsen, L. B. 39836 (1).
Brandbyge, J., & Asanza C, E. 32322 (8). Homeier, J. 257 (1).

Brandbyge, J., et al. 30477 (2), 33440 (1). Homeier, J., et al. 52(1).
Bristan, N. 501 (9). Huashicat, V. 376 (8), 586 (8), 848 (8), 1166 (8), 1338
Callejas, R., et al. 4283 (3), 8041 (9), 8135 (9), 9259 (8), 1572 (8), 2304 (8).
(9). Hurtado,F 653 (1), 683 (1), 853 (1).
Campos, J., et al. 3028 (3), 5474 (3). Hurtado, F, et al. 5 (1), 535 (1), 2753 (1).
Cardona, F., et al. 1053 (9). Jaramillo, J. 4052 (2), 7002 (5), 7003 (5), 7558 (5),
Cazalet, P. C. D., & Pennington, T. D. 5046 (5), 5183 14252 (8), 14453 (8).
(5). Jaramillo, J., & Coello, F. 3743 (2).
Cer?n,C. E. 587 (2), 642 (2), 3590 (2), 7693 (1), 7770 Jaramillo, J., & Grijalva, E. 12903 (1).
(1), 14288 (8). Jaramillo, J., & Winnerskjold, V. 5607 (8).
Cer?n, C. E., & Benavides, G. 6172 (5), 14114 (1), Jaramillo-Mejia., R., et al. 3572 (7).
14126 (8). Jativa, C, & Epling, C. 1062 (5), 1102 (5).
Cer?n, C. E., & Cer?n, M. 3151 (1), 4552 (2), 4650 Kayap, R. 844 (8), 1142 (8), 1291 (8), 1478 (8).
(2). Korning, J., & Thomsen, K. 47468 (1).
Cer?n, C. E., & Hurtado, F. 6677 (1). Leo Luna, M. 326 (3).
Cer?n, C. E., & Iguago, C. 5347 (2), 5384 (2). Little, Jr., E. L. 6152 (5).
Cer?n, C. E., & Montesdeoca, M. 8008 (2). Lugo S., H. 328 (1), 343 (1), 2081 (2), 3209 (1).
Clark, J. L., & Mailloux, M. 7852 (1). Macia, M. J., et al. 1263 (1), 3828 (1).
Clark, J.L., et al. 5193(1). McDaniel, S., & Rimachi Y, M. 20402 (6).
Cuatrecasas, J. 11147
(1), 13400 (7). McDowell, T., et al. 2103 (4).
Cuatrecasas, J., et al. 12581 (3). Miller, J. S., et al. 2459(1).
Delinks, T., et al. 284 (2). Mutis, J.C. 1025 (7), 2211 (7), 2232 (9).
D?az, C, et al. 10356 (3), 10605 (1). Neill, D., & Maries, R. 7010 (1).
Dik,A. 300(1). Neill, D., et al. 6384 (1), 8588 (1).
Dodson, C. H., & Gentry, A. 6318 (5), 10066 (5). 011gaard,B., et al. 57040(1).
Dodson, C. H., & Tan, K. 5340 (5). Palacios,W 1089 (1), 1390 (2), 1834 (1), 2600 (5),
Dressler, R. L. 4339
(9), 4341 (9). 2980 (2), 3274 (1), 4163 (2), 4792 (1).
Dugand, A., & Jaramillo, R. 3873 (7). Palacios, W, et al. 844 (2), 871 (1), 7669 (1), 8702
Dwyer, J. D., & Correa A., M. D. 7997 (9). (8), 9504 (1).
Espinoza, S., & Coba, T. 394 (1). Pav?n, J. 36(1).
Foster, R., & Balde?n, S. 12812 (6). Pennington, T. D. 10620 (1), 10626 (1).
Foster, R., et al. 7738 (3). Pennington, T. D., et al. 14285 (5), 14300 (5), 14684
Foster, R. B. 2781 (9). (5).
Foster, R. B., & Smith, D. 9165 (3). P?rez-Arbel?ez, E. 752 (1).
Freir?, E., & Cerda, J. 373 (1). Pipoly,J. 16411 (6).
Freir?, E., & Cerda, P. 1 (1), 131 (2). Pitman, N. 481(1).
Freir?, E., & Santi, L. 3288 (1). Pitman, N., & Alvarado, A. 95 (1).
2008 NEOSPRUCEA 61
Ram?rez, J. G., et al. 5613

(1). Ule, E. 6712 (1).
Rodr?guez, E., et al. 1420 (8). van der Werff, H., et al. 16803 (3).
Rojas, R., et al. 180 (8). Vargas, H., et al. 588 (1).
Romero-Casta?eda, R. 5175 (5) Vargas, H., & Cerda, J. 648 (1), 681 (1), 708 (1).
Romoleroux, K., et al. 2259 (1). V?squez, R., & Jaramillo, N. 2742 (1).
Seibert, R. J. 465 (9). V?squez, R., & Quiaco, E. 24554 (8).
Silverstone-Sopkin, P., & Giraldo, J. 6427 (3). V?squez, R., et al. 12176 (1), 22129 (8), 22469 (8),
Smith, D. N. 4429 (3). 22494 (8), 26725 (3), 27535 (3).
Smith, D. N., & V?squez S., S. 4545 (3), 4760 (3). Vodicka, M. R. 859 (5).
Spruce, R. 4897 (1). von Sneidern, K. 1350 (1).
St?hl, B., et al. 4338 (2). Watson, J. B. 172 (5).
Stein, B. A. 3013 (1). Woytkowski, F. 8286 (3).
Steyermark, J. A., & Liesner, R. 118458 (3). Zak, V, & Espinoza, S. 4463 (1), 4688 (1), 4747 (1),
Suin, L., et al. 1973(1). 4963 (1), 5024 (1).
Sytsma, K., et al. 4262 (9). Zaruma, J. 583 (1), 730 (2).
Tipaz, G., et al. 434 (1), 2184 (5). Zaruma, J., & Arguello, A. 511 (1).
Tirado, M. 1914(1). Zuleta, J. 145 (1).
Tunqui, S. 951(8), 1101(8).
INDEXTO SCIENTIFICNAMES
Accepted names are in roman type; the main entry for each is in boldface. Synonyms are in italics.
Abatia Ruiz & Pav. 18 Eperua Aubl. 40
angeliana M. H. Alford 17, 60 Euphorbiaceae 22

canescens Sleumer 17, 60 Flacourtiaceae 1, 2, 3, 7, 11, 12, 21, 22
mexicana Standl.17, 60 Fuchsia 51
parviflora Ruiz & Pav. 17, 60 Hasseltia H.B.K. 18,21,55
spicata (Turcz.) Sleumer 17, 60 allenii M. H. Alford 17, 60
stellataLillol7, 60 floribunda H.B.K. 4, 5, 15, 16, 17, 60
Ahernia Merr. 55 ex Benth.)
grandiflora (Spruce Sleumer 23
glandulosa Merr. 17, 60
guatemalensis Warb. 17, 60
Ampelozizyphus Ducke 22
Hasseltiopsis Sleumer 5, 18
Aphaerema Miers
dioica (Benth.) Sleumer 15, 16, 17, 60
spicata Miers 17, 60
Homalium Jacq.
Azara Ruiz & Pav. 18
racemosum Jacq. 5, 17, 60
alpina Poepp. & Endl. 17, 60 40
Leguminosae
celastrina D. Don 17, 60
Loganiaceae 22
lanceolata Hook. f. 17, 60
Lunania Hook. 21, 22
salicifolia Griseb. 17, 60
Macrothumia M. H. Alford 55
Ba?ara Aubl. 2, 5, 18,21
axilliflora Sleumer
kuhlmannii (Sleumer) M. H. Alford 17, 55, 60
17, 60
Malpighiales 1
domingensis Benth. 17, 60
Melastomataceae 1, 22
grandiflora Spruce ex Benth. 2, 23
Menispermaceae 22
guianensis Aubl. 4, 15, 16, 17, 18, 60
kuhlmannii 17 Neosprucea Sleumer 19-22
(Sleumer) Sleumer
minutiflora (A. Rich.) Sleumer 17, 60 grandiflora (Spruce ex Benth.) Sleumer 3,6,7, 8,9,
n?tida Spruce ex Benth. 17, 60 10, 12, 13, 14, 15, 16, 17, 18, 19, 20, 22,
regia Sandwith 17, 60 23-29, 30, 33, 47, 48, 51, 52, 60, 61
tomentosa Clos 17, 60 kuhlmannii Sleumer 55
umbraticola Arechav. 17, 60 melastomatoides M. H. Alford 6, 7, 9, 10, 11, 12,
vanderbiltii Urb. 17, 60 13, 14, 15, 16, 17, 18, 20, 21, 23, 29, 30-33,
Elaeocarpaceae 1 51,60,61
montana Cuatrec. 6, 9, 10, 11, 12, 15, 16, 17, 18, Pleuranthodendron L. O.Williams 21, 55
19, 20, 21, 22, 23, 33-37, 47, 51, 60, 61 lindenii (Turcz.) Sleumer 17, 60
paterna M. H. Alford 6, 7, 8, 9, 10, 12, 13, 15, 16, Prockia L. 5, 18,21
17, 18, 19, 20, 21, 22, 23, 37-40, 46, 60, 61 costaricensis Standl. 17, 60
pedicellata Little 6, 7, 10,12,15,16,17,18,19, 20, crucis L.15, 16, 17, 60

22, 23, 40-43, 60, 61 flava H. Karst. 17, 60
rimachii M. H. Alford et al. 8, 10, 12, 15, 16, 18, pentamera A. H. Gentry 17, 60
19, 20, 21, 22, 23, 37, 40, 44-46, 61 Prockieae 1, 2, 5
sararensis Cuatrec. 2, 10, 11, 15, 16, 18, 20, 22, 23, Prockiinae 2
36, 40, 46-47, 55, 61 Rhamnaceae 22
sucumbiensis Cuatrec. 3, 24, 29, 33 Salicaceae 1, 7, 8, 10, 11, 12, 18, 21
tenuisepala M. H. Alford 10, 11, 15, 16, 18, 21, 22, Scolopia Schreb.
23, 29, 33, 36, 48-51, 61 spinosa (Roxb.) Warb. 17, 60
wilburiana M. H. Alford 2, 10, 11, 12, 15, 16, 18, Spruceanthus Sleumer
2, 19
19, 20, 21, 22, 23, 36, 47, 51, 52-55, 61 grandiflorus (Spruce ex Benth.) Sleumer 23
Ochnaceae 11 Spruceanthus Verd. 3, 19
Onagraceae 51 Strychnos L. 22
Oxalidales 1 Tiliaceae 1, 3, 22
Passifloraceae 10, 11, 14 Turneraceae 4
Pineda Ruiz & Pav. 5, 18 Violaceae 10
incana Ruiz & Pav. 15, 16, 17, 60 Xylosma G. Forst.
ovata M. H. Alford & Zmarzty 17, 60 bahamense (Britton) Standl. 17, 60

American Society of Plant Taxonomists Is Collaborating With JSTOR To Digitize, Preserve and Extend Access To Systematic Botany Monographs

Uploaded by

Copyright:

Available Formats

You might also like

American Society of Plant Taxonomists Is Collaborating With JSTOR To Digitize, Preserve and Extend Access To Systematic Botany Monographs

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

American Society of Plant Taxonomists Is Collaborating With JSTOR To Digitize, Preserve and Extend Access To Systematic Botany Monographs

Uploaded by

Copyright:

Available Formats

Revision of Neosprucea (Salicaceae)

Author(s): Mac H. Alford

provided; all species, except N. sararensis, are illustrated.

Resumen. El g?nero neotropical Neosprucea (Salicaceae), incluido anteriormente en Flacourtiaceae,

Neosprucea (Salicaceae) is a neotropical genus of nine species of small to medium

The first known collections of Neosprucea were gathered in Colombia ("New

The first collection of Neosprucea to receive detailed

Species Circumscription. Specimens were sorted into morphologically similar enti

versions of that used by many museum taxonomists, exemplified by Cronquist (1988),

Anatomy and Morphology. Wood anatomy of Neosprucea was investigated by cutting

DNA. DNA was extracted from approximately 0.02 g of field-collected, silica-dried

Phylogenetic Matrices. Two phylogenetic matrices were generated: (1) amorpholog

species of Neosprucea were included in the matrix of morphological

Table 1. Variable anatomical features of the wood of Neosprucea.

Length of Length of #-seriate

N. grandiflora simple or 1-6 1-11 1-120 1-3

(Alford3091) simple 1-5 (-7) 1-28 1-21 1-2

(Alford3129) simple 1-5 1-15 (+?) 1-5 (+?) 1-2

(Alford3149) simple 1-8 1-12 1-22 (-30) 1-2

FIG. 1.Wood of Neosprucea.

If a ray is heterogeneous, it consists of short to long series of upright ray parenchyma,

Habit. Species of Neosprucea are small treelets to medium-sized trees, occasionally

FIG. 2. Vegetative and floral morphology of Neosprucea. A. Acrodromous venation of N. melastoma

Inflorescence. The inflorescence of Neosprucea is a determinate thyrse (=thyrsoid),

arrangement has resulted in different interpretations of the pedicel. In Flacourtiaceae (both

Androecium.The androecium consists of numerous stamens ([30-] 60-400), in some

Table 2. Pollen of Neosprucea: variation in shape and size.

N. grandiflora spherical 22-24 20-22

(Campos et al. 3028)

N. paterna subprolate 36-40 28-34

Gynoecium. The gynoecium of Neosprucea consists of one superior paracarpous

1. Leaf venation: pinnate = 0; actinodromous =

N. melastomatoides, N. tenuisepala), characterized by hirsute filaments, well-developed

85? W 80? 75? 70? 65? 55?

FIG. 7. Distribution of Neosprucea.

Treelets or trees to 30 m tall, sometimes reproductively mature when as small as

sometimes more densely pubescent; bracteoles lanceolate to triangular to ovate, some

Neosprucea is distinctive in Salicaceae (and among all former Flacourtiaceae sensu

Key to the Species of Neosprucea

tenuisepala); southern Colombia, Ecuador, and Peru.

through any pubescence. 8. N tenuisepala

Alternative Key to the Species of Neosprucea

usually a raceme; leaves 5-16.5 (-20.1) cm long. 3. N. montana (uncommon variant)

long; pulvini at base and apex of petiole usually present. 9. N. wilburiana

olate-triangular (uncommonly ovate); leaves coriaceous or subcoriaceous, glossy and

1. Neosprucea grandiflora (Spruce ex Bentham) Sleumer, Notizbl. Bot. Gart. Berlin

4897 (holotype: K!; isotypes: B-destroyed, BM! BR-photo! G! KW-photo! P!

Trees, small to large, to 30 m tall. Trunk to 60 cm dbh; bark whitish, smooth.

Additional Specimens Examined. Colombia. Caqueta: Florencia, [01?37'N, 75?37'W], P?rez-Ar

W 80? 75? 70?

FIG. 9. Distribution of Neosprucea grandiflora.

Neosprucea grandiflora has coriaceous to subcoriaceous elliptic to narrowly elliptic

2. Neosprucea melastomatoides M. H. Alford, sp. nov.?Type: Ecuador. Napo:

Arbor parva. Folia elliptica, subtus molliter pubescentia, breviter acuminata, ad

short-tomentose, trichomes sometimes in lines. Receptacle densely pubescent. Androe

FIG. 11. Distribution o? Neosprucea melastomatoides.

3. Neosprucea montana Cuatrecasas, Trop. Woods 101: 20. 1955.?Type: Colombia.