A new species of Henriettea (Melastomataceae) from the Sierra de Baoruco, the Dominican Republic
ABSTRACT Henriettea uniflora, which is known only from a diverse moist montane forest in the vicinity of Loma Trocha de Pey (or “Monteada Nueva”), Loma
Pie de Palo, and Loma Remigio, the easternmost peaks of the Sierra de Baoruco, is described and illustrated. It is compared
to species of the Henriettea squamulosa complex, especially H. squamulosa and H. ciliata. The species of this complex are characterized by an indumentum of ferruginous, stellate-lepidote hairs.
- SourceAvailable from: Frank Almeda[Show abstract] [Hide abstract]
ABSTRACT: With nearly 2,000 species, all restricted to the Neotropics, Miconieae represent the largest radiation within Melastomataceae. Generic limits and relationships within the tribe have long been contested. Our molecular phylogenetic analyses of the tribe, based on sequences of nrITS and ndhF, resulted in consistent support for a clade comprising Bellucia, Loreya, Henriettea, and Henriettella, which is neither nested within nor sister to the Miconieae. Furthermore, and based upon a series of presumably synapomorphic anatomical and morphological characters, e.g. megastyloid crystals, absence of both medullary and cortical vascular bundles, plinerved leaves, axillary or cauliflorous inflorescences, we consider three additional genera, Kirkbridea, Llewelynia, and Myriaspora, to belong to this clade, a newly recognized tribe, Henrietteeae. These seven genera had formerly been assigned to the Miconieae based on their having exappendiculate anthers (except Kirkbridea) and berry fruits. Henriettella and Llewelynia are relegated to synonymy under Henriettea (25 new combinations are made under that genus: Henriettea boliviensis, H. bracteosa, H. caudata, H. duckeana, H. fissanthera, H. glabra, H. cogniauxiana, H. goudotiana, H. heteroneura, H. hondurensis, H. ininiensis, H. lawrancei, H. lundellii, H. maguirei, H. manarae, H. odorata, H. ovata, H. prancei, H. ritnosa, H. steyermarkii, H. tachirensis, H. tobagensis, H. tovarensis, H. trachyphylla, H. williamsii), and Loreya and Myriaspora are synonymized under Bellucia (with nine new combinations under that genus: Bellucia egensis, B. gracilis, B. klugii, B. nigricans, B. ovata, B. riparia, B. strigosa, B. subandina, B. wurdackiana). A description of Henrietteeae along with a key to and brief descriptions of its constituent genera are provided, in addition to necessary new combinations.Systematic Botany 01/2010; 35(4):783-800. · 1.29 Impact Factor
A new species of Henriettea (Melastomataceae) from the Sierra
de Baoruco, the Dominican Republic
WALTER S. JUDD1,4, JAMES D. SKEAN JR.2, DARIN S. PENNEYS1,
FABIAN A. MICHELANGELI3
1Department of Botany, University of Florida, 220 Bartram Hall, P.O. Box 118526, Gainesville,
FL 32611, USA; e-mail: ; e-mail: firstname.lastname@example.org
2Department of Biology, Albion College, Albion, MI 49224, USA; e-mail: email@example.com
3The New York Botanical Garden, Bronx, NY 10458, USA; e-mail: firstname.lastname@example.org
4Author for correspondence; email: email@example.com
Abstract. Henriettea uniflora, which is known only from a diverse moist montane
forest in the vicinity of Loma Trocha de Pey (or “Monteada Nueva”), Loma Pie de
Palo, and Loma Remigio, the easternmost peaks of the Sierra de Baoruco, is described
and illustrated. It is compared to species of the Henriettea squamulosa complex,
especially H. squamulosa and H. ciliata. The species of this complex are character-
ized by an indumentum of ferruginous, stellate-lepidote hairs.
Key Words: Melastomataceae, Henriettea, Dominican Republic, Hispaniola, Sierra
de Baoruco, megastyloids.
In the course of fieldwork in connection
with a phylogenetic analysis of Miconieae
(Melastomataceae), an undescribed species of
Melastomataceae was collected on the Loma
Trocha de Pey, one of the easternmost peaks
of the Sierra de Baoruco, Dominican Republic.
This species has unusual inflorescence and leaf
morphology and was not readily assignable to
a genus. Thus, molecular phylogenetics and
leaf anatomical studies were undertaken in
order to determine its phylogenetic (and thus
generic) placement within the family. The
specimens are referable to Henriettea DC.
(including Henrietella Naudin; see Judd,
1989) on the basis of their leaves with
megastyloid crystals (Baas, 1981), axillary
flowers, petals with a dorso-apical tooth, and
stellate-peltate hairs with a central projection,
interpreted as a reduction of the elongate
multicellular hairs with radiate to stellate-
globular basal portion, which are characteristic
of the genus (Wurdack, 1986; Fig. 135; Judd,
1989). Within this genus, its affinities appear
to be with those species with a ferruginous
stellate-lepidote indumentum, and this hypoth-
esis is supported by a preliminary phylogenet-
ic analysis based on ITS sequences.
The new species, Henriettea uniflora Judd,
Skean, Penneys & Michelangeli is described
below and compared with its putative rela-
tives, especially H. squamulosa (Cogn.) Judd
and H. ciliata (Urb. & Ekman) Alain. These
three species (and their relatives) are remark-
able and easily separated from other Antillean
species of Henriettea because of their ferru-
ginous, stellate-peltate hairs. The group, here
called the H. squamulosa complex, is consid-
ered to be monophyletic based on the
preliminary hypothesis that the distinctive
lepidote indumentum of these species is
Henriettea probably is most closely related
to Llewelynia Pittier and Kirkbridea Wur-
dack, and these three genera may be sister to
a clade containing Loreya DC., Bellucia Raf.,
and Myriaspora DC (Penneys et al., 2004).
Together these genera comprise a monophy-
letic group, which receives support from
morphology (Judd, 1989; Penneys, et al.,
2004; putative synapomorphies: megastyloid
crystals present, anomocytic stomata, exclu-
sively septate fibers, plinervy, axillary or
cauliflorous inflorescences, petals pubescent
or at least with a few hairs near the apex,
Brittonia, 60(3), 2008, pp. 217–227
© 2008, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 30 September 2008
anther appendages short or lacking, inferior
ovaries, and berry fruits) and DNA sequences
(Michelangeli et al., 2004; Penneys et al.,
2004). These genera traditionally have been
placed in the large tribe Miconieae, but recent
molecular analyses support their placement
in a new tribe (Michelangeli et al., 2004;
Penneys et al., 2004), which has not yet
been formally described. The phylogenetic
placement of this clade and relationships
within it are currently under study by the
Methods and Materials
In the description below, plant height and
flower color were taken from information
given on specimen labels or observed in the
field; floral measurements were taken from
liquid-preserved or rehydrated material; other
measurements were taken from dried herbar-
Stem, leaf, and petiolar anatomy were
assessed in material of Skean 4298 preserved
in 70% ethanol using the phloroglucinol-
hydrochloric acid technique outlined by
In order to assess the generic placement of
the new species, ITS sequences for 13 species
of Henriettea (incl. Henriettella), Loreya, and
Bellucia, along with 14 other species of
Melastomataceae, including seven Miconieae
(Table I) were included in a phylogenetic
analysis. These species were chosen to
provide a broad sampling across the morpho-
logical diversity within the putative new tribe
Henrietteeae (Penneys et al., 2004), along
with representatives of Miconieae, especially
Calycogonium DC., a genus that has been
confused with Henriettea (Judd, 1986). The
trees were rooted using Mouriri gleasoniana
Standl. and two species of Pternandra Jack
(Table I), based on the relationships inferred
in Michelangeli et al. (2004).
Flowering and vegetative material was
collected in silica gel. The species were
sequenced for internal transcribed spacer
regions of nuclear ribosomal DNA including
LIST OF TAXA, VOUCHERS (WITH HERBARIUM ACRONYMS), AND GENBANK ACCESSION NUMBERS FOR SPECIES INCLUDED IN
DNA-BASED CLADISTIC ANALYSIS OF HENRIETTEA AND PUTATIVE RELATIVES.
Taxa VouchersGenBank accession numbers
Astronia smilacifolia Triana
Bellucia pentamera Naudin
Blakea trinervia L.
Calycogonium grisebachii Triana
Calycogonium reticulatum (Cogn.) Judd & Skean
Charianthus alpinus (Sw.) R. A. Howard
Clidemia hirta (L.) D. Don
Graffenrieda latifolia (Naudin) Triana
Henriettea martiusii (DC.) Naudin
Henriettea spruceana Cogn.
Henriettea squamulosa (Cogn.) Judd
Henriettea succosa DC.
Henriettea uniflora Judd, et al.
Henriettea flavescens (Triana) Baill.
Henriettella rimosa Wurdack
Henriettella tuberculosa Donn. Sm.
Loreya arborescens (Aubl.) DC.
Loreya mespiloides Miq.
Loreya spruceana Triana
Loreya subandina Wurdack
Meriania macrophylla (Benth.) Triana
Miconia howardiana Judd, Salzman, & Skean
Miconia laevigata (L.) D. Don
Miconia mirabilis (Aubl.) L. O. Williams
Mouriri gleasoniana Standl.
Pternandra coerulescens Jack
Pternandra echinata Jack
Topobea parasitica Aubl.
Clausing 189 (MJG)
Penneys 1868 (FLAS)
Penneys 1629 (FLAS)
Becquer 82261 (HAJB)
Skean 4128 (MICH)
Penneys 1301 (FLAS)
Michelangeli 806 (NY)
Penneys 1303 (FLAS)
Michelangeli 412 (NY)
Michelangeli 390 (NY)
Skean 3807 (MICH)
Michelangeli 685 (NY)
Judd 8094 (FLAS)
Baraloto 2827 (FLAS)
Michelangeli 684 (NY)
Almeda 7663 (CAS)
Baraloto 2823 (FLAS)
Baraloto 2824 (FLAS)
Penneys 1854 (FLAS)
Alford 3107 (BH)
Michelangeli 829 (NY)
Williams s. n. (FLAS)
Penneys 1317 (FLAS)
Penneys 1301 (FLAS)
Penneys 1752 (FLAS)
Chen 543 (FLAS)
Chen 542 (FLAS)
Penneys 1821 (FLAS)
portions of the 18S gene, the entire ITS1,
5.8S cistron, and ITS2 regions. Total genomic
DNA was extracted using a modified CTAB
procedure of Doyle and Doyle (1987); am-
plification and sequence processing proce-
dures were as outlined in Judd (2007), except
that cycle sequencing was preformed using
7.5 μl reactions including 1 μl of PCR
product template and 6.5 μl of master mix.
PAUP* 4.0b4a was used to reconstruct
phylogenetic relationships. A maximum par-
simony, heuristic method of analysis was
conducted, with equally weighted, unordered
characters, gaps treated as missing, addition
sequence random, with 1000 replicates, TBR,
MaxTrees setting 10000, and MulTrees on.
The strength of support for relationships
found in the analysis was assessed using
bootstrap support (Felsenstein, 1985). For
each bootstrap analysis, a search was per-
formed with 1000 resampling replicates,
TBR, with MulTrees in effect.
Henriettea uniflora Judd, Skean, Penneys &
Michelangeli, sp. nov. Type: Dominican
Republic. Prov. Barahona: Monteada Nueva
region (peak is Loma Trocha de Pey), S on
Cabral-Polo Road, then ca. 7.8 km SE on
“riverbed road” and dirt road to “Cortico”
and silica mine (open degraded area),
disturbed cloud forest, 1260-1290 m, 31
May 2006 (fl, imm fr), W. S. Judd, J. D.
Skean, Jr., J. R. Abbott & R. E. Judd 8094
(holotype: FLAS; isotypes: GH, JBSD, MO,
NY, S, US).(Fig. 1)
Species haec ab Henriettea squamulosa (Cogn.) Judd
differt floribus sessilibus, solitariis (non inflorescentiis
pedunculatis, 3 floris), bracteolis 5–10 mm longis (non
2.5–5 mm longis) et lobis calyciris longioribus, foliis
anguste ovatis vel ellipticis (non ellipticis vel obovatis),
margine ciliata (non integra), et trichomatibus plusminusve
stellatis (non squamosis). Henrietteae ciliatae (Urb. &
Ekman) Alain affinis, a qua imprimis differt foliis
angustioribus, apice acuminato vel breviter acuminato
(non rotundato vel obtuso et mucronato), trichomatibus
dimorphis (non monomorphis).
Shrub or small tree to 5 m tall, with Rauh’s
architectural model. Indumentum of multicel-
lular, ferruginous stellate-peltate scales (these
often with a slightly raised central region and
a fringe of radiating cells) and multicellular,
stout non-glandular setae 0.7-2.5 mm long
(along the leaf margins). Young twigs slightly
quadrangular, with the interpetiolar face (in
relation to the distally adjacent node) slightly
concave, and the adjacent sides slightly
convex, becoming terete with age, the indu-
mentum of moderate to dense, appressed,
peltate scales, some of these falling with age;
internodes 0.4–10.2 cm long, the proximal
the longest, and becoming shorter distally.
Leaves with petiole 2.5–8(–12) mm long, the
indumentum similar to that of the twigs; blade
narrowly ovate to elliptic, 1.8–6.3(–9.4)×
0.45–3 cm, 2.1–7 times longer than wide, flat,
coriaceous, the apex acute, acuminate, or
shortly acuminate, with the tip of the leaf
having upturned margins, forming a mucro
with an adaxial pocket (acarodomatium), the
base narrowly acute to cuneate, the margin
plane to sometimes slightly revolute, entire
and ciliate, due to presence of stout non-
glandular setae, these sometimes deciduous
with age; venation acrodromous, suprabasal,
with prominent midvein and 4 secondary
veins, with the 2 conspicuous secondary veins
placed 0.55–3 mm from margin, 2 very
inconspicuous secondary veins ± intramarginal,
and numerous percurrent tertiary veins orient-
ed subperpendicular to midvein, the tertiary
veins separated by composite-intertertiary
veins, higher-order veins reticulate; adaxial
surface green, the indumentum initially of
numerous peltate scales but these quickly
deciduous so surface nearly glabrous at matu-
rity, the midvein slightly to strongly impressed,
the 2 major secondary veins slightly impressed
to flat, the 2 intramarginal secondary veins,
tertiary, and higher-order veins flat, the surface
smooth and shiny when fresh, but appearing
wrinkled and with minute projections (when
dry) due to the presence of numerous mostly
vertically oriented, megastyloid crystals, their
ends subcuticular; abaxial surface pale green,
densely covered with peltate-scales (when
young) but becoming only moderately covered
with such hairs at maturity (with hairs not
obscuring the leaf surface), the hairs of two
sizes, i.e., large, pale-ferruginous scales 0.15–
0.3 mm across and smaller, dark-ferruginous
scales 0.04–0.07 mm across, the midvein
prominently raised, major secondary veins
prominently to slightly raised, minor second-
ary veins, tertiary veins, and higher-order
veins flat. Inflorescences reduced to a solitary
flower, axillary, associated with 1–3 leaf-
JUDD ET AL.: HENRIETTEA (MELASTOMATACEAE)
FIG. 1. Henriettea uniflora. A. Habit (Judd 8094).
apices, showing apical acarodomatia (Judd 8094). D. Ferruginous, stellate-peltate hair (Judd 8094). E. Leaf, cross-
section; showing palisade chlorenchyma, two veins, and several megastyloid crystals (Skean 4298). F. Young fruit
(Skean 4298). G. Flower, in longitudinal section, the petals and stamens removed (Judd 8094). H. Petal, adaxial and
lateral views (Skean 4298). I. Petal (removed from nearly mature flower bud), abaxial and adaxial views (Judd 8094).
J. Stamens (Skean 4298). (Drawn from specimens at FLAS.)
B. Leaves, adaxial and abaxial surfaces (Judd 8094). C. Leaf
bearing nodes in the distal portion of the
twigs; each flower with two persistent brac-
teoles, these ovate to obovate, 5–16×2.6–
4.2 mm, the apices shortly acuminate, with
an apical pocket domatium on the adaxial
surface (formed from the upturned leaf mar-
gins), the indumentum similar to that of the
leaves. Flowers perfect, actinomorphic, nearly
sessile. Hypanthium broadly conic, the free
portion 0.5–1.3 mm long, the outer surface
with dense, pale-ferruginous peltate-scales,
the inner surface glabrous, slightly to clearly
16-ridged, with the ridges extending onto apex
of ovary, the apices of the ridges not or only
very slightly projecting. External calyx lobes
closely associated with internal lobes, each
external lobe at the apex of the internal lobe
and forming an extension to the length of the
lobe, but well demarked from the internal lobe
(when viewed on the adaxial surface) because
the internal calyx lobes are glabrous adaxially,
while the external calyx lobes are densely
lepidote adaxially; external calyx lobes 4,
extending 0.7–1.6 mm beyond the apex of
the internal lobes, narrowly triangular, with
acute apex, indumentum of dense peltate
scales both adaxially and abaxially; internal
calyx lobes 4, 1.4–2.5×1.6–4 mm, triangular,
densely lepidote abaxially but glabrous adax-
ially, the apex acute to shortly acuminate, the
margin entire; calyx tube 0.4–1.1 mm long.
Petals 4, imbricate in bud, ± asymmetrical,
ovate-triangular, 3.5–7.2×2-3.4 mm, glabrous
except for a few scales at the apex of the outer
petals (in bud), white, apically green-tinged;
apex acute, with minute apical projection;
margin entire. Stamens 8, the anther ellipsoi-
dal, 1.5–2 mm long, glabrous, pale yellow,
obscurely sagittate at base, fertile the entire
length, and opening by 2 confluent, dorso-
apical pores, the filament terete, 2–3 mm long,
glabrous, white. Ovary 4-locular, fully inferior,
obconic, 1.8–2.7×2.9–4 mm, with a glabrous
and slightly to moderately ridged apical portion
(within hypanthium), and articulated at base of
the style, with axile placentation, the placentae
extending into the locules; style terete, 3.5–
5.7 mm long, glabrous; stigma ± truncate.
Berries obconic, 3–4 mm long, 4.5–5.5 mm in
diam., reddish (immature), with dense ferrugi-
nous peltate scales, with adaxial surface of the
calyx lobes also turning red. Seeds angular-
obovoid, 0.6–0.8 mm long; testa ± smooth, the
cells ± isodiametric, with thickened anticlinal
Distribution and ecology.—Henriettea
uniflora is endemic to Hispaniola and known
only from the easternmost peaks of the Sierra
de Baoruco from 1200 to 1290 m (Fig. 2).
The type locality is usually referred to as
“Monteada Nueva” in reference to a nearby
coffee finca, but this forested region actually
occupies the summit of Loma Trocha de Pey,
ca. 18° 2′ 20.1″ N, 72° 57′ 35.4″ W. The
species has also been collected from the
nearby peaks Loma Remigio and Loma “Pie
Pol” (= Pie de Palo). These peaks are
separated from the western part of the moun-
tain range by a slight depression, the Hoyo
del Pelembito, and thus their high-elevation
flora is fairly isolated from that farther west in
the Sierra de Baoruco. Henriettea uniflora is
an uncommon shrub to small tree of the
diverse, but in places quite disturbed, moist
montane forests of these peaks. Associated
melastomes include Clidemia umbellata (Mill.)
L. O. Williams, Henriettea barkeri (Urb. &
Ekman) Alain, Leandra lima (Desr.) Judd
& Skean, Leandra limoides (Urb.) Judd &
Skean, Mecranium ovatum Cogn., Meriania
involucrata (Desr.) Naudin, Miconia alainii
Judd & Skean, M. dodecandra Cogn., M.
howardiana Judd, Salzman & Skean, M.
subcompressa Urb., M. tetrastoma Naudin,
Sagraea gracilis (Alain) Alain, and Tibouchina
longifolia (Vahl) Baill. Common trees and
shrubs of this moist forest include Alchornea
latifolia Sw., Beilschmiedia pendula (Sw.)
Hemsl., Brunellia comocladiifolia Humb.
subsp. domingensis Cuatc., Cecropia peltata
L., Cestrum sp., Clusia clusioides (Griseb.)
D’Arcy, Cyathea sp., Didymopanax tremulum
Krug & Urb., Eugenia sp., Gomidesia lindeni-
ana Berg, Gyrotaenia myriocarpa Griseb.,
Hieronyma domingensis Urb., Lobelia rotun-
difolia Juss. ex A. DC., Lyonia stahlii Urb.
var. costata (Urb.) Judd, Magnolia hamori
R. A. Howard, Myrsine coriacea (Sw.) R. Br.
ex Roem. & Schult., Ocotea spp., Oreopanax
capitatus (Jacq.) Decne. & Planch., Persea
krugii Mez, Piper umbellatum L., P. aduncum
L., Psychotria pubescens Sw., and Weinmannia
pinnata L. For more detailed description of
the vegetation of the easternmost peaks of the
Sierra de Baoruco see Guerrero (1993). The
diverse and highly endemic flora of the Sierra
JUDD ET AL.: HENRIETTEA (MELASTOMATACEAE)
de Baoruco is also treated in Hager & Zanoni
(1993) and García et al. (2001). The description
of Henriettea uniflora adds to the species of
Melastomataceae that are known to be endemic
(or nearly endemic) to the eastern peaks of the
Sierra de Baoruco; others include Mecranium
ovatum, which also occurs farther west, in the
mountains south of Puerto Escondido (Skean,
1993), Miconia alainii (Judd & Skean, 1994b),
M. howardiana (Judd et al., 1995), and
Sagraea gracilis (Liogier, 1971). A striking
tree of these forests, which is also endemic, is
Magnolia hamori (Howard, 1948; Guerrero,
1993). These forests, unfortunately, are not
protected from agricultural development.
Phenology.—The species is likely in bloom
from May through August.
Etymology.—The specific epithet refers to
the reduced axillary inflorescences, each of
which consists of a single, 2-bracteolate axis
that ends in a terminal flower.
Additional specimens examined. DOMINICAN
REPUBLIC. PROV. BARAHONA: Sierra de Baoruco,
Loma Remigio, 18° 5′ N, 71° 10.5′ W, 1200 m, 31 Aug
1999 (fl, fr) F. Jiménez, B. Peguero & A. Veloz 3006
(FLAS, JBSD, NY, US); Sierra de Baoruco, Loma Trocha
de Pey, 18° 02′ 20.1″ N, 72° 57′ 35.4″ W, 6.5 km on rd
above jct of Polo-Cabral Rd & “riverbed rd,” 1260-
1290 m, 31 May 2006 (fl) J. D. Skean, W. S. Judd, J. R.
Abbott & R. E. Judd 4298 (FLAS, JBSD, MICH, NY, S,
US); Sierra de Baoruco, Loma “Pie Pol” (Pie de Palo, on
map), La Guasára de Barahona, 18° 10′ N, 71° 12′ W,
1250 m, 25 Mar 1987 (st), T. Zanoni, J. Pimentel & R.
García 38651 (FLAS, JBSD, NY).
Vegetative anatomy.—As is typical of Mela-
stomataceae the stems have a ring of xylem
with phloem positioned both externally and
internally. However, unlike members of the
Miconieae, and in agreement with the report of
Van Tieghem (1891), they lack medullary
vascular bundles. Perivascular fibers are also
lacking, but there is a dense layer of more or
less isodiametric, lignified idioblasts in the
cortex. The nodes are unilacunar/unifascicular,
but the vascular bundle divides within the
petiole base, forming several bundles in the
petiole. The leaves have dorsoventral blades
with an epidermis and hypodermis, palisade
(of usually one cell layer) and spongy meso-
phyll; the midvein and petiole have 5 to 7
amphicribral vascular bundles, which are
arranged in a ring. The bundles are not
associated with fibers, but scattered lignified
idioblasts occur in the parenchyma tissue of
the midvein. The most distinctive feature of
the leaf blade is the presence of numerous
FIG. 2. Distribution of Henriettea uniflora on Hispaniola.
megastyloid crystals. These are usually ar-
ranged vertically (when leaf is viewed in
cross-section) with one end facing the adaxial
side and the other end facing the abaxial leaf
surface, but a few crystals were observed to be
oriented horizontally within the palisade chlor-
enchyma (Fig. 1E).
Molecular analysis.—For the entire ITS
region, 791 nucleotide sites were unambigu-
ously aligned and included in the analysis. Of
these sites, 432 were constant, 125 were
variable and non-informative, and 234 were
phylogenetically informative. The heuristic
search resulted in 12 most parsimonious trees
with a length of 707 steps (CI=0.70; RI=
0.76); the strict consensus tree (with branch
support assessed by bootstrap analysis) is
presented in Figure. 3.
Our results, like those of Michelangeli et
al. (2004), indicate that Henriettea, Loreya,
and Bellucia form a clade that is distinct from
the rest of the Miconieae, supporting the
FIG. 3. Strict consensus of 12 most parsimonious trees from heuristic analysis of ITS sequences of selected
species of Henriettea and relatives, with Pternandra and Mouriri as outgroups. Bootstrap values above 50% are
shown above the line.
JUDD ET AL.: HENRIETTEA (MELASTOMATACEAE)
placement of these genera (and their close
relatives, see above) in a distinct tribe
(Penneys et al., 2004). The authors currently
are investigating the limits and relationships
of this new tribe in greater detail.
The species of Henriettea (incl. Henriet-
tella) form a clade (bootstrap: 60% BS;
Fig. 3). Within this clade, H. uniflora is
placed as sister to H. squamulosa (98% BS).
Both species have ferruginous stellate-peltate
scales, and are representative of a group of
nine Antillean endemics with a distinctive
ferruginous-lepidote indumentum (see also
Judd, 1986). Although Henriettea uniflora is
unusual within the genus in having solitary,
axillary flowers because most species of
Henriettea have fasciculate inflorescences
produced from nodes on the older part of
the shoot (from which the leaves have fallen),
our DNA-based preliminary phylogeny clear-
ly indicates that H. uniflora is nested within
the phylogenetic structure of Henriettea s.l.
We are confident in placing this species
within Henriettea and hypothesizing a close
relationship to members of the H. squamu-
Finally, we note that our molecular results
(Fig. 3) support the earlier transfer (Judd,
1986) of Calycogonium squamulosum Cogn.
to Henriettea, and its recognition as H.
squamulosa (Cogn.) Judd. Calycogonium
DC. is actually a genus of Miconieae with
sympodial growth (i.e., showing Leeuwen-
berg’s model; Hallé, Oldman & Tomlinson,
1978) and terminal inflorescences, tissues
with druse crystals, stems with medullary
vascular bundles, conspicuous external calyx
lobes, and acute or acuminate to rounded
petals lacking an external tooth. It is closely
related to several Antillean taxa, such as
Tetrazygia Rich. ex DC. and Charianthus D.
Don (Fig. 3; Michelangeli et al., 2004).
General comparisons.—In addition to
Henriettea uniflora, the H. squamulosa com-
plex includes the Cuban species, H. acunae
(Alain) Alain, H. ekmanii (Urb.) Alain, and H.
squamata (Alain) Alain; the Hispaniolan spe-
cies, H. barkeri (Urb. & Ekman) Alain, H.
ciliata, H. megaloclada (Urb. & Ekman) Alain,
and H. reflexa (Urb. & Ekman) Alain; and the
Puerto Rican species, H. squamulosa (Alain,
1957; Judd, 1986; Liogier, 1995, 2000). The
stellate-peltate scales of H. uniflora (Fig. 1D)
closely match those of the above-mentioned
species (see Judd, 1986, Fig. 1c, f, j-m).
Several of these species have apiculate leaf
apices with the upturned margins forming an
apical pocket, likely functioning as an acar-
odomatum (Fig. 1C). In addition to H. uniflora,
we have seen such apices in H. barkeri, H.
ciliata, H. squamulosa, and H. reflexa.
Within this putative clade, i.e., the H.
squamulosa complex, Henriettea uniflora is
most readily compared with two species. It is
similar to H. ciliata because both species have
ciliate margins, i.e., the margin with scattered,
elongate multicellular hairs (and these often
deciduous). The other seven species within the
H. squamulosa complex have entire margins.
But, H. uniflora also is similar to H. squamu-
losa in having leaves with dimorphic peltate
scales abaxially (which do not obscure the
abaxial epidermis), flowers consistently borne
in leaf axils and associated with a pair of
conspicuous and persistent bracteoles, and
similarly positioned megastyloid crystals, all
of which are potential synapomorphies.
Henriettea uniflora and H. squamulosa are
unusual in that the leaf blade has numerous
megastyloid crystals that are mainly arranged
vertically (when leaf is viewed in cross-section)
with one end facing the adaxial side and the
other end facing the abaxial leaf surface. This
girder-like arrangement of the megastyloid
crystals (Fig. 1E) in these two species leads to
the adaxial epidermis, when dried, having a
wrinkled appearance and developing numerous
minute, pointed projections. Characteristically,
in Henriettea, the megastyloid crystals are
arranged horizontally within or just above the
palisade chlorenchyma of the leaf, as seen in H.
barkeri, H. ciliata, and H. megaloclada (as well
as numerous non-lepidote species of the genus).
Henriettea uniflora is easily distinguished
from H. squamulosa because its flowers are
more or less sessile and solitary (Fig. 1A; vs.
in mostly 3-flowered, short- to long-stalked
dichasial inflorescences; Judd, 1986, Fig. 1a),
with a pair of ovate to obovate bracteoles,
5–16 mm long (vs. a pair of narrowly
triangular bracteoles, 2.5–5 mm long). Like
H. uniflora, the flowers of H. squamulosa are
4-merous, the petals are white, and have only
a few apical peltate scales; the external calyx
lobes of H. uniflora, however, are evident,
and extend beyond the apex of the internal
calyx lobes (Fig. 1G) while those of H.
squamulosa are obsolete (Judd, 1986,
Fig. 2a). Vegetatively, H. uniflora can be
distinguished by its narrowly ovate to elliptic
leaves with ciliate margins (Fig. 1A, B; vs.
elliptic to obovate leaves with entire, non-
ciliate margins; Judd, 1986, Fig. 1a, b).
Leaves of H. uniflora are acute, acuminate,
or shortly acuminate at the apex, while those
of H. squamulosa are usually obtuse to
rounded or retuse. Finally, the ferruginous
scales of H. uniflora have radiating individual
arms (i.e., individual cells; Fig. 1D) while in
those of H. squamulosa the radiating elon-
gated cells are fused into a flat, thin layer
(Judd, 1986, Fig. 1c). Henriettea squamulosa
is endemic to Puerto Rico, occurring in
montane forests of the Cordillera Central
and Sierra de Luquillo.
Henriettea uniflora can be distinguished
from H. ciliata by its usually narrower leaves
(length/width quotient: 2.1–7 vs. 1.8–2.9) that
are acute or acuminate, to shortly acuminate
(vs. rounded to obtuse-mucronate) at the apex.
In H. uniflora the peltate scales on the abaxial
leaf surface are dimorphic, with some hairs
large and pale-ferruginous scattered among
more numerous smaller, dark-ferruginous
hairs; the abaxial epidermis is not obscured
by the scales. In contrast, in H. ciliata the
hairs are more or less monomorphic, and not
strongly contrasting in color (although they do
vary from pale to orange-ferruginous); they
densely cover the abaxial leaf surface, which is
thus totally obscured. Henriettea ciliata grows
in moist forests in the Cordillera Central and
Sierra de Neiba, Dominican Republic, and the
Massif des Matheux, Haiti.
It is useful here to compare Henriettea
uniflora with all of the species of the H.
squamulosa complex by means of the fol-
lowing dichotomous key.
Key to species of the Henriettea squamulosa complex
1. Abaxial leaf surface with dimorphic peltate scales, i.e., large, pale-ferruginous scales and smaller, dark-ferruginous
ones; inflorescences or flowers consistently borne in leaf axils, with solitary flower or dichasium having a pair of
conspicuous bracteoles, these present at anthesis.
2. Flowers solitary, ± sessile, with pair of ovate to obovate bracteoles 5-16 mm long; leaf margin ciliate; leaf apex
acute, acuminate or shortly acuminate................................................................................................... H. uniflora
2. Flowers mostly in 3-flowered, short-to long-stalked dichasial inflorescences, with a pair of narrowly triangular
bracteoles 2.5–5 mm long; leaf margin entire; leaf apex usually obtuse to rounded or retuse............... H. squamulosa
1. Abaxial leaf surface with ± monomorphic scales, although they may vary in the length of the central projection;
inflorescences borne on older nodes below the leafy portion of the shoot, or with some inflorescences in leaf axils,
but not exclusively on leafy portion of stem; flowers fasciculate, lacking bracteoles and with only minute, quickly
3. Leaves with ciliate margins...................................................................................................................... H. ciliata
3. Leaves with entire margins.
4. Leaves with 3 to 5 longitudinal veins.
5. Hairs on abaxial leaf surface stellate-peltate with a variously developed central projection, at least some
hairs with a conspicuous, elongate-conical structure; leaves with 3 to 5 longitudinal veins; flowers 4- or
5-merous............................................................. H. barkeri [and the possibly conspecific H. megaloclada]
5. Hairs on abaxial leaf surface stellate-peltate, flat or with only a minute central raised central portion, i.e.,
lacking an elongate central projection; leaves with 3 longitudinal veins; flowers 5-merous.
6. Abaxial leaf surface densely ferruginous-lepidote, the epidermis obscured............................... H. ekmanii
6. Abaxial leaf surface sparsely ferrugineous-lepidote, the epidermis visible................................. H. reflexa
4. Leaves with only 1 longitudinal vein.
7. Leaves 4–6 cm long, 0.8–1.7 cm wide, narrowly oblong-obovate................................................... H. acunae
7. Leaves 13–18 cm long, 4–9 cm wide, elliptic.............................................................................. H. squamata
JUDD ET AL.: HENRIETTEA (MELASTOMATACEAE)
Species concepts.—Henriettea uniflora is
likely monophyletic, as evidenced by its
solitary flowers that are associated with a
pair of enlarged bracteoles. The ciliate leaf
margins probably are an additional autapo-
morphy (and evolved in parallel with those of
H. ciliata). Thus, H. uniflora is a cladospe-
cies, satisfying the definition of the apomor-
phic species concept (Donoghue, 1985;
Mishler & Theirot, 2000). Because H. uni-
flora can be distinguished from its closest
relatives by consistent differences in several
characteristics, it also satisfies the diagnostic
species concept (Davis & Nixon, 1992;
Wheeler & Platnick, 2000). Finally, H. uni-
flora is reproductively isolated from its
putative closest relatives, H. squamulosa and
H. ciliata, because their ranges are allopatric,
fitting the biological species concept (Grant,
We thank Norris Williams, Keeper (FLAS),
and Kent Perkins, Collection Manager, for
assistance in processing specimen loans. We
are indebted to the curators of JBSD, S, and
NY for their loans of herbarium material.
Special thanks are given to Milcíades Mejía,
Francisco Jiménez, Brígido Peguero, Teodoro
Clase, and other herbarium personnel at the
Jardín Botánico Nacional, Santo Domingo
(JBSD) for their support of fieldwork in the
Dominican Republic. We also thank J.
Richard Abbott and Reuben E. Judd for their
valuable assistance in the field. We thank
Frances Combs for her beautiful illustration
(Fig. 1), and Tom Zanoni and Carmen Ulloa
for their helpful comments on an earlier
version of this paper. This research was
supported by NSF Grant DEB-0515636.
Alain, Hno. 1957. Flora de Cuba. Vol. 4. Contribuciones
Ocasionales del Museo de Historia Natural del
Colegio “De La Salle” 10: 1–556.
Baas, P. 1981. A note on stomatal types and crystals in
the leaves of Melastomataceae. Blumea 27: 475–479.
Davis, J. I. & K. C. Nixon. 1992. Populations, genetic
variation, and the delimitation of phylogenetic spe-
cies. Systematic Biology 41: 421–435.
Donoghue, M. J. 1985. A critique of the biological
species concept and recommendations for a phyloge-
netic alternative. Bryologist 88: 172–181.
Doyle, J. J. & J. L. Doyle. 1987. A rapid DNA isolation
procedure for small quantities of fresh leaf tissue.
Phytochemical Bulletin 19: 11–15.
Felsenstein, J. 1985. Confidence limits on phylogenies:
an approach using the bootstrap. Evolution 39: 783–
García, R., M. Mejía, B. Peguero & F. Jiménez. 2001.
Flora endemic de la Sierra de Baoruco, Republica
Dominicana. Moscosoa 12: 9–44.
Grant, V. 1981. Plant speciation. Second edition.
Columbia University Press, NY.
Guerrero A., A. E. 1993. Magnolia hamori, la flora y la
vegetación asociadas en la parte oriental de la Sierra
de Baoruco, República Dominicana. Moscosoa 7:
Hager, J. & T. A. Zanoni. 1993. La vegetacion natural
de la Republica Dominicana: una nueva clasificacion.
Moscosoa 7: 39–81.
Hallé, F., R. A. A. Oldeman & P. B. Tomlinson. 1978.
Tropical trees and forests: an architectural analysis.
Howard, R. A. 1948. The morphology and systematics
of the West Indian Magnoliaceae. Bulletin of the
Torrey Botanical Club 75: 335–357.
———. 1974. The stem-node-leaf continuum of the
Dicotyledoneae. Journal of the Arnold Arboretum 55:
Judd, W. S. 1986. Taxonomic placement of Calycogo-
nium squamulosum (Melastomataceae: Miconieae).
Brittonia 38: 238–242.
———. 1989. Taxonomic studies in the Miconieae
(Melastomataceae). III. Cladistic analysis of axil-
lary-flowered taxa. Annals of the Missouri Botanical
Garden 76: 476–495.
———. 2007. Revision of Miconia sect. Chaenopleura
(Miconieae, Melastomataceae) in the Greater Antil-
les. Systematic Botany Monographs 81: 1–235.
——— & J. D. Skean, Jr. 1994b. Miconia alainii
(Melastomataceae: Miconieae), a new species from
Hispaniola. Novon 4: 112–115.
———, V. T. Salzman & J. D. Skean, Jr. 1995.
Taxonomic studies in the Miconieae (Melastomata-
ceae). VII. Miconia howardiana, a new species from
Hispaniola. Brittonia 47: 414–421.
Liogier, A. H. 1971. Novitates Antillanae. V (1)
Miscellaneous new species from the Dominican
Republic. Phytologia 22: 163–174.
———. 1995. Descriptive flora of Puerto Rico and
adjacent islands, Vol. 4. Melastomataceae to Lenti-
bulariaceae. Editorial de la Universidad de Puerto
Rico. Río Piedras, Puerto Rico.
———. 2000. La Flora de la Hispañola, Vol. 9.
Melastomataceae. Jardín Botánico Nacional “Dr.
Rafael Ma. Moscoso” & Instituto Tecnológico de
Santo Domingo (INTEC). Santo Domingo, Domini-
Michelangeli, F. A., D. S. Penneys, J. Giza, D. Soltis,
M. H. Hills & J. D. Skean, Jr. 2004. A preliminary
phylogeny of the tribe Miconieae (Melastoma-
taceae) based on nrITS sequence data and its
implications on inflorescence position. Taxon 53:
Mishler, B. D. & E. C. Theirot. 2000. The phylogenetic
species concept (sensu Mishler and Theirot): mono-
phyly, apomorphy, and phylogenetic species con-
cepts. Pages 44–54. In: Q. D. Wheeler & R. Meier,
editors. Species concepts and phylogenetic theory: a
debate. Columbia University Press, New York.
Penneys, D. S., F. A. Michelangeli, W. S. Judd & J. D.
Skean, Jr. 2004. Henrietteeae, a new tribe of
Neotropical Melastomataceae. Botany 2004. Abstract
Skean, J. D., Jr. 1993. Monograph of Mecranium
(Melastomataceae-Miconieae). Systematic Botany
Monographs 39: 1–116.
Van Tieghem, M. P. 1891. Classification anatomique
des Mélastomacées. Bulletin de la Société Botaniqué
de France. 38: 114–124.
Wheeler, Q. D. & N. I. Platnick. 2000. The phyloge-
netic species concept (sensu Wheeler and Platnick).
Pages 55–69. In: Q. D. Wheeler & R. Meier, editors.
Species concepts and phylogenetic theory: a debate.
Columbia University Press, New York.
Wurdack, J. J. 1986. Atlas of hairs for neotropical
Melastomataceae. Smithsonian Contributions to Bot-
any, No. 63: 1–80.
JUDD ET AL.: HENRIETTEA (MELASTOMATACEAE)