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Taxonomic revision of the Greater Antillean Pseudolima clade of Miconia (Miconia sect. Krugiophytum: Miconieae: Melastomataceae)

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The Pseudolima clade (Miconia sect. Krugiophytum) consists of two Puerto Rican species, M. karlkrugii and M. krugiana and one Hispaniolan species, M. inaequipetiolata. All three species display morphological characters (e.g., bulla-based hairs) that resemble those found in Miconia sect. Lima, and for that reason had been considered to be putative members of the Lima clade. Molecular phylogenetic analyses, presented here, suggest that these taxa actually are not most closely related to members of the Lima clade, and thus it appears likely that the characters seemingly linking these species with the Lima clade either are symplesiomorphic or convergent. These three species are representative of the taxonomic difficulties regarding determination of clade affinities (and generic limits) using morphological characters, which is a common problem in Miconieae. We provide a taxonomic revision of these three species including illustrations, distribution maps and descriptions. Resumen El clado Pseudolima (Miconia secc. Krugiophytum) consta de dos especies de Puerto Rico, M. karlkrugii y M. krugiana, y una de La Española, M. inaequipetiolata. Las tres especies muestran caracteres morfológicos (e.g., tricomas expandidos en la base) que se asemejan a los que se encuentran en Miconia sect. Lima y por esa razón han sido considerado como miembros putativos del clado Lima. Los análisis de filogenética molecular presentados aquí sugieren que estas especies están más relacionados a otros clados que al clado Lima, por lo que los caracteres semejantes a los del clado Lima probablemente son plesiomórficos o convergentes. Estas tres especies demuestran las dificultades taxonómicas con la delimitación de las afinidades de clados (y límites genéricos) tomando en cuenta solo la morfología, un problema muy común en la tribu Miconieae. Proveemos una revisión taxonómica de estas tres especies que incluye ilustraciones, mapas de distribución y descripciones.
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Taxonomic revision of the Greater Antillean Pseudolima clade
of Miconia (Miconia sect. Krugiophytum: Miconieae: Melastomataceae)
LUCAS C. MAJURE
1,2
,WALTER S. JUDD
1,2
,AND FABIÁN A. MICHELANGELI
3
1
Department of Biology, University of Florida, 220 Bartram Hall, P.O. Box 118525, Gainesville, FL 32611,
USA; e-mail: lmajure@ufl.edu; e-mail: wjudd@botany.ufl.edu
2
Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL 32611-7800,
USA; e-mail: lmajure@ufl.edu; e-mail: wjudd@botany.ufl.edu
3
Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458-5126, USA; e-mail:
fabian@nybg.org
Abstract.ThePseudolima clade (Miconia sect. Krugiophytum) consists of two Puerto
Rican species, M. karlkrugii and M. krugiana and one Hispaniolan species,M.
inaequipetiolata. All three species display morphological characters (e.g., bulla-based hairs)
that resemble those found in Miconia sect. Lima, and for that reason had been consideredto be
putative members of the Lima clade. Molecular phylogenetic analyses, presented here, suggest
that these taxa actually are not most closely related to members of the Lima clade, and thus it
appears likely that the characters seemingly linking these species with the Lima clade either are
symplesiomorphic or convergent. These three species are representative of the taxonomic
difficulties regarding determination of clade affinities (and generic limits) using morpholog-
ical characters, which is a common problem in Miconieae. We provide a taxonomic revision of
these three species including illustrations, distribution maps and descriptions.
Key Words: Calycogonium, Caribbean, Hispaniola, Leandra,Ossaea, Puerto Rico.
Resumen.ElcladoPseudolima (Miconia secc. Krugiophytum) consta de dos especies de
Puerto Rico, M. karlkrugii yM. krugiana, y una de La Española, M. inaequipetiolata.Lastres
especies muestran caracteres morfológicos (e.g., tricomas expandidos en la base) que se
asemejanalosqueseencuentranenMiconia sect. Lima y por esa razón han sido considerado
como miembros putativos del clado Lima. Los análisis de filogenética molecular presentados
aquí sugieren que estas especies están más relacionados a otros clados que al clado Lima, por
lo que los caracteres semejantes a los del clado Lima probablemente son plesiomórficos o
convergentes. Estas tres especies demuestran las dificultades taxonómicas con la delimitación
de las afinidades de clados (y límites genéricos) tomando en cuenta solo la morfología, un
problema muy común en la tribu Miconieae. Proveemos una revisión taxonómica de estastres
especies que incluye ilustraciones, mapas de distribución y descripciones.
The tribe Miconieae (Melastomataceae) con-
sists of ca. 1800 species distributed throughout
the Neotropics (Michelangeli et al., 2008). Most
species occurring in the Caribbean region are the
products of four introductions from continental
Central and South America, which subsequently
formed four major radiations: Mecranium,
Miconia sect.Sagraea,Miconiasect.
Chaenopleura, and the Caribbean clade
(Michelangeli et al., 2008). Members of the
Caribbean clade of Miconieae exhibit substantial
morphological diversity and the species of this
group have traditionally been placed in numerous
genera (e.g., Calycogonium DC., Charianthus D.
Don, Clidemia D. Don, Conostegia D. Don,
Leandra Raddi, Miconia Ruiz & Pav., Ossaea
DC., Pachyanthus A. Rich. and Tetrazy gi a Rich.
ex DC.). Of these, Calycogonium,Pachyanthus,
and Tetraz yg ia have been considered Caribbean
endemics or near endemics (i.e., one species of
Pachyanthus is found in Belize, and Te tr az ygia
also is encountered in Florida, USA; Wunderlin &
Hansen, 2011; Bécquer Granados 2012).
Molecular phylogenetic analyses have shown that
Brittonia, DOI 10.1007/s12228-014-9349-x
ISSN: 0007-196X (print) ISSN: 1938-436X (electronic)
© 2014, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
all of these aforementioned genera but
Charianthus are polyphyletic (Bécquer-
Granados et al. 2008;Goldenbergetal.,2008;
Martin et al., 2008; Michelangeli et al., 2008).
Thus, as currently circumscribed, these genera
are undiagnosable; they are recognized by
homoplasious characters. In order to render them
monophyletic, at least 16 genera within the
Caribbean clade would need to be recognized
based on current taxon sampling in phylogenetic
analyses (Goldenberg et al., 2008; Michelangeli
et al., 2008). However, extreme generic splitting
would also have taxonomic effects within other
clades of the tribe. Many of these genera would be
monotypic, either because they are not resolved in
any major clade or because they are morpholog-
ically divergent from and/or show no
synapomorphic morphological features with the
other members of their particular clade (Majure &
Judd, unpubl. data). The already large genus
Miconia, with around 1057 species (Goldenberg
et al., 2013), is especially problematic and would
have to be split into numerous poorly diagnosable
genera based on our current knowledge of the
phylogeny (Goldenberg et al., 2008;
Michelangeli et al., 2008). Thus, recent work
has suggested the recognition of these taxa in a
broadly circumscribed Miconia (Ionta et al.,
2012; Judd & Ionta, 2013). Therefore, we recog-
nize the three species treated in this work within
Miconia s.l.
Taxono mi c Histo ry o f Miconia sect.
Krugiophytum
Miconia sect. Krugiophytum (Cogn.) Majure
& Judd consists of three species from the
Greater Antilles, Miconia karlkrugii Majure &
Judd,M.krugiana(Cogn.) Majure & Judd and
M. inaequipetiolata Majure & Judd. Miconia
karlkrugii and M. krugiana areendemicto
Puerto Rico, while M. inaequipetiolata is en-
demic to Hispaniola (Haiti and Dominican
Republic).
Miconia sect. Krugiophytum was originally de-
scribed by Cogniaux (1886) under the genus
Calycogonium,andMiconia karlkrugii (as
Calycogonium krugii Cogn.) was the only species
treated within the section. Cogniaux (1891)orig-
inally placed Miconia krugiana under Ossaea
sect. Glaziophytum Cogn. along with a number
of other distantly related species, such as
members of Miconia sect. Lima (Miconia
argentimuricata Majure & Judd) and many other
mostly South American taxa, some of which are
in Leandra s.s. and the Secundiflorae clade (e.g.,
Ossaea amygdaloides (DC.) Triana,
O. angustifolia (DC.) Triana, O. capillaris (D.
Don) Cogn., O. confertiflora (DC.) Triana;
Michelangeli et al., unpubl. data). Miconia
inaequipetiolata has never been circumscribed
within any generic section.
Miconia karlkrugii was first described as
Calycogonium krugii (Cogniaux, 1886),
based on material collected by Paul Sintenis
(18471907), but was transferred later to
Leandra (Judd & Skean, 1991), and then
recently recognized as Ossaea krugii
(Cogn.) Axelrod (Axelrod, 2011;
Michelangeli & Bécquer Granados 2012).
Miconia krugiana, also based on a collection
by Sintenis, was originally described as
Ossaea krugiana Cogn. (Cogniaux, 1891)
but was later treated as Leandra krugiana
(Cogn.) Judd & Skean (Judd & Skean,
1991;Liogier,1995). Likewise, the closely
related Miconia inaequipetiolata,firstcol-
lected by Erik L. Ekman (18831931) in
theMassifdelaSelle,Haiti,wasfirstde-
scribed as an Ossaea (Urban & Ekman,
1929) and then recognized under Leandra
(Judd & Skean, 1991;Liogier,2000). All
three taxa were later transferred to Miconia
by Majure & Judd (2013).
Miconia inaequipetiolata, M. karlkrugii and
M. krugiana have often been considered close-
ly related to (Majure & Judd, 2013), if not
forming part of the Lima clade (Cogniaux,
1891, in part) based on morphological char-
acters. All of these taxa exhibit bulla-based
hairs on most surfaces of the plant, including
leaf adaxial and abaxial surfaces (Fig. 1),
petioles, stems, hypanthia and even floral
parts, such as ovary apices and petal abaxial
surfaces. These three taxa have never been
considered to be closely related and thus
never have been treated together.
In this paper we carry out a phylogenetic re-
construction for a portion of the Caribbean clade
of Miconieae in order to delimit the clade bound-
aries of these three taxa and determine their clos-
est relatives. We show for the first time that these
three species most likely form a clade (informally
referred to here as the Pseudolima clade) that is
BRITTONIA [VOL
FIG.1. Photos of Miconia krugiana (A, B)andMiconia karlkrugii (C, F). A)BranchofM. krugiana showing leaf adaxial
surface bulla-based hairs and glomerulate inflorescence. B)FlowerofM. krugiana. C)FlowerofM. karlkrugii (note: some petal
apices showing herbivore damage). D) Immature fruit of M. karlkrugii with calyx tube tearing. E) Maturing fruit of M. karlkrugii.
F) Mature fruit of M. karlkrugii showing dark red color. Aand Bfrom Skean 3820,C-F from Michelangeli 2021.
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
not most closely related to the Lima clade. We
then provide a taxonomic revision for the three
species of this clade, for which we provide the
formal name, Miconia sect. Krugiophytum.
Materials and Methods
TAXON SAMPLING AND PHYLOGENETIC ANALYSES
Leaf material of Miconia krugiana (Skean
3820:FLAS)andM. inaequipetiolata (Clase
6960: FLAS) was sampled from herbarium
specimens and extracted using a modified
CTAB extraction (Doyle & Doyle, 1987). The
plastid intergenic spacers accD-psa1 (Shaw
et al., 2007), psbK-psbL (Reginato et al.,
2010), and the nuclear ribosomal loci ITS
(Michelangeli et al., 2004) and ETS
(Michelangeli, unpubl. data; Majure et al.,
2014) were amplified and sequenced for their
inclusion into a larger dataset (F. Michelangeli
et al., unpubl. data) of members of the
Caribbean clade (Michelangeli et al., 2008).
The larger dataset includes members of the
Calycopteris (=Miconia sect. Calycopteris;
Judd et al., 2014), Calycogonium (glabrate-
domatial species, see Judd & Skean, 1991),
Lima (=Miconia sect. Lima; Majure & Judd,
2013), Paralima (discussed here),and
Pseudolima clades. Miconia blancheana Urb.
and M. umbellata (Mill.) Judd & Ionta were
used as outgroup taxa based on Michelangeli
et al. (2008) and Majure et al. (2014).
Sequences were edited using Sequencher
4.2.2 (Gene Codes, Ann Arbor, Michigan,
USA) and manually aligned in SeAl v. 2.0
(Rambaut, 2007). All data were combined after
preliminary analyses of each region showed no
well-supportedincongruence (i.e., hard
incongruence; Seelanen et al., 1997)among
data sets (Johnson & Soltis, 1998). Maximum
parsimony (MP) analyses were carried out in
PAUP* 4.0 (Swofford, 2002) by conducting
1000 random addition sequence replicates,
and support was evaluated by running 1000
nonparametric bootstrap (bs) pseudoreplicates,
each with 10 addition sequence replicates.
Maximum likelihood (ML) analyses were car-
ried out using RAxML (Stamatakis, 2006)un-
der the GTR + Γmodel of molecular evolution
and undertaking 1000 nonparametric bootstrap
pseudoreplicates. All sequence data are depos-
ited in GenBank (see Appendix 1).
HYPOTHESIS TESTING
Based on our MP analyses that yielded no bs
support for the Pseudolima clade (including
M. inaequipetiolata), we used Mesquite v. 2.73
(Maddison & Maddison, 2010) to construct five
different constraint trees to test the monophyly of
a clade consisting of M. inaequipetiolata, M.
krugiana, and M. karlkrugii. The five constraint
trees were as follows: 1) M. inaequipetiolata sis-
ter to the Lima clade, 2) M. inaequipetiolata sister
to the glabrate-domatial Calycogonium clade, 3)
M. inaequipetiolata sister to the Calycopteris
clade, 4) M. inaequipetiolata sister to the
M. krugiana/M. karlkrugii +Calycopteris clade,
and 5) M. inaequipetiolata sister to the
M. krugiana/M. karlkrugii clade (forming the
Pseudolima clade). We then used MP in PAUP*
and the Templeton test (Templeton, 1983)to
compare the length and statistical differences
among the constraint and observed trees.
MORPHOLOGICAL DATA
For our taxonomic revision, we examined a
total of 65 collections of the three species includ-
ed here. All together, we measured or observed 88
variable morphological characters of these spe-
cies. Phenological, distribution, and habitat data
were retrieved from herbarium specimens, when
available. Specimen measurements are based on
the methodology of Judd (2007). Type material
that was only seen online is given with the acces-
sion number from that institution.
Results
We were only able to amplify psbK-psbL and
ITS for M. krugiana, as a result of apparent DNA
degradation (only small molecular weight DNA
fragments were visible on a 1% agarose gel). In
contrast, DNA from Miconia inaequipetiolata
was amplified using all four loci. In MP analyses,
Miconia krugiana was resolved as sister to
M. karlkrugii,andM. inaequipetiolata, although
not statistically supported (bs <50), was sug-
gested to be sister to the M. krugiana-M.
karlkrugii clade. Miconia krugiana and
M. karlkrugii also formed a clade in ML
analyses (bs= 76), but their relationship to
M. inaequipetiolata was not supported (the
placement of M. inaequipetiolata was unresolved).
The M. krugiana-M. karlkrugii-M. inaequipetiolata
clade (Pseudolima clade), as resolved in MP
BRITTONIA [VOL
analyses, was nested within a more inclusive
clade, including the Calycopteris and glabrate-
domatial Calycogonium clades, which was sister
to the Lima clade (Fig. 2). Relationships among
the Calycopteris, glabrate-domatial
Calycogonium and Pseudolima clades were
unresolved.
In our maximum parsimony analyses, in the 14
most parsimonious trees recovered (all with a tree
length of 303), M. inaequipetiolata, M. krugiana,
and M. karlkrugii always formed a clade, al-
though there was no statistical support (bs) for
this grouping. All trees recovered based on one of
the five constraints where the Pseudolima clade
was rendered non-monophyletic were longer than
any of the most parsimonious trees (tree length
303 vs. 304305). Trees resulting from the
constraint of a monophyletic Pseudolima clade
were the same length (L=303) as the most parsi-
monious trees. However, there was no statistically
significant difference among the trees
representing a monophyletic Pseudolima group
and competing hypotheses.
Discussion
Our phylogeny suggests that
M. inaequipetiolata, M. krugiana,and
M. karlkrugii form a clade that is more closely
related to the glabrate-domatial Calycogonium
and Calycopteris clades than it is to the Lima
clade (Fig. 2). Our hypothesis tests show there
to be no significant difference in alternative to-
pologies, but they do suggest (based on MP) that
FIG.2. Maximum parsimony 50% majority rule phylogeny of the Pseudolima clade and closely related members of
Caribbean Miconieae. Bootstrap values are given for both MP and ML (MP/ML), respectively, above branches.
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
M. inaequipetiolata, M. krugiana,and
M. karlkrugii form a clade.It is also clear from
our results that the three species are likely not
mostly closely related to members of Miconia
sect. Lima.
In contrast to our phylogenetic results, Miconia
inaequipetiolata, M. karlkrugii and M. krugiana
have been considered closely related to members
of Miconia sect. Lima (e.g., M. argentimuricata,
M. lima (Desr.) M. Gómez, M. pedunculata
Majure & Judd, M. ottoschmidtii (Urb.) Majure
& Judd; see Cogniaux, 1891; Majure & Judd,
2013) because of their conspicuous bulla-based
hairs, which can be found throughout the stems,
leaves, hypanthia (see Fig. 1), and even at the
petal apex of M. inaequipetiolata. Consequently,
it appears that the striking bulla-based hairs
shared by these taxa, Miconia sect. Lima, and
other Caribbean taxa, such as the mostly Cuban
Paralima clade (e.g., Ossaea glomerata (Naud.)
Triana, Ossaea ovatifolia Urb.; Fig. 2)are
homoplasious. However, more resolution is need-
ed along the backbone of the tree to more con-
vincingly determine directionality of morpholog-
ical evolution in the Caribbean clade
(Michelangeli et al., 2008; Michelangeli et al.
unpubl. data).
Putative synapomorphies of the Pseudolima
clade are not immediately obvious. Miconia
karlkrugii has inflorescences reduced to a single
large flower (Fig. 1C), while M. krugiana pro-
duces condensed, cymose inflorescences, with
the individual flowers sessile to very shortly ped-
icellate (Fig. 1A, B), and these cymes resemble
glomerulate heads on long peduncles. Miconia
inaequipetiolata produces open, cymose inflores-
cences with flowers on long pseudopedicles (Fig.
3A). Miconia karlkrugii has subulate or narrowly
elliptic anthers with apical to dorsal pores;
M. krugiana produces ovate or broadly elliptic
anthers with dorsally-oriented pores, and
M. inaequipetiolata produces ovate or narrowly
elliptic anthers with dorsal pores. Miconia
karlkrugii produces large, glabrous petals, while
M. krugiana produces smaller petals with a min-
ute apical hair. Miconia inaequipetiolata produces
small petals with notched apices from which one
well-developed hair is produced (Fig. 3). Also,
M. inaequipetiolata and M. krugiana produce
poorly-developed, large bulla-based hairs
surrounded by poorly-developed small bulla-
based hairs on the leaf adaxial surface (these hairs
do not completely fill the leafareoles; Figs. 1A,3,
7), while M. karlkrugii produces well-developed
bulla-based hairs on the adaxial leaf surface that
almost or nearly completely fill the leaf areoles
(Figs. 1C, D,5).
A morphological character linking these three
species is the terminal inflorescences. These in-
florescences are often borne on shoots (produced
from lateral buds) that produce 13 nodes before
terminating in an inflorescence. These three spe-
cies, as seen in numerous specimens, produce
lateralbudsthatshowasynchronousgrowthpat-
terns, wherein one lateral bud produces a larger
and more dominant branch that eventually be-
comes the primary stem, after the termination of
the previous primary stem in an inflorescence.
The other lateral bud either aborts or produces a
smaller stem (with 13 leaf pairs) that also
terminates in an inflorescence. Both
M. inaequipetiolata and M. krugiana have
pendant inflorescences. The pendant
inflorescences and somewhat sympodial growth
form of these taxa likely led Urban and Ekman
(1929)dodescribeM. inaequipetiolata as having
both terminal and axillary inflorescences, al-
though they are not truly axillary. The growth
form described above is not generally exhibited
in the Lima clade, except for the basal-most taxon
in our analyses, M. jashaferi Majure & Judd; most
members of the Lima clade have shoots with
obviously terminal inflorescences and that contin-
ue growth through the development of a pair of
axillary buds (associated with leaf pair immedi-
ately below the inflorescence, and forming
branches of more or less equal dominance). This
growth architecture also is not characteristic of
other members of the Caribbean clade, except
for Miconia sect. Calycopteris (Judd et al.,
2014), a putative close relative (and potentially
the sister group; Majure et al., 2014)ofthe
Pseudolima clade. Both M. krugiana and
M. inaequipetiolata have pendant, cymose inflo-
rescences and are also strongly anisophyllous,
with the smaller leaves 1/4-1/3 the size of the
larger, at least at some nodes. All three species
have a crown of long, needle-like, bulla-based
hairs at the apex of the ovary (and base of the
style), a character also shared by most members of
the Lima clade (Majure & Judd, 2013), and all
three species have well-developed bulla-based
hairs on the adaxial leaf surface. Miconia
inaequipetiolata and M. krugiana have apical
petal hairs, although these are more poorly devel-
oped than most members of the Lima clade, and
BRITTONIA [VOL
those hairs in the Lima clade are mostly subapical
or produced in the medial portion of the petal.
Miconia karlkrugii and M. krugiana share
more features with one another than either species
does with M. inaequipetiolata. Both species pro-
duce clavate-dendritic hairs on the leaf petiole,
adaxial leaf surface in between bulla-based hairs,
at the base of bulla-based hairs along the leaf
FIG.3. Miconia inaequipetiolata. A) Habit. B) Close-up of lower leaf surface. C) Leaf abaxial surface. D) Close-up of leaf
abaxial surface. E)Flower.F) Petal. G)Stamen.H) Fruit longitudinal section. I) Fruit cross section. A,Ifrom Clase 6960,B, D
from Clase 4087,E,Gfrom Liogier 18150,F,Hfrom Clase 4055.
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
margins, and at the apices of the calyx tube, as
well as the upper margin of the calyx lobes. Also,
both species generally have strongly clawed
petals (Figs. 1,5&7). Miconia inaequipetiolata,
however, produces filiform or gland-headed fili-
form hairs in many of the same locations where
the other two species produce clavate-dendritic
hairs.
It is not surprising that M. inaequipetiolata,
given its putative phylogenetic position as sister
to the M. karlkrugii-M. krugiana clade (Fig. 2), is
found to be morphologically more divergent. We
have noticed that the earliest diverging species
within many clades (within the Caribbean clade)
are often morphologically quite divergent from
the remaining members of their respective clades
(e.g., M. jashaferi exhibits a number of morpho-
logical characters that may either be
symplesiomorphic or apomorphic but that differ
from other members of the Lima clade; Majure
et al., in prep.).
The Pseudolima clade
According to our phylogenetic reconstruction,
these three species likely are closely related, con-
stituting the Pseudolima clade. As Cogniaux
(1886) already had proposed a sectional name
in Calycogonium for Miconia karlkrugii,we
have adopted that sectional name and transferred
it to Miconia:Miconia sect. Krugiophytum,in-
cluding both M. krugiana and M. karlkrugii,and
provisionally M. inaequipetiolata. The closest
relatives of the Pseudolima clade, as discussed
above, are the Calycopteris and the glabrate-
domatial Calycogonium clades. All three clades
form a more inclusive clade, which is sister to the
Lima clade (Fig. 2). The Calycopteris clade can
easily be distinguished by globular stellate hairs,
strongly four-lobed ovaries, calyx teeth flattened
parallel to the floral radii, and seeds with a dis-
tinctive bulging raphe (Judd & Majure, 2014;
Bécquer et al. 2014;Juddetal.,2014). The
glabrate-domatial clade of Calycogonium is
most easily recognized by reduced inflores-
cences, the reduction in the indumentum of
the abaxial leaf surfaces, and more or less
volcano-like acarodomatia formed from the
union of hairs in the axils of the innermost
secondary veins and primary vein (Judd &
Skean, 1991), although the two subsequent sis-
ters to this clade, Calycogonium turbinatum Urb.
& Ekman and Miconia moensis (Britton) Alain,
do not have well-developed domatia, and
Miconia moensis does not have reduced inflores-
cences (as mentioned above). As noted above,
however, the species of the Pseudolima clade are
mosteasilyconfusedwiththoseoftheLima
clade(seekey).
Chromosome numbers are completely un-
known in the Pseudolima clade.
Miconia sect. Krugiophytum (Cogn.) Majure &
Judd, comb. nov. Calycogonium sect.
Krugiophytum Cogn., Bot. Jahrb. Syst. 4:
277. 1886. Type: Miconia karlkrugii Majure
&Judd(=Calycogonium krugii Cogn.).
Ossaea sect. Glaziophytum Cogn., Monogr. Phan.
7: 1048. 1891. Type, here designated,
Miconia krugiana (Cogn.) Majure & Judd
(=Ossaea krugiana Cogn.). We typify
Ossaea sect. Glaziophytum,asnotypewas
selected by Cogniaux (1891)andM. krugiana
well matches the protologue.
Evergreen shrubs 13 m tall, stems terete, lacking
longitudinal ridges, stem indumentum of multicel-
lular, bulla-based hairs, these either granulate or
long, ascending or spreading, nodal line present or
absent. Leaves opposite, decussate, anisophyllous to
± isophyllous, blade elliptic, coriaceous, base acute
to slightly rounded, apex acute, rounded or acumi-
nate, margin serrate with teeth generally covered in
one bulla-based hair, secondary veins 2 pairs, arising
from the base or suprabasal (especially the inner pair
of secondary veins), tertiary veins percurrent, mostly
subperpendicular to the midvein, often connected
by quaternary veins, the midvein, secondary and
tertiary veins flat adaxially and raised abaxially;
indumentum of strongly-developed bulla-based
hairs nearly filling the areoles or not, and also with
sessile glandular hairs on both surfaces,
acarodomatia absent or inconspicuous.
Inflorescence terminal, often appearing as arising
from sympodial growth as a result of asynchronous
lateral bud growth, where one lateral bud generally
grows more vigorously and thus is larger than the
other, and the apical meristem of the primary shoot
aborts after flowering, thus the more vigorous lateral
bud takes the place of the primary shoot, inflores-
cence expanded with numerous flowers on long
pseudopedicels or nearly sessile and forming
glomerulate heads, or flowers solitary, pendant or
erect, bracts small and linear or broad and folia-
ceous. Flowers 56 merous, actinomorphic to
slightly zygomorphic. Hypanthium terete, slightly
BRITTONIA [VOL
to strongly constricted above the ovary, the outer
surface covered in dense, elongate or granulate
multicellular hairs, these mostly bulla-based, and
also with sessile or stalked, glandular hairs, adax-
ial surface with sessile to short-stalked, glandular
hairs. Calyx lobes triangular or oblong, calyx tube
tearing or not between the lobes, calyx teeth
terete, exceeding the length of the calyx tube,
covered in bulla-based hairs. Petals ovate, elliptic
or obovate, clawed at the base or truncate-round-
ed, asymmetric or symmetric, apex with multicel-
lular hair or such hair absent. Stamens 1012,
isomorphic, filaments glabrous, anthers ovate or
slightly oblong, without a dorsal bump, with one
dorsally or apically-oriented pore. Ovary 36loc-
ular, inferior, apically flat or acute, with multicel-
lular, elongate hairs, placentation axile, placenta
deeply intruded, ovules numerous, style glabrous,
linear, not dilated in the middle, slightly deflexed
to one side of the flower or straight, stigma
punctate. Berries globose, bright red or potentially
purple-black at maturity. Seeds obpyramidal or
obovate, testa smooth, raphe dark, flat, extending
the length of the seed.
Key to the species of Miconia sect. Krugiophytum, with a comparison of Miconia sect.
Krugiophytum and Miconia sect. Lima
1. Inflorescences obviously terminal, petiole adaxial surface with bulla-based hairs only, flowers 45(6) merous, multicellular
hairs on petals, when present, produced mediallyor subapically, Cuba, Hispaniola, Jamaica ..1. Miconia sect. Lima (not treated
here)
1. Inflorescences terminal but oftentimes appearing lateral or on short shoots as a result of the unequal growth of lateral buds
(where one lateral bud produces one shoot generally larger than the other lateral bud, producing a pseudosympodial growth
form), petiole adaxial surface with mixed clavate-dendritic or filiform hairs and bulla-based hairs, flowers 56 merous,
multicellular hairs on petals, when present, produced apically, Hispaniola and Puerto Rico ......... 2. Miconia sect. Krugiophytum
2. Bulla-based hairs on adaxial leaf surface, dense and broadlyexpanded at the base, nearly covering areoles, and/or the hairs
compressed dorsi-ventrally, inflorescences glomerulate cymes or reduced to one flower, petals clawed, carpels 46, Puerto
Rico ........................................................................................................................................................................................... 3
3. Inflorescences reduced to one flower, calyx tube regularly tearing longitudinally, flowers 25.2 cm wide (from petal apex
to petal apex), leaf apices rounded to broadly acute .................................................................................. 2.M. karlkrugii
3. Inflorescences composed of 49 flowers forming a condensed cyme, calyx tube not regularly tearing longitudinally,
flowers 0.750.78 cm wide, leaf apices acuminate ..................................................................................... 3. M. krugiana
2. Bulla-based hairs on adaxial leaf surface sparse, not greatly expanded at the base, not covering areoles, the hairs elongate
with attenuate apices, not compressed dorsi-ventrally, inflorescences expanded cymes, flowersnumerous, petalsnot clawed,
carpels 3, Hispaniola .............................................................................................................................. 1. M. inaequipetiolata
1. Miconia inaequipetiolata Majure & Judd, J.
Bot. Res. Inst. Texas. 7: 268. 2013. Ossaea
inaequidens Urb. & Ekm. Ark. Bot. 22A, 17:
63. 1929. Leandra inaequidens (Urb. &
Ekm.) Judd & Skean, Bull. Florida Mus.,
Biol. Sci. 36: 61. 1991. Type: Haiti, Massif
de la Selle, Croix-des-Bouquets, Badeau, at
Trou-á-lSau, 1000m, 15 Mar 1927, (sterile,
infl. branches only), E. L. Ekman H7860 (ho-
lotype: S; isotypes: K [K000329545], NY).
Non Miconia inaequidens (DC.) Naudin.
(Fig. 3)
Evergreen shrub, 12.5 m tall; stems round
in cross section, not ridged, internodes 0.3
8.7 cm long, stem indumentum of long, ap-
pressed or spreading, ascending bulla-based
hairs 0.21.6 mm long; nodal line absent.
Leaves opposite, decussate, narrowly elliptic,
1.510.1 × 0.73.6 cm, anisophyllous with
larger leaves to twice as large as smaller leaves
in a pair, grading to ± isophyllous, bases acute
to slightly rounded, apex acute to truncate,
apex often with the primary vein dividing into
23 divisions, venation acrodromous, leaves 5
veined, innermost pair of secondary veins aris-
ing 0.52.1 mm from the leaf base, 1.57mm
from the leaf margin at widest part of leaf,
tertiary veins 2.46.8 mm apart, leaf margins
appearing erose as a result of large marginal
teeth terminating in bulla-based hair and small-
er bulla-based hair produced between teeth,
adaxial leaf surface with sparse, poorly devel-
oped bulla-based hairs not entirely filling are-
oles, erect to reflexed, larger bulla-based hairs
surrounded by diminutive bulla-based hairs,
primary and secondary veins impressed, the
primary veins covered in narrower bulla-based
hairs than those of the lamina, those hairs erect,
tertiary veins inconspicuous, quaternary veins
not visible, sessile or short-stalked glandular
hairs also present, filiform to poorly developed
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
clavate-dendritic hairs produced from in be-
tween bulla-based hairs towards the leaf base
along the primary vein, abaxial surface pitted
as a result from upper leaf surface hairs, lamina
covered with sparse, erect, narrowly bulla-
based hairs, hairs mostly spreading along the
raised primary and secondary veins, sessile,
glandular hairs present throughout lamina and
along veins, tertiary veins conspicuous, raised,
generally connected by conspicuous, raised
quaternary veins, inconspicuous hair tuft
acarodomatia often present along the primary
vein in the axils of the secondary and tertiary
veins; petioles 0.21.8 cm long, covered in
ascending and slightly spreading, narrow
bulla-based hairs and sessile to short-stalked
glandular hairs in between bulla-based hairs.
Inflorescences terminal, open cymes, often
appearing axillary as a result of overtopping
growth of lateral shoots, 18 flowered, 1.8
3.8 × 1.22.4 cm, peduncles 00.7 cm long,
inflorescence bracts 1.32.7 mm long, narrowly
ovate, proximal inflorescence branches 5
11 mm long, bracteoles 1.12×0.2mm,nar-
rowly ovate. Flowers 1.52.5 mm wide, actino-
morphic, pedicellate, pedicels 0.71.5 mm
long; hypanthium 3.54.5 mm long, globose,
slightly constricted below the torus, covered in
bulla-based hairs to 1.8 mm long, and with
sparse, sessile to long-stalked glandular hairs,
free portion of hypanthium 0.81.2 mm long;
calyx lobes 56, 1.43.5 × 1.11.8 mm, adax-
ial surface with sessile to short-stalked, glandu-
lar hairs, abaxial surface covered in bulla-based
hairs, margin often with long-stalked glandular
hairs, calyx tube not tearing, 0.20.4 mm long,
with sparse, sessile to short-stalked, glandular
hairs along the adaxial surface of the calyx
tube, bulla-based hairs on the abaxial surface,
the apex (both abaxial and adaxial surfaces) of
tube with long-stalked, glandular hairs, calyx
teeth round in cross section, 1.54.4 × 0.2
0.4 mm, covered in bulla-based hairs and
sparse, sessile, glandular hairs; petals 56,
2.63.4 × 1.92.3 mm, pinkish to white, ovate,
with one margin ± ruffled, apices acute and
notched with a slightly bulla-based hair 0.4
0.5 mm long, produced at the petal apex, base
rounded; stamens 1012, filaments 0.91.1 mm
long, color unknown, glabrous, anthers 1.6
1.7 mm long, color unknown, elliptic, base
rounded, anther thecae 1.41.5 mm long, with
a single, dorsally oriented pore; style 3 mm
long, glabrous, linear or slightly expanded in
the middle; stigma punctate; ovary 3 locular,
2.13.5 × 2.93.1 mm, apex flattened or slight-
ly convex, pubescent, with delicate, bulla-based
hairs, crown present, of needle-like hairs.
Berries purple-black?, ca. 7.5 × 6 mm, globose.
Seeds obpyramidal, 0.40.5 mm long, testa
smooth, light brown, raphe light brown, ex-
tending the length of the seed.
Phenology.This species has been collected in
flower from June through September and in bud
in March.
Distribution.Miconia inaequipetiolata is
endemic to southern Hispaniola and is
known from the Massif de la Selle, Haiti,
and Sierra de Baoruco, Dominican Republic
(Fig. 4), from 7501250 m in elevation. It
most commonly occurs in moist, mixed
broadleaved-Pinus occidentalis Sw. forests
on calcareous soils.
Additional specimens examined: DOMINICAN
REPUBLIC. Prov. Barahona: Sierra de Bahoruco, municipio
La Ciénaga, paraje Las Filipinas, cañada La Vasila, 18°0657N,
71°0627W, 26 Jun 2005, Clase 4055 (FLAS, JBSD), Sierra de
Bahoruco, municipio La Ciénaga, paraje Las Filipinas, cañada
La Vasila, 18°0657N, 71°0627W, 11 Aug 2005, Clase 4087
(MO). Prov. Independencia: Sierra de Bahoruco, Municipio
Duvergé, Sección Puerto Escondido, zona fronteriza entre El
Aguacate y Sapotén, carretera internacional, yendo desde Puerto
Escondido hacia El Aguacate, UTM 19 214645E, 2027569N, 1
Sep 2011, Clase 6960 (FLAS, JBSD). Prov. Pedernales: Sierra
de Bahoruco, approx. 8 a 9 km al N de Las Mercedes; en el lugar
llamado El Mogote,18°09N, 71°40Oeste, 13 Mar 1993,
García 4746 (FLAS, JBSD), Hoyo de Pelempito, Bahoruco
Mts., 38 Jul 1971, Liogier 18150 (NY, UCMM). Aceitillar,
Cayo, 2425 Jul 1973, Liogier 19632 (JBSD, NY, UCMM),
26 km N of Cabo Rojo, 18°06N, 71°38W, 16 July 1992,
Thompson 10456 (FLAS).
Miconia inaequipetiolata is most likely sister
to M. karlkrugii and M. krugiana (Fig. 1),and
M. krugiana appears to be phenetically transition-
al between M. inaequipetiolata and M. karlkrugii.
For example, Miconia krugiana shares the short,
stout, and well developed bulla-based hairs of
M. karlkrugii, but also has a cymose inflores-
cence, the small flowers, and a single, apical hair
on the petals, as in M. inaequipetiolata.
Miconia inaequipetiolata is a rare species
known from only a handful of collections. There
is very little morphological variation within the
species and no morphological discontinuities
BRITTONIA [VOL
have been observed among the Dominican
Republic and Haitian collections.
2. Miconia karlkrugii Majure & Judd, J. Bot.
Res. Inst. Texas. 7: 268. 2013. Calycogonium
krugii Cogn. Bot. Jahrb. Syst. 4: 278. 1886.
Leandra krugii (Cogn.) Judd & Skean, Bull.
Florida State Mus., Biol. Sci. 36: 61. 1991.
Ossaea krugii (Cogn.) F.S. Axelrod, J. Bot.
Res. Inst. Texas 34: 214. 2011. Ossaea krugii
(Cogn.) Michelangeli & Bécquer in
Acevedo-Rodríguez & Strong. Smithsonian
Contributions to Botany. 98. 554. 2012.
comb. superfl. Type: Puerto Rico, Maricao
in declivibus montis Alegrillo.26 Nov
1884, [fl.], P. Sintenis 293 (holotype: BR
[BR0000005193479]); isotypes: BR, CORD
[CORD00003589], G-DC [G00317710],
GH, GOET [GOET007892], K
[K000535771], MPU [MPU013629], S,
US). Non Miconia krugii Cogn. (Fig. 5)
Evergreen shrubs, 12.5 m tall; stems
roundincrosssectionormoderatelyrectan-
gular in very young shoots, not ridged, in-
ternodes 0.26.4 cm long, stem indumentum
of ascending, appressed, bulla-based hairs to
0.6 mm long; nodal line present. Leaves
opposite, decussate, broadly elliptic, rotund,
orobovatewithobtusetowidelyacuteapi-
ces, and acute to rounded bases, strongly
anisophyllous with the larger leaf roughly
twice the size of the smaller leaf in a pair,
or grading to isophyllous, 15.8 × 0.753.5
cm, 5-nerved, the outer most pair mostly
intramarginal, venation acrodromous, veins
impressed on the adaxial surface and raised
on the abaxial surface, those of the adaxial
surface covered in bulla-based hairs, the in-
nermost pair of secondary veins arising from
05 mm from the leaf base, 27mmfrom
the leaf margin at the widest point of the
leaf, tertiary veins 1.63.5 mm apart com-
monly with quaternary veins joining them,
leaf margins serrate as a result of large ar-
cuate, bulla-based hairs, adaxial leaf surface
covered with bulla-based hairs, clavate-
dendritic hairs arising from between bulla-
based hairs, especially toward the leaf base,
although produced throughout the leaf sur-
face, sessile glands also produced from be-
tween bulla-based hairs, abaxial leaf surface
covered with bulla-based hairs, these mostly
flattened, sessile, glandular hairs throughout
abaxial leaf surface, as well as along major
veins, clavate-dendritic hairs produced from
the bases of the enlarged, bulla-based hairs
that are produced along the leaf margin,
domatia absent; petioles 0.10.7 cm, covered
in bulla-based hairs and often with clavate
dendritic hairs produced from between the
bulla-based hairs. Inflorescence terminal, re-
duced to one flower, sessile, or with pedicels
to 1 mm; floral bracts foliaceous, obovate to
FIG.4. Distribution of M. inaequipetiolata on Hispaniola.
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
FIG.5. Miconia karlkrugii. A) Habit. B) Adaxial leaf surface. C) Close-up of adaxial leaf surface. D) Abaxial leaf surface. E)
Close-up of abaxial leaf surface. F) Flower longitudinal section. G) Petal. H)Stamen.I) Fruit. J) Fruit longitudinal section. K)
Seed. A, D and I, J from Skean 3749,Bfrom Croat 60928,Cand Efrom Stimson 1260,FH, and Kfrom Wagner 1456.
BRITTONIA [VOL
oblanceolate, lanceolate, or linear 213 ×
0.58mm.Flowers25.2 cm wide, actino-
morphic or zygomorphic; hypanthium 5
8 mm long, globose, greatly constricted just
below the torus, covered in bulla-based, awl-
shaped, ascending hairs to 2 mm long, the
free portion of the hypanthium 1.63.2 mm
long; calyx lobes 3.54.9 × 2.73.1 mm,
oblong, covered in bulla-based hairs
abaxially, flattened, linear hairs produced
adaxially, these grading into flattened,
clavate-dendritic hairs produced along the
upper lobe margin and sometimes with
short-stalked, glandular hairs, calyx tube 3
4.1 mm long, tearing regularly corresponding
to lobes as fruit matures, lateral and apical
margins membranaceous, with the same hairs
as calyx lobes, although with a tuft of clavate-
dendritic hairs at the apex, these seen on both
halves of the tube after tearing of the calyx; calyx
teeth 59.5 × 0.50.9 mm, linear, round in cross
section or moderately flattened perpendicular to
the floral axis, covered in bulla-based hairs; petals
5-7, 1423 × 9.516 mm, white, glabrous, or
occasionally with a few short-stalked glandular
hairs on the abaxial surface towards the medial
portion of the petal, elliptic to obovate, mod-
erately to strongly clawed, asymmetric with
one margin ruffled (forming plications); sta-
mens 12, filaments 4.86.7 mm long, white,
glabrous, anthers 4.36.4 mm long, yellow,
subulate, the sterile portion extending into
two lateral, basal lobes surrounding the fila-
ment, anther thecae 3.95.9 mm long, opening
by 1 apical or slightly dorsally-oriented pore;
style 814 mm, glabrous, linear; stigma punc-
tate; ovary 46locular,47 × 2.78 mm, apex
acute, pubescent, covered in delicate, bulla-
based hairs, crown present of dense, needle-
like hairs. Berries bright red, globose, 910 ×
6.511 mm. Seeds obpyramidal to obovate,
0.91.1 mm long, testa smooth, with a dark
colored raphe extending the length of the seed.
Phenology.Miconia karlkrugii flowers
throughout the year.
Distribution.Miconia karlkrugii is restrict-
ed to the Cordillera Central of Puerto Rico
from 580875 m in elevation and is most
common in the Maricao State Forest. It also
has been collected in Susúa State Forest and
once in the municipality of Pañuelas (Fig. 6).
This species has been well collected, al-
though it is not very widespread. It is most
commonly found growing in serpentine soils
of moist, broadleaved, montane forests.
Additional specimens examined: PUERTO RICO.
Mpio. Maricao: Maricao St. Forest, 31 Dec 1961, Bro. Alain
9059 (A, NY, US), Maricao St. Forest, 26 Jun 1962, Bro. Alain
9216 (NY), Rio de Maricao, 2 Apr 1913, Britton 2433 (F, GH,
NY, US), Maricao St. Forest, along Hwy. 120, 18°9 N,
66°59'W, 3 Apr 1985, Croat 60928 (F, MICH, MO, US),
Maricao National Forest, 2 Apr 1964, Duke 7365 (MO, US),
Maricao, 1 Jul 1913, Hioram s.n. (NY), above Maricao, 17 Apr
1905, Horne s.n. (NY), Cordillera Central, Reserva Forestal
Maricao, near forest headquarters, just off of Rt. 120m along
trail paststone house, S of Maricao (ca. 8 km S of jct. of Rt. 120
& 105), 16 Jan 1992, Judd 6111 (FLAS), Monte del Estado,
Maricao, 24 Mar 1980, Liogier 30515 (NY), Maricao St.
Forest, 2729 May 1989, Liogier 36797 (NY), Bosque de
Maricao, Maricao, 21 Oct 1994, Liogier et al. 86 (NY),
Maricao Forest, 16 Mar 1959, Martorell et al. s.n. (MO,
NY), Maricao State Forest Rt. 120, km 15.3, by casa de piedra,
15 Nov 2012 Michelangeli 2021 (NY, SJ, UPR), Maricao,
Barrio Maricao Afuera, Maricao State Forest Reserve, Rd.
120, km 17.7, SE side of rd., N of Alto del Descanso,
18°09'76"N, 66°59'72"W, 18 May 1996, Skean 3749
(MICH), Maricao, 28 Apr 1935, Sargent 396 (US), Maricao,
19 Jul 1915, Stevens 8799 (NY), Maricao, 29 Feb 1968,
Wagner 1381 (DUKE). Mpio. Pañuelas: Adjuntas, in monte
cerrote, 27 May 1886, Sintensis 4435 (A, F, GH, MSC, NY).
Mpio. Sáb ana Grande: Maricao St. Forest Reserve, along Rt.
362 beyond Campamento and along Sendero Santana, 600
650m, 6 Jun 1992, Axelrod 4610 (NY), Maricao St. Forest
Reserve, along Rt. 362 beyond Campamento and along
Sendero Santana, 8 Feb 1992, Axelrod 3974 (NY), Maricao
to Monte Alegrillo, 3 Apr 1913, Britton 2547 (F, GH, MO, NY,
US), PR 120 ca. 7 km N of Sabana Grande, 27 Mar 1971,
Burch 3468 (DUKE, MO, NY), Maricao Insular Forest, 26 Jul
1935, Gerhart 615 (NY), Reserva Forestal Maricao, along
Hwy.120,4.34.5 mi S of jct. with Hwy. 105; 5.5 mi S of
Maricao, 30 Nov 1981, Hansen 9509 (MO), roadside at
Maricao preserve, 2 Jun 1970, Hill 196 (NY), Sangre (sic)
Grande to Maricao, 2230 Jan 1963, Howard 15363 (A,
DUKE), Maricao Insular Forest, S side of Cordillera Central,
near rd. 120 at km 9, hm 6., 20 Jun 1965, Stimson 1260
(DUKE, GH, MICH, MO, MSC, US), Maricao Insular
Forest, 1 May1945, Wadsworth s.n. (MO, NY), km 8.8 on
Rt. 120, Maricao Forest, 5 Apr 1968, Wagner 1456 (A,
DUKE), km 8.2 on Rt. 120 in Maricao Forest, 3 May1968,
Wagner 1505 (DUKE), km 1011, Rt. 120, km 15.4 in Maricao
Forest, 21 Mar 1969, Wagner 1832 (A, DUKE). Mpio. San
Germán: Reserva Forestal Maricao on Rt. 120, 2 Apr 1968,
Ahles 70715 (GH), Maricao St. Forest Reserve, old rd. near
stone house, 16 Jan 1992, Axelrod 3679 (NY), Maricao,
18°8'N 66°62'W, 11 Mar 2004, Christenhusz 3516 (S),
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
Maricao St. Forest, 17 Jun 1980, Liogier 30741 (NY), 15 Apr
1936, Nelez 788 (NY), Maricao St. Forest, Barrio Cain Alto,
along Guamá trail 0.20.3 km due SE of Campamento Buena
Vista, 23 Jun 1992, Proctor 48060 (MO), Maricao, Barrio Caín
Alto, Maricao Forest, ca. 100 m away from Casa de Piedra,
18°09'05"N 66°45'55"W, 28 Nov 1992, Reyes 8 (MO), Barrio
Hocunuce Alto, Maricao Forest Reserve, trail SSW out of
Capamento Buena Vista, 18°8'93"N 66°59'49"W, 14 Jun
1997, Skean 3808 (FLAS). Mpio. Sabana Grande-San
Germán: Cordillera Central, Reserva Forestal Maricao, S of
Maricao, Sendero Santana, starting near recreation area and
going to peakof Cerro Santana,boundary of Municipio of San
German and Sabana Grande, 16 Jan 1992, Judd 6116 (FLAS,
NY), Cordillera Central, Reserva Forestal Maricao, S of
Maricao, boundary of Municipio of San German and Sabana
Grande, peak and ridge of Cerro Santana, 600m, 16 Jan 1992,
Judd 6126 (FLAS, NY), Maricao Bosque Insular de Maricao,
Las Tetas de Cerro Gordo, along boundary with Mpio. de San
Germán, near observ. tower, 10 Jul 1959, Webster 8863 (GH,
MICH, US). Mpio. San Germán-Sabana Grande-Maricao:
Yauco, Bosque Susúa Alta, Bosque Estatal de Susúa, 6 Apr
1991, Garcia 3457 (MAPR, UPRRP), in the Reserva Maricao,
along Rt. 120 at the DNR station, 22 Jun 1986, Taylor 7102
(DUKE).
Miconia karlkrugii is sister to M. krugiana
(Fig. 1). Miconia karlkrugii is not highly vari-
able morphologically, although certain popula-
tions exhibit smaller leaf and flower sizes,
which may be correlated with habitat differ-
ences (e.g., Burch 3468, collected in scrub over
shale substrate).
3. Miconia krugiana (Cogn.) Majure & Judd J.
Bot. Res. Inst. Texas. 7: 269. 2013. Ossaea
krugiana Cogn. Monog. Phan. 7: 1048.
1891. Leandra krugiana (Cogn.) Judd &
Skean, Bull. Florida State Mus., Biol. Sci.
36: 61. 1991. Type: Puerto Rico, Adjuntas,
in sylva juxta flumen ad Junco, 24 June
1886, (bud), P. Sintenis 4642 (holotype: BR
[BR519349]; isotypes: A, CAS
[CAS0003714], CORD [CORD00003622],
F, GOET [GOET008048], G-DC
[G000328231], GH, K [K000329533-
K000329534], M [M0165768], MO, MSC,
NY, S, US) (Fig. 7)
Evergreen shrubs, 1.53 m tall; stems rect-
angular in cross section when young becoming
round in age, internodes 1.28.9 cm long;
nodal line present; stem indumentum of short,
triangular to awl-shaped, ascending, appressed
hairs to 0.6 mm long. Leaves opposite, decus-
sate, broadly elliptic to ovate, bases rounded to
acute, apex acuminate, strongly anisophyllous
with larger leaves roughly four times the size
grading to 1/3 larger than the size of the
FIG.6. Distribution of M. karlkrugii (closed circles), and M. krugiana (open circles) on Puerto Rico.
BRITTONIA [VOL
FIG.7. Miconia krugiana. A)Habit.B)Leaf.C) Leaf adaxial surface close-up. D) leaf abaxial surface close-up. E)Flower.F)
Petal abaxial surface. G) Petal adaxial surface. H)Stamen.I) Immature fruit. J) Fruit longitudinal section. B, C from Woodbury
s.n., A, DJfrom Skean 3820.
MAJURE ET AL.: MICONIA (MELASTOMATACEAE)2014]
smaller leaf in a pair, 211.7 × 1.26.3 cm, 7-
veined, with the outermost pair of veins
intramarginal, margins slightly serrate through
the development of stout arcuate, bulla-based
hairs, veins impressed on the adaxial surface
and raised on the abaxial leaf surface, inner-
most pair of secondary veins arising 1.7
10 mm above leaf base, 3.811 mm from leaf
margin at midleaf, tertiary veins 36.5 mm
apart, connected by quaternary veins; adaxial
leaf surface covered with longitudinally com-
pressed bulla-based hairs, these of large bulla-
based hairs surrounded by small bulla-based
hairs, not completely filling the leaf areoles,
thus the lamina clearly visible, sessile to short
stalked, glandular hairs present throughout the
surface of the lamina, clavate-dendritic hairs
produced from between the bulla-based hairs,
especially along the primary vein and towards
the leaf base, abaxial leaf surface covered with
sparse, bulla-based hairs, as well as sessile
glandular hairs, clavate-dendritic hairs pro-
duced at the base of the bulla-based hairs
along the leaf margin, domatia absent; petioles
0.52.4 cm long, covered in appressed, ascend-
ing bulla-based hairs, and clavate dendritic
hairs produce on the adaxial petiole surface
and densely so in the petiole axils.
Inflorescences terminal, produced on short
shoots which are soon overtopped by lateral
growth, long pedunculate forming a condensed
cyme often appearing as a glomerulate head,
of 49 flowers, 0.93.3 × 0.61.4 cm, pedun-
cle 0.92 cm long, bracts 1.83mm long,
ovate, bracteoles 0.51.3 × 0.20.3 mm, nar-
rowly ovate; pedicels 00.5 mm long; flowers
± zygomorphic with the anthers mostly all to
one side and the style on the opposite side,
hypanthium 2.63.2 mm long, globose, cov-
ered in large arcuate, bulla-based hairs to
1 mm long, not constricted below the torus;
calyx 56 merous; calyx lobes 0.81.4 × 1.1
1.3 mm; calyx tube 0.30.5 mm long, not
tearing; calyx teeth 1.21.9 × 0.6 mm, round
in cross section; petals 56, 3.94.9 × 2.83.7
mm, white to pink or red, obovate, asymmet-
ric, clawed, with one apical hair, appearing as
acaplike structure, to 0.1 mm long, produced
from the abaxial surface, and one to several
stalked, glandular hairs on the medial-abaxial
portion of the petal; stamens 1012; filaments
1.61.8 mm long, white?, glabrous; anthers
yellow, ovate, 11.2 mm long; anther thecae
0.81 mm long, opening by one dorsally ori-
ented pore; style 3.13.4 mm long, glabrous,
linear or slightly expanded in the middle; stig-
ma punctate; ovary 5 locular, 1.53.5 × 2.6
4.1 mm, apex pubescent with delicate, bulla-
based hairs, apex obtuse, crown present of
needle-like hairs. Berries ca. 4 × 5.2 mm,
color unknown, globose. Seeds obpyramidal,
0.71.1 mm long, testa smooth, raphe light
brown, extending the length of the seed.
Phenology.Miconia krugiana flowers from
May through August, although specimens are not
available from September through November, so
flowering time may extend into later months as
well. Material collected from December through
early March was sterile.
Distribution.This species is endemic to
Puerto Rico and is only known from the
Cordillera Central from 7501250 m in ele-
vation. It has primarily been collected in the
Parque del Estado Toro Negro but also is
known from the eastern part of the
Cordillera Central in Cayey (Fig. 6). This
species is more widely distributed than
M. karlkrugii but is less common and more
poorly known. Miconia krugiana occurs in
dense, moist, broadleaved, montane forests
commonly near streams.
Additional specimens examined: PUERTO RICO:
Mpio. Adjuntas: Arroyo de los Corchos, between Adjuntas
and Jujuya, 13 Mar 1915, Britton 5239 (NY), Cayey, Banks of
a stream, Guavate, 14 Dec 1963, Liogier 10400 (NY),
Adjuntas, Mt. Guilarte, 16 Jan 80, Liogier 30280 (NY),
Adjuntas, Guilarte, 27 May 1961, Woodbury 4881 (NY, US).
Mpio. Ciales: Toro Negro, 17 May 1905, Duke 7620T (MO),
Ciales, Toro Negro, 20 Aug 1994, Liogier 37342 (NY), Ciales,
Toro Negro, picnic area, near top oftrail to tower, 20 Feb 1961,
Woodbury s.n. (NY, P), Ciales, Toro Negro, Jun 1975,
Woodbury s.n. (NY). Mpio. Jayuya: Cerro de Punta, 23 Feb
1983, Liogier 33927 (MO, NY). Mpio. Orocovis: Doña
Juana, Toro Negro Mts., 10 Jul 1962, Alain 9421 (A, NY).
Mpio. Ponce: Barrio Anon, Toro Negro State Forest, Rt. 143,
km 15.6 near Cerro de Punta (turn off for this at km 18),
18°10'00"N 66°35'81"W, 15 Jun 1997, Skean 3820 (FLAS),
Ponce, S slopes of Cerro de Punta, 8 Jul 1959, Webster 8795
(GH, US). Mpio. Villalba: Central Mts. above Villalba, 12 Jun
1935, Holdridge 502 (NY).
Miconia krugiana is sister to M. karlkrugii
(Fig. 2). Miconia krugiana is homogeneous mor-
phologically, other than showing minor variation
in leaf size across populations.
BRITTONIA [VOL
Acknowledgments
We thank M. Parks for the beautiful illustra-
tions presented here, K. Neubig (FLAS) for help
with lab work, J.D. Skean, Jr. (Albion College)
and F. Axelrod (UCRRP) for information regard-
ing M. karlkrugii and M. krugiana.Wealsothank
numerous herbaria (A, DUKE, F, GH, JBSD,
MICH, MO, MSC, NY, S, US) for loans used in
our revision and UCMM for access to their col-
lections, as well as K. Perkins, T. Linder, and N.
Williams (FLAS) for handling loan materials. J.
Dan Skean, Jr. and Eldis Bécquer (HAJB) provid-
ed helpful comments on an earlier version of this
manuscript, and J. Dan Skean, Jr. provided photos
of M. krugiana. This work was supported by a
grant from the National Science Foundation
(BSR-0818399).
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APPENDIX 1. Voucher specimens used in our
phylogenetic analyses along with repository and
associated GenBank numbers for ETS, ITS, accD-
pasI and psbK-psbI. Missing data are represented
with an em dash ().
Calycogonium glabratum DC., Skean 4151 (FLAS), ,
EU055645, , KF821795; Calycogonium rhamnoideum
Naudin, Skean 5000 (FLAS), KF820605, KF821414,
KF819885, KF821806; Calycogonium turbinatum Urb. &
Ekman, Judd 8123, (FLAS), KF820592, KF821417,
KF819888, KF821810; Clidemia barbeyana Cogn.,
Michelangeli 1482 (NY), KF820622, KF821423, KF819901,
KF821825; Clidemia wrightii Griseb., Becquer 83852
(HAJB), KF820716, KF821470, KF819971, KF821916;
Miconia argentimuricata Majure & Judd, Becquer 86122
(HAJB), KF821302, KF821749, KF820496, KF822487;
Miconia asperifolia (Naud.) Majure & Judd, Judd 8300
(FLAS), KF821279, KF821732, KF820475, KF822465;
Miconia blancheana Urb., Clase 4190 (JBSD), KF821322,
EU055871, KF820511, KF822505; Miconia granulata (Urb.)
Majure & Judd, Becquer 82266 (FLAS), KJ933938,
KJ933987, KJ933895, KJ934040; Miconia heterophylla
(Naudin) M. Gómez, Skean 4233 (FLAS), KF820597, ,
KF819878, KF821798; Miconia hispidula (Cogn.) Judd,
Bécquer & Majure, Skean 3151 (FLAS), KF820598,
EU055648, KF819879, KF821799; Miconia inaequipetiolata
Majure & Judd, Clase 6960 (FLAS), KJ418739, KJ418734,
KJ418744, KJ418729; Miconia jashaferi Majure & Judd,
Becquer 85489 (HAJB), KF821304, EF418913, KF820498,
KF822489; Miconia karlkrugii Majure & Judd, Fritsch s.n.
(SPF), KF821293, KF821742, KF820488, KF822479;
Miconia krugiana (Cogn.) Majure & Judd, Skean 3820
(FLAS), ,KJ933991,, KJ934044; Miconia lima (Desr.)
M. Gómez, Majure 4334 (FLAS), KJ418740, KJ418735, ,
KJ418730; Miconia microphylla (C. Wright) M. Gómez,
Becquer 84641 (HAJB), KF820600, KF821411, KF819881,
KF821801; Miconia moensis (Britton) Alain, Michelangeli
1484 (NY), KF821126, KF821657, KF820338, KF822313;
Miconia norlindii (Urb.) Majure & judd, Becquer 83858
(HAJB), KF821317, , KF820508, KF822500; Miconia
ottoschmidtii (Urb.) Majure & judd, Becquer 82434 (FLAS),
KF821303, KF821750, KF820497, KF822488; Miconia
pedunculata Majure & Judd, Skean 4103 (FLAS),
KF821307, KJ418738, KF820501, KF822492; Miconia
umbellata (Mill.) Judd & Ionta, Judd 8084 (FLAS),
KF820711, EU055674, KF819967, KF821911; Miconia
vegaensis (Cogn.) Judd, Bécquer & Majure, Skean 4330
(FLAS), KF820715, , KF819970, KF821915; Ossaea
baracoen sis Borhidi & O. Muniz, Michelangeli 1470 (NY),
KF821281, KF821734, KF820477, KF822467; Ossaea
brachystemon Urb., Becquer 83871 (HAJB), KF821283,
KF821736, KF820479, KF822469; Ossaea hirsuta (Sw.)
Triana, Judd 8311 (FLAS), KF821292, KF821741,
KF820487, KF822478; Ossaea ovatifolia Urb., Becquer
82485 (FLAS), , KJ418733, KJ418743, KJ418728.
BRITTONIA [VOL
... Krugiophytum (Cogn.) Majure & Judd (Majure et al., 2015b); (vii) Miconia sect. Lima Majure & Judd (Majure & Judd, 2013); (viii) Miconia sect. ...
... With the recognition of a single genus within Miconieae , and the fact that most traditional sections of Miconia have been shown to be polyphyletic , a new taxonomic framework for the tribe has been presented in parts: new sections have been recognized or recircumscribed isolately (e. g. Judd and Ionta 2013;Judd et al., 2014aJudd et al., , 2014bMajure et al. 2015aMajure et al. , 2015bGoldenberg et al., 2018;Skean et al. 2018), but the bulk of the species have yet to be assingned to a infrageneric classification. After weighing the options of defining the entire clade as a section (with series or subsections within it) or as a supersection with three sections within it as done by Naudin (1850), we chose the latter. ...
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The following guide is aimed at aiding in the curation of herbarium collections in Melastomataceae, with an emphasis on the New World species. It contains a summary of the taxonomic realignments at the tribal and generic level within Neotropical taxa of Melastomataceae, as well as some general comments for other groups. A table with a generic linear sequence is also provided, as well as tables with new and synonymized New World genera since 2005 and all currently accepted species. Lastly, a table with the synonyms of over 1000 accepted Neotropical species that have been impacted by these generic realignments is also provided.
... To study the evolution of floral morphology and symmetry, we generated the most densely sampled phylogeny of the Miconieae to date. The molecular data set (5584 sequences) comprised sequences available in GenBank from previously published studies (Michelangeli et al. 2004Goldenberg et al. 2008;Martin et al. 2008;Reginato et al. 2010;Gamba and Almeda 2014;Kriebel et al. 2015;Majure et al. 2015aMajure et al. , 2015bMeirelles et al. 2015;Reginato and Michelangeli 2016), sequences available in GenBank but not used to date in any published studies, and 1025 sequences (18.4% of all sequences) newly generated for this work (app. 1; apps. 1, 2A, 2B are available online). ...
Chapter
Miconieae are the largest tribe in the family Melastomataceae with ca. 1900 described species. The Greater and Lesser Antilles harbor an impressive number of species from the Miconieae, which occur throughout the islands. Four major clades of Miconia are represented in the Antilles, the Caribbean, Chaenopleura, Mecranium, and Sagraea clades, corresponding to five in situ radiations. The Caribbean clade consists of ca. 160 spp., the Chaenopleura clade (= Miconia sect. Chaenopleura) of ca. 55 spp., Mecranium clade (= Miconia sect. Sagraeoides) of ca. 25 spp., and the Sagraea clade (= Miconia sect. Sagraea) of ca. 45 endemic spp. and is represented by two separate radiations in the Antilles. Likewise, numerous widespread continental species (ca. 45) from disparate subclades across Miconieae occur throughout the islands (presumably originating from many long-distance dispersal events), further increasing the Melastome diversity there. Each of these major clades is briefly characterized, and the evolutionary/phylogenetic patterns within each are discussed. Based on a phylogenetic framework, we discuss the patterns of diversification among the islands, as well as some of the biogeographic patterns of these fascinating radiations within the islands. We incorporate known data regarding ecological pressures and morphological evolution that likely have aided the diversification of this clade principally throughout the mountainous areas of the islands. Three new combinations are made in Miconia for Mecranium integrifolium, Miconia neibensis (Skean) Skean, Judd & Majure, comb. nov., Miconia neibensissubsp.alainii Skean, comb. nov., andMiconia neibensissubsp.integrifolia Skean, comb. nov.
Chapter
The tribe Miconieae is a strictly New World group of ca. 1900 species of mostly shrubs and small trees, but also including herbs, epiphytes, and climbers. The taxonomic history of the group is very complex with anything from 28 to 17 genera recognized in the tribe. Generic delimitations have often relied on a few morphological characters that are poorly defined, unevenly applied, and highly homoplasious. Thus, not surprisingly, most genera have been resolved as nonmonophyletic and rendered the highly species-rich Miconia as wildly paraphyletic. Attempts to find consistently stable clades across the tribe, that are also reciprocally monophyletic and diagnosable for the entire tribe have failed, leading most specialists to recognize a single genus within the tribe: Miconia. In this chapter, we summarize the taxonomic history of the tribe and the current status of phylogenetic and taxonomic studies in it. We then present the reasoning to recognize Miconia as the sole genus in Miconieae. Under our circumscription, Miconia becomes the seventh largest genus of flowering plants and notably the only one among the 10 largest genera of angiosperms that is restricted to the Neotropics.
Article
A new section, Miconia sect. Menendezia, is recognized, characterized, and assessed phylogenetically. The section includes three species: M. angustifolia, M. biflora, and M. urbanii. All three have fruits with constricted hypanthia, and traditionally were included within an extremely non-monophyletic Tetrazygia. The monophyly of the section is supported by molecular analyses and morphology (i.e., calyx teeth flattened parallel to the associated lobe and equaling to much longer than the calyx lobes, and possibly also hypanthium constricted and calyx tube regularly tearing between the lobes). In addition, the inflorescence branches are flattened, although this condition is only poorly developed in M. angustifolia. Miconia urbanii (incl. M. stahlii) and M. biflora are morphologically distinctive (i.e., as evidenced by their strongly flattened inflorescence axes, flowers with an elongate hypanthium and calyx lobes, large petals that are undulate on one side, and fully inferior, slightly to strongly 4-lobed ovary) and these characters are likely synapomorphic for the putative M. urbanii + M. biflora clade, which was recognized as the genus Menendezia by N. L. Britton. Species descriptions, nomenclatural information, specimen citations, and eco-geographical characterization for each of the three species are presented, along with an identification key. Section Menendezia is well supported as a subclade within the Caribbean clade of Miconia, a morphologically heterogeneous clade of ca. 160 species (out of a total of some 1900 species in the genus). Finally, the names Calycogonium biflorum, Tetrazygia angustifolia, T. stahlii, and T. urbanii are lectotypified.
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Chapter
Numerous DNA regions representing the nuclear and both organellar genomes are now available for comparative sequencing in plants (see Chapter 1); in addition, morphological and chemical data can also be obtained for phylogenetic analyses. With such a diversity of potential data sets available and the relative ease with which DNA sequences can be obtained, the acquisition of multiple data sets for the same suite of taxa is straightforward. As a result, the number of groups for which multiple data sets is available is increasing rapidly. Although it is readily apparent that multiple data sets are needed for estimating phylogenetic relationships reliably, it is also recognized that different genes may, in fact, possess different branching histories (see Chapter 10). Consequently, incorporating multiple data sets into phylogenetic studies is not a casual undertaking. Essential tasks in the analysis of multiple data sets include assessing congruence between different phylogenetic trees and data sets, and ascertaining whether multiple data sets should be combined into a single data matrix prior to phylogenetic reconstruction.
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