Revision of the Lima clade (Miconia sect. Lima, Miconieae, Melastomataceae) of the Greater Antilles

Abstract Miconia sect. Lima is an entirely Greater Antillean clade that consists of 19 known species of shrubs and small trees, which were previously recognized under the polyphyletic genera Leandra and Ossaea. The highest species richness in the clade is represented on Cuba (10 species), followed by Hispaniola (8 species) and then Jamaica (1 species). Here we present a taxonomic revision of the clade based on the study of species in the field, herbarium specimens, as well as a DNA-based phylogeny reconstruction. The Lima clade most likely originated on Cuba and then spread to Jamaica once and Hispaniola multiple times. Species of this clade can be recognized by the well developed bulla-based hairs of the adaxial leaf surface, as well as the clavate-dendritic hairs produced along the primary, secondary and tertiary veins of the adaxial leaf surface, mostly towards the leaf base, terminal inflorescences, acute petal apices, slightly bulla-based hairs produced subapically along the petal abaxial surface, and anthers with a dorso-basal appendage and a single, dorsally oriented pore. Descriptions, synonymies, along with distribution maps and illustrations/figures, are given for each species. Miconia pagnolensis sp. nov. is newly described in this revision.


Introduction
Tribe Miconieae (Melastomataceae) is a widely distributed, species rich (ca. 1800 species), Neotropical clade of trees and shrubs, which is most diverse in montane regions of Central and South America and the West Indies. Th e group has been divided into numerous genera, with ca. 16 recognized until only recently ). Phylogenetic analyses have shown that with few exceptions (e.g., Mecranium Hook.f.; see Skean 1993), all of these genera are polyphyletic , Michelangeli et al. 2004Kriebel et al. 2015, Reginato and Michelangeli 2016 and undiagnosable, if recognized in the broad sense as currently circumscribed (e.g., Leandra Raddi s.l.; . Generic limits in this group have long been considered problematic and arbitrary (Cogniaux 1891, Gleason 1932, Macbride 1941, Wurdack 1972, Judd 1989, Judd and Skean 1991. Th e Lima clade (i.e., Miconia sect. Lima Majure & Judd: Miconieae: Melastomataceae) as treated here, is a group of 19 known species that is restricted to the Greater Antilles (excluding Puerto Rico) and occurs mostly in high elevation, montane forests. Th e island of Cuba contains the largest number of endemic species (10), while 8 species are endemic to Hispaniola and only one species is found on Jamaica. No species are shared between or among islands. Species within this clade are distinctive in their production of bullabased hairs on most parts of the plant. Th e adaxial leaf surfaces are often nearly or entirely covered in these hairs ( Fig. 1A-B, D), giving the leaves the appearance of lizard or toad skin (Majure and Judd 2013a). Th e majority of species within this clade previously have been recognized under either Leandra or Ossaea DC. (Cogniaux 1886, Urban 1921, 1923, 1926-27, 1929, 1931Judd and Skean 1991, Liogier 2000, Michelangeli and Bécquer 2012, a result of the presence of acute petals produced by almost all members of the group. However, it is clear that Leandra and Ossaea are polyphyletic , Michelangeli et al. 2004, and thus the most judicious decision regarding generic circumscription proposed has been to recognize these taxa, as well as most other taxa in the Miconieae (Ionta et al. 2012, under an expanded Miconia s.l. (Ionta et al. 2012, Majure and Judd 2013a-b, Majure et al. 2013-c, Judd et al. 2014a,b, Michelangeli and Goldenberg 2016, which would include species formerly placed in the genera Calycogonium DC., Charianthus D. Don, Clidemia D. Don, Conostegia D. Don, Killipia Gleason, Leandra, Maieta Aubl., Mecranium, Necranium Britton, Ossaea, Pachyanthus A. Rich., Pleiochiton Naudin ex A. Gray, Tetrazygia Rich. ex DC., and Tococa Aubl. Miconieae can be recognized by their berry fruit, poorly developed anther appendages, the presence of druse crystals and associated lack of megastyloid crystals in various plant parts , Ionta et al. 2012, and lack of two pairs of large, decussate bracts subtending each fl ower, as in tribe Blakeeae (Penneys and Judd 2013).
Th e striking bulla-based hairs produced along the adaxial leaf surface, although best developed in the Lima clade, also are found in two other closely related clades, the Pseudolima (Miconia sect. Krugiophytum (Cogn.) Majure et al. 2014b) and Paralima clades (Miconia sect. Echinata Judd, Bécquer & Majure;Fig. 2). Th e production of these bulla-based hairs may represent a retained pleisiomorphy in the Lima, Paralima and Pseudolima clades, although character reconstructions using parsimony are equivocal . Having strongly bulla-based hairs that entirely or nearly completely cover the areoles, however, is a synapomorphy of a subclade (with reversals within it) of the Lima clade . Th e Lima clade can be diff erentiated from the Pseudolima clade by the growth architecture. Members of the Pseudolima clade produce terminal infl orescences that often appear axillary as a result of their defl ection to a lateral position from the rapid growth of lateral buds. Th e Lima clade has infl orescences that are clearly terminal . Th e Paralima clade is easily distinguished from the Lima clade by the production of globular stellate hairs on the stem, abaxial leaf surface and hypanthium, and several species of the Paralima clade produce truly axillary infl orescences or both axillary and terminal infl orescences (e.g., M. glomerulifl ora Judd, Bécquer & Majure, M. ovatifolia Judd, Bécquer & Majure;Majure et al. 2015, Judd et al. in prep).

Taxonomic history
Th e taxonomic history of the Lima clade has been one of unstable generic classifi cation beginning with Miconia lima (Desr.) M.Gómez. Miconia lima was the fi rst species described in this clade and was named by Desrousseaux (1797), as Melastoma lima Desr. He did not cite a detailed geographic locality but merely mentioned that the collection was made by Martin s.n. in Sainte Domingue (i.e., Haiti). He mentions that the upper leaf surface of the species appeared similar to that of a fi le, and hence the specifi c epithet "lima" was used. Th e species was then transferred to Clidemia in 1828 (de Candolle 1828), Sagraea in 1852 (Naudin 1852), Calycogonium in 1866 (Grisebach 1866) and then to Ossaea by Triana (1871-72). Gómez (1894) later transferred the species to Miconia, and Judd and Skean (1991) made the transfer to Leandra. Grisebach (1866) incorrectly cited a specimen from Cuba collected by C. Wright (189) as the type; this specimen (i.e., Wright 189) was later described as Ossaea cubana Alain (Alain 1955). Miconia asperifolia (Naudin) Majure & Judd was the second species of the clade to be described (Naudin 1852), as Clidemia asperifolia Naudin, apparently from a specimen sent to Naudin from Joseph Hooker (Naudin 1952). Hooker at the time had employed William Purdie to collect plants in Jamaica from 1843-1844 (Sinha 1972), who collected the fi rst known specimen of the species. It was then transferred to Oxymeris (Triana 1871-72) and then to Ossaea (Triana 1871-72). Cogniaux (1891) described material of the same species as Leandra eggersiana Cogn., which was later transferred to Ossaea by Urban (1921). It should be noted that MacFayden had already described M. asperifolia, as Clidemia hirsuta Macfad., however, his manuscript (Vol. 2 of his Flora of Jamaica) was apparently not eff ectively published before his untimely death. It is usually considered to have been unfi nished and thus not yet intended for publication; only incomplete copies of the work exist (Stafl eu and Cowan 1981, see also Porter-Utley 2015) other than one complete copy at BM. Incomplete copies of the fl ora were sent out by the British Museum of Natural History to HH, K, MICH, and NY (see Stafl eu and Cowan 1981). Two species collected by Charles Wright in Cuba, Miconia argentimuricata Majure & Judd and M. cubacinerea Majure & Judd, were the next species to be described by Grisebach (1866), as Calycogonium muricatum Griseb. and Clidemia cinerea Griseb., respectively. Miconia jashaferi Majure & Judd was then described by Britton and Wilson (1920) as Ossaea shaferi Britton & Wilson. Urban (1923, 1926, 1927, 1929, and Ekman (1929, 1931) then described a series of new species based on material primarily collected by the Swedish botanist Erik L. Ekman (1883Ekman ( -1931 Judd (2013a-b) and Majure et al. (2014a) subsequently described four more species from the clade from Haiti (M. phrynosomaderma Majure & Judd), the Dominican Republic (M. paralimoides Majure & Judd) and Cuba (M. bullotricha Bécquer & Majure, M. hirtistyla Majure & Judd), and one new species from Massif de la Hotte, Haiti, is described herein. Majure and Judd (2013a) transferred all of the previously described species in this clade to Miconia based on our current understanding of the polyphyletic nature of the genera in which they had been placed , Michelangeli et al. 2004, Majure et al. 2014c.

Morphology
Habit and stems. Species within the Lima clade form evergreen shrubs to small trees ranging in height from 0.5 to 5 m tall. Th e height of several species, however, is completely unknown. Stems are rectangular or cylindrical in cross section and lack longitudinal ridges (Fig. 9M).
Leaves. Leaves are opposite, decussate and generally slightly anisophyllous. Th e blades are elliptical, ovate or lanceolate (narrowly ovate), with crenulate or dentate margins that are generally obscured by bulla-based hairs ( Fig. 1E; see Indumentum below); the margins may be fl at or slightly revolute. Venation is acrodromous with the 1-3 pairs of secondary veins arching towards the leaf apex; these are basal to suprabasal. Tertiary veins are percurrent, more or less perpendicular to the midvein and often connected by quaternary veins. Leaf shape in certain species is quite useful for species delimitation, such as the elliptical-rhombic leaves of Miconia marigotiana, which are unique in the group.
Indumentum. Most plants parts within this group are at least to some degree covered in bulla-based hairs. Th ese bulla-based hairs may be lacrimal as in M. pedunculata (Fig. 1B), volcaniform as in M. argentimuricata, M. lima, M. limoides, M. marigotiana, M. paralimoides and several other species (Fig. 1D), or granulate in shape as in M. norlindii and M. granulata (Fig. 7B). Th e degree of bulla-based hair coverage, shape and placement or orientation can be useful for identifying species. Certain species show complete leaf areole coverage, on the adaxial surface, by volcaniform, bulla-based hairs (Fig. 1D), as in M. lima, M. limoides, M. marigotiana and M. paralimoides. Conversely, M. jashaferi, M. hirtistyla and M. cubacinerea have poorly-formed, slender, lacrimalshaped bulla-based hairs that cover most, but not all, of the adaxial leaf surface. Clavatedendritic hairs are common on most species at the leaf blade bases (where the petiole enters the leaf) and are most easily observed on developing leaves. Miconia jashaferi, however, exhibits fi liform hairs at this position. Clavate-dendritic hairs can commonly be found also at the apices of the calyx tube in most species (but see Morphological Evolution below). Most species within the Lima clade exhibit tranluscent or colored glandular hairs, which may be present on most portions of the plant body (stem, both leaf surfaces, hypanthium). Th ese are smaller than and occur between (at the base) or below the bulla-based hairs. Domatia composed of bulla-based or linear hairs are common in certain species, generally at the union of the primary and secondary veins but can also be found at the unions of the tertiary and primary, as well as tertiary and secondary veins (e.g., M. asperifolia, M. hybophylla). Th e directionality of the bulla-based hairs of the stem of specifi c species is very useful for species delimitation. For example, only M. limoides and M. phrynosomaderma exhibit downward oriented hairs on the stem surface.
Infl orescences. Infl orescences are terminal in all species of the Lima clade and are formed typically from 3-fl owered dichasia; infl orescences are generally quickly surpassed by rapidly growing, lateral shoots. Th e fl owers may be pedicellate or sessile and thus form open to relatively dense, and even slighly capitate (e.g., M. jashaferi; Fig. 3A) cymose infl orescences. Bracts subtending 3-fl owered dichasia are either granulate, lacrimal or in some taxa (e.g., M. pedunculata; Fig. 22A) foliose. Bracteoles are similar in form to the bracts.
Hypanthium and calyx lobes. Th e hypanthia are mostly 4-5 lobed, or mostly speherical (Fig. 12H). However, slight lobing is often obscured by the production of dense bulla-based hairs. Calyx lobes are triangular with acute to acuminate apices (Fig. 3E). Th e calyx teeth may be equal or longer than the calyx lobes and are terete ( Fig. 3E) and often refl exing when in fruit.
Corollas. Corollas are actinomorphic or nearly so, with 4-5 ovate to obovate or oblong petals; these are symmetric or slightly assymetric with acute to acuminate apices. Th e petal apices often exhibit 1-several bulla-based hairs on the abaxial surface (e.g., Fig. 14B). Petals are white, rose, red, purple or white with a purple tinge abaxially and are generally refl exed in fl ower.
Stamens. Stamens are 8-10 with glabrous fi laments. Th e anthers are white to yellow and may or may not have a dorso-basal appendage; they typically have one, dorsally-inclined pore (Fig. 3C).
Gynoecium. Th e gyneocium in the Lima clade ranges from 2-7 carpellate, with M. marigotiana being the only species with two carpels (Fig. 25G). Th e ovary is inferior with axile placentation, with the placentas generally deeply intruded into the ovary locule. Th e upper portion of the ovary may be glabrous or pubescent. Th e style is straight or slightly curved with a punctiform stigma. Only M. hirtistyla exhibits a pubescent style (Fig. 5F), while the rest of the species have glabrous styles. However, a crown of needle-like hairs often is present at the base of the styles (along ovary apex) of most species (e.g., Fig. 3E).
Fruit. Fruit in the Lima clade are multi-seeded, mostly spherical berries that range in color from purple, purple-black to black (Figs 9I,N,15F,19C,27D). Th ere is still much missing data for fruit color and size for numerous taxa within this group, as fruit are not well-preserved in herbarium specimens, and many specimens lack fruit altogether.
Phylogenetics and evolution  reconstructed a phylogeny of the Lima clade using fi ve plastid intergenic spacers (accD-psaI, psbK-psbI, rpl32-trnL, trnH-psbA, trnV-ndhC) and two nuclear, ribosomal loci (ETS and ITS). Th ey included 11 of the 19 species of the Lima clade, i.e., all species for which material was available in silica from fi eld collections or from "recently" collected herbarium material (e.g., within the last 10 years). Th ey were able to include two accessions per species for six of the included species (Fig. 2). Other closely related members of the Caribbean clade were included as outgroup taxa (i.e.,  Judd & Ionta). Th e phylogeny presented here is based on . Th e Lima + Calycopteris + Paralima + Pseudolima + Calycodomatia clade is referred to as the Sandpaper clade, because of the many species of the group that have bulla-based hairs on the upper leaf surface (see Fig. 2).
Th e Lima clade is most closely related to the Reducens clade, composed of Miconia sect. Calycopteris, sect. Krugiophytum and sect. Calycodomatia. Most of the members of those other three sections have reduced infl orescences and lack the conspicuous bullabased hairs of the Lima clade (except for sect. Krugiophytum). Th ey also have conspicuous to reduced globular-stellate hairs, again, except for sect. Krugiophytum, whose members possess sessile-glandular hairs like those of the Lima clade .
Within the Lima clade, Miconia jashaferi is resolved as sister to the remaining species, although other putative close relatives of M. jashaferi have not been sampled (e.g., M. cubacinerea, M. hirtistyla, M. tentaculicapitata). It is most likely that these four species form a subclade sister to the rest of the Lima clade, as they have the putative synapomorphies of leaf-like bracts and bracteoles, long, fi liform hairs on the adaxial calyx lobe and calyx tube apex (homologs of clavate-dendritic hairs on other species of the clade; , the lack of dorso-basal anther appendages and 3-locular ovaries. In our phylogeny, the fi rst two diverging members of the Lima clade, M. jashaferi and M. norlindii, are Cuban species, as are other putative relatives of M. jashaferi not included in our analyses (i.e., M. cubacinerea, M. hirtistyla, M. tentaculicapitata). Th e M. lima complex of Hispaniola (sensu Majure and Judd 2013a,b), composed of M. lima, M. limoides, and M. paralimoides in our phylogeny, are phylogenetically adjacent, however, M. paralimoides apparently is more closely related to M. pedunculata than to either M. lima or M. limoides, contrary to relationships initially suspected based on morphology alone (Majure and Judd 2013b). Miconia limoides is sister to M. lima, and the Cuban M. ottoschmidtii is unresolved in the same subclade as well (i.e., the Bulliformes subclade; . Th us, M. ottoschmidtii and M. pedunculata must be included in what Majure and Judd (2013a) termed the M. lima complex, which we refer to here as the Bulliformes subclade. Th e Bulliformes subclade is restricted to Hispaniola and Cuba. Miconia marigotiana and M. phrynosomaderma also are likely members of the Bulliformes subclade, although they have not been recollected since the type material was collected by Ekman. Th ey are linked to the Bulliformes subclade by their bulla-based hairs on the adaxial leaf surface almost completely fi lling the leaf areoles, as well as the descending stem hairs in M. phrynosomaderma (found otherwise only in M. limoides of the Bulliformes subclade). Another subclade is formed by M. argentimuricata, M. asperifolia, M. granulata and M. pagnolensis (i.e., the Acuminata subclade) represented by members from Cuba, Haiti, and Jamaica. Miconia cubana and M. hybophylla also are most likely members of this subclade, as they produce large, open infl orescences (i.e., in M. cubana and M. hybophylla, as in M. asperifolia and M. argentimuricata). Th ey also tend to have less well-developed bulla-based hairs on the upper leaf surface, similar to other members of the Acuminata clade. Miconia hybophylla also is linked to M. asperifolia by the production of well-developed domatia in the axils of the primary and secondary, as well as the primary and tertiary and secondary and tertiary veins. Miconia bullotricha may also form part of the Acuminata subclade in that it produces defl exed infl orescences, which are similar to the partially defl exed infl orescences in M. granulata, demonstrates a similar leaf shape (narrowly ovate), as do both M. granulata and M. argentimuricata in many specimens, and occurs over serpentine soils, as do both M. argentimuricata and M. granulata. Miconia argentimuricata, M. asperifolia, M. jashaferi, M. lima, M. norlindii, M. ottoschmidtii, and M. pedunculata are likely cladospecies (sensu Donoghue 1985), as multiple accessions of each of these species form clades in our phylogeny Fig. 2). We were unable to assess the monophyly of the rest of the species, as multiple accessions were unavailable for phylogenetic analysis. However, considering the morphological apomorphies uniting the populations of most of the other members of the clade, they also likely represent cladospecies (see Judd 2013a-b, Majure et al. 2014a).

Biogeography
Ancestral area reconstruction, as carried out in , suggests that the Lima clade evolved in the northeastern mountains of Cuba (Sierra de Baracoa, Moa, and Cristal) and then subsequently spread to other mountain ranges in Cuba (Sierra Maestra, Trinidad Mts. and Pinar del Río), as well as to Hispaniola (likely four times) and Jamaica (once). Th ere were three separate movements into the Sierra Maestra of Cuba represented by M. argentimuricata, M. norlindii and M. ottoschmidtii  .
Th e Lima clade appears to have evolved primarily on serpentine soils of Cuba and then dispersed and adapted several times to limestone-derived soils, soils mixed with limestone, or other substrates (e.g., clay soils in M. asperifolia; . A number of taxa within the Lima clade are still mostly restricted to serpentine soils (M. bullotricha, M. granulata, M. jashaferi), or at least occur in parts of their range on serpentine (M. argentimuricata, M. ottoschmidtii).

Morphological evolution
Th ere are very few morphological synapomorphies for the Lima clade, but the production of clavate-dendritic hairs on the upper leaf surface is a synapomorphy of a subclade of the group (only excluding M. jashaferi in our analyses), as is the production of dilated styles (excluding only M. jashaferi and M. norlindii, the two basalmost taxa in our phylogeny). Th e production of very well-developed bulla-based hairs on the upper leaf surface that mostly fi ll the areoles also is a synapomorphy of a subclade (excluding M. jashaferi and M. norlindii), and the production of sessile, glandular hairs is a synapomorphy for the entire clade. Th e sessile-glandular hairs on the upper and lower leaf surfaces are homologs to the globular-stellate hairs produced in close relatives (Calycopteris and Calycodomatia clades;  and have also evolved in the Pseudolima clade, as well as a few members of the Paralima clade. Th us, although the sessile-glandular hair type is synapomorphic for the Lima clade, it is not a unique synapomorphy within the Sandpaper clade . Suites of characters that can be used to diagnose members of the Lima clade include multicellular, bulla-based hairs occuring over most surfaces of the plant, including the ad-and abaxial leaf surface, petioles, stem, infl orescence axes and hypanthia; sessile-pigmented glands on the lower leaf surface, as well as on the hypanthium; deciduous, dendritic hairs on adaxial leaf surface arising from the lamina between the bulla-based hairs, these long stipitate and fl attened or clavate at the apex; ovate or oblong-triangular petals with slightly bulla-based, subapical hairs, these usually extending past the acute apex of the petal. Other characters unifying most species in this clade are anthers with a dorso-basal appendage and dorsally-oriented single pore, hypanthia that are constricted below the torus, and which are often slightly 4-lobed, styles that are slightly expanded (dilated) in the middle, and obpyramidal seeds with a smooth testa and dark raphe extending the length of the seed.

Ploidy
Chromosome numbers are unknown in this group.

Species concepts
Criteria for the recognition of species in this group are based upon morphological cohesiveness or the morphological-phenetic/diagnostic species concepts (Judd 1981, Wheeler and Platnick 2000, Judd 2007, as well as phylogenetic or apomorphic species concepts (Donoghue 1985, Judd 2007, thus unifying our knowledge regarding the evolutionary history of a species along with its morphological characteristics (i.e., producing a more unifi ed species concept; DeQueiroz 2007). A species thus consists of populations of individuals that can be morphologically diagnosed from other populations based on synapomorphies or suites of characters, and when the information is available, also form part of the same lineage or clade. Species consisting of metaspecies complexes and resulting from the invasion of new niches by closely related and subsequently divergent populations (peri-or parapatric speciation, or peripherial isolate speciation) would appear to be a highly likely in this clade, considering the restriction of many of these taxa to high elevation mountain ranges (see Judd et al. 2015a for an example of this). A given population evolving within a metapopulation could easily diverge from the rest of the lineage via niche specialization or other processes leading to allopatric speciation. However, within the Lima clade, we do not have direct evidence of speciation via metaspecies lineages; all of the species included in phylogenetic analyses with multiple accessions appear to form clades. So it appears that the most common mode of speciation in this group is through cladogenesis, which may be brought about through the restriction of many species to specifi c ecological niches (e.g., high elevation cloud forests on limestone-derived, clay soils, or other substrates or low elevation sclerophyllous forests on serpentine soils). Th us, most species within the Lima clade have at least a few apomorphic characters (uniting their populations), as well as quantitative character diff erences, which may or may not be apomorphies. We do not have any information regarding ploidy in this group, so we cannot determine whether or not hybridization and subsequent genome duplication may have played a role in the origin and subsequent evolution of any of the species.

Materials and methods
A total of 894 herbarium specimens were observed and measured to determine morphological variability among and within species of the Lima clade. Herbaria from which specimens were studied are A, ALBC, BR, F, FTG, G, GH, HAJB, IJ, JBSD, JE, K, MICH, MO, NY, S, UCMM, and US. Abbreviations follow Th iers (2012). Acronyms and citation of collection numbers for Cuban specimens follows Regalado Gabancho et al. (2008). Altogether, more than 100 quantitative and qualitative morphological char-acters were analyzed from herbarium specimens, photos, alcohol preserved material, or from live plants in the fi eld. Phenological, distributional and ecological data were derived mostly from specimen label data or observations taken in the fi eld. Measurements for reproductive characters (e.g., hypanthium length, calyx tube, lobe and tooth length, ovary length and width, etc.) were based on Judd (2007) and were taken from herbarium material rehydrated via boiling for several minutes in water mixed with a detergent (see Judd 2007). Illustrations were based on herbarium specimens. Typifi cations of species described by Urban are carried out, in part, using the criteria of Bécquer (2012). However, as Urban never actually explicitly stated the repository of type material, and there is potential confusion as to where holotypes may have originally been deposited, since all specimens of Melastomataceae were lost in the allied bombings of Berlin during the second World War, we have chosen to lectotypify those species for clarity (see also McNeill 2014).
Although Alain and Liogier both refer to Henri Alain Liogier, we maintain those entries seperately, as Liogier used only the name, Bro. Alain, while collecting in Cuba, but generally used Henri Alain Liogier (or Alain Henri Liogier) while collecting in Hispaniola after he left the Catholic religious order. Collections made by Erik Ekman in Hispaniola are preceeded with an H, as his collection numbers were started anew during his work in Haiti and the Dominican Republic (see also Index to numbered collections). Description. Evergreen shrubs to small trees; young stems terete, elliptic or slightly rectangular in cross section, lacking longitudinal ridges, the indumentum of dense bullabased hairs, these long appressed, spreading, or recurved, or short and granulate. Leaves opposite, slightly anisophyllous; blade elliptical, ovate, or lanceolate, the margin crenulate to dentate, these crenulations/dentations obscured by large bulla-based hairs, which slightly fold over the leaf margin, producing in some cases a moderately revolute margin, the indumentum of adaxial leaf surface typically of broad bulla-based hairs and more or less fi lling the areoles, although sometimes these hairs relatively narrow and wide-spaced, not fi lling the areoles, with long-stemmed, clavate-dentritic hairs produced along the primary, secondary and tertiary veins from between the bulla-based hairs, and also sessile to short-stalked glandular hairs present on all parts of the lamina (between bulla-based hairs), the abaxial leaf surface variously covered by narrow bulla-based hairs, these either long and well developed or short and granulate, these appressed, spreading, or erect, the lamina with sparse, sessile glands, the venation acrodromous, with secondary veins arching toward leaf apex, 1 to 3 pairs, basal to suprabasal, tertiary veins percurrent, more or less perpendicular to the midvein, sometimes mostly obscured by bulla-based hairs on the adaxial leaf surface, connected by quaternary veins, the primary, secondary, tertiary and quaternary veins mostly impressed on the adaxial surface and raised on the abaxial surface, domatia present or absent, occurring at the junctions of primary, secondary and tertiary veins, forming a pocket-like structure in the axils of the primary and innermost secondary veins or formed from a tuft of hairs in the vein axils. Infl orescences terminal, although often surpassed by the rapid growth of axillary shoots, the fl owers in 3-fl owered dichasia, sessile, subsessile or pedicellate, thus forming open cymes or sessile and nearly headlike clusters. Flowers 4-5(7)-merous, mostly actinomorphic or nearly so; hypanthium 4-5-lobed, the lobes sometimes obscured by retrorse or antrorse bulla-based hairs, bulla-based hairs long and well developed, or granulate, hypanthium also with sessile glands; calyx lobes triangular, acute to acuminate, often covered by sessile glands throughout the adaxial surface or such glands restricted to the apex of the adaxial surface, abaxial surface covered in bulla-based hairs and more or less sessile glands; calyx teeth ca. equal to or longer than calyx lobes, terete, mostly refl exed in fruit, covered in long and well developed, or granulate bulla-based hairs, sessile glands present or absent; calyx tube often with long stemmed, clavate-dendritic hairs produced from apex along the margin, sessile glands more or less present on adaxial surface, abaxial surface covered in bulla-based hairs; petals ovate to obovate or slightly oblong, symmetric or asymmetric, white, red, rose, purple, or white with purple tinge abaxially, apices acute to acuminate, with moderately bulla-based hairs produced from the abaxial surfaces just below the petal apex and occasionally from the medial portion of the petal as well; stamens 8-10, not geniculate, the fi laments glabrous, the anthers with or without a small dorso-basal appendage and a single, dorsally inclined pore; style straight to moderately curved, generally expanded in the middle, the stigma punctiform; ovary 2-5(7) locular, more or less inferior, with axile placentation, the placenta intruded into each locule, the ovary apex without a collar but commonly with a crown of multicellular hairs, the upper portion of the ovary pubescent (bulla-based hairs) to mostly glabrous (i.e., with only crown hairs present). Fruit a globose and slightly 4-or 5-lobed berry, purple-black at maturity. Seeds angular, obpyramidal, obovoid to obovoid-falcate, with a linear to oblong, dark colored raphe that extends the length of the seed; testa smooth; appendage absent.

Miconia
Miconia sect. Lima is a clade of 19 species restricted to the Greater Antilles (excluding Puerto Rico).

Type. Based on Ossaea shaferi Britton & Wilson
Description. Evergreen shrub, 1-1.5 m tall; stems round in cross section, not ridged, the internodes 0.4-9.5 cm long, stem indumentum of spreading to descend-ing bulla-based hairs, the apices recurved upwards, hairs to 2.3 mm long; nodal line absent. Leaves opposite, decussate, ovate or elliptic, 2.9-12.8 × 1.8-4.7 cm, slightly anisophyllous, apex acute to acuminate, leaf base acute to rounded, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the outermost usually intramarginal, secondary veins mostly basal, the innermost pair suprabasal, produced 2-13 mm from leaf base, positioned 3-7 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 2-6 mm apart at midleaf, impressed, but clearly visible on the adaxial leaf surface, intertertiary veins present, usually prominent, tertiary veins often joined by quaternary veins; adaxial leaf surface densely covered in bulla-based hairs giving the upper leaf a velvety appearance, these mostly fi lling the leaf areoles, widest hair bases to 1.1 mm, apices of bulla-based hairs mostly recurved, young leaf adaxial surface not producing long-stemmed, clavatedentritic hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bullabased hairs, these erect, those along the primary, secondary, and tertiary veins larger than hairs on the clearly visible lamina, lamina with a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile to short stipitate, glandular hairs produced primarily along major and minor veins but also occasionally throughout the lamina; petioles 0.4-1.3 cm long, covered in bulla-based hairs on both surfaces, these spreading with apices recurved upwards. Infl orescences terminal, pendant, forming mostly 5-6 fl owered glomerules, only occasionally branching, 1.2-3.9 × 1.4-2.7 cm, the peduncle 0.2-2.3 cm long, proximal infl orescence branches absent to 5.5 mm long, pedicels absent; bracts foliaceous, oblong to ovate, 5.7-11 mm long; bracteoles fl at, foliaceous, ovate to broadly ovate or rotund, 2.9-3.7 × 1-1.7 mm, the margins and apex with long bulla-based hairs and fi liform gland-headed hairs, with sessile, glandular hairs along the adaxial surface and bulla-based hairs covering the abaxial surface, the adaxial surface black in herbarium specimens. Flowers 4-6-merous, sessile; hypanthium 3.4-6.5 mm long, short-oblong to globose, unlobed, slightly constricted below the torus; free portion of the hypanthium 1-1.3 mm long, abaxial surface covered in bulla-based hairs to 2.5 mm long, and occasional, sessile, glandular hairs near the bases of the bulla-based hairs, adaxial surface (i.e., free portion) with few, short, linear hairs and sessile, glandular hairs; calyx teeth 4-6, 6.8-7.5 × 0.2-0.4 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes more or less triangular, apex acute to rounded, 0.9-1.7 × 1-1.2 mm, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs, as well as fi liform hairs adaxially, these often expanded and fl attened at the apex or gland-headed; calyx tube not tearing, 0.2-0.4 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially, fi liform hairs produced from the apex of the tube; petals 4-6, white, broadly elliptic to obovate, 5.3-5.6 × 2.1-2.7 mm, with an acute or acuminate apex, without or with one slightly bulla-based hair produced abaxially just below the apex to 0.1 mm long; stamens 8-12, fi laments 2.2-2.6 mm long, glabrous, anthers 1.8-2 mm long, with one dorsally oriented pore, anther thecae 1.5-1.7 mm long, without a dorso-basal appendage; style 5.4-6 mm long, glabrous, not or only slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, linear to elongate-triangular (needle-like) hairs, which are notably longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 2-5.3 × 1.9-4.3 mm, apex fl at (not upraised), with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded pla-centa, 3-locular; berries globose, slightly 4-lobed, purple at maturity, 7.5 mm long (including calyx tube), 7.3 mm wide; seeds 0.8-1.6 mm long, obpyramidal, often falcate, testa smooth, light brown, raphe black, smooth, extending the length of the seed.
Phenology. Miconia jashaferi has been collected fl owering and in immature fruit from July through August. Individuals have been collected in mature fruit in November and December. Distribution (Fig. 4). Miconia jashaferi is known from the mountains of northeastern Cuba, including the Baracoa, Moa and Toa regions, as well as the Sierra de Cristal.
Ecology. Miconia jashaferi grows in thorny, xerophytic scrub and semidry montane rainforest on serpentine soils from 110 -800 m in elevation. Some associated melastomes include Miconia baracoensis Urb., Pachyanthus reticulatus Britton & P.Wilson, Miconia moensis (Britton) Alain, Mecranium integrifolium Triana subsp. alainii Skean and Miconia walterjuddii Bécquer & Michelangeli. Conservation status. Miconia jashaferi is a locally widespread species in eastern Cuba ( Fig. 4) and is known from numerous locations from a range of elevations and forest types. Bécquer (2007) regarded the species as threatened; the species is found in both Parque Nacional Alejandro de Humboldt and Parque Nacional Pico Cristal. Hoewever, widespread mining practices around the Moa-Toa region may negatively aff ect localized populations of this species. Likewise, populations are highly fragemented and occupy less than 500 km 2 , so we suggest this species have a preliminary conservation assessment of endangered.
Discussion. Miconia jashaferi is resolved as sister to the rest of the Lima clade ( Fig. 2) in our phylogenetic analyses. However, three other species likely form a clade with M. jashaferi (M. cubacinerea, M. hirtistyla, and M. tentaculicapitata). All four of these species share the characters of large foliaceous bracts subtending the fl owers, long calyx teeth, long, fi liform hairs produced on the adaxial surface of the calyx tube, 3-carpellate ovaries and relatively large fruit compared with the rest of the members of the clade. Miconia jashaferi has often been confused with Miconia ovatifolia (Urb.) Judd, Bécquer & Majure (=Ossaea ovatifolia Urb.) of the Paralima clade , as the adaxial leaf indumentum is somewhat similar, consisting of poorly developed bulla-based hairs (Figs 3, 9A-D). Th ese two taxa are easily separated by hair type, in that M. ovatifolia has globular-stellate hairs on most surfaces of the plant and axillary, as well as obviously terminal infl orescences.
Th e stem indumentum of Miconia jashaferi initially is reddish or purplish and then quickly turns white, in what appears to be the death of the hairs produced, which are eventually shed entirely.   Description. Evergreen shrub (height unknown); stems round in cross section, not ridged, the internodes 0.4-3.3 cm long, stem indumentum of bulla-based hairs to 1.6 mm long, these shaggy, spreading to slightly descending; nodal line absent. Leaves opposite, decussate, ovate to elliptic, not falcate, 1.6-8.2 × 1.4-3.9 cm, slightly to strongly anisophyllous (larger leaves at a node to twice as large as the smaller leaf), dark brown when dried, apex broadly acute, base broadly acute to rounded, margin dentate, the dentations obscure, each covered in one large bulla-based hair, venation acrodromous, 7-veined, the midvein and 3 pairs of arching secondary veins, secondary veins mostly basal, the innermost pair, suprabasal, produced 3-9 mm from leaf base, positioned 2.5-11 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.5-4.1 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface with primary, secondary and tertiary veins impressed, quaternary veins obscure, abaxial surface with all veins conspicuously raised; adaxial leaf surface covered in well developed but narrow bulla-based hairs mostly but not entirely covering the leaf areoles, widest hair bases to 0.8 mm, apices of bulla-based hairs mostly erect to recurved, sessile, glandular, hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bullabased hairs, these mostly erect with undulate apices, those along the primary, secondary, and tertiary veins spreading and larger than hairs produced throughout the lamina, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile, black, glandular hairs produced along all major and minor veins, domatia of multicellular, elongate hairs, abundant in axils of primary and secondary veins, as well as the axils of the primary and secondary with tertiary veins; petioles 0.4-1.8 cm long, covered in spreading, bulla-based hairs on both surfaces. Infl orescences terminal, cymose, 2-5 fl owered, fl owers mostly produced in glomerulate clusters, 1.3-2.4 × 1.2-3.8 cm, the peduncle 0.6-1.3 cm long, proximal infl orescence branches 0.8-1.1 mm long, pedicels absent; bracts ovate to elliptic, foliaceous, 5-17 mm long; bracteoles foliaceous, elliptic, 2.8-4.3 × 1.7-2.1 mm, covered in bulla-based hairs marginally and abaxially and glabrous abaxially or with fi liform hairs towards the base. Flowers 6-merous, sessile. Hypanthium 2.6-3.2 mm long, short-oblong to globose, unlobed, slightly constricted below the torus, free portion of the hypanthium 1-1.4 mm long, abaxial surface covered in bullabased hairs to 2.3 mm long, and occasional, sessile, glandular hairs near the bases of the bulla-based hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 6, 4.5-4.6 × 0.2-0.4 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes 6, more or less triangular, apex acute, 1-1.4 × 1-1.5 mm, covered in bulla-based hairs abaxially and gland-headed, fi liform hairs adaxially; calyx tube not tearing, 0.3-0.5 mm long with bulla-based hairs abaxially and sessile, glandular hairs, as well as fi liform, gland-headed hairs adaxially and along the apex of the tube; petals 6, most likely white, elliptic to obovate, 5.7-6.6 × 2.7-3.1 mm, with an acuminate apex, only slightly to conspicuously clawed, with one slightly bulla-based hair produced abaxially, subapically, or in some cases, marginally, to 0.1 mm long; stamens 12; fi laments 3.8-4.1 mm long, glabrous, anthers 2.2-2.6 mm long, ovate, with one apically oriented pore, anther thecae 2-2.5 mm long, anthers without a dorso-basal appendage; style 3.8-4.4 mm long, pubescent with slightly bulla-based hairs, oblong to only slightly dilated in the middle, collar absent, style subtended by multicellular, linear to elongate-triangular (needle-like) hairs, which grade into the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1.2-2.8 × 1.5-2.5 mm, apex convex, pubescent with bulla-based hairs, placentation axile, placenta apparently not deeply intruded, 3-locular; berries not seen, mature seeds not seen.
Phenology. Miconia hirtistyla was collected in bud, at anthesis, and in immature fruit in March and July.
Ecology Conservation status. Miconia hirtistyla is mostly known from the very, well protected forests of Parque Nacional Pico Turquino. Although the species has not been collected recently, and we know nothing regarding its reproductive biology or population numbers, it is most likely not threatened by anthropogenic disturbance and habitat loss. However, at present we must categorize the species as data defi cient, owed to the lack of data. Fieldwork will be necessary to appropriately assess the conservation status of this species.
Discussion. Although generally confused with M. jashaferi, M. hirtistyla is most phenetically similar to M. cubacinerea, and the two species may be sister taxa. Th e two species diff er in abaxial leaf surface indumentum and the degree of pitting on the abaxial leaf surface, calyx teeth length (i.e.  Description. Evergreen shrub, 2 m tall; stems round in cross section, not ridged, the internodes 0.4-2.4 cm long, stem indumentum of spreading, bulla-based hairs to 1.7 mm long; nodal line absent. Leaves opposite, decussate, elliptic, 3.9-7.2 × 1.7-3.6 cm, slightly anisophyllous, apex acute to obtuse, base acute to rounded, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, secondary veins mostly suprabasal, the innermost pair produced 0.4-0.8 mm from leaf base, positioned 0.25-0.6 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 0.2-0.48 mm apart at midleaf, impressed and inconspicuous on the upper leaf surface, intertertiary veins present, conspicuous on lower leaf surface, tertiary veins often joined by quaternary veins; adaxial leaf surface covered in poorly developed, bulla-based hairs, widest hair bases to 0.5 mm, apices of bulla-based hairs mostly recurved, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs at the leaf base and apex, sessile, glandular hairs produced throughout the lamina between the bulla-based hairs; abaxial leaf surface sparsely covered in poorly developed bulla-based hairs, these erect, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina easily visible, sessile, glandular hairs produced throughout the lamina between the bulla-based hairs; domatia of multicellular, elongate hairs prominent in the axils of the primary and secondary and primary and tertiary veins; petioles 0.3-1.1 cm long, covered in spreading, bulla-based hairs on both surfaces. Infl orescences terminal, 1-3 fl owered, fl owers produced in glomerulate clusters, 0.8-1.2 × 1-1.4 cm, the peduncle absent to 0.1 cm long, pedicels absent; bracts elliptic to rotund, 3-6.5 mm long; bracteoles elliptic to rotund, 3-4 × 2-2.4 mm, covered abaxially in bulla-based hairs, mostly glabrous abaxially but with bulla-based hairs produced at the base. Flowers 5-merous, sessile; hypanthium ca. 3.4 mm long, oblong, unlobed, barely constricted below the torus, free portion of the hypanthium ca. 1.7 mm long, abaxial surface covered in bulla-based hairs to 3.2 mm long; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 5, 5.7-6.2 × 0.6-0.7 mm, spreading, covered in bulla-based hairs, as well as a few clavate-dendritic hairs; calyx lobes triangular, apex rounded, ca. 0.6 × 1.5 mm, covered in bulla-based hairs abaxially and fi liform hairs at the apex adaxially; calyx tube not tearing, ca. 0.3 mm long with bulla-based hairs abaxially and fi liform hairs at the apex adaxially; petals 5, size at anthesis unknown, only seen in bud, white (according to Wright 2483), with one slightly bulla-based hair produced abaxially, subapically, hairs to 0.4 mm long; stamens 10; fi laments (immature) ca. 2.5 mm long, anthers (immature) ovate, 2-2.1 mm long, with one apically oriented pore, anther thecae 1.7-1.9 mm long, anthers lacking dorso-basal appendage; style (immature) ca. 3.5 mm long, glabrous, only slightly dilated in the middle, collar absent, style subtended by multicellular, linear (needle-like) hairs, which are the same as the hairs of the ovary apex, stigma punctate; ovary ca. 2.3 × 2.2 mm, apex convex, pubescent with linear hairs, placentation axile with shallowly intruded placenta, 3-locular; berries globose, 3.8 mm long (immature and including calyx tube), 7 mm wide; seeds not seen.

Miconia cubacinerea
Phenology. Th e type collection of Miconia cubacinerea was gathered in bud in June.
Distribution (Fig. 4). Miconia cubacinerea is only known from the Elemento Natural Destacado Yunque de Baracoa region of Cuba, based upon the type gathering and one other collection from that area.
Ecology. Little information is available regarding the ecology of this species. However, the vegetation of Yunque de Baracoa is primarily evergreen sclerophylous shrubland over limestone. Miconia cubacinerea likely forms part of the very humid interior karstic formation of Yunque de Baracoa,  Conservation status. Th is species in only known from the type specimen and one other gathering from 1960, thus we consider that is it data defi cient, as it has not been collected from the Yunque de Baracoa area since that time. More fi eldwork will be necessary to fully evaluate the conservation status of M. cubacinerea.
Discussion. Miconia cubacinerea is most phenetically similar to Miconia hirtistyla, but the two species diff er in leaf indumentum, the degree of pitting on the abaixal leaf surface, calyx teeth length, petal form (i.e., non-clawed in M. cubacinerea), and style indumentum (i.e., glabrous in M. cubacinerea); see also discussion under M. hisrtistyla.

Miconia tentaculicapitata
Phenology. Specimens fl owering, with immature and mature fruit have been collected in July and August. Distribution (Fig 4). Miconia tentaculicapitata is restricted to the Sierra Maestra of Cuba. It has been collected primarily in two localities, surrounding the Yara River (along Arroyo el Cristo) in the Granma province and from Loma del Gato in the Santiago de Cuba province.
Ecology. Very little information is available regarding the ecology of M. tentaculicapitata, but collections of Ekman (14748, 14823) were both from along streams ("on rocks along the river"), so this species likely inhabits moist, rocky outcrops in riparian habitats.
Conservation status. Insuffi cient data are available for a critical evaluation of the conservation status of M. tentaculicapitata, thus we suggest a designation of data deficient (DD) according to IUCN (2012) criteria. However, the areas of the Sierra Maestra from where the type collection was made (Ekman 14748) are contained within Turquino National Park, and the species also is found with the Reserva Ecológica (RE) Loma del Gato-Monte Líbano. Th erefore, this species is likely protected from habitat loss and any major anthropogenic disturbance.
Discussion. Miconia tentaculicapitata is most likely closely related to M. jashaferi, M. hirtistyla, M. cubacinerea, as the four species share several morphological characters (e.g., foliaceous bracts and bracteoles, long, fi liform hairs on the adaxial calyx tube surface, and 3 locular ovaries). Th e specimens Linden 2102 (BR, K, P), which are the basis for M. lima in Cuba (Alain 1957), were misidentifi ed and actually are M. tentaculicapitata. However, one of the two specimens of Linden 2102 at BR (13239718) (1) Type. Based on Ossaea norlindii Urb.
Phenology. Miconia norlindii has been collected in fl ower from November through February.
Distribution (Fig 8). Miconia norlindii is restricted to the Sierra Maestra and Sierra de Imías of eastern Cuba.
Ecology. Miconia norlindii occurs in wet montane rainforest and cloud forests occasionally along streams from 800-1800 m. Some associated melastomes include Pachyanthus pedicellatus Urb., Miconia Bécquer (2007) considered M. norlindii to be threatened, however, regarding the number of localities for the species and its presence in protected areas, such as Parque Nacional Pico Turquino, Parque Nacional Pico Bayamesa, Paisaje Natural Protegido La Gran Piedra and la Reserva Ecológica Loma del Gato-Monte Líbano, it may be best to categorize this species as near threatened. Although the species is widespread, it occurs in areas highly vulnerable to climate change (high montane forests) and populations are few.   Type. Based on Clidemia asperifolia Naudin Description. Evergreen shrub, 2-5 m tall; stems round in cross section, not ridged, the internodes 0.5-3.5 cm long, stem indumentum of ascending, appressed bulla-based hairs to 0.7 mm long; nodal line present. Leaves opposite, decussate, ovate to elliptic, 1.8-11.6 × 1.05-4.7 cm, slightly anisophyllous, apex acute to acuminate, base acute, cuneate, to slightly rounded, the margins dentate, dentations covered in one large bulla-based hair, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, secondary veins mostly basal, the innermost pair suprabasal, produced 1.1-18 mm from leaf base, positioned 2.1-9.8 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.8-5.5 mm apart at midleaf, intertertiary veins occasionally present, tertiary veins often joined by conspicuous, quaternary veins; adaxial leaf surface covered in dorsally compressed, bulla-based hairs, widest hair bases to 1.2 mm, apices of bulla-based hairs mostly recurved towards the leaf margin, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bulla-based hairs, these ascending, appressed, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina clearly visible, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile, glandular hairs produced throughout the lamina and along veins, domatia of tufts of multicellular hairs produced in the axils of the primary and secondary, primary and tertiary and secondary and tertiary veins; petioles 0.5-3.3 cm long, covered in ascending, appressed, bulla-based hairs on both surfaces. Infl orescences terminal, 19-57 fl owered, fl owers mostly produced in glomerulate clusters, 2.4-8.2 × 1.6-6.4 cm, the peduncle 0.3-3.2 cm long, proximal infl orescence branches 4-23 mm long; bracts oblong to ovate, 0.6-1.8 mm long; bracteoles narrowly ovate with an attenuate apex, 0.6-0.7 × 0.2-0.3 mm, appearing as enlarged bulla-based hairs, occasionally with smaller bulla-based hairs towards the base of the bracteoles. Flowers 5-merous, sessile or with pedicels to 0.3-0.8 mm long; hypanthium 1.3-2 mm long, short-oblong to globose, 5-lobed, strongly constricted below the torus, free portion of the hypanthium 0.5 mm long, abaxial surface covered in bulla-based hairs to 0.4 mm long, and sessile, glandular hairs between the bullabased hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx  teeth 0.3-1.3 × 0.2-0.5 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes triangular, apex acute, 0.5-1 × 0.7-1.8 mm, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs adaxially; calyx tube not tearing, 0.4-0.8 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 5, white, elliptic, 5.5-5.7 × 1.3-3.3 mm, with an acute apex and membranous margin, with one to four slightly bulla-based hairs produced abaxially, just below the apex, to 0.7 mm long; stamens 10; fi laments 2-2.2 mm long, glabrous, anthers 1.3-2.4 mm long, with one dorsally oriented pore, anther thecae 1.1-2 mm long, anthers with a dorso-basal appendage 0.1-0.2 mm long; style 5.2-5.4 mm long, glabrous, slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, linear to elongate-triangular (needle-like) hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1.8-4.5 × 2.9-5.6 mm, apex fl at, pubescent with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded placenta, 5 locular; berries globose, 5-lobed, purple at maturity, 3-5.5 mm long (including calyx tube), 4.1-5.8 mm wide, seeds 0.5-0.7 mm long, obpyramidal, often falcate, testa smooth, light brown, raphe light brown, smooth, extending the length of the seed.
Phenology. Miconia asperifolia has been collected in fl ower and fruit from February through August.
Distribution (Fig 11). Miconia asperifolia is found in the Blue and John Crow Mountains of Jamaica.
Conservation status. Miconia asperifolia is a widespread species, which also occurs in the protected Blue Mountains National Park, so the species should be considered stable.
Discussion. Miconia asperifolia is resolved as sister to M. granulata in a subclade consisting of Cuban species. However based on morphological similarity, M. asperifolia appears likely to be sister to M. hybophylla, a rare Haitian species not included in our phylogenetic analysis (see below under discussion about M. hybophylla). Both species have similar adaxial leaf surfaces with well-formed by slightly compressed bulla-based hairs composed of larger hairs generally surrounded by smaller hairs. Both species also have numerous well-formed domatia in the axils of the primary and secondary, primary and tertiary, and secondary and tertiary veins, as well as large, expanded, cymose infl orescences (Figs 10, 15G-J). Populations of M. asperifolia diff er slightly between the John Crow and Blue Mountains with those in the John Crow Mountains tending to have larger leaves and less well developed bulla-based hairs on the upper leaf surface, however, these characters form a gradient from one geographic area to the next and represent only minor intraspecifi c variation. Shade forms of M. asperifolia often have poorly formed hairs on the upper leaf surface (Fig. 15I), which resemble a putatively closely related Cuban species, M. cubana (Fig. 12).
William Purdie was assigned by J.D. Hooker to collect plants in Jamaica during the years 1843-1844 (Sinha 1972). Naudin (1852) states that his new species, Clidemia asperifolia, was brought to his attention by Hooker based on material sent from Jamaica ("In insula Jamaica, loco haud indicato. Planta a celeberrimo Hooker communicate."). Th us, we conclude that the specimens collected by Purdie are the type material and have been used as the lectotype (see above).    (5) Type. Based on Ossaea hybophylla Urb. Description. Small evergreen tree (height unknown); young stems purplish, round to slightly quadrangular with rounded angles in cross section, not ridged, the internodes 0.6-4.3 cm long, stem indumentum of granulate, bulla-based hairs to 0.3 mm long, these spreading; nodal line present but inconspicuous. Leaves opposite, decussate, broadly elliptic, 2.2-4.5 × 1.6-2.6 cm, slightly anisophyllous, margins with conspicuous spine-tipped hairs, these spreading (especially at base of leaf) to appressed or recurved along the leaf margin, apex acute, base acute often asymmetrical, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the out-ermost pair of secondary veins ocasionally intramarginal, mostly basal, the innermost pair, suprabasal, asymmetrical or symmetrical, produced 2.5-6.5 mm from leaf base, positioned 2.2-4.3 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.8-3.3 mm apart at midleaf, intertertiary veins rarely present and inconspicuous when present, tertiary veins often joined by quaternary veins; adaxial leaf surface covered in bulla-based hairs, widest hair bases to 0.7 mm, apices of bulla-based hairs mostly erect or recurved towards the leaf margin, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs, especially toward the base of the leaf; abaxial leaf surface with sparse bulla-based hairs, these mostly erect to spreading, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina clearly visible, olive green or occasionally purplish, not deeply pitted, sessile, glandular hairs produced throughout the lamina, conspicuous domatia produced as tufts of hairs at the junction of the primary and secondary veins, as well as at the junctions of the primary and secondary veins with the tertiary veins; petioles 0.5-1.1 cm long, purplish, adaxial surface of petiole with spreading, multicellular hairs to 1.2 mm long, the rest of the petiole covered in short 0.1-0.4 mm spreading, bulla-based hairs. Infl orescences terminal, pyramidal, purplish, with up to 41 fl owers, fl owers produced in cymose clusters, 1.7-4 × 2-3.1 cm, the peduncle 0.1-1.4 cm long, proximal infl orescence branches 6-13 mm long, bracts oblong to narrowly ovate, 0.6-1.5 mm long; bracteoles narrowly ovate, 0.4-0.8 × 0.2-0.25 mm, appearing as large bulla-based hairs. Flowers 4-merous, pedicels 0.5-0.8 mm long; hypanthium 1.3-2.7 mm long, short-oblong to globose, 4-lobed, slightly constricted below the torus; free portion of the hypanthium ca. 0.5 mm long, abaxial surface covered in dorsi-ventrally compressed or erect bulla-based hairs to 0.1 mm long, and sessile, glandular hairs between the bulla-based hairs, adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 0.5-1 × 0.3-0.6 mm, spreading with straight or recurved apices, having the general appearance of a large bulla-based hair and often with a large bulla-based hair at the base; calyx lobes triangular, apex acute, 0.9-1.2 × 1.4-1.7 mm, covered in bulla-based and sessile, glandular hairs abaxially and sessile, glandular hairs adaxially; calyx tube not tearing, 0.3-0.5 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially, clavate-dendritic hairs produced at the apex of the calyx tube; petals 4, most likely white, 2-3 × 1.7-1.8 mm, ovate, apex acute, with one slightly bulla-based hair produced just below the apex on the abaxial surface, to 0.3 mm long; stamens 8; fi laments 1.2-1.3 mm long, glabrous, anthers 1.1-1.3 mm long, with one dorsally oriented pore, anther thecae 0.9-1.1 mm long, anthers with a dorso-basal appendage 0.2 mm long; style 3.5-4.3 mm long, glabrous, not or only slightly dilated in the middle (mostly oblong), collar absent, style subtended by an inconspicuous crown of triangular hairs (longer than those on the rest of the ovary apex), which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1.3-4 × 1.9-2.4 mm, 4-lobed, apex truncate, pubescent with triangular, bulla-based hairs, placentation axile with deeply intruded placenta, 4-locular; immature berries globose, 4-lobed, color at maturity unknown, but probably more or less purple, 2.5-2.7 mm long (including calyx tube), 3.1-3.3 mm wide, seeds 0.4-0.5 mm long, obpyramidal, testa smooth, raphe smooth, extending the length of the seed.
Phenology. Flowers at anthesis, as well as buds and immature fruit were present on the type collection, which was gathered in March.
Ecology. Nothing is known regarding the ecology of this species. Conservation status. Insuffi cient data are available for determining the conservation status of this species, although M. hybophylla is very likely endangered as a result of forest clearing for subsistence agriculture and charcoal production in west-central Haiti.
Discussion. Miconia hybophylla is likely sister to M. asperifolia and can be easily recognized as a smaller, more compact version of M. asperifolia, as compared with other species in the Lima clade. Th e two species also share very, well developed domatia in the axils of the primary and secondary veins, as well as the axils of the tertiary with primary and tertiary with secondary veins.
Phenology. Th is species has been collected in bud in August, in fl ower and mature fruit in November, immature fruit in December, as well as mature fruit in April.
Distribution (Fig 8). Miconia granulata is restricted to the northern portion of eastern Cuba in the Guantánamo and Holguín provinces in the areas with moderate elevation, from Baracoa to Moa.
Conservation status. Bécquer (2007) considered Miconia granulata to be threatened, as it is known from relatively few locations and only occurs in one protected area, Parque Nacional Alejandro de Humboldt. We propose a preliminary conservation status of Critically Endangered owed to ongoing and future mining practices in the zone where it occurs, as well as the fact that M. granulata is not an abundant species in any part of its range.
Discussion. Based on phylogenetic analyses, M. granulata is resolved in a clade wtih M. argentimuricata and M. asperifolia and is sister to M. asperifolia (Fig. 2), a Jamaican species. It is the only member of this clade with 4-merous fl owers and occasionally pendulous infl orescences (Figs 7B, 9K-P), however, those characters are also shared with M. bullotricha, which may be closely related to M. granulata (see below under M. bullotricha). Miconia granulata also has very poorly developed bulla-based hairs on the upper leaf surface, as compared to its closest relatives (Fig 9 K, P).
Phenology. Miconia cubana was collected in bud, fl ower and immature fruit on March 16, if our interpretation of the label data is correct. However, no petals were seen on the type specimens (other than in bud), where only styles and a couple of stamens were present. Distribution (Fig. 13). Miconia cubana may be restricted to Pinar del Río, Cuba, if our interpretation of Wright's collections is correct. Th ree separate labels were distributed by A. Gray of the type material collected by Wright, which at the time was considered to be another species, Miconia asperifolia (see Alain 1955). Th e sheet of the isotype of M. cubana at GH (GH00713104) actually is composed of branches from two species, M. cubana and M. norlindii. Miconia cubana is the branch in the upper left hand corner on that sheet. It is presumed that the two other labels associated with that specimen actually pertain to M. norlindii, as they both refer to eastern Cuba date from the material of M. cubana. Th erefore, we interpret the Isabel María locality to actually pertain to M. cubana, contrary to Howard and Wright (1988)  Type. Based on Calycogonium muricatum Griseb. Description. Evergreen shrub, to 1.5 m tall; stems round in cross section, not ridged, the internodes 0.5-10.9 cm long, stem indumentum of ascending, bulla-based hairs to 2.2 mm long; nodal line absent. Leaves opposite, decussate, broadly to narrowly elliptic, 2.7-9.3 × 1.4-4.5 cm, slightly anisophyllous, apex acute to acuminate, base rounded to acute, margins dentate, dentations covered in well-developed bullabased hair, venation acrodromous, 5-7-veined, the midvein and 3 pairs of arching secondary veins, the outermost intramarginal, secondary veins mostly basal to suprabasal, the innermost pair suprabasal, produced 2.5-17 mm from leaf base, positioned 2.3-10 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.9-5.6 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface covered in well-developed bulla-based hairs completely covering the leaf areoles, widest hair bases to 2.1 mm, apices of bulla-based hairs mostly recurved toward the leaf margin, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in sparse bulla-based hairs, these spreading to erect, often crisped, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina clearly visible, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile, glandular hairs produced throughout the lamina and along veins; petioles 0.2-2.1 cm long, covered in ascending to spreading, bulla-based hairs on both surfaces. Infl orescences terminal, fl owers mostly produced in glomerulate clusters, 3-42 fl owered, 3.3-7.6 × 1.5-9.8 cm, the peduncle 0.4-2.2 cm long, proximal infl orescence branches 8-26 mm long; bracts narrowly ovate with a long attenuate apex, 1.2-2.2 mm long; bracteoles narrowly ovate with a long attenuate apex, 1.4-2 × 0.3-0.4 mm, glabrous. Flowers 5-merous, sessile or with pedicels to 0.5-0.9 mm long; hypanthium 3.2-3.4 mm long, short-oblong to globose, unlobed to slightly 5-lobed, but lobing mostly obscured by bulla-based hairs, slightly constricted below the torus, free portion of the hypanthium 0.5-0.6 mm long, abaxial surface covered in bulla-based hairs to 2.7 mm long, and sessile, glandular hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs developing into androecial fringe; calyx teeth 2.2-2.7 × 0.4-0.6 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes triangular, acute to rounded at apex, 0.4-2.2 × 0.8-2.5 mm, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs adaxially; calyx tube not tearing, 0.2-0.8 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 5, 3.6-4.5 × 1.9-2.7 mm, white, ovate or elliptic with an acute apex and membranous margin, with 3-4 slightly bulla-based hairs produced abaxially, just below the apex, to 2.7 mm long; stamens 10; fi laments 2.2-2.9 mm long, glabrous, anthers 1.2-2 mm long, with one dorsally oriented pore, anther thecae 1.1-1.8 mm long, anthers with a dorso-basal appendage 0.2-0.25 mm long; style 3.3-3.8 mm long, glabrous, slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, linear to elongate-triangular (needle-like) hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 2.4-3 × 3.8-4.5 mm, apex fl at to slightly rounded, pubescent with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded placenta, 5-locular; berries globose, purple to violet at maturity, 5-6 mm long (including calyx tube), 5.3-7 mm wide, seeds 0.6-0.65 mm long, obpyramidal, slightly falcate, testa smooth, light brown, raphe dark amber, smooth, extending the length of the seed. Phenology. Miconia argentimuricata has been collected in fruit from December through July and in fl ower from November through April. Distribution (Fig 11). Miconia argentimuricata is restricted to eastern Cuba and can be found in both the Sierra Maestra (including Loma del Gato) and the Baracoa-Moa-Sierra de Cristal regions.
Ecology. Miconia argentimuricata occurs from 600-1400 m in elevation and can be found on granitic soils (Sierra Maestra) or on serpentine soils in the northern part of its distribution (Moa region). Th e species occurs in lower elevation "manacales" (wet, montane rainforest; see Borhidi 1996) in the northern part of its range and higher elevation "monte frio" in the southern part of its range.
Conservation status. Considering the wide range of M. argentimuricata and that it occurs in protected areas in both parts of its range (Parque Nacional Alejandro de Humboldt, Parque Nacional Pico Turquino, Reserva Ecológica Loma del Gaato-Monte Líbano), this species should be considered stable and possibly of least concern. However, some populations in the Moa region may be under threat from mining operations, and Bécquer (2007) suggested the species was near threatened.
Discussion. Populations of Miconia argentimuricata from the Sierra Maestra are readily distinguished from those of the Baracoa and Moa regions by their more robust growth form (broadly elliptic leaves, larger infl orescences), as well as their darker, copper rather than silver color of the leaf adaxial surface (in herbarium specimens). Th e leaves are narrowly elliptic to often ovate with narrowly acute to acuminate apices in those specimens from the Baracoa-Moa-Sierra de Cristal region. However, morphological characters are suffi ciently cohesive between the two regions to place those geographically separated populations within a single species.
Although Howard and Wright (1988) provided a microfi che of the lectotype of Miconia argentimuricata (as Calycogonium muricatum), he did not lectotypify the species explicitly in writing, and thus his lectotypifi cation was incomplete. We have fully lectotypifi ed the species with the specimen from GOET here. As specimens collected by Wright were often assigned species numbers by Asa Gray, as well as Wright's collection numbers, one of the isolectotypes here subsequently has a diff erent number on the label (Wright 682) than that given in the protologue (Wright 2485) for the species (see also discussion by Bécquer 2012 Fig. 16 Type. CUBA. Prov. Guantánamo: Palenque. Bernardo. Sierra del Frijol, loma Bernardo, 800-900 m, 21 May 1983, J. Bisse, C. Beurton, H. Dietrich, J. Gutiér-rez, L. Lepper, R. Dolmus, E. Köhler, R. Rankin, I. Arias, HFC-49930  Description. Evergreen shrub (height unknown); stems round in cross section, not ridged, the internodes 1.1-2.4 cm long; stems densely covered in bulla-based hairs with strongly to narrowly dilated bases, to 0.3 mm long, the hairs spreading to descending with apices recurved upwards, young stem hairs often dark purple in color; nodal line inconspicuous, present. Leaves opposite, decussate, elliptic to ovate-elliptic, often slightly falcate, 4.2-6 × 1-2.2 cm, often slightly anisophyllous, yellowish when dried; apex narrowly acute; base rounded to broadly cuneate or abruptly cuneate; margin revolute, dentate, the dentations obscure, each covered in one large, bulla-based hair, venation acrodromous, 3 (-5)-veined, 1 primary vein and 1 (rarely 2) pairs of suprabasal secondary veins, often asymmetrical at union with midvein, produced 2-6 mm from the leaf base, positioned 0.7-3 mm in from margin at widest part of blade, the tertiary veins percurrent, more or less perpendicular to midvein, 2-3 mm apart at mid-leaf, intertertiary veins present, often joined by quaternary veins; adaxial leaf surface with primary and secondary veins impressed, tertiary veins fl at to slightly impressed, remaining veins fl at, abaxial surface with primary, secondary and tertiary veins raised, the higher order veins more or less fl at to slightly raised (i.e., clearly visible to more or less obscure); adaxial leaf surface completely covered in erect bulla-based hairs, these fully expanded at the base, thus the lamina obscured, widest hair bases to 1.5 mm wide, hair apices acute to truncate, sometimes slightly recurved toward the leaf margin, sessile, glandular hairs occurring between the bases of bulla-based hairs; abaxial leaf surface nearly completely covered with bulla-based hairs with strongly to narrowly dilated bases, the lamina areoles not completely fi lled, the hairs along the epidermis erect with apices recurved or not, veins completely covered by spreading to erect hairs mostly with narrowly dilated bases and recurved apices, sessile, glandular hairs occurring throughout the lamina, as well as along veins; acarodomatia inconspicuous, of multicellular, linear hairs present in the axils of the primary and secondary, as well as primary and tertiary veins; petiole 5-8 mm long, covered in spreading bulla-based hairs, those of the adaxial surface slightly longer and narrower than those of the abaxial surface and recurved towards to the leaf blade. Infl orescences terminal, well-developed to reduced cymes of 3-13 fl owers, 2-3.5 × 1.8-3.4 cm, the peduncle 0.7-1.4 cm long, usually conspicuously refl exed at base, thus the entire infl orescence pendant, the proximal infl orescence branches 0.5-1 cm long; bracts oblong to narrowly ovate, 1.1-2 mm long; bracteoles narrowly ovate, ca. 0.5-0.7 × 0.2-0.3 mm. Flowers 4-merous, pedicels 0-1 mm long. Hypanthium ca. 1.6 × 2.8 mm, globose, slightly constricted below torus, abaxial surface covered in granulate, bulla-based hairs with dilated bases and attenuate to truncate apices, to 0.5 mm long, and sessile, glandular hairs, the free portion of hypanthium 0.5-0.7 mm long, adaxial surface longitudinally ridged and covered by bulla-based hairs; calyx teeth 1.75-2.2 × 0.5 mm, linear and terete, recurved upon maturation, covered in bulla-based hairs; calyx lobes ca. 1 × 1.3 mm, triangular, apices acute, with bulla-based hairs abaxially and sessile, glandular hairs produced adaxially; calyx tube not tearing, ca. 0.4 mm long, with bulla-based hairs abaxially, sessile, glandular hairs adaxially and clavate- dendritic hairs produced at the apex; petals 4, immature (i.e., only seen in bud), ovate to elliptic, with acute apices, apices with one, slightly bulla-based hair produced subapically, hair to 0.5 mm long; stamens 8 (immature), fi laments glabrous, anthers ovate, with a welldeveloped dorso-basal appendage and one apically-oriented pore (the pore position could be an artifact of level of maturity); style (immature) dilated in the middle, subtended by a short crown of multicellular hairs, these only slightly longer than the surrounding bullabased hairs on the ovary apex; stigma punctate; ovary ca. 1.4 × 2.4 mm, apex fl at, covered in bulla-based hairs, 4 locular, with axillary placentation, the placenta deeply intruded into locule; berries (immature) globose, ca. 3-3.4 × 3 mm, color at maturity unknown, but probably more or less purple; seeds (immature) 0.2-0.6 mm long, obpyramidal, testa smooth, light brown, raphe extending the length of the seed, dark brown.
Phenology. Plants with buds and young fruits have been collected in May. Distribution (Fig. 11). Miconia bullotricha is restricted to the mountains of the Guantánamo province, Cuba.
Conservation status. We do not have extensive knowledge of numbers of individuals per population or the reproductive biology of this species, so the conservation status of M. bullotricha cannot be critically evaluated at this time and should be considered data defi cient. More fi eldwork is imperative before its status can be assessed.
Discussion. Although  compared M. bullotricha to the very phenetically similar M. ottoschmidtii (Figs 12E-H, 17), and several characters overlap with that species, it is possible that M. bullotricha may be more closely related to members of the Acuminata subclade , specifi cally M. granulata. Miconia bullotricha shares the infl orescence structure (Fig. 9O, 16)   Type. Based on Ossaea ottoschmidtii Urb.
Description. Evergreen shrub, 1-1.5 m tall; stems round in cross section, not ridged, the internodes 0.5-4.9 cm long, stem indumentum of spreading, granulate, bulla-based hairs to 0.4 mm long; nodal line present. Leaves opposite, decussate, rarely ovate to mostly narrowly elliptic, 2.4-7.9 × 1-3.1 cm, slightly anisophyllous, apex acute to slightly acuminate, base acute, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the outermost sometimes intramarginal, secondary veins mostly basal to suprabasal, the innermost pair suprabasal, produced 0.8-25 mm from leaf base, positioned 1.6-4.7 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.4-4.3 mm apart at midleaf, intertertiary veins rarely present, tertiary veins usually joined by conspicuous quaternary veins; adaxial leaf surface covered in well-developed, dorsally compressed bulla-based hairs, widest hair bases to 1.8 mm, these mostly to entirely covering the leaf areoles, apices of bulla-based hairs mostly erect, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bulla-based hairs, these erect along the lamina, spreading along the primary and secondary veins, and recurved towards the leaf margin along the tertiary veins, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina visible, appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile, glandular hairs produced throughout the lamina and along the veins, domatia present, consisting of tufts of bulla-based hairs, at the axils of the primary and secondary, as well as primary and tertiary veins; petioles 0.3-2.1 cm long, covered in granulate, bulla-based hairs on the abaxial surface and long, spreading, bulla-based hairs on the adaxial surface. Infl orescences terminal, cymose, fl owers mostly produced in glomerulate clusters, 19-25 fl owered, 2.2-6.5 × 1.3-4 cm, the peduncle 0.1-1.9 cm long, proximal infl orescence branches 3-20 mm long; bracts oblong to ovate, 0.6-1.7 mm long; bracteoles ovate, 0.3-0.7 × 0.2-0.6 mm. Flowers 4-merous, with pedicels 0.2-2 mm long; hypanthium 1.4-2.3 mm long, short-oblong to globose, strongly 4-lobed, clearly constricted below the torus, free portion of the hypanthium 0.5-0.9 mm long, abaxial surface covered in bulla-based hairs to 0.3 mm long, and sessile, glandular hairs near the bases of the bulla-based hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 0.4-0.8 × 0.2-0.4 mm, erect to spreading, covered in bulla-based hairs; calyx lobes 0.2-1.1 × 1.2-1.5 mm, triangular, broadly acute to rounded at apex, covered in bulla-based hairs abaxially and sessile, glandular hairs adaxially; calyx tube not tearing, 0.4-0.8 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 4, 2.9-3 × 1.4-1.7 mm, white, ovate to elliptic, with an acute apex, with 1-2 slightly bulla-based hairs produced abaxially, just below the apex, to 0.5 mm long; stamens 8; fi laments 1.3-1.6 mm long, glabrous, anthers 1.4-1.5 mm long, with one dorsally oriented pore, anther thecae 1.1-1.2 mm long, anthers with a dorso-basal appendage 0.3 mm long; style 3.5-3.7 mm long, glabrous, not or only slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, linear to elongate-triangular (needle-like) hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1.4-2 × 2.2-2.5 mm, strongly 4-lobed, apex fl at, pubescent with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded placenta, 4-locular; berries globose, 4-lobed, purple at maturity, 2.7-3.6 mm long (including calyx tube), 3-4.8 mm wide, seeds 0.4-0.6 mm long, obpyramidal, testa smooth, light brown, raphe dark brown to black, smooth, extending the length of the seed.
Phenology. Miconia ottoschmidtii has been collected in fl ower from January through July and in fruit from April through August.
Distribution (Fig. 13). Miconia ottoschmidtii occurs in northern Cuba in Pinar del Río (Pan de Guajaibón), Mountain of Guamuhaya of central Cuba, the Sierra Maestra of southern/eastern Cuba and Sierra Cristal.
Conservation status. Th is species is widespread throughout Cuba occurring in Parque Nacional Pico Turquino, Parque Nacional Pico Cristal, Reserva Ecológica Lomas de Banao, Reserva Ecológica Pico San Juan, Parque Nacional Topes de Collantes and Elemento Natural Destacado Pan de Guajaibón, among other non-protected areas. However, the species is not abundant throughout most of its distribution, and is only known from one collection each from Pico Cristal and Pan de Guajaibón. Th e habitat for this species has been reduced in size and quality, thus, we propose a category of endangered for M. ottoschmidtii.
Discussion. Miconia ottoschmidtii is the only Cuban member of an otherwise Hispaniolan clade containing M. lima, M. limoides, M. paralimoides and M. pedunculata (Fig. 2), as well as several other putative members of the clade (M. phrynosomaderma, M. marigotiana, also Hispaniolan endemics). Of these taxa, M. ottoschmidtii is most phenetically similar to M. lima but can be diff erentiated from that species by the size of nearly all parts of the plant, with M. lima being the larger of the two species. Also, M. lima has broadly elliptic leaves, whereas those of M. ottoschmidtii are narrowly elliptic (length/width ratio 1.3-4.6 cm in M. ottoschmidtii and 1.5-2.2 cm in M. lima). Several specimens of M. ottoschmidtii exhibit longer hairs along the stems and hypanthia than the typical granulate hairs generally exhibited by the species (i.e., Ekman 15794, 15912, Bécquer HAC-82434), and these plants are distributed more or less throughout the distribution of M. ottoschmidtii. It should be noted that the specimen, Bécquer HAC-82434, is the northernmost and westernmost collection of the species in Cuba from Pinar del Río, Pan de Guajaibón, and is disjunct from other collections to the southeast in Guamuhaya montains. Th is population exhibits less well developed bullabased hairs on the adaxial leaf surface and has larger and more broadly elliptic leaves (3.05-7.9 × 1.6-3.1 mm vs. 2.4-6.8 × 1-2.2 mm) than most populations from other   Description. Evergreen shrub, 1.5-5 m tall; stems round in cross section, not ridged, the internodes 0.3-6.3 cm long, stem indumentum of granulate to long, attenutate bulla-based hairs to 1.5 mm long, these ascending, appressed; nodal line present. Leaves opposite, decussate, elliptic to slightly obovate, 0.7-5.7 × 0.42-3.2 cm, slightly anisophyllous, apex acute, acuminate or obtuse, base acute to rounded, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the outermost sometimes intramarginal, secondary veins mostly basal to suprabasal, the innermost pair suprabasal, produced 0.5-10 mm from leaf base, positioned 1.2-6 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.4-3.9 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface densely covered in well-developed bulla-based hairs, these covering the leaf areoles, widest hair bases to 2.7 mm, apices of bulla-based hairs mostly erect to recurved towards to the leaf margin, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bulla-based hairs, these erect to spreading, oftentimes ascending along the primary and secondary veins, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina visible to nearly obscured, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface, sessile, glandular hairs produced throughout the lamina and along all primary, secondary and tertiary veins; petioles 0.3-2.1 cm long, covered in ascending, bulla-based hairs on both surfaces, those on the adaxial surface slightly longer. Infl orescences terminal, 5-43 fl owered cymes, 1.9-6.5 × 1-6 cm, the peduncle absent to 2.5 cm long, proximal infl orescence branches 3-23 mm long; bracts oblong to ovate with an attenuate apex, 1.3-2.7 mm long; bracteoles 0.8-2.2 × 0.2-0.6 mm, narrowly ovate with an attenuate apex, glabrous or with a few bulla-based hairs. Flowers 4-5-merous, with pedicels to 0.9 mm long; hypanthium 2-3.3 mm long, short-oblong to globose, 4-5-lobed, but lobing mostly obscured by bulla-based hairs, slightly constricted below the torus, free portion of the hypanthium 0.5-1 mm long, abaxial surface covered in bulla-based hairs from 0.8-1.6 mm long, and occasional, sessile, glandular hairs near the bases of the bullabased hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 1.2-3.2 × 0.3-0.6 mm, ascending, spreading or defl exed, covered in bulla-based hairs; calyx lobes 1-1.5 × 1.5-2 mm, more or less triangular, apex acute to obtuse, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs, and occasionally, clavate-dendritic hairs adaxially; calyx tube not tearing, 0.3-0.7 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially, clavate-dendritic hairs often produced at the tube apex; petals 4-5, red to pink or pinkish-red, elliptic, 3-4.3 × 1.9-2.6 mm, with an acute apex and slightly membranous margin, with one or two slightly bulla-based hairs produced abaxially, just below the apex, often two bulla-based hairs nearly identical and appearing as one hair with a cordate base, to 1.5 mm long; stamens 8-10; fi laments 1.2-2.1 mm long, glabrous, anthers 1.4-1.9 mm long, with one dorsally oriented pore, anther thecae 1.1-1.6 mm long, anthers with a dorso-basal appendage 0.25-0.4 mm long; style 4.7-4.9 mm long, glabrous, slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, elongate-triangular (needle-like) hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1. 5-2.5 × 2.3-3.4 mm, apex fl at, pubescent with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded placenta, 4-5-locular; berries globose, 4-5-lobed, purple to purple-black at maturity, 2-3.5 mm long (including calyx tube), 2-3.5 × 3-4.1 mm wide, seeds 0.6-0.8 mm long, obpyramidal, testa smooth, light brown, raphe light brown, smooth, extending the length of the seed.
Phenology. Miconia lima has been collected in fl ower and fruit from Februrary throught November.
Distribution (Fig 21). Miconia lima is the most widespread species of all of the members of the Lima clade occurring widely throughout Hispaniola on almost all mountain ranges from 550 to 2000 m in elevation. It is known from the Massif de la Selle, Massif du Cahos, and Massif de Matheux in Haiti and the Sierra de Baoruco, Sierra de Neiba, Cordillera Central and Cordillera Septentrional in the Domincan Republic (Fig. 22). Miconia lima was cited by Alain (1957) Ionta et al., Miconia subcompressa Urb. and Miconia zanonii Judd et al. See Ciferri (1936) for more information about the species occurrence and associate species in the Cordillera Septentrional (Pico Diego de Ocampo).
Conservation status. Miconia lima is the most widespread species of the Lima clade on Hispaniola and does not appear to be in any immediate danger from anthro- pogenic disturbance in most parts of its range. However, localized populations are likely under intense pressure from forest clearing for farming and charcoal production in Haiti (e.g., Massif de la Selle) and the Dominican Republic, so we consider the species as vulnerable.
Discussion. Miconia lima forms part of the M. lima species complex comprised of M. lima, M. limoides, M. marigotiana, M. ottoschmidtii, M. paralimoides, M. pedunculata and M. phyrnosomaderma. Miconia lima is perhaps the most taxonomically complex species of the entire Lima clade. Th e species varies greatly throughout its range but shows geographic cohesion in morphological features, i.e., plants from specifi c geographic regions can be distinguished from those of other regions. For instance, populations from Massif de la Selle, Haiti tend to have smaller leaves than other populations, and populations in the Cordillera Septentrional, Dominican Republic have longer calyx teeth than most other populations of the species.
Populations in the Monteada Nueva region of the Dominican Republic show potential introgession with M. limoides, as they have spreading to spreading-ascending stem hairs (instead of strongly ascending stem hairs), strongly angular, bulla-based hairs on the adaxial leaf surface, and dense, bulla-based hairs on the abaxial leaf surface, mostly or nearly obscuring the lamina, as in M. limoides (e.g., Howard 12300, Judd 5183, 8083, Liogier 11661, Majure 5960, Zanoni 18919, 30470, 38638; see Populations of M. lima from the Loma Quita Espuela region of the Cordillera Septentrional, Dominican Republic are unique within the species in having densely long-shaggy pubescent stems (with hairs to 2.8 mm long), infl orescence axes, and hypanthia, as well as adaxial leaf surfaces with well-developed bulla-based hairs with long, attenuate apices and calyx teeth up to 5 mm long (i.e., Abbott 2184, García 5222). Partial DNA sequence data from an herbarium specimen (García 5222) (5) Type. Based on Ossaea limoides Urb. Description. Evergreen shrub, 1.5-5 m tall; stems round in cross section, not ridged, the internodes 0.3-6.3 cm long, stem indumentum of descending to spreading, bulla-based hairs to 1.6 mm long, these with apices curved upwards; nodal line present. Leaves opposite,decussate,elliptic, cm, slightly anisophyllous, apex acute, acuminate, to slightly rounded, base acute, cuneate, to rounded, margins dentate to crenulate, dentations (crenulations) covered in a large bulla-based hair, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the outermost sometimes intramarginal, secondary veins mostly basal to suprabasal, the innermost pair suprabasal, produced 1.2-7 mm from leaf base, positioned 0.9-8 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.3-3.2 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface covered in well developed bulla-based hairs, widest hair bases to 1.8 mm, apices of bulla-based hairs mostly erect to recurved towards the leaf margin, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bulla-based hairs, these erect, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina and spreading to descending with the apices recurved upwards, the lamina completely obscured by bulla-based hairs, lamina appearing as a series of pits from depressions of the bullabased hairs produced from the upper leaf surface and slightly raised intertertiary veins, sessile, glandular hairs produced from between the bulla-based hairs; petioles 0.4-2.2 cm long, covered in descending to spreading, bulla-based hairs on both surfaces, the apices recurved upwards. Infl orescences terminal 3-33 fl owered cymes, fl owers mostly produced in glomerulate clusters, 0.9-10 × 1.4-5 cm, the peduncle 0.7-3.7 cm long, proximal infl orescence branches 3-15 mm long; bracts oblong to ovate often with an attenuate apex, 1.5-4 mm long; bracteoles narrowly ovate 1.4-3.5 × 0.4-0.8 mm, with an attenuate apex. Flowers 4-merous, sessile or with pedicels to 0.5 mm long; hypanthium 2.7-3.8 mm long, globose, 4-lobed, but lobing mostly obscured by bulla-based hairs, slightly constricted below the torus; free portion of the hypanthium 0.7-1.1 mm long, abaxial surface covered in bulla-based hairs from 1-3 mm long, and occasional, sessile, glandular hairs near the bases of the bulla-based hairs, adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 2.1-3.3 × 0.4-0.8 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes more or less triangular 1.3-2.5 × 1.4-2.6 mm, apex broadly acute, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs adaxially; calyx tube not tearing, 0.7-1.2 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 4, white to rose colored, elliptic, 4-4.1 × 2.4-2.6 mm, with an acute apex and membranous margin, with one or two slightly bulla-based hairs produced abaxially, just below the apex, to 3.5 mm long; stamens 8; fi laments 1.5-2 mm long, glabrous, anthers 1.2-1.4 mm long, with one dorsally oriented pore, anther thecae 1.1-1.3 mm long, anthers without a dorso-basal appendage or with a short appendage to 0.1 mm long; style 4-4.8 mm long, glabrous, not or only slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, bulla-based hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1. 5-2.8 × 2.5-4.3 mm, apex fl at to convex, pubescent of bulla-based hairs, placentation axile with deeply intruded placenta, 4-locular; berries globose, 4-lobed, purple at maturity, 3.9-7 mm long (including calyx tube), 4.5-9 mm wide, seeds 0.6-0.8 mm long, obpyramidal, testa smooth, light brown, raphe black, smooth, extending the length of the seed.
Phenology. Miconia limoides has been collected in fl ower and fruit from January through August. Distribution (Fig 23). Miconia limoides only occurs in the southern peninsula of Hispaniola from western Haiti in the Massif de la Hotte (from one collection; Clase 4132; see Judd et al. 2015b) to southwestern Dominican Republic in the Sierra de Baoruco (Fig. 19). Only two collections (Ekman 5462,9519) exist from the Massif de la Selle, Haiti from the Port-au-Prince area at Morne Malanga. It is likely that this species occurs more widely in the eastern part of the Massif de la Selle and other parts of Massif de la Hotte.
Conservation status. Miconia limoides is likely endangered in the Massif de la Selle, Haiti (where it is only known from two Erik Ekman specimens), from habitat loss and the species is not protected in the Sierra de Baoruco of the Dominican Republic, where it is also threatened from anthropogenic disturbance. Miconia limoides is only known from one collection in Massif de la Hotte from Parc National Pic Macaya (Clase 4132), and the area from where it was collected was heavily disturbed from local farming practices, so we categorize the species as endangered.
Discussion. Miconia limoides forms part of the Miconia lima complex (Majure and Judd 2013b) and is likely most closely related to M. phrynosomaderma considering the morphological similarities of descending stem hairs with upward, recurved apices and broadly elliptical leaves. Phylogenetic analyses not including material of M. phrynosomaderma place M. limoides as sister to M. lima, a species with which it is sympatric in parts of its distribution and potentially forms hybrids (see discussion under M. lima; Fig. 20C Description. Evergreen shrub (height unknown); stems round in cross section, not ridged, the internodes 1-3.2 cm long; stem indumentum of bulla-based hairs 0.4-1.2 mm long, these mixed with some hairs having strongly dilated bases and oth- ers only narrowly dilated at the base, the hairs apressed-retrorse along stem or slightly spreading with apices recurved; nodal line present, made up of triangular bulla-based hairs to 2 mm long. Leaves broadly elliptic, 2.4-4.3 × 2-3 cm, often slightly anisophyllous, purplish when young; base rounded to acute; apex broadly acute; venation acrodromous, 5-veined, i.e., with midvein and 2 pairs of arching secondary veins, the innermost pair of secondary veins, mostly symmetrical to subsymmetrical at union with midvein 1.5-5 mm above the leaf base, positioned 2.5-5.5 mm in from margin at widest point of blade, the tertiary veins more or less perpendicular to midvein, 2.4-3.5 mm apart at mid-leaf, tertiary veins sometimes joined by quaternary veins; adaxial surface covered in bulla-based hairs, these not meeting at the base, thus the lamina visible between the hairs, i.e., lamina areoles are not completely fi lled, widest hair bases to 1.8 mm wide, apices of bulla-based hairs mostly erect to slightly spreading, the young leaf adaxial surface with ephemeral, long-stemmed, clavate-dentritic hairs, these sometimes fl attened at the apex, arising from between the bases of bulla-based hairs along the primary and secondary veins toward the base of the leaf, and with subsessile to short stalked glandular hairs along the lamina between bulla-based hairs; abaxial leaf surface covered with bulla-based hairs, although the lamina clearly visible, also with bulla-based hairs covering the primary, secondary, tertiary, and quaternary veins, the lamina covered in sessile glands, also with depressions formed from the bullabased hairs on the adaxial leaf surface; petiole 0.4-1.2 cm long, covered in bulla-based hairs, these spreading to retrorse and recurved on adaxial surface and mostly appressedretrorse on abaxial surface. Infl orescences terminal, well-developed to reduced cymes of 3-15 fl owers, 1.7-3.9 cm long, 2.2-5.1 cm across, the peduncle 0.1-0.7 cm long, the proximal branches 0.7-1.7 cm long, and pedicels 0.6-1 cm long; bracts narrowly ovate, 2-3 mm long; bracteoles narrowly ovate, 2-2.2 × ca. 0.2 mm, occasionally with bulla-based hairs at base; nodes of infl orescence with mixed bulla-based hairs and long-stemmed, dentritic-clavate hairs, similar to those found at the base of young leaves. Flowers 4-merous; hypanthium 3.1-4 mm long, 5-5.2 mm wide, more or less spherical, slightly 4-lobed, although lobing mostly obscured by bulla-based hairs 0.9-2.5 mm long, the free portion of hypanthium 0.3 mm long, slightly constricted below the torus, both abaxial surface and base of bulla-based hairs with dark, sessile glands, adaxial surface with sessile-glandular hairs; calyx teeth 2.2-3.3 × 0.4-0.7 mm, linear and terete, recurved upon maturation, covered in long, bulla-based hairs; calyx lobes 4 more or less triangular, 1.3 × 1.6 mm, apex acute, with sessile glands near the apex adaxially and bulla-based hairs abaxially; calyx tube 0.4 mm long; petals 4, white, but purplish on the abaxial surface, ovate, 5.1-5.2 × ca. 3 mm, the apex acuminate, margins membranous and entire, clawed at base, with two bulla-based hairs at the apex on the abaxial surface, these 2-3 mm long; stamens 8, the fi laments 1.7-1.9 mm long, the anthers 1.4-1.5 mm long, with dorso-basal appendage and a single, dorsally inclined pore, the thecae 1.1 mm long. Style ca. 4.3 mm long, dilated at the center, with punctate stigma, subtended by a crown of long, multicelular hairs, these slightly longer than the surrounding bulla-based hairs on the apex of the ovary; ovary ca. 3.2 × 4.8 mm, apex fl at to convex, pubescent with bulla-based hairs. Berries globose, ca. 5 mm long, ca. 6.5 mm wide, blue-black (to purple-black?) at maturity. Seeds (immature) ca. 0.9 mm long, sickle-shaped.
Phenology. Th is species was collected in fl ower and fruit in May. Distribution (Fig 23). Miconia phrynosomaderma is only known from the Massif du Nord, at Morne Belle Terre, Departement du Artibonite, Haiti.
Ecology. Th is species occurs on metamorphic rock at 1050 m elevation, but no information regarding plant community is available.
Conservation status. Miconia phrynosomaderma is likely endangered due to the intense pressure from habitat destruction, as a result of current subsistence farming practices and charcoal production in the Massif du Norde, Haiti. However, there is insuffi cient data to determine actual conservation status of the species. Th us we categorize the species as data defi cient.
Discussion. Miconia phrynosomaderma is most likely most closely related to M. limoides and shares the descending stem hairs with upwardly recurved apices, as well as leaf shape (broadly elliptical; Figs 18,26D). However, the two species diff er in ad-and abaxial leaf surface indumentum, as well as petal base (clawed  Evergreen shrub, 2-3 m tall; stems round in cross section, not ridged, the internodes 0.9-2.6 cm long, stem indumentum of ascending, appressed, moderately bulla-based hairs to 1.6 mm long; nodal line absent. Leaves opposite, decussate, narrowly elliptic to rhomboid, 2.3-6.6 × 0.9-2.3 cm, slightly anisophyllous, apex acute, base acute to slightly rounded, venation acrodromous, 5-veined, the midvein and 2 pairs of arching secondary veins, the outermost intramarginal, secondary veins mostly basal to suprabasal, the innermost pair suprabasal, produced 2.1-9.3 mm from leaf base, po-sitioned 1.0-2.7 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.0-2.3 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface densely covered in well developed bulla-based hairs, these covering the leaf areoles, widest hair bases to 1.8 mm, apices of bulla-based hairs mostly recurved toward the leaf margin, young leaf adaxial surface producing occasional long-stemmed, clavate-dentritic hairs along the primary veins towards the leaf base, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs; abaxial leaf surface covered in bulla-based hairs, these appressed and ascending along the primary, secondary, and tertiary veins, and erect to spreading on those of along quaternary veins and lamina, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina mostly obscured by bulla-based hairs, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface and slightly raised intertertiary veins, sessile to short-stalked, glandular hairs produced from between the bulla-based hairs on the lamina; petioles 0.7-1.1 cm long, covered in ascending, appressed, bulla-based hairs on both surfaces. Infl orescences terminal, 9-26 fl owered cymes, fl owers mostly produced in glomerulate clusters, 2.3-4.1 × 1.0-5.6 cm, the peduncle 0.6-2.5 cm long, proximal infl orescence branches 7-19 mm long; bracts narrowly ovate with an attenuate apex, 1.9-2.1 mm long; bracteoles narrowly ovate, 1.1-1.4 × 0.2-0.22 mm, with an attenuate apex, both bracts and bracteoles only diff erentiated from bulla-based stem hairs by size, otherwise identical, glabrous or basally with bulla-based hairs. Flowers 5-merous, sessile or with pedicels to 0.3 mm long; hypanthium (immature) ca. 2 mm long, short-oblong, unlobed, slightly constricted below the torus, free portion of the hypanthium ca. 0.5 mm long, abaxial surface covered in bulla-based hairs from 1.1-1.3 mm long, and occasional, sessile, glandular hairs near the bases of the bulla-based hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 1-1.3 × 0.4-0.5 mm, ascending or spreading, covered in bulla-based hairs; calyx lobes more or less triangular, 0.8-1.2 × 0.5-0.6 mm, apex acute, covered in bulla-based hairs abaxially and sessile, glandular hairs adaxially; calyx tube apparently not tearing, 0.4-0.6 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 4-5, white, ovate, ca. 1.2 × 0.5 mm (immature), with an acute apex and membranous margin, with one large bulla-based hair produced abaxially, just below the apex, to 0.7 mm long; stamens 8-10; fi laments (immature) ca. 1.2 mm long, glabrous, anthers (immature) 1-1.1 mm long, with one dorsally oriented pore, anther thecae (immature) 0.9-1 mm long, anthers with a dorso-basal appendage to 0.1 mm long; style ca. 0.9 mm long, glabrous, not dilated in the middle, collar absent, style apparently not subtended by a crown of hairs, stigma punctate; ovary ca. 1.2 × 1.1 mm, apex fl at to concave, apparently glabrous, placentation axile with deeply intruded placenta, 2-locular; berries not seen, seeds not seen.
Phenology. Plants have only been collected in June, and these were in bud, and appear to be just before anthesis.
Distribution (Fig 21). Haiti, Dept. du Sud Est, Massif de la Selle, Marigot, Bassin Chotard. Ecology. Virtually nothing is known regarding the ecology of this species, but it was collected at 1750 m in elevation, so was likely growing in a moist montane forest over limestone.
Conservation status. Miconia marigotiana should be considered endangered, as the part of Massif de la Selle north of Marigot from where the species was collected is under intense pressure from habitat destruction, as a result of current subsistence farming practices and charcoal production. However, we do not have suffi cient data to determine the species' current conservation status and thus consider it data defi cient.

Discussion.
Miconia marigotiana is likely very closely related to Miconia lima and M. limoides, however, it diff ers from both species in leaf shape, as well as the 2-carpellate ovaries.
Specimens examined. Th is species is known only from the type gathering by Ekman. Description. Evergreen shrub, 1-3 m tall; stems round in cross section, not ridged, the internodes 0.4-8.9 cm long, stem indumentum of bulla-based hairs 0.1-1.3 mm long, these ascending (antrorse) appressed, mostly arcuate, making the stem appear smooth; nodal line present, with larger bulla-based hairs than those on the rest of the stem. Leaves opposite, decussate, broadly elliptic to obovate, 2.7-5.5 × 1.7-3.8 cm, slightly anisophyllous, apex widely acute to rounded, base acute to rounded, venation acrodromous, 5-7-veined, the midvein and 2-3 pairs of arching secondary veins, the outermost intramarginal, secondary veins mostly basal to slightly suprabasal, the innermost pair, more or less asymmetrical at union with midvein, produced 0.3-4 mm from leaf base, positioned 3-7 mm in from margin at widest point of blade, tertiary veins percurrent, more or less perpendicular to midvein, 1.5-2.9 mm apart at midleaf, intertertiary veins present, tertiary veins often joined by quaternary veins; adaxial leaf surface covered in well developed bulla-based hairs completely fi lling the areoles, bases of bullabased hairs strongly angular (mostly 4-5 angular) produced from the separation of one hair from another, widest hair bases to 2.2 mm, apices of bulla-based hairs mostly erect, young leaf adaxial surface producing long-stemmed, clavate-dentritic hairs along the primary, secondary, and tertiary veins from between the bulla-based hairs, sessile, glandular hairs produced along the primary, secondary, tertiary, and quaternary veins between the bulla-based hairs, especially toward the base of the leaf; abaxial leaf surface covered in bulla-based hairs, these strongly appressed, those along the primary, secondary, and tertiary veins larger than hairs produced throughout the lamina, the lamina completely covered in bulla-based hairs and thus obscured, lamina appearing as a series of pits from depressions of the bulla-based hairs produced from the upper leaf surface and slightly raised intertertiary veins, sessile, glandular hairs produced from between the bulla-based hairs; petioles 0.4-1.7 cm long, covered in appressed-ascending, bulla-based hairs on both surfaces. Infl orescences terminal, of 5-15 fl owered, condensed-short cymes, fl owers mostly produced in glomerulate clusters, 0.9-2.8 × 1.2-2.8 cm, the peduncle 0.05-1.5 cm long, proximal infl orescence branches 2-8 mm long, pedicels absent to 1.2 mm long; bracts oblong to ovate, 2.6-4.9 mm long; bracteoles narrowly ovate, 1.7-3.8 × 0.3-0.6  mm. Flowers 4-5(6)-merous, sessile or with pedicels to 1.2 mm long, when 4 or 5-merous, sometimes with one or two calyx teeth apparently aborted; hypanthium 2.9-3.5 mm long, short-oblong to globose, 4-lobed, but lobing mostly obscured by bulla-based hairs, slightly constricted below the torus, free portion of the hypanthium 0.8-1.1 mm long, abaxial surface covered in bulla-based hairs from 0.8-2.6 mm long, and occasional, sessile, glandular hairs near the bases of the bulla-based hairs; adaxial surface (i.e., free portion) covered in small, bulla-based hairs; calyx teeth 2.9-4.2 × 0.8-0.9 mm , ascending or spreading, covered in bulla-based hairs; calyx lobes more or less triangular, 1.3-1.6 × 1.5-2 mm, apex acute, covered in bulla-based hairs abaxially and sessile, sparse, glandular hairs adaxially; calyx tube not tearing, 0.1-0.4 mm long with bulla-based hairs abaxially and sessile, glandular hairs adaxially; petals 4-5(6), red to violet-red, elliptic, 4.5-7 × 2.6-3.6 mm, with an acute apex and membranous margin, with one or two slightly bulla-based hairs produced abaxially, just below the apex, to 2.8 mm long; stamens 8-10 (12); fi laments 1.8-2.8 mm long, glabrous, anthers 1.5-1.6 mm long, with one dorsally oriented pore, anther thecae 1.2-1.4 mm long, anthers with a dorso-basal appendage 0.18-0.3 mm long; style 5.0-5.3 mm long, glabrous, not or only slightly dilated in the middle, collar absent, style subtended by a crown of multicellular, linear to elongate-triangular (needle-like) hairs, which are slightly longer than the surrounding bulla-based hairs of the ovary apex, stigma punctate; ovary 1.6-2.2 × 2.4-3.4 mm, apex concave, pubescent with bulla-based hairs, except for the linear or elongate-triangular hairs forming crown, placentation axile with deeply intruded placenta, 4-locular; berries globose, slightly 4-lobed, purple (to purple-black) at maturity, 4-10 mm long (including calyx tube), 5.5-9 mm wide, seeds 0.8-1mm long, obpyramidal, often falcate, testa smooth, light brown, raphe dark brown, smooth, extending the length of the seed.

Miconia paralimoides
Phenology. Miconia paralimoides has been collected in bud, fl owering, and in immature fruit from September through April, and has been collected in mature fruit from February through April. Distribution (Fig 23). Miconia paralimoides is restricted to the Cordillera Central, Dominican Republic.
Ecology. Miconia paralimoides occurs in humid, broadleaved mixed forests or pine-dominated cloud forests from 1000-1950 m in elevation. Miconia paralimoides occurs with Miconia pedunculata and Miconia lima in part of its range, although it is unknown if all three species are sympatric in a single area. Other associated melastomes include Mecranium puberulum Cogn., Meriania involucrata, Miconia umbellata, and Sagraea fuertesii (Cogn.) Alain.
Conservation status. Certain populations of M. paralimoides are located inside either national parks or scientifi c reserves (i.e., Parque Nacional Jose del Carmen Ramirez, Reserva Científi ca Ébano Verde), however, the species also is under threat from habitat fragmentation and loss outside of those areas. We therefore suggest this species be given a preliminary conservation assessement of vulnerable.
Phenology. Th is species has been collected in fl ower and fruit from May through December. However, fl ower buds are present on the type specimen, which was collected in February, as well as on Majure 6053 also collected in Februrary, so this species likely fl owers throughout most of the year. Distribution (Fig 21). Miconia pedunculata is restricted to the Cordillera Central, Dominican Republic.
Conservation status. Although this species apparently can tolerate a range of anthropogenic disturbance (according to label data) and also occurs within a scientifi c reserve in a portion of its distribution, it is of limited distribution in the Cordillera Central, Dominican Republic, and not terribly abundant where it is found. Th us, we assign a preliminary status of vulnerable to M. pedunculata.
Discussion. Th e northern populations of M. pedunculata (e.g., those at the Reserva Científi ca Ébano Verde) are morphologically diff erentiated from the southern populations (e.g., from Prov. Peravia), and can be recognized by the less well-developed bullabased hairs on the adaxial leaf surface, and their more densely pubescent abaxial leaf surfaces. However, the minor morphological diff erences in these two geographically disjunct groups of populations appear to merely represent variation within a single species. Accessions of M. pedunculata from both of these regions form a well-supported clade in phylogenetic analyses of the group (Majure et al. 2015; Fig. 2).  Description. Evergreen shrub, 0.5-0.9 m tall, stems round in cross section, not ridged, the internodes 0.3-0.67 cm long, stem indumentum of ascending, appressed bulla-based hairs, the longest to 0.6 mm long, nodal line present. Leaves oppostie, decussate, elliptic, small, 0.9-2.3 × 0.5-1.2 cm, base acute, apex obtuse to acute, venation acrodromous, 5-veined, the midvein and two pairs of arching secondary veins, outermost pair of secondary veins basal, intramarginal, innermost pair suprabasal to basal, produced 0.4-1.4 mm from the base, 1.1-2 mm in from the margin at widest point of leaf blade, tertiary veins percurrent, more or less perpendicular to the primary vein, 0.9-1.7 mm apart at midleaf, intertertiary veins obscure but present, tertiary veins often joined by obscure quaternary veins; adaxial leaf surface covered in well developed bulla-based hairs, these completely covering the leaf areoles, the largest 1.6 mm wide at the base, clavate-dendritic hairs prominent in young leaves along the primary, secondary, tertiary and quaternary veins, especially toward the base, abaxial leaf surface covered in bulla-based hairs, those along the primary and secondary veins larger than those along the epidermis and tertiary veins, mostly ascending to appressed, prominent pits formed from the adaxial surface hairs, sessile, glandular hairs throughout the lamina; petioles 1.5-4.5 mm long, covered in ascending, appressed bulla-based hairs. Infl orescences terminal, structure indiscernable (only immature infl orescence seen).

Miconia pagnolensis
Phenology. Immature infl orescences can be seen on the type specimen collected in late June. Distribution (Fig 21). Th is species is restricted to the Massif de la Hotte, Haiti, and is only known from the type locality.
Etymology. Th e specifi c epithet, pagnolensis, refers to the type locality, Morne Bois Pagnol, and the only place that this species has been collected. It should be noted that there are diff ering spellings for the type locality. Th e word is derived from the French, Espagnol (meaning Spanish). Th e Haitian Creole spelling for the site is Panyòl, although it is also spelled Pangnol and Pagnol (J. Timyan, pers. comm.). We have chosen to use the spelling Pagnol for the site and thus the specifi c epithet, pagnolensis.
Ecology. Th is species has been collected in a moist, broadleaved forest on karst limestone at around 1200 m in elevation and is associated with Abarema sp. Conservation status. Th e conservation status of this species is unknown, however, the location of this collection is surrounded by degraded, remnant moist tropical forest, most of which has been cleared for subsistence farming. Th us, M. pagnolensis is likely endangered by habitat loss. However, we suggest a preliminary status of data deffi cient until future fi eldwork can further illuminate population sizes of the species.
Discussion. Material of this new species has been collected only once by Joel Timyan (i.e., Timyan 27, FLAS). According to phylogenetic analyses , this species is phylogenetically distinctive and most closely related to members of the Acuminata subclade of the Lima clade (Fig 2). Miconia hybophylla is the only other putative member of the Acuminata subclade in Haiti. Additionally, vegetative morphology certainly places M. pagnolensis within the Lima clade (Fig. 26F). Th e type specimen shows phenetic similarity with M. lima, but the shrubs are smaller than M. lima (0.5-0.9 vs. 1.5-5 m tall), with mostly smaller leaves (0.9-2.3 × 0.5-1.2 cm vs. 0. 7-5.7 × 0.42-3.2 cm), with obtuse to acute (vs. acute or acuminate) apices. Miconia pagnolensis also occurs outside of the range of M. lima, which, in the southern peninsula of Haiti, is only known from the Massif de la Selle. We formally name this species, despite its being known only from an essentially vegetative specimen, because our analysis of molecular data clearly indicates the phylogenetic distinctiveness of the taxon, and because its vegetative morphology is diagnostic (allowing it to be distinguished from the phenetically similar M. lima). Th e specifi c epithet refers to the type locality -which is quite inacessable. We think it is important to have a formal name for this rare taxon, as it may be years before it is re-collected (unfortunately, like several other species of sect. Lima, which are only known from the type gathering), and knowledge of its existence will stimulate searches. and American University of Les Cayes) for help with fi eldwork in Haiti, Joel Timyan (Société Audubon Haïti) for material from Massif de la Hotte, Fabián Michelangeli (NYBG) for photos of several Cuban species, Dan Skean for photos and information about collections, and Gretchen Ionto for taking the SEM images. Financial support for this work was provided by a grant from the National Science Foundation to F. Michelangeli and co-PIs (BSR-0818399) and the Desert Botanical Garden.