Miconia bullotricha and M. hirtistyla, two new species of Miconia sect. Lima (Miconieae, Melastomataceae) from eastern Cuba

Abstract We describe two new species in Miconia sect. Lima, Miconia bullotricha Bécquer & Majure and Miconia hirtistyla Majure & Judd, from eastern, Cuba. We also provide illustrations and distribution maps for the two species, as well as a key to members of the Lima clade on Cuba.


Key to the species of Miconia sect. Lima on Cuba
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 present, inconspicuous. 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, ± 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 ± 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; domatia inconspicuous, of multicellular, tufts of 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 fl owers produced in 3-7 fl owered dichasia, 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, glabrous or with small bulla-based hairs at base, bracteoles generally resembling one large, bulla-based hair. Flowers perfect, actinomorphic, 4-merous, with 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 bullabased hairs; calyx teeth 1.75-2.2 × 0.5 mm, linear and terete, recurved upon maturation, covered in bulla-based hairs; calyx lobes ± triangular, apex acute, ca. 1 × 1.3 mm, 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, (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 well-developed 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 bulla-based hairs on the ovary apex; stigma punctate; ovary ca. 1.4 × 2.4 mm, apex fl at, with bulla-based hairs, 4 locular, with axillary placentation, the placenta deeply intruded into locule; berries (immature) globose, ca. 3-3.4 × 3 mm; 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. Etymology. Th e specifi c epithet " bullotricha " refers to the well-developed bullabased hairs on the adaxial leaf surface. Although bulla is Latin in origin, we base formation of our compound epithet on the Greek rules for connecting vowels. Th us, we use "o" here instead of "i", as we fi nd " bullotricha " to be more euphonious than " bullitricha ." Th e connecting vowel "o" has had widespread usage in classical Latin based on the large infl uence of Greek (Stearn 1966).
Conservation status. We do not have extensive knowledge of population level numbers of individuals or the reproductive biology of this species, so the conservation status of M. bullotricha cannot be critically evaluated at this time. More fi eldwork is imperative to assess the status of this species. However, deforestation has occurred in the surrounding areas from where M. bullotricha is known, and thus, the species most likely should be considered threatened by habitat loss and other anthropogenic disturbances.  Discussion. Miconia bullotricha likely belongs to a subclade within the Lima clade that contains the phenetically similar, Cuban endemic, M. ottoschmidtii , as well as other members of the M. lima complex (Majure et al. 2013b, Majure et al. unpubl. data). Th ese species are recognized by their very well developed bulla-based hairs on the upper leaf surface (which mostly cover the leaf areoles; Fig. 1), as well as expanded, pyramidal infl orescences consisting of cymose clusters of fl owers subtended by highly reduced bracts. Th e only exception to this is M. pedunculata Majure & Judd of the Cordillera Central, Dominican Republic that has widely spaced bulla-based hairs on the adaxial leaf surface, which do not completely fi ll the leaf areoles, and fl owers that are subtended by foliaceous bracts.
As mentioned above, Miconia bullotricha is phenetically most similar to M. ottoschmidtii . However, the two species can be easily distinguished by stem and leaf indumentum, where M. bullotricha has the stem indumentum generally with apices attenuate and strongly recurved upwards, (as opposed to granulate with the apices truncate or short attenuate in M. ottoschmidtii ; although it should be noted that central and northern Cuban populations of M. ottoschmidtii have a tendency towards stem hairs with longer, attenuate apices that may be recurved upwards), and erect bullabased hairs throughout the lamina and along the tertiary veins on the leaf abaxial surface (while M. ottoschmidtii has spreading to appressed bulla-based hairs throughout the lamina and along the tertiary veins of the leaf abaxial surface). Miconia bullotricha usually produces entirely pendant infl orescences (Fig. 1), in contrast to the erect infl orescences of M. ottoschmidtii and the rest of the members of the subclade (however, several species often produce pendant basal infl orescence branches, including M. ottoschmidtii ), and has longer calyx teeth than M. ottoschmidtii (1.75-2.2 vs. 0.4-0.8 mm). Also, M. bullotricha has globose fl oral buds, while M. ottoschmidtii exhibits quadrangular fl oral buds.
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, 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, ± 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 bulla-based 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 (i.e., foveolate), sessile, black, glandular hairs produced along all major and minor veins, domatia of tufts of multicellular, linear 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, bullabased hairs on both surfaces. Infl orescences terminal, cymose, 2-5 fl owered, 1.3-2.4 × 1.2-3.8 cm, the fl owers mostly produced in glomerulate clusters, 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 perfect, actinomorphic, 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 bulla-based 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 bullabased hairs; calyx lobes 6, ± triangular, apices acute, 1-1.4 × 1-1.5 mm, covered in bullabased 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 (i.e., with scattered, slightly bulla-based hairs), oblong to only slightly dilated in the middle, collar absent, style subtended by multicellular, linear to elongatetriangular (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, with bulla-based hairs, placentation axile, placenta apparently not deeply intruded, 3-locular; berries not seen, mature seeds not seen.
Distribution and habitat. Miconia hirtistyla is only known from the western Sierra Maestra, Cuba (Fig. 2), where it occurs in montane rainforest, pine forest and elfi n forest on rocky soils at elevations of 700-1800 m. Associated melastomes include Miconia argentimuricata , Miconia norlindii and Miconia nystroemii Urb.
Phenology. Miconia hirtistyla was collected in bud, at anthesis, and in immature fruit in March and July.
Etymology. Th e specifi c epithet " hirtistyla " refers to the pubescent style of this species (Fig. 3). Within the Lima clade, M. hirtistyla is the only species that demonstrates this character.
Conservation status. Miconia hirtistyla is mostly known from the very well protected forests of Turquino National Park. Although the species has not been collected since 1978, and we know nothing regarding its reproductive biology or population numbers, it is most likely not threatened by anthropogenic disturbance and habitat loss, at least in the areas immediately surrounding the park. Fieldwork will be necessary to appropriately assess the conservation status of this species. Discussion. Miconia hirtistyla is the only species in the Lima clade known to possess pubescent styles and it is one of two species that exhibits clawed petals (e.g., M. phrynosomaderma Majure & Judd; Majure and Judd 2013a, a putatively distantly related species). Both characters are likely autapomorphies of M. hirtistyla, because mo rphology suggests that M. phrynosomaderma is more closely related to M. limoides and relatives (e.g., well-developed bulla-based hairs on leaf adaxial surface, open, expanded, cymose infl orescences, presence of anther dorso-basal appendages; see Majure and Judd 2013). Miconia hirtistyla is most likely closely related to M. jashaferi (Fig. 4), with which it had been confused, as well as M. cubacinerea and M. tentaculicapitata . All of these species have condensed infl orescences, leaf-like bracts and bracteoles, broad, oblong to obovate petals, a crown of long, needle-like hairs on the ovary apex and surrounding the style or merely needle-like hairs produced throughout the ovary apex, long, fi liform, eglandular or gland-headed hairs along the calyx lobe adaxial surface and apex of calyx tube, long calyx teeth, as well as "shallowly" intruded placenta (versus deeply intruded placenta as in most other species of the Lima clade). All four species also lack a dorsobasal anther appendage, the presence of which otherwise is a common feature in the clade (Majure and Judd 2013a). Miconia hirtistyla diff ers from all three of these species by the presence of pubescent styles and clawed petals and from M. jashaferi and M. cubacinerea by hypanthium shape (short oblong to globose in M. hirtistyla vs. narrowly oblong to cylindrical in the latter two species). Th e species also diff ers from M. jashaferi in infl orescence structure (erect vs. pendant infl orescences; Figs 3, 4), leaf shape (elliptic to ovate with broadly to narrowly acute apices rather than mostly ovate with acuminate apices), anther size (2.2-2.6 mm long in M. hirtistyla vs. 1.8-2 mm long in M. jashaferi ; Figs 3, 4) and shape (ovate with anther thecae continuous with sterile portion of anther vs. elliptic with anther thecae discontinuous with sterile portion of anther; Figs 3, 4), as well as having apically oriented anther pores instead of dorsally oriented pores. Miconia hirtistyla diff ers from M. cubacinerea in the pubescence of the abaxial leaf surface, in that M. cubacinerea has a clearly visible epidermis as a result of a sparser indumentum, while the epidermis of M. hirtistyla is mostly concealed by dense bulla-based hairs. Likewise, the primary, secondary, tertiary, and quaternary veins of M. hirtistyla are densely clothed in spreading bulla-based hairs, however, in M. cubacinerea the veins are easily seen, as the bulla-based hairs are less dense. Th e abaxial leaf surface of M. cubacinerea also is densely covered in sessile, glandular hairs, while that of M. hirtistyla has sparse, glandular hairs. Th e lamina of the abaxial leaf surface of M. hirtistyla is conspicuously, deeply pitted (as a result of the bulla-based hairs on the upper leaf surface to 0.5 mm deep), while that of M. cubacinerea is not deeply pitted (i.e., the pits are only superfi cial to <0.1 mm deep). Th e two species also diff er in calyx teeth length (4.5-4.6 mm in M. hirtistyla vs. 5.7-6.2 mm in M. cubacinerea ), and by the lack of clavate-dendritic hairs on the leaf adaxial surface and calyx teeth in M. hirtistyla .
Lastly, M. hirtistyla diff ers from M. tentaculicapitata by the less well-developed bulla-based hairs on the leaf adaxial surface, spreading to descending stem hairs, and lack of clavate-dendritic hairs on the leaf adaxial surface , as opposed to the well developed bulla-based hairs covering the leaf adaxial surface areoles, the ascending-appressed stem hairs, and presence of clavate-dendritic hairs on the leaf adaxial surface of M. tentaculicapitata .
Miconia hirtistyla, and the less phenetically similar, M. tentaculicapitata, are found in the western Sierra Maestra, while those species that are more phenetically similar to M. hirtistyla , i.e., M. cubacinera and M. jashaferi, are found in northeastern Cuba in the Sierra de Baracoa and Sierra de Moa regions; M. jashaferi also is found in the southern part of Sierra de Cristal.
Miconia hirtistyla is most likely a cladospecies (Donoghue 1985), as indicated by the putative autapomorphies of pubescent styles and clawed petals. Th e species also adheres to the morphological/phenetic species (Judd 2007) and diagnostic species concepts (Wheeler and Platnick 2000).