Research Article |
Corresponding author: Kenneth J. Wurdack ( wurdackk@si.edu ) Academic editor: Ricardo Kriebel
© 2019 Kenneth J. Wurdack, Fabián A. Michelangeli.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Wurdack KJ, Michelangeli FA (2019) Systematics and relationships of Tryssophyton (Melastomataceae), with a second species from the Pakaraima Mountains of Guyana. PhytoKeys 136: 1-21. https://doi.org/10.3897/phytokeys.136.38558
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The systematics of Tryssophyton, herbs endemic to the Pakaraima Mountains of western Guyana, is reviewed and Tryssophyton quadrifolius K.Wurdack & Michelang., sp. nov. from the summit of Kamakusa Mountain is described as the second species in the genus. The new species is distinguished from its closest relative, Tryssophyton merumense, by striking vegetative differences, including number of leaves per stem and leaf architecture. A phylogenetic analysis of sequence data from three plastid loci and Melastomataceae-wide taxon sampling is presented. The two species of Tryssophyton are recovered as monophyletic and associated with mostly Old World tribe Sonerileae. Fruit, seed and leaf morphology are described for the first time, biogeography is discussed and both species are illustrated.
Guyana, leaf anatomy, Melastomataceae, molecular phylogeny, seeds, Sonerileae
The flora of Guyana contains about 290 taxa of Melastomataceae in 40 genera, including three small endemic genera, Maguireanthus Wurdack, Ochthephilus Wurdack and Tryssophyton Wurdack (
Tryssophyton has been little studied since its description and collections have remained few. A 2012 expedition in the Pakaraima Mountains to reach the botanically unexplored ca. 1700 m summit of Kamakusa Mountain yielded new collections of the type species, T. merumense Wurdack, at lower elevations and a novel new species at the summit, which is described herein. We also provide further observations on the morphology and relationships of both species. Kamakusa Mountain and its vicinity at the wet, eastern edge of the Pakaraima Mountains have yielded many new plant taxa from the two expeditions (1960, 2012) that have traversed this remote region and it deserves further scientific exploration (
In order to ascertain the phylogenetic position of both species of Tryssophyton, we sequenced plastid rbcL and/or ndhF following protocols and primers used in previous broad studies of the Melastomataceae (i.e.
Scanning electron microscopy (SEM) used a Zeiss EVO MA15 (Carl Zeiss SMT, Inc., Peabody, Massachusetts) at 6–12 kV after sputter-coating herbarium specimen seeds or critical point dried (CPD) field-fixed (in ethanol) leaves with 25 nm of C + Au/Pd using a Leica EM ACE600 (Leica Microsystems GmbH, Wetzlar, Germany). Leaves were examined for venation after clearing in ethanol or 2.5% sodium hydroxide. For internal structure, they were hand cut for SEM or, for light microscopy (LM), they were paraffin-embedded, sectioned at 10 μm, stained with toluidine blue O and examined with a Zeiss Universal Compound Microscope. The anatomy of the delicate leaves was more intact with CPD and SEM than after traditional thin sectioning from the same starting material, thus our observations are largely based on SEM.
Melastomataceae are resolved as a strongly supported (BP 100) family with mixed backbone support, which of relevance here was notably weak (BP 60 or less) amongst the deepest nodes of Sonerileae (Fig.
Differs from Tryssophyton merumense in 4-verticillate, ovate, petiolate leaves, versus 10–16 verticillate, oblanceolate-lanceolate, subsessile leaves.
Guyana. Cuyuni-Mazaruni Region: Summit of Kamakusa Mtn. (i.e. on top of 4th escarpment of four); impenetrable elfin forest to 3 m, extremely dense and wet, rich in epiphytes, with Bonnetia (2 spp.), Brocchinia cf. tatei, Malpighiaceae, Melastomataceae, Cyperaceae spp., Weinmannia, Ilex cf. retusa, 5°52'50.9"N, 60°6'11.7"W, 1691 m elev., 8 June 2012 (fl.), K. Wurdack 5865 with E. Tripp, A. Radosavljevic, and J. Ralph (holotype: BRG!, isotype: US-3731242!).
Habit perennial, rhizomatous herbs; rhizomes persistent, fleshy, horizontal, to 10 cm long, 3–6 mm diam., rarely branched, bearing 1–2 leafy, erect, aerial stems per axis and older cup-shaped (collar to 0.3 mm high and centre sunken) aerial stem scars; adventitious roots fine, fibrous. Indument sparse on aerial stems, leaves, pedicels, hypanthium and sepal margins; trichomes simple, to 0.1 mm long, glandular, reddish. Aerial stems 8.5–11.5 cm tall, 0.9–2 mm diam., terete, purple, slightly flared at rhizome attachment, summit with cluster of leaves and often 1 secondary branch 1–2 cm long and crowned by another cluster of leaves. Leaves 4-verticillate, opposite pairs sub-equal in size, simple, petiolate, exstipulate; petioles 2–5 mm long (of slightly subequal length within whorl), ca. 1 mm diam.; lamina 2.1–5.2 × 1.0–1.9 cm, length:width ratio 2.2–3.7:1 (mean = 3.0, SD = 0.50, n = 11), symmetrical, ovate to lanceolate, membranous, apex acuminate, base cuneate, margin minutely serrate, with 6–12 teeth per side (2–5 teeth/cm); teeth 0.5–0.9 × 0.1 mm, first-order, spacing regular, projecting 0.2–0.3 mm from margin, sinus shape rounded, distal flank concave and proximal flank straight, apex long-attenuate; leaf tip with 1–3(13) adaxial scales, 0.6–1 × 0.1 mm, similar to attenuate teeth apices. Venation suprabasal acrodromous, with one pair of major (costal) secondaries 1/2 of the gauge of the midvein, joining the midvein 0.5–1 mm above the leaf base; and one pair of intramarginal secondaries < 1/3 of the gauge of the midvein (poorly defined from tertiary thickness), joining < 0.5 mm from base, traversing 0.2–0.8 mm from margin, distally fading into exterior tertiary loops; up to 12 interior tertiary veins per side, quaternaries random reticulate. Inflorescence terminal, pedunculate, bearing 1–4 flowers; peduncle 2.2–2.6 cm long, 0.5–0.7 mm diam. mid-length, terete, flaring at base, purple; bracts persistent, lanceolate, 1–1.3 × 0.3–0.4 mm, apex apiculate, margin entire. Flowers 4-merous, bisexual, pedicellate; pedicels 8.5–11.5 mm long, 0.4–0.6 mm diam. mid-length, terete, purple; hypanthium at anthesis ca. 2.5 mm × 2–2.5 mm (excluding calyx), campanulate, obscurely costate with thin ribs; calyx lobes 4, in bud narrowly triangular, ascending, protective of young corolla, at anthesis spreading, 0.8–1 mm tall, broadly triangular above short calyx tube ca. 0.4 mm high; petals ca. 9.5 × 5 mm, margins entire, in vivo red outside and pink inside. Stamens 8, incurved in bud with tips extending into hypanthium below point of filament attachment, slightly anisomorphic, with the antesepalous whorl larger than the antepetalous, glabrous; filaments 5–6 (antesepalous) or 4 (antepetalous) mm long, 0.2 mm diam., linear, pink in vivo; thecae 4.5 or 3.5 mm long, basifixed, connective basally prolonged, dilated below thecae, forming a thickened annulus that is more or less ventrally bilobate, thickening 0.8 or 0.5–0.6 mm diam., yellow in vivo; anther 0.1–0.2 mm wide at terminal pore. Ovary superior, ca. 1.8 × 1.3–1.5 mm, glabrous. Style ca. 10 mm long, 0.2 mm diam., curved, glabrous; stigma punctiform, minutely papillose. Capsule ovate, ca. 3 × 3.5 mm, crowned by persistent calyx. Seeds ovoid, ca. 0.7 × 0.4 mm (immature and partly collapsed), sparsely papillose, brown.
The specific epithet is derived from quadri- (Latin, four) and folium (Latin, leaf) and refers to the 4-verticillate leaves.
Tryssophyton quadrifolius is only known from the summit of Kamakusa Mountain where it was encountered as an infrequent epiphyte on moss and lichen covered branches and trunks of large shrubs. Scattered small plants with single stems were observed sporadically along the main trail transect cut along the north-south orientated narrow summit. The few reproductive plants were in a more sheltered spot on the western edge of the summit on a branch trail that was cut to a cliff edge for a view. Flowers and young fruit were collected in June. The summit of Kamakusa Mountain is covered by low montane evergreen cloud forest, 3–4 m tall, with dense thickets dominated by Bonnetia tepuiensis Kobuski & Steyerm. and B. roraimae Oliv. (Bonnetiaceae), Byrsonima pachypoda W.R. Anderson (Malpighiaceae), Miconia acutifolia Ule and M. silicicola Gleason (Melastomataceae), Raveniopsis microphyllus K. Wurdack (Rutaceae) and species of Weinmannia L. (Cunoniaceae) on a peat substrate overlying sandstone. Vascular epiphytes included species of Utricularia L. (Lentibulariaceae), Bromeliaceae and Orchidaceae.
While the upper part of Kamakusa Mountain is presently pristine and undisturbed, the new species is a delicate epiphytic herb with few reproductive plants (2 of 15 aerial stems collected had buds, flowers and/or fruit) in an area of extremely limited montane habitat. The species is vulnerable to climate and land use changes, such as regional gold mining (see
Illustration of Tryssophyton quadrifolius (A–G) and T. merumense (H, I). A Habit (right view in flower) B young flower with anthers inflexed C flower with anthers erect and petals fallen D shorter antepetalous anther, dorsal E longer antesepalous anther, dorsal F longer anther, lateral G young fruit H habit I capsular, 3-merous fruit. Sources: A–G Wurdack 5865 H Radosavljevic 165 I Wurdack 5870 (all US).
Guyana. Cuyuni-Mazaruni Region, Partang River, Merume Mtns., Merume Mt.; common on mossy logs in forest, 1140 m elev., 4 Jul 1960 (fl., fr.), S.S. Tillett, C.L. Tillett, & R. Boyan 43988 (holotype: US-2343844!; isotypes: K-000329332!, NY-00245868!).
It should be noted that both the US and NY sheets are each marked as the holotype. However, the protologue clearly states that the US specimen is the holotype, even citing the sheet number and has that designation in J. Wurdack’s handwriting. The NY sheet has “holotype” merely typed on the label. Thus, there is no need to lectotypify this name and the NY specimen should be considered as an isotype.
The genus is combined from tryssos (Greek, dainty or delicate) and phyton (Greek, plant) and refers to the plant habit. The specific epithet refers to Merume Mountain where the type was collected.
Guyana. Cuyuni-Mazaruni Region: Pakaraima Mountains, upper Karowrieng River at Maipuri Falls; mixed bryophyte, pteridophyte, herb community; sandstone boulders, white sand, large cave behind falls, 5°41'N, 60°13'W, 575–600 m elev., 13 Oct 1992 (fl.), B. Hoffmann 2939 (NY!, US!). 2nd and 3rd escarpments (of four) of Kamakusa Mt., upper west-facing slopes below summit, rich forest with Licania, Ebenaceae, tree ferns, Arecaceae, 5°52'55.2"N, 60°6'34.5"W, 1330 m elev., 8 June 2012 (fl., fr.), K. Wurdack 5870 (US!). Potaro-Siparuni Region: Mt. Wokomung, easternmost pinnacle of massif, scrub forest on sandstone and peat, with Guadua, Euterpe, and Sphagnum, 5°5'34.4"N, 59°50'13.3"W, 1524 m elev., 13 Jul 2003 (fl.), H.D. Clarke 10822 (NY!, US!). Mt. Ayanganna, east slope, plateau above second escarpment, growing on mossy tree trunks and roots, 5°22'28"N, 59°58'06"W, 1340 m elev., 16 Mar 2014 (fl.), A. Radosavljevic 165 (US!).
The five collections of T. merumense span a 90 km section of the central Pakaraima Mountains, but further exploration is likely to expand its range into similar habitats. The species was only recently discovered (Radosavljevic 165) on the slopes of relatively well-explored Mount Ayanganna, the highest mountain (2041 m) wholly within Guyana. At mid-elevations on the western slopes of Kamakusa Mountain, it occurred (Wurdack 5870) as scattered, rarely-reproductive individuals on rotting logs and peaty-humus zones around the bases of trees.
The mountainous area north of the village of Imbaimadai and including Kamakusa Mountain has been variously mapped as the Merume Mountains. However, exactly what corresponds to the peak “Merume Mountain” within the region and indicated as the type locality of T. merumense, is unclear. Field notes (Bassett Maguire Field Collections, vol. 19, Archives of The New York Botanical Garden) reveal that during 11 Jun–16 Jul 1960, after leaving Imbaimadai, the collecting team, led by Stephen Tillett entered the Kamakusa Mountain area from along the Partang River. After reaching Partang Falls, they ascended into the uplands following existing trails, which were probably made by gold-miners or “pork-knockers.” It is likely that “Merume Mountain” of Tillett et al. is equivalent to Kamakusa Mountain, but details referring to lower elevations, southeast ridge, southeast side and cliffs do not indicate they reached summit where T. quadrifolius was collected.
The broader relationships which we recovered within Melastomataceae largely agree with those from prior studies using the underlying sequence data (i.e.
The more robust placement of Tryssophyton within Sonerileae has clearer implications for taxonomy and historical biogeography. When originally describing Tryssophyton, J.
The great majority of the species in the Sonerileae-Dissochaeteae complex are found in the Old World. The exceptions are Tryssophyton and the Neotropical genera Boyania and Phainantha, resolved as successive branches, with poor support, at the base of the clade. Boyania contains two trailing (stoloniferous) or climbing herbaceous species with disjunct distributions. One of them, Boyania ayangannae, also grows in the same region as Tryssophyton, while the second species, Boyania colombiana, is restricted to the easternmost slopes of the Colombian Andes (
Although reproductive features (flowers and fruit) and overall habit (rhizomatous herbs) are nearly identical between the two species, Tryssophyton quadrifolius is vegetatively easily distinguished by fewer leaves per stem and differences in leaf architecture (see Diagnosis; Figs
Macromorphology of Tryssophyton merumense (A–C, G–K) and T. quadrifolius (D–F). A habit, growing with bryophytes and downward pointing red buds B flowering plant, adaxial C Flowering plant showing bicoloured petals, abaxial D part of type collection in vivo just before pressing E whole leaf venation, abaxial F venation close-up showing major veins and reticulate quaternaries, abaxial G Dehisced 4-merous fruit, with interior persistent fimbriate placenta remnants (f), axial (tips of 8 triangular valves partly broken) H fruit, with seeds, fimbriae (f) and carpel septa, transverse view I whole leaf venation and crystal druses (white spots), adaxial J close-up of leaf tip with marginal teeth and non-vascularised scales (s), adaxial K phyllotactic arrangement with leaf attachment scars surrounded by darkened glandular trichomes, lateral view with main stem at bottom, peduncle at top. Sources: A–C, I–K Radosavljevic 165 D–F Wurdack 5865 G, H Wurdack 5870 (all US).
While both Tryssophyton species grow on Kamakusa Mountain, they do not appear to be sympatric (K. Wurdack, personal observation) and their local habitats differ in elevation, exposure and vegetation type. Tryssophyton quadrifolius would appear poorly adapted from an ecophysiology perspective as a summit endemic, with relatively large thin leaves, compared with much of the associated montane vegetation which has reduced sclerophyllous and/or coriaceous physiognomies as adaptations to cold and exposure (see
Micromorphology of Tryssophyton merumense. A Leaf glandular trichome, adaxial B leaf with marginal teeth, erect adaxial scales and glandular trichomes, lateral view of distal part C leaf, with single file palisade parenchyma (p), loose spongy parenchyma (s) and stomata (st) beneath intercellular spaces, close-up of transverse view D leaf scale, transverse view near adaxial scale base E stomata, abaxial F leaf, transverse view G seed, lateral view with raphal zone near bottom. Sources: A–F Radosavljevic 165 G Wurdack 5870 (all US).
Amongst reproductive features, the buds of both Tryssophyton species are red due to a pigmented layer of the outer (abaxial) exposed parts of the petals, which then open to reveal lighter inner surfaces. Tryssophyton merumense has white inner (adaxial) petal surfaces and filaments, leaving the exterior strikingly bicoloured where the petals overlapped in bud (giving them a distinctive “candy-cane” pattern) and were still evident on the holotype (Fig.
Fruit and placenta morphology are remarkably diverse in Melastomataceae and phylogenetic evidence indicates complexity and homoplasy in fruit type evolution (
The seeds of Tryssophyton merumense are 0.6–0.7 by 0.4–0.5 mm, ovoid, lack appendages and are minutely papillose with sinuous interdigitating testa cell patterning (Fig.
We thank Alice Tangerini for the botanical illustration (Fig.
Sources for plastid data used in the phylogenetic analysis of Tryssophyton. Ordered as: Taxon, Voucher (new data only), GenBank numbers for ndhF, rbcL and rpl16, respectively. Dash (-) indicates missing data, asterisk (*) indicates taxa that presently appear under a synonym in GenBank and bold indicates new data.
Outgroups: Alzatea verticillata Ruiz & Pav., AF215591, -, AY151598. Eugenia uniflora L., AF215592, AF294255, AF215627. Myrtus communis L., AF215593, AF294254, AF215628. Olinia ventosa (L.) Cufod., AF215594, AF215546, -. Penaea mucronata L., AF270756, AJ605090, AF222782. Rhynchocalyx lawsonioides Oliv., AF270757, AF215547, AF215631. Melastomataceae: Aciotis purpurascens (Aubl.) Triana, AF215561, -, AF322231. Allomaieta ebejicosana Lozano, JF831961, JF831986, JF832012. Allomaieta grandiflora Gleason, JF831962, JF831987, JF832013. Allomaieta hirsuta (Gleason) Lozano, JF831963, JF831988, JF832014. Allomaieta pancurana Lozano, JF831967, JF831993, JF832017. Allomaieta villosa (Gleason) Lozano, JF831969, JF831994, JF832019. Allomaieta zenufanasana Lozano, JF831970, JF831995, JF832020. Alloneuron ulei Pilg., JF831971, JF831996, JF832021. Arthrostemma ciliatum Pav. ex D.Don, AF215562, AF215522, AF215605. Astronia smilacifolia Triana ex C.B.Clarke, AF215549, -, AF215596. “Behuria comosa R.Tavares, Baumgratz & R.Goldenb.”, JQ899111, JQ899084, JQ899060. Behuria glutinosa Cogn., JQ899112, JQ899085, JQ899061. Bellucia aequiloba Pilg., JF831972, JF831997, JF831997. Bellucia arborescens (Aubl.) Baill.*, EU711377, JQ626318, JF832035. Bellucia grossularioides (L.) Triana, EU711372, EU711385, JF832023. Bellucia mespiloides (Miq.) J.F.Macbr.*, GU968822, KF781623, -. Bellucia pentamera Naudin, AF215578, KF781624, AF215615. Bellucia spruceana (Benth. ex Triana) J.F.Macbr.*, GU968823, KF781625, -. Bertolonia margaritacea Naudin*, JQ899130, AF215512, JQ899080. Bertolonia mosenii Cogn., JF831973, JF831998, JF832024. Blakea gracilis Hemsl., JF831974, JF831999, JF832025. Blakea multiflora D.Don*, JQ899132, JQ899107, JQ899082. Blakea schlimii (Naudin) Triana, EU711373, EU711386, JF832026. Blakea trinervia L., AF215555, AF215516, AF215600. Blakea watsonii (Cogn.) Penneys & Almeda, JQ899133, JQ899108, JQ899083. Blastus borneensis Cogn. ex Boerl., AF215585, -, AF215621. Blastus cochinchinensis Lour., KX066244, KP094575, KM521849. Blastus pauciflorus (Benth.) Guillaumin, KX066245, KP095022, KM521850. Boyania colombiana Humberto Mend., -, JQ899086, JQ899062. Brasilianthus carajensis Almeda & Michelang., KX765168, KX765169, KX765170. Bredia fordii (Hance) Diels, KT354883, KT354892, KM521851. Bredia sessilifolia H.L.Li, -, KP094838, -. Calvoa grandifolia Cogn., -, AY667151, AY660632. Cambessedesia eichleri Cogn., JQ899113, JQ899087, JQ899063. Cambessedesia espora (A.St.Hil. ex Bonpl.) DC., JQ899114, JQ899088, JQ899064. Cambessedesia hilariana (A.St.Hil. ex Bonpl.) DC., JQ899115, JQ899089, JQ899065. Cambessedesia membranacea Gardner, AY553782, -, AY553775. Castratella piloselloides (Bonpl.) Naudin, AY553783, AY553779, AY553774. Centradenia inaequilateralis (Schltdl. & Cham.) G.Don, AF215563, EU711387, AF215606. Chalybea macrocarpa (Uribe) Penneys & Morales-P, JQ899121, JQ899095, JQ899071. Comolia microphylla Benth., JF831975, JF832000, JF832028. Desmoscelis villosa (Aubl.) Naudin, EU711374, EU711389, JF832029. Dichaetanthera arborea Baker, AF272800, -, AF294470. Dichaetanthera asperrima Cogn., AF215564, AF215523, AF215607. Diplectria divaricata (Willd.) O.Ktze, AF215556, AF270746, AF215601. Dissochaeta bracteata (Jack) Blume, AF289369, -, AF294471. Dolichoura spiritusanctensis Brade, JQ899116, JQ899090, JQ899066. Driessenia glanduligera Stapf, AF215586, AF270749, AF215622. Eriocnema fulva Naudin, AY553781, AY553777, AY553772. Fordiophyton brevicaule C.Chen, KT354884, KT354893, KM521852. Fordiophyton chenii S.Jin Zeng & X.Y.Zhuang, KT354886, KT354895, KM521854. Fordiophyton cordifolium C.Y.Wu ex C.Chen, KT354885, KT354894, KM521853. Fordiophyton faberi Stapf, KU208089, KU208090, KM521855. Fordiophyton huizhouense S.Jin Zeng & X.Y.Zhuang, KT354887, KT354896, KM521856. Fordiophyton peperomiifolium (Oliv.) C.Hansen, KT354888, KT354897, KM521857. Fordiophyton zhuangiae S.Jin Zeng & G.D.Tang, KX066246, -, KX037425. Graffenrieda latifolia (Naudin) Triana, AM235411, AM235644, AM235447. Graffenrieda moritziana Triana, EU055944, EU711390, JF832031. Graffenrieda rotundifolia (Bonpl.) DC., AF215576, AF215532, AF215613. Henriettea martiusii (DC.) Naudin, EU711375, EU711391, JF832032. Henriettea patrisiana DC., JF831977, JF832002, JF832033. Henriettea ramiflora (Sw.) DC., GU968811, KF781627, -. Henriettea succosa (Aubl.) DC., GU968815, KF781628, -. Henriettea tuberculosa (Donn. Sm.) L.O.Williams*, GU968816, KF781629, -. Heterocentron elegans (Schltdl.) Kuntze, AF272804, AY456135, AF325926 Heterocentron subtriplinervium (Link & Otto) A.Braun & C.D.Bouché, AF215566, AF270747, AF210374. Heterotis fruticosa (Brenan) Veranso-Libalah & G.Kadereit, AF272802, -, AF210377. Heterotis rotundifolia (Sm.) Jacq.-Fél., AF215565, U26323, AF270745. Huberia consimilis Baumgratz, JQ899117, JQ899091, JQ899067. Huberia peruviana Cogn., JQ899118, JQ899092, JQ899068. Lavoisiera cordata Cogn., AF215582, AF215540, AF210371. Lavoisiera pulchella Cham., EU711376, EU711392, JF832034. Macairea radula (Bonpl.) DC., EU711378, EU711394, JF832036. Macrocentrum cristatum (DC.) Triana, AM235412, AM235645, AM235448. Macrocentrum repens (Gleason) Wurdack, AF215551, AF215513, AF215598. Medinilla alternifolia Blume, AF289374, -, AF322229. Medinilla humbertiana H. Perrier, AF215557, AF215517, AF215602. Melastoma beccarianum Cogn., AF272805, AM235646, AM235449. Melastoma candidum D.Don, AB436365, GQ436728, AF215608. Melastoma dodecandrum Lour., AF272808, GQ436727, -. Melastoma malabathricum L., AF272810, AF270748, AB436376. Melastoma sanguineum Sims, AF270754, HQ415218, AB436378. Melastoma tetramerum Hayata, AB436364, -, AB436373. Memecylon durum Cogn., AM235408, AM235641, AM235444. Memecylon edule Roxb., AF215574, AF215528, AF215609. Meriania macrophylla (Benth.) Triana, AM235414, AM235647, AM235450. Meriania nobilis Triana, AF215577, AF215533, AF215614. Meriania phlomoides (Triana) Almeda, EU055971, EU711395, JF832037. “Merianthera bullata R.Goldenb., Fraga & A.P.Fontana”, JQ899129, JQ899104, JQ899078. Merianthera burlemarxii Wurdack, JQ899122, JQ899096, JQ899072. “Merianthera parvifolia R.Goldenb., Fraga & A.P.Fontana”, JQ899127, JQ899102, JQ899077. Merianthera pulchra Kuhlm., JQ899124, JQ899098, JQ899073. Merianthera sipolisii (Glaz. & Cogn.) Wurdack, JQ899126, JQ899100, JQ899075. “Merianthera verrucosa R.Goldenb., Fraga & A.P.Fontana”, JQ899125, JQ899099, JQ899074. Miconia bicolor (Mill.) Triana*, EU056130, KX397981, -. Miconia calycina Cogn., EU056001, JF832003, JF832038. Miconia cubatanensis Hoehne, EU056020, -, -. Miconia dodecandra (Desr.) Cogn., EU056026, EU711396, JF832039. Miconia donaeana Naudin, AM235415, AM235648, AM235451. Miconia fasciculata Gardner, EU056033, -, -. Miconia mayeta (D. Don.) Michelang.*, AF215581, AF215537, AF215618. Miconia petiolaris (Schltdl. & Cham.) Michelang., AM235410, AM235643, AM235446. Miconia rubra (Aubl.) Mabb.*, AF215579, AF215535, AF215616. Miconia secunmexicana G.Ocampo & Almeda*, AF215580, AF215536, AF215617. Miconia tococa (Desr.) Michelang.*, AM235417, AM235650, -. Miconia urbani (Cogn.) Cogn.*, AF270753, AF215538, AF215619. Monochaetum calcaratum (DC.) Triana, AF215568, AF215524, AF210372. Monolena primuliflora Hook. f., AF215552, AF215514, AF270743. Mouriri crassifolia Sagot, -, FJ038111, -. Mouriri guianensis Aubl., AF215575, AF215529, AF215610. Mouriri helleri Britton, AF322230, AF270752, AF215611. Nepsera aquatica (Aubl.) Naudin, AF215569, JQ592692, AF210373. Osbeckia chinensis L., AF215570, AF215525, AF210378. Osbeckia stellata Buch.-Ham. ex Ker Gawl., AF272818, U26330, -. Oxyspora paniculata (D.Don) DC.*,, KX527089, MH722293. Phainantha laxiflora (Triana) Gleason, JF831980, JF832006, JF832043. Phainantha shuariorum C.Ulloa & D.A.Neill, JF831981, JF832007, JF832044. Phyllagathis gymnantha Korth., AF215590, -, AF215626. Phyllagathis hispidissima (C.Chen) C.Chen, KT354889, KT354899, KM521858. Physeterostemon fiaschii R.Goldenb. & Amorim, EU711379, EU711397, JF832045. Physeterostemon jardimii R.Goldenb. & Amorim, EU711381, EU711399, JF832046. “Physeterostemon thomasii Amorim, Michelangeli & R.Goldenb.”, EU711383, EU711401, JF832047. Pternandra coerulescens Jack, AF215558, AF215518, AF322232. Pternandra echinata Jack, AF215559, AF215520, AF270744. Pternandra multiflora Cogn., AF215560, -, AF215603. Pterolepis glomerata (Rottb.) Miq., AF215571, AF215526, AF210376. Rhexia mariana L., AF272819, KJ773817, AF323723. Rhexia virginica L., AF215587, MG248427, AF215623. Rhynchanthera grandiflora (Aubl.) DC., AF215584, AF215542, AF210369. “Rupestrea johnwurdackiana (Baumgratz & D’El Rei Souza) Michelang., Almeda, & R.Goldenb.”, KM373899, KM373900, KM373901. Salpinga maranonensis Wurdack, JF831982, JF832008, JF832048. Salpinga secunda Schrank & Mart. ex DC., EU711384, EU711402, JF832049. Sarcopyramis parvifolia Merr. ex H.L.Li, -, KX527237, -. Siphanthera paludosa (DC.) Cogn., -, AY553780, AY553776. Tibouchina grossa (L.f.) Cogn., JF831983, JF832009, JF832050. Tibouchina longifolia (Vahl) Baill., AF215572, JQ592704, AF210375. Tibouchina urvilleana (DC.) Cogn., AF272820, U26339, AF322234. Tigridiopalma magnifica C.Chen, KT354891, KT354900, KM521859. Triolena amazonica (Pilg.) Wurdack, JF831984, JF832010, JF832051. Triolena paleacea (Triana) Almeda & Alvear, JF831976, JF832001, JF832030. Triolena pustulata Triana*, JQ899135, JQ899110, AF215599. Tristemma littorale Benth., -, AY667150, AY660631. Tristemma mauritianum J.F.Gmel., AF272821, -, AF322233. Tryssophyton merumense Wurdack, A. Radosavljevic 165 (US), MK284234, MK284232, -. Tryssophyton quadrifolius K.Wurdack & Michelang., K. Wurdack 5865 (US), -, MK284233, -. Warneckea membranifolia (Hook. f.) Jacq.-Fél., AF331711, KC628335, -. Wurdastom cuatrecasasii (Wurdack) B.Walln., JF831985, JF832011, -.