Phytokeys

Phytokeys

Published by Pensoft Publishers

Online ISSN: 1314-2003

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Print ISSN: 1314-2011

Disciplines: Plant Sciences

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Molecular phylogenetic tree recovered by the ML of the reduced alignment for matK, ndhF, rbcL, and PHYC presented by Davis and Anderson (2010) with the taxonomic sampling reduced to one or three terminals according to the accepted genera in the present study. Left tree shows branch lengths recovered. Right tree shows the names of families, subfamilies, and tribes coloured according to the current classification recognised in this study. Bootstrap support values from the ML presented on the nodes. Black circles represent homoplasies, and red circles represent synapomorphies/autapomorphies recovered for each lineage. Numbers above circles represent character numbers from Suppl. material 2, while the numbers below the circles represent character states from the same file. Galphimieae: flower of Galphimia gracilis Bartl. by M.O.O. Pellegrini. Acmanthereae : flower of Pterandra pyroidea L. by R.F. Almeida. Byrsonimeae: flower of Byrsonima sericea DC. by R.F. Almeida. Acridocarpeae: flower of Acridocarpus longifolius (G.Don) Hook.f. by E. Bidault. Mcvaughieae: flower of Mcvaughia sergipana Amorim & R.F.Almeida by R.F. Almeida. Barnebyeae: flower of Barnebya harleyi W.R.Anderson & B.Gates by F. Flores. Ptilochaeteae: flower of Ptilochaeta bahiensis Turcz. by R.F. Almeida. Bunchosieae: flowers of Tristellateia madagascariensis Poir. and Bunchosia glandulifera (Jacq.) Kunth by N. Rakotonirina and R.F. Almeida, respectively. Hiraeeae: flower of Hiraea restingae C.E.Anderson by R.F. Almeida. Hiptageae: flowers of Niedenzuella multiglandulosa (A.Juss.) W.R.Anderson, Hiptage benghalensis (L.) Kurz, Christianella multiglandulosa (Nied.) W.R.Anderson, Alicia anisopetala (A.Juss.) W.R.Anderson, and Callaeum psilophyllum (A.Juss.) D.M.Johnson all by R.F. Almeida. Malpighieae: flowers of Amorimia andersonii R.F.Almeida, Mascagnia cordifolia (A.Juss.) Griseb., Aspidopterys wallichii Hook.f., Triaspis macropteron Welw. ex Oliv., and Madagasikaria andersonii C.Davis by F. Michelangeli, M.O.O. Pellegrini, N. Singh, E. Bidault, and C.C. Davis, respectively. Gaudichaudieae: flowers of Bronwenia megaptera (B.Gates) W.R.Anderson & C.Davis, Diplopterys lutea (Ruiz ex Griseb.) W.R.Anderson & C.Davis, Stigmaphyllon paralias A.Juss., Peixotoa hispidula A.Juss., Camarea axillaris A.St.-Hil., and Mamedea pulchella (Griseb.) R.F.Almeida & M.Pell. by R.F. Almeida, M.O.O. Pellegrini, M.O.O. Pellegrini, R.F. Almeida, R.F. Almeida, and N. Taniguti, respectively.
Line drawings and scanning electron micrographs of common types of malpighiaceous hairs A T-shaped with a short base (i.e., foot) B T-shaped with a long base C T-shaped with a very reduced base and branches with spiked cell wall D T-shaped with a very reduced base and branches with smooth cell wall E Y-shaped with a long base and two equally long branches F Y-shaped with a long base, one long and one very reduced branch G Y-shaped with a very reduced base H T-shaped with reduced base and laterally flattened (i.e., scaly) I, J detail of a velutine indumentum comprising Y-shaped hairs K, L detail of a tomentose indumentum comprising T-shaped hairs with long bases M, N detail of a sericeous indumentum comprising T-shaped hairs with very reduced bases O detail of the spikes on the cell wall of a hair branch P detail of the rugae on the cell wall of a hair branch (all line drawings and SEMs by R.F. Almeida).
Phyllotaxis, stipules, and petioles of MalpighiaceaeA branch with opposite leaves of Bronwenia megapteraB branch with decussate leaves of Verrucularina glaucophyllaC branch with alternate leaves of Stigmaphyllon angustilobumD branch with verticillate leaves of Pterandra pyroideaE interpetiolar stipules of Mascagnia cordifoliaF epipetiolar stipules of Byrsonima intermediaG free stipules of Hiraea hatschbachiiH connate stipules of Peixotoa catarinensisI transverse section of a circular petiole J transverse section of a plane-convex petiole K transverse section of a canaliculate petiole L leaf with very reduced petiole of Byrsonima basilobaM leaf with short petiole of Banisteriopsis adenopodaN leaf with long petiole of Stigmaphyllon caatingicolaO alternate glands on the petiole of Banisteriopsis membranifoliaP opposite to alternate glands on the petiole of Schwannia mediterraneaQ subopposite glands on the petiole of Banisteriopsis membranifoliaR discoid and sessile gland of Banisteriopsis laevifoliaS cupuliform and stalked glands of Banisteriopsis adenopoda (line drawings and photographs A–C, G, I–K, L–O, Q–S by R.F. Almeida; D by C. Silva, E, F, H, P by M.O.O. Pellegrini).
Leaf blades of MalpighiaceaeA leaf with sagittate base of Stigmaphyllon ciliatumB leaf with rounded base of Banisteriopsis adenopodaC leaf with cordate base of Stigmaphyllon blanchetiiD leaf with cuneate base of Banisteriopsis vernoniifoliaE leaf with obtuse base of Stigmaphyllon paraliasF leaf with truncate base of Stigmaphyllon gayanumG leaf with oblique base of Stigmaphyllon lanceolatumH leaf with attenuate base of Acmanthera minimaI leaf with entire margin of Stigmaphyllon caatingicolaJ leaf with 3-lobed margin of Stigmaphyllon caatingicolaK leaf with 5-lobed margin of Stigmaphyllon angustilobumL leaf with crenate margin of Stigmaphyllon crenatumM leaf with ciliate margin of Stigmaphyllon ciliatumN leaf with dentate margin of Stigmaphyllon vitifoliumO leaf with plane blade margin of Banisteriopsis membranifoliaP leaf with revolute blade margin of Verrucularina glaucophyllaQ rounded leaf apex of Tetrapterys phlomoidesR mucronate leaf apex of Banisteriopsis magdalenensisS emarginate leaf apex of Hiraea cuiabensisT cuspidate leaf apex of Banisteriopsis adenopodaU acuminate leaf apex of Mamedea harleyiV acute leaf apex of Banisteriopsis membranifolia (photographs A–D, F, G, I, L, O, P, T–V by R.F. Almeida; E, M by M.O.O. Pellegrini; H by F. Farronay, Q by G.A. Dettke, N by A.C. Dal Col, R by C. Baez, S by I.L. Morais).
Inflorescence architecture of MalpighiaceaeA inflorescence evolution in Malpighiaceae according to Anderson (1981)B 1-flowered cincinnus of Niedenzuella lasiandraC Thyrse of 1-flowered cincinni of Niedenzuella lasiandra (line drawings modified from Anderson 1981; photographs A modified from Anderson 1981; B, C by M.O.O. Pellegrini).

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A new classification system and taxonomic synopsis for Malpighiaceae (Malpighiales, Rosids) based on molecular phylogenetics, morphology, palynology, and chemistry

May 2024

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343 Reads

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Thais Alves-Silva

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Aims and scope


PhytoKeys is a peer-reviewed, open-access, rapidly disseminated journal launched to accelerate research and free information exchange in taxonomy, phylogeny, biogeography and evolution of plants.

Phytokeys publishes papers in systematic botany containing taxonomic or floristic data on any taxon of any geological age from any part of the world. To respond to the current trends in linking biodiversity information and synthesising the knowledge through technology advancements, PhytoKeys also publishes papers across other taxon-based disciplines, such as ecology, molecular biology, genomics, evolutionary biology, paleontology, biodiversity informatics, and others.

Extensive floristic overviews on a group in a country or larger region are welcome. Short floristic contributions may be considered if they are based on significant or unexpected discovery. Regular floristic contributions may eventually be published in special issues devoted to a region/country.

Recent articles


Two new species of Paraphlomis (Lamiales, Lamiaceae) from limestone karsts in Guangdong Province, China
  • Article

June 2024

Wan-Yi Zhao

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Qin-Dai Xiong

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Rang-Min Wu

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Qiang Fan

Paraphlomis qingyuanensis and P. baiwanensis (Lamiaceae), two new species from the limestone area in Guangdong Province, China, are described. Morphologically, both species belong to P. ser. Subcoriaceae C.Y. Wu & H.W. Li. A close relationship between the two new and P. subcoriacea was revealed by molecular phylogenetic analyses based on ETS and ITS. Further morphological and population genetic evidence indicated that they are distinct species in Paraphlomis . According to the IUCN Red List Categories and Criteria, P. qingyuanensis and P. baiwanensis were assessed as Endangered (EN) and Deficient (DD), respectively.

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Eriotheca paganuccii (Bombacoideae, Malvaceae), a new endangered species from montane forests in the Atlantic Forest of Bahia, northeastern Brazil

June 2024

Jefferson Carvalho-Sobrinho

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Aline C. da Mota

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Laurence J. Dorr

A new species of Eriotheca (Bombacoideae, Malvaceae) from montane wet forests in the Atlantic Forest of Bahia, northeastern Brazil, is described and illustrated. It is known from only three populations situated between 750 m and 850 m in elevation on mountain summits and categorized as Endangered (EN) based on IUCN criteria. Eriotheca paganuccii is distinct from all congeners by the combination of coriaceous to strongly coriaceous leaves and remarkable few-seeded, globose to subglobose woody capsules that contain scanty kapok and the largest seeds known in the genus to date. The affinities of E. paganuccii to morphologically similar species as well as the importance of obtaining phenologically complete collections are discussed.


Figure 1. Plants and habitat of Astragalus liuaiminii Z. Z. Yang and Q. R. Liu A Mr. Aimin Liu and the distribution area B habitat C plant D side view of the plant.
Figure 3. Astragalus liuaiminii sp. nov. A plants B standard C keel D wings E leaf, adaxial view F mature saccate calyx G half calyx removed exposing immature fruit H pistils and stamens.
Astragalus liuaiminii, a new species of Astragalus (Fabaceae) from Xinjiang, China
  • Article
  • Full-text available

June 2024

A new species, Astragalus liuaiminii Z. Z. Yang & Q. R. Liu (Fabaceae), is described and illustrated from Xinjiang Province, China. The new species is close to A. wenquanensis S. B. Ho, but differs from the latter by leaves having a single leaflet (vs. 3–5 leaflets), and inflorescences with 1–2 flowers (vs. inflorescences with 5–7 flowers). It is also similar to A. monophyllus Maxim in leaf shape, but differs by its calyx expanding to become saccate and totally enveloping the pod (vs. calyx tubular, and ruptured by pod after flowering).


Figure 1. Bayesian 50% majority-rule consensus tree of the combined dataset of the Caucasian Campanula species based on three plastid markers (trnK/matK, petD and rpl16). Values above nodes indicate posterior probabilities (bold) and maximum likelihood bootstrap support (italic), values below nodes indicate maximum parsimony jackknife support, values in square brackets indicate conflicting topologies. Sample designations include the taxon name, DNA lab code, and ISO (international organization for standardization) country code, in case of the Russian part of the Caucasus also the TDWG (Biodiversity Information Standards) code of the territory (n.d. -sample not documented); for DNA lab codes and ISO codes of collapsed terminals with multiple accessions, see Suppl. materials 1, 2. Abbreviations in species names: A. -Adenophora, As. -Asyneuma, Az. -Azorina, C. -Campanula, G. -Githopsis, M. -Michauxia, Mu. -Musschia, P. -Petromarula, Ph. -Phyteuma. Sample designations in bold (in black) indicate species from the core Scapiflorae clade, sample designations in blue indicate Scapiflorae group members phylogenetically distant from the core clade. Subdivision of the Scapiflorae clade by numbered terminal clades (1-17) and geographical distribution of the lineages.
The polyphyletic Caucasus-centred Campanula subg. Scapiflorae (Campanulaceae) revisited with a newly circumscribed C. sect. Tridentatae for its core clade

June 2024

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20 Reads

Campanula L. is among the genera with the highest number of endemics in the Caucasus ecoregion. A group of attractive alpine and subalpine perennial rosette plants with short single-flowered stems centred in the Caucasus has been treated as Campanula subg. Scapiflorae or at other ranks, with considerably varying circumscription and classification. Molecular phylogenetic analysis of three plastid DNA regions (trnK/matK, petD, rpl16) of a strongly extended sampling, comprising 23 of the 27 commonly accepted taxa (85%) with 330 accessions built on and guided by the results of our previous study of the group, confirmed the polyphyly of C. subg. Scapiflorae in any of its circumscriptions. The core clade of the group comprises exclusively endemics and near-endemics of the Caucasus and is treated here as C. sect. Tridentatae in a revised circumscription. The phylogenetic relationships of the disparate other elements of the Scapiflorae group are outlined.


Molecular and morphological evidence supports the resurrection of Chrysosplenium guangxiense H.G.Ye & Gui C.Zhang (Saxifragaceae)

June 2024

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3 Reads

Chrysosplenium guangxiense H.G.Ye & Gui C.Zhang was first described as a new species in 1994 but later synonymized in the Flora of China treatment with C. glossophyllum H.Hara. Plastid genomes and nrDNA sequences were used to infer the phylogenetic relationships of selected taxa in Chrysosplenium . Our phylogenetic analyses revealed that C. guangxiense belongs to sect. Alternifolia, is closely related to Chrysosplenium hydrocotylifolium H.Lév. & Vaniot but distant from C. glossophyllum . Morphologically, C. guangxiense could be easily distinguished from C. glossophyllum by having robust rhizomes, basal leaves with a long cuneate base and fewer teeth in the margin, curled sepal margins, and red, larger seeds. It could also be easily distinguished from C. hydrocotylifolium by possessing long elliptic leaves and a long cuneate leaf base. Along with the phylogenetic studies, the complete plastid genome of C. guangxiense was also reported. The plastid genome was 154,004 bp in length and comprised two inverted repeats (IRs) of 28,120 bp, separated by a large single-copy of 80,646 bp and a small single-copy of 17,118 bp. A total of 111 functional genes were discovered, comprising 78 protein-coding genes, 29 tRNA genes, and four rRNA genes. Based on assessment of morphological and molecular data Chrysosplenium guangxiense H.G.Ye & Gui C.Zhang is resurrected from C. glossophyllum H.Hara at species level. A global conservation assessment classifies C. guangxiense as Vulnerable (VU).


Figure 5. The distribution of Cyrtomium adenotrichum (red circle) in Guangxi, China.
Main morphological differences amongst Cyrtomium adenotrichum and C. nephrolepioides, C. obliquum, C. sin- ningense and C. calcis.
Cyrtomium adenotrichum (Dryopteridaceae), a new species from Guangxi, China

Cyrtomium adenotrichum Y. Nong & R.H. Jiang (Dryopteridaceae), a new species from Guangxi, China, is described and illustrated. This new species is similar to C. nephrolepioides (Christ) Copel., C. obliquum Ching & K. H. Shing ex K. H. Shing, C. sinningense Ching & K. H. Shing ex K. H. Shing and C. calcis Liang Zhang, N.T.Lu & Li Bing Zhang in having erect rhizomes, dense, leathery lamina and rounded sori, but it can be easily distinguishable by its stipe sparsely glandular, base obvious oblique, basiscopic base truncate, acroscopic base auriculate or ovate.


Figure 1. Type specimens of Lasiostoma bredemeyeri Schult. & Schult.f A lectotype conserved in the Jacquin herbarium at W (Bredemeyer s.n.; W barcode W0078191) B, C isolectotype conserved in the Willdenow herbarium at B (Bredemeyer s.n.; B barcodes BW02865000 and BW02865010, respectively).
Figure 2. A-C sheet deposited in the type photograph collection of the New York Botanical Garden (Negative No. 1163; sheet without barcode or access number and not available online) containing a photograph of the isolectotype of Lasiostoma bredemeyeri Schult. & Schult.f. conserved in the Willdenow herbarium at B (Bredemeyer s.n.; Fig. 1B,C) and two overlapping notes made by Krukoff A overview; note the detail of the original label containing the description "Lasiostoma glabra, corollis fauce glabris" (at center) and Krukoff's identification tag made in 1975 (at right) B detail of the uppermost typewritten note made by Krukoff C detail of the basalmost handwritten note made by Krukoff (all images made by R. B. Setubal).
Figure 3. Comparison of labels on four specimens of Rouhamon pedunculatum A.DC A lectotype showing original label "Schomburgk 482" (G barcode G00132188) B-D specimens of R. pedunculatum without original labels and containing alternative numbers B specimen with the alternative number "792" (F Catalog No. 620082; not available online; image from R. B. Setubal) C, D specimens with the numbers "482" and "792.B" written juxtaposed (K barcodes K000573484 and K000573485, respectively).
Figure 4. Comparison of labels on four type specimens of Strychnos trinitensis Griseb A lectotype (H. Crueger s.n., GOET barcode GOET005464) B-D isolectotypes B K barcode K000573430 C TRIN Catalog No. 258 D TRIN Catalog No. 1529.
Clarifying the nomenclature of Strychnos bredemeyeri and Lasiostoma (Loganiaceae)

June 2024

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14 Reads

Strychnos (Loganiaceae, Gentianales) is a large and pantropical genus of woody plants, ethnobotanically important as a source of many toxic alkaloids, including strychnine. Unfortunately, the status of numerous names at various ranks of Strychnos remains unresolved, including that of many specific or infraspecific taxa in the Neotropics. In this study, we address Strychnos bredemeyeri (basionym Lasiostoma bredemeyeri ), a species described in 1827 based on type material collected in Venezuela during the poorly documented Austrian Märter expedition (1783–1788). Strychnos bredemeyeri is an unarmed liana with solitary tendrils and axillary inflorescences that occurs in Neotropical rainforests and savannas in Brazil, Guyana, Trinidad and Tobago, and Venezuela. We clarify here the nomenclatural status of Lasiostoma Schreb., an illegitimate and superfluous genus currently in synonymy under Strychnos , and its former species Lasiostoma bredemeyeri [= Strychnos bredemeyeri ]. Also, we lectotypify S. pedunculata and S. trinitensis , both taxa currently synonyms of S. bredemeyeri .


Table 3 .
Accepted species, subspecies, varieties, and synonyms within Erica across the four major global checklists.
Curating an online checklist for Erica L. (Ericaceae): contributing to and supporting global conservation through the World Flora Online

June 2024

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12 Reads

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1 Citation

To support the work of the Global Conservation Consortium for Erica and update the Erica checklist in the World Flora Online (WFO), we have curated the taxonomic backbone in the WFO by expanding it to include updated nomenclatural information from the International Plant Name Index, missing names present in the World Checklist of Vascular Plants (WCVP), the Botanical Database of Southern Africa (BODATSA), and from the “International register of heather names” database, a data source not readily available online. The result is the most robust database of Erica names to date, including 851 species, 111 subspecies, 244 varieties, and 2787 synonyms, which is a reliable reference for initiatives such as the Erica identification aid, conservation prioritisation, and gap analyses. We disambiguate common orthographic variants within the database and present an overview of these. We also comment on the correct orthography of E. heleophila Guthrie & Bolus and E. michellensis Dulfer and the validity of E. tegetiformis E.G.H.Oliv. are discussed, and the use of E. adunca Benth. for a South African species rather than E. triceps Link, which is here regarded as insufficiently known and of uncertain application, is clarified.


Lappula effusa D.H.Liu & W.J.Li, sp. nov. A habitat B habit C, D flower morphology E inflorescences F fruit G spreading calyx in fruit.
Type specimens of L. himalayensis, L. tadshikorum and L. effusaA holotype of L. himalayensis (PE00029615!) B lectotype of L. tadshikorum (LE 140!) C holotype of L. effusa (XJBI00135936!).
Morphological comparisons of L. himalayensis, L. effusa and L. tadshikorum. L. himalayensisA stem indumentum B gynobase C fruit lateral view D fruit polar view E nutlet abaxial view (with short glochids) F nutlet lateral view (with long glochids) G nutlet lateral view (with short glochids) H nutlet adaxial view. L. effusaI stem indumentum J gynobase K fruit lateral view L fruit polar view M nutlet abaxial view (with short glochids) N nutlet abaxial view (with long glochids) O nutlet lateral view P nutlet adaxial view. L. tadshikorumQ stem indumentum R gynobase S fruit lateral view T fruit polar view U nutlet abaxial view (with short glochids) V nutlet abaxial view (with long glochids) W nutlet lateral view X nutlet adaxial view. Scale bar represents 0.5 mm.
Morphological comparisons of L. effusa L. himalayensis and L. tadshikorum.
Lappula effusa (Boraginaceae), a new species from Xinjiang, China

June 2024

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7 Reads

Lappula effusa D.H.Liu & W.J.Li, a new species of Boraginaceae from Xinjiang, China, is described and illustrated in this study. The new species is morphologically similar to Lappula himalayensis and L. tadshikorum. However, it can be distinguished from the compared species by several characteristics, such as: stem single, erect, frequently branched at middle and above, densely spreading hispid, hairs discoid at base; corolla white or blue; fruit compressed, heteromorphic nutlets with two rows of marginal glochids, nutlets acute ovoid, disc narrowly ovate-triangular. The diagnosis of the new species is supported with comprehensive investigation including photographs, detailed description, notes on etymology, distribution and habitat, conservation status, as well as comparisons with morphologically similar species.


Impatiens karenensis Chit Soe Paing & Ruchis A habit B flower, front view C flower, side view D inner lateral sepals E outer lateral sepals F–H lower sepal I–J dorsal petal K lateral united petals L ovary, pedicel and bract M fruit (from Chit Soe Paing 002). Drawn by S. Ruchisansakun.
Impatiens karenensis Chit Soe Paing & Ruchis. in vivoA flower, front view B flowers, side view C habit. Photographed by Chit Soe Paing.
The distribution of Impatiens karenensis Chit Soe Paing & Ruchis. (SimpleMappr, Shorthouse 2010).
Impatiens karenensis (Balsaminaceae), a new tiny flowered species from Myanmar

June 2024

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42 Reads

Impatiens karenensis (I. sect. Semeiocardium) from Kayin State, Myanmar is described and illustrated here. It is most similar to I. micromeris, but differs in having lower petals with outer margins strongly undulate in the lower half (vs. lower petals entire), apex of upper petals acute to obtuse (vs. apex rounded), short stout spur, ± as long as the depth of lower sepal, ca. 2.5 mm long (vs. long attenuate spur, twice as long as the depth of lower sepal, ca. 5 mm long). Its conservation status is also assessed as Critically Endangered.


Revisiting the infrageneric classification of Garcinia L. (Clusiaceae): an updated sectional name and new, legitimate names in Garcinia for Allanblackia gabonensis (Pellegr.) Bamps and A. parviflora A.Chev.

June 2024

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6 Reads

In a recent publication dealing with the sectional-level taxonomy of Garcinia, an illegitimate superfluous sectional name and two illegitimate homonyms were published. Herein we choose a legitimate sectional name, Garcinia section Rheediopsis Pierre, for the superfluous name Garcinia section Rheedia (L.) S.W.Jones ex P.W.Sweeney; and create two new legitimate names in Garcinia for Allanblackia gabonensis (Pellegr.) Bamps and A. parviflora A.Chev.


Morphological and habitat diversity in the family Plumbaginaceae. Limonieae: AAcantholimon pterostegium Bunge BArmeria pungens (Brot.) Hoffmanns. & Link CBakerolimon plumosum (Phil.) Lincz. DCeratolimon feei (Girard) M.B.Crespo & Lledó ECeratolimon weygandiorum (Maire & Wilczek) M.B.Crespo & Lledó FLimoniastrum monopetalum (L.) Boiss. GLimonium bonduellei (T.Lestib.) Kuntze HLimonium virgatum (Willd.) Fourr. Photos A by Hossein Akhani B, F, G by Mario Martínez-Azorín C by Sergio Ibáñez D, E by José Quiles H by Konstantina Koutroumpa.
Morphological and habitat diversity in the family Plumbaginaceae. Limonieae: ALimonium thymoides (Girard) M.B.Crespo BPsylliostachys leptostachya (Boiss.) Roshkova CPsylliostachys spicata (Willd.) Nevski DSaharanthus ifniensis (Caball.) M.B.Crespo & Lledó. Plumbagineae: EDyerophytum africanum (Lam.) Kuntze FPlumbago auriculata Lam. GPlumbago europaea L. Photos A, E, G by Mario Martínez-Azorín B, C, F by Hossein Akhani D by José Quiles.
Topological incongruence in the sister relationships of the three tribes in Plumbaginaceae, using Polygonaceae as outgroup A plastid cpDNA tree (rbcL, trnL-F, matK; Lledó et al. 2001, 2005a; Koutroumpa et al. 2018) B nrDNA tree, ITS (see Suppl. material 2), and 353 low copy nuclear loci (Baker et al. 2022).
Phylogenetic relationships of the major clades in Limonium and the corresponding infrageneric units, following Koutroumpa et al. (2018). The size of triangles is proportional to the number of species assigned to the different sections and the “Mediterranean lineage”.
A taxonomic backbone for the Plumbaginaceae (Caryophyllales)

June 2024

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123 Reads

A taxonomic backbone of the Plumbaginaceae is presented and the current state of knowledge on phylogenetic relationships and taxon limits is reviewed as a basis for the accepted taxon concepts. In total, 4,476 scientific names and designations are treated of which 30 are not in the family Plumbaginaceae. The Plumbaginaceae are subdivided in three tribes with 26 genera and 1,179 accepted species. Two subgenera, 17 sections, two subsections and 187 infraspecific taxa are accepted. At the species and infraspecific level 2,782 synonyms were assigned to accepted taxa, whereas 194 names were excluded from the core checklist (i.e., unplaced taxa, infrageneric subdivisions with still uncertain application, names of verified uncertain application, invalid horticultural names, excluded names from other families, other excluded designations, and unresolved names). The EDIT Platform for Cybertaxonomy was utilized as the tool to compile and manage the names and further taxonomic data under explicit taxon concepts. Secundum references are given in case taxon concepts were taken from the literature, whereas this study serves as reference for newly circumscribed taxa. The family’s division into the tribes Aegialitideae, Limonieae, and Plumbagineae departs from earlier two-subfamily classifications, prompted by recent phylogenetic findings that challenge the subfamilial affinity of Aegialitis. The genus Acantholimon was extended to include Gladiolimon, as currently available phylogenetic and morphological data support this merger. In Limonium, all accepted species could be assigned to sections and subsections or the “Mediterranean lineage”, respectively, making use of the phylogenetic distribution of their morphological characters and states. A new combination and/or status is proposed for Dyerophytum socotranum, Limonium thymoides, Limonium × fraternum, Limonium × rossmaessleri, and Limonium sect. Jovibarba. Special attention is given to nomenclatural issues, particularly for Statice nomen ambiguum to resolve the names under accepted names. The use of artificial groupings like “aggregates”, “complexes” and “species groups” in alpha-taxonomic treatments is discussed. The taxonomic backbone will receive continued updates and through the Caryophyllales Taxonomic Expert Network, it contributes the treatment of the Plumbaginaceae for the World Flora Online.


Carex recondita Muñoz-Schüler, Martín-Bravo & Jim.Mejías A, B aerial and frontal view of the plant and a spike C, D habit E two of the authors looking for more specimens of C. recondita on its habitat, the road can be seen at the upper-left portion of the picture F habitat of C. recondita, showing dominant Azorella ruizii (green cushions) and Anarthrophyllum cumingii (prostrate shrub with orange-yellow flowers). Photos by P M-S (A–E) and MTKA (F).
Detailed comparison between C. recondita and C. austroamericanaA spikes of C. recondita (Aa) and C. austroamericana (Ab) B utricles of C. recondita (Ba) and C. austroamericana (Bb) C glumes of C. reconditaD rachillas of C. recondita (Da) and C. austroamericana (Db). Note that the rachilla of C. recondita is attached to the utricle base E achenes of C. recondita (Ea) and C. austroamericana (Eb).
Phylogram resulting from BI analysis using MrBayes of the multiaccession matrix for Carex subg. Psyllophorae. Branch support is not indicated when PP=1.00/BS = 100. Posterior probability (PP) support is given above branches while MLBS support is given below branches. Asterisks above branches indicate BI 100 > PP ≥ 0.95. Bold thick branches indicate nodes supported by MLBS ≥ 80 and/or BIPP ≥ 0.95. ML support BS ≤ 80 is given below branches. The newly sequenced voucher of C. recondita is displayed in bold red letters. Tip labels include the geographical origin of the specimen using TDWG level 3 regions abbreviations (“botanical countries”; Brummitt 2001).
Occurrence map of Carex sect. Junciformes species cited in this work A occurrence map of five Patagonian species of Carex sect. Junciformes. Species locations are depicted in colored circles and some cities are represented as black squares B detailed map of the area of occurrence of C. recondita (light-green circles). Another species occurring in the area, C. argentina, is depicted by red circles. The highlighted region corresponds to the Yerba Loca Nature Sanctuary. Altitude was represented by a color palette and contour lines, representing elevation every 200 meters. For orientation purposes, some landmarks were depicted on the map.
Carex recondita Muñoz-Schüler, Martín-Bravo & Jim.Mejías (Carex section Junciformes Kük., Cyperaceae), a new sedge species from the Andes of central Chile

June 2024

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67 Reads

Carex section Junciformes is one of the most diverse groups of the genus in South America, consisting of approximately 30 species. Here we describe a new species, Carex recondita, belonging to this section. We studied its placement within a molecular phylogeny of the group and found it to constitute an independent lineage. The new species is morphologically very close to C. austroamericana, from southern Patagonia, despite being phylogenetically divergent to the rest of Patagonian species of sect. Junciformes. So far, this species is known only from a few specimens recently collected in its type locality, despite growing in a well-collected area in the Andes of Metropolitana Region of Santiago, the most populated administrative region of Chile. We provide a detailed morphological description, comments on its relationship with other Southern Cone species of sect. Junciformes and relevant ecological notes.


Lectotype of Linaria bimaculata (Cout.) Farminhão & Carapeto (Palhinha & Mendes s.n., LISU33258).
Overview of Linaria bimaculata (Cout.) Farminhão & Carapeto A habit (ascending form) and habitat B underside of flower with visible stripes C habit (erect form) D flowers with erect-patent corolla tube, immaculate palate and stripes visible on the underside of flower bud E flower with reticulate palate F seeds (A. Moller s.n., COI), scale bar 1 mm. Photos by M. Hansch (A), D. Frade (B, E), V. Dvořák (C), J. Neiva (D) and A. Coelho (F).
Distribution of Linaria algarviana Chav., Linaria bimaculata (Cout.) Farminhão & Carapeto, Linaria spartea (L.) Chaz. and Linaria viscosa (L.) Chaz. in the Algarve, in southwesternmost Iberia. Plain symbols indicate observation records and dotted symbols indicate herbarium specimens.
Overview of Linaria algarvianaA habit B dark purple morph (Aljezur) typical of the westernmost populations C light purple morph (Loulé) D flower with erect-patent corolla tube (Loulé) E pink morph F bicolorous morph (Portimão) G yellow morph (Portimão). Photos by V. Achterberg (A, B), J. Neiva (C), J.T. Tavares (D), A.J. Pereira (E) and S. Lobo Dias (F, G).
A new combination for a neglected member of Linaria subsect. Versicolores (Plantaginaceae, Antirrhineae) endemic to the Algarve, Portugal

June 2024

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17 Reads

Linaria bimaculatacomb. et stat. nov., from the overlooked Central Algarve plant endemism centre, is here lectotypified and redescribed as a full species based on Linaria viscosa var. bimaculata, which was historically misidentified under allopatric L. spartea and L. viscosa. Traditional herbarium taxonomy and citizen science observations were combined to document the geographical range of the four species of Linaria subsect. Versicolores in the Algarve and amend an identification key for the Iberian clade of this subsection. Geographical patterns and morphological similarity suggest a sister relationship between L. bimaculata and L. algarviana, unveiling a new possible example of parallel speciation linked to a purple to yellow shift in corolla colour. Besides the yellow flowers, L. bimaculata differs from L. algarviana in the more elongate fertile stems and the invariably erect-patent corolla tube. It is assessed as Vulnerable (VU) according to the IUCN Categories and Criteria.


RADseq phylogeny of Hawaiian Myrsine based on the min16 dataset. Samples with newly-generated sequence data are highlighted in bold. Symbols at branches represent bootstrap support (bs) values (*: maximum bs; +: bs of 90 or higher; °: bs of 70 or higher; -: bs < 70).
Myrsine cirrhosa Lorence & K.R.Wood A habit, fruiting branch B leaf showing cirrhose apex and detail of intramarginal venation C inflorescence in bud D inflorescence, flowers at anthesis E flower at anthesis, view from apex F flower at anthesis, view from base G pistil H mature drupe. A, B drawn from Wood et al. 835 (PTBG), C drawn from Wood & Query 12824 (PTBG), D–G drawn from Wood et al.18139 (PTBG), H drawn from Perlman & Wood 12747 (PTBG). Illustration by Robin Jess.
Myrsine cirrhosaA habit showing leaves with characteristic undulate margins and cirrhose apex (from Kamo‘oloa headwater below Kapalaoa Kaua‘i, Wood & Query 12824) B twig with leaves and flowers (from Wai‘ahi, Kaua‘i Wood 18139) C open exposed wind-swept summit ridges of Wai‘ale‘ale, Kaua‘i representing the habitat for Myrsine cirrhosa. All photos by K.R. Wood.
Distribution map with dots representing known locations for three Myrsine species on Kaua‘i, Hawaiian Islands.
Myrsine cirrhosa (Primulaceae), a distinctive new shrub species from Kaua‘i, Hawaiian Islands

June 2024

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23 Reads

Myrsine cirrhosa Lorence & K.R.Wood (Primulaceae), a new single-island endemic shrub species from Kaua‘i, Hawaiian Islands, is described and illustrated. Notes on its distribution, ecology and conservation status are included. The new species is known from an area with ca. 45 individuals, where it is restricted to the remote central windward region of Kaua‘i in open bogs and along open windy ridges. Suggested IUCN Red List status is CR (Critically Endangered). It differs from its Kaua‘i congeners by its longer petals and narrowly elliptic leaves with strongly undulate margins and tendril-like apex. Phylogenetic analysis using RADseq data supports the recognition of this new species.


Rhododendron leishanicum and R. oligocarpumA isotype of R. leishanicumB holotype of R. oligocarpumC, D paratype from Leigong Mountain and Maoer Mountain of R. oligocarpum, respectively.
Comparison of Rhododendron leishanicum and R. oligocarpumA, D, G, J, MR. leishanicum from Leigong mountain B, E, H, K, NR. oligocarpum from Fanjing mountain C, F, I, L, OR. oligocarpum from Maoer mountain A–C branchlets with flowers D–F adaxial and abaxial surface of leaf G–I corolla with deep purple basal spots J–L pedicel, Calyx, ovary and style indumentum M–O stamens side view.
A new synonym of Rhododendron leishanicum (subg. Hymenanthes) from China (Ericaceae)

June 2024

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2 Reads

Based on a critical examination of type specimens, images of living plants, and the literature has shown Rhododendron oligocarpum to be conspecific with R. leishanicum. Although slight variations in corolla colour exist amongst different populations of R. oligocarpum, it does not serve as a key distinguishing trait. Therefore, we reduced R. oligocarpum to a synonym of R. leishanicum, and recommend placing it in Subsection Maculifera.


Petrocodon liboensisA habitat B, C flowering plant D adaxial and abaxial surfaces of leaf blade E cymes, and bracts (inset) F flower in front view G flower in side view H, I opened corolla J stamens, and anthers (inset) K pistil, and cross section of ovary (inset) L capsules, and seeds (inset) (Photographs by Sheng-Hu Tang).
Petrocodon luteoflorusA habitat B flowering plant C adaxial and abaxial surfaces of leaf blade D cymes, and bracts (inset) E flower in front view F flower in side view G calyx segments H opened corolla I stamens J pistil, and cross section of ovary (inset) K capsules L seeds (Photographs by Sheng-Hu Tang).
Petrocodon liboensis (Gesneriaceae), a new species from Guizhou, China

June 2024

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11 Reads

Petrocodon liboensis Sheng H.Tang & Jia W.Yang is a new species of Gesneriaceae from Guizhou, southwestern China. The new taxon has a pale-yellow corolla and is most similar to P. luteoflorus. However, it differs from the latter by having a urceolate (vs. cannulate) corolla tube, an abaxial corolla lip 0.8–1.1 mm (vs. 2–2.2 mm) long, and filaments 1.5–1.7 mm (vs. ca. 7 mm) long that are straight (vs. S-shaped or geniculate near the middle). The new taxon is assessed as “Data Deficient” (DD) according to the IUCN standards.


Distribution map of Amalophyllon in Ecuador. Localities with yellow circles =Amalophyllon clarkii Boggan & L.E.Skog, green circles =A. divaricatum (Poepp.) Boggan, L.E.Skog & Roalson, and red circles =A. miraculum J.L.Clark.
Amalophyllon miraculum J.L.Clark A abaxial view of leaf B front view of flower C lateral view of flower D pendent habit featuring rosette of leaves E adaxial view of leaf. (A, E from J.L. Clark et al. 16805; B, C, D from J.L. Clark et al. 16634). Photos by J.L. Clark.
Amalophyllon clarkii Boggan & L.E.Skog A abaxial view of leaf B front view of flower C lateral view of flower D erect habit featuring evenly spaced and rosette-forming leaves E adaxial view of leaf. (A–E from J.L. Clark 13101). Photos by J.L. Clark.
Amalophyllon miraculum (Gesneriaceae), an exceptionally small lithophilous new species from the western Andean slopes of Ecuador

June 2024

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68 Reads

Recent exploratory field expeditions to the western slopes of the Ecuadorian Andes resulted in the discovery of a new species of Amalophyllon (Gesneriaceae). Amalophyllon miraculum J.L.Clark, sp. nov. is described from two localities in the Centinela region in the Santo Domingo de los Tsáchilas province. The new species is differentiated from congeners by the pendent habit, basal rosette of leaves, leaf blades with deeply serrate margins, and miniature size. Based on IUCN guidelines, a preliminary conservation status is assigned as Critically Endangered (CR).


Paepalanthus magnusA habit B leaf apex C spathe, detail of the opening D capitula in dorsal (left) and ventral (right) view E involucral bract F floral bract G staminate flower in lateral view H staminate flower with sectioned corolla, exposing the stamens and pistillodes I pistillate flower in lateral view J pistillate flower with petals and sepals distended, exposing the gynoecium K seed with numerous appendices along the periclinal walls. Illustration by Klei Souza based on the holotype (P.M. Gonella et al. 3402).
Paepalanthus magnusA habitat at Pico do Pinhão, with the Pico da Bela Adormecida (Pico do Padre Ângelo) in the background B habit among grasses and quartzitic rocks C rosette in detail D leaves, showing ciliate margin and striate abaxial surface E the base of the leaves, showing the adaxial surface and a scape enclosed by a spathe emerging from a leaf axil F spathe opening G capitulum in posterior view evidencing the involucral bracts H capitulum, lateral view I capitulum, frontal view. A by Lucian Medeiros B–I by PMG.
Paepalanthus magnus. SEM micrograph of the seed coat (from the holotype, P.M. Gonella et al. 3402).
Distribution maps A distribution map of the new species and compared taxa cited in the text B distribution map of P. magnus at Serra do Padre Ângelo region, with other landmarks of the region indicated.
Conservation threats to Paepalanthus magnusA 35-year fire record (1985–2020) in the region of Serra do Pinhão, part of Serra do Padre Ângelo B fire record in the year 2020 C land use of the region. Data on fire and land use from MapBiomas (2024). A, B Map data ©2024 Google.
A missing piece is found: a new species of Paepalanthus (Poales, Eriocaulaceae) and the puzzling relations of the campos rupestres mountaintop floras of eastern Minas Gerais, Brazil

June 2024

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82 Reads

Paepalanthus is a diverse genus characteristic of the campos rupestres, a megadiverse vegetation found on mountaintops of mainly quartzitic mountain ranges of central-eastern Brazil. Recent efforts on prospecting the biodiversity of Serra do Padre Ângelo, a small mountain complex in eastern Minas Gerais, yielded several new plant and animal species, highlighting the urgency of conservation actions towards this still unprotected area. Here, we describe yet another new species found in the campos rupestres of these mountains, Paepalanthus magnus, a mountaintop microendemic species morphologically similar to taxa found in the Espinhaço Range, over 200 km distant, a biogeographic pattern shared by several other species. The affinities of the new species are discussed, and we provide illustrations, photographs, and SEM photomicrographs of the seed. We also discuss the conservation status of the species, which is preliminarily assessed as Critically Endangered, reinforcing the urgent need to address the conservation of the unique biodiversity of Serra do Padre Ângelo.


Map visualizing the only known occurrences of Crotalaria menglaensis S.A.Rather in Mengpeng village of Xishuangbanna Dai Autonomous Prefecture, Yunnan, China (red dot).
Phylogenetic hypothesis of the genus Crotalaria based on the concatenated matrix including matK and nrITS sequences constructed via maximum likelihood as implemented in IQ Tree. Bootstrap values are printed above the branches. Since Bayesian analyses resulted in almost the same topology, only the ML tree made is presented here. The new species Crotalaria menglaensis S.A.Rather was marked in red. The names on the right side of the phylogeny correspond to the infrageneric classification of the genus Crotalaria by Le Roux et al. (2013).
Scatter plot visualizing Dim1 and Dim2 from the principal component analyses based on the assembled morphological trait variables and accessions of the three species nested in the Incanae clade (see Fig. 2), namely, C. incana L., C. bracteata Roxb. ex DC. and the new species Crotalaria menglaensis S.A.Rather. Dim1 explained 44.3% of the variation, whereas DIM2 explained 40.2%. The vectors corresponded to KL = keel length, SW - standard width, SEW - seed width, and SEL - seed length.
Crotalaria menglaensis S.A.Rather A habit B plant twigs with leaves and flowers C inflorescence with flowers D inflorescences with flowers and fruits E flower in dorsal, lateral, and ventral views F calyx showing the dorsal and ventral surfaces G standard adaxial surface H standard abaxial surface with paired planar callosity pairs at the base with white silky pubescence I wing petals with prominent cavae and a distinct claw J adaxial and abaxial surfaces of keel petals, beak not twisted, pubescence along the margins from the middle to the base of the keel petal K anthers monodelphous, 10 dimorphic anthers (common to all the species within the genus) L gynoecium showing the ovary, style, and stigma M pod in ventral, dorsal, and lateral views N pod splitted longitudinally with young seeds.
Variance in the contributions of morphological trait variables as determined by principal component analysis.
Molecular, morphological, and morphometric evidence reveal a new, critically endangered rattlepod (Crotalaria, Fabaceae/Leguminosae, Papilionoideae) from tropical China

June 2024

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23 Reads

Here, we describe a new species of Crotalaria L. discovered in Mengla County, Xishuangbanna Dai Autonomous Prefecture, Yunnan, China. The new species, Crotalaria menglaensis S.A.Rather, was confirmed by identifying diagnostic morphological characteristics, performing principal component analyses of phenotypic traits, and phylogenetic analyses based on nuclear ITS and plastid matK sequences. Phylogenetic analyses recovered the two accessions of the new species to be sister to C. bracteata Roxb. ex DC. In turn, these two species formed the sister clade to the two accessions of C. incana L. The morphometric analyses revealed that all three species were distinct, while the analyses of distinctive characters enabled unambiguous distinction of the new species by its growth habit, leaflets, flower structure and pod morphology. In contrast to the two related species, the new species is currently known only from ca. 100 mature individuals. Thus, this species is considered to be critically endangered.


Impatiens yingjingensis (Balsaminaceae), a new species from Sichuan, China

June 2024

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42 Reads

This study describes Impatiens yingjingensis X.Q. Song, B.N. Song & Biao Yang, sp. nov., a new species collected from the Yingjing area of the Giant Panda National Park. This new species is distributed at an altitude of 1400–2100 m, with a plant height of 30–130 cm. The flowers are purple-red or light purple red, with 3–9 flowers on each inflorescence and the dorsal auricle of the lateral united petals is thread-like and about 2 cm long, differing significantly from other species of Impatiens. Furthermore, molecular data, as well as micro-morphological evidence under SEM (of pollens), also support the establishment of the new species.



Alseodaphnopsis maguanensis is conspecific with A. hokouensis (Lauraceae) based on morphological and molecular evidence

June 2024

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5 Reads

Based on both morphological and molecular evidence, it is confirmed that Alseodaphnopsis maguanensis is conspecific with A. hokouensis. Hence, Alseodaphnopsis maguanensis is treated as a synonym of A. hokouensis here. The conservation status of Alseodaphnopsis hokouensis is also re-evaluated according to the IUCN Red List Categories and Criteria in this study.


Seven new species of Rinorea (Violaceae) from the Neotropics

June 2024

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15 Reads

Over the course of revising the genus Rinorea (Violaceae) from Colombia, field observations and herbarium studies revealed seven new species. Several of the new species described here belong to species complexes that required examination of herbarium material from across the Neotropics. Each of the new species described here have oppositely arranged leaves and belong to Rinorea sect. Pubiflorae, a section restricted to the Neotropics. Two new species are segregated from the R. ovalifolia species complex: Rinorea chiribiquetensis from Chiribiquete National Park in the Colombian Amazon and Rinorea stevensii from the Orinoco River near the border of Colombia and Venezuela. Two new species are segregated from the Rinorea hirsuta species complex: Rinorea galeanoae-bernalii and Rinorea cogolloi, both from the eastern slopes of the Andean Central Cordillera along the mid-Magdalena River Valley in Colombia. From the widely distributed R. pubiflora species complex, we segregated one new species, Rinorea callejasii, from southeast Panama and the Chocó in Colombia. In addition to these five new taxa segregated from widely distributed species complexes, we discovered two previously unknown species with affinities to other Neotropical Rinorea. Rinorea aymardii is described from the Alto Orinoco-Casiquiare Biosphere Reserve in Venezuela and most closely resembles R. melanodonta from Colombia. Rinorea betancurii is segregated from R. macrocarpa and occurs in the Amazonian Regions of Colombia, Brazil, Peru and Venezuela. In this study, we provide descriptions, illustrations and distribution maps of the new species and make preliminary assessments of the risk of extinction using the IUCN Red List Categories and Criteria. We also furnish an identification key to the species of Rinorea sect. Pubiflorae in Colombia.


Thismia malayanaA flowering plant B flower, side view C flower, view from above D flower, longitudinal section E three stamens, outer view F tepal with terminal appendage, adaxial view G ovary, longitudinal section H style and stigma I involucral bract, adaxial view J leaf, adaxial view. All drawn by Mohamad Aidil Noordin from spirit material, Siti-Munirah FRI 101705 (A–C); FRI 101701 (D–J).
Thismia malayanaA flowering plant A1 floral tube, inner surface A2 annulus and stamen filaments, view from inside B inflorescence with anthetic flower and several young fruits B1 style and stigma B2 annulus, top view C flower, side view D, E stamens, view from inside and from outside, E1 stamen supraconnectives: one pair of club-shaped inwards-pointing, one pair of acute outwards-pointing, and one central appendage F stamen supraconnectives, apical view G stamen tube, view from below H, H1 fruit after dehiscence, top view, H2 seeds I shoot base with roots. Photos by Siti-Munirah (A1–I) and Hardy-Adrian (A) from FRI 101701 (A), FRI 101702 (F, G, I), FRI 101703 (E1), FRI 101705 (B, B2, H, H1, H2) & FRI 101710 (A1, A2, B1, C, D, E). Images not to scale (see dimensions in description and Figs 1, 3).
Thismia malayana with scales (the finest grade is 0.5 mm) A side view B top view C the size compared to the 20-sen coin (23.59 mm in diameter). Photos by Hardy-Adrian from uncollected plants.
Habitat (in situ) of Thismia malayana in Ulu Bendul RP in Gunung Angsi FR (A, B) and the Tengku Hassanal WR (C–E) AThismia malayana at its habitat, which is located right next to the main trail to Gunung Angsi B Siti-Munirah showing the habitat of T. malayanaC path to Lata Bujang and Gunung Benom D the plants growing on rotten wood E Mohamad-Shafiq observed a Thismia malayana in its habitat. Photos by Siti-Munirah (A, B) and Mohamad-Shafiq (C–E).
Distribution of Thismia malayana (black circle) in Peninsular Malaysia: the type locality in Ulu Bendul RP, Gunung Angsi FR in the state of Negeri Sembilan and the Tengku Hassanal WR, Temerloh in the state of Pahang.
Thismia malayana (Thismiaceae), a new mycoheterotrophic species from Peninsular Malaysia

May 2024

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145 Reads

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1 Citation

A new species of the mycoheterotrophic genus Thismia from Malaysia is described and illustrated. Thismia malayana introduced here was found in two localities: in the lowlands of Gunung Angsi Forest Reserve, Negeri Sembilan, and in the hilly dipterocarp forests of Gunung Benom in Tengku Hassanal Wildlife Reserve, Pahang. Thismia malayana falls into the section Thismia subsect. Odoardoa, as it has creeping vermiform roots and free equal tepals. It is characterised by the following taxonomically important features: a sepia-brown, urceolate-curved floral tube, free equal tepals with terminal appendages, prominent bright yellow annulus and bright violet-blue stamens each bearing five appendages (one pair of club-shaped inwards-pointing, one pair of acute outwards-pointing, and one central appendage). According to the categories and criteria of the IUCN Red List, T. malayana is provisionally classified as Vulnerable.


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