Research Article |
Corresponding author: Rafael F. de Almeida ( dealmeida.rafaelfelipe@gmail.com ) Academic editor: Alexander Sennikov
© 2024 Rafael F. de Almeida, Isa L. de Morais, Thais Alves-Silva, Higor Antonio-Domingues, Marco O. O. Pellegrini.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
de Almeida RF, de Morais IL, Alves-Silva T, Antonio-Domingues H, Pellegrini MOO (2024) A new classification system and taxonomic synopsis for Malpighiaceae (Malpighiales, Rosids) based on molecular phylogenetics, morphology, palynology, and chemistry. PhytoKeys 242: 69-138. https://doi.org/10.3897/phytokeys.242.117469
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Malpighiaceae has undergone unprecedented changes in its traditional classification in the past two decades due to several phylogenetic studies shedding light on the non-monophyly of all subfamilies and most tribes and genera. Even though morphological characters were used to reconstruct the last molecular generic phylogeny of Malpighiaceae, a new classification system has never been proposed for this family. Based on a comprehensive review of the last twenty years of published studies for this family, we propose a new classification system and provide a taxonomic synopsis for Malpighiaceae based on molecular phylogenetics, morphology, palynology, and chemistry as a baseline for the systematics, conservation, and taxonomy of this family worldwide. Malpighiaceae currently comprises two subfamilies (Byrsonimoideae and Malpighioideae), 12 tribes ( Acmanthereae , Acridocarpeae trib. nov., Barnebyeae trib. nov., Bunchosieae trib. nov., Byrsonimeae, Galphimieae, Gaudichaudieae, Hiptageae, Hiraeeae, Malpighieae, Mcvaughieae trib. nov., and Ptilochaeteae trib. nov.), 72 genera (incl. Mamedea gen. nov.), and 1,499 accepted species (715 of which are currently under some kind of extinction threat). We present identification keys for all subfamilies, tribes, and genera, a full morphological description for the proposed new genus, the re-circumscription of ten genera alongside the needed new combinations, the proposition of several new synonyms, the typification of several names, and notes on the taxonomy, distribution, conservation, and ecology up to the genus rank. Morphological plates are also provided to illustrate the immense diversity of morphological traits used in the new classification and synopsis.
Conservation, Elatinaceae, integrated monography, phylogeny, systematics, taxonomy
Malpighiaceae (Malpighiales) is a family of flowering plants currently comprising 75 genera and ca. 1,350 species of trees, shrubs, subshrubs, and lianas distributed in tropical and subtropical regions of the world (
In the past two decades, Malpighiaceae has gone through unparalleled changes in its traditional classification as a direct result of several molecular phylogenetic studies (
Molecular phylogenetic tree recovered by the ML of the reduced alignment for matK, ndhF, rbcL, and PHYC presented by
As a result of over ten years of integrative studies worldwide, and based on a comprehensive review of the last twenty years of published studies for this family, we present a new classification system for Malpighiaceae based on molecular phylogenetics, morphology, palynology, and chemistry. We also present a taxonomic synopsis, including descriptions or diagnoses for the two subfamilies and 12 tribes currently recognised, besides identification keys for subfamilies, tribes, and genera. Some genera are re-circumscribed, while others are proposed, together with the necessary new combinations. Several taxonomic changes are proposed for subfamilial, tribal, generic, and species ranks. A checklist of all currently accepted species of this family is also presented alongside their extinction threat risk (Suppl. material
We sampled one to three species and all 72 accepted Malpighiaceae genera recognised by us in the taxonomic treatment. The aligned matrix for matK, ndhF, rbcL, and PHYC from
The analyses of morphological data were based on specimens from the following herbaria: ALCB, AMAZ, ASE, BAH, BM, BHCB, BOTU, CEN, CEPEC, CESJ, CGMS, COL, CPAP, CVRD, CTES, CUZ, EAC, ESA, F, FLOR, FUEL, FURB, FZB, G, GH, GUA, HAS, HB, HCF, HEPH, HISA, HRB, HRCB, HSJRP, HST, HUCP, HUCS, HUEFS, HUEM, HUESC, HUFG, HUFU, HUPG, HURB, HUT, HUVA, IAC, IAN, ICN, INPA, IPA, JPB, L, LIL, K, MAC, MBM, MBML, MG, MICH, MO, MOSS, MPU, NY, OUPR, P, PACA, PAMG, PEUFR, PMSP, R, RB, RBR, RFA, PRE, S, SI, SMF, SP, SPF, SPSF, TEPB, U, UB, UEC, UFP, UFMS, UFMT, UFRN, UPCB, US, USZ, W, VIC, and VIES (acronyms according to Thiers, continuously updated). Indumentum terminology and structure shapes follow
All ten major informal phylogenetic clades of Malpighiaceae recognised in the taxonomy section were well-supported by bootstrap values ranging from 60–100% (Fig.
Despite Madagasikaria causing the non-monophyly of Rhynchophora (Fig.
Malpighia glabra L.
Trees, shrubs, subshrubs (erect, monopodial or scandent) or lianas, monoecious, rarely functionally dioecious or androdioecious, perennial; hairs unicellular, foot present, conspicuous or not, 2-branched (malpighiaceous), T-, Y- or V-shaped, rarely acicular or stellate, branches straight, undulate, or curled, surface smooth, rough or spiny (Fig.
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 Malpighiaceae A branch with opposite leaves of Bronwenia megaptera B branch with decussate leaves of Verrucularina glaucophylla C branch with alternate leaves of Stigmaphyllon angustilobum D branch with verticillate leaves of Pterandra pyroidea E interpetiolar stipules of Mascagnia cordifolia F epipetiolar stipules of Byrsonima intermedia G free stipules of Hiraea hatschbachii H connate stipules of Peixotoa catarinensis I 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 basiloba M leaf with short petiole of Banisteriopsis adenopoda N leaf with long petiole of Stigmaphyllon caatingicola O alternate glands on the petiole of Banisteriopsis membranifolia P opposite to alternate glands on the petiole of Schwannia mediterranea Q subopposite glands on the petiole of Banisteriopsis membranifolia R discoid and sessile gland of Banisteriopsis laevifolia S 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 Malpighiaceae A leaf with sagittate base of Stigmaphyllon ciliatum B leaf with rounded base of Banisteriopsis adenopoda C leaf with cordate base of Stigmaphyllon blanchetii D leaf with cuneate base of Banisteriopsis vernoniifolia E leaf with obtuse base of Stigmaphyllon paralias F leaf with truncate base of Stigmaphyllon gayanum G leaf with oblique base of Stigmaphyllon lanceolatum H leaf with attenuate base of Acmanthera minima I leaf with entire margin of Stigmaphyllon caatingicola J leaf with 3-lobed margin of Stigmaphyllon caatingicola K leaf with 5-lobed margin of Stigmaphyllon angustilobum L leaf with crenate margin of Stigmaphyllon crenatum M leaf with ciliate margin of Stigmaphyllon ciliatum N leaf with dentate margin of Stigmaphyllon vitifolium O leaf with plane blade margin of Banisteriopsis membranifolia P leaf with revolute blade margin of Verrucularina glaucophylla Q rounded leaf apex of Tetrapterys phlomoides R mucronate leaf apex of Banisteriopsis magdalenensis S emarginate leaf apex of Hiraea cuiabensis T cuspidate leaf apex of Banisteriopsis adenopoda U acuminate leaf apex of Mamedea harleyi V 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 Malpighiaceae A inflorescence evolution in Malpighiaceae according to
Compound inflorescences of Malpighiaceae A thyrse of 1-flowered cincinni of Byrsonima sericea B corymb of 1-flowered cincinni of Mascagnia cordifolia C umbel of 1-flowered cincinni of Banisteriopsis argyrophylla D line drawing of an inflorescence of Stigmaphyllon angustilobum showing 1-flowered cincinni arranged in umbels (1st order inflorescence), arranged in dichasia (2nd order inflorescence), arranged in a thyrse (3rd order inflorescence) E photograph of the inflorescence branch of Stigmaphyllon angustilobum (photographs B, C by M.O.O. Pellegrini; A, E by R.F. Almeida; line drawing D by K. Souza).
Flowers of Malpighiaceae A side view of the flower of Amorimia coriacea showing the cincinnus peduncle with a bract at base and two bracteoles at apex B detail of the 2-glandular bracteoles of Glicophyllum cardiophyllum C details of the glandular margin of the bracteole of Christianella multiglandulosa D bracteoles of Mezia araujoi concealing the floral bud at pre-anthesis (floral pedicel is absent) E floral bud of Niedenzuella multiglandulosa, showing a very short peduncle with a bract at base and two bracteoles at apex F flower of Banisteriopsis laevifolia showing pedicel with bract and bracteoles at base (peduncle absent) G plane and patent bract and bracteoles of Alicia anisopetala H cucullate bract and bracteoles of Dicella bracteosa I deflexed bract and bracteoles of Dicella nucifera J eglandular sepals concealing petals at pre-anthesis in Thryallis longifolia K 1-glandular calyx of Hiptage benghalensis L 10-glandular calyx of Camarea axillaris M 8-glandular calyx of Christianella multiglandulosa showing the multi-glandular margin of sepals N erect sepals of Galphimia australis O revolute apex of sepals of Byrsonima basiloba P revolute and reflexed sepals of Thryallis longifolia Q floral diagram of a Malpighiaceae flower with sepals in green, sepal glands in red, lateral petals in yellow, posterior petal in brown, androecium in blue (connectives) and white (pollen sacs), and gynoecium in pink (diagram and photographs B, E, F, H, K, L, N, O, Q by R.F. Almeida; A–D, G, M by M.O.O. Pellegrini; I by Amaury Jr.; J, P by J.V. Santos).
Sepals, glands, and petals of Malpighiaceae flowers A sepals with rounded apex of Hiptage benghalensis and sessile sepal gland B sepals with acute apex of Christianella multiglandulosa and sessile sepal glands C sepals with acute apex of Byrsonima gardneriana and sessile sepal glands D stalked sepal glands of Heladena multiflora E white petals of Acmanthera latifolia F pale yellow glands of Bunchosia glandulifera G yellow glands of Mcvaughia sergipana H pale green glands of Bronwenia megaptera I green glands of Camarea axillaris J yellow and deflexed petals of Byrsonima sericea K white glands of Acmanthera parviflora L pink glands of Heteropterys rubiginosa M brown glands of Amorimia pellegrinii N red glands of Niedenzuella poeppigiana O yellow and patent petals of Ptilochaeta bahiensis P oval petal limb of Galphimia gracilis Q elliptic petal limb of Heteropterys oberdanii R obovate petal limb of Bronwenia megaptera S pubescent petal surface of Diplopterys bahiana T orange-red petals of Tetrapterys phlomoides U entire margin of the petal of Heteropterys oberdanii V erose margin of the petal of Bronwenia megaptera W fimbriate margin of the petal of Schwannia schwannioides X dentate margin of the petal of Peixotoa hispidula Y glandular-fimbriate margin of the petal of Alicia anisopetala Z lilac petals of Mascagnia lilacina (A, C, F–H, I, J, L–S, U–W by R.F. Almeida; B, N, T, Y by M.O.O. Pellegrini; D by A. Francener; E by R. Goldenberg; Z by O.J.A. Ayala).
Androecium and gynoecium of Malpighiaceae A stamen of Banisteriopsis multifoliolata showing filament in green, anther in orange, glandular connective in blue, and pollen sacs in yellow B scanning electron micrograph of an anther of Banisteriopsis multifoliolata C SEM of a pollen grain of Banisteriopsis multifoliolata showing colpi in blue and endoaperture in red D heteromorphic stamen ring with 10 fertile stamens of Banisteriopsis multifoliolata E homomorphic stamen ring of Bronwenia megaptera with 10 fertile stamens F heteromorphic stamen ring of Peixotoa hispidula showing 5 fertile stamens and 5 staminodes in yellow G 6 fertile and free stamens of Schwannia hexandra H gynoecium of Banisteriopsis multifoliolata showing ovary in lilac, styles in blue and stigmas in yellow I uncinate apex of styles of Amorimia septentrionalis J truncate style apex of Amorimia rigida K subulate apex of style of Byrsonima sericea L 3 parallel and erect styles of Bronwenia megaptera M single curved style of Schwannia hexandra N 3 divergent styles of Stigmaphyllon blanchetii with foliate apices O 3 erect, slightly curved, and pubescent styles of Diplopterys lutea P 3 divergent styles of Stigmaphyllon lalandianum with reduced foliate apices Q 3 divergent styles of Stigmaphyllon glabrum without foliate apex (all line drawings by K. Souza; SEMs by R.F. Almeida; all scales: 1 mm, except for the pollen grain: 10 µm).
Types of fleshy fruits, nuts, and smooth to setose schizocarpic fruits in Malpighiaceae A transversely sliced drupe of Bunchosia maritima showing seeds B transversely sliced nut of Dicella nucifera showing the seed C nuts of Dicella bracteosa D nuts of Dicella macroptera E green drupe of Byrsonima blanchetiana F cream-coloured drupe of Byrsonima ligustrifolia G green drupe of Byrsonima melanocarpa with concrescent sepals H orange drupe of Bunchosia glandulifera I orange drupe of Bunchosia maritima J red drupe of Malpighia glabra K reddish-orange drupe of Malpighia mexicana L green drupe of Malpighia fucata M green and twisted drupe of Mcvaughia sergipana N striated drupe of Burdachia prismatocarpa O smooth mericarp of Heladena multiflora P smooth mericarp of Galphimia gracilis Q smooth and immature mericarp of Verrucularina glaucophylla R green and smooth mericarps of Acmanthera latifolia S smooth mericarps of Thryallis longifolia with concrescent sepals T setose mericarps of Tricomaria usillo U setose mericarps of Lasiocarpus ferrugineus V setose mericarp of Camarea axillaris W setose mericarp of Echinopterys eglandulosa (photographs A, B by Amaury Jr.; C, D by A. Assis; E, H, I, M, Q, V by R.F. Almeida; F by S.E. Martins; G by N. Bigio; J by P. Acevedo-Rodriguez; K, L by M.R. Pace; N by L.S.B. Calazans; O by A. Francener; P by M.O.O. Pellegrini; R by R. Goldenberg; S by J.V. Santos; T by I. Specogna; U by A. Nuno; W by S. Carnaham).
Types of winged schizocarpic fruits in Malpighiaceae A single dorsal winged mericarp of Diplopterys pubipetala showing lateral wings B detail of part of the winged mericarp of Banisteriopsis argyrophylla C, D 1 dorsal winged mericarp(s) of Heteropterys byrsonimifolia E winged mericarp of Peixotoa catharinensis showing lateral winglets F winged mericarp of Amorimia candidae with two lateral wings more developed than the reduced dorsal wing G winged mericarp of Lophopterys floribunda H winged mericarp of Carolus chasei with two lateral wings more developed (dorsal wing absent) I winged mericarp of Mascagnia sepium with 1 lateral orbicular wing J winged mericarp of Tetrapterys phlomoides with 4 free, lateral wings (superior ones longer, inferior ones shorter) K winged mericarp of Glicophyllum cardiophyllum with 4 free, lateral wings (superior ones shorter, inferior ones longer) L winged mericarp of Niedenzuella acutifolia with 4 free, equalling lateral wings M winged mericarp of Hiptage benghalensis with three free, lateral wings more developed (photographs A by A. Popovkin; B, E, I, J by M.O.O. Pellegrini; C, D, F, H, K, L by R.F. Almeida; G by G. Shimizu; M by G. Cahyadi).
Malpighiaceae is here circumscribed with two subfamilies, 12 tribes, 72 genera, and 1,499 species accepted (Table
Malpighiaceae Juss. | |
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Byrsonimoideae W.R.Anderson | Malpighioideae Burnett emend. R.F.Almeida |
Acmanthereae W.R.Anderson | Acridocarpeae R.F.Almeida |
3 genera, 23 species | 2 genera, 38 species |
Byrsonimeae W.R.Anderson | Mcvaughieae R.F.Almeida |
3 genera, 181 species | 3 genera, 12 species |
Galphimieae Nied. | Barnebyeae R.F.Almeida |
3 genera, 40 species | 1 genus, 2 species |
Ptilochaeteae R.F.Almeida | |
3 genera, 10 species | |
Bunchosieae R.F.Almeida | |
5 genera, 122 species | |
Hiraeeae A.Juss. emend. R.F.Almeida | |
5 genera, 105 species | |
Hiptageae DC. emend. R.F.Almeida | |
17 genera, 377 species | |
Malpighieae DC. emend. R.F.Almeida | |
13 genera, 253 species | |
Gaudichaudieae Horan emend. R.F.Almeida | |
14 genera, 337 species |
Identification keys for all subfamilies, tribes, and genera are presented, alongside a full morphological description for the proposed new genus, the recircumscription of ten genera accompanied by the needed new combinations, the proposition of several new synonyms, typification of miscellaneous names and notes on conservation, distribution, ecology, and taxonomy up to the genus rank.
1 | Posterior petal eglandular (Fig. |
Byrsonimoideae |
– | Posterior petal glandular (Fig. |
Malpighioideae |
Byrsonima Rich. ex Kunth.
Posterior petal eglandular, fertile stamens 10 (occasionally 6–10 in Diacidia), pollen 3-aperturate, zonoaperturate, colporate, styles apex subulate, mericarps smooth (i.e., never winged, or setose), chromosome number n = 6, presence of macrolactams and sulfenyl compounds.
The subfamily Byrsonimoideae currently comprises the original three tribes published by
1 | Leaf veins camptodromous (Fig. |
Acmanthereae |
– | Leaf veins brochidodromous (Fig. |
2 |
2 | Leaves eglandular; bracteoles eglandular (Fig. |
Byrsonimeae |
– | Leaves glandular (except in Verrucularina); bracteoles glandular (Fig. |
Galphimieae |
Acmanthera (A.Juss.) Griseb.
Stipules absent, leaf veins camptodromous, sepals enclosing petals in buds, carpels free, styles ventrally to subapically inserted on ovaries, presence of diazanapaphthalenes, propargyl-type 1,3-dipolar organic compounds, absence of benzopyrans, lactams, lignam glycosides, pyrimidine nucleosides, pyrimidine nucleotides, saccharolipids, sulfenyl compounds.
Acmanthereae
currently comprises only three accepted genera (Acmanthera, Coleostachys, and Pterandra) and 23 species (15 threatened species; Suppl. material
1 | Leaves lanceolate; flowers sessile, sepals eglandular, anthers poricidal | Coleostachys |
– | Leaves elliptic, ovate to obovate; flowers pedicellate, sepals glandular, anthers rimose | 2 |
2 | Flowers arranged in thyrses; sepals 2(–many)-glandular, petals 5(–7), glabrous, anthers with dorsal projections |
Acmanthera (Figs |
– | Flowers arranged in umbels; sepals 0–2-glandular, petals 5, pubescent, anthers with lateral projections |
Pterandra (Fig. |
≡ Pterandra sect. Acmanthera A.Juss., Ann. Sci. Nat. Bot., ser. 2, 13: 328. 1840.
Acmanthera latifolia (A.Juss.) Griseb.
Acmanthera currently comprises seven accepted species (four threatened species; Suppl. material
Coleostachys genipifolia A.Juss.
Coleostachys is represented by a single species (not threatened; Suppl. material
French Guiana: Cayenne., s.d., Martin s.n. (P-JU barcode P00671745!; isolectotypes: BR barcode BR0000008577450!, F barcode V0062669F!, K barcode K000427026!, MICH barcode MICH1102137!, P barcodes P02428718!, P02428719!, P02428720!, P02428721!, RB barcode 540728!).
Pterandra pyroidea A.Juss.
Pterandra currently comprises 15 accepted species (11 threatened species; Suppl. material
≡ Byrsoniminae Nied. in Engler, Nat. Pflanzenr. 92: 17, 28. 1928.
Byrsonima Rich. ex Kunth.
Stipules epipetiolar, leaves smaller than inflorescences, at least one petal cucullate at anthesis, presence of hydroxy acids and derivatives, imidolactams, keto acids and derivatives, organic phosphoric acids and derivatives, organofluorides, absence of oxanes.
Byrsonimeae currently comprises only three accepted genera (Blepharandra, Byrsonima, and Diacidia) and 181 species (57 threatened species; Suppl. material
1 | Lateral petals deflexed at anthesis, anther connectives expanded, anthers without stiff hairs; mericarps drupaceous |
Byrsonima (Figs |
– | Lateral petals patent at anthesis, anther connectives inconspicuous, anthers glabrous or with soft hairs; mericarps dry | 2 |
2 | Leaf base usually cordate, leaf apex rounded to emarginate; cincinni 2–3-flowered, sepals coriaceous, not accrescent in fruit, petals white, pink to red, anthers pubescent | Blepharandra |
– | Leaf base rounded, leaf apex acute to acuminate; cincinni 1-flowered, sepals membranous, accrescent in fruit, petals yellow, anthers glabrous | Diacidia |
= Callyntranthele Nied., Index Lect. Lyceo Braunsbergiensis 1897: 4. 1897. Type species: Callyntranthele angustifolia (Kunth) Nied. [≡ Blepharandra angustifolia (Kunth) W.R.Anderson].
Blepharandra hypoleuca (Benth.) Griseb.
Blepharandra currently comprises six accepted species (one threatened species; Suppl. material
= Alcoceratothrix Nied., Arbeiten Bot. Inst. Königl. Lyceums Hosianum Braunsberg 1: 45. 1901. Type species: Alcoceratothrix rugosa (Benth.) Nied. (≡ Byrsonima rugosa Benth.).
Byrsonima spicata (Cav.) DC.
Byrsonima currently comprises 164 accepted species (49 threatened species; Suppl. material
= Sipapoa Maguire, Mem. New York Bot. Gard. 8: 124. 1953. ≡ Diacidia subg. Sipapoa (Maguire) W.R.Anderson, Mem. New York Bot. Gard. 32: 63. 1981. Type species: Sipapoa kunhardtii Maguire [≡ Diacidia kunhardtii (Maguire) W.R.Anderson].
Diacidia galphimioides Griseb.
Diacidia currently comprises 11 accepted species (seven threatened species; Suppl. material
≡ Galphimiinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 53, 69. 1890.
Galphimia Cav.
Peduncle of cincinni present, floral buds with petals keeled, anther projections laterally inserted on thecae, presence of naphthopyrans and oxazinanes.
Galphimieae currently comprises only three accepted genera (Galphimia, Spachea, and Verrucularina) and 40 species (20 threatened species; Suppl. material
1 | Trees; leaves as long as the inflorescences; inflorescences pendulous, bracteoles glandular, glands stalked; filaments not changing colour at post-anthesis | Spachea |
– | Shrubs to subshrubs; leaves shorter than the inflorescences; inflorescences erect, bracteoles eglandular; filaments changing colour at post-anthesis | 3 |
2 | Leaves many-glandular; cincinni 1-flowered; calyx 0–5-glandular, when present secreting nectar, anthers smooth, unappendaged |
Galphimia (Figs |
– | Leaves eglandular; cincinni 1–3-flowered; calyx 10-glandular, glands secreting oil, anthers with 2 verrucose appendages at apex |
Verrucularina (Figs |
= Thryallis L., Sp. Pl., ed. 2: 554. 1762, nom. rej. ≡ Vorstia Adans., Fam. Pl. 2: (23). 1763, nom. superfl. Type species: Thryallis brasiliensis L. [≡ Galphimia brasiliensis (L.) A.Juss.].
Galphimia glauca Cav.
Galphimia currently comprises 26 accepted species (13 threatened species; Suppl. material
= Lophanthera A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 328. 1840, syn nov. Type species: Lophanthera kunthiana A.Juss., nom. superfl. [≡ Spachea longifolia (Kunth) R.F.Almeida & M.Pell.].
= Spachea sect. Meckelia Mart. ex A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 326. 1840 ≡ Meckelia (Mart. ex A.Juss.) Griseb. in Martius, Fl. Bras. 12(1): 25. 1858. Type species: Spachea tricarpa A.Juss.
= Andersoniella C.Davis & Amorim, Harvard Pap. Bot. 25(1): 51–56. 2020, nom. illeg., non Andersoniella K.J.F.Schmitz (1897) ≡ Andersoniodoxa C.Davis & Amorim, Phytotaxa 470(1): 121–122. 2020, syn. nov. Type: Andersoniodoxa spruceana (Nied.) C.Davis & Amorim [≡ Lophanthera spruceana (Nied.) R.F.Almeida & M.Pell.].
Spachea elegans (G.Mey.) A.Juss.
Spachea was described by
In the expanded circumscription presented here, Spachea includes 12 species (five threatened species; Suppl. material
≡ Lophanthera hammelii W.R.Anderson, Brittonia 35: 37. 1983 ≡ Andersoniella hammelii (W.R.Anderson) C.Davis & Amorim, Harvard Pap. Bot. 25: 53. 2020 ≡ Andersoniodoxa hammelii (W.R.Anderson) C.Davis & Amorim, Phytotaxa 470: 121. 2020.
≡ Lophanthera lactescens Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 103. 1925.
≡ Galphimia longifolia Kunth in F.W.H. von Humboldt, A.J.A. Bonpland & C.S. Kunth, Nov. Gen. Sp. 5: 173. 1822 ≡ Lophanthera longifolia (Kunth) Griseb. in C.F.P.von Martius & auct. suc. (eds.), Fl. Bras. 12(1): 25. 1858.
≡ Lophanthera marcelae W.R.Anderson, Acta Bot. Mex. 109: 37 (2014) ≡ Andersoniella marcelae (W.R.Anderson) C.Davis & Amorim, Harvard Pap. Bot. 25: 53. 2020 ≡ Andersoniodoxa marcelae (W.R.Anderson) C.Davis & Amorim, Phytotaxa 470: 121. 2020.
≡ Lophanthera pendula Ducke, Trop. Woods 50: 34. 1937.
≡ Lophanthera spruceana Nied., Arbeiten Bot. Inst. Königl. Lyceums Hosianum Braunsberg 5: 30. 1914 ≡ Andersoniella spruceana (Nied.) C.Davis & Amorim, Harvard Pap. Bot. 25: 55. 2020 ≡ Andersoniodoxa spruceana (Nied.) C.Davis & Amorim, Phytotaxa 470: 121. 2020.
≡ Verrucularia A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 327. 1840, nom. illeg., non Verrucularia Shur. Figs
Verrucularina glaucophylla (A.Juss.) Rauschert (≡ Verrucularia glaucophylla A.Juss.).
Verrucularina is a replacement name for Verrucularia A.Juss. since the latter is a posterior homonym of Verrucularia Suhr, a genus previously assigned to algae but currently belonging to Bryozoa. The genus currently comprises two accepted species (one threatened species; Suppl. material
Malpighia L.
Posterior petal glandular, 2–10 fertile stamens, pollen 3–12-aperturate, zono- to pantoaperturate, porate or colporate, styles capitate, uncinate, truncate, expanded or rarely subulate, stigmas usually lateral, nuts or mericarps, frequently winged or setose, rarely smooth, chromosome number n = 9–10, presence of dithiols, furanoid lignans, organic phosphoric acids and derivatives, and propargyl-type 1,3-dipolar organic compounds.
Aside from subfamily Byrsonimoideae, all previously proposed subfamilies are recovered nested within Malpighioideae, making it non-monophyletic. Furthermore, most of these subfamilies are non-monophyletic on their own since they were traditionally circumscribed based on fruit morphology (especially dry vs. fleshy) and the presence or absence of mericarp wings. Therefore, in our current circumscription, subfamily Malpighioideae comprises nine main lineages of mostly Neotropical genera of Malpighiaceae (including Burdachia, Glandonia, and Mcvaughia, which were previously placed by
1 | Stipules absent; leaves alternate (Fig. |
Acridocarpeae |
– | Stipules present; leaves opposite (Fig. |
2 |
2 | Cincinni 2–7-flowered (Fig. |
3 |
– | Cincinni 1-flowered (Fig. |
4 |
3 | Shrubs or subshrubs; leaves distributed along the branches (Fig. |
Mcvaughieae |
– | Trees; leaves congested at the apex of the branches (Fig. |
Barnebyeae |
4 | Leaves margin revolute when young (Fig. |
Ptilochaeteae |
– | Leaves margin plane when young (Fig. |
5 |
5 | Stipules inconspicuous (Fig. |
Hiptageae |
– | Stipules conspicuous (Fig. |
6 |
6 | Stipules epipetiolar (Fig. |
7 |
– | Stipules interpetiolar (Fig. |
8 |
7 | Leaf apex eglandular, tertiary veins reticulate; styles straight, parallel, apex capitate, stigma terminal (Fig. |
Bunchosieae |
– | Leaf apex glandular, tertiary veins scalariform; styles curved, divergent, apex uncinate, stigma lateral (Fig. |
Hiraeeae |
8 | Flowers arranged in umbels (thyrses in Bronwenia); fertile stamens 2–3–4–5–6–10; mericarps with a well-developed dorsal wing (larger than the lateral wings, when present), lateral wings reduced to absent (always smaller than the dorsal wing), free (Fig. |
Gaudichaudieae |
– | Flowers arranged in corymbs (thyrses in Amorimia and Ectopopterys); fertile stamens 10; drupes (Malpighia; Fig. |
Malpighieae |
Acridocarpus Guill. & Perr.
Lianas, shrubs to treelets; thyrses, many-flowered, cincinni 1-flowered, bracts 1-glandular, peduncle absent, bracteoles eglandular; sepals glandular, nectariferous; posterior petals 2, margin crenate, eglandular; connectives eglandular, anthers poricidal, pollen 3-zonosyncolporate; styles reflexed in fruits; mericarps 1-winged, dorsal wing more developed, chromosome number n = 9, presence of diazanaphthalenes, isoflavonoids, oxacyclic compounds, absence of tetrahydrofurans.
Acridocarpeae currently comprises only two accepted genera (Acridocarpus and Brachylophon) and 38 species (20 threatened species; Suppl. material
1 | Leaf apex rounded, acute or mucronate; corolla rotate, petals patent, margin not entire; dorsal wing well-developed; Africa to Western Asia | Acridocarpus |
– | Leaf apex caudate; corolla campanulate, petals erect, margin entire; dorsal wing very reduced; Southeastern Asia | Brachylophon |
= Heteropterys sect. Anomalopterys DC., Prodr. 1: 592. 1824 ≡ Anomalopterys (DC.) G.Don, Gen. Hist. 1: 647. 1831. Type species: Anomalopterys spicata G.Don [= Acridocarpus smeathmanii (DC.) Guill. & Perr.].
= Rhinopteryx Nied., Nat. Pflanzenfam. 3(4): 352. 1896. Type species: Rhinopteryx spectabilis Nied. [≡ Acridocarpus spectabilis (Nied.) Doorn-Hoekm.].
Acridocarpus plagiopterus Guill., Perr. & A.Rich.
Acridocarpus currently comprises 36 accepted species (19 threatened species; Suppl. material
Brachylophon curtisii Oliv.
Brachylophon currently comprises two accepted species (one threatened species; Suppl. material
Mcvaughia W.R.Anderson.
Trees, shrubs to subshrubs; thyrses, cincinni 1–7-flowered, bracteoles 1-glandular; pollen 4-zonocolporate (3-zonocolporate in Glandonia); drupes, epicarp striated, presence of linear 1,3-diarylpropanoids, and the absence of dithiols, indoles and derivatives.
Mcvaughieae currently comprises three accepted genera, Burdachia, Glandonia, and Mcvaughia, and 12 species (five threatened species; Suppl. material
1 | Pedicel straight at pre-anthesis, lateral petals yellow, fertile stamens 7, staminodes 3, anthers horseshoe-shaped, ovary 1-locular, styles straight at apex, stigma lateral; fruit pubescent |
Mcvaughia (Figs |
– | Pedicel circinate at pre-anthesis, lateral petals pink or white, fertile stamens 10, staminodes absent, anthers straight, ovary 3-locular, styles bent at apex, stigma terminal; fruit glabrous. | 2 |
2 | Stipules connate in epipetiolar pairs, persistent; inflorescences deflexed; floral buds globose, lateral petals pink, filaments glabrous, connective expanded, locule apex rounded, shorter than the connective |
Burdachia (Fig. |
– | Stipules connate in interpetiolar pairs, deciduous; inflorescences erect; floral buds pyramidal, lateral petals white, filaments pubescent, connective inconspicuous, locule apex acute, longer than the connective | Glandonia |
= Tetrapodenia Gleason, Bull. Torrey Bot. Club 53: 289. 1926. Type species: Tetrapodenia glandifera Gleason (= Burdachia sphaerocarpa A.Juss.).
Burdachia prismatocarpa A.Juss.
Burdachia comprises only six currently accepted species (one threatened species; Suppl. material
≡ Tetrapodenia glandifera Gleason, Bull. Torrey Bot. Club 53: 289. 1926 ≡ Burdachia sphaerocarpa var. glandifera (Gleason) W.R.Anderson, Mem. New York Bot. Gard. 32: 139. 1981.
≡ Burdachia prismatocarpa var. loretoensis W.R.Anderson, Mem. New York Bot. Gard. 32: 143. 1981.
Glandonia macrocarpa Griseb.
Glandonia comprises only three currently accepted species (one threatened species; Suppl. material
Mcvaughia bahiana W.R.Anderson.
Mcvaughia comprises only three currently accepted species (all threatened species; Suppl. material
Barnebya W.R.Anderson & B.Gates.
Trees; thyrses, cincinni 2–3-flowered; pollen 4-zonoporate; mericarps 1-winged, dorsal wing more developed, presence of diarylheptanoids, keto acids and derivatives, oxazinanes, absence of benzopyrans, furanoid lignans, glycerophospholipids, lignan glycosides, naphthalenes, naphthopyrans, propargyl-type 1,3-dipolar organic compounds, pteridines and derivatives, tetrahydrofurans.
Barnebyeae currently comprises a single genus, Barnebya, and two accepted species (one threatened species; Suppl. material
Barnebya dispar (Griseb.) W.R.Anderson & B.Gates.
Barnebya comprises two currently accepted species (one threatened species; Suppl. material
Ptilochaeta Turcz.
Treelets to shrubs; thyrses reduced, 4-flowered; pollen 8-zonocolporate, styles apex geniculate to truncate; mericarp winged or setose, presence of 2-aryl-benzofuran flavonoids, dibenzyl-butane lignans, isoflavonoids, oxacyclic compounds, oxanes, pyrrolidines, thiocarbonyl compounds, absence of organothiophosphorus compounds, thiophenes.
Ptilochaeteae currently comprises three accepted genera, Dinemandra, Lasiocarpus, and Ptilochaeta, and ten currently accepted species (one threatened species; Suppl. material
1 | Sepals with stipitate glands, posterior petal glandular, fertile stamens 2 or 8; mericarps winged; arid and desert areas of Argentina and Chile | Dinemandra |
– | Sepals eglandular, posterior petal eglandular, fertile stamens 10; mericarps setose; seasonally dry forests of Argentina, Bolivia, Brazil, Paraguay, and Mexico | 2 |
2 | Plants dioecious, sepal apex erect, petals narrowly elliptic, style apex expanded; Mexico |
Lasiocarpus (Fig. |
– | Plants monoecious; sepal apex convolute, petals widely elliptic to obovate, style apex truncate; Argentina, Bolivia, Brazil, Paraguay |
Ptilochaeta (Fig. |
= Dinemagonum A.Juss., Arch. Mus. Hist. Nat. 3: 585. 1843, syn. nov. Type species: Dinemagonum bridgesianum A.Juss. [= Dinemandra gayana (A.Juss.) R.F.Almeida & M.Pell.].
Dinemandra ericoides A.Juss. ex Endl.
Dinemandra and Dinemagonum were traditionally distinguished from each other based exclusively on their fruit morphology, with Dinemandra presenting dominant lateral wings and Dinemagonum presenting a dominant dorsal wing. Nonetheless, both genera are strongly supported as sister based on molecular data, being further morphologically supported by stalked sepal glands basally connate forming pairs (
≡ Dinemagonum gayanum A.Juss., Arch. Mus. Hist. Nat. 3: 585. 1843.
Lasiocarpus salicifolius Liebm.
Lasiocarpus comprises four currently accepted species (one threatened species; Suppl. material
Ptilochaeta bahiensis Turcz.
Ptilochaeta comprises only three currently accepted species (no threatened species; Suppl. material
= Thryallidinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 53, 67. 1890, syn. nov. Type genus: Thryallis Mart., nom. cons.
Bunchosia Rich. ex Kunth.
Trees, shrubs or lianas; thyrses, cincinni 1-flowered; pollen 4–12-pantoporate (colporate in Echinopterys and Heladena); styles free, rarely connate, parallel; stigma terminal, capitate; drupes or mericarps smooth or winged, presence of azolidines, benzodioxoles, organochlorides, quinolizines, absence of organic carbonic acids and derivatives, organic phosphoric acids and derivatives.
Bunchosieae currently comprises five accepted genera, Bunchosia, Echinopterys, Heladena, Thryallis, and Tristellateia, and 122 species (68 threatened species; Suppl. material
1 | Lianas; leaves glandular at or along margin | 2 |
– | Trees, shrubs or scandent shrubs; leaves eglandular or glandular at base | 3 |
2 | Floral buds smooth, sepals 2-glandular, glands pedunculate, petal margin fimbriate to denticulate, anthers rimose, styles 3; mericarps smooth or setose; Neotropics |
Heladena (Figs |
– | Floral buds keeled, sepals eglandular, petal margin entire, anthers poricidal, styles 1; mericarps winged; Paleotropics | Tristellateia |
3 | Leaves eglandular; stamen filaments pubescent; mericarps setose |
Echinopterys (Fig. |
– | Leaves glandular at base; stamen filaments glabrous; mericarps smooth or drupaceous | 4 |
4 | Trees or erect shrubs; inflorescence, flowers and fruits with malpighiaceous hairs, bracteoles glandular, not surrounding floral buds; sepals 2-glandular, erect at anthesis, anthers connivant; mericarps drupaceous |
Bunchosia (Figs |
– | Scandent shrubs; inflorescence, flowers and fruits with stellate hairs, bracteoles eglandular, surrounding floral buds; sepals eglandular, deflexed at anthesis, anthers divergent; mericarps smooth |
Thryallis (Figs |
= Malacmaea Griseb., Linnaea 13: 248. 1839. Type species: Malacmaea fluminensis Griseb. [= Bunchosia maritima (Vell.) J.F.Macbr.].
Bunchosia odorata (Jacq.) DC.
Bunchosia comprises 93 currently accepted species (46 threatened species; Suppl. material
= Bunchosia sect. Coelostylis A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 325. 1840 ≡ Coelostylis (A.Juss.) Kuntze, Revis. Gen. Pl. 1: 87. 1891, nom. illeg., non Coelostylis Torr. & A.Gray. Type species: Coelostylis glandulosa Kuntze [= Echinopterys eglandulosa (A.Juss.) Small].
Echinopterys lappula A.Juss. [= Echinopterys eglandulosa (A.Juss.) Small].
Echinopterys comprises only two currently accepted species of shrubs or lianas endemic to the seasonally dry tropical forests of Mexico (
= Henlea Griseb., Abh. Königl. Ges. Wiss. Göttingen 9: 37. 1860, syn. nov., nom. illeg., non Henlea H.Karst. ≡ Henleophytum H.Karst., Fl. Columb. 1: 158. 1861. Type species: Henleophytum echinatum (Griseb.) Small [≡ Heladena echinata (Griseb.) R.F.Almeida & M.Pell.].
= Malpigiantha Rojas Acosta, Cat. Hist. Nat. Corrientes: 55. 1897. Type species: Malpigiantha volubilis Rojas Acosta [= Heladena multiflora (Hook. & Arn.) Nied.].
Heladena multiflora (Hook. & Arn.) Nied.
Similar to Dinemandra and Dinemagonum, Heladena and Henleophytum are strongly supported as sister by molecular data, being exclusively distinguished by their fruit morphology (Heladena having smooth mericarps and Henleophytum having setose mericarps). However, both genera share unique stalked peltate sepal glands, added to hairy petals, weakly coherent but soon separating styles, and stigmas elliptic and geniculate. Thus, we also propose the expansion of Heladena to include two currently accepted species (one threatened species; Suppl. material
≡ Henlea echinata Griseb., Abh. Königl. Ges. Wiss. Göttingen 9: 37. 1860 ≡ Henleophytum echinatum (Griseb.) Small in Britton & al., N. Amer. Fl. 25: 149. 1910.
≡ Hemsleyna Kuntze, Revis. Gen. Pl. 1: 88. 1891.
Thryallis longifolia Mart.
Thryallis comprises five currently accepted species (one threatened species; Suppl. material
= Zymum Noronha ex Thouars, Hist. Vég. Îsles Austral. Afriq.: 69. 1808. Type species: Zymum madagascariense Spreng. (= Tristellateia madagascariensis Poir.).
= Platynema Wight & Arn., Edinburgh New Philos. J. 15: 179. 1833. Type species: Platynema laurifolium Wight & Arn. (= Tristellateia australasiae A.Rich.).
Tristellateia madagascariensis Poir.
Tristellateia comprises 21 species of lianas endemic to rainforests and seasonally dry tropical forests of Madagascar (19 threatened species; Suppl. material
Hiraea Jacq.
Lianas; leaf blades with apex glandular; thyrses, many-flowered; pollen 4–12-pantocolporate (porate in Psychopterys); styles with apex uncinate, stigma lateral; mericarps winged, 2 lateral wings more developed than the dorsal, usually butterfly-shaped, presence of piperidines, absence of benzofurans, benzopyrans, dithiols, furanoid lignans, hydroxy acids and derivatives, naphthopyrans, pteridines and derivatives, pyrimidine nucleosides.
Hiraeeae currently comprises five accepted genera, Adelphia, Excentradenia, Hiraea, Lophopterys and Psychopterys, and 105 species (54 threatened species; Suppl. material
1 | Flowers arranged in thyrses | 2 |
– | Flowers arranged in umbels | 4 |
2 | Inflorescence branches longitudinally costate; sepals 1-glandular, posterior petal shorter than laterals; mericarps with 2 V-shaped lateral wings |
Lophopterys (Fig. |
– | Inflorescence branches smooth; sepals 2-glandular, posterior petal equalling or longer than laterals; mericarps with 2 butterfly-shaped lateral wings | 3 |
3 | Leaves apex glandular; bracteoles glandular; petals yellow, deflexed, margin fimbriate, posterior petal longer than laterals, styles apex uncinate | Adelphia |
– | Leaves apex eglandular; bracteoles eglandular; petals white, patent, margin dentate to erose, posterior petal equalling laterals, styles apex capitate | Psychopterys |
4 | Stipules at base of petioles; posterior petal with fimbriae two times longer than those from the laterals; mericarps with lateral wings connate at base | Excentradenia |
– | Stipules at middle or apex of petioles; all petals with equally long fimbriae; mericarps with lateral wings free |
Hiraea (Figs |
Adelphia hiraea (Gaertn.) W.R.Anderson.
Adelphia comprises four currently accepted species (two threatened species; Suppl. material
Excentradenia adenophora (Sandw.) W.R.Anderson.
Excentradenia comprises four currently accepted species (two threatened species; Suppl. material
Hiraea reclinata Jacq.
Hiraea comprises 81 currently accepted species (43 threatened species; Suppl. material
= Dolichopterys Kosterm., Recueil Trav. Bot. Néerl. 32: 279. 1935. Type species: Dolichopterys surinamensis Kosterm. [≡ Lophopterys surinamensis (Kosterm.) Sandwith].
Lophopterys splendens A.Juss.
Lophopterys currently comprises seven accepted species (two threatened species; Suppl. material
Psychopterys dipholiphylla (Small) W.R.Anderson & S.Corso.
Psychopterys comprises nine currently accepted species (five threatened species; Suppl. material
= Banisterieae DC., Prodr. 1: 584. 1824, syn. nov. ≡ Banisteriinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 52, 60. 1890. Type genus: Banisteria L., nom. rej. (= Heteropterys Kunth).
= Tricomarieae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 52, 66. 1890, syn. nov. Type genus: Tricomaria Gillies ex Hook. & Arn.
Hiptage Gaertn.
Treelets, shrubs or lianas; thyrses, multi-flowered; pollen 4–12-pantocolporate (porate in Hiptage and some Heteropterys); nuts or mericarps winged, with 2–4-wings, butterfly, Y or X-shaped, rarely setose, absence of organic phosphonic acids and derivatives.
Hiptageae currently comprises 17 accepted genera, Alicia, Callaeum, Carolus, Chlorohiptage, Christianella, Dicella, Flabellaria, Flabellariopsis, Glicophyllum, Heteropterys, Hiptage, Jubelina, Malpighiodes, Mezia, Niedenzuella, Tetrapterys, Tricomaria, and 377 species (163 threatened species; Suppl. material
1 | Styles 1–2; mericarps with 3 free lateral wings or setose | 2 |
– | Styles 3; mericarps with 1–2–4 free lateral wings | 3 |
2 | Style 1, apex truncate, stigma terminal; mericarps with 3 free lateral wings; Africa and Asia |
Hiptage (Figs |
– | Styles 2, apex uncinate, stigma lateral; mericarps setose; South America (Argentina) |
Tricomaria (Fig. |
3 | Petals green, styles shorter than the filaments; mericarps with 1 lateral wing; Asia (Vietnam) | Chlorohiptage |
– | Petals white, lilac, yellow, orange or red, styles longer than the filaments; mericarps with 2–4 free lateral wings; Africa or Americas | 4 |
4 | Sepals deflexed, stigma terminal; Africa | 5 |
– | Sepals erect, stigma lateral; Americas | 6 |
5 | Leaves glandular at margin, petiole with 2–3 gland pairs; stigma capitate | Flabellaria |
– | Leaves glandular near or along margin, petiole eglandular; stigma truncate | Flabellariopsis |
6 | Petals glabrous to glabrescent | 7 |
– | Petals densely pubescent | 10 |
7 | Sepals not enclosing petals in bud, filaments usually glabrous | 8 |
– | Sepals enclosing petals in bud, filaments usually pubescent | 9 |
8 | Flowers arranged in thyrses, corymbs or umbels, inflorescences never arranged in dichasia; mericarps with 1 dominant dorsal wing |
Heteropterys (Figs |
– | Flowers arranged in umbels, inflorescences arranged in dichasia; mericarps with 4 dominant lateral wings |
Tetrapterys (Figs |
9 | Leaves glandular near or along margin, petioles eglandular; flowers arranged in umbels, 4-flowered, secondarily arranged in dichasia, bracteoles elliptic; mericarps with 2 dominant lateral wings, connate at base | Malpighiodes |
– | Leaves glandular at margin, petioles usually with 1 gland pair; flowers arranged in thyrses, many-flowered, solitary, bracteoles triangular; mericarps with 2–4 dominant lateral wings, free |
Niedenzuella (Figs |
10 | Bracteoles leaf-like; sepals enlarged in fruit; nuts |
Dicella (Figs |
– | Bracteoles minute; sepals not enlarged in fruit; schizocarps | 11 |
11 | Flowers arranged in umbels, 4-flowered, secondarily arranged in dichasia | 12 |
– | Flowers arranged in thyrses, many-flowered, solitary or grouped but never secondarily arranged in dichasia | 14 |
12 | Sepals deflexed at anthesis, anterior lateral petals deflexed at anthesis, posterior lateral petals patent at anthesis; mericarps with several lateral winglets, vertically inserted between lateral wings and the dorsal wing | Jubelina |
– | Sepals erect to patent at anthesis, lateral petals deflexed at anthesis; mericarps without lateral winglets, when present (in Mezia) horizontally inserted between the lateral wings and the dorsal wing | 13 |
13 | Bracteoles not enclosing floral bud; connectives inconspicuous; lateral wings free | Callaeum |
– | Bracteoles enclosing floral bud; connectives expanded; lateral wings connate at base |
Mezia (Fig. |
14 | Bracts, bracteoles, sepals and petals glandular at margin; mericarps with acicular (unbranched) hairs |
Christianella (Figs |
– | Bracts, bracteoles, sepals and petals eglandular at margin; mericarps with 2-branched hairs | 15 |
15 | Stipules interpetiolar; petioles eglandular; petal margin fimbriate, anthers pubescent |
Carolus (Fig. |
– | Stipules epipetiolar; petioles glandular; petal margin glandular, erose or dentate, anthers glabrous | 16 |
16 | Petioles with 2–4 gland pairs; bracteoles eglandular; lateral petals erect; mericarps bearing 2 dominant lateral wings, usually connate at base |
Alicia (Figs |
– | Petioles eglandular or with 1 gland pair; bracteoles glandular; lateral petals patent; mericarps bearing 2–4 dominant lateral wings, free |
Glicophyllum (Figs |
Alicia anisopetala (A.Juss.) W.R.Anderson.
Alicia comprises only two currently accepted species (no threatened species; Suppl. material
= Cabi Ducke, Arq. Serv. Florest. 2(1): 13. 1943. Type species: Cabi paraensis Ducke [= Callaeum antifebrile (Griseb.) D.M.Johnson].
Callaeum nicaraguense (Griseb.) Small.
Callaeum comprises 11 currently accepted species (five threatened species; Suppl. material
Carolus chlorocarpus (A.Juss.) W.R.Anderson.
Carolus comprises eight currently accepted species (five threatened species; Suppl. material
Chlorohiptage vietnamensis T.V.Do, T.A.Le & R.F.Almeida.
Chlorohiptage comprises a single species (one threatened; Suppl. material
Christianella mesoamericana (W.R.Anderson) W.R.Anderson.
Christianella comprises five currently accepted species (two threatened species; Suppl. material
Dicella bracteosa (A.Juss.) Griseb.
Dicella comprises seven currently accepted species (one threatened species; Suppl. material
Flabellaria paniculata Cav.
Flabellaria comprises a single currently accepted species of liana endemic to rainforests, savannas, and seasonally dry tropical forests of Africa (
Flabellariopsis acuminata (Engl.) R.Wilczek.
Flabellariopsis comprises a single currently accepted species (not threatened; Suppl. material
Glicophyllum chamaecerasifolium (A.Juss.) R.F.Almeida.
Glicophyllum comprises 28 currently accepted species (four threatened species; Suppl. material
≡ Hiraea argentea A.Juss., Fl. Bras. Merid. 3: 17. 1833 ≡ Tetrapterys jussieuana Nied. in Engler, Nat. Pflanzenr. 93: 169. 1928 ≡ Glicophyllum jussieuanum (Nied.) R.F.Almeida, Nordic J. Bot. 39(1)-e02876: 15. 2021.
= Banisteria L., Sp. Pl.: 427. 1753, nom. rej. Type species: Banisteria brachiata L. [≡ Heteropterys brachiata (L.) DC.].
= Banisteria sect. Holopetalon Griseb., Linnaea 13: 199. 1839 ≡ Holopetalon (Griseb.) Rchb., Deut. Bot. Herb.-Buch: 207. 1841. Type species: Banisteria patens Griseb. [≡ Heteropterys patens (Griseb.) A.Juss.]
= Clonodia Griseb. in Martius, Fl. Bras. 12(1): 26. 1858. Type species: Clonodia verrucosa Griseb. (= Heteropterys racemosa A.Juss.).
= Atopocarpus Cuatrec., Webbia 13: 454. 1958. Type species: Atopocarpus papillosus Cuatrec. (= Heteropterys racemosa A.Juss.).
= Skoliopteris Cuatrec., Webbia 13: 451. 1958. Type species: Skoliopteris lehmanniana (Nied.) Cuatrec. [= Heteropterys complicata (Kunth) W.R.Anderson & C.Davis].
Heteropterys purpurea (L.) Kunth.
Heteropterys comprises 166 currently accepted species (75 threatened species; Suppl. material
= Gaertnera Schreb., Gen. Pl., ed. 8[a]. 1: 290. 1789, nom. rej. Type species: Gaertnera indica J.F. Gmel. [= Hiptage benghalensis (L.) Kurz].
= Molina Cav., Diss. 9: 435. 1790. Type species: Molina racemosa Cav. [= Hiptage benghalensis (L.) Kurz].
= Succowia Dennst., Schlüssel Hortus Malab.: 32. 1818, nom. illeg., non Succowia Medik. Type species: Succowia fimbriata Dennst. [= Hiptage benghalensis (L.) Kurz].
Hiptage madablota Gaertn. [= Hiptage benghalensis (L.) Kurz].
Hiptage comprises 47 currently accepted species (39 threatened species; Suppl. material
= Sprucina Nied., Arbeiten Bot. Inst. Königl. Lyceums Hosianum Braunsberg 3: 18. 1908. Type species (designated here): J. grisebachiana W.R.Anderson.
Jubelina riparia A.Juss.
No names have ever been published under the generic name Sprucina, but the collection cited in the protologue (Spruce 2853) refers to J. grisebachiana W.R.Anderson. Thus, J. grisebachiana is here designated as the type of Sprucina under Art. 10.2 Ex. 2 (
Malpighiodes spruceana Nied. [=Malpighiodes bracteosa (Griseb.) W.R.Anderson].
Malpighiodes comprises four currently accepted species (one threatened species; Suppl. material
= Stenocalyx Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 393. 1858, nom. illeg., non Stenocalyx O.Berg. (1856). Type species: Stenocalyx involutus Turcz. [= Mezia includens (Benth.) Cuatrec.].
Mezia araujoi Schwacke ex Nied.
Mezia comprises 15 currently accepted species (eight threatened species; Suppl. material
= Aenigmatanthera W.R.Anderson, Novon 16: 173. 2006. Type species: Aenigmatanthera lasiandra (A.Juss.) W.R.Anderson [≡ Niedenzuella lasiandra (A.Juss.) R.F.Almeida].
Niedenzuella poeppigiana (A.Juss.) W.R.Anderson.
Niedenzuella currently comprises 18 accepted species (three threatened species; Suppl. material
= Adenoporces Small in Britton & al., N. Amer. Fl. 25: 128. 1910. Type species: Adenoporces buxifolius (Cav.) Small (≡ Tetrapterys buxifolia Cav.).
Tetrapterys inaequalis Cav.
Tetrapterys comprises 56 currently accepted species (18 threatened species; Suppl. material
≡ Tetrapterys discolor var. andina Nied., Verz. Vorles. Königl. Lyceum Hosianum Braunsberg 1909–1910: 42. 1909.
Tricomaria usillo Gillies ex Hook. & Arn.
Tricomaria comprises a single currently accepted species (no threatened species; Suppl. material
≡ Malpighiinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 53, 71. 1890.
= Aspidopteryinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 52, 53. 1890, as “Aspidopteridinae”, syn. nov. Type genus: Aspidopterys A.Juss. ex Endl.
= Mascagniinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 52, 55. 1890, syn. nov. Type genus: Mascagnia (Bertero ex DC.) Bertero.
= Rhynchophoreae Arènes, Notul. Syst. (Paris) 12: 135. 1946, syn. nov. Type genus: Rhynchophora Arènes.
Malpighia L.
Treelets, shrubs or lianas; thyrses or corymbs; pollen 4–12-pantocolporate (porate in the Paleotropical species); styles with apex uncinate to truncate, stigma lateral; mericarps winged, 1–2-wings, butterfly-shaped to orbicular, rarely drupaceous or with dorsal wing more developed than lateral ones, presence of (3’–>5’)-dinucleotides and analogues, piperidines, absence of benzofurans, furanoid lignans, imidolactams, lignan glycosides.
Malpighieae currently comprises 13 accepted genera: Amorimia, Aspidopterys, Calcicola, Caucanthus, Diaspis, Digoniopterys, Ectopopterys, Madagasikaria, Malpighia, Mascagnia, Microsteira, Rhynchophora, and Triaspis, and 253 species (157 threatened species; Suppl. material
1 | Plants androdioecious; flowers actinomorphic, sepals eglandular, stigmas terminal | 2 |
– | Plants dioecious (androdioecious in Triaspis); flowers zygomorphic, sepals glandular, stigmas lateral (terminal in Triaspis) | 8 |
2 | Style apex truncate; continental Africa, Arabian Peninsula, and Asia | 3 |
– | Style apex with projections (1–2-lobed); Madagascar | 5 |
3 | Flowers arranged in umbels; flower buds obovoid, petal margin entire, reflexed, filaments slightly longer than sepals; Asia | Aspidopterys |
– | Flowers arranged in corymbs; flower buds ovoid to oblongoid, petal margin fimbriate to lobed, patent, filaments shorter or two times longer than sepals; Africa and Arabian Peninsula | 4 |
4 | Leaves spirally-alternate, glabrous; petal margin fimbriate, limb base obtuse, 2-carpellate | Diaspis |
– | Leaves opposite, tomentose; petals margin undulate, limb base sagittate, 3-carpellate | Caucanthus |
5 | Leaves spirally-alternate, up to 5 mm wide; umbels 1-flowered; petals narrowly spatulate, abaxially completely densely sericeous, style apex long-lobed | Digoniopterys |
– | Leaves opposite, at least 1 cm wide (mostly much wider); umbels 4–many-flowered; petals elliptic to orbicular, glabrous or abaxially sparsely sericeous along the keel, style apex shortly-lobed | 6 |
6 | Stipules enlarged, leaf-like, persistent; flowers in thyrses; mericarps with wings fused into an orbicular wing | Madagasikaria |
– | Stipules reduced, triangular, persistent to deciduous; flowers in umbels; mericarps with lateral wings fused into a single apical geniculate wing or a Y-shaped wing | 7 |
7 | Ovary bearing conspicuous initials for lateral wings and dorsal crest on each carpel, visible even in young flowers; mericarps dehiscent, lateral wings fused into a Y-shaped wing | Microsteira |
– | Ovary lacking initials for wings or crests; mericarps indehiscent, lateral wings fused into a single apical geniculate wing | Rhynchophora |
8 | Plants monoecious; stipules connate, leaf-like; bracteoles inserted at middle or below peduncle apex; floral buds keeled; sepals 1–2-glandular, glands very reduced, secreting nectar, petal margin long-fimbriate, limb base sagittate; Paleotropics | Triaspis |
– | Plants dioecious; stipules free, triangular; bracteoles inserted at peduncle apex (except in Calcicola); floral buds smooth (except in few Mascagnia spp.); sepals 2-glandular, glands large, secreting oil, petal margin entire, limb base obtuse, cuneate or rounded; Neotropics | 9 |
9 | Sepals deflexed at anthesis, connectives bearing large glands, style apex lobed; mericarp with a dominant dorsal wing | Ectopopterys |
– | Sepals erect at anthesis, connectives bearing inconspicuous glandular tissue, style apex truncate; mericarp with dominant lateral wings or wings greatly reduced and fleshy at maturity | 10 |
10 | Lianas; flowers arranged in thyrses or corymbs, bracteoles 1–6-glandular, rarely eglandular | 11 |
– | Shrubs to treelets; flowers arranged in umbels, bracteoles eglandular | 12 |
11 | Flowers arranged in thyrses, bracteoles 2–6-glandular; floral buds smooth; petals yellow, turning orange to red at post-anthesis, pubescent; mericarp with lateral wings free, coriaceous |
Amorimia (Figs |
– | Flowers in thyrses or corymbs, bracteoles 0–1-glandular; floral buds keeled; petals white, pink or lilac, if yellow not turning orange to red at post-anthesis; glabrous; mericarp with lateral wings fused into an orbicular wing, membranous |
Mascagnia (Figs |
12 | Leaves with stiff, spine-like (generally urticating) hairs; bracteoles inserted at peduncle apex; ovary glabrous; mericarps indehiscent, fleshy, dorsal and lateral wings much reduced, free, fleshy at maturity |
Malpighia (Fig. |
– | Leaves with soft hairs; bracteoles inserted at peduncle middle; ovary pubescent; mericarps dehiscent, dry, lateral wings conspicuous, fused | Calcicola |
Amorimia rigida (A.Juss.) W.R.Anderson.
Amorimia comprises 15 currently accepted species (eight threatened species; Suppl. material
Aspidopterys elliptica (Blume) A.Juss. ex Endl.
Aspidopterys comprises 24 currently accepted species (ten threatened species; Suppl. material
Calcicola parvifolia (A.Juss.) W.R.Anderson & C.Davis.
Calcicola comprises only two currently accepted species (no threatened species; Suppl. material
= Caucanthus sect. Eriocaucanthus Nied., Bull. Herb. Boissier, sér. 2, 4: 1010. 1904 ≡ Eriocaucanthus (Nied.) Chiov., Ann. Bot. (Rome) 10: 29. 1912. Type species: Caucanthus argenteus Nied.
Caucanthus edulis Forssk.
Caucanthus comprises only two currently accepted species (no threatened species; Suppl. material
Diaspis albida Nied.
Diaspis comprises a single currently accepted species (no threatened species; Suppl. material
Digoniopterys microphylla Arènes.
Digoniopterys comprises a single currently accepted species (one threatened species; Suppl. material
Ectopopterys soejartoi W.R.Anderson.
Ectopopterys comprises a single currently accepted species (no threatened species; Suppl. material
Madagasikaria andersonii C.Davis.
Madagasikaria comprises a single currently accepted species (one threatened species; Suppl. material
= Rudolphia Medik., Malvenfam.: 111. 1787. Type species: Rudolphia edulis Medik. (= Malpighia urens subsp. urens).
Malpighia glabra Plum. ex L.
Malpighia comprises 110 currently accepted species (85 threatened species; Suppl. material
≡ Hiraea [unranked] Mascagnia Bertero ex DC., Prodr. 1: 585. 1824.
= Triopterys L., Sp. Pl.: 428. 1753, nom. rej. Type species: Triopterys jamaicensis L. [= Mascagnia lucida (Kunth) W.R.Anderson & C.Davis].
Mascagnia americana Bertero [= Mascagnia macradena (DC.) Nied.].
Mascagnia comprises 48 currently accepted species (19 threatened species; Suppl. material
Microsteira curtisii Baker.
Microsteira comprises 27 currently accepted species (all threatened species; Suppl. material
= Calyptostylis Arènes, Notul. Syst. (Paris) 12: 131. 1946. Type species: Calyptostylis humbertii Arènes (= Rhynchophora phillipsonii W.R.Anderson).
Rhynchophora humbertii Arènes.
Rhynchophora comprises only two currently accepted species (all threatened species; Suppl. material
= Umbellulanthus S.Moore, J. Bot. 58: 220. 1920. Type species: Umbellulanthus floribundus S.Moore (≡ Triaspis mooreana Exell & Mendonça).
Triaspis hypericoides Burch.
Triaspis comprises 19 currently accepted species (five threatened species; Suppl. material
≡ Gaudichaudioideae A.Juss. ex C.V.Morton, Taxon 17: 318. 1968.
= Sphedamnocarpinae Nied. in Engler & Prantl, Nat. Pflanzenfam. III, 4: 52, 59. 1890, syn. nov. Type genus: Sphedamnocarpus Planch. ex Benth. & Hook.f.
Gaudichaudia Kunth.
Lianas, shrubs to subshrubs; umbels, rarely thyrses, usually 4-flowered; pollen 4–12-pantocolporate (porate in Stigmaphyllon subg. Ryssopterys, Philgamia, and Sphedamnocarpus); mericarps winged, 1-winged, dorsal wing more developed, rarely reduced, presence of macrolactams, absence of biotin and derivatives, sulfenyl compounds.
Gaudichaudieae currently comprises 14 accepted genera, Aspicarpa, Banisteriopsis, Bronwenia, Camarea, Cottsia, Diplopterys, Janusia, Mamedea, Mionandra, Peixotoa, Philgamia, Schwannia, Sphedamnocarpus, and Stigmaphyllon, and 336 species (154 threatened species; Suppl. material
1 | Petiole or leaf base glands ellipsoid, sunken; flowers arranged in thyrses, secondarily arranged in thyrses; sepal glands decurrent into the pedicel, stamens homomorphic |
Bronwenia (Figs |
– | Petioles or leaf base glands discoid or cupuliform, not sunken; flowers arranged in corymbs or umbels, solitary or secondarily arranged in dichasia or thyrses, rarely solitary thyrses; sepals not decurrent, stamens heteromorphic | 2 |
2 | Leaves apex long-acuminate; petals abaxially pubescent, styles pubescent; mericarps with dominant lateral wings |
Diplopterys (Figs |
– | Leaves apex emarginate, rounded, obtuse, acute, short-acuminate or acuminate; petals abaxially glabrous, styles glabrous; mericarps with a dominant dorsal wing, lateral wings reduced or absent | 3 |
3 | Branches with scale-like hairs; leaves long-petiolate, rarely short-petiolate; flowers arranged in corymbs or umbels, 5–many-flowered, peduncle curved; styles divergent and lyrate, apex expanded (foliaceous), rarely reduced |
Stigmaphyllon (Figs |
– | Branches without scale-like hairs; leaves short-petiolate; flowers arranged in umbels or thyrses, 1–4-flowered, peduncle straight; styles parallel and straight, apex truncate or cylindrical | 4 |
4 | Cincinni sessile or short-pedunculate; styles 3 | 5 |
– | Cincinni long-pedunculate; style 1(–2–3), if styles 2–3 then carpels slightly rotated so that no carpel aligns with the anterior sepals and posterior petal, and mericarps with dominant lateral wings | 9 |
5 | Stipules minute, free; flowers chasmogamous, sepal apex straight, fertile stamens 10, staminodes absent | 6 |
– | Stipules expanded, fused or bifid; flowers chasmogamous or cleistogamous, sepal apex revolute or involute along margins, fertile stamens 5, staminodes 3–5 | 8 |
6 | Flowers zygomorphic, sepals at anthesis bent towards the centre of the flower, connectives glandular; Neotropics |
Banisteriopsis (Figs |
– | Flowers actinomorphic, sepals erect at anthesis, connectives eglandular; Africa | 7 |
7 | Petals yellow; mericarp dorsal wing well-developed; Africa and Madagascar | Sphedamnocarpus |
– | Petals white; mericarp dorsal wing absent or very reduced; Madagascar | Philgamia |
8 | Stipules connate at base or up to the middle (i.e., bifid); flowers arranged in umbels, 1-flowered, bract and bracteoles absent; sepals free, completely revolute or involute along margins, antherodes filiform, minute, styles apex truncate to slightly expanded | Mionandra |
– | Stipules connate (i.e., entire); flowers arranged in umbels, 4-flowered, bract and bracteoles present; sepals connate at base, revolute only at apex, antherodes globose, conspicuous, styles apex capitate |
Peixotoa (Figs |
9 | Flowers chasmogamous, fertile stamens 2, staminodes 3, antherodes absent, carpels syncarpic | Cottsia |
– | Flowers chasmogamous or cleistogamous, fertile stamens 3–4–5–6, staminodes 0–2, antherodes present, carpels syncarpic at base and apically apocarpic | 10 |
10 | Flowers enantiostylous, petal margin long-fimbriate, fertile stamens 6, style curved |
Schwannia (Figs |
– | Flowers non-enantiostylous, petal margin entire, erose, denticulate or dentate, rarely short-fimbriate, fertile stamens 3–4–5, style straight | 11 |
11 | Fertile stamens 5, staminodes absent; mericarp dorsal wing developed | Janusia |
– | Fertile stamens 3–4(–5), staminodes present; mericarp dorsal wing absent or reduced to a crest | 12 |
12 | Branches herbaceous, annual, with acicular hairs; leaf blades reduced and narrow, margin usually revolute; fertile stamens 4, homomorphic |
Camarea (Figs |
– | Branches woody, perennial, with malpighiaceous hairs; leaf blades usually expanded and broad, margin plane; fertile stamens 3(–5), heteromorphic | 13 |
13 | Leaf base never with filamentous or tooth-like projections; flowers arranged in 1–4-flowered umbels, peduncles absent to reduced, without associated reduced leaves; antherodes equalling or larger than anthers (reduced to an apical swelling in M. harleyi and M. lanata), pubescent (glabrous in M. harleyi), usually red to orange at post-anthesis, 2 posterior carpels rotated so that all face the posterior petal; mericarp wings reduced to crests to teeth; central to southern South America |
Mamedea (Fig. |
– | Leaf base generally with a filamentous or tooth-like projection at each side of the blade; flowers arranged in 3–4-flowered umbels, peduncles long, with associated reduced leaves; antherodes smaller than anthers (few species with all anthers fertile), glabrous, yellow at post-anthesis, carpels slightly rotated so that no carpel aligns with the anterior sepals and posterior petal; mericarp lateral wings dominant (often fused), sometimes all wings equally developed into wings or reduced to crests; North and Central America | Aspicarpa |
= Acosmus Desv., J. Bot. Agric. 3: 229. 1816. Type species: Acosmus pruriens Desv. (=Aspicarpa hirtella Rich.).
= Gaudichaudia Kunth, Nov. Gen. Sp. (quarto ed.) 5: pl. 445. 1821, syn. nov. Type species: Gaudichaudia cynanchoides Kunth [≡ Aspicarpa cynanchoides (Kunth) Hassl.].
= Gaudichaudia [unranked] Tritomopterys A.Juss. ex Endl., Gen. Pl. 1058. 1840 ≡ Tritomopterys (A.Juss. ex Endl.) Nied., Arbeiten Bot. Inst. Königl. Lyceums Hosianum Braunsberg 4: 28. 1912. Type species (designated here): Gaudichaudia confertiflora A.Juss. [≡ Aspicarpa confertiflora (A.Juss.) R.F.Almeida & M.Pell.].
= Rosanthus Small in Britton & al., N. Amer. Fl. 25: 131. 1910. Type species: Rosanthus subverticellatus (Rose) Small [≡ Aspicarpa subverticillata (Rose) Hassl.].
Aspicarpa hirtella Rich.
In its current circumscription, Aspicarpa (now including Gaudichaudia) comprises 27 species (ten threatened species; Suppl. material
Most of the morphological diversity found in Aspicarpa s.lat. (especially the production of cleistogamous flowers and variation in the number of style number) might be attributed to polyploidy events (
≡ Gaudichaudia andersonii S.L.Jessup, Madroño 49: 254. 2002.
≡ Gaudichaudia arnottiana A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 252. 1840.
≡ Gaudichaudia chasei W.R.Anderson, Contr. Univ. Michigan Herb. 16: 68. 1987.
≡ Gaudichaudia confertiflora A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 252. 1840.
≡ Hiraea cycloptera DC., Prodr. 1: 586. 1824.
≡ Gaudichaudia filipendula A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 252. 1840.
≡ Gaudichaudia implexa S.L.Jessup, Madroño 49: 247. 2002.
≡ Gaudichaudia intermixteca S.L.Jessup, Madroño 49: 251. 2002.
≡ Gaudichaudia krusei W.R.Anderson, Contr. Univ. Michigan Herb. 16: 69. 1987.
≡ Gaudichaudia mcvaughii W.R.Anderson, Contr. Univ. Michigan Herb. 16: 72. 1987.
≡ Hiraea oxyota DC., Prodr. 1: 586. 1824.
≡ Gaudichaudia palmeri S.Watson, Proc. Amer. Acad. Arts 21: 421. 1886.
≡ Gaudichaudia symplecta S.L.Jessup, Madroño 49(4): 253. 2002.
≡ Gaudichaudia synoptera S.L.Jessup, Madroño 49(4): 251. 2002.
≡ Gaudichaudia zygoptera S.L.Jessup, Madroño 49: 249. 2002.
Banisteriopsis argentea (Kunth) C.R.Rob. [= Banisteriopsis muricata (Cav.) Cuatrec.]
Banisteriopsis comprises 65 currently accepted species (24 threatened species; Suppl. material
≡ Banisteriopsis malifolia var. appressa B.Gates, Fl. Neotrop. Monogr. 30: 79. 1982.
≡ Banisteriopsis martiniana var. subenervia Cuatrec., Webbia 13: 501. 1958.
≡ Banisteriopsis pulchra var. glabrata B.Gates, Fl. Neotrop. Monogr. 30: 109. 1982.
Bronwenia ferruginea (Cav.) W.R.Anderson & C.Davis.
Bronwenia comprises 13 currently accepted species (four threatened species; Suppl. material
≡ Banisteriopsis acapulcensis var. llanensis B.Gates, Fl. Neotrop. Monogr. 30: 46. 1982 ≡ Bronwenia acapulcensis var. llanensis (B.Gates) W.R.Anderson & C.Davis, Contr. Univ. Michigan Herb. 25: 141. 2007.
≡ Banisteria maracaybensis A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 285. 1840 ≡ Banisteriopsis cornifolia var. maracaybensis (A.Juss.) W.R.Anderson, Contr. Univ. Michigan Herb. 20: 15. 1995 ≡ Bronwenia cornifolia var. maracaybensis (A.Juss.) W.R.Anderson & C.Davis, Contr. Univ. Michigan Herb. 25: 143. 2007.
≡ Banisteriopsis cornifolia var. standleyi B.Gates, Fl. Neotrop. Monogr. 30: 44. 1982 ≡ Bronwenia cornifolia var. standleyi (B.Gates) W.R.Anderson & C.Davis, Contr. Univ. Michigan Herb. 25: 143. 2007.
= Camarea sect. Cryptolappa A.Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 254. 1840 ≡ Cryptolappa (A.Juss.) Kuntze, Revis. Gen. Pl. 1: 88. 1891. Type species: Camarea affinis A.St.-Hil.
Camarea ericoides A.St.-Hil.
Camarea comprises eight currently accepted species (three threatened species; Suppl. material
= Janusia sect. Metajanusia Nied., Verz. Vorles. Königl. Lyceum Hosianum Braunsberg 1912–1913: 50. 1912 ≡ Gaudichaudia sect. Erostratae Chodat, Bull. Soc. Bot. Genève, sér. 2, 9: 100. 1917, nom. superfl. ≡ Aspicarpa sect. Metajanusia (Nied.) Hassl., Annuaire Conserv. Jard. Bot. Genève 20: 212. 1918. Type species: Janusia gracilis A.Gray [≡ Cottsia gracilis (A.Gray) W.R.Anderson & C.Davis].
Cottsia scandens Dubard & Dop [= Cottsia californica (Benth.) W.R.Anderson & C.Davis].
Cottsia comprises four currently accepted species (one threatened species; Suppl. material
= Jubistylis Rusby, Mem. New York Bot. Gard. 7: 273. 1927. Type species: Jubistylis mollis Rusby [= Diplopterys lutea (Ruiz ex Griseb.) W.R.Anderson & C.Davis].
Diplopterys paralias A.Juss. [=Diplopterys pauciflora (G. Mey.) Nied.]
Diplopterys comprises 31 currently accepted species (11 threatened species; Suppl. material
= Peregrina W.R.Anderson, Syst. Bot. 10(3): 303. 1985, syn. nov. Type species: Peregrina linearifolia (A.St.-Hil.) W.R.Anderson [≡ Janusia linearifolia (A.St.-Hil.) A.Juss.].
Janusia guaranitica (A.St.-Hil.) A.Juss. ex Endl.
With the reestablishment of Schwannia, the recognition of Peregrina as independent of Janusia based only on the subshrub habit (vs liana) and laterally flattened stigmas (vs rounded stigmas) unnecessarily inflates the number of genera in Malpighiaceae without providing any taxonomic or systematic benefits. Since Janusia and Peregrina share non-enantioustilous flowers and androecia with five fertile stamens, without staminodes, we choose to return Peregrina to Janusia. In its current sense, Janusia comprises only two currently accepted species (one threatened species; Suppl. material
Mamedea pulchella (Griseb.) R.F.Almeida & M.Pell.
Mamedea can be recognised by its erect shrub to subshrub habit, present xylopodium, leaves entire at base, umbels, 1–4-flowered, peduncle usually absent or reduced, not bearing reduced leaves, sepals bent inwards between the petals at anthesis, petals fimbriate, androecium with 3 fertile stamens, anthers glabrous to pubescent, staminodes 2, antherodes present or not, when present larger than the fertile anthers, glabrous to pubescent, usually red to orange at post-anthesis, 2 posterior carpels rotated so that all face the posterior petal, mericarps with dorsal and lateral wings reduced to ribs or teeth, and with a rugose nut, chromosome number n = (20–)40.
Shrubs to subshrubs . Roots fibrous, woody near the xylopodium. Xylopodium present, small to large. Branches erect, slender, woody to herbaceous, sometimes brittle, sericeous to glabrescent; internodes inconspicuous to elongated. Stipules interpetiolar, minute, free to connate, sericeous or distally glabrous, deciduous or persistent. Leaves opposite or decussate; petioles short, sericeous, tomentose, lanate or glabrescent, eglandular; lamina entire, elliptical, lanceolate to ovate, velutinous, sericeous, lanate or tomentose, base cuneate or rounded, margin entire, apex acute, obtuse, rounded or mucronate; venation eucamptodromous or brochidodromous, secondary veins strongly ascending and subparallel. Umbels solitary, axillary, (1–)2–4-flowered, sessile to pedunculate; inflorescence leaves not reduced; bract alternate, minute, plane, persistent, sericeous to glabrous, eglandular, persistent; cincinni (1–)2–4, alternate, 1-flowered, pedunculate; bracteoles opposite, minute, plane, persistent, sericeous to glabrous, eglandular, persistent. Flowers chasmogamous or cleistogamous, bisexual, zygomorphic, hypogynous; pedicel elongated, longer or shorter than the peduncle, sparsely sericeous, tomentose, velutinous or glabrescent; sepals 5, free valvate in bud, erect in bud, bent inwards between the petals at anthesis, triangular to broadly ovate, sericeous or tomentose, apex acute, the anterior eglandular and narrower, the lateral 4 biglandular, the glands green, yellowish-green, dark red, or reddish-purple, secreting oil, in fruit persistent, somewhat accrescent, enclosing nutlets until maturity; petals 5, imbricate in bud, yellow to orange-yellow at anthesis, glabrous or abaxially sparsely tomentose, limb plane, margin short-fimbriate, basal fimbriae mostly tipped with tiny glands, posterior petal with claw slightly thicker, sometimes with a pair of glands near the limb, limb slightly broader than the 4 lateral ones; androecium with 5 stamens, filaments free or connate at base with adjacent filaments, fertile stamens 3, opposite anterior and posterior-lateral sepals, heteromorphic, filaments stout, glabrous, anthers rimose, glabrous or locules tomentose at apex, connective glandular; staminodes 2, opposite anterior-lateral sepals, homomorphic, filaments slender, antherode equalling or larger than anthers (reduced to an apical swelling in M. harleyi and M. lanata), glandular, pubescent (glabrous in M. harleyi); ovary superior, 3-carpellate, carpels syncarpic, the posterior 2 rotated so that all face the posterior petal, minutely puberulent, style 1, basal, straight, glabrous, borne low on inner face of anterior carpel, stigma terminal, truncate, held above anthers or at the same level at anthesis. Schizocarp with 3 mericarps, dorsal and lateral wings reduced to ribs or teeth, glabrous to velutine; carpophore absent. Chromosome number n = (20–)40.
The genus name honours the Brazilian botanist Dra Maria Candida Henrique Mamede (b. 1956), friend, colleague, and long-time contributor to the Brazilian Malpighiaceae.
Mamedea currently comprises seven accepted species (one threatened species; Suppl. material
≡ Aspicarpa boliviensis Nied., Meded. Rijks-Herb. 19: 72. 1913.
≡ Aspicarpa harleyi W.R.Anderson, Contr. Univ. Michigan Herb. 16: 55. 1987.
≡ Camarea pulchella Griseb. in Martius, Fl. Bras. 12(1): 105. 1858.
≡ Camarea lanata Chodat, Mém. Soc. Phys. Genève 31(2): 20. 1892 ≡ Aspicarpa schininii W.R.Anderson, Contr. Univ. Michigan Herb. 16: 59. 1987.
≡ Camarea salicifolia Chodat, Arch. Sci. Phys. Nat., sér. 3, 24: 500. 1890.
≡ Aspicarpa sericea Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 68. 1879.
≡ Aspicarpa uruguariensis Nied., Verzeichnis Vorles. Konigl. Akad. Braunsberg 1912/13: 62. 1912.
= Brittonella Rusby, Bull. Torrey Bot. Club 20: 429. 1893. Type species: Brittonella pilosa Rusby [= Mionandra camareoides Griseb.].
= Cordobia Nied., Verzeichnis Vorles. Konigl. Akad. Braunsberg 1912–13: 41. 1912. Type species: Cordobia argentea (Griseb.) Nied. [≡ Mionandra argentea Griseb.].
= Gallardoa Hicken, Physis (Buenos Aires) 2: 101. 1916. Type species: Gallardoa fischeri Hicken [≡ Mionandra fischeri (Hicken) R.F.Almeida]
Mionandra camareoides Griseb.
Mionandra comprises four currently accepted species (one threatened species; Suppl. material
Peixotoa glabra A.Juss.
Peixotoa comprises 29 currently accepted species (18 threatened species; Suppl. material
Philgamia hibbertioides Baill.
Philgamia comprises four currently accepted species (all threatened species; Suppl. material
≡ Fimbriaria A.Juss., Fl. Bras. Merid. (quarto ed.) 3(22): 63. 1833, nom. illeg., non Stackh. (1809), nec Nees ex Steud. (1824).
Fimbriaria elegans A.Juss. [= Schwannia mediterranea (Vell.) R.F.Almeida & M.Pell.].
Despite being the oldest available name for this genus, Fimbriaria A.Juss. is illegitimate for being a later homonym to Fimbriaria Stackh. (Rhodomelaceae, Rhodophyta). This is unaffected by the posterior rejection of Fimbriaria Stackh. against Odonthalia Lyngbye. Furthermore, even if this rejection made “Fimbriaria” available as a generic name, Fimbriaria Nees ex Steud. (Aytoniaceae, Marchantiophyta) still has priority over the Malpighiaceae name. Therefore, Schwannia is the earliest available name for this genus.
Schwannia comprises 14 currently accepted species (seven threatened species; Suppl. material
≡ Janusia christianeae W.R.Anderson, Contr. Univ. Michigan Herb. 16: 80. 1987.
≡ Janusia diminuta R.Sebast. & Mamede, Hoehnea 41(1): 121. 2014.
≡ Banisteria hexandra Vell., Fl. Flum.: 188. 1825.
≡ Banisteria mediterranea Vell., Fl. Flumin.: 191. 1829.
≡ Janusia occhionii W.R.Anderson, Contr. Univ. Michigan Herb. 16: 84. 1987.
≡ Schwannia paraensis R.Sebast. & Mamede, Hoehnea 41(1): 123. 2014.
≡ Janusia prancei W.R.Anderson, Contr. Univ. Michigan Herb. 16: 87. 1987.
≡ Janusia schwannioides W.R.Anderson, Contr. Univ. Michigan Herb. 15: 133–135, f. 14. 1982.
= Tricomariopsis Dubard, Compt. Rend. Hebd. Séances Acad. Sci. 145: 1190. 1907. Type species: Tricomariopsis madagascariensis Dubard (= Sphedamnocarpus dubardii R.Vig. & Humbert ex Arènes).
= Banisterioides Dubard & Dop, Rev. Gén. Bot. 20: 356. 1908. Type species: Banisterioides madagascariensis (Baker) Dubard & Dop (= Sphedamnocarpus multiflorus Nied.).
Sphedamnocarpus angolensis (A.Juss.) Oliv.
Sphedamnocarpus comprises ten currently accepted species (nine threatened species; Suppl. material
= Brachypterys A.Juss. in Deless., Icon. Sel. Pl. 3: 20. 1838. Type species: Brachypterys australis A.Juss. (= Stigmaphyllon paralias A.Juss.).
= Ryssopterys Blume ex A.Juss. in Deless., Icon. Sel. Pl. 3: 21. 1838. Type species: Ryssopterys timoriensis (DC.) Blume ex A.Juss. [≡ Stigmaphyllon timoriense (DC.) C.E.Anderson].
Stigmaphyllon auriculatum (Cav.) A.Juss.
Stigmaphyllon comprises 119 currently accepted species (60 threatened species; Suppl. material
The phylogenetics of Malpighiaceae has been the subject of at least 17 different studies based on plastid and nuclear markers over the last two decades. Eight of these studies focused on the family as a whole, trying to sample its main morphological or phylogenetic groups (
In fact, different lines of evidence, besides DNA, support our new classification system. Pollen grain morphology also recovered different pollen types in Malpighiaceae that characterise both subfamilies and most of the tribes recognized in our study (
Finally, the previously mentioned non-monophyly of Rhynchophora and Madagasikaria is a minor issue that will be easily solved by additional taxonomic and genomic sampling in future studies in the family. Additionally, the uncertain placement of Ectopopterys within tribe Malpighieae can only lead to its placement in another currently recognised tribe or to the proposition of a new tribe to accommodate this peculiar monospecific genus. Thus, the integrity and phylogenetic confidence of the new classification system proposed here for Malpighiaceae will remain strong, further advancing the taxonomic knowledge of this important family of flowering plants.
After over two decades of phylogenetic studies in Malpighiaceae and almost three centuries of taxonomic work, we finally have the proposition of the first classification system with a monophyletic recircumscription of all currently accepted genera and an updated species list for this family. A total of two subfamilies, 12 tribes, 72 genera, and 1,499 species are accepted in this study for Malpighiaceae worldwide as a solid basis for future systematic and taxonomic study in this family. Even though generic circumscriptions are now monophyletic and re-circumscribed, generic relationships within its most species-rich tribes (i.e., Gaudichaudieae, Hiptageae, Hiraeeae, and Malpighieae) still need further phylogenetic/omic studies for better statistical support and proposition of a subtribal system. Taxonomic revisions are urged for most genera of this family, especially the most species-rich ones [i.e., Heteropterys (166 spp.), Byrsonima (164), Bunchosia (93), Hiraea (81), Tetrapterys (56), Mascagnia (48), Hiptage (47), Acridocarpus (36), Glicophyllum (28), Aspicarpa (27), Aspidopterys (24), Triaspis (19), Niedenzuella (18), Schwannia (14), Bronwenia (13), Spachea (12), Sphedamnocarpus (10), Psychopterys (9), Carolus (8), Mamedea (7), Jubelina (6), Christianella (5), Adelphia (4), Excentradenia (4), Malpighiodes (4), Ptilochaeta (3), Brachylophon (2), Calcicola (2), Caucanthus (2), Echinopterys (2), Heladena (2), and Janusia (2)]. Finally, special attention is urgently needed to the taxonomy of the long-neglected and highly threatened African and Asian Malpighiaceae.
We would like to thank the staff of the visited herbaria for their support with herbarium specimens and images; Alexandre Sennikov, Jose Maria Cardiel and two anonymous reviewers for their constructive criticism on an earlier version of the manuscript; and Amaury Jr., A. Assis, A.C. Dal Col, A. Francener, A. Popovkin, C. Baez, C.C. Davis, C. Silva, E. Bidault, F. Farronay, F. Flores, F. Michelangeli, G. Cahyadi, G.A. Dettke, G. Shimizu, I. Specogna, Juvante’s, K. Souza, L.S.B. Calazans, M.R. Pace, N. Bigio, N. Rakotonirina, N. Singh, N. Taniguti, O.J.A. Ayala, P. Acevedo-Rodriguez, R. Goldenberg, S. Carnaham, and S.E. Martins for allowing us to use their beautiful photographs and line drawings in our study.
The authors have declared that no competing interests exist.
No ethical statement was reported.
RFA was sponsored by Programa de Desenvolvimento Científico e Tecnológico Regional CNPq/FAPEG (grants #317720/2021-0 and #202110267000867) and UEG/PrP, Termo de Fomento do Recurso Pró-Programa UEG 2022 (grant #21/2022).
Conceptualization: RFA, MOOP. Data curation: ILM, TAS. Formal analysis: HAD, RFA, MOOP. Funding acquisition: ILM. Methodology: RFA, HAD, MOOP. Resources: ILM. Software: HAD. Validation: MOOP. Writing - original draft: RFA. Writing - review and editing: HAD, TAS, MOOP, ILM.
Rafael F. de Almeida https://orcid.org/0000-0002-9562-9287
Isa L. de Morais https://orcid.org/0000-0001-8748-9723
Thais Alves-Silva https://orcid.org/0009-0001-0760-6019
Higor Antonio-Domingues https://orcid.org/0000-0001-9405-1930
Marco O. O. Pellegrini https://orcid.org/0000-0002-8783-1362
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Checklist of accepted species names of Malpighiaceae
Data type: xlsx
Morphological matrix including 31 scored and coded characters used in the phylogenetic optimization
Data type: xlsx