Malpighiaceae Juss., Gen. Pl.

de Almeida, Rafael F., de Morais, Isa L., Alves-Silva, Thais, Antonio-Domingues, Higor & Pellegrini, Marco O. O., 2024, A new classification system and taxonomic synopsis for Malpighiaceae (Malpighiales, Rosids) based on molecular phylogenetics, morphology, palynology, and chemistry, PhytoKeys 242, pp. 69-138 : 69-138

Malpighiaceae Juss., Gen. Pl. : 252. 1789 nom. cons.

Figs 2 , 3 , 4 , 5 , 6 , 7 , 8 , 9 , 10 , 11

Type species.

Malpighia glabra L.

Description.

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. 2 ). Roots fibrous or tuberous, generally becoming woody with age. Xylopodium present or not. Branches woody, rarely herbaceous, lenticelate or not. Stipules present, rarely absent, inter- or epipetiolar, minute to expanded, free to connate, pair of stipules free or connate, deciduous or persistent (Fig. 3 E – H ). Leaves opposite, decussate, rarely verticillate, ternate, subopposite or alternate (Fig. 3 A – D ), petiolate; petioles short to very long (Fig. 3 L – N ), circular, plane-convex to canaliculate (Fig. 3 I – K ) in cross-section, usually glandular (Fig. 3 O-S ); blades simple, entire, rarely lobed (Fig. 4 I – K ), usually glandular, margin plane to revolute (Fig. 4 O, P ), entire, sometimes dentate, crenate or lobulate, glabrous, ciliate or pubescent (Fig. 4 I – N ). Inflorescences solitary or compound, terminal to axillary, pedunculate, rarely sessile (Fig. 5 A ); flowers arranged in a 1 – multi-flowered cincinnus (Fig. 5 B ), cincinni opposite to alternate, usually pedunculate (Fig. 5 C ), rarely sessile, solitary or arranged in 2–5 - degrees of ramification into thyrses, corymbs, umbels, or dichasia (Fig. 6 ); leaves associated with the inflorescences similar to vegetative leaves but reduced in size (Fig. 6 D ); cincinni bract 1, minute to expanded, plane or concave, persistent, rarely deciduous, glandular or eglandular (Fig. 7 A – I ); bracteoles 2, opposite or alternate, usually inserted at the apex of peduncles, rarely subapical, medial or basal, minute to expanded, plane or concave, persistent, rarely deciduous, glandular or eglandular (Fig. 7 A – I ). Flowers chasmogamous, rarely cleistogamous, bisexual, rarely unisexual, zygomorphic, rarely actinomorphic, hypogynous, rarely perigynous (Fig. 7 Q ); pedicel well-developed, rarely absent or inconspicuous, straight, rarely circinate (Fig. 7 D – F ); sepals 5 (– 7), free to connate at base, imbricate in bud, rarely valvate, erect, rarely deflexed, apex erect to revolute, persistent in fruit, usually not accrescent in fruit, rarely accrescent, abaxially (0 –) 1–2 (– many) - glandular, sometimes the anterior sepal eglandular, rarely all sepals eglandular (Fig. 7 J – L ), glands multicellular, usually sessile (Fig. 8 C ), rarely pedunculate (Fig. 8 C ), secreting oil (Fig. 8 B ), rarely nectar (Fig. 8 A ); petals 5 (– 7), free, imbricate in bud, rarely valvate, clawed, patent (Fig. 8 O ) or deflexed (Fig. 8 J ), usually yellow (Fig. 8 F – J ), sometimes white (Fig. 8 C, E, K ), pink (Fig. 8 A, L, Y ), orange (Fig. 8 M ) or red (Fig. 8 T ), rarely green, lilac (Fig. 8 Z ) or purple, usually changing colour after pollination, the posterior petal usually differing from the 4 lateral ones in size, colour, shape and / or presence of glands, limb generally crumpled in bud, plane, concave or cucullate at anthesis, margin entire (Fig. 8 U ), erose (Fig. 8 V ), denticulate (Fig. 8 X ), fimbriate (Fig. 8 W ), or glandular-fimbriate (Fig. 8 Y ); androecium with (3–5 – 6 –) 10 stamens (Fig. 9 D – G ), in two whorls, fertile stamens (2–6 –) 10 (Fig. 9 E ), staminodes 0 (– 2–5; Fig. 9 F, G ), filaments free (Fig. 9 G ) or connate (Fig. 9 D, E ) at base, rarely connate up to the middle, short to long, glabrous (Fig. 9 E ), rarely pubescent (Fig. 9 F ), homo- (Fig. 9 E ) or heteromorphic (Fig. 9 D ), anthers basifixed (Fig. 9 A ), connectives minute or expanded (Fig. 9 A, B ), usually glandular (secreting non-volatile oils, rarely essential oils; Fig. 9 B ), with or without an apical projection, glabrous (Fig. 9 E ) to pubescent (Fig. 9 D ), thecae 2, parallel, rarely divergent at base and connivant at apex, apex free, rarely confluent, rimose (Fig. 9 A ), rarely poricidal; gynoecium superior, (1–2 –) 3 - carpelate (Fig. 9 H ), all fertile, rarely 1 abortive, carpels syncarpic, rarely basally syncarpic and apically apocarpic, locules 1 - ovulate, ovules pendulous, anatropous, styles (1–2 –) 3 (Fig. 9 L, M ), free, rarely connate, basal, lateral, subapical or apical, straight (Fig. 9 L ), curved (Fig. 9 O ) or lyrate (Fig. 9 N, P ), apex subulate (Fig. 9 K ), cylindrical (Fig. 9 L ), laterally flattened, truncate (Fig. 9 J ) or uncinate (Fig. 9 I ), rarely expanded (Fig. 9 N, P ), stigmas terminal (Fig. 9 L, M ) or lateral (i. e., facing the centre of the flower or the posterior petal; Fig. 9 I, N, P ), subulate (Fig. 9 K ), punctiform (Fig. 9 K ), capitate (Fig. 9 L ) or crateriform (Fig. 9 I ). Fruits dry (Fig. 10 O – W ) or fleshy (Fig. 10 A – N ), schizocarps (Fig. 10 O – W ), nuts (Fig. 10 B – D ) or drupes (Fig. 10 A, E – N ), glabrous or pubescent, mericarps (1–2 –) 3, indehiscent or splitting at maturity, smooth (Fig. 10 O – S ), setose (Fig. 10 T – W ), or winged (Fig. 11 A – M ), when winged mericarps with 1 dorsal (Fig. 11 A – E ) and / or 1 – several lateral (Fig. 11 F – M ) wings, free (Fig. 11 F – H, J – M ) or connate (Fig. 11 I ). Seeds 1 per locule (Fig. 10 A ), globose or ovoid (Fig. 10 A ), smooth (Fig. 10 A ) to rugose, without endosperm; embryo curved, bent or spiralled. Chromosome number n = 6–10.

Notes.

Malpighiaceae is here circumscribed with two subfamilies, 12 tribes, 72 genera, and 1,499 species accepted (Table 1 ; Suppl. material 1). From this total, 60 genera and 715 species are currently under some kind of extinction threat ( Bachman et al. 2024), representing 84.5 % of the accepted genera and 47.82 % of the species in our study (Suppl. material 1). Most of Malpighiaceae ’ s diversity is confined to the American continent, with 55 genera (53 endemic) and 1,274 species (1,272 endemic), and just 15 genera (13 endemic) and 125 species (123 endemic) in Africa, seven genera (four endemic) and 84 species (77 endemic) in Asia, and four genera (all not endemic) and 21 species (19 endemic) in Oceania (Suppl. material 1). Most threatened species are found in the Americas (564 species), with 86 threatened species in Africa, 57 in Asia, and 12 in Oceania (Suppl. material 1). Five African genera, more specifically endemic to Madagascar (i. e., Digoniopterys , Madagasikaria , Microsteira , Philgamia , and Rhynchophora ) stand out, with all their current accepted species under some kind of extinction threat (Suppl. material 1). Mcvaughia was the only American genus to present all its species under some kind of extinction threat (Suppl. material 1). Malpighia was the American genus with most species under some kind of extinction threat (Suppl. material 1). Hiptage was the genus with highest number of threatened species in Asia, and Stigmaphyllon was the most threatened in Oceania (Suppl. material 1).

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.

Figures

Figure 2. 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).

Figure 3. 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).

Figure 4. 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).

Figure 5. Inflorescence architecture of Malpighiaceae A inflorescence evolution in Malpighiaceae according to Anderson (1981) B 1 - flowered cincinnus of Niedenzuella lasiandra C 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).

Figure 6. 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 (1 st order inflorescence), arranged in dichasia (2 nd order inflorescence), arranged in a thyrse (3 rd 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).

Figure 7. 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).

Figure 8. 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).

Figure 9. 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).

Figure 10. 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).

Figure 11. 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).