Eight Segregates from the Neotropical Genus Mascagnia (Malpighiaceae)

Home , Adelphia (plant), Alicia (plant), Byrsonima, Carolus (plant), Christianella, Heladena

William R. Anderson University of Michigan Herbarium, 3600 Varsity Drive, Ann Arbor, Michigan 48108-2287, U.S.A. [email protected]

ABSTRACT . In the light of morphological and molec- nilles) W. R. Anderson, N. caracasana W. R. ular evidence that the genus Mascagnia is poly- Anderson, N. castanea (Cuatrecasas) W. R. Anderson, phyletic, eight segregate genera are described, N. glabra (Sprengel) W. R. Anderson, N. leucosepala discussed, and illustrated, the necessary new combi- (Grisebach) W. R. Anderson, N. lucida (A. Jussieu nations are proposed, and five new species are in A. St.-Hilaire) W. R. Anderson, N. mater-dei described (Amorimia camporum, A. kariniana, A. (Cuatrecasas) W. R. Anderson, N. metensis (Cuatre- septentrionalis, A. velutina, and Niedenzuella caraca- casas) W. R. Anderson, N. mogoriifolia (A. Jussieu in sana). Keys to genera and species are provided. The A. St.-Hilaire) W. R. Anderson, N. multiglandulosa genera and species are: Adelphia W. R. Anderson [A. (A. Jussieu) W. R. Anderson, N. peruviana (Niedenzu) hiraea (Gaertner) W. R. Anderson, A. macrophylla W. R. Anderson, N. poeppigiana (A. Jussieu) W. R. (Rusby) W. R. Anderson, A. mirabilis (W. R. Anderson, N. sericea (A. Jussieu in A. St.-Hilaire) W. Anderson) W. R. Anderson, A. platyrachis (Triana & R. Anderson, N. stannea (Grisebach) W. R. Anderson, Planchon) W. R. Anderson]; Aenigmatanthera W. R. N. suaveolens (A. Jussieu) W. R. Anderson, N. Anderson [A. doniana (Grisebach in Martius) W. R. warmingiana (Grisebach) W. R. Anderson]. Lecto- Anderson, A. lasiandra (A. Jussieu) W. R. Anderson]; types are designated for the following names: Alicia W. R. Anderson [A. anisopetala (A. Jussieu in Heladena hassleriana Niedenzu in Chodat & Hassler, A. St.-Hilaire) W. R. Anderson, A. macrodisca (Triana Hiraea anisopetala A. Jussieu in A. St.-Hilaire, Hiraea & Planchon) W. R. Anderson]; Amorimia W. R. glabra Sprengel, Hiraea heteropetala A. Jussieu, Anderson [A. amazonica (Niedenzu) W. R. Anderson, Hiraea multiflora Grisebach, Hiraea poeppigiana A. A. camporum W. R. Anderson, A. concinna (C. V. Jussieu, Hiraea pubiflora A. Jussieu in A. St.-Hilaire, Morton) W. R. Anderson, A. exotropica (Grisebach in Hiraea renidens A. Jussieu, Hiraea rigida A. Jussieu Martius) W. R. Anderson, A. kariniana W. R. in A. St.-Hilaire, Hiraea sericea A. Jussieu in A. St.- Anderson, A. maritima (A. Jussieu) W. R. Anderson, Hilaire, Malpighiodes Niedenzu, Malpighiodes spru- A. pubiflora (A. Jussieu in A. St.-Hilaire) W. R. ceana Niedenzu, Mascagnia sect. Pleuropterys Grise- Anderson, A. rigida (A. Jussieu in A. St.-Hilaire) W. bach in Martius, Mascagnia subsect. Sericopetalis R. Anderson, A. septentrionalis W. R. Anderson, A. Niedenzu, Mascagnia coriacea Grisebach in Martius, velutina W. R. Anderson]; Carolus W. R. Anderson Mascagnia exotropica Grisebach in Martius, Mascag- [C. anderssonii (W. R. Anderson) W. R. Anderson, nia jamaicensis Urban & Niedenzu, Mascagnia multi- C. chasei (W. R. Anderson) W. R. Anderson, C. glandulosa Niedenzu in Chodat & Hassler, Mascagnia chlorocarpus (A. Jussieu) W. R. Anderson, C. dukei parnahybensis Glaziou, Mascagnia sericans Niedenzu (Cuatrecasas & Croat) W. R. Anderson, C. renidens in Chodat & Hassler, Tetrapterys fraxinifolia A. (A. Jussieu) W. R. Anderson, C. sinemariensis Jussieu, Tetrapterys guilleminiana A. Jussieu, Tetra- (Aublet) W. R. Anderson]; Christianella W. R. pterys lancifolia A. Jussieu, Tetrapterys ligustrifolia Anderson [C. glandulifera (Cuatrecasas) W. R. Niedenzu, Tetrapterys lucida A. Jussieu in A. St.- Anderson, C. mesoamericana (W. R. Anderson) W. Hilaire, Tetrapterys martiana Niedenzu, Tetrapterys R. Anderson, C. multiglandulosa (Niedenzu in Chodat mogoriifolia A. Jussieu in A. St.-Hilaire, Tetrapterys & Hassler) W. R. Anderson, C. paludicola (W. R. multiglandulosa A. Jussieu, Tetrapterys punctulata A. Anderson) W. R. Anderson, C. surinamensis (Koster- Jussieu in A. St.-Hilaire. mans) W. R. Anderson]; Malpighiodes Niedenzu [M. Key words: Adelphia, Aenigmatanthera, Alicia, bracteosa (Grisebach in Martius) W. R. Anderson, M. Amorimia, Carolus, Christianella, Malpighiaceae, guianensis (W. R. Anderson) W. R. Anderson, M. Malpighiodes, Mascagnia, Niedenzuella. leucanthele (Grisebach in Martius) W. R. Anderson, M. liesneri (W. R. Anderson) W. R. Anderson]; and Almost 20 Neotropical genera of Malpighiaceae Niedenzuella W. R. Anderson [N. acutifolia (Cava- produce samaras with lateral wings. The first such

NOVON 16: 168–204. PUBLISHED ON 26 JULY 2006. Volume 16, Number 2 Anderson 169 2006 Segregates from Mascagnia

genus to be described was Triopterys L. (1753), a small within Mascagnia as possible. Another former stu- West Indian group with Y-shaped samaras. Next came dent, Charles Davis, is now continuing that collabo- Hiraea Jacquin (1760), which had two-winged butter- ration. The resulting papers (Cameron et al., 2001; fly-shaped samaras, followed by Tetrapterys Cava- Davis et al., 2001, 2002) have provided dramatic nilles (1790) with an X-shaped samara, and then proof that Mascagnia sensu Niedenzu is indeed highly Mascagnia (Bertero ex DC.) Colla (1824), with an polyphyletic and must be divided up. In this paper I orbicular samara. Because the samara of Triopterys initiate that process by describing eight segregate was so distinctive, few used that name for the many genera and proposing the necessary new combinations lateral-winged species that soon began to be de- so that those names will be available for floristic scribed; the next-oldest name Hiraea was the one treatments now in preparation. employed for most species that did not fall into The morphological terminology used in the keys Triopterys or the equally distinctive Tetrapterys. and descriptions below conforms to that discussed in However, Hiraea is itself distinctive, with characters Anderson (1981). of the stipules and inflorescence that make it easy to The synonymies below are somewhat incomplete. segregate from the great diversity in this group of For fuller synonymies, see my database (Malpighia- plants. Adrien de Jussieu (1843) retained a broad ceae Nomenclature) on the website of the University concept of Hiraea, but he divided his Hiraea into of Michigan Herbarium (http://herbarium.lsa.umich. sections Hiraea and Mascagnia. Grisebach (1858) edu/). took the next step, restricting Hiraea to the species The following key to genera accounts for most of the treated as section Hiraea by Jussieu and placing all species Niedenzu treated under Mascagnia in his the remaining species in a broad Mascagnia. 1928 monograph. The numbers in parentheses Niedenzu (1928) followed Grisebach, with a restricted following leads refer to the species in Niedenzu’s Hiraea and an inclusive Mascagnia, attempting to treatment. Two of Niedenzu’s species are not deal with the variation in the latter by dividing the accounted for in this key: (1) his 35, Mascagnia genus into subgenera and sections. When I started to fluminensis (Grisebach) Grisebach in Martius, is now study the family at the New York Botanical Garden in referable to Heteropterys fluminensis (Grisebach) W. 1971, I soon realized that Niedenzu’s Mascagnia was R. Anderson; Niedenzu did not confuse Heteropterys an omnium-gatherum that could only be characterized HBK and Mascagnia, but he did not see the as all the Neotropical Malpighiaceae with lateral- diagnostic fruits of that species; (2) his Sp. Incert. 6, winged samaras, 10 fertile stamens, and internal Mascagnia spicigera (Turczaninow) Niedenzu in stigmas that do not fit into any of the well-delineated Engler, is referable to Gaudichaudia cynanchoides segregates such as Triopterys, Hiraea, Tetrapterys, HBK. Mascagnia s. str. is keyed, but not treated Jubelina A. Jussieu, and Mezia Niedenzu. More than further in this paper, nor are the genera Callaeum, a third of the species Niedenzu assigned to Mascagnia mentioned above, and Clonodia Grisebach in Martius, (his species 3–17) were morphologically coherent and which is related to Heteropterys. included the type of Mascagnia, so it was clear that a large bloc of species could be segregated relatively KEY TO THE GENERA INCLUDED UNDER MASCAGNIA BY NIEDENZU easily and that group would keep the name Mascag- (1928) nia. However, many species included by Niedenzu 1a. Lamina bearing glands in abaxial surface, between and others described since 1928 did not hold together midrib and margin. in one or even a few equally obvious segregates. It 2a. Stipules interpetiolar. seemed inevitable that if one started to split 3a. Lateral wings of samara membranous, Mascagnia, the result would be the recognition of as with strongly reticulate veins, in most species continuous at base and 6 many as a dozen genera, and I did not want to do that circular in outline (cleft to nut at base until I was sure those groups were not a diverse but in 2 species, chartaceous and strongly monophyletic clade. dissected in 2 species, completely lost in One of the most divergent groups was treated by 1 species); petals mostly glabrous (very Johnson (1986) under the name Callaeum Small. For rarely bearing a few abaxial hairs); fruit subtended by a smooth 3-lobed disc the rest, I decided to seek help from the then- (Niedenzu, 1928: species 3–5, 7–10, emerging field of molecular systematics. When, in the 12–14, 16, 17, Spp. Incert. 1, 5, 7) . . . late 1980s, I persuaded my former student, Mark ...... Mascagnia Chase, to initiate with me a molecular survey of 3b. Lateral wings of samara chartaceous, with many fine parallel veins, cleft to Malpighiaceae, one of my principal goals was to nut at base and butterfly-shaped or (in resolve the problem of Mascagnia, so I made a point of 1 species) continuous at base; petals including in our study set as many of the groups abaxially 6 densely hairy; fruit not 170 Novon

subtended by a smooth disc (23, 30, 31, stigma transversely ex- Sp.Incert.2)...... IV.Amorimia panded, elliptical; stip- 2b. Stipules epipetiolar or lacking. ules epipetiolar (36– 4a. Flowers borne ultimately in umbels or 39)...... Callaeum corymbs of 4(–8), or in pairs; petals 10b. Anthers all 6 alike; yellow turning red in age; stigmas stigma circular or lat- terminal or nearly so (21, 22) ...... erally compressed; stip- ...... VII. Malpighiodes ules interpetiolar or epi- 4b. Flowers borne ultimately in short to petiolar. elongated pseudoracemes; petals lilac, 11a. Petals abaxially pink, white, or some combination of those densely sericeous; colors; stigmas internal. anthers glabrous 5a. Sepals 6 completely concealing or bearing a few petals throughout enlargement of hairs; stipules in- bud, strongly recurved in anthesis; terpetiolar (often petals abaxially unwinged and deciduous) (25, 28, densely tomentose or subsericeous; 29)...... V.Carolus mericarps with membranous or 11b. Petals glabrous or chartaceous lateral wings well de- bearing a few hairs; veloped(18,19)...... III.Alicia anthers abaxially 5b. Sepals leaving petals visible during sericeous; stipules enlargement of bud, appressed in borne on petiole, anthesis; petals abaxially winged at or slightly above and glabrous or bearing a few hairs; the base, persistent mericarps with wings replaced by (24, Sp. Incert. 4) irregular coriaceous outgrowths . . II. Aenigmatanthera (1,2,32)...... Clonodia 1b. Lamina bearing glands on or embedded in margin, or eglandular. I. Adelphia W. R. Anderson, gen. nov. TYPE: 6a. Sepals bearing a row of small marginal glands, Adelphia hiraea (Gaertner) W. R. Anderson. these sessile or subsessile to long-stalked, clavate or capitate (20) ...... VI.Christianella Lianae lignosae; petiolus plerumque prope apicem 6b. Sepals without marginal glands. biglandulosus glandulis protuberantibus; lamina eglandulosa 7a. Sepals longer than petals in bud, imbri- vel margine in dimidio distali glandulis minimis instructa; cated and concealing petals until anthe- stipulae interpetiolares; inflorescentia pseudoracemus vel sis or in a few species separating to panicula elongata floribus plerumque 2, 3, vel 4 in quoque expose petals in enlarging bud, strongly nodo portatis; pedunculi floriferi bene evoluti; 1 bracteola revolute in anthesis (26, 27) ...... cujusque paris eglandulosa, altera uniglandulifera glandulo ...... VIII. Niedenzuella excentrico; pedicellus in alabastro parum circinatus; sepala 7b. Sepals much exceeded by petals in per anthesin plusminusve appressa; petala glabra, 4 lateralia enlarging bud, appressed or recurved lutea, posticum luteum vel luteum rubrumque, fimbriatum; only at apex in anthesis. carpella fere distincta in ovario; styli arcuati vel lyrati, 8a. Shrubs, never twining; flowers borne stigmate introrso, apice dorsaliter truncati, apiculati, vel singly or in umbels of 2–4 in axils of breviuncinati; samara alis lateralibus apice basique usque ad full-sized leaves; fruit subtended by nucem incisis (alis lateralibus rudimentariis in A. mirabili), a smooth 3-lobed disc (6, 11, 15) nuce subglobosa...... Mascagnia s.l. 8b. Woody vines, occasionally described as shrubby; flowers many Woody vines. Petiole usually biglandular above in axillary or terminal inflores- middle, the glands when present protuberant, usually cences, often pseudoracemes or large and knoblike; lamina eglandular or bearing very panicles; fruit not subtended by small marginal glands, often drawn out into vascular- a smooth disc. ized cilia, on distal half, especially near apex; tertiary 9a. One bracteole of each pair bearing 1 large eccentric gland; veins weakly or not scalariform; stipules very small, petiole usually bearing 2 large triangular, borne on stem beside base of petiole. knoblike glands near apex, Inflorescence an open, elongated, axillary or terminal sometimes eglandular; styles 6 panicle or pseudoraceme with the flowers borne singly bowed or lyrate (33, 34, Sp. Incert. 3) ...... I. Adelphia or (most often) in 2s, 3s, or 4s; floriferous bracts 9b. Both bracteoles eglandular; eglandular; floriferous peduncle well developed; petiole eglandular or bearing bracteoles borne at apex of peduncle, one of each glands, the glands if present pair bearing 1 bulging eccentric abaxial gland; not especially large; styles straight or bowed. pedicel slightly circinate in bud. Sepals ovate or 10a. Posterior 3 anthers small- broadly rounded, separating to expose petals during er than the other 7; enlargement of bud, 6 appressed in anthesis, the Volume 16, Number 2 Anderson 171 2006 Segregates from Mascagnia

lateral 4 or all 5 abaxially biglandular; corolla clade. If so, we can reconsider these genera then; for bilaterally symmetrical, the posterior petal long- now it seems best to recognize these very different- fimbriate and strongly differentiated from the lateral looking groups as separate genera. 4; petals yellow or the posterior yellow and red, Adelphia comprises four species of the West Indies, glabrous; androecium bilaterally symmetrical; sta- Mesoamerica, and western South America. mens 10, all fertile, glabrous; filaments distinct or only slightly connate at very base, longer opposite KEY TO THE SPECIES OF ADELPHIA sepals, shorter opposite petals; anthers 6 alike, 1a. Petiole of larger leaves (20–)25–60 mm long; mostly larger on longer filaments; carpels nearly lamina of larger leaves (21–)23–30 6 (11–)13– distinct in ovary; styles inserted low on ventral face of 19 cm, eglandular; samara (55–)60–80 mm wide, carpels, 6 bowed or lyrate, the anterior mostly shorter 6 orbicular, or the lateral wings rudimentary; and less strongly bent than the posterior 2; apex of posterior petal proximally glandular-fimbriate, distally eglandular-fimbriate; sepals abaxially styles with a large internal stigma and dorsally densely and evenly appressed-tomentose. truncate, apiculate, or bearing a hook up to 0.5 mm 2a. Samara with the lateral wings rudimentary, 1– long. Fruit dry, breaking apart into samaras separating 2 mm wide, the dorsal wing larger than the from a short pyramidal torus; samara orbicular to laterals;AmazonianPeru...... 3.A. mirabilis butterfly-shaped with lateral wings dominant, borne 2b. Samara with the lateral wings well developed, 25–40 mm wide, 50–72 mm high, the dorsal on upper edge of nut, membranous or chartaceous wing much smaller than the laterals; Bolivia with many fine strongly parallel veins, cleft to nut at andAmazonianPeru...... 2.A. macrophylla base and apex, the margin undulate or erose (lateral 1b. Petiole of larger leaves 6–15 (rarely –27) mm wings rudimentary and the small dorsal wing domi- long; lamina of larger leaves 9–19(–26) 6 4–10 (–12.5) cm, eglandular or bearing very small nant in A. mirabilis); dorsal wing smaller and distinct marginal glands, often drawn out into vascular- from lateral wings at both apex and base; intermediate ized cilia, on distal half, especially near apex; winglets absent; ventral areole circular, partially samara 15–55 mm wide, 6 butterfly-shaped; surrounded by an irregular callose thickening. posterior petal eglandular-fimbriate all around margin; sepals abaxially glabrous or proximally The molecular studies published to date (Cameron sericeous, ciliate on margin. 3a. Lateral wings of samara 5–10 mm wide, 10– et al., 2001: 1856; Davis et al., 2001: 1839, and 2002: 25 mm high, coriaceous; Pacific coast of 6835) consistently place this genus [represented by Colombia...... 4.A. platyrachis Mascagnia hippocrateoides (Triana & Planchon) 3b. Lateral wings of samara 12–25 mm wide, 15– Niedenzu] in a clade with Excentradenia W. R. 40 mm high, membranous; Jamaica, southeastern Mexico, Central America, and north- Anderson, Hiraea, an undescribed genus [represented western South America ...... 1.A. hiraea by Mascagnia dipholiphylla (Small) Bullock], and (in Davis et al., 2002) Lophopterys A. Jussieu in 1. Adelphia hiraea (Gaertner) W. R. Anderson, Delessert, but mostly without strong support or good comb. nov. Basionym: Triopterys hiraea Gaertner, resolution of the relationships among the five genera. Fruct. 2: 169, pl. 116. 1790. Mascagnia hiraea The morphology suggests to me that this genus is (Gaertner) Fawcett & Rendle, Fl. Jamaica 4: 240. probably sister to Excentradenia, hence the name 1920. TYPE: Jamaica. W. Wright s.n. (holotype, Adelphia, from the Greek word for sister, adelphe. BM Banks Hb.). Figure 1. Adelphia and Excentradenia are alike in most characters, including the eccentric bracteole glands Hiraea hippocrateoides Triana & Planchon, Ann. Sci. Nat. for which Excentradenia was named (Anderson, 1997). Bot., Se´r. 4, 18: 328. 1862. Mascagnia hippocrateoides However, they have dramatically different inflores- (Triana & Planchon) Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum Hosianum Braunsberg 3: 24. 1908. cences, that of Excentradenia being a short, compact TYPE: Colombia. Vicinity of Anapoima, J. Triana s.n. raceme of 3–7(–9) umbels of four flowers each, with (holotype, COL 17026; isotype, BM [MICH, WRA neg. the floriferous peduncles very short to absent. The 81-2-10]). sepals in Excentradenia are elongated, and revolute in Mascagnia jamaicensis Urban & Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum Hosianum Braunsberg 3: 25. anthesis; the tertiary leaf veins are strongly scalari- 1908. TYPE: Jamaica. W. Harris 9862 (lectotype, form; and except for a few populations of E. designated here, NY; duplicates, BM, K). adenophora (Sandwith) W. R. Anderson the lateral Of the five syntypes, all from Jamaica, Harris 9862 wings of the samara are continuous at the base. The is the most complete and representative. inflorescence in Excentradenia probably resulted from Mascagnia excelsa Standley & Steyermark, Publ. Field condensation of an ancestral inflorescence resembling Mus. Nat. Hist., Bot. Ser., 23: 168. 1944. TYPE: that of Adelphia, so it is possible that Excentradenia Guatemala. Izabal: Rı´o Dulce, J. A. Steyermark 39456 will eventually be found to nest within the Adelphia (holotype, F).

Volume 16, Number 2 Anderson 173 2006 Segregates from Mascagnia

3. Adelphia mirabilis (W. R. Anderson) W. R. androecium radially symmetrical; stamens 10, all Anderson, comb. nov. Basionym: Mascagnia fertile; filaments connate at base, straight, alike but mirabilis W. R. Anderson, Contr. Univ. Michigan somewhat longer opposite sepals than opposite petals, Herb. 14: 19. 1980. TYPE: Peru. San Martı´n: glabrous or abaxially sparsely to densely sericeous, Puente Colombia to Shapaja, T. C. Plowman especially distally; anthers alike, abaxially 6 densely 6019 (holotype, MICH). white-sericeous on the dark connective, at least proximally; carpels connate their whole length in 4. Adelphia platyrachis (Triana & Planchon) W. R. ovary; styles stout, straight, erect to somewhat Anderson, comb. nov. Basionym: Hiraea platy- spreading distally, with large terminal or internal rachis Triana & Planchon, Ann. Sci. Nat. Bot., circular or laterally compressed stigmas, dorsally Se´r. 4, 18: 328. 1862. Mascagnia hippocrateoides rounded at apex. Fruit dry, breaking apart into var. platyrachis (Triana & Planchon) Niedenzu, samaras separating from a short pyramidal torus; Arbeiten Bot. Inst. Ko¨nigl. Lyceum Hosianum samara butterfly-shaped with lateral wings dominant, Braunsberg 3: 24. 1908. Mascagnia platyrachis chartaceous with many fine strongly parallel veins, (Triana & Planchon) Cuatrecasas, Webbia 13: trapezoidal, cleft to nut at base and apex, the margin 374. 1958. TYPE: Colombia. Choco´: Isthmus of coarsely dentate or lobed, occasionally deeply and San Pablo, J. Triana s.n. (holotype, COL not irregularly lobed; dorsal wing small, distinct at base seen; isotypes, BM, G, K, P). and apex, extended forward at apex between lateral wings; raised ribs or irregular outgrowths present II. Aenigmatanthera W. R. Anderson, gen. nov. between dorsal and lateral wings; ventral areole ovate. TYPE: Aenigmatanthera lasiandra (A. Jussieu) The name Aenigmatanthera comes from the Latin W. R. Anderson. words for riddle or mystery (aenigma) and anther Lianae lignosae vel frutices; lamina eglandulosa vel (anthera). It refers to the odd stamens and especially aliquot glandulis minutis marginalibus instructa; stipulae the anthers, one of the important differences distin- minutae prope basim petioli portatae; bracteae bracteolaeque guishing this genus from Niedenzuella W. R. Ander- parvae eglandulosae; sepala quam petala multo breviora in alabastro; petala lutea, glabra vel abaxialiter paucipilifera; son. filamenta abaxialiter glabra vel sparsim vel dense sericea; I have had great difficulty in placing these two antherae abaxialiter plus minusve sericea; styli recti et erecti closely related species. The combination of marginal vel distaliter recurvati, stigmate terminali vel introrso; leaf glands, epipetiolar stipules, short sepals leaving samara alis lateralibus apice basique usque ad nucem incisis. the nearly glabrous yellow petals well exposed in the enlarging bud, and a butterfly-shaped samara ex- Woody vines, occasionally described as shrubs. cludes their assignment to any of the other genera Leaves opposite; petiole eglandular or biglandular at recognized here. They most resemble Niedenzuella, or above middle; lamina eglandular or bearing several which has decussate pseudoracemes, marginal leaf tiny glands embedded in the margin; stipules very glands, tiny epipetiolar stipules, and yellow petals small (sometimes apparently lacking), triangular, that are glabrous or thinly sericeous. Moreover, the borne on petiole at or slightly above base, persistent. samaras of Aenigmatanthera lasiandra sometimes Inflorescence an axillary or terminal pseudoraceme show irregular dissection of the lateral wings, as in or, more commonly, a panicle of pseudoracemes, the Niedenzuella stannea (Grisebach) W. R. Anderson. flowers 6 decussate, sometimes crowded distally to Grisebach also recognized this similarity, treating A. form a corymb or umbel; bracts and bracteoles lasiandra as a variety of Tetrapterys mogoriifolia A. eglandular, short, narrowly triangular, 6 spreading, Jussieu in A. St.-Hilaire [5 Niedenzuella mogoriifolia persistent or deciduous in fruit, the bracteoles borne (A. Jussieu in A. St.-Hilaire) W. R. Anderson]. at or somewhat below apex of peduncle; floriferous However, the sepals in Niedenzuella are very different peduncle developed, much shorter than pedicel. from those of Aenigmatanthera. They are papery and Sepals triangular or ovate, only moderately exceeding elongated, much exceeding the glands, being imbri- the glands, much exceeded by the petals in enlarging cated over the petals during much of the enlargement bud, appressed in bud, appressed to slightly recurved of the bud, then becoming revolute in anthesis. The at apex in anthesis, the lateral 4 abaxially biglandular anthers in Niedenzuella are short and glabrous or bear and the anterior eglandular or all 5 biglandular; a few hairs on the locules, not elongated and densely corolla bilaterally symmetrical, the posterior petal sericeous on the connective as in Aenigmatanthera. erect, its claw thicker than in lateral petals; petals The carpels in Niedenzuella are connate only at the small, yellow, glabrous or bearing a few straight base and distally distinct, whereas in Aenigma- appressed hairs abaxially in center, shallowly erose; tanthera they are connate their whole length. Another 174 Novon

genus that resembles Aenigmatanthera is Carolus W. 259. 1840. Tetrapterys mogoriifolia var. lasian- R. Anderson, but that has interpetiolar stipules, dra (A. Jussieu) Grisebach in Martius, Fl. Bras. densely sericeous petals, and glabrous or subglabrous 12(1): 81. 1858. Mascagnia lasiandra (A. anthers. There is still no published molecular Jussieu) Niedenzu, Verz. Vorles. Ko¨nigl. Lyceum evidence on the relationships of Aenigmatanthera. Hosianum Braunsberg 1909/10: 56. 1909. Unless and until DNA sequences indicate a different TYPE: Brazil. Mato Grosso: Cuiaba´, Mart. Herb. disposition of these two species, it seems best to Fl. Bras. s.n. [657] (holotype, P; isotypes, G, L]. recognize their unique combination of characters by Figure 2. giving them generic status. Hiraea nitens S. Moore, Trans. Linn. Soc. London, Ser. 2 Bot., Lowrie (1982: 123–124) noted the peculiarity of the 4: 328. 1895. Mascagnia nitens (S. Moore) Niedenzu in pollen in this group of plants: ‘‘Mascagnia lasiandra is Engler, Pflanzenr. IV, 141: 123. 1928. TYPE: Brazil. Mato Grosso: Serra da Chapada, S. Moore 74 (holotype, remarkable among clonodioids for its regularity: BM). perfectly colpoidal, ora always centered in colpi, colpi always perpendicular to the equator, no Aenigmatanthera lasiandra is common in northern branching or fusing of the colpi to speak of. I would and eastern Bolivia (Nee 31847, MICH) and south- place this pollen type very close to byrsonimoids, central Brazil, and rare in southern Amazonas, Brazil especially Mcvaughia which I find to be virtually (Prance et al. 14200, MICH), and Para´ (Maciel et al. indistinguishable from it.’’ Mcvaughia W. R. Ander- 782, MICH). son has four broad fusiform colpi; both species of Aenigmatanthera have four or five similar colpi. III. Alicia W. R. Anderson, gen. nov. TYPE: Alicia Aenigmatanthera is a genus of two South American anisopetala (A. Jussieu in A. St.-Hilaire) W. R. species. Anderson. Hiraea sect. Hiraeostachys Grisebach, Linnaea 13: 242. KEY TO THE SPECIES OF AENIGMATANTHERA 1839. LECTOTYPE, designated by Morton (1968: 320): Hiraea anisopetala A. Jussieu in A. St.-Hilaire 1a. Lamina of larger leaves 3–9 6 1–4 cm, initially [5 Alicia anisopetala (A. Jussieu in A. St.-Hilaire) W. loosely sericeous, soon glabrate or persistently R. Anderson]. sericeous on midrib; pseudoracemes always con- Mascagnia subsect. Sericopetalis Niedenzu, Arbeiten Bot. densed to umbels or very tight corymbs of 2–8 Inst. Ko¨nigl. Lyceum Hosianum Braunsberg 3: 15. (mostly 4–6) flowers; filaments abaxially sparsely 1908. LECTOTYPE, designated here: Mascagnia sericeous to glabrate; anthers 1–1.5 mm long, anisopetala (A. Jussieu in A. St.-Hilaire) Grisebach in straight in anthesis, abaxially 6 densely sericeous Martius [5 Alicia anisopetala (A. Jussieu in A. St.- proximally, less hairy distally; northeastern Brazil Hilaire) W. R. Anderson]. (Maranhaõ and Ceara´)...... 1.A. doniana 1b. Lamina of larger leaves 8–16(–21) 6 (3–)4–9 Lianae lignosae; lamina in pagina abaxiali aliquot (–11) cm, soon glabrate; pseudoracemes mostly 6 glandulis parvis impressis instructa; stipulae parvae epipe- elongated (i.e., with measurable internodes be- tiolares prope basim petioli portatae; sepala valvata, 4 tween pairs of flowers), mostly comprising 6–14 lateralia biglandulifera; petala alba, alba et rosea, rosea, flowers; filaments abaxially densely and persis- vel lilacina, abaxialiter dense tomentosa vel subsericea; tently sericeous distally; anthers 1.5–2 mm long, stamina glabra, inter se similaria; styli recti, inter se recurved in anthesis, abaxially densely and similares, apice dorsaliter rotundati, truncati, vel apiculati; persistently sericeous their whole length; Bolivia samara alis lateralibus apice distinctis, basi plerumque and south-central Brazil, rarely in southern continuis. Amazonas and Para´,Brazil...... 2.A. lasiandra Woody vines. Petiole bearing 2–4(–8) small glands in 2 rows; lamina bearing few to many small glands 1. Aenigmatanthera doniana (Grisebach in Mar- impressed in abaxial surface in a row parallel to but tius) W. R. Anderson, comb. nov. Basionym: set in from the margin; stipules minute, triangular, Mascagnia doniana Grisebach in Martius, Fl. borne on petiole at or slightly above base. In- Bras. 12(1): 94. 1858. TYPE: Brazil. Maranhaõ, florescence a terminal and lateral panicle with the G. Don 85 (holotype, BR). flowers borne decussate or distally irregularly in short to elongated pseudoracemes; floriferous peduncle well This species in known only from northeastern Brazil developed; bracteoles borne between middle and apex [Maranhaõ and Ceara´ (Fernandes & Matos s.n. [EAC of peduncle or near apex. Sepals valvate, completely 11640], MICH)]. concealing petals during enlargement of bud, recurved in anthesis, the lateral 4 biglandular, the 2. Aenigmatanthera lasiandra (A. Jussieu) W. R. anterior usually eglandular; corolla bilaterally sym- Anderson, comb. nov. Basionym: Hiraea lasian- metrical to almost radial; petals white, white and pink, dra A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, 13: pink, or lilac, spatulate, abaxially unwinged and

176 Novon

samaras separating from a short pyramidal torus; 1. Alicia anisopetala (A. Jussieu in A. St.-Hilaire) samara suborbicular or transversely elliptical with W. R. Anderson, comb. nov. Basionym: Hiraea lateral wings dominant, membranous or chartaceous anisopetala A. Jussieu in A. St.-Hilaire, Fl. Bras. with distal looping anastomoses, cleft to nut at apex, Merid. 3: 15. 1832 [1833]. Mascagnia anisope- usually continuous at base (rarely cleft to nut at base), tala (A. Jussieu in A. St.-Hilaire) Grisebach in the margin usually entire or undulate; dorsal wing Martius, Fl. Bras. 12(1): 95. 1858. TYPE: Brazil. small, distinct at apex, distinct at base or very Near Sebastiano´polis [Rio de Janeiro], A. St.- abruptly confluent with lateral wing; ventral areole Hilaire Catal. D no. 766 (lectotype, designated ovate. Lombello and Forni-Martins (2002: 246) here, P [MICH, WRA neg. 81-21-19]; isotypes, reported chromosome counts of n 5 30 and 2n 5 P). Figure 3. 60 for Alicia anisopetala (as Mascagnia anisopetala); I have not had the opportunity to verify the identifi- Hiraea macrocarpa Chodat, Arch. Sci. Phys. Nat. III 24: 500. cation of their voucher. 1890. TYPE: Paraguay. Asuncio´n, B. Balansa 2408 (holotype, G [F neg. 24285]; isotypes, K, P). I am happy to name this genus in honor of my friend Mascagnia nobilis C. V. Morton, Proc. Biol. Soc. Wash. Alicia Lourteig (1913–2003), co-author of the first 45: 51. 1932. TYPE: Peru. Loreto: Yurimaguas, E. P. Killip & A. C. Smith 27557 (holotype, US; isotype, account of the Malpighiaceae of Argentina (O’Donell NY). & Lourteig, 1943). During my 1981 visit to study the great wealth of Malpighiaceae in the herbarium of the Muse´um National d’Histoire Naturelle in Paris, 2. Alicia macrodisca (Triana & Planchon) W. R. Dr. Lourteig was unfailingly helpful and hospitable, Anderson, comb. nov. Basionym: Hiraea macro- making my stay in Paris both pleasant and profitable. disca Triana & Planchon, Ann. Sci. Nat. Bot., Se´r. 4, 18: 326. 1862. Mascagnia macrodisca In the molecular trees published to date (Cameron et al., 2001; Davis et al., 2001, 2002), Alicia was (Triana & Planchon) Niedenzu, Arbeiten Bot. ¨ represented by A. anisopetala (as Mascagnia anisope- Inst. Konigl. Lyceum Hosianum Braunsberg 3: tala). The best-resolved of those trees, that of Davis et 16. 1908. Mascagnia anisopetala var. macro- al. (2002), places Alicia with 90% bootstrap support in disca (Triana & Planchon) Kostermans, Meded. a clade with Callaeum and Malpighiodes Niedenzu (as Bot. Mus. Herb. Rijksuniv. Utrecht 25: 5. Mascagnia bracteosa Grisebach in Martius). All three 1936. TYPE: Colombia. Meta: Villavicencio, J. genera have lateral-winged samaras and epipetiolar Triana 5568-6 (holotype, COL; isotypes, BM, G, stipules. Alicia is distinguished from Callaeum and K, MO). Malpighiodes by the combination of glands in the IV. Amorimia W. R. Anderson, gen. nov. TYPE: abaxial leaf surface, flowers in pseudoracemes, and Amorimia rigida (A. Jussieu in A. St.-Hilaire) W. densely hairy white/pink/lilac petals that are com- R. Anderson. pletely concealed by the sepals during enlargement of the bud. In A. anisopetala and some populations of A. Mascagnia sect. Pleuropterys Grisebach in Martius, Fl. Bras. macrodisca (Triana & Planchon) W. R. Anderson the 12(1): 91. 1858. LECTOTYPE, designated here: petals are strongly dimorphic, with the posterior petal Mascagnia pubiflora (A. Jussieu in A. St.-Hilaire) much larger than the four lateral petals, which makes Grisebach in Martius [5 Amorimia pubiflora (A. Jussieu in A. St.-Hilaire) W. R. Anderson]. those plants quite distinctive. Alicia consists of two species widespread in South Lianae lignosae; petiolus plerumque eglandulosus, rarius America. apice biglandulosus; lamina plerumque 2–multis glandulis parvis impressis in pagina abaxiali inter costam et marginem instructa (glandulae marginales in A. concinna); stipulae KEY TO THE SPECIES OF ALICIA interpetiolares; inflorescentia pseudoracemus vel panicula 1a. Lamina abaxially densely and persistently tomen- elongata; bracteae floriferae magnae (2–8 mm longae), tose (rarely glabrescent in age), the hairs stalked, patentes, saepe lanceolatae et glandulis submarginalibus with their crosspieces non-parallel, straight to prope basim instructae; petala abaxialiter 6 dense pilosa (in twisted; petals strongly dimorphic, the posterior A. concinna parce sericea), lutea vel aetate rubescentia; petal much larger than the lateral petals, often carpella omnino connata in ovario; styli recti et erecti vel nearly twice as long and much broader; samara distaliter recurvati, stigmate introrso, apice dorsaliter tomentose or loosely sericeous to glabrate, 2.8–5 rotundati, truncati, acuti, vel breviuncinati; samara alis (–6.5) cm across, mostly less than 4 cm ...... lateralibus apice usque ad nucem incisis, basi continuis vel ...... 1.A. anisopetala usque ad nucem incisis. 1b. Lamina abaxially sericeous to glabrate, with the hairs sessile, straight, and parallel; petals subequal Woody vines, sometimes described as shrubs. to strongly dimorphic; samara tightly sericeous to Leaves usually opposite, sometimes subopposite or glabrate,4–7.5cmacross...... 2.A. macrodisca alternate; petiole eglandular or biglandular near apex;

178 Novon

lamina eglandular or bearing 2–many small glands Amorimia is easily recognized by its leaf glands impressed in abaxial surface between midrib and usually on the abaxial surface (not the margin), margin, on very margin at base in Amorimia concinna interpetiolar stipules, yellow abaxially hairy but usually somewhat set in from margin in other petals, large spreading gland-bearing bracts, and species; stipules very small, triangular, borne on stem straight erect styles. The genus is not represented between petioles. Inflorescence with the flowers in any published molecular phylogeny. If one decussate to irregularly inserted in elongated ter- considers the morphological characters that seem to minal or axillary pseudoracemes or panicles; bracts be most informative in this group of genera (position large (2–8 mm long) and spreading, often lanceo- of stipules and leaf glands, exposure of petals in late and short-petiolate, often bearing 1(2) submar- bud, degree to which carpels are connate in ovary), ginal or marginal glands on each side near base; it would seem to resemble most closely Mascagnia floriferous peduncle 6 well developed; bracteoles s. str. Mascagnia differs from Amorimia in its like bracts but narrower, shorter, and less often glabrous petals, small mostly eglandular bracts, bearing glands, borne at or somewhat below apex of membranous samara with a prominent reticulum of peduncle. Sepals leaving petals exposed during arching anastomoses and the lateral wing usually enlargement of bud, appressed or revolute in anthe- continuous at base, and the fleshy disc that subtends sis, the lateral 4 bearing large paired abaxial the samaras. glands (occasionally the 2 glands adjacent to Amorimia is a genus of South America, comprising anterior sepal absent), the anterior eglandular (rarely ten species. bearing 1 or 2 small glands); corolla bilaterally symmetrical, the posterior petal erect, its claw KEY TO THE SPECIES OF AMORIMIA usually longer and thicker and its limb usually 1a. Petals adaxially glabrous, or hairy only on claw, or smaller, the lateral 4 petals spreading to reflexed; bearing only a few scattered hairs distally on limb. petals yellow or yellow turning red-orange in age 2a. Lamina cordate or subcordate at base; south- centralBrazil...... 7.A. pubiflora (especially the claws), abaxially densely velutinous, 2b. Lamina cuneate or rounded at base; western tomentose, or sericeous, at least on proximal half South America. (only thinly sericeous in A. concinna), adaxially 3a. Petals only sparsely sericeous abaxially glabrous or densely to sparsely hairy on distal in center of limb, otherwise glabrous; bracts linear, eglandular; leaf glands half; limb of petals entire or slightly erose; stamens marginal; Colombia ...... 3.A. concinna 10, all fertile; filaments connate at base, straight, 3b. Petals densely and evenly hairy over the subequal or longer opposite sepals; anthers alike, whole abaxial surface; bracts lanceolate, glabrous or variously hairy; carpels connate their usually biglandular near base; leaf whole length in ovary; styles terete or flattened glands borne in abaxial surface, at least the distal glands set in from margin, the laterally toward apex, erect and straight to recurved, glands at base of lamina sometimes subequal or the anterior somewhat shorter, slenderer, almost marginal. and more recurved than the posterior 2; apex of 4a. Lamina originally thinly sericeous styles dorsally rounded or truncate or acute to short- below, soon glabrate or sericeous only on midrib with straight ap- hooked, the stigma internal. Fruit dry, breaking apart pressed hairs; southwestern Amazo- into samaras separating from a short or moderately nia...... 1.A. amazonica high pyramidal torus; samara butterfly-shaped to 4b. Lamina at least originally velutinous depressed-elliptical with lateral wings dominant, below, the hairs persistent or the lamina glabrescent at maturity. chartaceous with many fine parallel veins, cleft to 5a. Petiole of larger leaves (5–)7– nut at apex, continuous at base or cleft part-way or 10 mm long; sepals mostly completely to nut, the margin entire or undulate to appressed in anthesis; posterior coarsely dentate; dorsal wing small, distinct at apex petal strongly differentiated from lateral petals, with a much and base or confluent with lateral wings at base; nut longer claw and smaller limb; almost always smooth between lateral and dorsal limb of lateral petals 7.5– wings; ventral areole broadly ovate or rotund to very 8.5 6 6–7 mm; lateral wings narrowly elliptical. of samara 30–42 mm high, 20– 30 mm wide; dorsal wing of The name Amorimia honors my Brazilian friend and samara extending to base of nut colleague, Andre´ M. Amorim (b. 1966), who has or nearly so; Guayas, Ecua- dor...... 5.A. kariniana begun a most promising career in the study of 5b. Petiole of larger leaves up to Malpighiaceae, concentrating at present on the large 6 mm long, mostly shorter; and difficult genus Heteropterys. sepals revolute at apex in Volume 16, Number 2 Anderson 179 2006 Segregates from Mascagnia

anthesis; posterior petal only 1. Amorimia amazonica (Niedenzu) W. R. Ander- moderately differentiated from son, comb. nov. Basionym: Mascagnia amazonica lateral petals, the claw thicker and slightly longer, the limb Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum somewhat smaller; limb of lat- Hosianum Braunsberg 8: 59. 1926. TYPE: eral petals 5.5–7.5(–8) 6 4– Brazil. Amazonas: Seringal S. Francisco, Rio 6 mm; lateral wings of samara Acre, E. Ule 9478 (holotype, B{; fragment of (11–)14–30 mm high, (6–)10– holotype, NY; isotypes, G, K, L). 18 mm wide; dorsal wing of samara usually extending at This species is known from southwestern Amazonia most to middle of nut, from there to base represented only [Peru (Foster 11965, MICH), Bolivia (Steinbach 339, by a rib; Cajamarca and San MICH), and adjacent Brazil]. Martıń, Peru . . . . 2. A. camporum 1b. Petals adaxially abundantly hairy distally on limb; eastern Brazil and adjacent Argentina [A. rigida 2. Amorimia camporum W. R. Anderson, sp. nov. complex]. TYPE: Peru. Cajamarca: San Ignacio, Distr. 6a. Lamina densely velutinous on both sides, the Chirinos, entre La Catagua y Tabloń, 5u199S, hairs V-shaped, persistent below, eventually 78u479W, 550–650 m, 9 Feb. 1996 (fl), J. deciduousabove;MinasGerais,Brazil.... Campos & O. Dıáz 2490 (holotype, MICH)...... 10.A. velutina 6b. Lamina initially sericeous or appressed-to- Liana; petiolus 2–6 mm longus; lamina foliorum majorum mentose on both sides with 6 appressed hairs, 5–11 6 3–6.7 cm, basi cuneata vel rotundata, abaxialiter soon glabrate on both sides or with some hairs pertinaciter velutina vel interdum glabrescens; flores in persistent below. pseudoracemis 5–18 cm longis 10–40-floris portati; bracteae 7a. Lamina with some hairs long-persistent 2–7 mm longae, lanceolatae, plerumque glanduliferae; below, especially on midrib, to eventually sepala plerumque apice 6 revoluta in anthesin; petala lutea, glabrescent; northeastern Brazil (Ceara´, abaxialiter dense subsericea vel velutina, adaxialiter glabra; Paraı´ba,Pernambuco)...... petala lateralia ungue 1–2 mm longo, limbo 5.5–7.5(28) 6 ...... 9.A. septentrionalis 4–6 mm; petalum posticum ungue 2–2.5 mm longo, limbo 5– 7b. Lamina very soon quite glabrate; eastern 7.5 6 3.5–5 mm; antherae glabrae; samara alis lateralibus Brazil from Bahia south to Rio Grande do discretis, (11–)14–30 mm altis, (6–)10–18 mm latis; ala Sul, and Misiones, Argentina. dorsali (1–)3–6 mm alta, (1–)2 mm lata, plerumque usque ad 8a. Ventral areole of samara narrowly medium nucis evoluta. elliptical or ovate, (2–)2.2–3.5 times as long as wide; sepals appressed in Woody vines; stems persistently short-velutinous or anthesis; styles dorsally rounded or glabrate in age. Leaves opposite; petiole 2–6 mm truncate at apex ...... 8. A. rigida long, velutinous, eglandular or biglandular near apex; 8b. Ventral areole of samara broadly lamina of larger leaves 5–11 6 3–6.7 cm, elliptical ovate to rotund, 1–1.8 times as long as wide; sepals recurved or or ovate, cuneate or rounded at base, acute, obtuse, revolute at apex in anthesis; rounded, or abruptly short-acuminate and often styles dorsally acute or apiculate at apiculate at apex, eglandular or bearing (1) several apex. small impressed abaxial glands in a row parallel to 9a. Samara 4.3–6 cm high and 5– margin but set in from it up to 5 mm, initially densely 8 cm wide; lamina usually biglandular at or just above velutinous on both sides with very short V-shaped base and eglandular distally; hairs, soon or eventually glabrate above, the abaxial bracts subsessile, the broad surface mostly persistently velutinous but occasional- petiole up to 0.5 mm long; ly glabrescent in age, the lateral veins 4–7 on each leaves often alternate or sub- side of midrib, the reticulum 6 prominent on both opposite, sometimes opposite ...... 6.A. maritima surfaces, especially above; stipules 0.3–0.5 mm long, 9b. Samara up to 3.3 cm high and black, very narrowly triangular, borne on stem 4.8 cm wide, mostly smaller; between petioles, persistent or deciduous. Flowers lamina often bearing 2(–4) borne in open pseudoracemes 5–18 cm long, contain- glands well above base and sometimes also biglandular at ing 10–40 flowers with the flowers decussate or just above base; bracts proximally and in no consistent arrangement distally, notably petiolate, the slender the pseudoracemes axillary or terminal, the axes petiole often 1 mm long; leaves (including peduncles and pedicels) velutinous; bracts mostly opposite, sometimes and bracteoles persistent or deciduous during en- subopposite; Parana´, Santa Catarina, and Rio Grande do largement of fruits; lowest bracts sometimes transi- Sul, Brazil, and Misiones, Ar- tional from vegetative leaves, more distal bracts 2– gentina...... 4.A. exotropica 7 6 (0.7–)1–2.5 mm, lanceolate, spreading, usually 180 Novon

bearing 2(–4) marginal glands near base, velutinous; which usually extends only to the middle of the nut, peduncle 2–11 mm long; bracteoles like bracts but not to the base as in A. pubiflora. Amorimia kariniana, smaller and sometimes eglandular, mostly borne at of Ecuador, differs from A. camporum in having apex of peduncle; pedicel 4–10 mm long, somewhat a longer petiole, sepals mostly appressed in anthesis, inflated distally. Sepals 1–2 mm long beyond glands, the posterior petal strongly differentiated from the 1.5 mm wide, triangular or ovate, acute to obtuse at lateral petals, larger lateral petals, and larger samaras apex, at least somewhat revolute at apex in anthesis, with the dorsal wing extending to the base of the nut or abaxially densely subvelutinous or sericeous, adaxi- nearly so. ally glabrous, the abaxial glands 8 on 4 lateral sepals Paratypes. PERU. Cajamarca, San Ignacio: Distr. (rarely 1 gland on the anterior sepal), 1.8–3 mm long; Huarango, entre Puerto Tabalozo y Nueva Esperanza, petals yellow, abaxially densely subsericeous to Campos & Dıáz 2015 (MO); Distr. Huarango, Puerto velutinous on claw and limb, adaxially glabrous, the Ciruelo–Camino a Huarango, Campos & Dıáz 2658 (MICH); limb entire or sinuate; lateral petals with claw 1– Distr. Chirinos, ‘‘Las Juntas,’’ Campos & Dıáz 3266 (MICH); Chirinos, Las Juntas, Campos 4846 (MICH). San Martıń: 2 mm long, limb 5.5–7.5(28) 6 4–6 mm, orbicular Huallaga, entre Bellavista y Banõs, Ferreyra 4744 (MICH); or broadly obovate or elliptical; posterior petal Juanjui, alto Rıó Huallaga, Klug 4259 (MO, NY, US); moderately differentiated from laterals by its often Huinguillo, Woytkowski 7183 (MICH, MO); Juanjui, Woyt- thicker and slightly longer claw and somewhat smaller kowski 7200 & 7202 (MICH, MO). limb, the claw 2–2.5 mm long and limb 5–7.5 6 3.5– 5 mm, orbicular; stamens glabrous; filaments 1.3– 3. Amorimia concinna (C. V. Morton) W. R. 2.5 mm long, connate at base; anthers 0.8–1.1 mm Anderson, comb. nov. Basionym: Mascagnia long, alike, reflexed in anthesis, the connective brown concinna C. V. Morton, Publ. Carnegie Inst. or dark red; ovary 1.3–2 mm high, densely velutinous; Wash. 461: 130. 1936. Mascagnia dumetorum C. styles 1.4–1.8 mm long, flattened laterally toward V. Morton, Proc. Biol. Soc. Wash. 45: 53. 1932, apex, glabrous or proximally pilose, erect to divergent, nom. illeg., non M. dumetorum Grisebach, 1879. dorsally acute at apex, alike or the anterior slightly TYPE: Colombia. Bolı´var: Since´, F. W. Pennell shorter than the posterior 2 and often more divergent. 4033 (holotype, US; isotypes, GH, NY). Samara (1.5–)2–3.5 cm wide, butterfly-shaped with the flabellate lateral wings distinct at base and apex, 4. Amorimia exotropica (Grisebach in Martius) W. short-velutinous on nut and wings; lateral wings (11–) R. Anderson, comb. nov. Basionym: Mascagnia 14–30 mm high, (6–)10–18 mm wide, the margin exotropica Grisebach in Martius, Fl. Bras. 12(1): sinuate, erose, or coarsely dentate; dorsal wing (1–)3– 93. 1858. TYPE: Brazil. Rio Grande do Sul: near 6 mm high, (1–)2 mm wide, widest at apex extended Porto Alegre, H. Fox 19 (lectotype, designated between lateral wings and narrowing abruptly down- here, K). ward, usually extending up to half the distance to base and thereafter represented only by a rib (but rarely Of the two syntypes, both from near Porto Alegre, extending as a narrow crest almost to base), distinct Fox 19 seems to me to be the fuller, more repre- from lateral wings, entire; ventral areole ovate, 4.5– sentative collection. 7 6 2–3 mm. This species occurs in the southernmost states of Brazil [Parana´ (Cordeiro 1217, MICH), Santa Catarina Phenology. Collected with flowers and fruits from (Krapovickas & Cristo´bal 39391, MICH), and Rio January to April and in September. Grande do Sul] and adjacent Misiones, Argentina Ecology and distribution. Dry and wet forests at (Vanni et al. 2903, MICH). 400–800 m in Cajamarca and San Martıń, Peru. Etymology. The epithet of this species honors one 5. Amorimia kariniana W. R. Anderson, sp. nov. of the collectors of the type and four of the paratypes, TYPE: Ecuador. Guayas: Guayaquil, Cerro Azul, the Peruvian botanist Jose´ Ricardo Campos (b. 1955). 50 m, 22 Oct. 1958 (fl), G. Harling 3026 Amorimia camporum differs from the A. rigida (holotype, S; isotype, MICH). Figure 4. complex of eastern Brazil and adjacent Argentina by Liana; petiolus (5–)7–10 mm longus; lamina foliorum having its petals adaxially glabrous, and it is majorum 8.5–13 6 5–8.5 cm, basi cuneata vel truncata, immediately distinguished from A. amazonica, whose abaxialiter 6 pertinaciter velutina; flores in pseudoracemis range overlaps that of A. camporum, by the velutinous 7–18 cm longis 16–26-floris portati; bracteae 3–6 6 1– 3 mm, patentes, glanduliferae; pedunculi 2–7 mm longi; lamina. It differs from A. pubiflora, of south-central pedicelli 6–9 mm longi; sepala apice obtusa vel rotundata, Brazil, by its smaller petals, its lamina that is usually plerumque appressa in anthesin; petala lutea, abaxialiter cuneate at the base, and the dorsal wing of its samara, dense subsericea vel velutina, adaxialiter glabra vel apice Volume 16, Number 2 Anderson 181 2006 Segregates from Mascagnia

paucis pilis instructa; petala lateralia ungue 1.5–2 mm divergent, dorsally truncate at apex, alike or the longo, limbo 7.5–8.5 6 6–7 mm; petalum posticum ungue anterior slenderer, shorter, and more strongly re- 3.5–4 mm longo, limbo 6–6.5 6 4.5–5 mm; antherae glabrae; samara alis lateralibus discretis, 30–42 mm altis, curved than the posterior 2. Samara 4.5–5 cm wide, 20–30 mm latis, ala dorsali 12–15 mm alta, 4 mm lata, butterfly-shaped with the flabellate or depressed- usque ad basim nucis evoluta vel paene. elliptical lateral wings distinct at base and apex, velutinous on nut and wings; lateral wings 30–42 mm Vine with slender woody stems, occasionally high, 20–30 mm wide, the margin sinuate or erose; described as a shrub; stems loosely sericeous or dorsal wing 12–15 mm high, 4 mm wide, widest at velutinous, glabrate in age. Leaves opposite; petiole apex extended between lateral wings and narrowing (5–)7–10 mm long, sericeous or velutinous to gla- downward but with measurable width almost or quite brate, eglandular or biglandular between middle and to base of nut, distinct from lateral wings at base and apex; lamina of larger leaves 8.5–13 6 5–8.5 cm, apex, entire or sinuate; ventral areole ovate or broadly elliptical, cuneate to truncate at base, elliptical, 8–12 6 4–6 mm. abruptly short-acuminate at apex, eglandular or, mostly, bearing a row of several (up to 10) small Phenology. Collected with flowers in June, July, impressed abaxial glands parallel to margin but set in September, and October, and with fruits in February from it up to 3 mm or nearly marginal at base, initially and June. moderately or thinly velutinous on both sides with V- Ecology and distribution. Known only from Gua- shaped hairs (except often loosely sericeous on yas, Ecuador, at 50–200 m, in roadside thickets, pas- midrib), soon nearly glabrate above, at least some tures, and dry tropical forests, and along streams. hairs long-persistent below, especially on veins, the lateral veins 5–7 on each side of midrib, the reticulum Etymology. Amorimia kariniana is named in prominent on both surfaces; stipules 0.3–0.7 mm long, honor of Karin Weishaar Douthit (b. 1926), who drew triangular, borne on stem between petioles, persistent the figure of it and all the other beautiful illustrations or deciduous. Flowers borne in elongated, open in this paper. Karin has been drawing Malpighiaceae pseudoracemes 7–18 cm long, containing 16–26 for me and my colleagues for over 30 years, and I want flowers with the flowers 6 decussate proximally and to pay tribute to her outstanding service. Her eye is as in no consistent arrangement distally, the pseudora- sharp, and her hand as steady, as when she started cemes axillary or terminal, the axes (including working for me soon after I joined the faculty of the peduncles and pedicels) velutinous; bracts and University of Michigan in 1974. bracteoles persistent or deciduous during enlargement Amorimia kariniana resembles A. pubiflora in its of fruits; bracts 3–6 6 1–3 mm, lanceolate, spread- velutinous leaves and large flowers, but it differs in ing, abaxially biglandular near base, velutinous the lamina having a cuneate base and more glands especially abaxially; peduncle 2–7 mm long; brac- distributed in a longitudinal row, the mostly shorter peduncles and pedicels, and the posterior petal, which teoles like bracts but much smaller (especially is more differentiated from the lateral petals here than narrower) and mostly eglandular, borne at apex of in A. pubiflora. See the discussion under A. camporum peduncle; pedicel 6–9 mm long, somewhat inflated for the distinctions between that species and A. distally. Sepals 1.5–2 mm long beyond glands, 1.7– kariniana. 2.5 mm wide, ovate, obtuse to rounded at apex, appressed in anthesis (or slightly revolute at sides), Paratypes. ECUADOR. Guayas: vicinity of Guayaquil, abaxially densely velutinous or subsericeous, adaxi- Cerro Azul, Asplund 16617 & 17586 (S); E of Las Juntas, ally glabrous, the abaxial glands 8 on 4 lateral sepals, Fagerlind & Wibom 242 (S); Pedro Carbo, Haught 3070 2.5–3.5 mm long; petals yellow, abaxially densely (MICH, NY, US); Guayaquil, road to Aguas Piedras, Rowlee & Mixter 1109 (US); Guayaquil, Cerro Mirador de los Monos, subsericeous to velutinous on claw and limb, adaxially Rubio & Palacios 2445 (MICH, MO). glabrous or bearing a few scattered hairs near apex, the limb entire or sinuate; lateral petals with claw 6. Amorimia maritima (A. Jussieu) W. R. Ander- 1.5–2 mm long, limb 7.5–8.5 6 6–7 mm, broadly son, comb. nov. Basionym: Hiraea maritima A. obovate; posterior petal strongly differentiated from Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, 13: 259. laterals, the claw 3.5–4 mm long and limb 6–6.5 1840. TYPE: Brazil. Bahia: ‘‘in petrosis mariti- 6 4.5–5 mm, spatulate; filaments 2.5–3.5 mm long, mis,’’ P. Salzmann s.n. (holotype, G; isotypes, G, connate at base, glabrous or abaxially sparsely pilose; K, P). anthers 0.8–1.4 mm long, alike, glabrous, the connective brown; ovary 1.7–2 mm high, densely veluti- This species is known only from near the Atlantic nous; styles 2–2.2 mm long, terete or flattened coast in the Brazilian states of Bahia and Espı´rito laterally toward apex, sericeous proximally, erect to Santo (Do¨bereiner & Tokarnia 2009, MICH).

Volume 16, Number 2 Anderson 183 2006 Segregates from Mascagnia

7. Amorimia pubiflora (A. Jussieu in A. St.- Amorimia rigida is known from the Brazilian states Hilaire) W. R. Anderson, comb. nov. Basionym: of Minas Gerais, Bahia (Amorim 3565, MICH), Hiraea pubiflora A. Jussieu in A. St.-Hilaire, Fl. Espı´rito Santo (Do¨bereiner & Tokarnia 1765, MICH), Bras. Merid. 3: 14. 1832 [1833]. Mascagnia and Rio de Janeiro (Andreata 373, MICH). pubiflora (A. Jussieu in A. St.-Hilaire) Grisebach in Martius, Fl. Bras. 12(1): 91. 1858. TYPE: 9. Amorimia septentrionalis W. R. Anderson, sp. Brazil. Minas Gerais: near Mangahy, A. St.- nov. TYPE: Brazil. Ceara´: sine loc., 22 June Hilaire s.n. (lectotype, designated here, P [F neg. 1958 (fl/fr), J. Do¨bereiner 538 (holotype, MICH). 35631]; isotypes, P). Figure 6.

There are three type sheets in P, all annotated by Liana; lamina foliorum majorum 6–12(214) 6 3.3– Jussieu, probably all from the same gathering. The one 7.5 cm, elliptica vel parum ovata obovatave, plerumque I am designating lectotype is the fullest, most biglandulosa prope basim, sericea vel appresso-tomentosa representative specimen. mox glabrescens aliquot pilis abaxialibus saepe persistenti- bus; flores in pseudoracemis 6–25 cm longis 10–70-floris Mascagnia parnahybensis Glaziou, Bull. Soc. Bot. France 52 portati; bracteae 2–4.5(26) 6 1–2 mm, patentes, plerum- (Me´m. 3): 77. 1905. Mascagnia pubiflora var. grandi- que glanduliferae; sepala apice acuta, revoluta in anthesin; folia Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum petala lutea, utrinque tomentosa, subaequalia, ungue 0.5– Hosianum Braunsberg 3: 23. 1908. TYPE: Brazil. 1.5 mm longo, limbo 3.2–6 6 2–2.7 mm, basi cuneato; Goia´s: Rio Parnahyba, A. Glaziou 20757 (lectotype, antherae 0.8–1.5 mm longae, basi piliferae; styli apice designated here, P [MICH, WRA neg. 81-22-7]; dorsaliter acuti vel breviuncinati; samara alis lateralibus isotypes, G, K, P). discretis, (11–)20–33 mm altis, 8–20 mm latis, ala dorsali 7–15 mm alta, 1.5–5 mm lata, areola ventrali late ovata, 4.5– Glaziou 20757 was the only collection cited. There 7 6 4–5 mm. are two sheets in P, as well as duplicates in G and K. Vine with slender woody stems, sometimes de- It is appropriate to make one of the two P sheets scribed as a shrub; stems sericeous or appressed- lectotype, because Glaziou was associated with that tomentose to glabrate. Leaves opposite; petiole 3– herbarium, and the fuller of the two specimens is the 6 mm long, sericeous to glabrate, usually eglandular, one I have selected as lectotype. occasionally biglandular at very apex; lamina of larger leaves 6–12(214) 6 3.3–7.5 cm, elliptical 8. Amorimia rigida (A. Jussieu in A. St.-Hilaire) or slightly ovate or obovate, cuneate or rounded at W. R. Anderson, comb. nov. Basionym: Hiraea base, obtuse or acute or (usually) acuminate at apex, rigida A. Jussieu in A. St.-Hilaire, Fl. Bras. mostly bearing 1(2) small impressed glands on each Merid. 3: 14. 1832 [1833]. Mascagnia rigida (A. side of abaxial surface, at base or up to 1.5 cm above Jussieu in A. St.-Hilaire) Grisebach in Martius, base, between midrib and margin, initially sericeous Fl. Bras. 12(1): 92. 1858. TYPE: Brazil. Minas or appressed-tomentose on both sides, the hairs soon Gerais: S. Miguel de Jequitinhonha, A. St.- deciduous or some often persistent on abaxial surface, Hilaire Catal. B1 no. 1501 (lectotype, designated the lateral veins 4–7 on each side of midrib; stipules here, P [MICH, WRA neg. 81-22-15]; isotypes, 0.4–1.5 mm long, triangular, borne on stem between P). Figure 5. petioles, persistent or deciduous. Flowers borne in pseudoracemes 6–25 cm long, containing 10–70 The St.-Hilaire collection cited above is the only flowers with the flowers often decussate proximally potential type in P. There are three sheets, all and in no consistent arrangement distally, the annotated by Jussieu; I have selected the best material pseudoracemes axillary or arrayed in lateral or of those three to designate as lectotype. terminal panicles, the axes loosely sericeous or Mascagnia coriacea Grisebach in Martius, Fl. Bras. 12(1): velutinous; bracts and bracteoles mostly deciduous 92. 1858. Mascagnia rigida subsp. coriacea (Grisebach during enlargement of fruits; bracts 2–4.5(26) 6 1– in Martius) Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. 2 mm, lanceolate, often narrowed at base to a short Lyceum Hosianum Braunsberg 3: 19. 1908. TYPE: petiole, spreading, mostly bearing 1(2) bulging Brazil. Minas Gerais: Tejuco [Diamantina], G. Gardner 5394 (lectotype, designated here, K; duplicate, BM). marginal or submarginal glands on each side below middle, loosely sericeous especially abaxially; pedun- Grisebach cited two syntypes, Barbosa s.n. from Rio cle 0.5–4 mm long, loosely sericeous or velutinous; de Janeiro and Gardner 5394 from Minas Gerais. I bracteoles 1.5–2 6 0.6–1 mm, elliptical, mostly have not been able to locate the Barbosa collection. eglandular but sometimes bearing 2 small glands, The Gardner collection is a good representative of this borne at or somewhat below apex of peduncle; pedicel species, and can serve well as lectotype; the sheet at 3–5 mm long, loosely sericeous or velutinous. Sepals K is better material than the one at BM. 1.5–2.2 6 1.2–2 mm, triangular, acute at apex,

186 Novon

in age, borne on stem at base of petiole, persistent or has some appressed hairs persistent on the abaxial deciduous. Flowers borne in pseudoracemes 4–20 cm surface. It also differs in the samara; in A. rigida the long, containing 10–50 flowers with the flowers nut is sericeous, the wings are thinly sericeous to sometimes decussate and sometimes in no consistent glabrate, and the ventral areole is narrower, usually at arrangement, the pseudoracemes axillary or arrayed in least 2.2 times as long as wide. lateral or terminal panicles, the axes velutinous; Paratype. BRAZIL. Minas Gerais: Entre Itaobim e bracts and bracteoles persistent or deciduous late in Jequitinhonha, Shepherd et al. 4409 (NY). anthesis or during enlargement of fruits; bracts 3– 4(26) 6 1.5–3 mm, lanceolate, ovate, or deltate and often narrowed at base to a short petiole, spreading, V. Carolus W. R. Anderson, gen. nov. TYPE: Carolus usually bearing 1 or 2 small inframarginal to marginal chlorocarpus (A. Jussieu) W. R. Anderson. abaxial glands on each side near or below middle, Mascagnia sect. Dipterys Skottsberg, Kongl. Svenska Ve- velutinous or appressed-velutinous on both sides; tenskapsakad. Handl. 35(6): 7. 1901. TYPE: Mascag- nia chlorocarpa (A. Jussieu) Grisebach [5 Carolus peduncle 1–4 mm long, velutinous; bracteoles 2– chlorocarpus (A. Jussieu) W. R. Anderson]. 2.5 6 1–2 mm, like bracts but smaller and less Heladena sect. Hassleria Niedenzu, Arbeiten Bot. Inst. consistently glanduliferous, borne at apex of pedun- Ko¨nigl. Lyceum Hosianum Braunsberg 5: 15. 1914. cle; pedicel 3–5 mm long, velutinous. Sepals ca. TYPE: Heladena hassleriana Niedenzu in Chodat & 2 mm long and wide, rounded at apex, appressed in Hassler [5 Carolus chlorocarpus (A. Jussieu) W. R. Anderson]. anthesis, abaxially densely appressed-velutinous, adaxially sericeous proximally, the abaxial glands The name of this section was spelled Halleria in the 2–2.7 mm long; petals ‘‘yellow with an orange median protologue. Niedenzu (1928: 582) said that was stripe,’’ abaxially densely tomentose or velutinous on a typographical error and corrected the spelling to claw and limb, adaxially densely tomentose or Hassleria. velutinous on distal half of limb with hairs decurrent Lianae lignosae; lamina eglandulosa vel margine 22mul- on midrib, the limb elliptical, truncate or slightly tis glandulis parvis instructa; stipulae interpetiolares; hastate at base (especially in lateral petals), entire; inflorescentia pseudoracemus, umbella, vel panicula, flori- lateral petals with claw 1–1.5 mm long, limb 3.5– bus plerumque decussatis; bracteae bracteolaeque eglandu- 5 6 2.5–3.5 mm; posterior petal with claw 2–2.7 mm losae; sepala petalis in alabastro breviora, appressa per long, limb 3–4 6 2.3–3.5 mm; filaments 1.5–2 mm anthesin; petala lutea, abaxialiter dense sericea; antherae inter se similares, glabrae vel paucipiliferae; styli stigmate long, connate at base, glabrous; anthers 1.1–1.4 mm introrso; samara alis lateralibus apice basique usque ad long, each half with a basal tuft of hairs, the nucem incisis. connective dark red; ovary 1–1.5 mm high, densely Woody vines. Leaves opposite; petiole eglandular appressed-velutinous; styles 1.3–1.7 mm long, cylin- or bearing 2–4(–6) small glands; lamina eglandular or drical, initially erect but soon recurved, rounded or bearing 2–many tiny glands on or embedded in truncate dorsally and somewhat flattened laterally at margin; stipules very small, triangular, borne on stem apex, the anterior slenderer and shorter than the between petioles or beside base of petiole, caducous posterior 2. Samara (slightly immature?) ca. 2.5 cm or persistent. Inflorescence axillary or terminal, wide, butterfly-shaped with the flabellate lateral wings a pseudoraceme or umbel or a panicle of pseudo- distinct at apex and base, velutinous on nut, loosely racemes or umbels, the flowers mostly decussate; sericeous on wings; lateral wings 12–14 mm high, bracts and bracteoles eglandular, persistent; florifer- 10 mm wide, entire; dorsal wing 6–7 mm high, 3– ous peduncle often well developed, reduced or lacking 4 mm wide, coarsely dentate; ventral areole broadly in some populations. Sepals shorter than petals in bud ovate, ca. 5.5 6 3–3.5 mm. and leaving petals exposed during enlargement of Phenology. Collected with flowers and fruits in bud, appressed in anthesis, all 5 eglandular or the March and April. lateral 4 bearing large paired abaxial glands; corolla bilaterally symmetrical, the posterior petal erect and Ecology and distribution. Known only from two 6 different from the spreading lateral 4; petals yellow, collections from Minas Gerais, Brazil; in neither case abaxially densely sericeous on claw and limb except did the collectors record the elevation or habitat. near margin, adaxially glabrous; stamens 10, all Etymology. The epithet of this species refers to fertile; filaments connate in proximal 1/4–1/2, the persistently velutinous stems and leaves, which sometimes subequal but mostly of differing lengths, immediately distinguish it from the other species in often longest opposite anterior sepal and shortest the Amorimia rigida complex; those have the stems opposite posterior petal, the 2 opposite posterior- and leaves sericeous to soon glabrate, or the lamina lateral petals often thicker than others; anthers alike Volume 16, Number 2 Anderson 187 2006 Segregates from Mascagnia

or subequal, glabrous or sparsely hairy; carpels 3a. Lamina up to 7 6 3.5 cm, mostly small- connate their whole length in ovary; styles erect and er; inflorescence an unbranched axillary pseudoraceme; samara smooth between straight or somewhat bowed or spreading, alike, lateral wings, the dorsal winglet completely dorsally rounded to short-apiculate or short-hooked lacking or represented at most by a round- and laterally compressed or terete at apex, the stigma ed hump at apex; anthers glabrous, the internal. Fruit dry, breaking apart into samaras connectivebrownish...... 2.C. chasei 3b. Lamina of larger leaves mostly at least separating from a high pyramidal torus; samara 9 6 4 cm, often larger; inflorescence butterfly-shaped with lateral wings dominant, charta- usually compound, an axillary or termi- ceous with many fine parallel veins, cleft to nut at nal panicle of pseudoracemes; samara base and apex, the margin sinuate to coarsely dentate; bearing between lateral wings ca. 5 parallel 6 dissected winglets 1–2.5 mm dorsal wing(s) lacking or, if developed, small, distinct wide; anthers bearing a few hairs on at base and apex; ventral areole ovate to linear; locules, the connective black. . 5. C. renidens samara reduced or highly modified in some popula- 2b. Flowers borne ultimately in umbels of 4(–6); tions of Carolus sinemariensis. Panama and Pacific South America. 4a. Samara with a well-developed rectangu- I am happy to name this genus in honor of my friend lar or triangular dorsal wing between and colleague, Charles Cavender Davis (b. 1974), lateral wings, (2–)3–7 mm wide; styles dorsally short-apiculate at apex; pedun- whose molecular studies have already made a sub- cles 1–2.5 mm long; Pacific Ecuador and stantial contribution to our understanding of the Peru ...... 1.C. anderssonii systematics of the Malpighiaceae. 4b. Samara nearly smooth between lateral Carolus is distinguished by its interpetiolar stip- wings, the dorsal wing represented only by a longitudinal rib and a triangular ules, marginal leaf glands, densely sericeous yellow winglet up to 1.5 mm high and wide at petals, and butterfly-shaped samaras. Because they apex; styles with a dorsal hook at apex; both have interpetiolar stipules and hairy yellow peduncles mostly lacking, occasionally petals, this genus can be confused with Amorimia, but up to 0.5 mm long; Panama . . . 4. C. dukei 1b. Lamina abaxially glabrous or glabrate at maturity, that genus differs in having its leaf glands usually on rarely thinly sericeous or appressed-tomentose with the abaxial surface between the midrib and the the hairs never dense enough to completely hide margin, and in its large spreading gland-bearing epidermis. bracts. Aenigmatanthera is readily distinguished from 5a. Flowers borne in 6 elongated pseudoracemes of 6–14(–20), with internodes between pairs of Carolus by its epipetiolar stipules, glabrous or flowers longer than peduncles and the pairs 6 subglabrous petals, and anthers densely sericeous on evenly distributed; South America south of the the connective. In previously published molecular Amazon, from Bolivia and Paraguay to studies (Cameron et al., 2001; Davis et al., 2001, southeastern Brazil ...... 3.C. chlorocarpus 2002), Carolus was represented by C. chasei (W. R. 5b. Flowers crowded distally in umbels, corymbs, Anderson) W. R. Anderson (as Mascagnia chasei W. or dense pseudoracemes of 4–8(–12) with R. Anderson). The best-resolved of those trees is that internodes between pairs of flowers lacking or in Davis et al. (2002) in which C. chasei was placed in very short, mostly shorter than peduncles; Mexico to South America north of 4uS, and a clade comprising Dicella Grisebach, Tricomaria Lesser Antilles ...... 6.C. sinemariensis Hooker & Arnott, Heteropterys, Hiptage Gaertner, and Niedenzuella (as Mascagnia stannea (Grisebach) 1. Carolus anderssonii (W. R. Anderson) W. R. Niedenzu). That clade had no bootstrap support, nor Anderson, comb. nov. Basionym: Mascagnia did the position of Carolus in it. The only one of those anderssonii W. R. Anderson, Contr. Univ. genera that has butterfly-shaped samaras is Nieden- Michigan Herb. 22: 15. 1999. TYPE: Ecuador. zuella, which has long, imbricated sepals that conceal Loja: near Malacatos, G. Harling & L. Andersson the petals in the enlarging bud. 21477 (holotype, GB; isotype, US). Carolus comprises six species of Mexico, Central America, the Lesser Antilles, and South America. This species is known from Pacific Ecuador and Peru (Anderson, 1999: 16).

KEY TO THE SPECIES OF CAROLUS 2. Carolus chasei (W. R. Anderson) W. R. Ander- 1a. Lamina abaxially densely and persistently seri- son, comb. nov. Basionym: Mascagnia chasei W. ceous at maturity. R. Anderson, Contr. Univ. Michigan Herb. 19: 2a. Flowers borne ultimately in 6 elongated pseudoracemes of 4–20, with internodes de- 378. 1993. TYPE: Brazil. Bahia: S of Maraca´s, T. veloped between pairs of flowers and the pairs S. dos Santos et al. 3480 (holotype, CEPEC; 6 evenly distributed; eastern Brazil. isotypes, MICH, NY). 188 Novon

3. Carolus chlorocarpus (A. Jussieu) W. R. berg 3: 20. 1908, has been widely used for this plant Anderson, comb. nov. Basionym: Hiraea chloro- (Niedenzu, 1928: 111). It is a superfluous synonym for carpa A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, 13: Hiraea cinerea, because Niedenzu cited Bunbury’s 259. 1840. Mascagnia chlorocarpa (A. Jussieu) collection as his type. Grisebach in Martius, Fl. Bras. 12(1): 93. 1858. TYPE: Brazil. Region of Rio de Janeiro, Vauthier Hiraea heteropetala A. Jussieu, Arch. Mus. Hist. Nat. 3: 557. [21] (lectotype, designated by Anderson (1993: 1843. Tetrapterys heteropetala (A. Jussieu) Grisebach 379), P [F neg. 35628]; duplicates, G, K, P). in Martius, Fl. Bras. 12(1): 88. 1858. TYPE: Brazil. P. Claussen 33 (lectotype, designated here, P [F neg. Heladena hassleriana Niedenzu in Chodat & Hassler, Bull. 35603]). Herb. Boissier, Se´r. 2, 7: 294. 1907. TYPE: Para- guay. Rı´o Apa, E. Hassler 7837 (lectotype, designated As syntypes Jussieu cited Clausen and Sellow here, G; isotypes, A, BM, G, K, LIL, MICH, MO, NY, P). collections from Brazil, without numbers or detailed localities. In B there was a Sellow collection The sole type collection was Hassler 7837, but annotated by Jussieu (F. neg. 12731). In P there are because the holotype in B was destroyed it is two Clausen sheets, 33 and 77, both representing this necessary to designate one of the many isotypes as species. Only Clausen 33 was annotated by Jussieu, lectotype. I have selected one of the six sheets at G, for which reason that specimen is designated the one annotated by Niedenzu in 1907; four of the lectotype. others were not annotated by Niedenzu, and the fifth was not annotated by him until 1910. Banisteria riedeliana Regel, Index Sem. Horti Bot. Petropol. Carolus chlorocarpus is known from Bolivia (Gentry 17. 1855. Mascagnia riedeliana (Regel) W. R. et al. 73882, MICH) and Paraguay to southeastern Anderson, Contr. Univ. Michigan Herb. 14: 21. 1980. TYPE: Cultivated in Hort. Petropol. from seeds sent by Brazil. L. Riedel, from the base of Corcovado, Rio de Janeiro, Brazil (holotype, LE). 4. Carolus dukei (Cuatrecasas & Croat) W. R. Anderson, comb. nov. Basionym: Mascagnia 6. Carolus sinemariensis (Aublet) W. R. Anderson, dukei Cuatrecasas & Croat, Ann. Missouri Bot. comb. nov. Basionym: Banisteria sinemariensis Gard. 67: 908. 1980 [1981]. TYPE: Panama. Aublet, Hist. Pl. Guiane 1: 462. 1775. Mascag- Panama´: between Can˜asas and Sabalo, J. Duke nia sinemariensis (Aublet) Grisebach in Martius, 14468 (holotype, US; isotypes, MO, PMA not Fl. Bras. 12(1): 93. 1858. TYPE: French Guiana. seen). J. Aublet s.n. (holotype, BM). Figure 7.

5. Carolus renidens (A. Jussieu) W. R. Anderson, Malpighia volubilis Sims, Bot. Mag. 21: 809. 1805. comb. nov. Basionym: Hiraea renidens A. Byrsonima volubilis (Sims) DC., Prodr. 1: 581. 1824. Banisteria volubilis (Sims) Endlicher, Cat. Horti Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, 13: 260. Vindob. 2: 375. 1842. Mascagnia volubilis (Sims) 1840. Tetrapterys renidens (A. Jussieu) Grisebach Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum Hosia- in Martius, Fl. Bras. 12(1): 68. 1858. Mascagnia num Braunsberg 3: 22. 1908. TYPE: Cultivated, from renidens (A. Jussieu) W. R. Anderson, Contr. West Indies (holotype, K). Univ. Michigan Herb. 15: 136. 1982. TYPE: Hiraea schizoptera Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 36: 584. 1863. Mascagnia schizoptera (Turcza- Brazil. Sebastiano´polis [Rio de Janeiro], Vauthier ninow) Cuatrecasas, Webbia 13: 373. 1958. TYPE: St. 450 (lectotype, designated here, P [MICH, WRA Vincent. G. Caley s.n. (holotype, KW not seen; isotype, negs. 81-22-30 & 31]; isotypes, G, L, P). G). Diplopterys microcarpa Sandwith, Kew Bull. 1931: 183. Vauthier 450 was the single type collection. It is 1931. Mascagnia microcarpa (Sandwith) W. R. Ander- represented at P by three sheets, of which I have son, Mem. New York Bot. Gard. 32: 220. 1981. TYPE: selected as lectotype the best, most representative Guyana. Essequibo: Moraballi Creek near Bartica, N. specimen, which was annotated by Jussieu. Sandwith 289 (holotype, K; isotype, NY). Mascagnia hondensis C. V. Morton, Proc. Biol. Soc. Wash. 45: 52. 1932. TYPE: Colombia. Tolima: Honda, Bro. Hiraea cinerea Bunbury, Proc. Linn. Soc. London 1: 109. Ariste Joseph A371 (holotype, US). 1841. TYPE: Brazil. Rio de Janeiro, Corcovado, C. Bunbury s.n. (holotype, CGE not seen; isotypes, BR not This species is interpreted very broadly here. In seen, K not seen). that sense, it is known from Mexico (Wilbur & Wilbur The name Mascagnia metallicolor Niedenzu, Ar- 2394, MICH) to South America north of 4uS, and the beiten Bot. Inst. Ko¨nigl. Lyceum Hosianum Brauns- Lesser Antilles.

190 Novon

VI. Christianella W. R. Anderson, gen. nov. TYPE: rarely 1, narrow; ventral areole circular or broadly Christianella mesoamericana (W. R. Anderson) ovate. W. R. Anderson. It gives me great pleasure to name this genus in honor of Christiane Eva Seidenschnur Anderson (b. Lianae lignosae vel frutices; lamina eglandulosa vel in dimidio proximali aliquot glandulis parvis marginalibus 1944). Christiane’s excellent revisions in the Malpigh- instructa; stipulae parvae prope basim petioli portatae; iaceae and her achievements as editor of Systematic bracteae bracteolaeque eglandulosae vel glandulis parvis Botany Monographs have made her an important figure marginalibus instructae; bracteolae apice pedunculi portatae; in our generation of plant taxonomists. sepala aliquot glandulis parvis marginalibus instructa, his Christianella is distinguished by the small glands sessilibus, subsessilibus, vel stipitatis; petala lutea, abaxialiter tomentosa; filamenta valde heteromorpha; samara found on the margin of the sepals, and in most species membranacea, alis lateralibus apice distinctis, basi continuis on the margin of the bracteoles and sometimes the vel usque ad nucem incisis. bracts. Such marginal glands are found in a few other Woody vines, or shrubby in open habitats. Petiole genera of Malpighiaceae, but not in any of the other eglandular or bearing 2–10 large to small glands in 2 groups included in Mascagnia by Niedenzu. This rows; lamina eglandular or bearing several small group of species is also notable for the marginal leaf sessile glands on margin on proximal half; stipules glands, the yellow abaxially tomentose petals, the very small, triangular, borne on petiole near base. heteromorphic filaments, and the membranous sa- Inflorescence with flowers decussate to irregularly maras with the lateral wings cleft to the nut at the apex and continuous or cleft at the base. The stiff hairs on inserted in short to elongated pseudoracemes, termi- the samara wings are usually inserted at characteristic nal or lateral, single or (usually) grouped in panicles; dark spots. Christianella has not been included in any bracts and bracteoles eglandular or bearing several published molecular study. subsessile to long-stalked, clavate or capitate marginal glands; floriferous peduncle well developed; Christianella consists of five species of Central and bracteoles borne at apex of peduncle. Sepals elongat- South America. ed but separating to expose petals during enlargement of bud, appressed to spreading in anthesis, the lateral KEY TO THE SPECIES OF CHRISTIANELLA 4 bearing large paired abaxial glands, all 5 bearing 1a. Lamina sericeous to soon glabrate, the hairs sessile a row of small sessile or subsessile to long-stalked, and straight. 2a. Bracts and bracteoles 2–3 mm long, eglandu- clavate or capitate marginal glands; corolla bilaterally lar or with several tiny sessile or subsessile symmetrical, the posterior petal with a longer and marginal glands; marginal glands of sepals thicker claw than the lateral 4; petals yellow, sessile or subsessile; southwestern Brazil . . . abaxially densely tomentose, adaxially glabrous; ...... 4.C. paludicola androecium bilaterally symmetrical; stamens 10, all 2b. Bracts and bracteoles 3–7 mm long, with many long-stalked marginal glands; marginal fertile, glabrous; filaments connate at base, straight, glands of sepals long-stalked. strongly heteromorphic, longest and much thickened 3a. Bracts caducous; petiole of larger leaves opposite the 2 posterior-lateral petals, usually long 11–15 mm long; lamina of larger leaves but slender opposite the anterior sepal, shortest 5–7.2 cm wide; samaras orbicular, 35– 60 mm wide, the lateral wing continuous opposite the posterior petal; anthers subequal or at base, cleft to nut at apex; western larger on longer filaments; gynoecium bilaterally Amazonia ...... 1.C. glandulifera symmetrical; carpels very shortly connate in ovary; 3b. Bracts persistent or deciduous in fruit; anterior style straight and shorter than the others, petiole of larger leaves 2–8 mm long; the posterior 2 straight or bowed from the base; lamina of larger leaves 2.5–5 cm wide; samaras butterfly-shaped, 22–42 mm apex of styles dorsally truncate to short-hooked, wide, the lateral wing cleft to nut at base the stigma internal. Fruit dry, breaking apart into and apex; southeastern Mexico to Pana- samaras separating from a short obscure pyramidal ma...... 2.C. mesoamericana torus; samara suborbicular to butterfly-shaped, the 1b. Lamina densely and persistently tomentose, velu- lateral wing(s) dominant, membranous with reticulate- tinous, or woolly, especially abaxially, the hairs stalked. anastomosing venation, cleft to nut at apex, continu- 4a. Peduncle and pedicel very deeply woolly, 1– ous at base or cleft to nut, bearing stiff, usually 1.5 mm or more diam. including vesture; hairs basifixed or sub-basifixed hairs inserted at dark so dense on abaxial leaf surface as to make it spots, the margin undulate or coarsely dentate; dorsal difficult to distinguish the crosspiece of any given hair; bracteoles abaxially very densely wing small, extended forward at apex through gap spreading-woolly, adaxially tomentose, at least in lateral wing, distinct at base or confluent with on the distal half; sepals abaxially deeply lateral wing; intermediate winglets mostly lacking, spreading-woolly; petals abaxially densely Volume 16, Number 2 Anderson 191 2006 Segregates from Mascagnia

woolly; southern Bolivia, northern Paraguay, multiglandulosa var. surinamensis Kostermans, and southern Brazil . . . 3. C. multiglandulosa Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 25: 4b. Peduncle and pedicel densely tomentose or velutinous but only 0.4–0.7 mm diam. in- 5. 1936. Mascagnia surinamensis (Kostermans) cluding vesture; leaf hairs much less dense, W. R. Anderson, Contr. Univ. Michigan Herb. the crosspieces of individual hairs easi- 17: 53. 1990. TYPE: Suriname. Brownsberg, 23 ly distinguished; bracteoles abaxially ap- Sep. 1931, W. C. v. Emden (holotype, U; isotypes, pressed-tomentose, adaxially glabrous or tomentose only at very apex; sepals abaxially K, MICH). Figure 8. appressed-tomentose or subsericeous; petals abaxially moderately appressed-tomentose; This species is widespread in Amazonian South Amazonian South America . . . 5. C. surinamensis America, including Guyana and Suriname (Anderson 1990b: 53). 1. Christianella glandulifera (Cuatrecasas) W. R. Anderson, comb. nov. Basionym: Mascagnia VII. Malpighiodes Niedenzu, Verz. Vorles. Ko¨nigl. glandulifera Cuatrecasas, Webbia 13: 365. Lyceum Hosianum Braunsberg 1909/10: 31. 1958. TYPE: Colombia. Amazonas-Vaupe´s: Rı´o 1909. Tetrapterys subsect. Malpighiodes (Nie- Apaporis, R. E. Schultes & I. Cabrera 13208 denzu) Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. (holotype, US; isotypes, GH, NY). Lyceum Hosianum Braunsberg 4: 12. 1912. LECTOTYPE: Malpighiodes spruceana Niedenzu 2. Christianella mesoamericana (W. R. Anderson) [5 Malpighiodes bracteosa (Grisebach in Mar- W. R. Anderson, comb. nov. Basionym: Mascag- tius) W. R. Anderson]. Brazil, Amazonas: ‘‘in nia mesoamericana W. R. Anderson, Contr. Univ. vicinibus Barra [Manaus], prov. Rio Negro, coll. Michigan Herb. 16: 105. 1987. TYPE: Guate- R. Spruce Dec.–Martius 1850–51,’’ R. Spruce mala. Retalhuleu: Between Retalhuleu and [1093] (lectotype, designated here, M). Nueva Linda, P. C. Standley 87301 (holotype, F; isotype, US). Woody vines. Petiole eglandular or bearing 2–8 This species is known from southeastern Mexico, small glands in 2 rows; lamina usually bearing few to Guatemala, and Panama (Anderson, 1987: 105), and has many small glands impressed in abaxial surface in 1– recently been found in Costa Rica (Acosta 803,INB). 3 rows between midrib and margin; stipules minute, triangular, borne on proximal half of petiole or at junction of petiole and stem, or apparently lacking. 3. Christianella multiglandulosa (Niedenzu in Inflorescence a terminal or lateral compound di- Chodat & Hassler) W. R. Anderson, comb. nov. chasium or paniculate dichasium, strictly decussate, Basionym: Mascagnia multiglandulosa Niedenzu with the flowers borne in pairs or umbels or corymbs in Chodat & Hassler, Bull. Herb. Boissier, Se´r. 2, of 4(–8); floriferous peduncle well developed; 7: 284. 1907. TYPE: Paraguay. Near Valenzuela, bracteoles eglandular, broad and rounded (elliptical E. Hassler 6998 (lectotype, designated here, G; or obovate), 2.5–4 mm long, borne between middle isotypes, A, G, K, MO, NY). and apex of peduncle. Sepals valvate, completely The holotype sheet of Hassler 6998 was destroyed concealing petals during enlargement of bud, revolute at B. There are four isotypes at G; the lectotype is in anthesis, all 5 biglandular; corolla bilaterally the isotype that was annotated lectotype by W. R. symmetrical, the posterior petal somewhat different Anderson in 1993. from lateral 4; petals yellow turning red in age, This species is known from Paraguay and southern glabrous; stamens 10, all fertile; filaments ca. 1/2- Brazil (Anderson 1990b: 53) and southern Bolivia connate, straight, those opposite sepals slightly (Gutie´rrez et al. 1170, MICH). longer than those opposite petals; anthers alike, glabrous; carpels completely connate in ovary; 4. Christianella paludicola (W. R. Anderson) W. styles 6 straight, subequal, stout, truncate at apex R. Anderson, comb. nov. Basionym: Mascagnia with the stigma terminal or nearly so. Fruit dry, paludicola W. R. Anderson, Contr. Univ. Michi- breaking apart into samaras separating from a prom- gan Herb. 16: 106. 1987. TYPE: Brazil. inent pyramidal torus (in Malpighiodes bracteosa the Amazonas: Rio Puru´s, Rio Ituxi, G. T. Prance mericarps with the wings reduced to coriaceous or et al. 13918 (holotype, INPA; isotypes, K, MICH, corky outgrowths); samara suborbicular or transverse- MO, NY, U, US, VEN). ly elliptical, the lateral wing, if developed, dominant, membranous with many looping anastomoses, cleft to 5. Christianella surinamensis (Kostermans) W. R. nut at apex, continuous at base, the margin entire, Anderson, comb. nov. Basionym: Mascagnia undulate, or coarsely toothed; dorsal wing small, free

Volume 16, Number 2 Anderson 193 2006 Segregates from Mascagnia

from lateral wing at base, extended forward at apex vicinity of Manaus, Amazonas, Brazil . .... through gap in lateral wing; intermediate winglets ...... 1.M. bracteosa 1b. Abaxial leaf hairs stalked, the branches straight or none or 1–several seta-like structures as high as serpentine and often ascending. width of dorsal wing or smaller; ventral areole broadly 3a. Mature leaves very densely and persistently ovate. tomentose below; petiole usually biglandular near middle; filaments abaxially sparsely The name Malpighiodes refers to a resemblance sericeous; Guyana, Suriname, and French Niedenzu saw between the reduced fruits of Mal- Guiana ...... 2.M. guianensis pighiodes bracteosa and the fruits of Malpighia 3b. Mature leaves glabrate or thinly hispid below; petiole eglandular; filaments glabrous; western (Niedenzu, 1908: 18). That fruit is quite different Amazonas,Brazil...... 3.M. leucanthele from the membranous samaras of Malpighiodes guianensis (W. R. Anderson) W. R. Anderson and 1. Malpighiodes bracteosa (Grisebach in Martius) Malpighiodes leucanthele (Grisebach in Martius) W. W. R. Anderson, comb. nov. Basionym: Mascag- R. Anderson (the fruits of Malpighiodes liesneri (W. R. nia bracteosa Grisebach in Martius, Fl. Bras. Anderson) W. R. Anderson are not known), but many 12(1): 97. 1858. Hiraea bracteosa (Grisebach in other shared character-states tie these four species Martius) Sagot, Ann. Sci. Nat. Bot., Se´r. 6, 12: together into a coherent group. Among those similari- 187. 1881. Malpighiodes spruceana Niedenzu, ties are the glands embedded in the abaxial leaf Verz. Vorles. Ko¨nigl. Lyceum Hosianum Brauns- surface, dichasial inflorescence with decussate flow- berg 1909/10: 31. 1909. Diplopterys bracteosa ers, large rounded bracteoles, long valvate biglandular (Grisebach in Martius) Niedenzu, Arbeiten Bot. sepals concealing the petals in bud and revolute in Inst. Ko¨nigl. Lyceum Hosianum Braunsberg 4: anthesis, glabrous yellow petals turning red in age, 20. 1912. Diplopterys spruceana (Niedenzu) and terminal or subterminal stigmas. In Davis et al. Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. Lyceum (2002), this genus (represented by Mascagnia brac- Hosianum Braunsberg 4: 21. 1912. Jubelina teosa Grisebach in Martius) was placed with 90% bracteosa (Grisebach in Martius) Cuatrecasas, bootstrap support as sister to a clade containing Alicia Webbia 13: 446. 1958. Mascagnia heterocarpa anisopetala and Callaeum septentrionale (A. Jussieu) W. R. Anderson, Mem. New York Bot. Gard. 32: D. M. Johnson. The suite of characters given above 216. 1981, replacement name for Malpighiodes immediately distinguishes Malpighiodes from those spruceana Niedenzu, non Mascagnia spruceana two genera. Niedenzu. TYPE: Brazil. Amazonas: vic. of Barra The name Malpighiodes was introduced in a footnote [Manaus], R. Spruce [1093] (lectotype, designat- (Niedenzu, 1908: 18). In that publication the di- ed by Anderson (1990a: 31), M; duplicates, BM, agnosis was so minimal as to be unacceptable and no CGE, E, G, GH, K, LE, NY). Figure 9J. species or combinations were proposed, so even See Anderson, 1990a: 31, for a discussion of the though it is clear that Niedenzu intended to publish lectotypification of Mascagnia bracteosa. The M sheet a generic name I consider it not to have been validly that was designated in 1990 as lectotype of that name published in 1908. A year later (Niedenzu, 1909: 31), is designated above as lectotype of Malpighiodes he gave an expanded diagnosis and published three spruceana, the type of the genus. species; I am accepting that as the valid publication of the name Malpighiodes. Malpighiodes comprises four species of northern 2. Malpighiodes guianensis (W. R. Anderson) W. R. Anderson, comb. nov. Basionym: Mascagnia South America. guianensis W. R. Anderson, Mem. New York Bot. Gard. 32: 213. 1981. TYPE: Guyana. Rockstone, KEY TO THE SPECIES OF MALPIGHIODES H. A. Gleason 825 (holotype, NY; isotypes, K). 1a. Abaxial leaf hairs sessile, straight, appressed. Figure 9A–I. 2a. Hairs on lateral axes of inflorescence straight to twisted, loose, mostly white; bracts and This species is known from Guyana, Suriname, and bracteoles densely white-tomentose on both French Guiana (Anderson, 1981: 215). sides; bracteoles borne at apex of peduncle; fruit unknown; Amazonas, Venezuela ...... 4.M. liesneri 3. Malpighiodes leucanthele (Grisebach in Mar- 2b. Hairs on inflorescence axes straight, ap- tius) W. R. Anderson, comb. nov. Basionym: pressed, mostly golden; bracts and bracteoles Mascagnia leucanthele Grisebach in Martius, Fl. densely sericeous on abaxial side, thinly sericeous on adaxial side; bracteoles borne Bras. 12(1): 96. 1858. TYPE: Brazil. Amazonas: at or somewhat below apex of peduncle; between Barcelos and Sa˜o Gabriel, R. Spruce mericarp wings coriaceous and rudimentary; [2070] (isotypes, BM, K, M). 194 Novon

The holotype is not in GOET (J. Heinrichs, pers. Niedenzuella lucida (A. Jussieu) W. R. Anderson] comm.). If Grisebach annotated any of the isotypes, (lectotype, designated by Morton (1968: 321)). that should be considered the holotype. Lianae lignosae, interdum frutescentes; lamina margine glandulis parvis instructa vel eglandulosa; stipulae parvae epipetiolares, interdum absentes; inflorescentia panicula 4. Malpighiodes liesneri (W. R. Anderson) W. R. ex pseudoracemis 2–30-floris floribus plerumque decussatis Anderson, comb. nov. Basionym: Mascagnia constans; sepala tenuia, imbricata, petala 6 tegentia in liesneri W. R. Anderson, Contr. Univ. Michigan alabastro, per anthesin revoluta; petala flava, glabra vel abaxialiter parce sericea; filamenta basi connata, recta, inter Herb. 17: 51. 1990. TYPE: Venezuela. Amazo- se 6 aequalia; antherae glabrae vel sparsim sericeae, inter nas: San Carlos de Rı´o Negro, R. Liesner 9063 se aequales; carpella basi connata in ovario, distaliter (holotype, MO; isotypes, MICH, VEN). distincta; styli recti et erecti vel recurvati, stigmate introrso vel paene terminali, apice dorsaliter rotundati vel truncati; 6 VIII. Niedenzuella W. R. Anderson, gen. nov. samara alis lateralibus evolutis vel rudimentariis, usque ad nucem apice basique incisis, quoque dimidio trapezoideo TYPE: Niedenzuella poeppigiana (A. Jussieu) dentato vel lobato vel usque ad nucem inciso. W. R. Anderson. Woody vines, sometimes shrubby. Petiole eglan- Hiraea sect. Hiraeanthele Grisebach, Linnaea 13: 239. 1839. dular or bearing 2–4 or more glands; lamina Tetrapterys sect. Hiraeanthele (Grisebach) C. V. Morton, Taxon 17: 322. 1968. TYPE: Hiraea multiflora eglandular or bearing few to many small glands on Grisebach [5 Niedenzuella lucida (A. Jussieu) W. R. margin; stipules very small, triangular or subulate, Anderson] (lectotype, designated by Morton (1968: borne on petiole at base or distally, as high as middle 322)). of petiole, persistent, sometimes lacking. Inflores- Tetrapterys sect. Stenantaeris Grisebach, Linnaea 13: 235. cence a terminal or lateral panicle with the ultimate 1839. TYPE: Tetrapterys acutifolia Cavanilles [5 Niedenzuella acutifolia (Cavanilles) W. R. Anderson] branches short pseudoracemes containing 2–30 (lectotype, designated by Morton (1968: 322)). flowers, the flowers mostly strictly decussate, occa- Tetrapterys subsect. Isopterae A. Jussieu, Ann. Sci. Nat. Bot., sionally subopposite distally; floriferous bracts Se´r. 2, 13: 262. 1840. TYPE: Tetrapterys fraxinifolia A. eglandular or biglandular; floriferous peduncle lack- Jussieu [5 Niedenzuella acutifolia (Cavanilles) W. R. Anderson] (lectotype, designated by Morton (1968: ing or short to long; bracteoles borne at apex of 321)). peduncle or slightly below, usually eglandular, rarely Hiraea sect. Trilophopterys Grisebach, Linnaea 22: 24. 1849. bearing 1 or 2 small abaxial glands. Sepals long and TYPE: Hiraea poeppigiana A. Jussieu [5 Niedenzuella imbricated, mostly concealing petals throughout poeppigiana (A. Jussieu) W. R. Anderson] (lectotype, enlargement of bud (separating to expose petals in designated by Morton (1968: 322)). Tetrapterys sect. Schizopterys Grisebach in Martius, Fl. Bras. bud in a few species), thin and membranous toward 12(1): 87. 1858. Tetrapterys subsect. Schizopterys margin, revolute in anthesis, the lateral 4 abaxially (Grisebach in Martius) Niedenzu, Verz. Vorles. Ko¨nigl. biglandular and the anterior eglandular or all 5 Lyceum Hosianum Braunsberg 1909/10: 20. 1909. eglandular; corolla bilaterally symmetrical to nearly Tetrapterys ser. Schizopterys (Grisebach in Martius) radial, the posterior petal with its claw usually Niedenzu in Engler, Pflanzenr. IV, 141: 183. 1928. TYPE: Tetrapterys poeppigiana (A. Jussieu) Grisebach somewhat longer or thicker than the 4 lateral petals; in Martius [5 Niedenzuella poeppigiana (A. Jussieu) W. petals small, bright yellow, glabrous or abaxially R. Anderson] (lectotype, designated by Morton (1968: thinly sericeous, erose or subentire; androecium 323)). radially symmetrical or nearly so; stamens 10, all fertile; filaments connate at base, straight, alike or the Niedenzu (1928: 183) used the name Tetrapterys 2 opposite the posterior-lateral petals thicker than sect. Microphyllaris Niedenzu, Verz. Vorles. Ko¨nigl. others, glabrous or abaxially sericeous; anthers 6 Lyceum Hosianum Braunsberg 1909/10: 20. 1909; alike, glabrous or sericeous; carpels connate at base in that name is a superfluous synonym of Tetrapterys ovary, distally distinct; styles stout, straight and erect sect. Schizopterys. to distally spreading or recurved, with large internal to Tetrapterys subg. Architetrapterys Niedenzu in Engler & nearly terminal stigmas, dorsally rounded or truncate Prantl, Nat. Pflanzenfam. III, 4: 57. 1890. TYPE: at apex. Fruit dry, breaking apart into samaras or all Tetrapterys poeppigiana (A. Jussieu) Grisebach in Martius [5 Niedenzuella poeppigiana (A. Jussieu) W. wings more or less reduced or rudimentary and R. Anderson] (lectotype, designated by Morton (1968: replaced by crests or irregular outgrowths, the 323)). mericarps separating from a flat or short-pyramidal Tetrapterys subsect. Isopterys Niedenzu, Verz. Vorles. Ko¨nigl. torus; samara (when not reduced) butterfly-shaped or Lyceum Hosianum Braunsberg 1909/10: 22. 1909. Tetrapterys ser. Isopterys (Niedenzu) Niedenzu in X-shaped, the lateral wings dominant, chartaceous Engler, Pflanzenr. IV, 141: 184. 1928. TYPE: with many fine strongly parallel veins, cleft to nut at Tetrapterys multiflora (Grisebach) A. Jussieu [5 apex and base, each side trapezoidal with the margin

196 Novon

coarsely toothed or sometimes deeply and irregularly samara. Wherever one puts one of those species, the lobed, or divided to the nut into 2 elongated, 6 others will have to follow. Davis et al. (2002), placed parallel-sided wings; dorsal wing or crest small, Tetrapterys sect. Microphyllaris (as represented by distinct from lateral wings, often extended forward at Mascagnia stannea) in an unsupported clade with apex between lateral wings; irregular outgrowths Dicella, Tricomaria, Heteropterys, Mascagnia chasei present between dorsal and lateral wings in some [Carolus chasei], and Hiptage.Thattreegivesus species; ventral areole ovate. little help in understanding the relationships of Tetrapterys sect. Microphyllaris,butitdoesremove The name Niedenzuella honors the memory of Franz the group effectively from Mascagnia s. str. and also Josef Niedenzu (1857–1937), who was the second from Tetrapterys s. str., represented by T. discolor (G. specialist (following Adrien de Jussieu) to monograph Meyer) DC., which came out quite separately in the the entire family Malpighiaceae. Niedenzu has had his 2002 tree. There is no support in morphology for critics, but his 1928 treatment of the family in Das placing this group of species in Dicella, Tricomaria, Pflanzenreich remains an essential reference. He was Heteropterys, Carolus,orHiptage (see discussion never able to study the plants in the field, had very under Carolus), so I see no alternative to recognizing little material from some areas, and never got the them as a genus. It remains to be seen which type of opportunity to visit several critically important samara, butterfly-shaped or X-shaped, was ancestral herbaria, especially those in Paris and London. Given in this group. those limitations, he did amazingly well. Niedenzuella comprises 16 species of Central and In 1928 Niedenzu recognized as Tetrapterys sect. South America. Microphyllaris a group of 20 species, 18 of which share a suite of characters that distinguish them from KEY TO THE SPECIES OF NIEDENZUELLA other sections of Tetrapterys: tiny epipetiolar stipules, 1a. Lamina at maturity tomentose, thinly sericeous, or marginal leaf glands, elongated, very thin, imbricated glabrous below, the hairs never dense enough to sepals that more or less conceal the petals in bud and completely hide the epidermis. are revolute in anthesis, and carpels that are distinct 2a. Abaxial hairs on young leaves loose and distally in the ovary. (Two of the species Niedenzu serpentine to strongly twisted, the lamina persistently tomentose or glabrescent in age. included in that section, T. heteropetala and T. 3a. Samara with the nut mostly smooth renidens, are treated above as Carolus renidens and between lateral and dorsal wings, occa- will not be discussed further here.) Most of those 18 sionally bearing 1 or 2 elongated out- species have samaras with four lateral wings, quite growths; inflorescence composed of umbels or dense corymbs with the like those in other sections of Tetrapterys, so workers internodes between pairs of flowers, if from Cavanilles to Cuatrecasas have been content to developed, mostly shorter than peduncles leave them in that genus, which has traditionally been except occasionally the most proximal defined by that X-shaped samara. Also in the 1928 pair more distant from the others; eastern Ecuador, Amazonian Peru, and western work Niedenzu recognized in Mascagnia the species Bolivia ...... 11.N. peruviana M. sericans Niedenzu and M. stannea (Grisebach) 3b. Samara with several well-developed and Niedenzu, which have the same characters given obvious (although often low and rounded) above for Tetrapterys sect. Microphyllaris but butter- outgrowths between lateral and dorsal fly-shaped samaras, with the lateral wings dentate but wings; inflorescence composed of pseudoracemes with at least the more proxi- not divided to the nut; in doing this Niedenzu followed mal pair 6 well separated from the distal his common practice of assigning species to genera on pairs; southeastern Bolivia, Paraguay, the basis of their fruits, regardless of what the other and southern and eastern Brazil. characters might say about probable relationships. In 4a. Lamina narrowly to broadly elliptical and rounded or obtuse to acute 1981 I treated in Mascagnia three species of this at apex; eastern Brazil, found in the group (M. benthamiana (Grisebach in Martius) W. R. Planalto as far west as the Distrito Anderson, M. castanea (Cuatrecasas) W. R. Anderson, Federal. and M. poeppigiana (A. Jusssieu) W. R. Anderson) 5a. Petals abaxially sparsely to densely hairy; marginal leaf that have butterfly-shaped samaras or mericarps glands, if present, protuberant, derived from such samaras by reduction of the wings, often cylindrical; calyx glands, but I have since concluded that it was a mistake to if present, 1.5–2.5 mm long, separate those species from the others in Niedenzu’s narrowlyelliptical...... 10.N. multiglandulosa Tetrapterys sect. Microphyllaris; the unifying char- 5b. Petals glabrous; marginal leaf acters listed above are more significant than the glands discoid, not or hardly lobing or lack of lobing in the lateral wings of the protuberant; calyx glands, if Volume 16, Number 2 Anderson 197 2006 Segregates from Mascagnia

present, 1–1.3 mm long, broad- tic South America from ly ovate or obovate ...... Guyana to southeastern Bra- ...... 16.N. warmingiana zil, and with populations in 4b. Lamina 6 narrowly lanceolate, ta- Mato Grosso and Acre, as pered distally to a narrowly acute well as Bolivia, Peru, and or acuminate apex; southeastern Ecuador ...... 1.N. acutifolia Bolivia, southwestern Brazil (Mato 10b. Lamina mostly obtuse (to Grosso and Rondoˆnia), and southern rounded) at apex, sometimes Paraguay ...... 15.N. suaveolens acute, soon quite glabrate, 2b. Abaxial hairs on young leaves straight and 6 even on abaxial midrib; peti- strongly appressed, the lamina initially dense- ole mostly eglandular (rarely ly to sparsely sericeous below, at maturity biglandular), mostly soon gla- thinly sericeous to glabrate. brate; anthers sparsely to 6a. Samara with the nut mostly smooth densely pilose; coastal south- between lateral and dorsal wings, rarely eastern Brazil (Rio de Janeiro bearing 1 or 2 outgrowths. and Sa˜oPaulo)...... 7a. Petiole mostly bearing 2(–4) large ...... 9.N. mogoriifolia discoid glands 0.7–1.5 mm diam.; 1b. Lamina, even at maturity, persistently metallic- northern Venezuela . . 2. N. caracasana sericeous below (rarely patchily glabrescent in 7b. Petiole mostly eglandular, occasion- age), the vesture so dense as to completely hide the ally bearing 2 small glands up to epidermis. 0.5 mm diam.; southeastern Brazil. 11a. Samara tetrapteroid, i.e., bearing 4 6 equal 8a. Petals abaxially sericeous; lam- well-developed lateral wings. ina soon quite glabrous on both 12a. Samara nut smooth between dorsal and sides, usually eglandular but lateral wings; Meta, Colombia ...... rarely with impressed marginal ...... 8.N. metensis glands; coastal Bahia to Rio de 12b. Samara nut bearing outgrowths be- Janeiro ...... 4.N. glabra tween dorsal and lateral wings; south- 8b. Petals glabrous or nearly so, eastern Brazil, Paraguay, and north- occasionally with a few hairs; easternArgentina...... 13.N. sericea lamina abaxially 6 evenly and 11b. Samara mascagnioid, i.e., bearing 2 6 equal persistently sericeous or even- well-developed lateral wings, or with the tually glabrescent, the hairs lateral wings variously and irregularly often very small and difficult dissected and reduced. to see; lamina eglandular or 13a. Petals abaxially sericeous with many with 1–several prominent mar- hairs thinly scattered over much of ginal glands; Minas Gerais and the limb; samara nut bearing out- Espı´rito Santo to Santa Cata- growths between dorsal and lateral rina ...... 6.N. lucida wings; southeastern Brazil, Paraguay, 6b. Samara with several well-developed and andnortheasternArgentina...... obvious (although often low and rounded) ...... 13.N. sericea outgrowths between lateral and dorsal 13b. Petals glabrous; samara nut mostly wings, or with several prominent ribs at smooth (or at most rugose) between right angles to dorsal wing. dorsal and lateral wings, but some 9a. Lamina thinly but evenly and per- populations in Para´, Brazil, with sistently sericeous below at maturity several elongated intermediate out- (hairs sometimes difficult to see, growths; Central and South America. especially on older leaves, where 14a. Sepals separating to expose they may be flattened and scale- petals during enlargement of like); petals glabrous or bearing flower bud; anthers 0.5–1 mm a few hairs abaxially at base of long; lateral petals with limb limb; southeastern Brazil from Bahia 1.5–3.5 mm long; floriferous to Rio de Janeiro . . . 5. N. leucosepala bracts up to 1 mm long, 9b. Lamina soon glabrate below or with eglandular; pedicels up to some hairs persistent on base of 0.5 mm diam., sessile or raised midrib and nearby; petals mostly on peduncles up to 0.5(21) mm thinly but obviously sericeous abaxi- long. ally on limb; South America. 15a. Lamina abaxially very 10a. Lamina mostly acute or acu- tightly metallic-seri- minate at apex but sometimes ceous, with most hairs obtuse, usually bearing long very short, 0.2–0.4 mm hairs persistent on abaxial long; wings of samara base of midrib, even in with short tight hairs age; petiole bearing (1)2–4 like those of lamina, to glands or eglandular, 6 per- glabrate; marginal leaf sistently sericeous; anthers glands, if present, 0.4– glabrous or sericeous; Atlan- 0.8 mm diam.; limb of 198 Novon

lateral petals 2.5–3.5 mm Tetrapterys guilleminiana A. Jussieu, Ann. Sci. Nat. Bot., long; widespread in South Se´r. 2, 13: 263. 1840. TYPE: Brazil. Near Sebastiano´- America .... 12.N. poeppigiana polis [Rio de Janeiro]: C. Gaudichaud 959 (lectotype, 15b. Lamina abaxially relatively designated here, P-JU 11706+B [MICH, WRA neg. 81- loosely sericeous with the 17-17]; duplicate, P). outer, longer hairs 0.8– 1.2 mm long; wings of samara As syntypes Jussieu cited collections by Gaudi- with long, relatively loose chaud and Guillemin, both represented in P by hairs like those of lamina, to several sheets. As lectotype I have selected a speci- glabrate; marginal leaf glands 0.2–0.3 mm diam.; limb of men in the Jussieu Herbarium that was annotated by lateral petals 1.5–2.2 mm Jussieu and has both flowers and fruits. long; southeastern Peru and adjacent Brazil and Bolivia Tetrapterys lancifolia A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, ...... 7.N. mater-dei 13: 263. 1840. Tetrapterys acutifolia var. lancifolia (A. 14b. Sepals concealing petals almost Jussieu) Niedenzu, Verz. Vorles. Ko¨nigl. Lyceum throughout enlargement of flower Hosianum Braunsberg 1909/10: 27. 1909. TYPE: bud; anthers (0.8–)1–1.5 mm long; Brazil. Minas Gerais: ‘‘Ilhas de Aboithe´,’’ P. Claussen lateral petals with limb 3–5 mm s.n. (lectotype, designated here, P [F neg. 35605]; long; floriferous bracts at least isotypes, G, K, NY, P, P-JU). 1 mm long, often longer, bearing 2 large abaxial glands or eglandu- The Claussen collection cited above is the sole lar; pedicels 0.5–1 mm diam., sub- type. It is represented in P by several sheets, of which sessile or raised on peduncles up to I have selected as lectotype one that was annotated by 4 mm long. 16a. Lamina abaxially dark Jussieu. brown-sericeous; peduncles Bunchosia laxiflora Gardner, London J. Bot. 1: 168. 1842. 0.5–1.5 mm long; Amazo- TYPE: Brazil. Rio de Janeiro: 1837, G. Gardner 17 nas, Venezuela, and upper (holotype, K). Rio Negro, Brazil ...... Tetrapterys heteroalata Kralik, Denkschr. Kaiserl. Akad...... 3.N. castanea Wiss., Math.-Naturwiss. Kl. 79: 277. 1908. TYPE: 16b. Lamina abaxially golden- Brazil. Saõ Paulo: near the city of Saõ Paulo, Aug. or silvery-sericeous; pedun- 1901, R. Wettstein & V. Schiffner (holotype, WU not cles mostly 1–4 mm long; seen). Costa Rica, Panama, and Amazonian South America This species is widely distributed, being known south to Bolivia and Para- from Atlantic South America from Guyana (Jansen- guay...... 14.N. stannea Jacobs 3192, MICH) to southeastern Brazil, with populations also in Mato Grosso (Thomas et al. 1. Niedenzuella acutifolia (Cavanilles) W. R. 4100, MICH) and Acre (Lowrie et al. 701, MICH), as Anderson, comb. nov. Basionym: Tetrapterys well as Bolivia (Solomon 17711, MO), Peru (Gentry et acutifolia Cavanilles, Diss. 9: 433, pl. 261. al. 18588, MICH), and Ecuador (Gudinõ 1343, 1790. TYPE: French Guiana. Stoupy s.n. (holo- MICH). type, P-JU 11707+A [F neg. 37465]).

Tetrapterys punctulata A. Jussieu in A. St.-Hilaire, Fl. Bras. 2. Niedenzuella caracasana W. R. Anderson, sp. Merid. 3: 10. 1832 [1833]. TYPE: Brazil. Minas Gerais: nov. TYPE: Venezuela. Distrito Federal: Cerro Sumidor, A. Saint-Hilaire s.n. (lectotype, designated here, P [MICH, WRA negs. 81-23-13 & 14]; duplicate, Naiguata´, Fila de las Delicias, north side, P). 1500 m, 10 Apr. 1976 (fr), B. J. Manara s.n. (holotype, MICH). There are four potential lectotypes in P, representing two or three collections from Minas Gerais. I am Niedenzuellae acutifoliae similis, sed samara sine alulis designating as lectotype a specimen annotated by inter alas laterales et cristam dorsalem differt. Jussieu; it is a good representative of the species. Woody vine, the stem initially sericeous with Tetrapterys fraxinifolia A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, straight, tightly appressed hairs, soon glabrate. Petiole 13: 263. 1840. TYPE: Brazil. Near Sebastiano´polis 4–6 mm long, sericeous to glabrate, bearing between [Rio de Janeiro]: Vauthier 460 (lectotype, designated middle and apex 2(–4) raised, discoid glands 0.7– here, P [F neg. 35599]; isotypes, P, P-JU 11709). 1.5 mm diam.; lamina of larger leaves 7–8.5 6 2.5– Vauthier 460 was the sole type collection. It is 4 cm, elliptical or narrowly ovate, cuneate at base, represented in P by four sheets; as lectotype I am acute to obtuse at apex, coriaceous with the reticulum selecting the one in the general herbarium that was of veinlets visible to obscure in dried leaves, bearing annotated by Jussieu. 2–4 discoid marginal glands 0.7–1.3 mm diam. on Volume 16, Number 2 Anderson 199 2006 Segregates from Mascagnia

each side in proximal half, initially sericeous on both son, Mem. New York Bot. Gard. 32: 218. 1981. sides with straight, tightly appressed hairs but TYPE: Brazil. Amazonas: upper Rio Negro, R. E. glabrate at maturity or thinly sericeous below, Schultes & F. Lo´pez 10014 (holotype, US; especially on midrib, the principal lateral veins 4–6 isotypes, IAN, MICH, US). on each side; stipules 0.2–0.5 mm long, borne on petiole between base and middle. Inflorescence a short This species is known from Amazonas, Venezuela axillary panicle, sericeous to glabrate, comprising (Davidse et al. 17463, MICH), and upper Rio Negro, several short-stalked umbels or pseudoracemes of 2 or Brazil. 4 flowers; bracts 0.8–1.5 mm long; peduncles 2–5 mm long; bracteoles 1–1.5 mm long, borne between 4. Niedenzuella glabra (Sprengel) W. R. Anderson, middle and apex of peduncle; pedicels 7–9 mm long. comb. nov. Basionym: Hiraea glabra Sprengel, Sepals (in fruit) revolute at apex, abaxially sericeous, Neue Entdeck. 2: 154. 1821. Tetrapterys glabra adaxially glabrous, the anterior eglandular, the lateral (Sprengel) Grisebach in Martius, Fl. Bras. 12(1): 4 biglandular with the glands 2–2.5 mm long, 82. 1858. TYPE: Brazil. Espı´rito Santo: Between narrowly elliptical; petals not seen, presumably Vitoria and Bahia, F. Sellow II it. 107 (lectotype, yellow; anthers (seen only as remnants in fruit) designated here, NY). apparently glabrous; styles dorsally rounded or truncate at apex. Samara consistently bearing 4 lateral The specimen in NY that I am designating lectotype wings and a dorsal crest, loosely sericeous on nut, came from Niedenzu’s herbarium and originated in B, sericeous to glabrescent on wings; lateral wings 10– so it is the logical replacement for the B holotype that 17 6 7–10 mm, the upper and lower pairs subequal; was destroyed. The type was cited in the protologue as dorsal crest 2–2.5 mm wide; nut smooth or rugose but ‘‘Otto,’’ certainly a slip for Sellow. completely lacking outgrowths between lateral and This species is known from the Brazilian states of dorsal wings; torus 1.8 mm high. Bahia (Mori et al. 9642, MICH), Espı´rito Santo, and Rio de Janeiro (Gentry et al. 49471, MICH). Phenology. Collected with flower buds in May, and with fruits in February, April, and November. 5. Niedenzuella leucosepala (Grisebach) W. R. Ecology and distribution. Wet, evergreen forests Anderson, comb. nov. Basionym: Hiraea leuco- at 900–1500 m in the mountains above Caracas, on sepala Grisebach, Linnaea 13: 240. 1839. the north coast of Venezuela. Tetrapterys leucosepala (Grisebach) A. Jussieu, Etymology. The epithet caracasana refers to the Arch. Mus. Hist. Nat. 3: 531. 1843. TYPE: source of all the known collections. In most of the Southern Brazil: F. Sellow (holotype, B{ [F neg. characters that I can observe in this material it 12734]; isotype, M not seen]. resembles Niedenzuella acutifolia, which I have not Grisebach cited as type a Sellow collection from seen from Venezuela. Niedenzuella caracasana is southern Brazil, without number or precise locality. distinguished by its samara, which lacks the out- Niedenzu (1928: 194) said the collection was growths between dorsal and lateral wings that are Sellow II it. 88 from between Rio de Janeiro and always present in N. acutifolia. In addition, N. Campos, Sa˜o Paulo. He also cited what he considered caracasana has thick, often obtuse laminas bearing an isotype at M. The latter specimen might be very large discoid marginal glands and similar glands a candidate for lectotype of this name, but I do not on the petioles. When flowers of N. caracasana want to make that designation until I have verified its become available for study they may furnish addi- identity. tional distinguishing characters. Tetrapterys martiana Niedenzu, Verz. Vorles. Ko¨nigl. Paratypes. VENEZUELA. Aragua: Cordillera Interior, Lyceum Hosianum Braunsberg 1909/10: 29. 1909. entre El Paujı´ y El Socorro, hacia Cerro La Ojeden˜a, al sur de TYPE: Brazil. Rio de Janeiro: Fl. Itahipe, Mar.–Apr. El Consejo, Steyermark & Perkins 122040 (MICH). Distrito 1822, L. Riedel (lectotype, designated here, K [MICH, Federal: Cerro Naiguata´, Lomas de Las Delicias, entre WRA neg. 91-2-4]; duplicate, K). Quebrada de Basenilla y Quebrada Guayoyo, Steyermark 92097 (NY); north-facing slopes, 7 km W of Hacienda Niedenzu cited syntypes collected by Luschnath, Cocuizal, Quebrada Frontina, 7 Feb. 1967, Manara s.n. (NY). Martius, and Riedel (under Langsdorff). The Riedel collection of Mar.–Apr. 1822, is excellent and 3. Niedenzuella castanea (Cuatrecasas) W. R. representative material, and I have selected the better Anderson, comb. nov. Basionym: Heteropterys of the two sheets at K to serve as lectotype. castanea Cuatrecasas, Webbia 13: 475. 1958. This species occurs in southeastern Brazil from Mascagnia castanea (Cuatrecasas) W. R. Ander- Bahia (Amorim 3616, MICH) to Rio de Janeiro. 200 Novon

6. Niedenzuella lucida (A. Jussieu in A. St.- Janeiro], A. Saint-Hilaire s.n. (lectotype, desig- Hilaire) W. R. Anderson, comb. nov. Basionym: nated here, P [F neg. 35609]). Tetrapterys lucida A. Jussieu in A. St.-Hilaire, Fl. Bras. Merid. 3: 11. 1832 [1833]. TYPE: There are two sheets in P that were collected by St.- Brazil. Minas Gerais: Serra da Caraça, A. Saint- Hilaire from near Rio, both annotated with the Hilaire s.n. (lectotype, designated here, P [F neg. basionym by Jussieu. They appear to represent 35608]; isotypes, P). different gatherings, because one is in flower and immature fruit and the other is in old fruit. I believe There was a single type collection, represented in P they should be regarded as syntypes, and have by three specimens, all annotated by Jussieu. I have designated the better of them as lectotype. selected the best of the three to serve as lectotype. This species is known from coastal southeastern Hiraea multiflora Grisebach, Linnaea 13: 239. 1839. Brazil, in Rio de Janeiro and Saõ Paulo (Gentry & Tetrapterys multiflora (Grisebach) A. Jussieu, Arch. Zardini 49400, MICH). Mus. Hist. Nat. 3: 533. 1843. TYPE: Southern Brazil. [Rio de Janeiro]: F. Sellow [III it. B2249 c2287, fide 10. Niedenzuella multiglandulosa (A. Jussieu) W. Niedenzu, 1928] (lectotype, designated here, NY). R. Anderson, comb. nov. Basionym: Tetrapterys The NY sheet designated as lectotype to replace the multiglandulosa A. Jussieu, Ann. Sci. Nat. Bot., lost holotype is from the Niedenzu herbarium, and is Se´r. 2, 13: 264. 1840. TYPE: Brazil. Minas marked as coming from B. It may well be from the Gerais: Curvelo, Claussen s.n. (lectotype, desig- holotype; in any event, it is surely the best candidate nated here, P [MICH, WRA negs. 81-24-1 & 2]; for lectotype. isotypes, G, P, P-JU). Figure 10. Tetrapterys cuneifolia Kralik, Denkschr. Kaiserl. Akad. There are several Claussen sheets in P, all from Wiss., Math.-Naturwiss. Kl. 79: 278. 1908. TYPE: Brazil. Saõ Paulo: near Rio Grande between Santos and Minas Gerais, which is the locality Jussieu gave; they the city of Saõ Paulo, 1902, M. Wacket (holotype, WU may represent more than one collection. The two not seen). sheets from Curvelo represent the best candidates for Tetrapterys cuneifolia f. eglandulosa Kralik, Denkschr. lectotype, because they were both annotated by Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 79: 278. 1908. TYPE: Brazil. Saõ Paulo: near Campo Grande Jussieu. I have selected the better of them as between Santos and the city of Saõ Paulo, 1902, M. lectotype. Wacket (holotype, WU not seen). Tetrapterys adenodon Miquel, Linnaea 22: 548. 1849. TYPE: This Brazilian species is known from Minas Gerais Brazil. Minas Gerais: near Caldas, A. Regnell I 28 and Espı´rito Santo (Hatschbach 59748, MICH) to (holotype, U; isotypes, GH, K, MEL, NY). Santa Catarina (Occhioni 3749, MICH). 11. Niedenzuella peruviana (Niedenzu) W. R. Anderson, comb. nov. Basionym: Tetrapterys 7. Niedenzuella mater-dei (Cuatrecasas) W. R. multiglandulosa var. peruviana Niedenzu, Verz. Anderson, comb. nov. Basionym: Mascagnia Vorles. Ko¨nigl. Lyceum Hosianum Braunsberg mater-dei Cuatrecasas, Brittonia 11: 167. 1959. 1909/10: 29. 1909. TYPE: Peru. ‘‘Tambo TYPE: Peru. Madre de Dios: Iberia, R. E. Azalaya,’’ A. Weberbauer 1125 (holotype, B{ [F Schultes 6228 (holotype, US; isotypes, COL, neg. 12739]). GH, K, NY). This species is known from eastern Ecuador (Neill This species is known from southeastern Peru and 5976, MICH), Amazonian Peru, and western Bolivia adjacent Brazil (Pardo et al. 120, MICH) and Bolivia (Brummitt et al. 19359, MICH). (Rueda 947, MICH). 12. Niedenzuella poeppigiana (A. Jussieu) W. R. 8. Niedenzuella metensis (Cuatrecasas) W. R. Anderson, comb. nov. Basionym: Hiraea poep- Anderson, comb. nov. Basionym: Tetrapterys pigiana A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, metensis Cuatrecasas, Webbia 13: 419. 1958. 13: 260. 1840. Tetrapterys poeppigiana (A. TYPE: Colombia. Meta: Villavicencio, Bro. Jussieu) Grisebach in Martius, Fl. Bras. 12(1): Apollinaire s.n. (holotype, US). 87. 1858. Mascagnia poeppigiana (A. Jussieu) 9. Niedenzuella mogoriifolia (A. Jussieu in A. St.- W. R. Anderson, Mem. New York Bot. Gard. 32: Hilaire) W. R. Anderson, comb. nov. Basionym: 219. 1981. Lectotype collection, designated by Tetrapterys mogoriifolia A. Jussieu in A. St.- Anderson (1981: 219): Brazil. Amazonas: ‘‘Prov. Hilaire, Fl. Bras. Merid. 3: 11. 1832 [1833]. Rio Negro,’’ E. Poeppig 2799 (lectotype, desig- TYPE: Brazil. Near Sebastiano´polis [Rio de nated here, G; duplicates, MICH, NY, P, US, W).

202 Novon

Ann. Sci. Nat. Bot., Se´r. 2, 13: 264. 1840. TYPE: Banisteria discolor (Donnell Smith) Niedenzu in Engler, Brazil. Near Rio de Janeiro, A. St.-Hilaire catal. Pflanzenr. IV, 141: 451. 1928. TYPE: Costa Rica: San 1 Pedro de la Calabaza, A. Tonduz s.n. [CR 10924] A no. 380 (lectotype, designated here, P [F neg. (holotype, CR; isotype, US). 35612]; isotypes, P). Mascagnia sericans Niedenzu in Chodat & Hassler, Bull. Herb. Boissier, Se´r. 2, 7: 284. 1907. TYPE: Paraguay. The St.-Hilaire collection cited above was the sole Bellavista, E. Hassler 7825 (lectotype, designated here, type. It is represented in P by three sheets; I have G; isotypes, G, K, MICH, NY). selected the best of those, annotated by Jussieu, to serve as lectotype. The holotype sheet of Hassler 7825 was destroyed at B. There are four isotypes at G; the lectotype is Hiraea laurifolia A. Jussieu, Ann. Sci. Nat. Bot., Se´r. 2, 13: the isotype that was annotated lectotype by W. R. 260. 1840. Tetrapterys laurifolia (A. Jussieu) Grisebach Anderson. in Martius, Fl. Bras. 12(1): 88. 1858. TYPE: Brazil. Minas Gerais: Tejuco [Diamantina], C. F. P. Martius Mascagnia sericans subsp. buricana Cuatrecasas & Croat, s.n. (holotype, M; isotype, P-JU). Ann. Missouri Bot. Gard. 67: 912. 1980 [1981]. TYPE: Tetrapterys chalcophylla A. Jussieu, Ann. Sci. Nat. Bot., Se´r. Panama. Burica Peninsula, W of Puerto Armuelles, T. 2, 13: 264. 1840. TYPE: Brazil. Bahia: J. Blanchet Croat 22495 (holotype, MO; isotype, NY). 1062 (holotype, G). Tetrapterys glandulosa Grisebach in Martius, Fl. Bras. 12(1): This species is defined broadly here, and with 83. 1858. TYPE: Brazil. Minas Gerais: near Sa˜o that circumscription it is known from Costa Rica, Caetano, G. Gardner 4477 (holotype, K [MICH, WRA Panama, and Amazonian South America south to Boli- neg. 81-9-13]; isotypes, BM, K). Tetrapterys ligustrifolia Niedenzu, Arbeiten Bot. Inst. Ko¨nigl. via (Solomon & Urcullo 14137, MICH) and Para- Lyceum Hosianum Braunsberg 4: 14. 1912, non T. guay. ligustrifolia A. Jussieu in A. St.-Hilaire, 1833. TYPE: Brazil. Minas Gerais: 1845, J. F. Widgren (lectotype, designated here, NY; duplicates, LE, MICH, U). 15. Niedenzuella suaveolens (A. Jussieu) W. R. Anderson, comb. nov. Basionym: Tetrapterys Niedenzu had three syntypes, collected by Wid- suaveolens A. Jussieu, Ann. Sci. Nat. Bot., Se´r. gren, Lindberg, and Mose´n, all in Minas Gerais. I have 2, 13: 263. 1840. TYPE: Bolivia. Santa Cruz: seen only the Widgren collection, and am designating Santiago de Chiquitos, A. d’Orbigny 910 (holo- as lectotype the NY sheet that came from Niedenzu’s type, P; isotype, G). herbarium. This species is known from southeastern Brazil, This species is known from southeastern Bolivia, Paraguay (Zardini & Gamarra 55629, MICH), and southwestern Brazil (Mato Grosso [Hatschbach 63892, northeastern Argentina (Anderson 13612, MICH). MICH] and Rondoˆnia [Anderson 12156, MICH]), and southern Paraguay (Tressens et al. 4770, MICH).

14. Niedenzuella stannea (Grisebach) W. R. 16. Niedenzuella warmingiana (Grisebach) W. R. Anderson, comb. nov. Basionym: Heteropterys Anderson, comb. nov. Basionym: Heteropterys stannea Grisebach, Vidensk. Meddel. Dansk warmingiana Grisebach, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 1853: 46. 1854. Dansk. Naturhist. Foren. Kjøbenhavn 1875: Mascagnia stannea (Grisebach) Niedenzu, Verz. 135. 1875. Tetrapterys warmingiana (Grisebach) Vorles. Ko¨nigl. Lyceum Hosianum Braunsberg Niedenzu, Verz. Vorles. Ko¨nigl. Lyceum Hosia- 1912/13: 12. 1912. TYPE: Costa Rica. Monte num Braunsberg 1909/10: 20. 1909. TYPE: Aguacate, Nov. 1846, A. Ørsted (holotype, GOET Brazil. Minas Gerais: Lagoa Santa, J. Warming not seen; isotype, C). Figure 11. s.n. (holotype, C not seen [F neg. 21354]). Tetrapterys benthamiana Grisebach in Martius, Fl. Bras. Niedenzu (1928: 183) implied that Grisebach had 12(1): 88. 1858. Malpighiodes benthamiana (Grise- bach in Martius) Niedenzu, Verz. Vorles. Ko¨nigl. two syntypes, both collected by Warming in 1863, on Lyceum Hosianum Braunsberg 1909/10: 31. 1909. 14 Sep. and 5 Nov. If that is true, one should be Mascagnia benthamiana (Grisebach in Martius) W. selected as lectotype. R. Anderson, Mem. New York Bot. Gard. 32: 217. 1981. TYPE: Brazil. Para´: near Santare´m, Nov.– Mar. Acknowledgments. Karin Douthit drew the illus- 1849–50. R. Spruce s.n. (holotype, GOET not seen; trations with her usual skill and attention to detail. My isotypes, BM, CGE, E, G, GH, K, M [F neg. 19338], collaborators, Christiane Anderson and Charles Davis, NY). Banisteria cornifolia var. discolor Donnell Smith, Bot. Gaz. have given unfailing assistance and advice. Roy (Crawfordsville) 24: 391. 1897. Banisteriopsis discolor Vickery and Vicki Noble at BM sent me notes on (Donnell Smith) Small, N. Amer. Fl. 25: 132. 1910. and a scan of the type of Triopterys hiraea. Piet

204 Novon

Stoffelen at BR sent me a scan of the holotype of Cameron, K. M., M. W. Chase, W. R. Anderson & H. G. Mascagnia doniana and helpful replies to my Hills. 2001. Molecular systematics of Malpighiaceae: Evidence from plastid rbcL and matK sequences. Amer. questions on morphological details. Fernand Jacque- J. Bot. 88: 1847–1862. moud at G arranged for me to receive scans of their Cavanilles, J. 1790. Nona Dissertatio Botanica. Madrid. isotypes of Heladena hassleriana and Mascagnia Colla, L. A. 1824. Hortus Ripulensis. Torino. multiglandulosa. Jochen Heinrichs at GOET was most Davis, C. C., W. R. Anderson & M. J. Donoghue. 2001. helpful with information about types in the Grisebach Phylogeny of Malpighiaceae: Evidence from chloroplast ndhF and trnL-F nucleotide sequences. Amer. J. Bot. 88: herbarium. At M, Gu¨nter Gerlach and Alison Davies 1830–1846. arranged for me to receive scans of the type of Hiraea ———, C. D. Bell, S. Mathews & M. J. Donoghue. 2002. laurifolia, and Franz Schuhwerk directed me to BR Laurasian migration explains Gondwanan disjunctions: when I was seeking the type of Mascagnia doniana. Evidence from Malpighiaceae. Proc. Natl. Acad, U.S.A. Ju¨rgen Do¨bereiner and Carlos H. Tokarnia donated 99: 6833–6837. Grisebach, A. H. R. 1858. Malpighiaceae. Columns 1–124 in especially interesting specimens of plants belonging C. F. P. Martius (editor), Flora Brasiliensis, Vol. 12(1). to this group of genera. I am grateful to the curators of Leipzig. the following herbaria for sending to MICH gifts and Jacquin, N. J. 1760. Enumeratio Systematica Plantarum. T. loans of Malpighiaceae, and for their hospitality and Haak, Leiden. assistance when I visited their institutions: A, BM, C, Johnson, D. M. 1986. Revision of the Neotropical genus CEPEC, CGE, COL, CR, E, F, G, GB, GH, IAN, Callaeum (Malpighiaceae). Syst. Bot. 11: 335–353. Jussieu, Adrien de. 1843. Monographie de la famille des INPA, K, L, LE, LIL, M, MEL, MO, NY, P, S, U, US, Malpighiace´es. Arch. Mus. Hist. Nat. 3: 5–151, 255–616 VEN, W. pl. 1–23. Linnaeus, C. 1753. Species Plantarum, Vol. 1. Stockholm. Lombello, R. A. & E. R. Forni-Martins. 2002. Cytogenetics of twelve species of Malpighiaceae A. Jussieu from Literature Cited southeastern Brazil. Caryologia 55: 241–250. Anderson, W. R. 1980. Notes on Mascagnia in South Lowrie, S. R. 1982. The Palynology of the Malpighiaceae America. Contr. Univ. Michigan Herb. 14: 17–23. and Its Contribution to Family Systematics. Ph.D. Disser- ———. 1981. The botany of the Guayana Highland Part XI, tation. University of Michigan, Ann Arbor. [University Malpighiaceae. Mem. New York Bot. Gard. 32: 21–305. Microfilms #82-24999.] ———. 1987. Notes on Neotropical Malpighiaceae—II. Morton, C. V. 1968. A typification of some subfamily, Contr. Univ. Michigan Herb. 16: 55–108. sectional, and subsectional names in the family Mal- ———. 1990a. The taxonomy of Jubelina (Malpighiaceae). pighiaceae. Taxon 17: 314–324. Contr. Univ. Michigan Herb. 17: 21–37. Niedenzu, F. 1908. De genere Mascagnia. Arbeiten Bot. Inst. ———. 1990b. Notes on Neotropical Malpighiaceae—III. Ko¨nigl. Lyceum Hosianum Braunsberg 3: 3–29. Contr. Univ. Michigan Herb. 17: 39–54. ———. 1909. De genere Tetrapteryge. Verz. Vorles. Ko¨nigl. ———. 1993. Notes on Neotropical Malpighiaceae—IV. Lyceum Hosianum Braunsberg 1909/10: 3–56. Contr. Univ. Michigan Herb. 19: 355–392. ———. 1928. Malpighiaceae. Pp. 1–870 in A. Engler ———. 1997. Excentradenia, a new genus of Malpighiaceae (editor), Das Pflanzenreich, Vol. IV, 141. Wilhelm Engel- from South America. Contr. Univ. Michigan Herb. 21: 29–36. mann, Leipzig. ———. 1999. Notes on Neotropical Malpighiaceae—VII. O’Donell, C. A. & A. Lourteig. 1943. Malpighiaceae Contr. Univ. Michigan Herb. 22: 1–19. argentinae. Lilloa 9: 221–316 pl. 1–18.