Eight Segregates from the Neotropical Genus Mascagnia (Malpighiaceae)
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Eight Segregates from the Neotropical Genus Mascagnia (Malpighiaceae) 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,