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Taxonomic Revision of Amphiodon (Leguminosae, Papilionoideae, Brongniartieae)Author(s): Jose Eduardo Meireles and Ana Maria G. Azevedo TozziSource: Systematic Botany, 39(4):1150-1153. 2014.Published By: The American Society of Plant TaxonomistsURL: http://www.bioone.org/doi/full/10.1600/036364414X683967

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Systematic Botany (2014), 39(4): pp. 1150–1153© Copyright 2014 by the American Society of Plant TaxonomistsDOI 10.1600/036364414X683967Date of publication September 3, 2014

Taxonomic Revision of Amphiodon (Leguminosae, Papilionoideae, Brongniartieae)

Jose Eduardo Meireles1,3 and Ana Maria G. Azevedo Tozzi2

1Biology Department, Duke University, Durham, North Carolina 27708, U. S. A.2Departamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas, Caixa Postal 6109,

CEP: 13083-970, Campinas, Sao Paulo, Brazil.3Author for correspondence (jcm54@duke.edu)

Communicating Editor: Martin Wojciechowski

Abstract—The South-American genus Amphiodon Huber is reinstated and circumscribed to include a single species, A. effusus. The genus issegregated from Poecilanthe and is diagnosed by a vexillary stamen that is mostly adherent to the standard claw and a septate pod with seedsthat are transversally arranged. Poecilanthe ovalifolia is synonymized with A. effusus. Description, illustration, and geographical distributionfor A. effusus are provided.

Keywords—Amazonia, Fabaceae, Poecilanthe, South America.

A recent phylogeny of Poecilanthe Benth. s. l. (Leguminosae,Papilionoideae) revealed it to be paraphyletic, consisting ofthree well-supported independent clades (Meireles et al.2014). The subclade formed by the Amazonian species P. effusa(Huber) Ducke and P. ovalifolia Kleinhoonte, hereinafterreferred to as the Amphiodon clade, was shown to be sister toTabaroa L. P. Queiroz, G. P. Lewis & M. F. Wojc., and embed-ded in the tribe Brongniartieae (Cardoso et al. 2012; Meireleset al. 2014).Poecilanthe effusa was first described in the genus

Amphiodon (Huber 1909) as Amphiodon effusus Huber.Amphiodon was distinguished from Poecilanthe by havingdiadelphous stamens, bi-auriculated wing petals, andobovoid pods. Later, Ducke (1932) subsumed Amphiodoninto Poecilanthe, and proposed the new combinationP. effusa (Huber) Ducke. His decision was based on thefinding that the base of the vexillary filament was fused tothe staminal column (pseudo-monadelphous) and the wingpetals are only plicate at their lower base. Kleinhoonte(1925) had earlier described Poecilanthe ovalifolia fromSurinam, which Ducke (1932) observed to be very similarto P. effusa.Morphologically, the Amphiodon clade can be easily

distinguished from the remaining Brongniartieae by anumber of apomorphic characters, including the highlybranched panicles with a shortened primary axis, valvateaestivation of the calyx; the vexillary stamen mostly adher-ent to the standard claw, well-developed anther connec-tives, the internally septate pods with transversallyarranged seeds, and the colpate and suboblate pollen(Meireles and Tozzi 2007; Meireles and Tozzi 2008;Meireles et al. 2014; Souza et al. 2014). A key to the generaof the tribe Brongniartieae, including Tabaroa and theAmphiodon clade (as P.effusa + P.ovalifolia) is provided byQueiroz et al. (2010).The phylogenetic separation of the Amphiodon clade

from Poecilanthe in combination with the morphologicalcharacters cited above support the reinstatement of Amphiodonas a distinct genus. A morphological analysis also indicatesthat the diagnostic traits of Poecilanthe ovalifolia fall withinthe morphological variation of A. effusus. Thus, this workrevalidates the genus Amphiodon and synonymizes Poecilantheovalifolia with Amphiodon effusus. A description, illustration,and the geographical distribution for Amphiodon effususare provided.

Taxonomic Treatment

AMPHIODON Huber, Bol. Mus. Paraense Hist. Nat. 5: 398.1909. —TYPE: A. effusus Huber

Trees up to 20 m tall. Leaves alternate, pulvinate; stipulesminute, caducous. Leaflets 5–7 per leaf, alternate, sub-opposite or opposite; stipels persistent or caducous; second-ary venation brochidodromous. Inflorescences paniculate,axillary, cauliflorous, or pseudo-terminal, primary axis veryshort, secondary axes long, clustered or fasciculate. Flowersbilaterally symmetrical, pedicellate; bracteoles 2, minute,opposite, inserted at the base of the calyx. Calyx turbinate,sepals 5, basally fused and apically toothed, the upper 2 teethconnate for most of their length; hypanthium inconspicuous.Corolla papilionaceous, petals 5, glabrous, clawed; the wingpetals slightly longer than the keel, their upper base auricu-late; keel petals weakly adherent along part of their lowermargins. Androecium with 10 stamens, diadelphous, thevexillary stamen usually adherent to the base of the standardclaw or occasionally inserted on the hypanthium; stamenfilaments free apically, anthers sub-equal (the two types witha 2:1 length ratio), the shorter ones sub-dorsifixed (almostbasifixed), alternating with the longer basifixed anthers; con-nective conspicuous. Ovary sessile. Fruits dehiscent, inter-nally septate by a proliferation of papery tissue from theinner epidermis. Seeds with their broad faces transversallyorientated to the fruit length; post-chalazal branches reachingthe hilum; plumule villose, hypocotyl-root axis straight,shorter than the plumule.

AMPHIODON EFFUSUS Huber, Bol. Mus. Paraense Hist. Nat. 5:398. 1909.—TYPE: BRAZIL. Para: Faro, Serra do Dedal,in silvis, 3 Sep 1907, A. Ducke 8585 (holotype: MG!;isotype: BM). Poecilanthe effusa (Huber) Ducke, Bull.Mus. Hist. Nat. (Paris) ser. 2, 4: 733. 1932.

Poecilanthe ovalifolia Kleinhoonte, Recueil Trav. Bot. Neerl. 22:398. 1925.—TYPE: SURINAM. am Tapanahoniflusse,11 Nov 1918, J. W. Gonggrijp 4132 (holotype: U!; isotypes:IAN!, K!, NY!, US!), syn. nov.

Tree or treelet to 20 m; terminal branches slightly fissured;stipules caducous. Petiole (3–)5–9 cm long, rachis (3.5–)8.5–14 cm long, pulvinule 3–5 mm long, stipels caducous,rarely persistent. Leaflets elliptic to ovate, rarely suborbicularor narrowly elliptic, (5–)7–14(–19) + (2–)2.5–6(–8.5) cm;base obtuse to rounded, occasionally acute-cuneate; apex

1150

acuminate to distinctly cuspidate, membranaceous, chartaceousor rarely coriaceous, glabrous. Inflorescence 5–26 cm long,delicate (rarely robust), pubescent to sparsely tomentose,bract 0.6–0.8 mm long. Flower 7–10 mm long; pedicel 1–

3 mm long; calyx 4.7–6.3 mm long, sparsely to distinctlytomentose; corolla deep-red; standard petal obcordate towidely depressed-obovate, 7.7–9.5 mm long, base obtuseto acute, apex emarginate; wing petals elliptic-obovate,

Fig. 1. Amphiodon effusus. A. Flowering branch. B. Flower. C. Bracteole. D. Calyx opened out (inner face). E. Standard petal. F. Wing petal. G. Keelpetal. H. Stamens. I. Two anther types. J. Gynoecium, external and internal views. K. Closed fruit. L. Opened fruit showing the tranversal arangment ofthe seeds. M. Seed. Branch and flowers based on A. Ducke 16352 (MG). Fruit and seeds from D. C. Daly D867 (MG).

2014] MEIRELES AND TOZZI: REVISION OF AMPHIODON 1151

7–9.3 mm long, apex obtuse to rounded; keel petals elliptic-obovate to obovate, 6.6–7.8 mm long, apex obtuse torounded; stamens 5.7–7.3 mm long; basifixed anthers 0.8–1 mm long, sub-dorsifixed anthers 0.6–0.8 mm long;gynoecium ca. 7–7.4 mm long; ovules 8–9. Fruit obovoid,3.4–4.5 + 1.5–1.8 + 1–1.2 cm, apex rounded, base acute,slightly compressed in cross section. Seeds (1–)2–4(–5) perpod, widely elliptic, orbicular or slightly oblate, 9–13 + 7–12 + 3.5–6 mm; compressed or often plano-convex in cross-section; testa papery, dull, smooth. Figure 1.Distribution and Ecology—Recorded from Bolivia, Brazil,

French Guiana (Barneby and Heald 2002), Peru, and Surinam(Fig. 2). Widely distributed in Amazonian terra-firme (non-flooded) forest, especially in secondary forests and clearings.Phenology—Flowers recorded from March to November,

with most flowering specimens collected between June andOctober. Fruiting mainly from June to December, althoughtfruits were recorded as early as March.Vernacular Names—cumaru-de-rato, gema-de-ovo,

kloemansingi (for P. ovalifolia; Amshoff 1939).Representative Specimens Examined—BOLIVIA. Pando: 20 Jul 1978,

Meneces 761 (MG!); Nicolas Suarez, 3 Oct 1989, Beck et al. 19006 (MG!),28 Oct 1978, Meneces 803 (INPA!).

BRAZIL. Acre: Brasileia, 2 Jun 1980,Nelson 832 (INPA!, MG!, NY!, RB!);Rio Branco, Estrada Rio Branco/Brasileia km 42, 16 Oct 1980, Lowrie et al.559 (INPA!, MG!, RB!); Rio Macahuan, 22 Aug 1933, Krukoff 5605 (BM,NY!, SP!, U!, US!); Sena Madureira, 27 Sep 1980, Cid Ferreira & Nelson2553 (INPA!, MG!). Amapa: Contagem, Entre Porto Platon e Serra doNavio, 1976, Rosa 1322 (MG!); Mazagao, 21 Oct 1984, Daly et al. 3947(GH!, MG!, NY!); Rio Araguari, 29 Aug 1961, Pires et al. 50588 (IAN!,MG!, NY!, RB!, US!); Serra do Navio, 1961, Rodrigues 2922 (INPA!).Amazonas: Maues, Rio Urupadi, Igarape Quininha, 19 Jul 1983, Zarucchiet al. 3104 (INPA!, MG!); Novo Aripuana, Transamazonica a 400 km deHumaita, 3 May 1985, Cid Ferreira 6002 (INPA!); Parintins, Lago Uaicurapa,5 Sep 1932, Ducke s. n. (RB!). Maranhao: Buritirama, Rio Itacaiunas, 21 Jun1970, Pires 12201 (IAN!); Carutapera, Gurupiuna, 3 Nov 1986, Balee &Ribeiro 2786 (NY!); Moncao, Bacia do Rio Turiacu, 5 Jun 1985, Balee 971(MG!, NY!); Santa Ines, Rodovia p/ Acailandia, km 100, 6 Aug 1976,Pinheiro 25 (IAN!); Santa Luzia, Fazenda CVB, perto do km 130 da BR222, 24 Oct 1980,Daly et al. D752 (INPA!, MG!, NY!). Para: Acara, FazendaBorba Gato, 7 Nov 1980, Daly et al. D867 (F!, GH!, HRB!, IAN!, INPA!,MG!, NY!, US!); Altamira, Igarape Trindae, 3 Aug 1971, Cavalcante &M. Silva 2856 (MG!, RB!); Arumanduba, Serra da Arumanduba, 23 Jul1961, Egler & Irwin 45930 (IAN!, MG!,NY!, RB!, S, U!, UB!); Barcarena, VilaItupanema, Reserva ALBRAS, 17 Sep 1983, Cordeiro 1800 (IAN!); Belterra,5 Aug 1947, Black 47–1152 (IAN!); Braganca, Colonia 3 de outubro, 23 Aug1952, Pires 4125 (IAN!, NY!); Cachoeira, BR 22, Capanema - Maranhao,km 96, 27 Oct 1965, Prance 1716 (IAN!, NY!); Carajas, Serra dos Carajas,21 May 1982, Sperling et al. 5777 (NY!); Faro; Fordlandia, Regiao do RioTapajos, Sep 1931, Krukoff 1016 (BM!, NY!, S, U!); Ipixuna do Para, Km 129,

alem de Ipixuma, 6 Jul 1966, Duarte 9811 (RB!, RFA!, UB!); Itaituba, 3 Sep1902, Ducke 2972 (MG!); Itinga do Para, Fazenda Santa Rosa, 26 Oct 1980,Daly et al. D781 (INPA!, MG!); Jacunda, Remansao, 15 Apr 1981, Silva et al.1575 (INPA!, MG!); Lago Moiossu, 24 Jul 1959, Egler 1009 (IAN!, INPA!,MG!, NY!, UB!); Maraba, Carajas, Serra Norte, 6 Aug 1982, Maciel 766(INPA!, MBM!, MG!, NY!); Melgaco, 13 Oct 1991, Silva & Silva 2371(MG!, NY!), Flona de Caxiuana, 15 Feb 1991, Silva et al. 2221 (MG!); Moju,Campo experimental da Embrapa, km 30 da rodovia PA-150, 11 Mar1996, Ribeiro 2110 (IAN!); Monte Dourado, Rio Jarı, 5 Oct 1968, Silva 1117(IAN!, NY!); Oriximina, BR 163 a 6 km da Cachoeira Porteira, 20 Aug1986, Cid Ferreira et al. 7894 (F!, INPA!, MG!, NY!), Porto Trombetas, 3 Jul1995, Faria&Delmo Fonseca 1009 (HUEFS!, rb!); Rio Cumina-mirim, 13 Oct1913, Ducke 14977 (MG!, RB!); Rio Cachorro, Igarape Cabeca de Onca,25 Aug 1986, Cid Ferreira et al. 8020 (F!, inpa!, MG!, NY!); Rio Paru doOeste, 10 Sep 1980, Cid Ferreira et al. 2360 (INPA!, MG!, NY!, RB!); Ourem,Rio Guama, Jul 1953, Pires & Silva 4603 (IAN!, INPA!, NY!); Paragominas,Fazenda da FFT, area da Cikel, 29 May 2002, Ferreira 812 (IAN!); Peixe-Boi, 22 Jul 1907, Siqueira s. n. (MG!), E. de F. de Braganca, 23 Apr 1907,Siqueira 5575 (BM!, INPA!, K, MG!, RB!); Ponta de Pedras, Rio Itacaiuna,14 Jun 1949, Froes& Black 24484 (IAN!, SP!); Ponte Nova, Entre Ponte Novae Boa Vista, 19 Aug 1919, Ducke s.n. (RB!); Rio Capim, 17 Jun 1897, Huber733 (BM!, MG!, RB!, US!); Rio Jarı, 20 Jun 1969, Silva 2270 (IAN!, NY!); RioTapajos, 11 Sep 1916, Ducke 16480 (BM!, MG!); Rodovia Belem-Brasılia,km 94, 8 Sep 1959, Kuhlmann & Jimbo 208 (IAN!, MG!, NY!, SP!, SPF!, EUC!),Ipanema, 18 Aug 1916, Ducke 16352 (BM!, MG!, RB!); Muji dos Campos,13 Aug 1969, Silva & Sousa 2243 (MG!, RB!, S!), Muji dos Campos, 19 Aug1969, Silva & Sousa 2314 (mg!, NY!, RB!, S, US!); Sao Domingos do Capim,Rio Capim, 3 Jul 1974, Cavalcante 2973 (INPA!, MG!, NY!, RB!); Tome-Acu,Rio Acara, 29 Oct 1979, Silva & Pinheiro 5115 (MAC!, MG!, NY!); Tucuruı,3 Oct 1983, Revilla et al. 8437 (INPA!, NY!), 2 Jun 1980, Silva & Rosario 5319(INPA!, MG!, NY!, RB!, SPF!), Igarape Cagancho, 21 Aug 1980, Rodrigueset al. 10261 (INPA!, MG!, NY!, RB!); Ulianopolis, Fundacao Floresta Tropical,03–07 Jun 2000, Lobato 2566 (MG!). Rondonia: Abuna, Serra proximoa Namorado Novo, 5 Aug 1971, Prance et al. 14716 (INPA!, MG!, NY!);Alvorada d’Oeste, p/ Costa Marques, km 90, 25 Jun 1983, Silva 6464(INPA!, MG!); Ji-Parana, 4 May 1987, Cid Ferreira 9030 (INPA!, MBM!,NY!), estrada Cuiaba/Porto Velho, km 353, 26 Jun 1984, Cid Ferreira et al.4811 (F!, INPA!, MG!, NY!, RB!, UEC!), Porto Velho, reservatorio da UsinaHidreletrica de Samuel, 20 Jun 1986, Cid Ferreira et al. 7528 (F!, GH!,INPA!, NY!); Santa Barbara, 28 May 1982, Teixeira et al. 840 (F!, INPA!,MG!, NY!, RB!). Roraima: Boa Vista, Aug 1932, Capucho 388 (IAN!).

FRENCH GUIANA. Cayenne: 26 Jul 1990, Sabatier & Prevost 3423(NY!, U!), 26 Oct 1968, Oldeman 254 (NY); Reserve de Petit-Saut, Oct1997, Paget 2 (NY!). Saint Leurent duMaroni: Saul, Circuit de la montagneGgrand Fosses, km 0.485, 1971, Oldeman 4053 (NY!, U!).

PERU. Madre de Dios: Manu Province, Puerto Maldonado, Los AmigosBiological Station, 21 Aug 2002, Janovec & Maceda 2727 (BRIT, K!, MOL,USM); 28 Sep 2003, Maceda & Cornejo 953 (K!)

SURINAM. 26 May 1957, Schulz 7989 (AAU!, U!); Brownsberg NaturePark, 90 km S of Paramaribo, 24 Sep 1976, Mori & Bolten 8386 (NY!).

Comments—Amphiodon effusus is the only species in thegenus. As with other widespread species, A. effusus possessesconsiderable variation in vegetative morphology. Althoughmembranaceous to chartaceous leaflets and leaves lackingstipels (very early cacucous) are the most common morphol-ogies, robust leaves and stipels are occasionally encountered.

Poecilanthe ovalifolia has been differentiated from A. effususprecisely for its leaves bearing coriaceous leaflets and persistentstipels, robust inflorescence rachises and for its blue corolla.

As stated above, coriaceous leaflets and persistent stipelsare not diagnostic for P. ovalifolia. The putatively blue corollaof P. ovalifolia is also of limited taxonomic value and the colourrecord might well be incorrect. Information concerning petalcolor is provided only in the protologue and not on the spec-imen field label. These observations suggest that A. effususand P. ovalifolia are the same species and justifies the synon-ymy of Poecilanthe ovalifolia.

Poecilanthe ovalifolia is represented only by its type, collectedalong the Tapanahoni river in Surinam. Even though the spec-imens listed above as isotypes have B.W. instead of Gonggrijpas the collector, they most certaintly belong to the same collec-tion. The abbreviature B.W. stands for Boschwezen Suriname,

Fig. 2. Geographical distribution of A. effusus in northernSouth America.

1152 SYSTEMATIC BOTANY [Volume 39

the Forestry Bureau of Suriname, and many duplicatesof their material were sent out without crediting the individ-ual collector.

Acknowledgments. We are grateful to Matt Lavin, Gwilym Lewis,and Haroldo Cavalcante de Lima for their critical comments on earlierdrafts of this paper and to Samira Rolim for the illustration. The curatorsof the visited herbaria are also gratefully acknowledged. Financial sup-port was received from CNPq and FAPESP (proc. 06/50154-0).

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