Dicorynia paraensis Benth. var. uaupensis R.C. Koeppen. Brittonia

Falcão, Marcus José De Azevedo, Torke, Benjamin Marland & Mansano, Vidal De Freitas, 2022, A Taxonomic Revision of the Amazonian Genus Dicorynia (Fabaceae: Dialioideae), Phytotaxa 554 (1), pp. 1-31 : 19-24

Dicorynia paraensis Benth. var. uaupensis R.C. Koeppen. Brittonia 19(1): 55 (1967).

≡ Dicorynia uaupensis Spruce, Spruce, R. Pl. N ° 2772, nom. ined. ≡ Dicorynia uaupensis Spruce ex Benth. in Mart., Fl. Bras. 15(2): 81 (1870), nom. nud. TYPE: — BRAZIL. Amazonas: “Prope Panuré, ad Rio Uaupés, in sylvis ripariis. Arbor 60 pedalis, flores albi”, XI-1852, Spruce, R. 2772 K000555123. (Holotype: K!; Isotypes: K!, BR, F!, G, GH!, NY!, P!, RB!, TCD!, W!).

= Dicorynia paraensis Benth. var. breviflora (Benth.) R.C. Koeppen. Brittonia 19(1): 59 (1967). ≡ Dicorynia breviflora Benth. in Mart., Fl. Bras. 15(2): 82 (1870). TYPE: — BRAZIL. Amazonas: “Barra (= Manaus), Igapó do Rio Negro. Large tree, 60 ft × 4–6 ft with thick roughish bark”, II-1851, Spruce, R. 1306 (Lectotype, Step I. Koeppen 1967 Step II. designated here: K barcode 000264605!; Isolectotypes: F!, G!, GH!, K!, M, NY!, P!, RB!, TCD!, US!)..

= Dicorynia paraensis Benth. var. floribunda R.C. Koeppen. Brittonia 19(1): 57 (1967). ≡ Dicorynia floribunda Spruce, Spruce Pl. N ° 2135, nom. ined. ≡ Dicorynia floribunda Spruce ex Benth. in Mart., Fl. Bras. 15(2): 82(1870). nom. nud. Type: — Brazil: Amazonas: “Falls of São Gabriel, Igapó. Spreading tree, 30 × 1ft ”, II-IV-1852, Spruce, R. 2135. (Lectotype, Step I. Koeppen 1967 Step II. designated here: K barcode 000264598!; Isolectotypes: F!, FI! G, GH!, K!, M, NY!, P!, RB!, TCD!, W!)..

= Dicorynia paraensis Benth. forma parviflora Ducke. nom. ined. In sched, Ducke, A. s.n. RB20337 (RB).

Trees up to 28 (–40) m tall, very rarely shrubs. Leaves (10–) 14–31 cm long, leaflets 7 (9), very rarely 10 (some leaves of a single specimen), terminal leaflets narrowly elliptical to narrowly oblong to narrowly ovate, usually obtuse to truncate at base, very rarely cordate or cuneate, the abaxial face sparsely pubescent to glabrous, papillate or not, (5.5–) 8–12 (–16.7) × (2–) 3.5–5 (–7.6) cm, the length 2–3 (–4) times the width; petiolules (2–) 4–7 (–9) mm long; axillary buds non deciduous, generally laterally narrowly obovate, less commonly oblong or orbicular, acuminate to obtuse at apex, (2–) 3–5 (–7) × 1–3 (–4) mm, commonly apart from the point of leaf attachment, 1–4mm above it. Inflorescences ca. (9–) 13–28 × (4.5–) 6–12 (–20) cm, sericeous, golden to brown, secondary branches perpendicular to the central axis or ascending, rarely tertiary branches; indumentum of the external face of the sepals and petals white to golden to brown. Fruits 4.3–7 × 3–4 × 0.5 cm. (Figs. 3B–D; 4A-B, H-N; 6B, G; 7C).

Distribution, Habitat, and Ecology: — D. paraensis var. uaupensis occurs in southern Venezuela, eastern Colombia, and in the Brazilian states of Amazonas and, rarely, Amapá. The variety occurs mainly in the basins of the Negro (Guainia), Japurá (Caquetá), and Uatumã Rivers. Its distribution partially overlaps with varieties macrophylla and paraensis (Fig. 5). It is the only taxon within Dicorynia found so far in the Brazilian state of Amapá, although with great rarity, being observed in the present work as a new area of occurrence for the genus. Only one specimen was recorded in the region (H. Medeiros 3114 RB). Although such specimen is sterile, its leaves and axillary buds strongly resemble this variety. This specimen occurs considerably distant from the closest populations of var uaupensis in the Uatumã river basin, in the state of Amazonas, thus increasing its area of occurrence. As the region between these two populations is little sampled in botanical collections, we think it is likely that the variety occurs at least in the southern part of this apparent disjunction, including in the state of Pará, through the Amazon River basin. It is important to emphasize that the northernmost part of this disjunction corresponds to areas of higher elevation and less precipitation than those where records of the genus exist (Figs 5, 10). Therefore, we find it less likely that the species occurs in such regions. There is a single record of the variety from outside of the native distribution, a specimen collected on a farm in Rio de Janeiro in 1872 (Glaziou 1583). Whether the locality data are incorrect or the record represents a cultivated specimen is unclear. Cases of specimens of Amazonian species numbered by Glaziou and erroneously indicated as occurring in Rio de Janeiro are recurrent in herbaria. The former farm is now a highly urbanized part of the metropolitan area of Rio de Janeiro.

Variety uaupensis exhibits the broadest environmental tolerance, with some populations almost always associated with well-drained terra firme forest, such as in the upper Negro River basin, and others with seasonally inundated “várzea” or “igapo” forests, such as those along the Japurá River and the middle and lower Negro River. It is very rare, however, that the variety occupies multiple forest types in the same region, and where it co-occurs with var. macrophylla , it is essentially restricted to seasonally inundated forests, while the latter occurs almost always, in “terra firme” forests. In Negro River, where it co-occurs with D. paraensis var. paraensis , they share similar environments of “igapó” forests.

Etymology: —The name uaupensis is a reference to the Uaupés River, one of the main tributaries of the upper Negro River and along which Richard Spruce collected the type in 1852.

Phenology: —Flowering from September to February, more rarely from August to March; fruiting from February to August.

Conservation: — Dicorynia paraensis var. uaupensis has an estimated EOO of 1,025,074 km and is accessed as Least Concern, with the same reservations mentioned for the status of the whole species.

Nomenclatural Comments: —It was necessary to select second steps lectotypes for D. paraensis var. floribunda R.C. Koeppen and D. paraensis var. breviflora (Benth.) R.C. Koeppen. In both cases, Koeppen (1967) specified that the “type” was housed at K, but the two collections are represented in that herbarium by three duplicates each. The collections are, respectively, R. Spruce 2135, which was not seen by Koeppen and may represent three different collections from the same tree ( Koeppen 1967), and R. Spruce 1306, which was indicated by Bentham in the description of the basionym (1870) and later cited by Koeppen (1967). The duplicates selected as the lectotypes are two that bear extensive notes written by the collector Richard Spruce (K000264598; K000264605) (Article 9.6 of the Code: Turland et al. 2018).

As the varieties uaupensis , breviflora , and floribunda were published simultaneously by Koeppen (1967), they have equal priority under the Code ( Turland et al. 2018, article 11.5). Here we choose the name D. paraensis var. uaupensis for the recognized taxon, as the morphology of the type from the Uaupés River region best exemplifies the taxon. As var. floribunda appears to be an aberrant morphological variation and var. breviflora has this name due to Bentham’s observations of flower characters not supported here.

It should be noted that Koeppen (1967) considered D. paraensis var. floribunda and D. paraensis var. uaupensis to be new combinations based on the names that appeared as synonyms of D. paraensis in Flora Brasiliensis ( Bentham, 1870), citing their authorship as “(Spruce ex Benth.) R.C. Koeppen”, which is broadly used to date. However, both names coined by Richard Spruce, were not validly published since Bentham included them in the comments concerning D. paraensis , citing Spruce’s specimens that contained such names (see Turland et al. 2018, article 36.1). Thus, Koeppen inadvertently described two new varieties, and the correct authorship for both taxa are R.C. Koeppen, without any parenthetical authorship. A note regarding this problem was made by Kanchi Gandhi and is attached to the isotypes deposited in the NY herbarium.

Taxonomic Comments: —It can be distinguished from the typical variety by the generally greater number of leaflets (although there is a small overlap in this character), narrower leaflets, smaller petiolule length, and shape of the axillary buds. Differs from D. paraensis var. ingens by the generally smaller leaf size, smaller number of leaflets, narrower leaflets, base of the leaflets generally obtuse, the larger petiolule length, and shape of the axillary buds; and from D. paraensis var. macrophylla by the generally smaller leaf size, the smaller number of leaflets, the shorter leaflets, the smaller petiolules length, shape of the axillary buds, and habitat preference where the two varieties cooccur (Table 1).

The types of varieties uaupensis and breviflora are practically identical in number, size and shape of leaflets, inflorescences, flowers, etc. For Koeppen (1967) they are distinct by the petals abaxial face with whitish indumentum in variety uaupensis (vs. golden in variety breviflora ) and the leaflets abaxial face with non-papillate epidermis in var. uaupensis (versus papillate in var. breviflora ). Although whitish petals and, sometimes also sepals, are indeed characteristic of some populations of var. uaupensis , mainly in Japurá and Uatumã Rivers, such characters are not exclusive to this variety, they are rarely seen in specimens of other varieties with variable vegetative morphologies (A. Ducke s.n. RB24185). Besides that, individuals from the aforementioned populations of var. uaupensis and from other populations, such as those in Venezuela and the lower Negro River, have petals’ indumentum with intermediate colors from white to golden/brown (C.A. Cid Ferreira 3564; 3753), similar to those found in specimens cited as varieties breviflora and floribunda . Finally, different patterns in leaflet surface, including presence/absence of papillae could not be related to other morphological or geographic patterns through these populations.

In addition to these characters, leaflet venation in the type of variety breviflora is much more evident than in the type of var. uaupensis . This distinction also does not hold in comparisons of broader sets of specimens. Other purported differences in the flowers cited by Bentham (1870), as concluded by Koeppen (1967), are of little taxonomic value and were probably based on observation of underdeveloped flower buds. Finally, it should be noted that the geographical distributions of these two varieties as circumscribed by Koeppen completely overlap, further substantiating our decision to treat them as synonyms.

As for D. paraensis var. floribunda , the type is remarkable in its relatively small, revolute-margined leaflets. Koeppen (1967) also cited the following characters to differentiate it: pustules on the leaflets, the leaflet abaxial face papillate and glandular, relatively small flower buds, and the tip of the ovary tapering. However, most of these characters are exceptionally variable in D. paraensis , and even the two main features that characterize the type, small and revolute-margined leaflets, vary considerably, even among duplicates of the type, with one of them (R. Spruce 2135 K000264600) with leaflets of shape and size indistinguishable from those of several individuals of var. uaupensis . Tiny leaves are also found in individuals of other varieties at the bases or even in the middle of the inflorescences, being in reality leafy developed bracts. Besides that, several specimens of var. macrophylla , ingens and uaupensis present revolute margins (C.A. Cid Ferreira 6787). This character is more common in branches with smaller or less developed leaves, while this varies even among the duplicates of var. floribunda’ type. Concerning the presence of pustules in the leaflets, a character that, according to Koeppen, had not been observed in any other taxa than var. floribunda . We could observe that this is not such an unusual feature, being found in specimens of various vegetative morphological types (B. Krukoff 8635; A. Ducke 1011, among others).

Nevertheless, Ducke (1948) indicated that the former taxon, which he treated as a full species, D. floribunda , was also distinct in being a very large tree with a wide and flat crown. However, the subjectivity of this description and almost total absence of information on shape and size of tree crowns for most of the collected specimens of D. paraensis , along with the fact that Ducke did not describe the character for varieties uaupensis and breviflora , make the feature questionable for the delimitation of these taxa. Koeppen (1967) defined D. paraensis var. floribunda as scarse, based on the existence of only two specimens and on Spruce’s field notes, which collected a single specimen of this “ Dicorynia of small leaves” even after intense search in the region. After Koeppen’s work, only one other similar specimen was collected in the region, with leaflets of intermediate size between those of D. paraensis var. floribunda and D. paraensis var. uaupensis (M.G.M. Roosmalen 1377) . Thus, D. paraensis var. floribunda does not seem to be clearly defined by any character, and is thus synonymized here to D. paraensis var. uaupensis .

Koeppen (1967) indicated that var. floribunda and var. breviflora differ by the shape of the upper portion of the ovary, being tapered in the first. Even in the types, this characteristic is weakly distinguishable and is just one more minimal variation among several others found in Dicorynia gynoecium.

Additional Specimens Examined:— Brazil.— AMAPÁ: Pedra branca do Amapari, Parque Nacional das Montanhas do Tumucumaque, 1°14’14”N 52°25’30”W, Floresta ombrófila densa, 24-XI-2017, Medeiros, H. 3114 (RB);AMAZONAS: Novo Airão, praia de areia branca, abaixo do rio Negro, 2°38’5”S 60°55’35”W, floresta de igapó, 25-II-2000, Oliveira, A.A. 3557 (UNIP); braço de rio ao lado de Novo Airão, 2°37’1”S 60°57’52”W, 24-II-2000, Oliveira, A.A. 3546 (UNIP); rio Içana, 25-V-1975, Rosa , N.A. 386 (IAN); margem do rio Içana, adjacência da serra de Tunuhy, 14-XI-1945, Fróes, R.L. 21403 (IAN; NY; US); Villa Bittencourt, rio Japurá, margem esquerda, mata de várzea, 22-XI-1982, Amaral, I.L. 618 (INPA; MG; NY; RB); Novo Japurá, ao longo do rio, 1°54’S 67°1’W, 10-XI-1982, Cid, C.A. 3564 (INPA; MG; NY; R; RB; US); Villa Bittencourt, rio Japurá, igarapé Patoá, igapó, 19-XI-1982, Amaral, I.L. 574 (INPA; MG; NY; RB); Novo Japurá, margem direita, lago do Mapari, 10-XI-1982, Amaral, I.L. 401 (INPA; NY; RB; US); fronteira de Brasil e Colômbia, Novo Japurá, Villa Bittencourt, rio Apapóris, margem esquerda, igarapé preguiça, 1°14’S 69°25’W, 21-XI-1982, Cid, C.A. 3753 (INPA; MG; NY; RB; US); margem do rio Negro, próximo de Manaus, praia do Cajo, solo arenoso, 30-XI-1977, Coelho, L.F. 656 (INPA); Manaus, estrada do Aleixo, porto Mauá, 23-IV-1970, Rodrigues, W. 8819 (INPA); Manaus, Tarumanzinho, rancho 6 irmãos, 23-XI-2000, Mansano, V.F. 117 (UEC); Manaus, igarapé da Cachoeira Grande, silva riparia non inundabili, 25-V-1941, Ducke, A. 727 (IAN; MG; NY; R); 2-II-1930, Ducke, A. s.n. RB20337 (RB; US); Manaus, 20-III-1932, Ducke, A. s.n. RB24184 (RB; US); Manaus, 25-X-1929, Ducke, A. s.n. RB23319 (RB; US); Manaus, rio Negro, entrada do igarapé Tarumã-Açú, praia da Lua, 3°2’23”S 60°7’18”W, 16-VIII-2018, Falcão, M.J. 91 (RB); margem do igarapé do Tronco-Manaus, capoeira fechada, 20-II-1956, Chagas, J. 3471 (IAN; INPA; US); Manaus, Condomínio T. Loureiro, 3°2’39”S 60°6’19”W, 25-XI-1999, Oliveira, A.A. 3504 (UNIP); UHE Balbina, rio Uatumã, Base II, 22-IX-1988, Webber, A. 1250 (INPA); Presidente Figueiredo, canteiro de obras da UHE Balbina, 1°30’S 59°30’W, 11-IX-1986, Cid Ferreira, C.A. 8065 (MO; NY; US); Santa Isabel do rio Negro, 0°24’S 65°0’W, 16-VIII-1999, Roosmalen, M.G.M. 1377 (INPA); Santa Isabel, rio Negro, silva rarius inundabili, 8-III-1936, Ducke, A. s.n. RB35075 (INPA; NY; RB; US); rio Negro, boca do Miry, 1947, Fróes, R.L. 22867 (IAN; US); “Para” (probably actual Amazonas, rio Negro), prior 1785, probably L.C. Richard s.n. P02743988 (P); Colombia.— AMAZONAS: Resguardo Aduche, Correg. De Pto, Santander, 72°19’W 0°39’S, 1-XII-1993, Cárdenas, D. 4289 (COAH; MG); rio Caquetá, margen derecho, frente a Villa Azul, terraza baja, 6-IX-1989, van Andel, T. 233 (COAH; NY; U); 12-IX-1989, van Andel, T. 271 (COAH; NY; U); Pena Roja, varzea, 13-X-1993, Dulmen, A.V. 168 (COAH; L); 23-IX-1993, Dulmen, A.V. 160 (COAH; U); aproximadamente 540 m en dirección 20° de la margen izquierda del Río Caquetá en frente de la punta sur de la isla Yarumal, 1°7’51”S 71°32’33”W, 21-V-1997, Sánchez, M. 3261 (COAH); CAQUETÁ: Araracuara, El Engaño, 25-XI-1991, Restrepo, D. 599 (MO; NY); 8-XI-1991, Restrepo, D. 404 (NY); VAUPES: Estación Biológica Caparú, within 3 km of the norh bank of lago Taraira, 1°0’S 69°49’W, 8-III-1990, Defler, S. 727 (COAH; MO; US); Venezuela.— AMAZONAS: Atabapo, IV-1990, Yanez, M. 481 (MO); Atabapo, Piedra Sapo, rio Atacavi, 3°5’N 67°2’W, XI-1989, Velazco, J. 1005 (MO; NY; US); bosque de tierra firme de ladera, ca. 1.5 km al este de la boca del caño frente a Laja Viento, por el rio Guainia, 2°4’59”N 67°5’21”W, 1-IV-2000, Berry, P.E. 7460 (MO; NY); rio Guainia, riverine forest just south of Maroa, 28-XI-1953, Maguire, B. 36460 (F; NY; RB; U; US).