Martiodendron parviflorum (Amshoff) R.C. Koeppen, 1962

Falcão, Marcus José De Azevedo, Torke, Benjamin M., Garcia, Gabriel Santos, Silva, Guilherme Sousa Da & Mansano, Vidal De Freitas, 2023, A Taxonomic Revision of the Neotropical Genus Martiodendron (Fabaceae: Dialioideae), Phytotaxa 578 (1), pp. 11-56 : 43-52

Martiodendron parviflorum (Amshoff) R.C. Koeppen . Brittonia 14(2): 202 (1962).

≡ Martiusia parviflora Amshoff . Meded. Bot. Mus. Utrecht 52: 32 (1939).

Type:— Suriname: Sectie O. 30–35 m tall, 5-VI-1905, Boschbeheer 22 U00003287 (Lectotype designated by Koeppen & Iltis [1962: 202]: U!; Isolectotypes: WAG!). ( Figures 10–11 ) .

= Martiodendron elatum var. impressivenium W. Rodrigues . In sched.: Brazil: Amazonas : Estrada Manaus-Itacoatiara, km 108. Árvore de 15 m por 25 cm de diâmetro; flores amarelas; mata de terra firme e solo argiloso. 20-XI-1968, Coêlho, D. s.n. INPA25952 (INPA!).

Trees, (20–) 30–50 m tall, up to 70 cm in diameter, usually with large buttresses up to 4(–5) m tall; no exudate. Leaf rachis (4.5–) 7–16.5 cm long; petiole 1.1–4.1 cm long; petiolule 3–6 mm long; leaflets (4–)6–7(–8), the blades chartaceous, the terminal ones (7–)8–17 × 2.7–5.3(–8) cm, ca. 2–2.8(–3.5) times longer than wide, ovate to elliptical to oblong, slightly pubescent to glabrous, apex acuminate to cuspidate to acute, base obtuse to rounded or cordate; axillary buds ovate, (2–)2.7–4.3(–5) × 1.7–1.9(–2.5) mm, apex almost always acute, very rarely acuminate. Inflorescences thyrsoid, distichous, terminal, with an elongated primary and secondary axis from which cymes form, taller than wide, often more congested than other species, 9–25 × 5–25 cm. Flower buds 1–1.5(–1.8) cm long, strongly curved, usually falcate; sepals curved, 1–1.8 × 0.1–0.3 cm; petals (1–)1.5–2.1 × (0.3–) 0.5–1 cm; stamens 4 or 4+1 abaxial staminode; anthers 0.8–1.2 × 0.1–0.2 cm; anthers usually uneven, two larger laterals, two smaller adaxials, curved like the flower bud, densely pubescent from base to midportion; carpel 4–7 × 2 mm, fully pubescent, style 5–18 mm long. Fruits oblong to slightly asymmetrical, 12–17(–18) × 4–6.2 × 0.3–0.8 cm, ca. 2.3–3(–3.2) times longer than wide, red to vinaceous to purple, wings (0.7–) 1.2–2 cm wide in the middle portion, the dorsal and ventral wings equal or strongly unequal in width, seminiferous nucleus occupying the central part of the fruit, ca. 1.5–2 times wider than the widest of the two wings in the middle portion of the fruit.

Distribution, Habitat, and Ecology:— This species occurs in French Guiana, Suriname, and Brazil, in the states of Amapá, northern Pará, and northeastern Amazonas. It occurs almost always north of the Amazon River or, on the south bank, very close to it, in the region of Melgaço and Caxiuan ã in Pará ( Figure 15 ). It usually occurs in upland forests associated with large river basins, mainly the Amazon, Trombetas, and Jari Rivers, sometimes on the banks of the rivers, but never in floodable areas or campinaranas. It is usually found in clayey soils, more rarely in sandy-clay soils, and is the species of the genus most associated with very humid rainforests ( Figure 16 ), being quite large in height, with trees reaching 20– 50 m. These tall trees, less commonly found in M. elatum , are commonly found in M. parviflorum , with reports of 50 m trees in French Guiana (S.A. Mori 24247) and in the Brazilian states of Amazonas (J.G. Carvalho-Sobrinho 1581) and Pará (N.T. Silva 4628). This species has the largest buttresses in the genus, up to 5 m tall (J.G. Carvalho-Sobrinho 1581). Mori et al. (2002) indicate that the dispersal of the species would be anemochoric. The hypothesis proposed here agrees with the suggestion by Mori et al. (2002), that species associated with upland forests and with expanded fruit wings would have anemochoric dispersion, while M. excelsum , a species found in floodable areas and riverbanks and with reduced wings, it would have hydrochoric dispersion. Irwin (1966) mentions the frequent presence of ants in the inflorescences, an association also observed in the close related genus Dicorynia (Falc ã o et al., 2022).

Koeppen & Iltis (1962) mentioned the species only for the Guianas. Silva et al. (2005) cited for the first time the occurrence in Brazil in the states with occurrences confirmed here: Amazonas, Pará, and Amapá. However, the authors also mentioned the occurrence of the species in Rondônia, Maranh ã o, Piauí, Goiás, and Mato Grosso ( Silva et al., 2005), which is not correct. As the work referred solely to the Amazon basin, specimens from Maranh ã o, Piauí, Goiás, and Mato Grosso were not cited. Concerning the specimens from Rondônia, the authors cited only specimens identified here as M. elatum var. occidentale . Some of these specimens have even correct identification in labels as M. elatum made by the authors of the work ( Silva et al., 2005) (M.G. Silva 6127). Numerous wrongly identified specimens in several Brazilian herbaria are identified as M. parviflorum , emphasizing M. mediterraneum and M. elatum specimens.

Two specimens indicate a possible deciduousness for this species, in Suriname (J.P. Schulz 7970), and French Guiana (S.A. Mori 24247), losing leaves during flowering and fruiting, a phenomenon not mentioned in the other Martiodendron species. Several specimens of the other four species were observed in many different phenological phases and with no signal of caducifoly.Also, there are no registers of such character in any other herbarium specimen. Caducifoly is found in other Dialioideae taxa, like Apuleia (Falc ã o et al., in prep), Poeppigia densiflora (Falc ã o et al., 2021), a few species of Dialium ( Rojo, 1982) , and Koompassia ( Hou et al., 1996) .

Etymology:— The species was named by Amshoff from the Latin parvus (small) and flos (Flower), due to the smaller flowers compared to the other species of the genus. Although most specimens actually have the smallest flowers within the genus, this is a plastic feature. Some specimens have flowers of relatively common size for the genus, and some specimens of other species, mainly M. elatum have, in some cases, flowers also quite small.

Phenology:— It blooms from June to September, more rarely from May to November; fruits from August to November, sometimes losing the leaves before flowering/fruiting.

Uses:— The hardwood of this species is reported to be of local use for heavy construction in both Pará and Guianas.

Conservation:— An EOO of 632,824 km ² was estimated for M. parviflorum . The species is common in regions of Suriname (J.P. Schulz 7970), French Guiana ( Mori et al., 2002), and Amapá in Brazil (J.M. Irwin 4780), with mentions of the use of its timber being scarce. Although few specimens were collected after 2000, there are many specimens found in well preserved and/or protected areas such as Parc Amazonien de Guyane in French Guiana, Reserva Biológica Uatum ã, Estaç ã o Ecológica do Jari and Floresta Nacional Caxiuan ã in Brazil. Thus, based on IUCN criteria (2019), we suggest a preliminary least concern category for M. parviflorum .

Vernacular Names:— Muirá-Pixuna ( Brazil), Basmahonie, Boesimaoni, Bois d’amarante rouge, Tataboballi, Witte Pinto-Locus, and Witte-Purperhart ( Suriname and French Guiana). The name Tataboballi is also used in Suriname for the species Coutarea hexandra (Jacq.) K. Schum , a Rubiaceae .

Nomenclatural Comments: — Amshoff (1939) cited only one specimen in M. parviflorum ’s description, but did not discriminate which of the duplicates was to be considered the type, citing the herbaria U and P. However, the duplicate in P wasn’t cited by Koeppen & Iltis (1962) nor observed in the present work. Five extant duplicates of such collection were observed here: one in U and four in WAG. As Koeppen & Iltis (1962) cited only the specimen in U as type, we consider that the authors inadvertently accomplished a proper lectotypification (see Turland et al., 2018, Art. 7.11).

Taxonomic Comments:— Martiodendron parviflorum is a taller tree than M. excelsum , M. fluminense and M. mediterraneum , species that also occur in different habitats. It differs from all species in the genus by its shorter and ovate axillary buds with the apex acute, curved and commonly smaller flower buds and anthers, and the fruits with the seminal nucleus generally smaller compared to the wings. Together with M. excelsum , it differs from the other three species by having pubescent anthers, but the pubescence is more evenly distributed along the anther in M. parviflorum and generally more concentrated in the middle portion of the anther in M. excelsum . It has generally fewer leaflets than M. elatum , M. fluminense and M. mediterraneum , with those terminal leaflets having a smaller length × width ratio than the first species and being larger than the second and third species. Its leaflets are also generally narrower and with greater length × width ratio than in M. excelsum . It differs from M. fluminense by its long thyrsoid inflorescences and by having less stamens. It also differs from M. excelsum by its fully pubescent gynoecium. Its fruits are larger than those in M. excelsum and M. mediterraneum and have slightly smaller length × width ratio than those in M. elatum ( Table 1 ).

A specimen of INPA herbarium (Coêlho s.n. INPA25952), belonging to the species M. parviflorum , was misidentified as M. elatum , containing an indication by W. Rodrigues that would belong to a new variety, named in this label as M. elatum var. impressivenium W. Rodrigues. This binomial has never been validly described and the specimen does not present any morphological peculiarities. The curved flower buds are strongly characteristic of this species, bus in very rare cases, M. elatum can have similar curved flower buds (Capucho 477 IAN).

Selected List of Additional Specimens Examined: (21 of 51 analyzed specimens):— BRAZIL: Amapá: contagem entre porto Platon e serra do Navio. 15-XII-1976, Rosa , N. A. 1225 ( MG); rio Araguari , 1°40’N 51°56’W. Tree, 30 m high × 50 cm, 26-VIII-1961, Pires, J. M. 50498 ( IAN; MG; NY; US); rio Oiapoque: boca do rio Iaue , 2°53’N 52°22’W. Árvore, 30 m × 40-50 cm, 25-VIII-1960, Irwin, H. S. 47806 ( IAN; RB; MG; NY; US); rio Muturá, 0-10 km from its confluence with rio Oiapoque , 2°31-34’ N 52 °30-32’ W. 20-IX-1960, Irwin, H. S. 48386 ( IAN; MG; NY; US); Amazonas: Manaus: estrada Manaus-Itacoatiara, km 165. Árvore , 20 m × 35 cm, 6-VI-1973, Rodrigues, W. 9091 ( INPA; RB); estrada Manaus-Rio Branco. Árvore , 25 m × 40 cm, Coêlho, L. 1971 ( INPA); Presidente Figueiredo: Rebio Uatum ã, grade do PPBio. Árvore ca. 50 m alt. 11-VI-2007, Carvalho-Sobrinho, J. G. 1581 ( INPA; RB); Pará: Almeirim: estrada do Munguba, fazendinha. Árvore , 27 m, 33 cm DAP, 26-VI-1979, Silva, N. T. 5123 ( INPA; MG); Monte Dourado, estrada perimetral. Árvore , 40 m × 38 cm, 28-XI-1978, Santos, M. R. 429[42] ( INPA; MG; NY; RB); Jari, Monte Dourado. Árvore , 15 m × 20 cm, 15-VI-1969, Silva, N. T. 2191 ( INPA; IAN; NY; US); Melgaço: Flona Caxiuan ã, arredores da base física da estaç ã o científica Ferreira Pena, 1°44’15”S 51°27’19”W. Árvore, 20 m, 27-VIII-2014, Neves, D. M. 1960 ( NY; RB); Oriximiná, Porto Trombetas, área da mineraç ã o Rio do Norte , 1°27’S 56°22’W. Árvore jovem com 12 m de altura, DAP 14 cm, 14-XII-2007, Lima, H. C. 6809 ( RB); FRENCH GUIANA: Cayenne: near cachoeira Camaraua, about 3 km south of mouth of River Camopi , 3°10’N 52°19’W. 1-X-1960, Westra , L. Y. T. 48534 ( MG; IAN; K; NY; UB; US); Massif des Emerillons , 3°17’N 53°4’W. Arbre, 31-X-2007, Sabatier, D. 5352 ( US); Saint Laurent du Maroni: Riviere Grand Iaini, Bassin du Maroni. 12-VII-1990, Sabatier, D. 3237 ( NY); Saul , route de Bélizon, 3°37’N 53°12’W. Tree, 50 m, 28-IX-1995, Mori, S. A. 24247 ( MO; NY; P; US); Monts La Fumée , 3°37’N 53°12’W. Tree 13 m x 13.5 cm, 17-IX-1982, Boom, B. 1728 ( NY); SURINAME: Brokopondo: Bergendal , 7- XII-1921, B. W. 5531 ( US); Para: Zanderij. VII-1944, Stahel, G. s.n. RB750751 ( RB; U); Sipaliwini: Nicjerie River, Blche Marie Falls, forest 3 km SE of campo on west bank. Tree , 15-20 m × 30 cm, 19-VI-1965, Maas, P. J. M. 10887 ( F; NY; U; US); Mapane creek area. Tree , 32 m × 40 cm, V-1957, Schulz, J. P. 7970 ( NY; WAG).

List With Summary Data of Additional Specimens Examined: (30 of 51 analyzed specimens):— BRAZIL: Amapá: Irwin , H. S. 47705 ( IAN; MG; NY; UB; US); Amazonas: Rodrigues , W. 9115 ( INPA); Nascimento, J. R. M. s.n. NY 01161965 ( NY); Pereira, M. J. R. s.n. NY 01161966 ( NY); Assunção, P. A. C. L. s.n. NY 01161958 ( NY); Pará: Freitas , M. A. 906 ( MG); Silva , N. T. 2032 ( IAN; NY; US); 2825 ( IAN); 3289 ( IAN); 3312 ( IAN); 4628 ( MG; NY; US); Santos , M. R. 667 ( IAN; INPA; MG; NY); Carvalho , C. 167 ( MG); Silva, A. S. L. 2197 ( MG); 3047 ( MG); Oliveira, E. 4693 ( IAN); Soares , E. s.n. HSTM006066 ( HSTM); s.n. INPA171262 ( INPA); Faria, S. M. 643 ( RB); FRENCH GUIANA: Saint Laurent du Maroni: Boom , B. 2375 ( NY); Mori, S. 15596 ( NY); 15643 ( NY); 15658 ( NY); 15664 ( NY); 19195 ( NY; P); 22903 ( NY); SURINAME: Para: s.c. IAN38384 ( IAN); s.c. K000835081 ( K); Samuels, J. A. 243 ( L; NY; P; US); s.c. MO1575798 ( MO).

Figures

FIGURE 10. Martiodendron parviflorum (Amshoff) R.C. Koeppen. A. Branch with leaves and inflorescence; B. Detail of the petiolule and leaflet base; C. Detail of the axillary bud enveloped by the stipules; D. Part of an inflorescence subthyrsoid with flower buds in cymose subunits; E. Flower; F. Inner face of the sepal; G. Pubescent anther; H. Detail of the base of the stamen; I. Detail of the anther apex with the apical pore still closed; J. Detail of the receptacle with sepals, petals and four stamens removed, showing the abaxial staminode and pubescent carpel; K. Gynoecium; L Detail of the stigma; M. Fruit.A, D: Westra, L.Y. 48534 and Silva, N.T. 2191; B–C, F–L: Irwin, H.S. 47806; E, M: Mori, S.A. 24247. Drawn by M. Falcão. Scale bar. A, M: 2 cm; B–F: 3 mm; G–H, K: 1 mm; I–J, L: 0.5 mm.

FIGURE 11. Martiodendron parviflorum (Amshoff) R.C. Koeppen. A. Trunk of a specimen ca. 50 meters tall, with large buttresses; B. Inflorescence; C. Fruits; D. Branch with inflorescence and typical leaf with seven wide leaflets; E. Fruits; F. Detail of curved floral buds. Image authors: A–C. Carol Gracie and Scott Mori; D–F. Reflora: D: Silva, N.T. 3289; E: Mori, S. 24247; F: Silva, N.T. 2191. Scale bar. A: 50 cm; B, F: 1 cm; C–E: 5 cm.

FIGURE 15. Martiodendron distribution map. Topographic map of part of the Neotropics containing all analyzed specimens of the genus separated by species and varieties. Note the occurrence of most species in areas with elevations below 200 meters, with the exception of M. mediterraneum, which generally occurs in regions of elevation 200–600 meters. Source: NASA with modifications.

FIGURE 16. Martiodendron distribution related to precipitation. Average annual precipitation map in mm of part of the Neotropical region containing all analyzed specimens of the genus separated by species and varieties in the case of M. elatum. Note the distribution of M. parviflorum and M. elatum var. elatum in wetter areas (orange color), M. elatum var. occidentale and M. excelsum generally in intermediate areas (red color), and M. mediterraneum in drier areas (green color) to intermediate areas. The preferential environments of M. fluminense in relation to humidity, despite occurring in a region with lower average precipitation (red to green color), are restricted to small areas of coastal rainforests with higher humidity. Source: WorldClim with modifications.