Dicorynia guianensis Amshoff

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 : 11-13

Dicorynia guianensis Amshoff . Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 28–31 (1939).

TYPE: — SURINAME. Zanderij , 10-XII-1914, BW 452 U0003269 (Holotype: U!; Isotype: NY!) .

Medium to large trees, (9–) 22–45 (–50) m tall, trunk 14–100 cm in diameter, buttresses up to 1 m tall. Leaves (13–) 17–33 (–60) cm long, leaflets usually 7, very rarely 3, 5, 9, or 11, usually oblong, very rarely elliptical or ovate, acuminate to cuspidate at apex, base usually obtuse or truncate, very rarely cuneate or cordate, the abaxial face sparsely pubescent and papillate, rarely non-papillate, terminal leaflets (6–) 7–12 (–25) × (3–) 4–7 (–11) cm, the length (1.5–) 2–2.5 (–3.8) times the width; petiolules (2–) 4–6 (–8) mm long; axillary buds laterally obovate, oblong or orbicular, obtuse at apex, 2–4 × 2–3 mm, the distance between the bud and the leaf attachment point 0–1mm; terminal buds strongly capitate, orbicular at apex, 4–7 × 5–8 mm. Inflorescences (10–) 19–35 (–40) × (6.5–) 10–27 cm, sericeous, golden to brown; bracts 9–12 × 4–7 mm; pedicels 4–8 mm long. Flower buds 8–11 × 5–8 mm; sepals 8–11 × 5–8 mm; petals (7–) 10–16 × (6–) 10–13 mm, claws 2–4 mm long; filament of the longer stamen 4–7 × 1–3 mm, filament of the shorter stamen 2–3 × 1–3 mm, anther of the longer stamen with (6–) 8 sporangia, anther of the shorter stamen with 8–11 sporangia in the median to apical region, both 3–6 × 2–3 mm; carpel 4–5 × 2 mm, style 4–6 mm long, ovules 4–6. Fruit (4.4–) 6–8.2 × 2.2–4 × 0.4–0.5 cm, wing 3–8 mm wide; seeds 1.5–2 × 1.2–1.7 × 0.2–0.5 cm. (Figs. 1-2; 6A, C; 9E–G, I–J, L–M, P–Q).

Distribution, Habitat, Biogeographical Patterns, and Ecology: —The species occurs in French Guiana, Suriname, and rarely in Guyana. Some specimens from French Guiana are located in areas so close to the border with the Brazilian states of Amapá and Pará, that its occurrence in such areas does not seem unlikely, especially because this region is little botanically sampled. There are even reports of the species in the region of Massif of Mitaraka, basically on the border with Amapá ( Sabatier 2019) (Fig. 5).

Few records of the species were cited for Guyana, and all but one did not belong to Dicorynia , consisting of a sterile Papilionoideae ( T. van Andel 2124 U0088103) and several specimens of Martiodendron excelsum (Benth.) Gleason , another Dialioideae similar to Dicorynia in its leaves and fruits (K.M. Redden 5080 US; S. Mori 24350 U; B. Hoffman 1366 U; D. Clarke 1358 U), including all three specimens cited in the checklist of the Iwokrama Forest in central Guyana ( Clarke et al. 2001). They have very different flowers, and their fruits can be differentiated by Martiodendron having two expanded wings with a central and monospermic seminal nucleus, delimited by characteristic venation, and an elliptical seed with a gelatinous endosperm. The fruits of Dicorynia have one narrow wing, without a differentiated seminal nucleus, generally with one to three reniform to orbicular seeds, with gelatinous endosperm only in its margin. In addition, D. guianensis almost always has seven leaflets, while M. excelsum almost always has five leaflets (Falcão et al. unpubl. data, present work). There are also specimens of Dicorynia mistakenly indicated as coming from Guyana, due to the old collection labels tagged as “ Guyana ”. However, the correct location could be identified by location names as “route de Cayenne” or “Laurent du Maroni”, from French Guiana. In the checklist of the Mabura forest ( Ek 2003) and the useful tree guide in Guyana ( Polak 1992), the genus, known for its timber, is not cited. The only known Guyana specimen occurs considerably distant from the other individuals of D. guianensis in Suriname (Fig. 5). If this is due to lack of sampling or if the genus is very rare in Guyana is not known.

Dicorynia guianensis is usually found below 200m elevation on clayey or sandy soils of “terra firme” ombrophilous forests, less commonly in seasonally inundated areas (Fig. 5). At the same time, some varieties of D. paraensis are associated with seasonally flooded forests. D. guianensis is commonly an emergent tree, with many specimens reaching 40-45 meters tall; populations of this species appear to have relatively higher frequencies of tall trees than D. paraensis . D. guianensis is considered very common or dominant in parts of French Guiana and Suriname (Record and Hess 1943, Maas 1971, Mori et al. 2002, Jesel 2005). Its abundance permitted the industrial scale extraction of its timber until recent years. The flowers of D. guianensis might be large-bees pollinated ( Jesel, 2005). Some collectors like S. Mori (specimen 24340) pointed out that the flowers are very odoriferous with a pleasant scent and that the flowers are diurnal, opening at 5:30 a.m., generally with many trees of a region flowering at the same time, making the species quite conspicuous in the forest canopy. The fruits are widely consumed by Psittacidae birds such as those of the genera Pionus Wagler , Pionopsitta Bonaparte , and Pionites Heine. The fallen fruits are also consumed by insects and rodents. The seeds are commonly preyed on by insects, especially Lophopoeum carinatulum Bates ( Coleoptera , Cerambycidae ). Jesel (2005) postulated that, in addition to the zoochory, anemochory provides a limited fruit dispersal (not reaching great distances from the mother tree).

Etymology: —The species was named for the region where the first collections were made.

Phenology: —Flowering has been observed from November to February, fruiting from January to May. Comprehensive and long-term studies on the phenology and ecology of D. guianensis were carried out by Loubry (1993), Caron et al. (1998), and Jesel (2005). Bena (1960) mentioned that D. guianensis lose their leaves after fruiting, but this was not corroborated by any other literature about the genus, herbarium specimens, or several individuals of both species observed by us in different phenological states, including after fruiting.

Uses: —The timber of this species is considered to be of excellent quality, making it one of the most economically important genera of subfamily Dialioideae . The wood is exported from French Guiana and Suriname to markets worldwide; it is reported to be one of the most or even the most exported timber species from French Guiana (Record and Hess 1943, Koeppen 1967, Hammond et al. 1996, Jesel 2005). The wood is rich in silica and exceptionally hard (as in the three closest related genera: Apuleia Mart. , Distemonanthus Benth. , and Dialium ), making it highly resistant to rot and consumption by crustaceans and aquatic mollusks like the “gusano” or “shipworm” (family Teredinidae ). Because these properties it is widely used for shipbuilding and civil construction in general (Record and Mell 1924, Record and Hess 1943, Milanez and Mattos Filho 1959, Kukachka 1964, Koeppen 1967). Its resin can be used to make varnish ( Ducke 1948), although Koeppen (1967) indicated that such use appears to be rare.

Conservation: —The estimated EOO for D. guianensis is 152,312 km, and several authors have indicated that the species is common ( Hammond et al. 1996, Mori et al. 2002; Jesel 2005). Some populations are contained in protected areas, mainly in French Guiana, such as the Parc Amazonien de Guyane, where most of the forest is intact, and logging activities are generally well managed ( Gourlet-Fleury 2004, Guitet et al. 2012). Thus, we tentatively recommend the Least Concern category (LC) for the species. However, we stress that given high harvesting pressure from the timber industry and the unknown impacts of climate change on the species, populations of D. guianensis should be monitored throughout the range of the species. Jesel (2005) conducted comprehensive analyses on reproduction, seed dispersal, and population regeneration following logging.

Vernacular Names: —Through the Guianas, the species has common names mainly in French and in several indigenous languages, such as: Angélique, Angélique gris, Angélique rouge, Angélique blanc (these different names used to different “morphotypes” varying according with timber characters like color), Angélique franc, Angélique batard, Aisili, Bois angelique, Barakaroeballi, Basrakaroeballi, Barakaruballi, Basralocus, Basralokus, Basra loksi, Bastard locus, Dobono-so, Guyana teak, Gueli, Lébi, Lokus, Kabakally, Kire-oe, Kierou, Kiejere-oe, Kieereoe, Sienga pretoe, Sienxa pretoe, Siengdia peto, Siengdia apeto, Singapetou, Tamoene-kereoe, Teck de la Guyane, Teck de Guyane, Wamaradan, and Wéti.

Taxonomic Comments: —An anomalous D. guianensis specimen from French Guiana (A. Gentry 63030) has the second-largest leaflet observed in the genus. It is also the only mature specimen with trifoliolate leaves, an unstable character since it has other leaves presenting five leaflets. D. guianensis is morphologically homogenous, including in the same area of the anomalous specimen. So, this anomaly does not seem to be taxonomically relevant.

Additional Specimens Examined:—Guyana.— ESSEQUIBO: Puruni River , 16-V-1953, Forestry Dept. 7799 ( K); French Guiana.— CAYENNE: Approuague: Fleuve Approuague, riviere Arataye, saut Pararé, 24-IV-1976, Sastre, C. 4828 ( NY; P; U); ancient village Tortue , grande forêt derrièvere village, 28-I-1967, Oldeman 2391 ( P); montagne des Nouragues, bassin de l’Approuague, 4°5’N 52°42’W, 31-XII-1988, Loubry, D. 109 ( US); fleuve Approuague, riv. Arataye, sauts Parare, Barrier, S. 3959 ( MO); Camopi : forêt primaire près de Zidockville, haut Oyapock, 7-VIII-1980, Prévost, M. F. 931 ( P); Trois Sauts , forêt primaire Ipési, 17-V-1975, Grenand 948 ( P); Cayenne: new road to Brazil (route de l’est), 12 kms S of the bridge over the Compté River, Ca. 57 kms S of Cayenne, 9-I-1977, Mori, S. 8885 ( MO; NY; P; US), crique cabaret, 25-I-1966, Oldeman, B. 1950 ( IAN; NY); 12-II-1985, Sauvain, M. 223 ( P); Kourou: environs de la montagne des Singes, exploitation forestière des compagnos Réunis , 23-XII-1981, Granville 4977 ( P; US); Montagne des Chevaux : carrière de la SCC, 4°44’34”N 52°26’12”W, 5-II-2009, Tostain, O. 2347 ( P); St georges: Régina, entre pk 25,4 et 29,2, 4-XI-1998, Grenand 3057 ( NY; U); Route de Cayenne , km 8,750, 27-III-1956. Unknown collector 7421 ( NY); 18-I-1956, Bafog 7181 ( NY); 18-I-1956, Bafog 7184 ( NY); Sinnamary : piste de Saint-Elie, interfluve Sinnamary / Counamana , 5°20’N 53°0’W, 9-I-1991, Prévost, M. F. 2946 ( INPA; NY); 21-XII-1987, Feuillet, C. 4488 ( P); fleuve Sinnamary, sur de la crique Grégoire , au quatrème abattis, 3-V-1968, Oldeman B -1624 ( L; NY; P; US); Sinnamary River, above Petit Saut , 5°0’N 53°1’W, 29-VIII-1993, Mori, S. 23447 ( NY); SW de Sinnamary , piste de St Elie PK 17 ecerex, 15-VIII-1980, Granville 3442- A ( P); forêt Paracou, site expérimental C. T. F. T., bassin du Sinn Paraku, 5°20’N 52°55’W, 1-XII-1986, Sauvain, M. 740 ( P); future nouvelle RN 1, a 1km vers l´ouest a partir du pk 16 de la route d´acess vers petit saut, 5°10’N 52°53’W, 12-II-1988, Billiet, F. 4375 ( RB); Crique Paracou, dispositif sylvolab Paracou , 5°16’N 52°56’W, 31-X-1996, Hallé, F. 4420 ( U); route de Ste. Elie, parcelle arbocel, 25-IX-1977, Sastre, C. 6047 ( P); SW de Sinnamary, piste de St Elie, parcelle biomasse, III-1980, Lescure 889 ( NY; US); St. Elie research station of IRD, 5°18’12”N 53°2’53”W, 27-IV-2007, Redden, K. M. 5979 ( US); Route de L’Est , near the bridge over the Compte, 1976, Champagne, H. 84 ( NY); SAINT LAURENT DU MARONI: 6-III-1994, Angel, G. 57 ( U); 8-IV-1993, Angel, G. 129 ( L); 27-X-1992, Angel, G. 182 ( L); 14-IV-1961, Aubréville 235 ( P; US); Batteufeli s.n. P02743965 ( P); 1861, Mélinon 300 ( F; NY; P; US); Mélinon 333 ( P; R); 1862, Mélinon 593 ( P); Mélinon 618 ( P); 1865, Mélinon P00251023 ( P); Mélinon 28 ( P); Mélinon s.n. P00251020 ( P); P00251021 ( P); P00251032 ( P); P00251033 ( P); P900251042 ( P); P00251037 ( P); P00251038 ( P); P00251039 ( P); P02743943 ( P); P02743944 ( P); Service Forestier 6046 ( P); 6058 ( P); 1856, Sagout 97 ( P); 1874, Sagout 1311 ( P); 1859, Sagout 1211 ( NY; P); 1859, Sagout s.n. P00251024 ( P); 20-X-1938, Vaillant 35 ( P); XII-1919, Wachenhein, G. 50 ( P); 18-XI-1921, Wachenhein, G. 141 ( P); 13-VII-1921, Wachenhein, G. s.n. P00251030 ( P); Charvein : 29-I-1914, Benoist, R. 510 ( P); route de Charvein à l’Acarouany, km 3 coté sud er a 30 metres de la route, 17-XII-1953, Unknown collector P02771988 ( P); Route de Mana, Bafog 7195 ( NY); 21-XII-1955, Bafog 7113 ( NY); 4-I-1956, Bafog 7130 ( NY); 20-I-1956, Bafog 7186 ( NY); 20-I-1956, Bafog 7192 ( NY); 20-I-1956, Bafog 7194 ( NY); 20-I-1956, Bafog 7191 ( NY); 19-XII-1955, Bafog 7103 ( NY); Mana: route de mana, à 1 km après le pour de la crique Marget, à gauche er a 50 mètres de la route, 28-I-1954, Unknown collector P02771986 ( P); Maripasoula : forêt dense entre Maripasoula et Wacapou, 30-VIII-1961, Schnell, R. 11667 ( P); Saul, Monts La Fumée , 3°37’N 53°12’W, 16-IX-1982, Boom, B. 1710 ( NY); 21-IX-1982, Boom, B. 1781 ( NY); 24-IX-1982, Boom, B. 1810 (NY); Boom, B. 1823 ( NY); 8-X-1982, Boom, B. 1903 ( NY); 12-X-1982, Boom, B. 2003 ( NY); 21-X-1982, Boom, B. 2245 (NY); Boom, B. 2246 ( NY); 24-X-1982, Boom, B. 2324 (NY); Boom, B. 2345 ( NY); 25-X-1982, Boom, B. 2366 (NY); Boom, B. 2377 (NY); Boom, B. 2392 ( NY); Saul: near junction of main la Fumée trail with Antenne nord, 3°37’N 53°12’W, 28-III-1983, Mori, S. 15430 ( NY; US); vicinity of Eaux Claires , just S of Eaux Claires on the route de Bélizon, 3°37’N 53°12’W, 9-II-1993, Mori, S. 22890 ( MO; NY; U); 10-I-1996, Mori, S. 24340 ( L; NY; P; US); near the geographic center of the department of Guyane Française, plateau La Duoane and vicinity, 19-XII-1976, Mori, S. 8801 ( NY; P); Saul, forêt humide vers le Crique Limonode , 3-IX-1976, Raynal, A. 18533 ( NY; P); La Fumée mountain trail, between entrance and junction of now defunct Antenne Nord , 3°37’N 53°12’W, 8-I-1996, Mori, S. 24319 ( L; NY; P; US); forêt humide vers la crique Limonade , 9-III-1976, Raynal, A. 18553 ( P); rivière grand Inini, entre Grand Carbet et l’embouchure du Petit Inini, 10-IX-1970, Granville B-3755 ( P); 15-I-1976, Granville 2661- A ( P; US); upland tropical moist forest on lateritic soil, transect No. 5, 3°38’N 53°12’W, 21- VI-1988, Gentry, A. 63030 ( MO); Saint-Laurent: crique Voltaire, camp Voltaire , 5°3’N 54°5’W, 5-XII-1996, Cannell, R. 4509 ( P; U); Unknown collector P04449742 ( P); P00251036 ( P); P00251047 ( P); P00251048 ( P); P00251049 ( P); P00251050 ( P); P02743948 ( P); P02743949 ( P); P02771992 ( P); P02771995 ( P); P02743955 ( P); P02743956 ( P); P02743974 ( P); P03602353 ( P); Suriname.— BROKOPONDO: van Blommesteijn lake, Tonka Island , the almost northern arm, 14-IV-2008, Behari - Ramdas, J. 197 (L); along road to Affobakka, 1 km N of junction with Brokopondo Road , 13- II-1964, Lems, K. 5109 ( NY); MAROWJINE: via secta ab Moengo Tapoe ad Grote Zwiebelzwamp , 22-IX-1948, Lanjouw, J. 389 ( IAN; NY); PARA: Suriname rivier, Suhoza , 29-V-1947, Bosbeheer, L. 60 ( U); Suhoza , 10-VI-1947, Helstone, R. M. 45577 ( WAG; US); Zanderij, grounds of Land’s Bosbeheer , 11-II-1964, Lems, K. 5086 ( NY); Zanderij , Stahel, G. s.n. RB 756216 ( RB); 1-IX-1942, Stahel s.n NY2599 ( NY; IAN; MO); 1-IX-1948, Unknown collector MO3425232 ( MO); 20-III-1923, Unknown collector 6090 IAN 49927 ( IAN; NY); 1-II-1943, Unknown collector s.n. IAN 38284b ( IAN; F; NY); 11-IX-1916, Unknown collector F2037938 ( F); 1-II-1923, BW 2371 ( P); 2-XI-1918, Boswezen, B. W. 4065 ( L; NY; US); Zanderij, Unknown collector NY2574 ( NY); near Zanderij airfield, 14-XII-1946, Navy Project 245 ( F); SIPALIWINI: vicinity of Ulemari River , 13 km upstream from its confluence with Litani River , 3°13’17”N 54°15’31”W, 6-IV-1998, Hammel, B. 21377 ( L; NY; US); 2°46’05”N 54°51’15”W, 6-IV-1998, Evans, R. 2943 ( MO; US); 3°13’17”N 54°15’31”W, 4-IV-1998, Evans, R. 2812 ( QCNE; F; MO; NY; U; US); upper Suriname r., Gran Lio , 1.8 km S from Gran Lio end of Kajana aistrip, 3°53’1”N 55°34’29”W, 16-IX-2005, Hoffman, B. 6397 ( L); UNKNOWN DISTRICT / PROVINCE: 24-I-1907, Bosbeheer 44 ( L; IAN); 2-XII-1922, Forestry Bureau 6028 ( RB); 16-II-1923, Forestry Bureau 6073 ( U); 20-II-1924, Forestry Bureau 6371 ( MO; NY); 14-I-1952, Helstone, E. M. s.n. WAG1016583 ( WAG); 25-X-1926, Junker 501 ( L); XI-1959, Schulz, J. P. 8204 ( WAG); VI-1944, Unknown collector IAN 38284 ( IAN); 12- II-1915, Unknown collector 651 L1957775 ( L); 19-II-1918, Unknown collector 4198 MO1576407 ( MO; NY); 24- XI-1922, Unknown collector F1954879 ( F; NY; US); 11-II-1919, Unknown collector 4254 US1954853 ( US); 8-II-1919, Unknown collector 4237 F1769029 ( F); 15-II-1923, Unknown collector 6071 ( IAN; U); 22-II-1918, Unknown collector 4084 P251017 ( P).