Guibourtia chodatiana (Hassler) J. Léonard (1949: 401)

Pinto, Rafael Barbosa, Mansano, Vidal De Freitas, Sartori, Ângela Lúcia Bagnatori & Tozzi, Ana Maria Goulart De Azevedo, 2017, Revisiting The Taxonomic Diversity Of Guibourtia In The Neotropics (Leguminosae, Detarioideae), Phytotaxa 327 (2), pp. 130-140 : 132-136

Guibourtia chodatiana (Hassler) J. Léonard (1949: 401) . Copaifera chodatiana Hassler (1903: 119) . Pseudocopaiva chodatiana (Hassler) Dwyer (1944: 7). Type:— PARAGUAY. “Ad ripam minori in rivi cursus superiori fluminis Apa”,

November 1901 (fl.), E. Hassler 7984 (Lectotype: G381541, designated here, Isolectotypes: A, BM, F, G, GH, MICH,

MO, MPU, NY, P, S, UC). Fig. 2 .

Guibourtia chodatiana var. fruticosa (Hassler) J. Léonard (1949: 401) , syn. nov. Copaifera chodatiana var. fruticosa Hassler (1903: 119) . Type:— PARAGUAY. “In sabulosis humidis in regione cursus superioris fluminis Apa”, November 1901 (fl.), E. Hassler 8046 (Lectotype: G00381557, designated here, Isolectotypes: A, G, K, MICH, MPU, NY, P, RB, S, UC).

Copaiba kuntzei (Harms ex Kuntze) Kuntze (1893: 56) , syn. nov. Copaifera kuntzei Harms ex Kuntze (1893: 56) . Type:— BOLIVIA. Oto- Velasco, 200m, July 1892, O. Kuntze s.n. (Holotype: B [Fneg1513.], Isotypes: F, NY).

Trees up to 10 m tall; bark smooth, slightly striate; branchlets glabrous. Petioles dilated at base and apex, 1–2.7 cm long, tomentose to glabrescent; petiolules 2.1–3.4 mm long, tomentose to glabrescent; leaflet blades membranaceous, asymmetric, usually ovate, rarely wide-elliptic, 5–10 × 2.4–4.1 cm, the base obliquely cordate, asymmetric, the apex acuminate to rounded, the abaxial surface sparsely villous to glabrescent, with the midrib densely tomentose to glabrescent, the adaxial surface glabrous, tertiary venation slightly apparent to inconspicuous adaxially. Inflorescence a simple raceme or panicle, 2–8 cm long, axillary, the flowers 18–23 per axis, spirally arranged, the axis terete in cross section, 0.8–1.4 mm wide, puberulent to tomentose; bracts ovate, ca. 2 × 1.3 mm, tomentose on both sides, caducous. Pedicels terete in cross section, 1.4–1.7 × 0.5–0.6 mm, tomentose; bracteoles alternate, inserted near middle of pedicel, caducous; flower buds dark brown to black when dried, orbicular to ellipsoid, 3–3.5 × 2–2.5 mm, tomentose to glabrescent. Calyx green when fresh with white or cream hairs, dark brown to black when dried, densely tomentose internally, sparsely tomentose to glabrescent externally, sepals 4, subequal, elliptic, slightly verrucose, 2.7–3.8 × 1.9– 2.4 mm. Corolla lacking. Stamens 10, glabrous, the filaments 4–4.5 mm long, cream to white, the anthers cream, elliptic, 1.6–2 × 0.8–1 mm, glabrous. Gynoecium light green, tomentose at the base and along the sutures of the carpel; the stipe terete in cross section, 0.5–1.2 mm long, glabrous, the ovary D-shaped in outline, 1.2–1.5 × 1.2–1.5 mm, ovules 2, the style terete in cross section, 1.5–2.5 mm long, glabrous, the stigma punctiform. Fruits follicular, green when immature and dark brown to black when mature, minutely verrucose, glabrous, the body D-shaped in outline, with part of the style persistent, 2.4–3 × 1.3–2 cm. Seeds 1 per fruit, 1.8–2 × 0.9–1 cm, smooth, red to brown, ellipsoid, with a sub-apical depression on the funicular side, this covered by the white aril.

Phenology:— Flowering October to December, fruiting June to May.

Distribution and habitat:— Guibourtia chodatiana occurs in South America, in dry forests, riparian forests and seasonal forests, sometimes near rivers. In Pantanal occurs in the plateau as Serra do Urucum. This species is cultivated at the arboretum of the Botanical Garden of Rio de Janeiro and also in the Universidade Federal Rural do Rio de Janeiro —UFRRJ. ( Fig. 1 ).

Additional specimens examined: — BOLIVIA. Santa Cruz: Ñuflo de Cháves, Región de Lomerío, a 200m al sur de la plaza, Comunidad de Palmira , 16°45’09”S 61°56’05”W, 16 November 1994 (fl.), M. Toledo et al. 446 ( NY) ; Puerto Isabel, Baía Mandioré , 18°11’49,1”S 57°30’37,8”W, 23 November 2010 (fl.), T. S. Yule et al. 75 ( CGMS) . BRAZIL. Bahia: Campo Formoso, Estrada Alagoinhas-Água Preta , without date (fr.), L. Coradin et al. 6056 ( RB) ; Casa Nova, dunas do Rio São Francisco , 09°24’37”S 41°08’59”W, 04 July 2007 (fr.), G. Pereira-Silva et al. 9164 ( CEN; HUEFS) ; Casa Nova , 27 April 2001 (fr.), R. M. Harley et al. 54348 ( HUEFS) ; Casa Nova , 09°25’06”S 41°08’56”W, 18 April 2004 (fr.), T. S. Nunes et al. 1123 ( HUEFS) ; Casa Nova , 09°24’57”S 41°08’24”W, 05 July 2003 (fl.), L. P. Queiroz et al. 7920 ( HUEFS) ; Casa Nova , 09°24’37”S 41°08’54”W (fr.), L. P. Queiroz et al. 7418 ( HUEFS) ; Jequié, Morro da Torre , 13°53’27”S 40º07’20”W, 13 April 2007 (fr.), L. P. de Queiroz et al. 12910 ( HUEFS) ; Paramirim, Caminha Catuarama para Mateus , 13°17’50”S 42°14’44”W, 28 April 2007 (st.), A. A. Conceição et al. 1940 ( HUEFS) ; Rui Barbosa, Serra do Orobó, 12°17’59”S 40°29’28”W, 20 August 2008 (st.), J. Costa et al. 1299 ( HUEFS). Goiás: Edéia , 17°46’06”S 50°04’22”W, 02 April 2013 (fr.), C. E. B. Proença 4504 ( RB) ; Iaciara, Vão do Paranã, 13°58’21”S 46°58’50”W, 07 July 2013 (fr.), R. F. Haidar et al. 1769 ( CEN). Mato Grosso do Sul: Bonito, às margens do Rio Formoso , propriedade Jardim das Águas, 16 June 2000 (fr.), R. Constantino 10 ( CGMS) ; Bonito, Rancho do Netinho , 21°25’13”S 56°27’02”W, 14 August 2004 (fr.), J. L. Battilani 62 ( CGMS) ; Bonito, Serra da Bodoquena, Rodovia MS-382, 24 May 2002 (fr.), G. Hatschbach et al. 73394 ( MBM; RB) ; Bonito, Gruta de Nossa Sra. Aparecida , 29 August 1989 (fr.), P. Cunha s/n ( RB601744 ) ; Corumbá, Serra Grande, Planalto Residual do Urucum , 27 July 2004 (fr.), R. R. Silva & J. S. Velasquez 1222 ( COR; UEC) ; Corumbá, Estrada da Mina , 21 October 1994 (st.), S. M. de Faria & S. A. Rodrigues 823 ( RB) ; Corumbá, Estrada Corumbá-C. Grande , 22 May 1992 (fr.), S. M. de Faria s/n ( RB315481 ) ; Jardim, Próximo ao Rio Miranda , 21°28’03”S 56°08’11”W, 12 December 2001 (fl.), J. L. Battilani 100 ( CGMS) ; Ladário, Fazenda Urúba, 08 June 1994 (fr.), G. Hatschbach , V. Pott & J. M. Silva 60775 ( MBM) ; Ladário, Serra do Rabichão, Planalto Residual do Urucum , acesso pela fazenda Uruba , 17 November 2004 (fl.), R. R. Silva & M. V. Silva 1278 ( COR; UEC). Minas Gerais: Jacinto, 5–10km E, 12 February 1989 (fr.), G. Hatschbach & J. Cordeiro 52720 ( NY) ; Januária, about 1km E of Rio Pandeiros, 18 April 1973 (fl., fr.), W. R. Anderson 9115 ( NY; RB; U) ; Jequitinhonha, Mumbuca , 30 Oct 2014 (fr.), H. C. Lima 7860a ( RB) ; Santa Fé de Minas, RESEX Santa Fé de Minas, 16°39’44”S 45°26’25”W, 10 August 2014 (fr.), R. P. Athaídes & F. Haidar 14 ( CEN). Rio Grande do Norte: Serra Nova, 17 July 1991 (fr.), M. A. Figueiredo et al. 271 ( EAC). Rio de Janeiro: Itaguaí, Campus da UFRRJ, 14 June 1992 (fr.), A. L. Peixoto s/n ( RB322348 ). São Paulo: Campinas, Gramado ao lado do Instituto de Matemática da Unicamp , 22°48’56”S 47°04’05”W, 30 August 2013 (fr.), L. P. Queiroz & J. Semir 15826 ( HUEFS) ; Tanabí , 23 August 1941 (fr.), A. Gehrt s/n ( MBM167431 ; SP45855 ) . PARAGUAY. Amambay: Bella Vista, forest along Rio Apa , 24 March 1983 (fr.), J. E. Simonis et al. 191 ( U) ; Cerro Corá, Parque Nacional Cerro Corá , 14 December 1999 (fl.), M. S. Ferrucci et al. 1358 ( U) ; Cerro Corá , [without date] (fl.), M. Bernardi 18946 (?) ( U) ; Estancia Carumbé , 27 November 1969 (fl.), T. M. Pedersen 9423 ( L) .

Taxonomic comments: —Within Guibourtia , the species G. chodatiana is characterized mainly by its membranaceous leaflets, with the lamina ovate to wide-elliptic, and by its tomentose ovary ( Figure 2 ). Hassler (1903) first described G. chodatiana recognizing two varieties within it. He suggested that G. chodatiana var. fruticosa Chodat & Hassler (1903: 1047) differs from the typical variety in its shrubby habit, leaflet apex form (subacute-acuminate vs. obtuse-acuminate), and inflorescence length (20–25 mm vs. 40–50 mm). In our analysis, we found that these features show extensive and continuous variation among collections of G. chodatiana , thus providing no basis for recognizing infraspecific taxa.

Copaifera kuntzei Harms ex Kuntze was described based on a fruiting collection from Bolivia. Until that time, the only known bifoliolate species of the genus Copaifera was C. hymenaeifolia Moric. from Cuba and, because of this and the similar apetalous flowers, Kuntze (1893) included it in Copaifera even though he commented that the species had very few translucid glands on its leaves, in contrast to the other American species of Copaifera . Subsequently, Léonard (1949) synonymized Copaifera kuntzei under Guibourtia confertiflora . However, the morphology of the type collection differs from G. confertiflora in the size, shape and asymmetry of the leaflets, but the taxon fits squarely within our circumscription of Guibourtia chodatiana .

The two lectotypifications here proposed are necessary because, although Hassler cited collection numbers in his protologue, he did not specify in which herbarium his collections were housed, nor which duplicate should be considered as the holotype. We here propose as lectotypes specimens lodged in the herbarium where Hassler worked and where he deposited the top set of his collections.

Guibourtia hymenaeifolia (Moric.) J. Léonard (1949: 401) . Copaifera hymenaeifolia Moric. (1833: 529) . Copaiba hymenaeifolia (Moric.) Kuntze (1891: 172) . Pseudocopaiva hymenaeifolia (Moric.) Britton & Wilson (1929: 28) . Type:— CUBA. Quiebra hacha, [without date] (fl.), R. de la Sagra s/n (Lectotype: P00481483, designated here, Isolectotype: P00481481). Fig. 3 .

Trees up to 18 m tall; bark usually peeling; branchlets glabrescent. Petioles basally and apically swollen, 1.1–1.7 cm long, glabrous; petiolules 2.2–3.3 mm long, glabrous; leaflet blades chartaceous, strongly asymmetric, ovate to elliptic, 4.8–8.8 × 2–4.3 cm, the base obliquely cordate to sligthly rounded, asymmetric, the apex acuminate to cuspidate, glabrous on both the abaxial and adaxial surfaces, tertiary venation strongly conspicuous and salient on both sides. Inflorescence a simple raceme or panicle, axillary, the flowers 20–26 per axis, spirally arranged, the axis terete in cross section, 0.7–0.83 mm wide, sparsely puberulent; bracts caducous. Pedicels terete in cross section, 3–4 × 0.4–0.6 mm, glabrescent; bracteoles alternate, located at the middle of the pedicel, caducous; flower buds dark brown to black when dried, ellipsoid, 2.9–3.8 × 2–2.5 mm, glabrescent. Calyx dark brown to black when dried, densely velutinous internally and glabrescent externally, the sepals 4, subequal, elliptic, verrucose, 3.6–3.8 × 2–2.6 mm. Stamens 10, glabrous, the filaments 4.3–4.8 mm long, the anthers oblong to broadly-elliptic, 1.1–1.4 × 0.8–1 mm, glabrous. Gynoecium glabrous; the stipe terete in cross section, 1–1.2mm long, glabrous, the ovary D-shaped in outline, 1.2–1.8 × 1.1–1.3 mm, glabrous, ovules 2, the style terete in cross section, 2.5–2.8 mm long, glabrous, the stigma punctiform. Fruits not seen.

Phenology:— Flowering May to July, fruiting February to June.

Distribution and habitat:— The species is endemic to Cuba where it occurs in the provinces of Franqueville, Guantánamo, Holguín, and Las Villas. Little information is available on the habitat preferences of G. hymenaeifolia . It is recorded as growing in dry forests, sometimes near to rivers and in disturbed areas. ( Fig. 1 ).

Additional especimens examined: — CUBA. [Without locality], [without date] (fl.), R. de la Sagra s/n ( B100846994 ; G405535 ; K555226 ) ; [without locality], 1831, R. de la Sagra 138 ( G) ; [without locality], [without date] (fl.), R. de la Sagra 241 ( BM; L) ; [without locality], [without date] (fl.), R. de la Sagra 138 ( F). Franqueville , May 1865 (fl.), R. de la Sagra 240 ( K) ; Las Villas: Cienfuegos, Belmonte, Soledad , 6 February1928 (fr.), J. G. Jack 5019 ( HAC) ; Guantánamo, Cerca del Central San Antonio, 14 April 1940 (fr.), León & Victorin 17726 ( HAC). Holguín: Moa, Bosque este del aeropuerto, 26 June 1945 (fl., fr.), Clemente 4356 ( HAC) ; Moa, Cayoguán, Río Cromita , 25 July 1944 (fl.), Clemente & Alain 4096 ( HAC) ; Moa, Sierra de Moa , 20km W of Compania de Moa mil, July 1941 (fl.), R. A. Howard 5918 ( U) ; Quiebra hacha, [without date] (fl.), R. de la Sagra s/n ( P00252852 ). Oriente: Boyate , 05 July 1915 (fl.), E. L. Ekman 6203 ( U). Villa Clara: Santa Clara , 8 miles north of Soledad , 25 July 1941 (fl.), R. A. Howard & F. Gonzalez 6598 ( L; U) .

Taxonomic comments: —The geographical distribution of Guibourtia hymenaeifolia in Cuba is clearly disjunct from populations of G. chodatiana and G. confertiflora ( Fig. 4). Guibourtia hymenaeifolia also differs from G. confertiflora in its chartaceous (vs membranaceous) and strongly asymmetric (vs. symmetric) leaflet lamina. From G. chodatiana , it differs in its glabrous (vs tomentose) ovary. Moreover, the venation on the adaxial surface of the lamina in G. hymenaeifolia is much more prominent especially the tertiary and quaternary venation when compared to those of G. chodatiana and G. confertiflora .

The lectotypification was necessary here because Moricand (1833) neither cited the collection number of the type specimen, nor the herbarium where the specimen is housed. The only information about the original material is the name of the collector (R. de la Sagra). We here designate the Paris collection P00481483 of R. de la Sagra s.n. as the lectotype because the material best matches the description in the species protologue and is a well preserved specimen.

Figures

FIGURE 1. Distribution map of the Neotropical Guibourtia species.

FIGURE 2. Guibourtia chodatiana. A. branch with inflorescences; B. leaflet and detail of the abaxial surface; C. floral bud; D. flower; E. immature fruits; F. detail of the epicarp; G. dehiscent fruits; H. fruit with the exposed seed and aril; I. seed (A–C: R. R. Silva & M. V. Silva 1278 (UEC); D: Bernardi 18946 (U); E–I: R. R. Silva & J. S. Velásquez 1222 (UEC)) Drawn by Klei Souza.

FIGURE 3. Guibourtia hymenaeifolia. A. branch with inflorescences and immature fruits; B. leaflet and detail of the abaxial surface; C. floral bud; D. flower; E. detail of the glabrous ovary; F. immature fruit (A–F: E. L. Ekman 6203(U)) Drawn by Klei Souza.