oCoTea BaTaTa (LAuRACEAE), A NEW SPECIES fRoM BRAzIL peDro luís roDrIgues De Moraes1, 2 anD Matheus carvalho vergne1 Abstract. ocotea batata, a new species from the Brazilian Atlantic rain forest of Bahia and Espírito Santo States is described and illustrated. Its relationships within the genus are discussed. Resumo. ocotea batata, uma espécie nova da Mata Atlântica brasileira dos estados da Bahia e Espírito Santo é descrita e ilustrada. Suas relações dentro do gênero são discutidas. Keywords: Lauraceae, ocotea, new species, Bahia, Espírito Santo, Brazil In the course of preparing the treatments of Lauraceae for the flora of Bahia and for the Reserva Natural Vale, Linhares, Espírito Santo, Brazil, an undescribed species of ocotea Aubl. was encountered. Its description and illustrations are presented below. ocotea is the largest genus of Lauraceae in the Neotropics found from Mexico to Argentina. Rohwer (1993) estimated the existence of about 350 species, most of them occurring in tropical and subtropical America and about 50 in Madagascar, seven in Africa, and one in the Canary Islands. More recently, van der Werff (2011) estimated the existence of 350 to 400 species, but an additional 44 species have been described since then by several authors (Brotto and Baitello, 2012; van der Werff, 2012, 2013a, b, 2014; Assis and Santos, 2013), and another 21 will be published soon (H. van der Werff, pers. comm.). In Brazil, there are about 170 species recognized to date (156 in Quinet et al., 2010): at least 48 occur in Bahia, and 32 in Reserva Natural Vale, Espírito Santo. The genus is characterized by having paniculate-cymose inlorescences, lowers with nine 4-celled stamens with the locelli arranged in two superposed pairs and the fruits seated in a cupule (van der Werff, 2009). The genus is likely polyphyletic (Chanderbali et al., 2001) and in need of a revision. However, the large size of the genus makes a revision of the entire group dificult to accomplish, being beyond the scope of most botanists (van der Werff, 2014). The last revision of ocotea sensu Kostermans (1957), including Nectandra Rol. ex Rottb. and Pleurothyrium Nees, dates back to the lauraceae americanae of Mez (1889). Rohwer (1986) published a synopsis of the genus ocotea, proposing subdivision into smaller informal entities, which encompassed 29 groups of species sharing morphological afinities, and 54 species treated singly. No subsequent monographic treatments of such groups have been published, except for the study of the ocotea indecora (Schott) Mez group undertaken by Assis and Mello-Silva (2010). A synopsis of the Central American species was published (van der Werff, 2002), but the South American species are still less well known (Moraes and van der Werff, 2011). MaterIal anD MethoDs This study was based on literature review and Photographs of leaves were obtained with a digital morphological analysis of specimens deposited in the equipment faxitron X-ray (model LX-60 number following herbaria: ALCB, B, BAH, BHCB, BR, C, CEN, 120807305) coupled to a computer with the software CEPEC, CVRD, E, ESA, f, G, G-DC, GoET, Gzu, faxitron DX version 1.0, where the images were captured HAL, HB, HBG, HRB, HRCB, HuEfS, HuESBVC, by using an X-ray exposure time of 19 seconds at a voltage HuNEB, IAN, IBGE, INPA, IPA, K, KIEL, L, LE, LISu, of 30 kV. Photographs of leaf areoles were obtained with M, MBM, MBML, MEL, MG, Mo, NY, oXf, P, PEufR, a light microscope (Leica, M500) equipped with camera R, RB, SP, SPf, SPSf, TuB, u, uB, uEC, uESC, ufP, and software LAS. Leaf diaphanization was done according and VIES (acronyms according to Thiers, continuously to Moraes and Paoli (1999). Descriptive terminology of updated). Photographs of lower structures were obtained leaf venation follows Hickey (1973) and Coe-Teixeira with a stereomicroscope (Leica, M80) equipped with (1980). Abbreviations used in the text are as follows: a camera (Leica, IC80 HD), using the software LAS l. = lower; fr. = fruit; immat. fr. = immature fruit; (Leica Application Suite) Interactive Measurements. ster. = sterile. We thank the curators and staff of the herbaria we consulted (see list below) and of those that sent loans and gifts to HRCB. our sincere gratitude to Mario Tomazello filho (ESALQ/uSP) for making available the X-Ray equipment, to Henrique L. Ribeiro for producing the map, to Geovane S. Siqueira for all support and help during ield work in Reserva Natural Vale, and to Isabel C. L. da Cruz for the diaphanization of leaves. The senior author thanks CAPES for the grant 0781/2014 (Proc. 88881.030430/2013-01), and CNPq for the grant PQ2 (Proc. 304985/2012-0). The junior author thanks CNPq and CAPES for the M. Sc. scholarships. 1 universidade Estadual Paulista “Júlio de Mesquita filho,” Instituto de Biociências, Departamento de Botânica, Av. 24 A 1515, Bela Vista, Caixa Postal 199, 13506-900 Rio Claro, SP, Brazil; 2 Author for correspondence; pmoraes@rc.unesp.br Harvard Papers in Botany, Vol. 22, No. 1, 2017, pp. 61–70. © President and fellows of Harvard College, 2017 ISSN: 1938-2944, DoI: 10.3100/hpib.v22iss1.2017.n7, Published online: 30 June 2017 62 HARVARD PAPERS IN BoTANY VoL. 22, No. 1 noMenclature Ocotea batata P.L.R.Moraes & Vergne, sp. nov. TYPE: lowers: equal in shape to male lowers, larger, 2.8–2.9 × BRAzIL. Espírito Santo: Linhares, Reserva Natural Vale, 2.8–3.6 mm, subssesile; tube deeper, inside pubescent; 19˚11'11.1"S, 39˚54'40.6"W, 12 December 2012, bud, l. ♂, tepals 6, equal, spreading in anthesis, elliptic to ovate, about P. l. R. de Moraes & g. S. Siqueira 3528 (Holotype: HRCB 1.7–2.1 × 1.1–1.3 mm, tip obtuse, roundish, rarely acute, [59814]; Isotypes: CVRD, HBG). fig. 1. papillose, outside pubescent, inside slightly pubescent; ocotea batata is remarkable among the Brazilian staminodia 12 (whorls I, II, III, and IV), equal in shape to species of the genus and can be separated from them by a the stamens of male lowers, but smaller, clavate, glabrous, combination of a peculiar issured, yellowish bark, twigs pilose at the base; whorl I 0.7–0.8 mm long; whorl II 0.7– with lenticels, glabrous leaves above, pubescent below, 0.8 mm long; whorl III 0.8–1.3 mm long, glands globose, with barbellate-foveolate domatia in the axils of the primary 0.3–0.4 × 0.4–0.6 mm, stalk 0.1–0.3 mm long; whorl IV and secondary veins, very short, axillary, pubescent, fewliguliform, densely pubescent, 0.6 mm long; pistil glabrous, lowered inlorescences, fruit ellipsoid seated on a shallow, 1.2–1.7 mm long, style cylindrical, robust, twisted, 0.6 infundibuliform cupule. mm long, ovary urceolate, 0.9–1.0 × 0.7–0.9 mm, stigma Trees up to 7 m. Twigs terete, angular when young, discoid, large. Fruits dark purple in live material, 1.3–1.7 glabrous to glabrescent at the base, dense to sparse × 1.1–1.3 cm, ellipsoid, subglobose, exserted from the pubescent or sericeous near the tip, with long, straight and shallow, 0.7–0.9 × 0.5–0.9 cm, infundibuliform cupule (red appressed trichomes, lenticelled; terminal buds pubescent, in live material); the pedicel swollen, but shrunken in dried with yellowish trichomes. leaves alternate, 3.5–16.8 × 1.2– material. figs. 2–4. 6.2 cm, elliptic to ovate to obovate, chartaceous, glabrous Phenology: flowers collected from December to April. above, except for some sparse trichomes on midrib, Immature fruits collected from September to December; papillose, pubescent below, with ± short, straight, appressed mature fruits collected in December. trichomes, papillose, the base acute, short cuneate, Etymology: The common name of the new species asymmetric, the margin scleriied, lat, the apex shortly to has been coined by Domingos folli as “canela-batata,” in long acuminate; venation pinnate, eucamptodromous to allusion to the color of the bark of the trunk that resembles weak brochidodromous, areole development incomplete, the color of the peel of potato tubers (Solanum tuberosum randomly arranged, with branched veinlets; midrib and L.), which is called “batata” in Portuguese. We propose the secondary veins impressed above, raised below, secondary speciic epithet name batata, based on its common name, veins 3–5 on each side of the midrib; barbellate-foveolate here used as a noun in apposition, thus in reference to the domatia in the axils of secondary veins; petioles 0.4–1.4 peculiar color of the bark of the trunk, which is unusual for cm long, semiterete, lat to slightly canaliculate above, the known Brazilian species of ocotea. glabrescent in older leaves, densely pubescent in young Distribution and habitat: ocotea batata is known ones. Inlorescences pubescent, tyrsoid, much shorter than only from few locations of the Atlantic rain forest domain leaves, 0.5–1.5 cm long, few-lowered, mainly in the axils (fig. 5). It is relatively abundant in the understory of the of distal leaves, peduncle short. Flowers unisexual, white tabuleiro forest (lowland ombrophilous dense forest) of in live material, tube shallow, urceolate, with very short Reserva Natural Vale, Linhares, Espírito Santo, in an pedicels, sparsely pubescent, with long, appressed, straight altitudinal range of 28–65 m. It has been also collected to curled trichomes; male lowers: inside pubescent, pedicels in montane ombrophilous dense forest in Santa Lúcia 1.0–1.3 × 0.7–0.8 mm, fertile stamens 9 (whorls I, II and Biological Station, Santa Teresa, Espírito Santo, at 650 III) exserted, all 4-celled, 2.2–2.4 × 3.1–4.4(–5.0) mm, the m, and in the region of Arataca, southern Bahia, in an ilaments equal to shorter than the anthers, the outer six with altitudinal range of 500–800 m. It is also registered from the locelli introrse, the inner three with the locelli extrorsesmall fragments of submontane ombrophilous dense forest lateral and 2 globose glands at the base; tepals 6, equal, in the region of Itamaraju, at about 115 m. spreading in anthesis, about 1.7–2.7 × 1.3–1.8 mm, elliptic Additional specimens examined: BRAzIL. Bahia: to ovate, outside pubescent, with straight, appressed, short Arataca, Rod. Arataca/una, Serra do Peito de Moça, RPPN to long trichomes, inside glabrous to slightly pubescent, Palmeiras/IESB, trilha que leva ao topo da serra, 15˚10'27"S, the trichomes as the ones on outside, tip roundish, obtuse, 39˚20'22"W, 500–800 m, 18 December 2005, immat. fr., j. g. papillose; stamens of whorl I glabrous, 1.5–2.0 mm long, jardim et al. 4869 (CEPEC, NY, RB); Itamaraju, fibrasa, anthers ovate, tip truncate, subemarginate, 0.9–1.0 × 3 December 2014, l. ♀, fr., P. l. R. de Moraes et al. 4328 0.9–1.0 mm, ilaments slender, slightly pilose at the base, (HRCB, Mo). Espírito Santo: Linhares, Reserva Natural 0.6–1.0 mm long; stamens of whorl II equal to the whorl I, Vale, estrada Paraju km 0.15, 5 october 1982, immat. fr., slightly shorter, 1.3–1.8 mm long, anthers 0.9–1.0 × 0.7– D. a. Folli 401 (CVRD, ESA, Mo, SPSf); idem, BR 101 0.8 mm, ilaments 0.6–0.9 mm long; stamens of whorl III km 1.2, próximo ao Córrego Pau Atravessado, 3 April clavate, glabrous, 1.7–1.9 mm long, anthers rectangular, tip 1999, l. bud, D. a. Folli 3390 (CVRD, HRCB, HuEfS, truncate, subemarginate, papillose, 0.8–0.9 × 0.6–0.7 mm, Mo); idem, estrada Gávea, RfL-01/80 Bloco D Trat-5, ilaments slender, 0.8–0.9 mm long, glands globose, slightly 23 February 2004, l. bud, D. a. Folli 5033 (CVRD, ESA, shorter than the ilaments, 0.5–0.7 × 0.6–0.8 mm, stalk 0.2– HRCB, HuEfS, RB); idem, estrada Alameda 03, 3 March 0.3 mm long; stamimodia of whorl IV liguliform, densely 2005, l. ♂, D. a. Folli 5035 (CVRD, HRCB, HuEfS); pubescent, 0.4 mm long; pistillode glabrous, 1.8–2.0 mm idem, estrada Peroba Amarela, 19˚09'45.5"S, 40˚04'11.5"W, long, stipitiform, stigma discoid, well-developed; female 16 November 2006, immat. fr., a. quinet 1051 (CVRD, 2017 DE MoRAES AND VERGNE, NEW oCoTea (LAuRACEAE) fRoM BRAzIL fIgure 1. Holotype of ocotea batata P.L.R. Moraes & Vergne (Moraes 3528, HRCB). 63 64 HARVARD PAPERS IN BoTANY HRCB); idem, estrada Louro, 12 April 2010, l. ♂, g. S. Siqueira 541 (CVRD, HRCB); idem, estrada Ipê Amarelo, 12 February 2011, l. ♂, D. a. Folli 6765 (CVRD, HRCB); idem, 6 September 2011, ster., P. l. R. de Moraes et al. 3175 (HRCB); idem, 19˚07'01.6"S, 39˚55'05.3"W, 11 December 2012, l. ♀, P. l. R. de Moraes & g. S. Siqueira 3530 (CVRD, HBG, HRCB); idem, estrada Jueirana Vermelha, 10 September 2015, immat. fr., P. l. R de Moraes & M. C. Vergne 4957 (HRCB); Santa Teresa, Estação Biológica de Santa Lúcia, Valsugana Velha, 650 m, 19 March 1999, VoL. 22, No. 1 l., l. Kollmann & e. Bausen 2193 (MBML, SPf, uEC); Sooretama, Reserva Natural Vale, estrada Jueirana facão, 3 october 2014, immat. fr., g. S. Siqueira 1010 (CVRD, HRCB, RB); idem, Reserva Biológica de Sooretama, área da sede, 14 March 1972, l., D. Sucre 8693 (Mo, RB). Given the combination of morphological characters found in ocotea batata (Table 1), it is dificult to place the new species in any of the informal groups of species with unisexual lowers proposed by Rohwer (1986). It appears to it best in the ocotea dispersa group, which includes fIgure 2. Leaf venation of ocotea batata P.L.R. Moraes & Vergne (Moraes 3528). A, mature leaf, pinnate, weak brochidodromous; B, detail of reticulation, areoles incomplete, randomly arranged; C, detail of areoles incomplete, with branched veinlets. Bars = 1 cm (A); 0.33 cm (B); 0.5 mm (C). 2017 DE MoRAES AND VERGNE, NEW oCoTea (LAuRACEAE) fRoM BRAzIL 65 fIgure 3. ocotea batata P.L.R. Moraes & Vergne. A, trunk showing the peculiar yellowish bark; B, branch with inlorescences (Moraes 3528); C, female lower (Moraes 3530); D, immature fruit (Moraes 4957); E, mature fruit (Moraes 4328). 66 HARVARD PAPERS IN BoTANY VoL. 22, No. 1 fIgure 4. ocotea batata P.L.R. Moraes & Vergne. A–I. Male lower from Moraes 3528 (HRCB). A, lower; B, outer tepals, abaxial and adaxial surfaces; C, inner tepals, abaxial and adaxial surfaces; D, stamen of whorl I; E, stamen of whorl II; F, stamen of whorl III; G, gland; H, staminode of whorl IV; I, pistillode. J–R. Female lower from Moraes 3530 (HRCB). J, lower; K, outer tepals, abaxial and adaxial surfaces; L, inner tepals, abaxial and adaxial surfaces; M, staminode of whorl I; N, staminode of whorl II; O, staminode of whorl III; P, gland; Q, staminode of whorl IV; R, pistil. Bars = 1 mm (A, J); 0.5 mm (B, C, D, E, f, G, I, K, L, M, N, o, Q, R); 0.2 mm (H, P). 2017 DE MoRAES AND VERGNE, NEW oCoTea (LAuRACEAE) fRoM BRAzIL fIgure 5. Geographic distribution of ocotea batata P.L.R. Moraes & Vergne. 67 68 table 1. Comparative morphology and geographic distribution of ocotea batata and putative related species (States of Brazil: AM = Amazonas, AP = Amapá, BA = Bahia, ES = Espírito Santo, MG = Minas Gerais, PA = Pará, PR = Paraná, RJ = Rio de Janeiro, SC = Santa Catarina, SP = São Paulo). O. dispersa O. divaricata O. percurrens Habit Tree (7 m) Tree (16 m) Tree (18 m) Tree (35 m) Twig indument Dense to sparsely pubescent at the tip, glabrous to glabrescent at the base Pubescent at the tip, glabrescent at the base Glabrous to sparsely pubescent or pubescent Pubescent Twig lenticels Present Present Present at the base Absent Phyllotaxy Alternate Alternate to subopposite at tip Alternate Alternate Leaf form Elliptic to ovate to obovate Elliptic to obovate Wide-elliptic to obovate Elliptic Leaf size (cm) 3.5–16.8 × 1.2–6.2 3.5–13.4 × 1.7–5.2 2–20 × 1.6–15 4.8–13.7 × 2.3–5.1 Leaf indument Glabrous above, pubescent below Glabrous to glabrescent above, sparsely to dense pubescent below Glabrous above, glabrescent below Glabrous above, glabrescent to pubescent below With trichomes and pit With trichomes and pit With trichomes and pit With trichomes Petiole length (cm) 0.4–1.4 0.6–1.1 0.4–1.8 0.5–1.2 Petiole-cross section Slightly canaliculate Canaliculate Slightly canaliculate Non-canaliculate Glabrescent Glabrescent to pubescent Glabrous Pubescent Inlorescence indument Pubescent Pubescent Glabrous Pubescent Inlorescence size (cm) 0.5–1.5 2–3 5—10 4–7.2 Sparsely pubescent Dense to sparsely pubescent Glabrescent Pubescent 2.2–2.4 × 3.1–4.4(–5.0) 2–4 × 2–3 2.1–2.4 × 2–2.8 3.0–3.7 × 2.5–3 1.7–2.7 × 1.3–1.9 0.9–1.1 × 0.5–0.8 1.3–2 × 1–2 1.0–1.7 × 0.6–1.1 Domatia Petiole indument flower indument flower size (mm) ♂ Tepal size (mm) ♂ VoL. 22, No. 1 O. batata HARVARD PAPERS IN BoTANY CharaCters CharaCters O. dispersa O. divaricata O. percurrens outer stamens length (mm) ♂ 1.3–2.0 1–1.5 1.1–1.3 0.6–1.2 Inner stamens length (mm) ♂ 1.7–1.9 1.0–1.6 1.1–1.2 0.8–1.3 1.8 Absent to vestigial Absent to present: 0.3–0.4 1.1–1.8 flower size (mm) ♀ 2.8–2.9 × 2.8–3.6 2.1–3 × 2.1–2.2 1.9–2.2 × 1.9–2 2.8–3.7 × 2.9–3.4 Tepal size (mm) ♀ 1.7–2.1 × 1.1–1.3 0.9–1.4 × 0.6–1.1 1.2–1.3 × 0.9–1.2 0.9–1.2 × 0.6–0.8 outer staminodia length (mm) ♀ 0.7–0.8 0.6–0.8 0.7–0.8 0.4–0.5 Inner staminodia length (mm) ♀ 0.8–1.3 0.7–0.8 0.7–0.9 0.5–0.6 Glabrous Glabrous Glabrous Glabrous or nearly so Pistil length (mm) ♀ 1.2–1.7 1.6 1–1.1 1.5–1.6 Style length (mm) ♀ 0.6 0.5–0.6 0.04–0.09 0.5–0.7 ovary length (mm) ♀ 0.9–1.0 0.8–0.9 0.8–1.2 0.6–0.8 1.3–1.7 × 1.1–1.3 0.9–1.2 × 0.6–0.8 0.7–2 × 0.6–2.5 0.9–1.5 × 0.5–0.8 understory of lowland, submontane and montane ombrophilous dense forests understory of lowland and montane ombrophilous dense forests, and in montane ombrophilous mixed forest Submontane and montane ombrophilous dense forests, in semideciduous forests, and in restinga Northeastern part of the Amazon Basin, with disjunct distribution in the Atlantic forest in Bahia BA, ES ES, MG, PR, RJ, SC, SP BA, ES, MG, RJ, SP Brazil: AM, AP, BA, PA; french Guiana; Suriname Pistillode (mm) ♂ Pistil ♀ fruit size (cm) Habitat Geographical distribution DE MoRAES AND VERGNE, NEW oCoTea (LAuRACEAE) fRoM BRAzIL O. batata 2017 table 1 cont. Comparative morphology and geographic distribution of ocotea batata and putative related species (States of Brazil: AM = Amazonas, AP = Amapá, BA = Bahia, ES = Espírito Santo, MG = Minas Gerais, PA = Pará, PR = Paraná, RJ = Rio de Janeiro, SC = Santa Catarina, SP = São Paulo). 69 70 HARVARD PAPERS IN BoTANY VoL. 22, No. 1 The fruit specimen collected by jardim 4869 in Arataca, o. dispersa (Nees & Mart.) Mez, o. nutans (Nees) Mez, and Bahia was identiied by A. Amorim (CEPEC) and by o. glauca (Nees & Mart.) Mez, which are essentially Brazilian L. Assis (RB) as allied to ocotea divaricata (Nees) Mez, a species. Characteristic of this group are the relatively small species that belongs to the ocotea cernua (Nees) Mez group trees (up to 16 m), leaves about 5–15 cm long, shape varying proposed by Rohwer (1986). Although the leaves and fruits of from ± elliptic, ovate-elliptic to oblanceolate, inlorescences that specimen at irst glance generally would resemble those shorter than the leaves and only slightly branched, lower of o. divaricata in shape and size, a detailed examination of diameter about 3–4 mm, pistillode varying from relatively the exsiccate shows that the indument on the twigs, leaves well developed (stipitiform) to completely reduced and petioles, presence of barbellate-foveolate domatia, (vestigial), pubescent or glabrous, fruit elliptic seated on a and the axillary, short pedicelled fruits, developed from cup-shaped to almost hemispherical cupule, which usually no peduncle, closely match the other known specimens of keeps remnants of tepals on the single rim. Actually, o. o. batata. Since o. divaricata has inlorescences relatively batata resembles o. dispersa in the general vegetative large, often with an axis in zigzag, many-branched, manyaspect and some herbarium specimens have been wrongly lowered, usually glabrous to sparsely pubescent, its fruits determined as the latter (Kollmann 2193; Sucre 8693). Both species share features like twigs with lenticels, leaves of usually have relatively long pedicels developed from the almost same shape and size, barbellate-foveolate domatia in branches of former inlorescences, thus not straight from the axils of secondary veins, short inlorescences, pubescent, the axils of leaves or bracts like in o. batata. Moreover, and lowers short pedicelled. However, they differ in the o. batata could not be closely related to o. divaricata different indument of twigs and leaves: the trichomes of o. because their loral parts are quite distinct. dispersa are longer, usually denser, and ascending, while in ocotea batata could also be confounded at irst glance o. batata they are appressed. Additionally, the trunk of o. with o. percurrens Vicentini, a species with a disjunct batata has the bark issured and yellowish vs. smooth and distribution between the Amazon and Atlantic forest greyish in o. dispersa, and lowers relatively wider, fruits domains. Their leaves can be similar in shape and size larger, cupule infundibuliform, shallow, smooth, enclosing and in the general aspect of the dried material. However, only the base of the fruit vs. cup-shaped, usually 6-lobed, o. percurrens is a tall tree and also differs in the indument warty, enclosing about 1/3 of the fruit in the latter. The male covering the terminal buds and twigs, by erect and crisped lowers of o. dispersa have smaller stamens, stout ilaments, trichomes, petioles non canaliculate, barbellate domatia, absent or liguliform staminodia, and vestigial or absent smaller ilaments of stamens of male lowers, pistillode pistillode, while in o. batata the ilaments are relatively with stigma inconspicuous, and female lowers with tube slender, the liguliform staminodia are always present, and glabrous inside, for instance. the pistillode is stipitiform. lIterature cIteD assIs, l. c. s., anD r. De Mello-sIlva. 2010. Taxonomic and nomenclatural changes in the ocotea indecora group (Lauraceae). Novon 20(4): 377–380. assIs, l. c. s, anD M. f. santos. 2013. ocotea grandifructa (Lauraceae), a novelty from the Brazilian Atlantic forest. Novon 22(3): 265–270. brotto, M. l., anD J. b. baItello. 2012. uma espécie nova de Lauraceae da loresta atlântica do Brasil. 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