Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, 2022

de Moraes, P. L. R., Cardoso, D. B. O. S., Gregório, B. S. & Baitello, J. B., 2022, A REMARKABLE NEW GYNODIOECIOUS SPECIES OF OCOTEA (LAURACEAE) FROM THE BRAZILIAN ATLANTIC RAIN FOREST, AND ITS PHYLOGENETIC PLACEMENT IN THE OCOTEA COMPLEX, Edinburgh Journal of Botany 79 (1875), pp. 1-23 : 7-15

Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes , sp. nov.

Similar to Ocotea daphnifolia (Meisn.) Mez but differs by its leaves, nine stamens with 2-celled anthers and pistil with well-developed ovule in apparently hermaphroditic flowers, and nine conspicuous staminodes in pistillate flowers, with vestigial 2-celled anthers, which evince its gynodioecious breeding system. Pistils from hermaphroditic individuals have no sign of a reduction, and their sizes are similar to those of pistillate flowers. –

Type: Brazil, São Paulo, Cunha, Parque Estadual da Serra do Mar – Núcleo Cunha–Indaiá , trilha do Ribeirão Bonito , 4 xi 2015 (fl ♀, fr), Moraes et al. 5032 (holotype HRCB [69304-A and 69304-B]; isotype SPSF [53621]). Figures 2 , 3A, B, F, G , 4 , 5 , 6A, B .

1

Small tree, 3–16 m tall. Terminal buds golden silky-tomentose. Young branchlets minutely pubescent, becoming glabrous in older parts, terete, bright, usually smooth, often with micro-exfoliating bark, longitudinally fissured, lenticels relatively sparse and conspicuous. Petioles 0.3–1 cm long, glabrescent, blackish, longitudinally wrinkled. Leaves alternate, evenly distributed along branchlets, elliptic to often obovate, 4–12 × 2–5.5 cm, chartaceous, apex shortly acuminate to caudate, base acute to cuneate, upper surface smooth, glabrous, reticulum lax, midvein prominent, secondary and tertiary veins prominulous to immersed, lighter than the lamina; lower surface somewhat crumpled in dried leaves, sometimes rusty, glabrescent, trichomes appressed, almost restricted to the midvein and domatia, midvein more prominent at the base, secondary veins prominent; venation eucamptodromousbrochidodromous, 4–6(–7) secondary veins on each side, with pit-shaped domatia, usually in the axils of several secondary veins, often with slit-like openings, not protruding to the other side of the leaf. Inflorescences subterminal and in the axils of foliage leaves, shorter than their subtending leaves, narrowly paniculate, (9)12–15(36)-flowered, peduncles

2–4 cm long. Hermaphroditic flowers: whitish in living material, 2.4–4.8 × 1.6–2.6 mm, tepals erect, subequal, outer whorl 1.4–2.1 × 1.1–1.4 mm, inner whorl 1.2–1.6 × 1–1.2 mm, broadly elliptic, apex acute, base truncate, densely glandular dotted, both surfaces with reddish, short, straight and appressed trichomes, margins with short and twisted trichomes; receptacle shallow, 0.4–0.6 mm depth, c. 0.7 mm in diameter, glabrous to subglabrous inside; stamens of first and second whorl subequal in size and shape, 0.7–1.3 mm long, filaments conspicuous, narrow, slightly shorter than anthers, trichomes straight only at the base, anthers ± trapeziform, gland dotted, connective subpapillate, mainly on the apex, the latter truncate to obtuse, locelli introrse, elliptic to oval, occupying almost the entire length of the anther; stamens of third whorl 0.8–1.2 mm long, filaments slightly narrower and slightly shorter than anthers, the latter subrectangular to roundish-trapeziform, densely gland-dotted, locelli latrorse-extrorse, narrowly elliptic, connective extended, apex obtuse to truncate, filaments with two basal glands, 0.3–0.5 × 0.3–0.5 mm, irregularly rectangular, shortly stalked; staminodes of fourth whorl lacking; pistil lageniform, often gland-dotted, 1.2–1.6 mm long, ovary globose, 0.7–0.9 × 0.5–0.9 mm, style slender and slightly shorter than the ovary, 0.4–0.7 mm long, stigma robust, discoid, at the same level of apex of anthers; ovule well developed, 0.4–0.6 × 0.3–0.4 mm. Pistillate flowers: externally similar to the hermaphrodites, white in living material, 3.9–4.6 × 2.1–2.8 mm, tepals erect, subequal, outer whorl 1.2–1.8 × 0.9–1.1 mm, inner whorl 1.1–1.8 × 1–1.1 mm, broadly elliptic, apex acute, base truncate, densely glandular-dotted, indument as on the hermaphrodites, receptacle shallow, 0.4–0.6 × 0.5–0.6 mm, glabrous to subglabrous inside; staminodes of first and second whorl subequal in size and shape, 0.4–0.8 mm long, filaments short, ± as long as anthers; staminodes of third whorl 0.7–0.9 mm long, filaments with two basal glands, 0.2–0.4 × 0.3–0.5 mm, shortly stalked; staminodes of fourth whorl lacking; pistil lageniform, often gland-dotted, 1.3–1.6 mm long, ovary ellipsoid, 0.8–1 × 0.5–0.7 mm, style stout, shorter than ovary, 0.4–0.7 mm long, stigma robust, triangular, positioned above the apex of the anthers; ovule well developed, 0.5–0.7 × 0.2–0.3 mm. Berry globose-ellipsoid, 1.2–2 × 0.8–1.3 cm, borne on an incrassate pedicel that merges into a patelliform cupule, often with remnants of tepals on a single margin; pedicels and cupules red in living material, lenticellate.

Distribution. Ocotea bilocellata is currently known from only five municipalities on the north coast of the Brazilian state of São Paulo (a region encompassing the Paraíba river valley and the north coast itself, from Caraguatatuba to Ubatuba), within the Serra do Mar State Park, close to the border with Rio de Janeiro ( Figure 7 ).

Habitat and ecology. It is found as medium-sized trees in the understorey of montane ombrophilous forests of the Atlantic Forest domain, at an altitudinal range of 800 to 1150 m a.s.l. Flowering from October to February, and fruiting in April, July, and September to November.

Etymology. The specific epithet refers to the 2-celled stamens.

Proposed IUCN conservation category. São Paulo is the most populous state in Brazil, with more than 46 million people inhabiting an area of 248.219. 481 km 2 ( IBGE: https://www. ibge.gov.br/en/cities-and-states/sp.html). The new species Ocotea bilocellata occurs in well-protected areas of the Serra do Mar State Park , in three of its Conservation Nuclei , namely Cunha– Indaiá , Picinguaba and Santa Virgínia, and also in the Boracéia Biological Station. Although its oldest collection dates back to 1941, there are 22 specimens from six locations (sensu IUCN). Ocotea bilocellata has an estimated EOO of 1640.24 km 2 and minimal AOO of 32 km 2. Both estimates fall within the limits for Endangered (EN) status under criteria B1 (EOO <5000 km 2) and B2 (<500 km 2), but the number of locations is greater than five and falls within the Vulnerable (VU) category. However, its population is not severely fragmented, and there is no continuing decline in the number of mature individuals, which mean that it does not qualify it for criterion B. Moreover, because there are no plausible threats and the species is within well-protected areas, it meets category Least Concern (LC) ( IUCN, 2012; IUCN Standards and Petitions Committee, 2022).

Additional specimens examined. São Paulo: Cunha, Parque Estadual da Serra do Mar , Núcleo Cunha– Indaiá, bacia D, 19 vii 1989 (fr), Baitello 307 ( SPSF) ; Cunha, ao longo do rio Paraibuna , 12 xii 1996 (fl ☿), Ferretti et al. 30 ( ESA, SPSF) ; Cunha, Parque Estadual da Serra do Mar , Núcleo Cunha– Indaiá , trilha da nascente do rio Bonito, casa de Pedra–Indaiá, 16 xii 1996 (fl ☿), Ferretti et al. 107 ( ESA, SPSF, UEC) ; ibid., trilha Barra do Rio , 30 i 2004 (fl), Ivanauskas 5078 ( SPSF) ; ibid., trilha do Ribeirão Bonito , 4 xi 2015 (fl ☿), Moraes et al. 5029 ( HRCB), (fl ☿), Moraes et al. 5030 ( HRCB, MBM, SPSF), (fl ☿), Moraes et al. 5031 ( HRCB, MBM, SPSF) ; ibid., 25 x 2017 (fr; from fl ♀), Moraes et al. 5468 ( HRCB, MBM, SPSF) ; ibid., trilha do Rio Paraibuna , 26 x 2017 (fr), Taxonomia de Campo – Grupo Verde: Marcusso et al. 155 ( HRCB, RB, SPF) ; ibid., trilha do Rio Bonito , 23°19′31′′S, 44°49′55′′W, 25 x 2017 (bud, fr), Taxonomia de Campo – Grupo Azul: Luize et al. 102 ( BHCB, HRCB, RB, SPF). Natividade da Serra, Parque Estadual da Serra do Mar, Núcleo Santa Virgínia, trilha do Corcovado , 14 ii 2005 (fl ☿), Ivanauskas et al. 5190 ( SPSF). Paraibuna , Parque Estadual da Serra do Mar , mata atlântica, 800–900 m, 28 vi 2011 (ster), Stefani Jr. et al. 199 ( UEC). Salesópolis, Boracéia, Estação Experimental do IAC , 16 i 1941 (fl ☿), Lima s.n. ( ESA [491], IAC [6113], SP [48716]) ; ibid., Biritiba-Mirim , Est. Biol. Boracéia, 29 ix 1983 (fr), Custódio Filho 1603 ( SP, SPSF) ; ibid., Biritiba-Mirim, Est. Biol. Boracéia, 950 m, 19 iv 1986 (fr), Custódio Filho 2582 ( SP, SPSF). Ubatuba, Parque Estadual da Serra do Mar, Núcleo Picinguaba, Morro do Corisco , 1000 m, 16 iv 1997 (fr), Pedroni et al. 643 ( UEC), (ster), Pedroni et al. 640 ( UEC), (ster), Pedroni et al. 641 ( UEC), (ster), Pedroni et al. 642 ( UEC), (ster), Pedroni et al. 647 ( UEC), (ster), Pedroni et al. 649 ( SPSF, UEC) .

Figures

Figure 2. Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov. A and B, Holotype (Moraes et al. 5032, HRCB [69304-A and 69304-B]). Photographs: P. L. R. de Moraes.

Figure 3. Leaf abaxial surface of Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov., and O. daphnifolia: A and B, O. bilocellata, domatia in the axils of secondary veins (voucher: Moraes et al. 5468, HRCB); C–E, O. daphnifolia, axillary and extra-axillary domatia (vouchers: C, Moraes et al. 3209, HRCB; D and E, Moraes et al. 3364, HRCB); F and G, O. bilocellata, domatia (voucher: Moraes et al. 5032, HRCB); H, O. daphnifolia, domatium (voucher: Harley et al. 50206, MO). Scale bars: F, 250 μm; G, 125 μm; H, 500 μm. Photographs: A, B, F, G and H, P. L. R. de Moraes; C–E, J. G. Rohwer.

Figure 4. Pistillate flower of Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov. A, Flower; B and C, dissected flowers with tepals removed, showing staminodes of whorls I, II and III and pistils conspicuously larger than them; D, outer tepal, adaxial view; E, staminode of whorl I; F, staminode of whorl II; G, pistil; H, ovule inside the ovary. Voucher: Moraes et al. 5032 (HRCB). Scale bars: A, 1 mm; B–H, 0.5 mm. Photographs: P. L. R. de Moraes.

Figure 5. Hermaphroditic flowers of Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov.: A, flower; B–E, dissected flowers with tepals partially removed, showing stamens of whorls I, II and III and pistils with stigmas at the same level of apex of anthers; D, detail of pistil insertion on the floral tube; E, detail of stamens conspicuously with pollen grains; F and G, tepal of whorl I (adaxial and abaxial views, respectively); H, stamens of whorl I (left) and II (right); I, stamen of whorl III, clearly with pollen grains; J, gland; K and L, pistil and respective ovule; M and N, pistil and respective ovule. Vouchers: A–C, H, K, L, Moraes et al. 5029 (HRCB); D, I, J, M and N, Moraes et al. 5030 (HRCB); E, Moraes et al. 5031 (HRCB); F and G, Ferretti et al. 30 (ESA). Scale bars: A and C, 1 mm; B, D, E–H, J, K and M, 0.5 mm; I, L and N, 0.2 mm. Photographs: P. L. R. de Moraes.

Figure 6. Fruits of Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov., and O. daphnifolia: A and B, O. bilocellata (voucher: Moraes et al. 5468, HRCB); C and D, O. daphnifolia (voucher: Moraes et al. 3209, HRCB); E, O. daphnifolia (voucher: Moraes et al. 3239, HRCB). Photographs: A and B, P. Moraes; C–E, J. G. Rohwer.

Figure 7. Geographical distribution of Ocotea bilocellata Baitello, D.B.O.S.Cardoso & P.L.R.Moraes, sp. nov., and O. daphnifolia in the state of São Paulo, Brazil. Blue circles: O. bilocellata; red circles: O. daphnifolia; shaded area: Atlantic rain forest domain.