Research Article |
Corresponding author: Daniel Santamaría-Aguilar ( daniel.santamaria366@gmail.com ) Academic editor: Thomas L.P. Couvreur
© 2021 Daniel Santamaría-Aguilar, Laura P. Lagomarsino.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Santamaría-Aguilar D, Lagomarsino LP (2021) Two new species of Otoba (Myristicaceae) from Colombia. PhytoKeys 178: 147-170. https://doi.org/10.3897/phytokeys.178.64564
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Otoba is the third largest genus of Myristicaceae in the Neotropics with 12 species, nine of them native to Colombia. Two new species from the department of Antioquia, O. scottmorii sp. nov. and O. squamosa sp. nov., are described and illustrated. Otoba scottmorii occurs in humid, lowland forests, while O. squamosa occurs in premontane forest. Previously, Otoba scottmorii was confused with O. acuminata (which here is considered restricted to Costa Rica and Panama), while O. squamosa was confused with O. gordoniifolia. The similarities and differences between these and other species are discussed.
Antioquia, Chocó, Magnoliales, Neotropics, Parque Nacional Natural Las Orquídeas, taxonomy
Otoba (A. DC.) H. Karst. is one of six genera of Myristicaceae native to the Neotropics. Morphologically, the genus is relatively easy to recognize. Its species have sessile or short-stalked (e.g. O. gordoniifolia (A. DC.) A.H. Gentry; W. Devia et al. 2291, MO) malpighiaceous foliar trichomes; conduplicate vernation; staminate flowers with filaments fused in an elongated column (except O. novogranatensis Moldenke in South America) with fused or free anthers; globose to ellipsoid green fruits; and seeds usually covered by a white aril and marked by the presence of a lateral or apical gibba (
Otoba has been comprehensively treated twice in the last century. In the first treatment (
Species of Otoba showing anther long, vestiture on the ovary, fruit size, pericarp thickness, and distributions.
Taxon | Anther long (mm) | Ovary vestiture | Fruit size (cm) | Pericarp thickness (mm) | Distribution |
---|---|---|---|---|---|
O. acuminata (Standl.) A.H. |
0.45–0.8 | Pubescent | 1.9–2.5 (−2.8) × 1.7–2.2 (−2.4) | 1–2.1 | Costa Rica, Panama |
O. cyclobasis T.S. Jaram. & Balslev (2001: 563) ‡ | ca. 0.3 | Glabrous | 1.5–2.5 × 1.5–2 | 1–2 | Ecuador |
O. glycycarpa (Ducke) W.A. Rodrigues & T.S. Jaram. (2001: 446) ‡ | 0.5–0.6 | Pubescent | 3.4 × 3.4 | 5–8 | Colombia, Ecuador, Peru, Bolivia, Brazil |
O. gordoniifolia (DC.) A.H. |
0.7–1.5 | Pubescent | (3–) 5 × 4 | 3–5 | Colombia, Ecuador |
O. gracilipes (A.C. Sm.) A.H. |
0.6–0.8 | Glabrous | 3–3.5 × 2.5 | 3–4 | Colombia, Ecuador |
O. latialata (Pittier) A.H. |
0.2–0.3 (in Panama; pers. obs.) | Glabrous | 2.2–2.8 × 1.8–2.3 (in Panama pers. obs.) | 2‡ (ca. 1 in Panama pers. obs.) | Panama, Colombia |
O. lehmannii (A.C. Sm.) A.H. |
0.7–1.3 | Glabrous | 4–6 × 4–4.5 | 3–5 | Colombia |
O. novogranatensis |
1–1.5‡ (0.6–1.1 in Central America; pers. obs.) | Pubescent in South America‡; (always glabrous in Central America; pers. obs.) | 3–5.5 × 2.5–4‡ (2.3–3.5 × 2–2.5 in Central America; pers. obs.) | 2–5‡ ([1.4–] 2.1–2.3 in Central America; pers. obs.) | Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Venezuela |
O. parvifolia (Markgr.) A.H. |
0.3–0.7 | Glabrous | 2.3–3 × 1.5–2 | 1–2 | Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela |
O. squamosa | 0.5–0.7 | Glabrous | (2.5–) 3.4–3.6 × (1.9–) 2.8–3 | 2.6–3 | Colombia |
O. scottmorii | 0.2–0.4 | Unknown (speculated is glabrous) |
1.8–2.1 × 1.5–1.7 | 0.68–0.89 | Colombia |
O. vespertilio D. Santam. & J.E. Jiménez (2019: 371) | ca. 0.2–0.3 | Glabrous | 2.2–2.7 × 1.6–1.7 | 1.3–1.8 | Costa Rica, Panama |
Two new species of Otoba are here described. Both occur in the Antioquia department of Colombia in premontane (1330–1450 m elevation) and humid (20–80 and 410–730 m) forests. Currently, there are seven species registered in this department: O. acuminata, O. gordoniifolia, O. gracilipes, O. latialata, O. lehmannii, O. novogranatensis, and O. parvifolia (
Otoba often is abundant within their habitats (
Despite its abundance, our knowledge of floral biology of Otoba is very limited. The small flowers, which usually have a greenish-yellow or yellow perianth, are reported to have unpleasant odors (e.g., H. van der Werff 10764, MO), sometimes reported to smell of human semen (M.H. Grayum et al. 5502, MO). Small coleopterans are known pollen vectors for O. novogranatensis (
More is known about dispersal in Otoba. Fruits are generally aromatic and globose, with a green, often shiny pericarp and a seed covered by a white or, occasionally, reddish aril. These arils can be very sweet (
Examples of natural history of Otoba in Central America A, B Otoba novogranatensis, fruits (A) and seeds (B) accumulating on the ground in Costa Rica C, D Fruits of O. acuminata, inset seed covered by aril (C), which are eaten by Metachirus nudicaudatus (D) in Panama E, F Dyscophellus phraxanor (Hesperiidae), a herbivore of O. novogranatensis, as a larvae (E) and adult (F). Photos by Reinaldo Aguilar (A), Orlando Vargas (B) from https://sura.ots.ac.cr/florula4/index.php, Ángel Sosa-Bartuano (C, D), Daniel H. Janzen (E, F) from http://janzen.sas.upenn.edu/caterpillars/database.lasso.
Leaves of Otoba are also an important food source for animals. This includes mammals, such as the Caquetá titi (Plecturocebus caquetensis), which eats the young leaves of Otoba parvifolia (
In the course of herbarium investigations, we identified two new taxonomic species that we describe here. Descriptions are based on herbarium specimens observed at the following herbaria: CR (including ex INB), LSCR, LSU, MO, NO and USJ, as well as loaned material from: GH, NY, and US (acronyms follow
In the nomenclatural section for each new species, we cite both accession numbers and barcodes when present. Barcodes are included in square brackets and follow the format of a series of numbers preceded by the herbarium acronym (e.g., [04206141]); all other numbers correspond to accession numbers.
When the coordinates and/or elevation were not included on the herbarium label but were present in the TROPICOS database (
The preliminary conservation status of each new species was assessed using quantitative criteria recommended by the IUCN Red List (
Colombia. Antioquia: [Municipio] Mpio. Segovia, 24.5 km N of Remedios (17 km N of La Cruzada) on road to Zaragoza, hills side forest above río Poconé, 07°12'N, 074°48'W, [not elev.], 20 Jul. 1987 (♂ fl), W.W. Thomas & C.J. Castaño 5501 (holotype: NY! [04206141]; isotypes: INPA-159329 [digital image!], UPCB-35245 [n.v.]).
Otoba scottmorii is similar to O. cyclobasis from Ecuador. However, it differs in having smaller leaves (5.5–8.5 [–11.2] vs. 9–14 [–18] cm long) with an attenuate to acuminate apex (vs. cuspidate), inconspicuous marginal and secondary veins (vs. conspicuous), fewer lateral veins (4–7 vs. 14–17), straight axes of staminate inflorescences (vs. zig zag), a longer filament column ([1.3–] 1.5–1.6 vs. 1 mm long), larger fruits (1.8–2.1 vs. 1.5–2.5 mm long) with thinner pericarp (0.68–0.89 vs. 1–2 mm thick), and seeds that are gibbose at the apex (vs. near-basal and lateral gibba).
Tree (2–) 15–20 m tall × 45 cm diam., external and internal bark not described. Exudate once described as red in the bark, transparent and without specifying from where, or without exudate. Twigs 0.88–0.1 cm thick, terete to slightly flattened laterally, the external bark brown to grayish, with malpighiaceous trichomes that are 0.2–0.3 mm long, brown to ferruginous or sometimes glabrescent. Young foliar bud 0.6–1.2 cm long, densely pubescent. Leaves: petiole 0.5–1.1 × 0.055–0.1 cm, canaliculate, not winged; lamina 5.5–8.5 (–11.2) × 1.7–2.5 (–3.7) cm, elliptic; adaxial side glabrous, drying blackish, sometimes brown in young leaves, the surface muricate; abaxial side drying brown to whitish grayish, the surface wrinkled to muricate, sparsely pubescent to glabrescent, with malpighiaceous trichomes that are 0.2–0.4 mm long, sessile, and ferruginous, and squamate trichomes that are ca. 0.08 mm diam. with the central portion dark and sides lighter (sometimes these sides appearing absent), with whitish crystals; vernation line absent; midvein flat to very slightly ribbed on adaxial side, the same color as the surface, abaxially 0.2–0.4 mm wide, slightly raised, darker than the surface; secondary veins brochidodromous, the loops 0.1–0.3 cm from the margin (sometimes the loops not visible), lateral veins 4–7 per side, 3–5 veins per 3 cm, on adaxial side flat and visible, on abaxial side flat and not very conspicuous, arcuate distally, the marginal vein not visible on adaxial side, slightly visible on abaxial side; tertiary veins indistinct; base attenuate, not revolute; margin entire, not revolute; apex attenuate to acuminate, the acumen 0.6–1 (–2) cm long. Staminate inflorescence: axillary (only very young twigs) or supraaxillary, with 1–2 (–3) main axes, spiciform, these axes 1.5–4.3 cm × 0.41–0.1 mm, pubescent, the trichomes ferruginous to coppery, each axis compound with 2–3 (–6) fascicles of flowers, each fascicle with 1–5 (–8) flowers, alternate; bracts not seen; pedicel 3–5 mm long, pubescent; bracteoles absent. Staminate flowers: flower bud 1–2.5 × 0.6–0.1 mm, elliptic to lanceolate; perianth 2–2.5 mm long, yellow or orange (in fresh material), sub-membranaceous, connate by 0.5–0.7 (–1) mm; lobes 3 (4), 1.5–1.8 × 0.8–1.1 mm, without resinous punctuations or lines, pubescent outside, the trichomes ferruginous to coppery, inside glabrous, the apex acute to obtuse, without inflexed-apiculo, the edges flat or slightly turned inwards distally, without a swollen lobed ring; filament column (1.3–) 1.5–1.6 mm long, bowling pin-shaped, fleshy, connate, glabrous; anthers 3, 0.2–0.4 mm long, free, lanceolate to oblong, apex slightly incurved. Pistillate inflorescence and flowers: not seen. Infructescence 2–4 cm long, with a solitary fruit; pedicel 1.2–1.6 cm long. Fruits 1.8–2.1 × 1.5–1.7 cm, green when fresh, globose, surface glabrous, colliculate-rugose, sometimes with whitish lenticels (J. Brand 682), the line of dehiscence smooth, the base obtuse, apex apiculate, the acumen 0.1–0.3 cm long; pericarp 0.68–0.89 mm thick; seed ca. 1.6 × 1.5 cm, similar in shape to the fruit, brownish (in dry material), gibbose at the apex (ca. 0.3 mm wide), the testa ca. 0.4 mm thick; aril described once as red (E. Renteria 4680), brownish to yellowish translucent in dry material, dry and membranous in texture.
Otoba scottmorii A staminate flowering branch, with detail of leaf surface (detail below left) B leaf base, petiole and basal portion of inflorescence C detail of staminate inflorescence D flower bud E staminate flowers, entire (above right) and with one perianth lobe removed, showing the androecium (below center) and perianth lobe (above left) F androecium G stem with infructescence, including an immature, opening fruit H view of dehisced fruit (left, below G) showing the seed and aril (right) Otoba cyclobasis I staminate flowering branch showing leaves with brochidodromous venation, including detail of marginal vein J staminate flower with one perianth lobe removed, showing the androecium and perianth lobe (above left) K androecium. Illustration by Bobbi Angell, W.W. Thomas & C.J. Castaño 5501 (NY) (A–F), J. Brand & M. Narváez 682 (MO) (G, H), E. Narvaéz et al. 1072 (MO-2468293) (I–K).
Otoba scottmorii is recognized by a combination of leaf, inflorescence, and fruit traits. Its small leaf blades have long apices and thin petioles, and lack vernation lines on the abaxial surface. The staminate inflorescences are delicate with flowers on relatively thin, long pedicels; these flowers have sub-membranaceous perianth and lack a swollen-lobed ring, a bowling pin-shaped filament column, and anthers that are lanceolate to oblong. Finally, fruits are small with thin pericarp and membranous aril.
Is a pleasure to name this species after Dr. Scott A. Mori (1941–2020), a wonderful person and skilled botanist; a dedicated explorer of Central and South America humid forests (where this species occurs), especially in the Guianas and the Amazon basin; and an authority on Neotropical Lecythidaceae. His taxonomic and ecological publications gave great inspiration to the first author, as did Dr. Mori’s personal support. For an account of Dr. Mori’s legacy, see
Cuángare otobo (
Otoba scottmorii is known only from the humid forests in the Department of Antioquia in northwestern Colombia in the Municipios of Mutatá, Segovia, Tarazá, Turbo, and Valdivia (Fig.
Herbarium specimens of Otoba scottmorii have been collected with staminate flowers in May, July, and November, and with fruits in November. Pistillate flowers have yet to be observed.
Otoba scottmorii is Endangered following IUCN criteria B2a. Justifying this status, it is known from five localities, has an EOO of 5,341 km2, and an AOO of 20 km2. The regions where it occurs are threatened by high levels of deforestation (
Otoba scottmorii is similar to O. acuminata and O. vespertilio D. Santam. & J.E. Jiménez, from Costa Rica and Panama, and O. cyclobasis T.S. Jaram. & Balslev from Ecuador. They all have relatively small leaf blades of similar shape and thin petioles, and usually lack vernation lines (Figs
Taxon | O. scottmorii | O. acuminata | O. cyclobasis | O. vespertilio |
---|---|---|---|---|
Petiole length (cm) | 0.5–1.1 | (1.1–) 1.8–2.6 | 1–1.5‡ | 0.8–2 |
Leaf size (cm) | 5.5–8.5 (–11.2) × 1.7–2.5 (–3.7) (Fig. |
(6.5–) 8.5–15.5 × (2.7–) 4.4–6.1 (Fig. |
9–14 (–18) × 3–5‡ (Fig. |
5.3–12.5 × 2–3.5 (−4.8) (Fig. |
Lateral vein number | 4–7 | 8–12 | 14–17‡ | 6–8 |
Staminate flower pedicel length (mm) | 3–5 | 5–8 | 4–5‡ | 2–3.6 |
Staminate perianth length (mm) | 2–2.5 | 4.5–5.5 | ca. 2.5‡ | 2–3 |
Perianth lobe width (mm) | 0.8–1.1 | 1.5–2 | ca. 1 § | 0.5–1 |
Ring | Absent | Absent | Present‡ | Absent |
Filament column length (mm) and shape | (1.3–) 1.5–1.6, bowling pin | 2–2.7, cylindrical, ovoid, to pyriform | 1, cylindrical | 0.6–1.5, cylindrical |
Anther length (mm) and shape | 0.2–0.4 lanceolate to oblong | 0.45–0.8, oblong | ca. 0.3, globose‡ | ca. 0.2–0.3, globose to subglobose |
Ovary vestiture | Unknown | Pubescent | Glabrous‡ | Glabrous |
Fruit length and width (cm) | 1.8–2.1 × 1.5–1.7 (Fig. |
1.9–2.5 (−2.8) × 1.7–2.2 (−2.4) (Fig. |
1.5–2.5 × 1.5–2‡ (Fig. |
2.2–2.7 × 1.6–1.7 (Fig. |
Pericarp thickness (mm) | 0.68–0.89 | 1–2.1 | 1–2‡ | 1.3–1.8 |
Seed length and width (cm) | 1.6 × 1.5 | ca. 2.2 × 1.9 | 1–2 × 1–1.5‡ | 1.8–1.9 × ca. 1.7 cm |
Distribution and elevation | Colombia, 20–80 m and 410–730 m | Costa Rica, 100–900 m; Panama, 500–1000 | Ecuador, 150–300 m‡ | Costa Rica 0–200 m, Panama 100–800 m |
Comparison of leaf blades and fruits of O. acuminata (A L. Poveda et al. 3793, MO A1 from H.W. Churchill & G.C. de Nevers 4976 MO; immature fruit) O. scottmorii (B R. Callejas et al. 3440 MO B1–B2 from R. Callejas et al. 5789, MO) O. cyclobasis (C E. Narvaéz et al. 1072, MO C1–C2 from W.A. Palacios & M. Tirado 11318 MO) and O. vespertilio (D G. McPherson 13597, MO, juvenile leaf, D1 G. McPherson 12543, MO). Scale bars: leaves: 2 cm; fruits: 0.75 cm.
As mentioned in the introduction, Otoba scottmorii was confused with the Mesoamerican species O. acuminata, which we now consider to be endemic to the Caribbean slope of Costa Rica and Panama. The specimens referred to as O. acuminata from Colombia in recent floras or checklists (e.g.,
The type specimen of Otoba scottmorii and other collections here mentioned have been previously identified in herbaria and cited in literature as O. acuminata and O. gracilipes, and duplicates may have been distributed under these names.
Colombia. Antioquia: Mutatá, Sitio Rio Surambay, 12 km N de Mutata, 07°20'N, 076°30'W, 30–80 m, 21 Nov. 1987 (♂ fl), R. Callejas et al. 5752 (INPA digital image, MO, NY); ibid, 21 Nov. 1987 (fr), R. Callejas et al. 5789 (INPA digital image, MO, NY); Turbo, Carretera Tapón del Darién, Sector Río León-Lomas Aisladas, km 37, [07°39'11"N, 076°58'02"W], 20 m, 29 Nov. 1993 (fr), J. Brand & M. Narváez 682 (COL [n.v.], JAUM digital image, MO); río Cianurá, Paso de la Reina, 730 m, 13 Mar 1986 (fr), E. Renteria et al. 4680 (JAUM digital image); Tarazá, Corregimiento El 12, 210 kms, NE de Medellín, vía El 12-Barroblanco, km 3, 07°30'N, 075°20'W, [450 m], 09 Nov. 1987 (♂ fl), R. Callejas et al. 5509 (INPA digital image, NY); Valdivia, corregimiento Puerto Valdivia, km 5 de Puerto Valdivia hacia “El 12”, colecciones a lo largo del Río Pescado, 07°20'N, 075°20'W, 410 m, 14 May. 1987 (♂ fl), R. Callejas et al. 3440 (MO-2 sheets, NY).
Colombia. Antioquia: Urrao, Vereda Calles, Parque Nacional Natural “Las Orquídeas”, margen derecha Quebrada La Honda, 06°32'N, 076°19'W, 1330–1400 m, 08 May 1993 (fr), Á. Cogollo, R. Carmona, E. Álvarez 6197 (holotype: MO-05011088! [1500541]; isotypes: n.v.).
Otoba squamosa is similar to O. gordoniifolia from Colombia and Ecuador, and both species grow in montane forest in the Andes. However, it differs in its leaves with shorter petioles (1.7–2.7 [–3.8] vs. [3–] 5–7 cm long) and smaller lamina (6.7–14.5 vs. [13–] 24–34 cm long), staminate flowers with a perianth with a swollen ring (vs. without) and smaller anthers (0.5–0.7 vs. 0.7–1.5 mm long), pistillate flowers with a glabrous ovary (vs. pubescent), and fruits with thin pericarp (2.6–3 vs. 3–5 mm thick).
Otoba squamosa A staminate flowering branch, with detail of malpighiaceous trichomes on abaxial leaf surface (detail below right) B detail of staminate inflorescence C staminate flower, with detail of trichomes D open staminate flower E longitudinal section of staminate flower and androecium F pistillate flowering branch G portion of pistillate inflorescence, showing the perianth (left) and ovary after the perianth has fallen H ovary I closed and open fruit (left), showing the pericarp and aril. Illustration by Bobbi Angell. Á. Cogollo et al. 6279 (MO) (A–D), J. Pipoly et al. 17019 (MO) (E), J. Pipoly et al. 16798 (G), Á. Cogollo et al. 6197 (MO) (F, H, E).
Tree 8–18 (–24) m tall × 11.9–34.7 cm diam., external and internal bark not described. Exudate hyaline, oxidizing reddish, only reported from in flowers and fruits. Twigs 0.13–0.27 cm thick, terete to slightly flattened laterally, the external bark brown to blackish, with malpighiaceous trichomes 0.2–0.6 mm long, brown to ferruginous, the indument denser in young parts. Young foliar bud 1.6–4.3 (–5.8) cm long, densely pubescent. Leaves: petiole 1.7–2.7 (–3.8) × 0.1–0.2 (–0.25) cm, canaliculate, very short-winged; lamina 6.7–14.5 × 3.4–6 (–8) cm, elliptic, rarely widely elliptic; adaxial side glabrous, usually drying dark brown to blackish, the surface muricate-reticulate; abaxial side usually drying pale to dark brown, the surface muricate, sparsely pubescent, with malpighiaceous trichomes 0.3–0.6 mm long, sessile, ferruginous, and scale-like trichomes ca. 0.1 diam. with the central part dark, contrasting with the lighter sides, crystals generally absent and if present very few; vernation line imprints 2 parallel lines, 0.7–1.5 (–1.8) cm from the margin, the panel area 1.8–2.5 (–3.4) cm wide (in the central portion), the same color as the surface; midvein flat on adaxial side, the same color as the surface or blackish, abaxially 0.5–0.9 mm wide, raised, a little darker than the surface; secondary veins brochidodromous, the loops 0.2–0.3 cm from the margin, lateral veins 13–17 per side, (3–) 4–6 veins per 3 cm, on adaxial side slightly caniculate, on abaxial side flat to slightly raised, not very conspicuous, arcuate distally, the marginal vein not visible on adaxial side, slightly visible on abaxial side; tertiary veins indistinct; base acute to cuneate, not revolute; margin entire, not revolute; apex acute, the acumen 0.3–0.9 cm long. Staminate inflorescence: axillary and/or ramiflorus, with 1–2 main axes, spiciform, these axes 2–7 cm × 0.6–1.2 mm, pubescent, the trichomes ferruginous, each axis compound with 2–5 fascicles of flowers, each fascicle with 3–6 flowers, alternate; bracts ca. 1.5–1.6 × 1 mm (observed in very young inflorescences), densely pubescent outside, the trichomes ferruginous; pedicel 1.3–4 mm long, pubescent; bracteoles absent. Staminate flowers: flower bud 2–3 × 1–1.5 mm; perianth 3.5–4.7 mm long, yellowish to yellowish-green (in fresh material), fleshy (hardening near the base by the ring), connate by 1–1.7 mm; lobes 3 (4), (2–) 2.5–3.4 × (0.9–) 1.4–2 mm, without resinous punctuations or lines, pubescent outside, the trichomes ferruginous, inside glabrous, smooth to lightly spongy, the apex in some flowers with a minutely inflexed-apiculate, the margin edges slightly turned inwards, slightly wavy; ring present, 0.1–0.4 mm wide, lobed, smooth, or sometimes spongy; filament column 1.5–2.1 mm long, usually cylindrical, slightly narrow towards the apex, fleshy, glabrous; anthers 3 (4), 0.5–0.7 mm long, free, lanceolate to oblong, apex slightly incurved. Pistillate inflorescence: axillary, 1.4–3.2 cm long, pubescent, the trichomes ferruginous, each axis compound with 1–2 fascicles of flowers, each fascicle with 2–3 flowers, alternate; bracts not seen; pedicel 3–4 mm long, pubescent; bracteoles absent. Pistillate flowers: flower bud 3–4 × ca. 2 mm; perianth 3–5 mm long, the color and texture as in the staminate flower, connate by 1–1.5 mm; lobes 3 (4), 2–3.5 × 2–2.5 mm, without resinous punctuations or lines, pubescent outside, the trichomes ferruginous, inside glabrous; ring present; gynoecium 2–3 × 1.5–2.3 mm, glabrous, ovary sessile to short-stalked, ca. 0.6 mm long; stigma 2-lipped, subsessile; stigmatic lips ca. 0.6 mm long. Infructescence probably with one fruit (fruits separated from the axis in all specimens observed); pedicel ca. 1–1.1 cm long. Fruits (2.5–) 3.4–3.6 × (1.9–) 2.8–3 cm, green, globose, the surface glabrous, rugose, sometimes with whitish to brownish lenticels, the line of dehiscence smooth, the base obtuse, and sometimes getting narrower towards the pedicel, apex obtuse or acute, the acumen 0.8–1 cm long; pericarp 2.6–3 mm thick; seed (2.1–) 2.5–2.9 × (1.7–) 2.3–2.4 cm, similar in shape to the fruit, whitish or brown (in dry material), gibbose at the apex or nearly so, the testa 0.4–0.6 mm thick; aril described once as white (D. Sánchez et al. 1529), brownish to white-yellowish in dry material, waxy to dry in texture.
Otoba squamosa is recognized by a variety of leaf traits, including: squamate or scale-like indument mixed with malpighiaceous trichomes on the abaxial surface, lateral veins that are more or less conspicuous, forming a marginal vein, and vernation lines that parallel the midvein. Additionally, the staminate flowers have a perianth with swollen-lobed ring in the inner surface, a typically cylindrical filament column with lanceolate to oblong anthers and pistillate flowers that have a glabrous gynoecium. Finally, the fruits are relatively large, with thick pericarp.
The specific epithet refers to the squamate or scale-like indument that is present with more typical malpighiaceous trichomes on the abaxial surface of the leaf blades. The squamate or scale-like indument is not unique to this species; it is also present in most specimens of O. acuminata, O. glycycarpa, O. scottmorii, and O. vespertilio; also
None recorded.
Otoba squamosa is known from the Cordillera Occidental of Colombia, specifically in the municipalities of Frontino and Urrao in the Department of Antioquia (Fig.
Fertile herbarium specimens of Otoba squamosa have been collected with staminate flowers in May and December, with pistillate flowers in May, September, and December, and with fruits in March, May, September, October, and December.
Otoba squamosa is Endangered following IUCN criteria B2a. Justifying this status, it is known from only two localities and has an AOO of 4 km2; there are too few verified localities to reasonably estimate the EOO. The Andean forests of the Antioquia Department of Colombia where it occurs are particularly at risk for deforestation (
The collections now identified as Otoba squamosa were previously included under the concept of O. gordoniifolia (Fig.
Otoba squamosa | O. gordoniifolia ‡ | |
---|---|---|
Petiole length (cm) | 1.7–2.7 (–3.8) × 0.1–0.2 (–0.25) | (3–) 5–7 × 0.3–0.4 |
Leaf size (cm) | 6.7–14.5 × 3.4–6 (–8) | (13–) 24–34 × (6–) 8–12 |
Staminate perianth length (mm) | 3.5–4.7 | (2–) 4–6 |
Ring | Present | Absent |
Anthers length (mm) | 0.5–0.7 | 0.7–1.5 |
Ovary vestiture | Glabrous | Pubescent |
Fruit size (cm) | (2.5–) 3.4–3.6 × (1.9–) 2.8–3 | (3–) 5 × 4 |
Pericarp thickness (mm) | 2.6–3 | 3–5 |
Within Otoba, the swollen-lobed ring in the staminate perianth is shared between O. cyclobasis (and even gives this species its specific epithet;
In the “Key to the species (pistillate or fruiting plants)” from
The specimen [J.] Pipoly 17003 (MO nv), included in
Specimens of O. squamosa, including the type, have been previously identified in herbaria as O. gordoniifolia, and duplicates may have been distributed under these names.
The following sterile specimens from MO were collected in the same locality as fertile material or nearby (Parque Nacional Natural “Las Orquídeas”, Sector Calles); because they were not fertile, they were not used in the description presented above, though we believe that they correspond to this new species: Á. Cogollo et al. 6069, 6132, 6169, 6298, 6344, 6642, 6644, 6657, 6683, 6684, 6869, 6926, 6929, 7109, 7180, 7221, 7237, 7238, 7259, 7262, 7268, 7269; and J. Pipoly et al. 16663, 17022, 17033, 17042, and 17076.
Colombia. Antioquia: Frontino, Corregimiento Nutibara, cuenca alta del Río Cuevas, bosque al lado carretera a La Blanquita, 1100 m, 21 Sep. 1987 (♀ fl & fr), D. Sánchez et al. 1529 (MO); Urrao, Parque Nacional Natural “Las Orquídeas”, Sector Calles, margen derecha del Río Calles, 06°32'N, 076°19'W, 1420 m, 25 Mar. 1988 (fr), Á. Cogollo et al. 2573 (MO); Vereda Calles, Parque Nacional Natural “Las Orquídeas”, margen derecha Quebrada La Honda, 06°32'N, 076°19'W, 1330–1400 m, 03 May 1993 (♂ fl), Á. Cogollo et al. 6074 (MO); ibid, 08 May 1993 (♂ fl), Á. Cogollo et al. 6190 (K-2 sheets [n.v.], MO); ibid, 08 May 1993 (♀ fl), Á. Cogollo et al. 6198 (MO); ibid, 11 May 1993 (♂ fl), Á. Cogollo et al. 6279 (MO, NY); ibid, 11 May 1993 (♂ fl), Á. Cogollo et al. 6300 (K n.v., MO); Vereda Calles, Parque Nacional Natural “Las Orquídeas”, margen derecha del Río Calles, en el filo NW de la Cabaña de Calles, 06°32'N, 076°19'W, 1450 m, 15 Oct. 1993 (fr), Á. Cogollo et al. 6918 (MO); ibid, 18 Oct. 1993 (fr), Á. Cogollo et al. 7107 (MO); ibid, 08 Dec. 1993 (fr), Á. Cogollo et al. 7959 (GH digital image, K [n.v.], MO, NY); Las Orquídeas, Vereda Calles, Parque Nacional Natural Las Orquídeas, Quebrada Honda, 06°29'N, 076°14'W, 1300 m, 08 Dec. 1992 (fr), J. Pipoly et al. 16745 (MO); ibid, 1330 m, 08 Dec. 1992 (♀ fl), J. Pipoly et al. 16798 (MO); ibid, 10 Dec. 1992 (♂ fl), J. Pipoly et al. 16902 (MO); ibid, 1300 m, 11 Dec. 1992 (♂ fl), J. Pipoly et al. 17019 (K [n.v.], MO); Parque Nacional Natural Las Orquídeas, Vereda Calles, margen derecha del Río Calles, 06°32'N, 076°19'W, 1350–1450 m, 05 Dec. 1993 (fl bud), J. Pipoly et al. 17721 (MO).
The identification key below is modified from
1 | Abaxial leaf blades with vernation lines (e.g. 7A, 8B). | |
2 | Leaf blades 6.7–14.5 cm long, abaxial surface with sparse, scale-like and trichomes; perianth of staminate flowers with swollen-lobed ring; staminate flowers with anthers 0.5–0.7 mm long; fruits with pericarp 2.6–3 mm thick | O. squamosa |
2' | Leaf blades (13–) 24–43 cm long, abaxial surface with dense, scale-like and trichomes; perianth of staminate flowers without swollen-lobed ring; staminate flowers with anthers 0.7–1.5 mm; fruits with pericarp 3–5 mm thick. | |
3 | Pistillate flowers with pubescent ovaries | O. gordoniifolia |
3' | Pistillate flowers with glabrous ovaries | O. lehmannii |
1' | Abaxial leaf blades without vernation lines. | |
4 | Leaf blades with petioles less than 0.2 cm thick. | |
5 | Staminate inflorescence 6–7 cm long, flower bud 1.2–1.6 mm wide|, perianth ca. 3 mm long; filament column divided distally; anthers ca. 0.5 mm long | O. gracilipes¶ |
5' | Staminate inflorescence 1.5–4.3 cm long, flower bud 0.6–0.1 mm wide, perianth 2–2.5 mm long; filament column completely connate; anthers 0.2–0.4 mm long | O. scottmorii |
4' | Leaf blades with petioles up to (0.1–) 0.2 cm thick. | |
6 | Staminate flowers with filament column distinct almost to the base | O. novogranatensis |
6' | Staminate flowers with filament column connate, if distinct never to the base | |
7 | Leaf blades with petioles conspicuously winged; staminate inflorescence with the first fascicles of flowers 2.2–4 (–8) cm# from the base, fascicles of flowers arranged more or less in a zig-zag, fascicles well separated (1–1.5 cm apart#), pedicel filiform, flowers with membranaceous perianth | O. latialata |
7 | Leaf blades with petioles usually not winged, if winged, the wing very narrow; staminate inflorescence with the first fascicles of flowers 0.5–2 cm# from the base, fascicles of flowers arranged in a straight line, each fascicle in close proximity (0.5–1 cm apart#), pedicel stout, flowers with fleshy perianth. | |
8 | Abaxial leaf surface densely covered with scale-like trichomes; pistillate flowers with pubescent gynoecium; fruits with pericarp 5–8 mm thick | O. glycycarpa |
8' | Abaxial leaf surface sparsely covered with scale-like trichomes; pistillate flowers with glabrous gynoecium; fruits with pericarp 1–2 mm thick | O. parvifolia |
We would like to express our gratitude to the curators of the following institutions for allowing access to, and use of, their physical collections, loan material, and/or making digital images available: CR, EAP, F, HUH, INPA, JAUM, K, LSU, MEXU, MO, NO, NY, P, PMA, SCZ, US and USJ. Also, we are very grateful to R. Aguilar, I. Chacón, R. Gelis, W. Hallwachs, D.H. Janzen, J.E. Jiménez, R. Pérez, Á. Sosa-Bartuano, and O. Vargas for providing images; A. Arbelaez and Á. Idárraga-Piedrahíta, F. Barrie, and B.E. Hammel supplied images of collections deposited at JAUM, F and CR respectively, while A.K. Monro confirmed the presence of some specimens at Kew; as well to R. Aguilar, J.M. Chaves, I. Chacón, F. Oviedo, and J.A. Rosales for field observations. As always, we are in awe of B. Angell’s beautiful and detailed illustrations, which strikingly highlight even inconspicuous structures. To S. Oliveira, anonymous reviewer and the subject editor T.L.P. Couvreur for the comments and suggestions they provided which greatly improved this manuscript. Daniel Santamaría-Aguilar is very grateful to A. Arbelaez, G. Davidse, M.H. Grayum, M. Merello, O.M. Montiel, and J. Solomon for their help provided during herbarium visits at MO; to MO for providing substantial material as loans and donations; and to J. Kluse (LSU) for providing the support and resources necessary for this endeavor. This contribution would not have been possible without support from the Department of Biological Sciences and the Shirley C. Tucker Herbarium at Louisiana State University.