Plukenetia chocoensis, Cardinal-McTeague & Gillespie, 2020

11. Plukenetia chocoensis L.J.Gillespie & Card. -McTeag. sp. nov. TYPE: COLOMBIA. Valle del Cauca: Buenaventura, Bajo Colima, road to Junchaco Palmeras, [3.91666°, -77.20000°], 100 m, 10 July 1984, A.H. Gentry, M. Monsalve B. & D.A. Wolfe 47799 (holotype: MO! [accession 3265264, barcode

MO-1381711 ]; isotypes: COL [accession 540920, barcode

COL000397503; image!], F [accession 1938863, barcode

V0216686F; image!]).

Similar to P. penninervia Müll.Arg. but differs in its elongate narrowly oblong-elliptic adaxial basilaminar extrafloral nectaries, presence of abaxial laminar extrafloral nectaries, and longer inflorescences.

Monoecious vines to slender lianas; stems erect or twining; old stems to 1.2 1 cm in diam, bark dark purplish brown, coarsely striate; older flowering stems medium brown when dry, to; 5 mm in diam, faintly striate, puberulous to glabrate; younger stems faintly striate to striate, puberulous or pubescent. Leaves alternate, evergreen; stipules deltoid, 0.5–0.7 mm long, deciduous; petioles generally of uniform thickness and colour, 0.7–1.6 cm long, puberulous; blades simple, elliptic, oblong-elliptic, ovate-elliptic or obovate-elliptic (sometimes narrowly so), 8–16 3 3–6.5 cm, thick-chartaceous, drying shiny olive-green above, dull olive-green or pale olive-green with orange-brown veins below, both surfaces glabrous or glabrate with major veins glabrate to sparsely puberulous, base acute to obtuse, usually shortly attenuate around basilaminar extrafloral nectaries, margins serrulate with small glandular teeth, apex acuminate to cuspidate, tip 0.6–1.3 cm long; venation pinnate, primary vein 1, secondary veins 5–7 on each side of the midrib, brochidodromous, tertiary veins percurrent (rarely reticulate); stipels and glandular knobs absent at petiole apex; adaxial basilaminar extrafloral nectaries 2, narrowly oblong or narrowly oblong-elliptic, sometimes irregular in shape, 2–4 3 0.5–1(–1.6) mm; abaxial laminar extrafloral nectaries 1–8/side, 0.3–0.7 (–0.9) mm in diam, 0.4–2 mm from margin on distal 5/6–1/10 of blade, adaxial laminar nectaries absent. Inflorescences axillary, bisexual racemose thyrses, 0.5–2 cm long at the pistillate flower stage, elongating up to 6 cm in the later staminate stage, 1(2)/ axil; peduncle 0.1–0.2 mm long, elongating to 10 mm in later staminate and fruit stage; axes moderately or sometimes densely puberulous throughout; staminate bracts ovate-deltoid, 0.5–0.9 mm long, puberulous; pistillate bracts deltoid or ovate-deltoid, 0.7–1 mm long, puberulous; staminate and pistillate bracteoles similar but usually smaller; staminate flowers numerous, distal,;3–12/node in condensed to racemose cymules (to 10 mm long); pistillate flowers 1, basal, fallen or in fruit when staminate flowers are at anthesis. Staminate flowers: pedicel 3–4.2 mm long at anthesis, moderately puberulous; bud subglobose, apex rounded to obtuse, often mucronate; sepals 3, reflexed, elliptic or ovate-elliptic, 1–1.2 3 0.6–0.8 mm, apex obtuse, abaxial surface moderately puberulous; receptacle subglobose to globose, fully covered with anthers, on stalk to 0.3 mm long; nectary an interstaminal 3–6-segmented ring (often inconspicuous in bud); androecium subglobose, 0.4–0.5 3 0.5–0.6 mm, stamens 10–15, dimorphic, 3(4) basalmost with short filaments (0.2–0.3 mm long), alternating with sepals, emerging below and between the nectary segments and arching upwards, remaining 7–12 with filaments absent and densely packed on receptacle, anther sacs ellipsoid, dehiscing longitudinally. Pistillate flowers: pedicel 3–9 mm long (comprising cyme axis 1.5–6 mm 1 true pedicel 1.5–3.5 mm), moderately to densely puberulous, bracteoles 1/3 to 2/3 way along pedicel between cyme axis and true pedicel; sepals 4, triangular, 0.8–1.2 3 0.5–0.8 mm, sparsely puberulous to glabrate, margins hairy; ovary 4–lobed, 0.8–1.1 3 1.4–2 mm, puberulous or sparsely puberulous, lobes rounded and somewhat laterally compressed, 0.3–0.5 mm long, 0.8–1 mm wide at base, conspicuous wings or horns absent; styles 4, entirely connate into a thick cylindrical or obovoid-cylindrical column, 0.9–1.2 3 0.8–1 mm, base constricted or not, apex truncate-rounded, unlobed distally, glabrate; stigmas 4, round-deltoid to circular, each 0.5 mm long, smooth. Fruits 4-lobed capsules,; 1.5 cm wide, surface orange-brown, irregularly rugose-verrucose, glabrate, each carpel lobe subglobose, with a horizontally elongate or crescent-shaped tubercle, 2–3 mm wide, 0.5–0.6 mm high; pedicel; 18 mm long, including cyme axis 4–7 mm long, densely puberulous, true pedicel; 11 mm long, sparsely puberulous. Seeds (not well preserved, squashed) very broadly lenticular or subglobose?,; 7 mm in diam, with radial ridge, surface orange-brown to red-brown, with irregular cream splotches. Figure 7 View FIG .

Etymology —The specific epithet is derived from the Chocó, the Biogeographic Region where it occurs, and the Colombian department where many collections are from.

Distribution, Habitat, and Phenology —This species is known from the central Chocó and eastern Valle del Cauca departments of Colombia, west of the Andes ( Fig. 6 View FIG ). They are vines or slender lianas of tropical pluvial forest, growing in primary or disturbed primary forest, on shrubby vegetation, or along creeks from 10–100 m elevation. Flowering specimens were collected in January, February, May, and July; the single collection seen with fruit was collected in February. Field notes describe their flowers as green or brownish.

Discussion —Collections belonging to P. chocoensis were previously included within P. penninervia , but were noted as possibly representing a new species in the P. penninervia complex ( Gillespie 1993). This species complex is characterized by staminate flowers with dimorphic stamens (outer with filaments, inner with anthers sessile on a subglobose receptacle) and often with a segmented annular nectary, pistillate flowers with a stout-cylindrical stylar column and 6 elliptic leaf blades. Similar to P. penninervia in its leaf blade shape and presence of a single pair of basilaminar extrafloral nectaries, P. chocoensis differs in its larger, elongate, narrowly oblong or oblong-elliptic basilaminar nectaries (versus smaller and circular in P. penninervia ), the blade attenuate basally around the basilaminar nectaries (versus not attenuate), presence of abaxial laminar extrafloral nectaries (versus usually absent), and margins with glandular teeth (versus with distinct rimmed circular glands or with both circular glands and glandular teeth). Blade colour (in the dried state) also differs, with P. chocoensis blades drying olive-green, with darker orange-brown venation below, and P. penninervia blades typically drying grey-green, with paler cream or straw-coloured venation below. Petioles, stems, and inflorescence axes are moderately to densely short puberulous (versus sparsely to moderately puberulous to pubescent in P. penninervia ) and staminate stage inflorescences are longer (1.5–6 cm versus 1–2 cm). The remaining species in the P. penninervia complex ( P. multiglandulosa and P. supraglandulosa ) are also distinguishable from P. chocoensis . Plukenetia chocoensis differs from P. multiglandulosa in having a single pair of basilaminar glands, puberulous petioles, and abaxial leaf blades glabrous or glabrate with major veins glabrate to sparsely puberulous (versus multiple pairs of basilaminar glands, densely hirsute petioles, and hirsute abaxial blades), and differs from P. supraglandulosa in the absence of laminar glands on the adaxial blade surface (versus present).

The segmented annular nectary of P. chocoensis appears to be usually 3-segmented, with each segment horizontally elongate and often 2-lobed. Sometimes one or more of these segments are fully divided into two, resulting in up to 6 segments per nectary.

In contrast to its morphological similarity to P. penninervia , molecular data suggests a relationship with three Amazonian and Guianan species. A previous molecular phylogeny based on two plastid (matK, ndhF) and five nuclear (ETS, ITS, KEA 1 introns 11 and 17, TEB exon 17) markers resolved P. chocoensis (referred to as P. cf. penninervia ; Gentry et al. 47799) in a strongly supported clade with P. brachybotrya , P. loretensis , and P. verrucosa ( Cardinal-McTeague et al. 2019a; summarized in Fig. 1 View FIG ). This suggests that the P. penninervia complex is paraphyletic and that dimorphic stamens and stout-cylindrical styles may be plesiomorphic characters in sect. Penninerviae.

Several collections from Cerros del Cuchillo in the Urabá area of northern Chocó department (Cardenas 1675, 1748, 1903) are here referred to P. penninervia , but need further study. They differ from P. chocoensis in their elliptic basilaminar extrafloral nectaries, absence of abaxial laminar extrafloral nectaries, presence of distinct circular nectaries on the blade margin, longer pubescence on stems and petioles, longer narrowly triangular staminate bracts, and longer pistillate sepals.

Specimens Examined — Colombia. — CHOCÓ: Municipio de Quibdó, bosque frente al barrio Obrero , [5.708333°, -76.65389°], 24 May 1985, Espina & Garcia 1533 ( COL [accession 283420, barcode COL000395807 ; image], F [accession 1982641, barcode V0216685F; image], MO [accession 3292704, barcode MO-1381696 ]) ; Road to Lloró (under construction) 1–2 km SE of Yuto, ca. 30 km S of Quibdó , [5.53833°, -76.63472°], 80 m, 7 January 1979, Gentry & Renter´ ıa 23761 ( COL [accession 206142, barcode COL000395801 ; image], MO [accession 2717121, barcode MO-1381700 ]) ; Road (under construction) to Llora from Yuto , pluvial forest, along creek ca. 2 km E of Yuto, [5.53333°, -76.63333°], 50 m, 18 January 1979, Gentry & Renteria 24367 ( COL [accession 223108, barcode COL000395803 ; image), MO [accession 2716706, barcode MO-1381701 ] ). — VALLE DEL CAUCA: Costa del Pacifico, rio Naya, Puerto Merizalde , 5–20 m, 20–23 February 1943, Cuatrecasas 14006 (F [accession 1358438, barcode V0216684F; image], F [accession 1358485, barcode V0216683F; image], US [accession 2815114, barcode 01298527], US [accession 2815115, barcode 01298528]) ; Costa del Pacifico, rio Cajambre , 5–80 m, 5–15 May 1944, Cuatrecasas 17472 (F [accession 1358549, barcode V0216687F; image], F [accession 1358550, barcode V0194071F; image], US [accession 2817383, barcode 01298562]) ; Puerto Merizalde, R´ıo Nanay , forest behind town, [3.25000°, -77.46666°], 10 m, 22 Feb 1983, Gentry & Juncosa 40563 ( COL [accession 263437, barcode COL000395806 ; image], F [accession 1988829, barcode V0216682F; image], MO [accession 3031302, barcode MO-1381713 ]) ; Buenaventura, Bajo Calima, Pulpapel logging concession SE of Buenaventura, [3.95000°, -77.00000°], 100 m, 22 January 1986, Stein et al. 3260 ( MO [accession 3490284, barcode MO-1381712 ]) .

12. PLUKENETIA LORETENSIS Ule, Verh. Bot. Vereins Prov. Brandenburg View in CoL 50: 81. 1908. Apodandra loretensis (Ule) Pax & K.Hoffm. in A.Engler (ed.), Pflanzenr. IV, 147, IX (Heft 68): 21. 1919. TYPE: PERU. Iquitos, April 1903, E. Ule 6837 (holotype: B [destroyed]; lectotype designated here: L [accession 601470, barcode L 0137704; image!]; isolectotypes: F! [accession 767223, barcode V0042476F; fragment ex G], G! [barcode G00441994], HBG [barcode HBG-515850; image!], MG [barcode MG006665; image!]).

Apodandra corniculata Cardiel, Revista Acad. Colomb. Ci. Exact. 18(71): 469. 1993. Type: Colombia. Caquetá, Sierra de Chiribiquete. Campamento Norte. Recorrido por el cauce casi seco de un arroyo al NE del campamento, [1.11667°, -72.83333°], 350–500 m, 13 December 1990 [mislabeled as 1991 on sheets], S. Castroviejo, J.M. Cardiel, G. Galeano & F. González 12042 (holotype: COL [accession 365413, barcode COL000002059 ; image!]; isotypes: COAH, COL [accession 365412, barcode COL000002060 ; image!], G [barcode G00434095; image!], MA [barcodes MA 509696, MA 509696-2; images!], US [accession 3303795, barcode 00433407; image!]).

Notes —See Gillespie (1993) for species discussion and Gillespie and Armbruster (1997) for species description. We report one new addition to the distributions given in Gillespie (1993) and Gillespie and Armbruster (1997): Pará, Brazil (Ramos 1105).

The holotype was destroyed at B so a new lectotype is designated from the isotype housed at L. This specimen has the most abundant and intact floral and vegetative material among the available isotypes.

Here, we treat Apodandra corniculata as a synonym of P.loretensis . Cardiel Sanz (1993) distinguished this species from P. loretensis on the basis of having leaf blades with 5–7 secondary veins and staminate flowers with 20–25 stamens; however, these characters are within the range of variation observed in P. loretensis ( Gillespie and Armbruster 1997; new observations). Additionally, the collection date on the type labels of A. corniculata are mislabeled as 1991, but the expedition is reported elsewhere on the label and in the manuscript as occurring in 1990 ( Cardiel Sanz 1993).