Larnax toledoana Barboza & S. Leiva, 2014

Larnax toledoana Barboza & S. Leiva View in CoL , sp. nov. ( Figs. 3 G, H View FIGURE 3 , 9 View FIGURE 9 , 10 View FIGURE 10 )

Type:— ECUADOR. Zamora-Chinchipe: [Cerro Toledo], rumbo a Valladolid desde Yanganá, al costado de la ruta , 2690 m, 79°08'52.7''W, 04°26'30.7’S, 16 November 2011 (fl, fr), C. I. Orozco, G. E. Barboza, A. Orejuela & S. Leiva 3936 (holotype QCA! , isotypes CORD 0000676 ! , CORD 00006770 ! , CORD 00006771 ! , COL , HAO ).

Larnax toledoana differs from L. sagasteguii in its short equal calyx lobes (vs. long unequal calyx lobes in L. sagasteguii ), glabrous filaments (vs. pubescent filaments), non-mucronate anthers (vs. mucronate anthers), and in its widely subglobose fruiting calyx (vs. globose fruiting calyx).

Shrubs, widely branched, plagiotropic, 1–1.6 m tall. Stems terete, hollow; old stems nodose, 10–12 mm wide at base, purple, glabrescent (long simple 4–7-celled non-glandular trichomes), with longitudinal short fissures, and with abundant cream or pale tan lenticels; young stems green, sometimes purple above, always green below, densely villous, slightly sericeous, with long patent simple multicellular non-glandular trichomes; nodes usually deep purple. Leaves alternate, sometimes paired; petiole semi-terete, 1.7–2 cm long, purple above, green below; leaf blade entire, ovate to elliptic, 7.7–8.5 cm long, 3.4–4.1 cm wide, apex somewhat acute, base cuneate, oblique or asymmetric, membranous, slightly fleshy, dark-green and scabrous above, light-green, markedly reticulately nerved with prominent midvein and 4–6 secondary veins pairs below, homogeneously villous with the same trichomes of the stem and petioles on adaxial surface, and mainly on veins and margins on the abaxial surface; branched long non-glandular hairs occurring occasionally along veins below. Fascicles axillary, 4–5-flowered; pedicels 9–11 mm long, green, sometimes purple or dull purple above, filiform, pendent, villous, with long multicellular non-glandular and glandular trichomes. Flowering calyx 4.5–5 mm diameter in anthesis, green, sometimes purple or dull purple externally, dull purple the distal half and green the basal half internally, cupshaped, fleshy, with protruding principal veins, densely villous externally, with long transparent patent nonglandular and tiny glandular trichomes (stalk unicellular, ochraceous head 6-celled), glabrous internally; tube 2.3–2.5 mm long, 3.6–4.5 mm diameter; lobes minute, 0.3–0.5 mm long, 0.5–0.6 mm wide, shortly triangular, acute, erect, with papillae or 2–3-celled non-glandular trichomes at the apex. Corolla slightly campanulate before anthesis, clearly stellate in anthesis, (13–) 16–17 mm diameter, fleshy; tube greenish yellow, 3.4–3.8 mm long, 3.5–4 mm diameter, completely glabrous on both surfaces; lobes 7–9.2 mm long, 2.9–3.1 mm wide, triangular, greenish yellow with dull brownish red spots externally, greenish yellow with a few brownish red spots to almost completely brownish red, except for some greenish yellow stripes internally, the apex papillate and reflexed, margin ciliate, pubescent externally, with long transparent multicellular non-glandular simple and branched (bifurcate or trifurcate) trichomes along the veins, and short glandular trichomes scattered on the surface; glabrescent internally, with the same small glandular trichomes of outer calyx and corolla surface; upper edge of corolla tube with an annular ring of long transparent non-glandular trichomes. Stamens exserted, heterodynamous; filaments glabrous, cream, three longer (2.9–3.1 mm long), two shorter (2.4–2.6 mm long), filament base expansion auriculate, greenish cream, 1–1.1 mm long, auricle small and conspicuous; anthers 1.9–2.1 mm long, 1.7–2 mm wide, white, rarely dull lilac, widely ovoid, non-mucronate, thecae divergent, connective white, triangular, wide, and protruding. Ovary glaucous, glabrous, subgobose, 1.4–1.6 mm long, 1.2–1.4 mm wide, nectary annular, yellowish cream, occupying 20–30% of the ovary length; style 6–6.3 mm long, glabrous, green in the distal area, cream at the base, broadened at the apex, extending ca. 1 mm beyond the anthers; stigma clavate, subbilobate, dark green, 0.4–0.5 mm long, 0.8–1 mm diameter. Fruit a berry, globose, 8–11 mm diameter, bright orange when mature, green when immature, fleshy, glabrous, stone cells 21–23 per fruit, cream, variously shaped (triangular, oblong, polyhedric). Fruiting calyx accrescent, tightly enveloping the berry, (7–) 11–15 mm diameter, open at the apex, transparent or reddish orange apically, markedly 10-costate, pubescent with shorter rather than longer glandular trichomes (stalk 4–6-celled, head oblong ochraceous unicellular) mixed with transparent nonglandular trichomes mainly along veins; lobes conspicuous, short, <1 mm long, equal, triangular acute. Fruiting pedicels 12.5–15 mm long, green or purple, oblique to pendent, pubescent, sericeous. Seeds 60–75 per fruit, 1.7–2 mm long, 1.2–1.4 mm diameter, yellowish brown, reniform, glabrous; testa foveolate.

Etymology: — From the same place as the holotype, “Cerro Toledo” in Podocarpus National Park ( Ecuador).

Phenology: — Flowering and fruiting from April to December.

Distribution and Ecology: — Endemic to the Podocarpus National Park located in southern Ecuador (boundary between Zamora-Chinchipe and Loja Provinces) where it is distributed over an area of approximately 600 km 2 (GoogleEarth 2011; Fig. 4 View FIGURE 4 ). Larnax toledoana inhabits primary cloud forest between 2500–3200 m elev., nearby to streams and rivers.

Species conservation assessment: — According to IUCN criteria ( IUCN 2012), L. toledoana is considered as Vulnerable (VU) based on its extent of occurrence which is estimated to be less than 20,000 km 2 (Criterion B1) and the fact that it is known from less than 10 localities (B1a) and there are fluctuations in its area of occupancy (B1c) suggesting ongoing threats to this species.

Additional specimens examined: — ECUADOR. Loja: ca. 14 km east of Loja, eastern range of the Andes , 2850 m, 79°1'60''W, 4°0'0''S, 28 November 1965 (fl, fr), Knight 922 & 939 ( WIS!) GoogleMaps ; Parque Nacional Podocarpus, common along El Sendero de Lagunas del Compadre , wet montane forest along a quebrada, 2800 m, 79°08'W, 04°05'S, July 1995 (fl, fr), Sawyer 709 ( CORD!) GoogleMaps ; Parque Nacional Podocarpus , vegetación de bosque nublado, 2500–2910 m, 2 August 1986 (fl, fr), Jaramillo et al. 8766 ( QCA 95-13/66 !, 95-13/74 !, NY!) ; Parque Nacional Podocarpus, 5 km from the entrance. Humid montane forest on the "Nudo de Sabanilla" , 2730 m, 79°09'W, 4°07'S, 2 August 1986 (fl), JØrgensen 61386 ( QCA!) GoogleMaps ; Parque Nacional Podocarpus, Nudo de Cajanuma, alrededores del "Centro de Información" , 2850–3000 m, 79°09'W, 04°02'S, 19 December 1990 (fl, fr), Yánez & Gavilanés 88 ( QCA!) GoogleMaps ; Cerro Toledo, jeep track to “La Torre”, 10–12 km SE Yangana , montane forest, 3000–3200 m, 6 April 1985 (fl, fr), Harling & Anderson 23769 ( GB!, CORD!) . Zamora-Chinchipe: rumbo a Valladolid desde Yanganá , al costado de la ruta, 2573 m, 79°08'54.3''W, 04°28'21.4''S, 16 November 2011 (fl, fr), Orozco et al. 3942 ( CORD 00006765 !, 00006764 !, 00006763 !, COL, QCA, HAO!) GoogleMaps ; 500 m adelante del punto anterior, 2650 m, 79°08'52.6''W, 04°27'53.1''S, 16 November 2011 (fl, fr), Orozco et al. 3939 ( CORD 00006767 !, 00006766 !, COL, QCA, HAO!) GoogleMaps ; Yanganá, rumbo al Cerro Toledo, acceso a 700m de un puente de Yanganá pasando la segunda curva , 3024 m, 79°06'57''W, 04°23'1.9°S, 17 November 2011 (fl, fr), Orozco et al. 3949 ( CORD!, COL, HAO!) ; ruta Yanganá-Valladolid, en borde de bosque, base de Reserva Joco-Toco , 2647 m, 79°09’51.7’’W, 4°27’34.2’’S, 15 July 2012 (fl, fr), Deanna & Leiva 7 & 8 ( CORD 00006751 !, 00006752 !, 00006753 !, 00006754 !, 00006755 !) GoogleMaps ; Vilcabamba-Valladolid road, 21.4–27.8 km S of Yangana , disturbed montane forest with steep roadside slopes, 2560–2680 m, 79°08'W, 4°30'S, 21 April 1992 (fl, fr), Luteyn & Romoleraux 14527 ( QCA!) GoogleMaps .

Discussion: — Larnax toledoana can be distinguished from other species of the genus by the combination of short calyx lobes ( Fig. 10 D View FIGURE 10 ), pubescence on vegetative and reproductive organs ( Fig. 10 A–D View FIGURE 10 ), corolla colour ( Fig. 10 B–D View FIGURE 10 ), and the fruiting calyx colour and shape ( Fig. 10 A, C View FIGURE 10 ).

Larnax toledoana most closely resembles L. sagasteguii S. Leiva, Quipuscoa & N. W. Sawyer (1998: 86) , an endemic from eastern Ayabaca (Dept. Piura, Peru). Both species have an externally villous calyx with long transparent simple multicellular non-glandular trichomes, exserted and heterodynamous stamens, an orange mature berry tightly enclosed by the fruiting calyx, and young stems sometimes purple above, with long transparent simple non-glandular trichomes ( Fig. 10 A–E View FIGURE 10 , 17 G–J View FIGURE 17 ). Larnax toledoana differs from L. sagasteguii in its axillary flowers ( Fig. 10 A View FIGURE 10 ), calyx lobes ( Fig. 10 D View FIGURE 10 ), stamen pubescence, presence of mucronate anthers ( Fig. 10 B, D View FIGURE 10 , 17 G View FIGURE 17 ) and fruiting calyx shape and pubescence ( Fig. 10 C View FIGURE 10 , 17 J View FIGURE 17 ) as summarised in Table 7.

Larnax psilophyta N. W. Sawyer (1998: 74) is sympatric with L. toledoana , both inhabiting Podocarpus National Park ( Sawyer 1999) and can be distinguished according to the characters in Table 8. Larnax harlingiana is another geographically close species and differs from L. toledoana in several characters ( Table 9).

A specimen belonging to L. toledoana has been erroneously cited as neotype of L. suffruticosa ( Sawyer 1999) because no original material of this latter species had been located. Consequently many specimens of L. toledoana have frequently been wrongly annotated as L. suffruticosa (here synonymized with Deprea orinocensis ).