Danaea pumila Keskiniva & Tuomisto, 2024

Keskiniva, Venni & Tuomisto, Hanna, 2024, Danaea (Marattiaceae) keeps diversifying, part 1: eighteen new species, Willdenowia 53 (3), pp. 173-228 : 215-217

15. Danaea pumila Keskiniva & Tuomisto , sp. nov. (D. subg. Holodanaea) – Fig. 28.

Holotype: Panama, Colón, Teck Cominco Petanquilla mining concession , 08°49.7'N, 80°40.7'W, 362 m, 4 Dec 2007, van der Werff 22253 ( PMA! (119482); isotypes: MO! (2 duplicates: 6252745, 6046894), TUR! (624450)).

Diagnosis — Similar to Danaea humilis T. Moore , but D. pumila generally has shorter sterile leaves (19–32 cm vs (27–) 34–42 cm), fewer pinna-pairs (11–18 vs (16–) 20–25), shorter fertile leaves (19–27 cm vs 37 cm) with fewer pinna-pairs (10–17 vs 20–25), and terminal pinnae almost always replaced by a bud (vs terminal pinnae usually present). Genetically, species differs in locations 80 (C vs T), 764 (A vs G), 779 (A vs C), 1007 (T vs C), 1109 (G vs T) of atpB reference sequence; 80 (A vs C), 101 (A vs C), 108 (G vs C), 109 (T vs C), 158 (T vs G), 185 (G vs A), 390 (G vs A), 665 (G vs A), 758 (G vs A), 1060 (T vs A), 1148 (G vs T), 1178 (C vs T) of rbcL reference sequence; 218 (A vs G), 320 (G vs A), 399 (C vs T), 906 (C vs A) of trnL-F reference sequence; 213 (A vs T), 231 (G vs A), 257 (A vs C), 365 (G vs T), 388 (A vs G), 395 (T vs A), 396 (T vs A), 683 (C vs T), 714 (gap vs A or C), 737 (G vs A), 863 (G vs A) of rpl32 reference sequence.

Description — Rhizomes radial, decumbent to erect, 0.5–1.0 cm in diam., 4–13 cm long. Sterile leaves 19–32 cm long; petioles 6.2–11 cm long, with 1–2 nodes, not winged; laminae 10–22 × 3.4–6.9 cm, linear to lanceolate, paripinnate (or imparipinnate), 11–18 pinna-pairs, medial pinnae 0.8–1.0 cm apart, bicolorous, dark green adaxially, light green abaxially, laminar texture thin, rachises winged (sometimes only in distal part of lamina), wings to 0.2–1 mm wide; terminal pinnae 2.2–3 × 0.6– 1 cm, lanceolate, bases acute, apices 0.9–1.5 cm long, acuminate, margins of apices serrulate to serrate down to widest part of pinna; largest lateral pinnae 2.4–3.4 × 0.7–1 cm, 2.5–4.2 times as long as wide without apex, parallel-sided or widest below middle, slightly ascending, bases asymmetrical (obtuse proximally, obtuse, acute or truncate distally), apices 0–1.2 cm long, acute (to acuminate), margins of apices serrate to serrulate (serrations down to 2/3 of pinna length in distal pinnae); veins 13–20 per cm, variably simple or forked at costae. Fertile leaves 19–27 cm long; petioles 9.5–18 cm long, 1–2 nodes; laminae 7–12 × 1–4 cm, parallel-sided, paripinnate, 10–17 pinna-pairs; largest lateral pinnae 0.7–1.5 × 0.2–0.5 cm, linear-oblong, bases obtuse, apices obtuse to mucronate. Juveniles usually imparipinnate, laminae lanceolate, terminal pinnae lanceolate, lateral pinnae oblong to lanceolate, smallest pinnate juvenile leaves 2 cm long.

Distribution and habitat — Found in Colombia (Chocó, Norte de Santander, Antioquia) and Panama (Colón), in lowland wet rain forests from 250– 600 m. Fig. 8.

Conservation status — We place Danaea pumila in the Vulnerable (VU B1ab(iii)+2ab(iii)) category ( IUCN 2012). It has an Extent of occurrence of 2483 km 2 and an Area of occupancy of 20 km 2, which corresponds to the EN category, and is known from only eight locations, which corresponds to the VU category. One of the collections was from inside a protected area (Parque Nacional Natural Los Katíos in Colombia), but all other collection localities are outside protected areas (one locality was described as secondary forest), and half of the specimens were collected from inside the Minera Panama mining concession, where the area, extent and quality of suitable habitats were inferred to be suffering continuing decline from deforestation.

Etymology — The word Pumila is Latin for diminutive and refers to the small size of the plant in relation to other species of Danaea .

Remarks — Danaea pumila is a small species that morphologically most closely resembles D. humilis . It is smaller (sterile leaves 19–32 cm vs (27–) 34–42 cm, fertile leaves 19–27 cm vs 37 cm long), has fewer pinna-pairs (11–18 vs (16–)20–25), and adult leaves mostly have terminal pinnae replaced by a bud (vs present; however, the terminal pinna is mostly present in juveniles of D. pumila ). Danaea pumila grows at lower elevations than D. humilis (to 600 m vs 900–1600 m) and is found only west of the Andes (vs also on the Amazonian side of the Andes). Genetically the two species are clearly distinct.

Danaea pumila is smaller than D. chococola (sterile leaves 19–32 cm vs 38–48 cm long) and has fewer pinnae (11–18 vs (19–)23–26 pairs) that are bicolorous with abaxial side whitish (vs concolorous).

Danaea pumila differs from D. wendlandii in having acute to acuminate pinna apices (vs usually obtuse) that are serrulate to serrate (vs often crenulate). In addition, D. pumila has generally longer sterile leaves (19–32 cm vs 13–25 cm long), with generally more nodes on the petioles (1–2 vs 0–1), and its terminal pinnae, when present, are longer (2.2–3.0 cm vs 1.7–2.0 cm).

Danaea pumila differs from D. gracilis in having generally fewer pinna-pairs (11–18 vs 15–25) that are opaque and bicolorous (vs translucent and concolorous), wider (2.5–4.2 vs 3.0–6.2 times as long as wide without apex) and have acute apices (vs usually obtuse). If terminal pinnae are present, they are longer in D. pumila than in D. gracilis (2.2–3.0 cm vs max. 1.5 cm).

See Danaea nasua (described above) for comparison with that species.

Additional specimens examined — COLOMBIA: ANTIOQUIA: Vic. Planta Providencia, 28 kms SW of Zaragoza, valley of Río Anorí, 07°18'N, 75°04'W, 400–700 m, 6 Apr 1977, Alverson 383 (MO!, NY!, WIS!); Río Anorí valley near Planta Providencia, 360–600 m, Shepherd s.n. (WIS!) ; CHOCÓ: Riosucio Municipality. Parque Nacional Natural Los Katíos, 250 m, 31 May 1976, Forero 1726 (COL!, MO!) ; NORTE DE SANTANDER: Camp 84 on pipeline, 1800 (ft), 549 m, 16 Sep 1946, Foster 1715 (A!, COL!) . — PANAMA: COLÓN: MPSA Concession, Valle Grande, Sierra 19, 08°50'N, 80°41'W, 291 m, 17 May 2012, Hammel 26229 (MO!, PMA!, UC!); Teck Cominco Petanquilla mining concession, 08°49.9'N, 80°41.1'W, 296 m, 29 Nov 2007, van der Werff 22192 (MO!, PMA!, TUR!); mine site of Minera Panama, Valle Grande Road , 08°50'N, 80°41'W, 250 m, 9 Sep 2012, van der Werff 24420 (MO!, NY!, PMA!, UC!, US!).