Danaea andina Keskiniva & Tuomisto, 2024

Keskiniva, Venni & Tuomisto, Hanna, 2024, Danaea (Marattiaceae) keeps diversifying, part 1: eighteen new species, Willdenowia 53 (3), pp. 173-228 : 180-184

3. Danaea andina Keskiniva & Tuomisto , sp. nov. (D. subg. Holodanaea C. Presl) – Fig. 6, 7.

Holotype: Ecuador, Zamora-Chinchipe, Reserva Tapichalaca, sector Ventanillas , 04°29'S, 79°07'W, 2520– 2600 m, 17 Sep 2004, Lehnert 1278 ( UC! (1794187); isotype: GOET! (042670)).

Diagnosis — Resembles Danaea moritziana C. Presl , but is usually smaller (sterile leaves to 71 cm vs to 96 cm long, sterile pinnae 4.5–13 cm vs 7.8–20 cm long, fertile pinnae to 7 cm vs to 11 cm long), has variable rhizomes (vs always erect), and its veins are mostly forked at costa (vs often forked well above costae); genetically differs in locations 985 (T vs C) of atpB reference sequence; 80 (C vs A), 113 (C vs A), 885 (C vs A), 886 (G vs A), 1060 (A vs T), 1136 (C vs T), 1148 (T vs G) of rbcL reference sequence; 233 (A vs C), 248–252 (gap vs insertion of ATTAG), 265 (G vs A), 728–731 (gap vs AATA) of trnL- F reference sequence; 231 (A vs G), 257 (C vs A), 396 (A vs T), 424 (C vs A), 539 (A vs C), 709–710 (TT vs gap), 713 (C vs T), 737 (A vs G), 863 (A vs G) of rpl32 reference sequence. Similar to and co-occurs with D. cuspidopsis Keskiniva & Tuomisto , D. excurrens Rosenst. and D. betancurii A. Rojas. Differs from D. cuspidopsis in generally shorter fertile pinnae (4.2–7.0 cm vs 6.5–12 cm); shorter pinna apices (0.7–1.6 cm vs 1.7–3.2 cm) that are abruptly tapering (vs usually gradually tapering); generally shorter leaves (28–) 46–71 cm vs 54–120 cm); veins forked (can be simple); rhizomes creeping to erect (vs erect). Differs from D. betancurii in generally shorter fertile pinnae (4.2–7.0 cm vs 5.3–8.0 cm); generally wider pinnae (medial pinnae 2–6 vs 4–8 times as long as wide without apex); abruptly tapering pinna apices (vs gradually tapering); veins forked at costa (often simple or forked well above costae). Differs from D. excurrens in generally shorter fertile pinnae (4.2–7.0 cm vs 3.6–10.4 cm); generally longer rhizomes (to 21 cm vs to 10 cm long); acute pinna bases (vs obtuse); pinnae parallel-sided (vs often widest at base). Is unique among Danaea by having a deletion at positions 635–698 of rpl32 reference sequence and at positions 382–394 of trnL-F reference sequence.

Description — Rhizomes with leaf and root bases arranged radially, variously described as creeping to erect, 0.7–2.2 cm in diam., to 21 cm long. Sterile leaves (28–) 46–71 cm long; petioles 6–38 cm long, with 0–2 nodes, not winged or winged down to most distal node on petiole; laminae 19–43 × 7–23 cm, oblong, elliptic or (long-) lanceolate, imparipinnate (or paripinnate), 8–16 pinna-pairs, proximal pinnae often smallest and more distant, medial pinnae 1.4–3.5 cm apart, bicolorous, dark green adaxially, light green to green abaxially, texture thin, rachises entirely or at least distally winged, wings to 0.2–0.7 mm wide; terminal pinnae 3.6– 8.0 × 0.8–2.1 cm, lanceolate, bases acute, apices 1.1–2.8 cm long, (long-)acuminate, margins of apices crenulate to serrate; largest lateral pinnae 4.5–13.1 × 1.0– 2.3 cm, 2.2–5.9 times as long as wide without apex, parallel-sided, slightly falcate, bases asymmetrical (acute, obtuse or slightly auriculate proximally, acute distally), apices 0.7–1.6 cm long, attenuate to acuminate, margins of apices serrate (to crenate); veins 12–22 per cm, mostly forked at costae (sometimes partly forked above costae or partly simple). Fertile leaves 45–65 cm long; petioles 26–37 cm long, with 1 node; laminae 17–28 × 5–7 cm, long-elliptic, imparipinnate (or paripinnate with a terminal bud), 13–15 pinna-pairs; terminal pinnae c. 2.8 × 0.5 cm, linear-lanceolate, bases acute, apices cuneate; largest lateral pinnae 4.2–7.0 × 0.5–0.7 cm, linear-elliptic to linear-lanceolate, slightly falcate, bases asymmetrical (obtuse or acute proximally, obtuse or acute distally), apices 0.4–0.6 cm long, cuneate to acuminate, margins of apices crenulate. Juveniles with parallel-sided laminae, terminal pinnae oblong to lanceolate, lateral pinnae oblong (to elliptic), apices cuspidate to acuminate.

Distribution and habitat — Danaea andina is found in the tropical Andes and seems to be relatively abundant in Ecuador, where it occurs on both sides of the mountain chain at 1750–2600 m. It is also found in Colombia (Putumayo), and probably occurs in northern Peruvian Andes as well. Danaea andina has been found in montane rain forests, often in shady, moist and inclined spots such as stream ravines. Fig. 8.

Conservation status — We place Danaea andina in the Least Concern (LC) category ( IUCN 2012). It has a known Area of occupancy of 56 km 2, which corresponds to the EN category, and an Extent of occurrence of at least 51,331 km 2, which corresponds to the LC category. Danaea andina seems to be abundant in the Ecuadorian Andes, as it is known from at least 18 collections, and some of its occurrences are inside protected areas (Tandayapa Cloud Forest Reserve, Parque National Podocarpus, and Reserva Tapichalaca in Ecuador). There appears to be no imminent threat to all its subpopulations.

Etymology — This species is named for its distributional area in the Andes.

Remarks — Danaea andina is one of the few species of Danaea that grow on both sides of the Andes, and it is also among the ones that reach the highest elevations. It belongs to the species complex that was previously referred to D. moritziana , and its distribution overlaps with several other species that have recently been segregated from that complex ( D. cuspidopsis , D. excurrens , and D. betancurii ). Although many individuals of D. andina are easy to identify by their characteristic appearance, within-species morphological variation and its overlap with other species is considerable. Consequently, the morphological comparisons below are rather inconclusive; D. andina is most reliably identified based on its unique DNA.

Danaea andina differs from D. cuspidopsis (and also from the Mexican D. cuspidata ) in having generally shorter leaves (28–71 cm vs 54–120 cm long), shorter fertile pinnae (4.2–7.0 cm vs 6.5–12 cm), and shorter pinna apices in sterile pinnae (0.7–1.6 cm vs 1.7–3.2 cm long) that are abruptly tapering (vs often gradually tapering). In addition, the rhizomes of D. andina can be creeping to erect (vs always erect), its veins are mostly forked at the costa (vs sometimes mostly simple), and the terminal pinna is almost always present (vs often replaced by a bud in D. cuspidopsis ).

Danaea andina also resembles D. moritziana but is usually smaller (sterile leaves to 71 cm vs to 96 cm long, sterile pinnae 4.5–13 cm vs 7.8–20 cm long, fertile pinnae to 7 cm vs to 11 cm long), has more variable rhizomes (vs always erect), and its veins are mostly forked at the costa (vs often forked well above the costae). Under our current relatively narrow concept, D. moritziana is restricted to Venezuela and northern Colombia, and does not appear to have spread far enough south as to overlap with the distribution of D. andina .

Christenhusz (2010) considered Danaea betancurii as synonymous with D. moritziana , but Rojas-Alvarado (2013) argued that the two are distinct. We do not have DNA for D. betancurii , but the type of this species is distinctive with a simple venation and rather delicate pinnae with gradually tapering apices. We consider D. betancurii to be a distinct species, although it is variable in our current circumscription.

Danaea betancurii occurs on the Amazonian side of the Andes from Colombia to northern Peru and is usually found at lower elevations than D. andina (600–2050 m vs 1950–2600 m). The veins of D. andina are generally forked at the costa, unlike the simple veins in the type of D. betancurii (although the venation seems to be variable in D. betancurii ). In addition, D. andina has generally shorter fertile pinnae (4.2–7.0 cm vs 5.3–8.0 cm) and generally wider sterile pinnae (medial pinnae 3–6 vs 4–8 times as long as wide without apex) that have abruptly tapering pinna apices (vs gradually tapering). Danaea andina often differs from D. excurrens in having shorter fertile pinnae (to 7.0 cm vs to 10 cm), longer rhizomes (to 21 cm vs to 10 cm long), and lateral pinnae that are parallel-sided (vs often widest at base) and have acute bases (vs obtuse). However, both species are variable, and we do not have reliable criteria to morphologically separate between Ecuadorian specimens of D. andina and Bolivian specimens of D. excurrens . Because we do not have genetic material of either species from Peru, we are not sure about the identity of the Peruvian specimens. This renders the distributional limits between the two species uncertain as well.

Danaea andina differs from D. inaequilatera A. Rojas , which occurs in the lowlands of the Pacific coast, by having clearly bicolorous pinnae that are whitish beneath (vs almost concolorous), less regularly arranged sterile pinnae of more variable shape, and narrower fertile pinnae (0.5–0.7 cm vs 0.8–1.3 cm wide).

The juveniles of Danaea andina have short and broad, often squarish pinnae with short, abruptly tapering apices and long, rather slender radial rhizomes that have been variously described as creeping to erect. The juveniles of D. moritziana have rounder and more elliptic pinnae and the juveniles of D. excurrens usually have narrower pinnae than those of D. andina .

Additional specimens examined — COLOMBIA: PUTUMAYO: Above Sibundoy, 2500 m, 4 May 1939, Alston 8378 (BM!). — ECUADOR: LOJA: Trails to Quebrada Romerillos c. 5 km ENE of San Pedro de Vilcabamba, 04°14'S, 79°10'W, 2100–2200 m, 29 Nov 1994, Øllgaard 105897 (AAU, NY!, QCA!, TUR-2!, UC); Trails c. 5 km ENE of San Pedro de Vilcabamba, Loma and Quebrada El Trigal, 04°14'S, 79°10'W, 2050–2250 m, 2 Dec 1994, Øllgaard 106015 (AAU!, QCA!); MORONA-SANTIAGO: Road Plan de Milagro-Gualaceo, km 10.8, S of road, 03°00'S, 78°32'W, 2200–2250 m, 24 Nov 1997, Øllgaard 2740 (AAU!, QCA!); NAPO: Quijos Canton, Yanahyacu, 00°36'S, 77°53'W, 3 Aug 2015, Lehnert 3279 (VT!); Baezam along road leading to radio towers behind town, 77°53'S, 00°27'W, 2080 m, 24 Feb 2005, Moran 7540 (NY!, UC!); PICHINCHA: Bellavista; between Tandayapa and Mindo (old road Quito–Puerto Quito), 2300 m, 10 Sep 2004, Lehnert 1161 (GOET, QCA!, TUR!, UC!); Tandayapa Cloud Forest Reserve, 1750–1880 m, 12 Sep 2004, Lehnert 1214 (GOET, TUR!, QCA!); km 6 on Pela Gallo-Monte Cristi road, 00°03'N, 78°31'W, 2500 m, 26 May 1997, Navarrete 1949 (QCA!); TUNGURAHUA: W of Río Guambi, c. 7 km along road and trails S of Río Negro, 01°27'S, 78°15'W, 1780 m, 4 Feb 1998, Navarrete 3053 (QCA!); ZAMORA-CHINCHIPE: Estación Cientifica San Francisco, Quebrada 2, 03°58'S, 79°04'W, 2050 m, 13 Sep 2003, Lehnert 842 (QCA!, UC!); Reserva Tapichalaca, study plot B5, on N slope of Cerro Tapichalaca, 04°29'S, 79°07'W, 2645 m, 31 Oct 2003, Lehnert 1044 (QCA!, UC!); Parque National Podocarpus, Reserva Biologica San Fransisco, Q5, MATRIX plot L6 (old label Q6), 03°58'S, 79°04'W, 1940–1960 m, 10 Sep 2010, Lehnert 1796 (STU!); Parque Nacional Podocarpus , trail into Quebrada San Francisco, km 9.4 E of pass on Loja- Zamora road, 03°59'S, 79°06'W, 2000–2300 m, 25 Mar 1998, Øllgaard 2978 (AAU-3!, QCA!); Road Loja-Zamora, 13 km E of pass, just before junction with old road, 03°58'S, 79°05'W, 2030 m, 8 Mar 1989, Øllgaard 90898 (AAU!, QCA!); new road Loja-Zamora, km 13 E of pass, 03°58'S, 79°06'W, 2000 m, 14 Feb 1991, Øllgaard 98814 (AAU!, QCA!); area of Estación Cientifica San Francisco, road Loja-Zamora, c. 35 km from Loja, 03°58'S, 79°04'W, 2050 m, 4 Aug 2005, Werner 1732 (QCA!, UC!).