Danaea panamensis Keskiniva & Tuomisto, 2024

Keskiniva, Venni & Tuomisto, Hanna, 2024, Danaea (Marattiaceae) keeps diversifying, part 1: eighteen new species, Willdenowia 53 (3), pp. 173-228 : 209-211

12. Danaea panamensis Keskiniva & Tuomisto , sp. nov. (D. subg. Danaea ) – Fig. 24 , 25 .

Holotype: Panama, Panamá, Cerro Campana National Park , 08°41'N, 79°56'W, 850–900 m, 25 Oct 2005, Tuomisto 15161 ( PMA! (mounted on 2 sheets: 102072 & 102073, has separate rhizome); isotypes: AAU!, TUR! (mounted on 3 sheets), UC! (mounted on 2 sheets), Z! (mounted on 2 sheets)).

Diagnosis — Most closely related to Danaea latipinna Tuomisto & R. C. Moran , but differing in more pinna-pairs (12–16 vs 2–6); generally narrower pinnae (2.6– 3.5 cm vs 2.8–8.7 cm). Similar to D. nodosa (L.) Sm., but differing in thicker laminar texture; dark blue-green colour when dry (vs light yellow green); pinnae that are generally smaller (to 27 vs to 35 cm long, to 3.5 vs 5.8 cm wide) and narrower (6–12 vs 4–8 times as long as wide without apices); pinna apices that are usually gradually tapering (vs often abruptly tapering), generally shorter (to 3.3 cm vs to 4.0 cm long), and entire, sinuate or crenulate (vs often serrulate at shoulder of pinna); can have a petiolar node (vs no nodes in petiole). Differs from D. leussinkiana Christenh. in rhizomes with leaf bases in two rows (vs 3–5 rows); generally narrower pinnae (2.6–3.5 cm vs 3.1–4.9 cm wide); longer pinna apices (1.3–3.3 cm vs 0.9–1.5 cm); thinner laminar texture. Differs from D. pterorachis Christ in rhizomes with leaf bases in two rows (vs 3–5 rows); longer pinnae (21–27 cm vs 14–21 cm long); dark blue-green colour (vs green); thicker laminar texture. Differs from D. ampla Keskiniva & Tuomisto in more pairs of fertile pinnae (12–14 vs 4–7); generally more pinna-pairs (12–16 vs 4–13) that are narrower (2.6–3.5 cm vs 3.1–4.9) and parallel-sided (vs usually widest above middle). Danaea panamensis is genetically unique among Danaea in having a C (vs A) at position 126 of rpl32 reference sequence and differs from all other species of D. subg. Danaea by having an A (vs G) at position 481 of rpl32 reference sequence and a T (vs C) at position 695 of atpB reference sequence.

Description — Rhizomes creeping, dorsiventral with leaf bases in two rows, 2.5–3.5 cm in diam. when dry, 4–5 cm in diam. when fresh, to 40 cm long. Sterile leaves 121–157 cm long; petioles 66–92 cm long, with 0(–1) nodes, not winged; laminae 55–76 × 42–47 cm, (long-)obovate, imparipinnate, terminal pinnae shorter than second distal-most pinna-pair, 12–16 pinna-pairs, proximal pinna-pair more distant and slightly smaller, medial pinnae 2.4–4.0(–6.0) cm apart, almost concolorous, rather dark bluish green adaxially, often more yellowish toward margins, abaxial side uniformly green or brown, laminar texture relatively thick, rachises not winged or winged in distal part of lamina, wings to 0.7 mm wide; terminal pinnae 9.8–21 × 2.1–2.6 cm, lanceolate to oblong, bases acute, apices 1.5–3.0 cm long, (long-)acuminate, margins of apices entire, sinuate or crenulate; largest lateral pinnae 21–27 × 2.6–3.5 cm, 5.6–11.8 times as long as wide without apex, parallel-sided, bases symmetrical (obtuse or acute) or asymmetrical (obtuse or auriculate proximally, acute or obtuse distally), apices 1.3–3.3 cm long, acuminate (to abruptly caudate), margins of apices entire, sinuate to crenulate; veins 14–18 per cm, mostly forked at costae, sometimes above. Fertile leaves 122–145 cm long; petioles 72–89 cm long, lacking nodes; laminae 50–57 × c. 22 cm, 12–14 pinna-pairs, imparipinnate, long-elliptic to long-lanceolate, pinnae spaced proximally, crowded distally; terminal pinnae 6.2–9.8 × 1.3–2 cm, lanceolate, bases acute, apices acuminate; largest lateral pinnae 13–16 × 1.2–2.0 cm, oblong to elliptic, bases asymmetrical (obtuse or auriculate proximally, acute to obtuse distally), apices 1.3 cm long, acuminate, margins of apices entire or crenulate. Juveniles with terminal pinnae elliptic, oblong or lanceolate, lateral pinnae oblong when leaves are more than 10 cm long, bases asymmetrical (obtuse or auriculate proximally, acute distally), apices acuminate, smallest juveniles with elliptic pinnae, largest simple juvenile 4.4 cm long, smallest pinnate juvenile 3.9 cm long.

Distribution and habitat — Known from Panama from 70– 900 m. Danaea panamensis has been found on steep slopes and on the banks of a ravine and a stream channel. One locality mentions clay to loam soils and hilly terrain with ravines. Fig. 3 .

Conservation status — We place Danaea panamensis in the Least Concern (LC) category ( IUCN 2012). It has an Area of occupancy of 68 km 2, which corresponds to the EN category, and an Extent of occurrence of 25,571 km 2, which corresponds to the LC category. Danaea panamensis seems to be rather common, and is known from 24 collections. It has been collected in several protected areas in Panama (Barro Colorado Island, Bosque Protector Palo Seco, Cerro Campana National Park, Parque Nacional Soberanía, Parque Nacional San Lorenzo, and Parque Nacional Santa Fe) and there appears to be no imminent threat to all its subpopulations.

Etymology — The species is known only from Panama.

Remarks — In the field, Danaea panamensis and D. ampla were thought to represent D. media and D. nodosa , respectively, but genetic evidence now shows that there are three distinct species involved and that D. media is a synonym of D. nodosa (see D. ampla for more details). Danaea panamensis has a thicker laminar texture than D. nodosa and D. ampla and dries a rather dark bluish green (vs often yellowish light green), with a more yellowish area around the midrib adaxially (vs concolorous); it can also have a petiolar node (vs none). It generally differs from D. nodosa in having smaller pinnae (to 27 cm vs to 35 cm long, to 3.5 cm vs to 5.8 cm wide) that are narrower (6–12 vs 4–8 times as long as wide without apex) and in more gradually tapering apices (vs usually rather abruptly tapering) that are shorter (to 3.3 cm vs to 4.0 cm long), almost never with serrulations (vs often serrulate at the shoulder of the pinna). Danaea panamensis differs from D. ampla in having more pairs of fertile pinnae (12–14 vs 4–7) and generally more pairs of sterile pinnae (12–16 vs 4–13) that are also narrower (2.6–3.5 cm vs 3.1–4.9 cm wide, 5.6–12 vs 2.4–5.6 times as long as wide without apex) and parallel-sided (vs usually widest above the middle).

In leaf morphology, Danaea panamensis is similar to the Costa Rican D. pterorachis but differs in having rhizomes with two rows of leaves (vs 3–5 rows) and longer pinnae (21–27 cm vs 14–21 cm long) that have a darker bluish colour (vs brighter green) and a thicker texture.

We have not seen specimens from Panama that we could confidently identify as Danaea nodosa or D. pterorachis , and it is therefore possible that these two species do not overlap in distribution with D. panamensis , which seems to be endemic to, and rather common in, Panama.

Genetically Danaea panamensis is most closely related to D. latipinna of coastal Ecuador but differs morphologically in having more pinnae (12–16 vs 2–6 pairs) that are narrower (2.6–3.5 cm vs 4.8–8.7 cm wide, 6–12 vs 2–3 times as long as wide without apex). Danaea leussinkiana co-occurs with D. panamensis and shares the dark blue-green colour when dry, but D. panamensis has rhizomes with leaf bases in two rows (vs 3–5) and narrower pinnae (2.6–3.5 cm vs 3.1–4.9 cm wide, 5–7 vs 6–12 times as long as wide without apex) with longer apices (1.3–3.3 cm vs 0.9–1.5 cm) and a thinner laminar texture.

Danaea panamensis also co-occurs with D. alba , but is a smaller plant (sterile leaves 121–157 cm vs 160–200 cm long) with rhizomes having pinna apices in two rows (vs 3–5) and more pinna-pairs (12–16 vs 8–9) that are smaller (21–27 × 2.6–3.5 cm vs 33–37 × 5.2–5.5 cm), and dry to a darker bluish green (vs pale yellowish or greyish green). The shape of the juveniles of these species also differs, with D. panamensis having oblong pinnae (vs elliptic).

Additional specimens examined — PANAMA: BOCAS DEL TORO: Bosque Protector Palo Seco, Río Changuinola, Charco La Pava. At house of Mr. Reynaldo Abrego, 09°09'N, 82°30'W, 348 m, 3 Feb 2013, Ortiz 1204 (MO!); El Guabo District. Bosque Protector Palo Seco, Farm of Mr. Dario Araúz, 08°50'N, 82°11'W, 431 m, 6 Feb 2013, Zapata 3148 (MO!); Bosque Protector Palo Seco, Concesión Hidro Ecológica del Teribe, near Bonyic creek, Casa máquinas, 09°20'N, 82°38'W, 355 m, 29 Sep 2013, Zapata 3347 (MO-2!); CHIRIQUI: Canal Zone northwest of Gamboa along Pipeline Road, McAlpin 1393 (D!, F!); COLÓN: Cocle del Norte, Escorpio 02 helipad area, taking southern route, 08°56'N, 80°40'W, 39 m, 21 Jul 2012, Espinosa 6022 (MO-3!); Colón, Sierra Llorona, at P32, 09°21'N, 79°44'W, 285 m, 4 Mar 2008, Jones 920 (TUR!); Colón, Parque Nacional San Lorenzo, at P2, 09°20'N, 79°58'W, 137 m, 23 Apr 2008, Jones 1051 (PMA!, TUR!); Minera Panama, Point C24, Río San Lucas, 08°59'N, 80°34'W, 12 Mar 2010, Martínez 531 (MO!); Minera Panama, W13 helipad area, Río Caimito area, 08°58'N, 80°40'W, 69 m, 20 Oct 2010, Martínez 661 (MO!); Porto Bello, 5–200 m, 6–8 Apr 1911, Maxon 5750 (US-2!); Santa Rita, STRI tree plot 32, 09°21'N, 79°44'W, 400 m, 20 Oct 2005, Tuomisto 15142 (PMA!); PANAMÁ: Parque Nacional Soberanía, at P8, 09°10'N, 79°45'W, 215 m, 21 Jan 2008, Jones 635 (TUR!, US!); PN Soberanía, at P8, 09°10'N, 79°45'W, 215 m, 28 Jan 2008, Jones 683 (TUR!); PN Soberanía, at P8, 09°10'N, 79°45'W, 215 m, Feb 2 2008, Jones 735 (TUR!); PN Soberanía, at P20, 09°12'N, 79°46'W, 254 m, 8 Feb 2008, Jones 753 (TUR!); PN Soberanía, at P15, 09°10'N, 79°45'W, 90 m, 15 Feb 2008, Jones 802 (TUR!); PN Soberanía, at P19, 09°12'N, 79°46'W, 153 m, 25 Mar 2008, Jones 957 (TUR!, US!); PN Soberanía. Collected at P16, 09°08'N, 79°43'W, 182 m, 2 Apr 2008, Jones 982 (TUR!); Canal Zone, Pipeline Road, Agua Salud, 1 Nov 1972, Kennedy 1889 (PMA!, US!); Barro Colorado Island, 13 Feb 1934, Taylor 1328 (MICH-2!); Barro Colorado Island, close to beginning of Shannon Trail, 09°10'N, 79°50'W, 100–150 m, 23 Oct 2005, Tuomisto 15149 (PMA-2!, TUR!); VERAGUAS: Parque Nacional Santa Fe. Alto de Piedra. First branch of Río Mulabá. Highway to Guabal, close to bridge at km 4.45, 08°32'N, 81°08'W, 663 m, 16 Nov 2012, Ortiz 993 (MO-2!); Parque Nacional Santa Fe, Alto de Piedra, First branch of Río Mulabá, highway to Guabal, close to bridge at km 4.45, 08°32'N, 81°08'W, 676 m, 16 Nov 2012, Ortiz 1011 (MO!, PMA!).

Figures

Fig. 24. Danaea panamensis: A: medial fertile pinna; B: creeping rhizome seen from above; C: creeping rhizome seen from side; D: medial pinna apex; E: juvenile; F: sterile leaf; G: fertile leaf. – A, D, F, G: Tuomisto 15161 (TUR); B, C: Jones 1051 (TUR); E: Tuomisto 15149 (TUR). – Drawn by Venni Keskiniva.

Fig. 25. Danaea panamensis: A: field habit; B: whole plant. – A, B: Tuomisto 15149. – © Hanna Tuomisto.

Fig. 3. Distribution of types and paratypes of Danaea alba, D. ampla, D. antioquiana and D. panamensis.