Danaea cuspidopsis Keskiniva & Tuomisto, 2024

Keskiniva, Venni & Tuomisto, Hanna, 2024, Danaea (Marattiaceae) keeps diversifying, part 1: eighteen new species, Willdenowia 53 (3), pp. 173-228 : 187-191

5. Danaea cuspidopsis Keskiniva & Tuomisto , sp. nov. (D. subg. Holodanaea) – Fig. 11, 12.

Holotype: Costa Rica, Puntarenas, Monteverde Biological Station , 10°18.2'N, 84°47.5'W, 1500 m, 16 Jul 2000, Boyle 5971 ( NY!).

Diagnosis — Morphologically very similar to but genetically distinct from Danaea cuspidata Liebm. , differing in locations 390 (A vs G), 948 (A vs G), 1060 (T vs A) of rbcL reference sequence; 192 (A vs C), 233 (C vs A), 265 (A vs G), 399 (C vs T), 822 (C vs A), 249–253 (insertion of ATTAG vs gap), 729–732 (insertion of AATA vs gap) of trnL-F reference sequence; 170 (A vs T), 176 (A vs C), 231 (G vs A), 257 (A vs C), 396 (T vs A), 424 (A vs C), 539 (C vs A), 643 (G vs T), 678 (G vs T), 713 (T vs A), 737 (G vs A), and 867 (G vs T) of rpl32 reference

sequence. Generally differs from D. moritziana C. Presl in veins being simple or forked at costa (vs veins almost never simple and usually forked well above costae), veins more densely packed (17–22 vs 12–17 veins per cm), terminal pinnae often replaced by a bud (vs terminal pinnae always present), and gradually tapering pinna apices (vs abruptly tapering); genetically differs in location 192 (A vs C) of trnL-F reference sequence and location 678 (G vs T) of rpl32 reference sequence. Differs from D. andina Keskiniva & Tuomisto in generally having longer fertile pinnae (6.5–12 cm vs 4.2–7.0 cm); longer (1.7–3.2 cm vs 0.7–1.6 cm) and gradually tapering (vs usually abruptly tapering) sterile pinna apices; generally longer sterile leaves (54–120 cm vs 28–71 cm); veins sometimes mostly simple (mostly forked); rhizomes erect (vs can be creeping to ascending). Differs from D. robbinmoranii Keskiniva & Tuomisto in narrower wings (to 0.1–0.7 mm vs 0.7–2 mm wide) that cover only distal part of rachises (vs usually all of rachises and sometimes part of petioles as well); generally more pinna-pairs (13–20 vs 7–16) that are usually narrower (largest lateral pinnae 5–8 vs 3–6 times as long as wide without apex), and spaced more closely together (medial pinnae 1.4–2.5 cm vs 2.9–3.5 cm apart).

Description — Rhizomes erect, radial, 1.5–2.5 cm in diam., to at least 25 cm long. Sterile leaves 54–120 cm long; petioles 22–57 cm long, with 1–2 nodes, not winged; laminae 29–65 × 12–33 cm, 13–20 pinna-pairs, oblong to lanceolate, imparipinnate or paripinnate with a terminal bud, proximal pinna-pair sometimes clearly smaller than others and more distant, medial pinnae 1.4–2.5 cm apart, bicolorous, (dark) green (to brown) adaxially, light green (to light brown) abaxially, texture thin, rachises winged

in distal parts of internodes in distal part of lamina, wings to 0.1–0.7 mm wide; terminal pinnae 5.0–15 × 1.3–3.0 cm, lanceolate, bases acute, apices 1.5–3.0 cm long, acu- minate, margins of apices serrulate to serrate; largest lat- eral pinnae 8.0–16 × 1.9–2.4 cm, 5.0–7.8 times as long as wide without apex, parallel-sided, (slightly) ascending (or perpendicular to rachis), bases asymmetrical (ob- tuse or auriculate proximally, obtuse or acute distally), apices 1.7–3.2 cm long, (long-)acuminate, margins of apices serrulate to serrate; veins 17–22 per cm, usually forked at costa, only occasionally forked above costae or simple, rarely almost all simple. Fertile leaves 55–118 cm long; petioles 27–60 cm long, with 2 nodes; laminae 37–50 × 9–22 cm, oblong or lanceolate, imparipinnate or paripinnate with a terminal bud, 13–19 pinna-pairs; terminal pinnae 3.5–5.5 × 0.5–0.7 cm, linear-lanceolate, bases acute, apices acuminate to cuneate; largest lateral pinnae 6.5–12 × 0.5–1.0 cm, linear, slightly ascending or perpendicular to rachis, bases symmetrical (obtuse) or asymmetrical (obtuse distally, auriculate proximally), apices 0.4–1.0 cm long, acuminate to cuneate, margins of apices serrulate to serrate. Juveniles with lanceolate laminae, lateral pinnae oblong, rather narrow, apices acu- minate. Distribution and habitat — Abundant in Costa Rica, but also found from Panama (Chiriquí) and the Pacific side of the Andes in Colombia (Antioquia, Chocó , Risalda , Valle de Cauca) and Ecuador (Imbabura, Pichincha). Possibly extends to Nicaragua. Grows at elevations 550–2300 m, but usually found growing above 1000 m. It has been found in primary forests and disturbed secondary forests, in tall, montane forests and cloud forests, often on steep slopes and in ravines. Fig. 13.

Conservation status — We place Danaea cuspidopsis in the Least Concern (LC) cat- egory ( IUCN 2012). It has a known Area of occupancy of at least 128 km 2, which cor- responds to the EN category, and an Extent of occurrence of at least 534,564 km 2, which corresponds to the LC category. Danaea cuspidop- sis is abundant especially in Costa Rica, and it is known from 50 collections with oc- currences in several protected areas (Braulio Carillo National Park, Monteverde Nature Reserve, Tapantí Reserve, Parque Nacional La Amistad in Costa Rica, Tandayapa Cloud Forest Reserve in Ecuador). There appears to be no imminent threat to all its subpopulations.

Etymology — The name highlights the fact that Danaea cuspidopsis is a cryptic species morphologically very similar to but genetically clearly distinct from D. cuspidata.

Remarks — Danaea cuspidopsis is an intermediately sized Danaea with bicolorous leaves, pinnae with rather long acuminate apices, veins usually forked at the costa, and terminal pinnae often replaced by a bud. It is a cryptic species that we separate from D. cuspidata on the basis of genetic evidence: D. cuspidopsis is deeply embedded in a 9-species clade that does not contain D. cuspidata , this being resolved to another clade ( Keskiniva & al. 2024). Danaea cuspidopsis seems to grow to a larger size than D. cuspidata (sterile leaves to 120 cm vs to 86 cm long), but we have been unable to find reliable morphological characters to separate the two. We have genetically verified records of D. cuspidopsis from Costa Rica and of D. cuspidata from Mexico but do not know how either species is distributed in between. Danaea cuspidata has

often been considered synonymous with D. moritziana (e.g. Christenhusz 2010; Tuomisto & al. 2018), but on the basis of both genetic and morphological evidence we are now convinced they are distinct.

Another potential name from the same geographical area is Danaea muenchii Christ. The protologue of D. muenchii written by Rosenstock (1926) cites both Costa Rican and Mexican specimens. However, the protologue also quotes a letter by Christ which says that the diagnosis of the species can be found in Christ (1905), which in turn is based on a Mexican specimen collected by Münch from San Pablo, Chiapas, Mexico in 1904. A specimen corresponding to this information, Münch 159 (P), was mentioned as a holotype in Mickel & Beitel (1988), and this can be considered a lectotypification under Art 9.23 ( Turland & al. 2018). As we have identified all of the similar specimens from Mexico as D. cuspidata , we consider D. muenchii to be a synonym of that species.

Danaea cuspidopsis resembles D. betancurii , which is found on the Amazonian side of the Andes and between the Cordilleras in Colombia (vs D. cuspidopsis only on the Pacific side), but D. cuspidopsis is generally larger than D. betancurii (sterile leaves 54–120 cm vs 37–99 cm long) and almost always has wider pinnae (largest lateral pinnae 1.9–2.4 cm vs 0.9–2 cm wide, the type of D. betancurii having especially narrow pinnae) that are clearly bicolorous with the abaxial side whitish (vs sometimes almost concolorous). Rhizomes are always erect in D. cuspidopsis , whereas some specimens of D. betancurii seem to have a creeping to ascending rhizome.

Our current understanding is that Danaea moritziana has a mostly Venezuelan distribution and does not occur in the western side of the Andes, where D. cuspidopsis is common. Danaea cuspidopsis generally differs from D. moritziana in veins being simple or forked at the costae (vs veins almost never simple but usually forked well above the costae), a denser venation (17–22 vs 12–17 veins per cm), terminal pinnae often replaced by a bud (vs terminal pinnae always present), and gradually tapering pinna apices (vs abruptly tapering).

Danaea cuspidopsis differs from D. stricta Tuomisto & Keskiniva in having clearly bicolorous laminae (vs almost concolorous), narrower fertile pinnae (0.5–1.0 vs 1.0– 1.5 cm wide), falcate pinnae (vs strictly linear), medial pinna apices that are asymmetrical (vs symmetrical) and obtuse or acute (vs truncate), and a thinner lamina texture. In addition, the terminal pinna is often present in D. cuspidopsis (vs always replaced by a bud in D. stricta ), and the rhizome is generally shorter (up to 25 cm tall vs to 40 cm tall) and more slender (1.5–2.5 cm vs 2.2–4 cm in diam.).

Danaea cuspidopsis differs from D. inaequilatera in having generally narrower fertile pinnae (0.5–1.0 vs 0.8– 1.3 cm wide) and sterile pinnae that are whitish abaxially (vs almost concolorous), longer (medial pinnae 5.0–7.8 vs 3.7–4.7 times as long as wide without apex), asymmetrical at bases (vs usually symmetrical), and have generally longer apices (1.7–3.2 cm vs 0.5–2.0 cm long) that are gradually tapering (vs abruptly tapering).

See Danaea andina (described above) and D. robbinmoranii (described below) for comparison with those species.

Additional specimens examined — COLOMBIA: ANTIOQUIA: Angelópolis Municipality. Vereda Romeral, Hacienda La Argentina, Quebrada Las Animas, 06°08'N, 75°42'W, 2050–2130 m, 18 Nov 2005, Rodriguez 5604 (NY!); CHOCÓ: NW side of Alto del Buey, Trail along ridge from confluence of forks of Río Mutatá above Río Dos Bocas to top of Alto del Buey, 1450–1750 m, 9 Feb 1971, Lellinger 247 (US-3!); 0.3 km E of Ciudad Bolívar-Quibdo Road across suspension bridge at c. km 141, 750 m, 3 Apr 1971, Lellinger 881 (US-2!); RISARALDA: Corregimiento de Puerto de Oro, Vereda Chirrincha, banks of Río Aguita, 800–900 m, 13 Sep 1991, Fernández-Alonso 8993 (MO!); VALLE DEL CAUCA: El Tambo– Munchique, between Uribe and “La 81” (La Romelia), 1800 m, 12 Feb 2015, Kessler 14850 (HUA!, TUR!). — COSTA RICA: ALAJUELA: Vicinity of Coliblanco, 1950 m, 30 Apr–2 May 1906, Maxon 291 (NY!, US!); Zarcero, Smith H261 (F!); Viento Fresco, 1600–1900 m, 13 Feb 1926, Standley 47774 ( US!); CARTAGO: Tapantí Reserve, 1400–1700 m, 1982, Gómez 19287 (AAU, MO!); SE of Orosi, c. 2.2 km SSE of Purisil, above Finca La Concordia, in gorge next to house of uppermost portion of upper finca, 1800 m, 8–10 Aug 1970, Lellinger 1481 ( US!); 1923, Maxon 8151 ( US!); Turrialba, near Interamerican Institute, in Cervantes ravine, 550–650 m, 9–10 Mar 1953, Scamman 6992 (GH!, US!); along trail leading eastward into mountains from road into Tapantí Reserve c. 1 km S of Jct. of Quebrada Salto and Río Grande de Orosi, 09°43'N, 83°47'W, 1500–1800 m, 1 Feb 1986, Smith 2165 (CR!, NY-2!, UC!); El Munco, S of Navarro, 1400 m, 1924, Standley 33642 ( US!); Cerro de la Carpintera, Standley 35594 ( US!); GUANACASTE: SW (Pacific) slope of Cerro Cacao, Cordillera de Guanacaste, 10°55'N, 85°28'W, 1300–1450 m, 10 Aug 2007, Grayum 12581 (CR!, MO!); Liberia, Parque Nacional Guanacaste, Volcán Cacao, trail to top, 10°56'N, 85°27'W, 1450–1550 m, 8 Aug 2007, Rojas 7709 (MO!); La Cruz, Santa Cecilia c. 5 km from Estación Pitilla, Fila Orosilito, headwaters of Río Mena (Pitilla-Volcán Cacao route), 10°58'N, 85°27'W, 1100–1200 m, 8 Apr 2008, Rojas 8340 (MO!); HEREDIA: Braulio Carillo Park, Zurquí, 1700–2000 m, 1 Mar 1983, Gómez 20219 (MO!, UC!); Braulio Carillo National Park, 10°15'N, 84°10'W, 1830 m, 8 Nov 1986, Hennipman 6784 (AAU, BM!, CR, F!, K!, MO!, NY!, P!, U!, UC!, US!, Z!); Vara Blanca, between Poás and Barba Volcanoes, 1600–1700 m, 22 Mar 1953, Scamman 6991 (GH!, US!); Pcia. San José, along unnamed N fork of Río Zurquí, Cordillera Central, 10°04'N, 84°01'W, 1500–1600 m, 18 Jan 1986, Smith 1667 (AAU, CR!, MO!, NY!, UC-2!); S slopes of Cerro Zurquí, 5 km N of San Luis Norte, 10°03'N, 84°02'W, 1800 m, 20 Mar–4 Apr 1973, Stolze 1569 (F-2!); LIMÓN: Cordillera de Talamanca, Atlantic slope, canyon of Río Siní, 09°13'N, 82°59'W, 1800–1900 m, 15 Sep 1984, Davidse 29113 (MO!); PUNTARENAS: In and around Monteverde Nature Reserve, Chomogo Trail, 10°18'N, 84°47'W, 1450–1650 m, 31 Oct–2 Nov 1975, Burger 9780 (F-2!); Coto Brus, P. I. La Amistad, Cordillera de Talamanca. Estación Pittier. Trail to Río Gemelo, 09°01'N, 82°58'W, 1650 m, 30 Jan 1995, Fletes 26 (CR!, MO!); Upper Río Burú, 2010 m, 19 Aug 1983, Gómez 21429 (CR!, MO!, UC!); Upper Río Burú, 2010 m, 19 Aug 1983, Gómez 21753 (MO!, UC!); Vicinity of biological field station at Finca Wilson , 5 km S of San Vito de Java, 1200–1400 m, 4 Aug 1967, Mickel 3114 (NY!, US!); P. N. La Amistad. Cuenca Térraba-Sierpe, Estación Pittier, Cerro Gemelo, 09°03'N, 82°56'W, 2600–2700 m, 23 May 1996, Moraga 477 (CR!, MO!); Monteverde, Motas, 1967, Walter 5-1968 (MO-2!); SAN JOSÉ: San Cristobal Norte, 2000 m, 1969, Gómez 2397 (F!); Tarrazu, San Marcos, from street crossing to San Carlos 100 m toward San Marcos, 09°34'N, 84°10'W, 1700 m, 21 Sep 2004, Rojas 6104 (CR!, MO!); La Palma, 17 Mar 1910, Brade 508 (NY!); La Picada de San Antonio, Cerro de la Muralla de Socorro de San Ramón, 1200–1300 m, 22 Aug 1927, Brenes 5680 (F!); La Palma de San Ramón, 1275– 1275 m, 8 Mar 1930, Brenes 11907 (F!); 1928, Valerjo 306 ( US!); Monteverde, 28 Oct 1963, Walter 258 (MO!). — ECUADOR: IMBABURA: In vicinity of Río Verde, c. 5 km SW from village of Mani, Río Cachaco, 00°46'N, 78°28'W, 1300 m, 2 Jun 1980, Sperling 5035 (GH!, QCA!); PICHINCHA: Tandayapa Cloud Forest Reserve, 1750–1880 m, 2004, Lehnert 1215 (UC-2!, QCA!); Road El Paraiso–Saguangal, 11 km from El Paraiso, 00°12'N, 78°46'W, 1250 m, 2 May 1982, Øllgaard 37644 (AAU, QCA!); Road El Paraiso–Saguangal, 3 km from El Paraiso, 00°10'N, 78°46'W, 1500 m, 2 May 1982, Øllgaard 37782 (AAU, QCA!, UC!); Road to Mindo, 00°02'S, 78°44'W, 1350 m, 26 May 1997, Navarrete 1954 (AAU, QCA!). — PANAMA: CHIRIQUI:Vicinity of El Boquete, 1380 m, 1933, Bro. Maurice 667 ( US!); Vicinity of El Boquete, 1000–1500 m, 1918, Cornman 1082 ( US!); Boquete Region, Cerro Horqueta, 1900–2100 m, 30 Jul 1940, von Hagen 2166 (NY-2!); Between Alto de las Palmas and top of Cerro de la Horqueta, 2100–2268 m, 18 Mar 1911, Maxon 5518 (US-2!); Upper Caldera River, above El Boquete, 1450– 1650 m, 22–24 Mar 1911, Maxon 5595 (US-2!); Along Río Caldera (Boquete region) and on slope to E, c. 3.5 km NW of Bajo Mono, 08°50'N, 82°28'W, 1600 m, 8 Feb 1986, Smith 2506 (NY!, PMA-2!, UC-2!, US). Boquete, 08°50'12.7''N 82°27'34.3''W, 1800 m, 29 Jan 2016, Testo 994 (NY, PMA, VT).