Prescottia smithii Schltr., Repert., 2014

Azevedo, Cecília Oliveira De, Berg, Cássio Van Den & Barros, Fábio De, 2014, A revision of Prescottia (Orchidaceae: Orchidoideae, Cranichideae), Phytotaxa 178 (4), pp. 233-286 : 274-279

15. Prescottia stachyodes (Sw.) Lindl., Bot. Reg. 22: sub t. 1916: 1. 1836. [ Cogniaux 1895, Cogniaux 1907, Cogniaux 1909, Fawcett & Rendle 1910, Kränzlin 1911, Gale & Baldomero 1938, Hoehne 1945, Williams 1946, Renz 1948, Williams 1951, Hodge 1953, Williams 1956, Schweinfurth 1958, Dunsterville & Garay 1959, Correll 1965, Vargas 1965, Pabst 1966, Schweinfurth 1967, Adams 1972, Garay & Sweet 1974, Hamer 1974, Pabst & Dungs 1975, Garay 1978, Hamer 1984, Werkhoven 1986, Barros 1987, Ackerman 1989, Correa 1992, Cremers & Hoff 1992, Ackerman 1995, Gloudon & Tobisch 1995, McLeish et al. 1995, Sprunger 1996, Espejo-Serna & López-Ferrari 1998, Jørgensen & León-Yánez 1999, Ackerman 2000, Balick et al. 2000, Dix & Dix 2000, Nir 2000, Carnevali et al. 2001, Feldmann & Barré 2001, Stevens et al. 2001, Hammel et al. 2003, Rocha & Waechter 2006, Azevedo & van den Berg 2007a, Azevedo & van den Berg 2007b]; basionym: Cranichis stachyodes Sw., Prodr. : 120. 1788. [ Lindley 1836, Lindley 1840 a, Cogniaux 1895, Cogniaux 1909, Fawcett & Rendle 1910, Gale & Baldomero 1938, Williams 1946, Williams 1951, Hodge 1953, Williams 1956, Schweinfurth 1958, Dunsterville & Garay 1959, Pabst 1966, Schweinfurth 1967, Garay & Sweet 1974, Hamer 1974, Garay 1978, Hamer 1984, Ackerman 1989, Sprunger 1991, Correa 1992, Cremers & Hoff 1992, Ackerman 1995, McLeish et al. 1995, Sprunger 1996, Espejo-Serna & López-Ferrari 1998, Jørgensen & León-Yánez 1999, Ackerman 2000, Nir 2000, Carnevali et al. 2001, Johnson 2001, Stevens et al. 2001, Hammel et al. 2003, Rocha & Waechter 2006, Azevedo & van den Berg 2007a, Azevedo & van den Berg 2007b, Govaerts 2014].

Type :— JAMAICA. (Blue Mountains) Swartz s.n. (lectotype BM!, selected by Azevedo & van den Berg 2007a). Fig. 31 .

Heterotypic synonyms: Prescottia colorans Lindl., Bot. Reg. 22: t. 1916. 1836 [ Cogniaux 1895, Cogniaux 1909, Gale & Baldomero 1938, Williams 1956, Schweinfurth 1958, Dunsterville & Garay 1959, Pabst 1966, Hamer 1974, Pabst & Dungs 1975, Sprunger 1991, Cremers & Hoff 1992, McLeish et al. 1995, Sprunger 1996, Nir 2000, Govaerts 2014]. Type:— BRAZIL. Loddiges s.n. (holotype K-L!); Prescottia petiolaris Lindl., Bot. Reg. 22: t. 1916: 2. 1836 [ Schweinfurth 1958, Sprunger 1991, Cremers & Hoff 1992, Sprunger 1996]. Type:— PERU.Yambras (Bamba), Mathews 1875, Herb. Hookerianum (holotype K!, line drawing K-L!); Prescottia pellucida Lindl., Ann. Mag. Nat. Hist., Ser. 3, 1: 335. 1858. [ Cogniaux 1909, Gale & Baldomero 1938, Ackerman 2000]. Type (lectotype, here designated):— Cuba. Orientali: 1856–7, Wright s.n. (K-L!, the left collection); Prescottia longipetiolata Barb.Rodr. , Gen . Spec. Orchid. 1:177. 1877 [ Cogniaux 1895, Schweinfurth 1958, Pabst 1966, Pabst & Dungs 1975, Sprunger 1991, Cremers & Hoff 1992, Sprunger 1996, Johnson 2001, Govaerts 2014]. Type (lectotype, here designated):—Barbosa Rodrigues’ original drawing (plate t. 475) in the library of the Jardim Botânico do Rio de Janeiro [reproduction printed in Iconographie des orchidées du Brésil, Sprunger et al. (1996), 2: t. 84]; Prescottia paulensis Cogn. in Mart., Fl. Bras. 3(6): 548. 1906 [ Pabst 1966, Pabst & Dungs 1975, Sprunger 1991, Sprunger 1996, Johnson 2001, Goaerts 2014]. Type:— BRAZIL. São Paulo: Serra da Cantareira, Comm. Geogr. e Geol. S. Paulo 3052, comm. cl. Löfgren (holotype BR; isotype SP!); Prescottia smithii Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 52. 1920. [ Garay 1978, Azevedo & van den Berg 2007a, Govaerts 2014]. Type:— COLOMBIA. Magdalena: Santa Marta, above las Partidas, Smith 2277 (lectotype NY!; isolectotypes CM!; F!; GH!; K!; US!, selected by Azevedo & van den Berg 2007a); Prescottia longifolia Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 51. 1920 [ Azevedo & van den Berg 2007a]. Type:— COLOMBIA. Antioquia: c. 2000 m. Madero 120 (lectotype drawing of type AMES!, selected by Azevedo & van den Berg 2007a); Prescottia longipetiolata Schltr., Repert. Spec. Nov. Regni Veg. Beih. 8: 39. 1921. Type (lectotype, here designated):—Schlechter’s drawing [printed in Mansfeld 1929. Repert. Spec. Nov. Regni Veg. Beih 57: t. 75, f. 291]; Prescottia schlechteri Hoehne, Fl. Bras. 8 (12; 2): 113. 1945 (non Prescottia longipetiolata Barb.Rodr. 1877 ); Prescottia colorans var. macrophylla Hoehne, Revista Soc. Brasil. Agron. 8: 222. 1945. [ Govaerts 2014]. Type:—(location unknown, not in SP); Prescottia tepuyensis Carnevali & C. A. Vargas, Lindleyana 11: 236–238. 1996. Type:— VENEZUELA. Territorio Federal Amazonas: Rio Negro: Cerro Aracamuni, summit, Proa Camp, savanna with small to large patches of forest, especially along streams, in ravines and near edge of tepui, 01 o 32’ N, 65 o 49’ W, 1,400 m, 25 Oct 1987, “epiphyte on base of tree trunk. Flowers green. Unicate”, Liesner & Carnevali 22460 (holotype: VEN, photo!; isotype MO!); Prescottia villenaorum Christenson, Orchids 71: 618. 2002. Type:— PERU. San Martin: mountains near Moyobamba, February 2002, Christenson 2034 (holotype CUZ).

Terrestrial herbs. Leaves 1–4(–5), rosulate, petiolate; petiole green to rose-red (2.0–)6.0–35.0(–41.0) cm long; blade dark green to variegated, (3.5–)6.5–28.0(–30.5) × (2.0–)3.0–11.5(–15.8) cm, elliptic to ovate, apex acute, base sometimes hyaline-white, attenuate to obtuse, margin entire to serrulate. Inflorescence (4–)30–75(–160)-flowered; peduncle red to green (7.0–)20.0–60.0(–95.0) × 0.2–0.6(–1.3) cm; peduncle bracts (2–)3–6(–8), green to rose-red, (8.0–)20.0–50.0(–120.0) × (8.0–)10.0–15.0(–22.0) mm, oblong, apex acute; rachis green to rose-red (6.0–)15.0–25.0(– 44.0) × 0.4–1.5(–2.2) cm. Flower bracts rose-red to green, (3.0–)7.0–10.0(–17.0) × 2.0–3.0(–5.0) mm, lanceolate, apex acuminate to caudate; flower erect, whitish to green; dorsal sepal strongly revolute, (1.0–)3.3–4.6(–6.0) × (0.5–)1.0– 1.2(–1.5) mm, triangular to ovate, apex acute; lateral sepals strongly revolute, (2.0–)4.6–5.7(–6.6) × (0.6–) 1.2–1.6 mm, triangular to ovate, apex acute; petals strongly revolute, (1.0–)3.5–5.0(–5.4) × 0.4–0.5 mm, linear, apex acute; lip green, (2.0–)2.8–4.0(–6.0) × 2.5–3.0(–4.0) mm, inner surface glabrous. Column (0.8–) 1.6–2.7 mm long, glabrous.

Distribution:— This species occurs from North America ( Mexico), Central America, West Indies to South America ( French Guiana, Surinam, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia, Paraguay, Brazil, and Argentina). In Brazil, it has been cited for the states of Amazonas, Pará, Ceará, Pernambuco, Alagoas, Bahia, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul, Minas Gerais and Distrito Federal ( Fig. 32 ).

Selected Specimens Examined:— ARGENTINA. Buenos Aires, Trindade Hills, 17 March 1932, Jack 8628 ( GH) . BELIZE. Toledo: Vicinity of Doyle’s Delight, southen Maya Mountains , 11 December 1993, Allen 15406 ( MO) . BOLIVIA. La Paz: Muñecas , Consata , 08 April 1981, Luer s.n. ( SEL 81666) . BRAZIL. Alagoas: Quebrangulo, Parque Estadual da Pedra Talhada, Pedra D’agua, 24 July 1987, Rodrigues 1189 ( MAC) . Amazonas: Camanaus, 31 October 1971, Prance 15882 ( INPA, NY) . Bahia: Mucugê, Parque Municipal de Mucugê, mata do Zé Leandro , 16 August 2002, Azevedo 148 ( HUEFS) . Ceará: Guaramiranga , Sitio São Salvador, 13 September 1998, Castro 601 ( EAC) . Distrito Federal: Riacho Fundo, ca. 15 km SW of Brasília on road to Goiânia , 24 September 1965, Irwin 8609 ( F, MO, NY) . Espírito Santo: Castelo , Forno Grande, 15 August 2006, Azevedo 288 ( HUEFS) . Minas Gerais: Chapadão das Perdizes, cruzamento a vau do Rio Capivari , 06 April 2007, Azevedo 325 ( HUEFS) . Pará: São Felix do Xingu , a 20 km sul da Vila Central , 15 August 2001, Salles 2256 ( HEPH) . Paraná: Alexandra , 07 September 1910, Dusén 10193 ( F, S) . Pernambuco: Brejo da Madre de Deus , Propriedade Bituri, 05 February 1965, Andrade-Lima 65 4299 ( IPA) . Rio de Janeiro: Itatiaia, Parque Nacional de Itatiaia , em direção a parte alta, trilha para o Rebouças , 02 April 2007, Azevedo 318 ( HUEFS) . Rio Grande do Sul: São Leopoldo, Morro das Pedras , 20 October 1927, Dutra 945 ( HUEFS, ICN) . Santa Catarina: Blumenau, 1884, Schwacke 59 ( R) . COSTA RICA. Alajuela: SE slope of Vulcan Arenal between Fortuna and Palma , 25 July 1990, Luther 2812 ( SEL) . DOMINICA. Badineau: wooded slopes west of Morne Gay house, 24 March 1940, Hodge 2227 ( GH) . DOMINICAN REPUBLIC. Cordillera Central: Prov . La Vega, Parque Nacional J. A. Bermúdez : en la Loma “La Cotorra”, aprox. 45–60 mins. lejos de Los Tablones en el sendero al Pico Duarte, 16 January 1987, Zanoni 37759 ( NY) . ECUADOR. Pichincha: Rio Toachi on road from Quito-Santo Domingo, 15 July 1983, Hirtz 1084 ( SEL) . EL SALVADOR. Boqueron del Volcán San Salvador, 17 December 1974, Hamer 445 ( SEL) . GRENADA. Mt. Gilbert : NE of Mt. Maitland, 4–10 March 1979, Howard 18793 ( BM, GH, NY, US) . GUADELOUPE. Cascade Vauchelet : s.d., Quentin 646 ( P) . GUATEMALA. Alta Verapaz: Cubilquitz , 06 January 1906, Tuerckheim 8589 ( GH6366 , GH71571 , K, NY, US). FRENCH GUYANA. Mont Saint-Marcel : zone du sommet Sud, 20 July 2002, Granville 15362 ( CAY) . GUYANA. U. Takutu-U. Essequibo, Rupununi River, betw Kwattamang Landing & Rewa Village , 29 September 1997, Clarke 6793 ( NY, US) . HAITI . Mission: Fonds Varettes , 17–04 April– May 1920, Leonard 3871 ( NY, US) . HONDURAS . Cortes: NE shore of Lake Yojoa , 30 December 1952, Williams 18755 ( F, US) . JAMAICA. St.Andrew: Mt. Horeb , Fairy Glades , 01 February 1977, Podzorski 08 ( K) . MARTINIQUE. s.l., 1870, Hahn , 1361 ( BM, G, P, S) . MEXICO. Chiapas: Ocosingo , en estación Chajul, 14 December 1992, Martínez 25877 ( MO) . NICARAGUA. Castillo: Río San Juan, Reserva Indio-Maíz, a 8 km de la cabecera del Río Bartola , en dirección hacia el Cerro el Diablo, 03 January 1997, Rueda 5278 ( MO) . PANAMA. Chiriqui: Fortuna Dam area , N of reservoir, between Quebrada Bonito and Quebrada Franh , to E of road, 01 August 1984, Churchill 5888 ( MO) . PARAGUAY. Paraguai: Parque Nacional Ybycu’i , Road to Cesar Barrientos , 15 September 1988, Zardini 7207 ( MO) . PERU. Cusco: Urubamba, Machu Picchu, on a hillside above the Rio Mandor , 3 km from km 114 of the Urubamba railroad, plot #196, 21 September 1982, Peyton 1296 ( MO, SEL) . USA. Puerto Rico: Adjuntas, Cordillera Central , Barrio Saltillo , mountain slopes c. 1.4–2 km due SSW of Alto de la Bandera, 31 January 1987, Proctor 42977 ( US). Virgin Islands : St. John , Coral Bay Quarter , road from Bordeaux to Reef Bay , 22 February 1993, Acevedo-Rodríguez 5278 ( NY, US). ST. VINCENTS. s.l., March 1890, Smith 949 ( NY) . SURINAM. Inselberg Talouakem - Massif des Tumuc-Humac , 08 August 1993, Granville 12141 ( US) . TRINIDAD. West Indies: Blanchissene Road near 10 1/ 2 mile post near a ridge, 01 May 1925, Broadway 5668 ( K) . VENEZUELA. Aragua: Henri Pittier National Park , Pico Guacamaya, ridge extending E from peak, 15 February 1990, Edwards 224 ( K) .

Habitat:— Terrestrial in thick leaf litter in shade in wet broadleaf forests, 0–3,600 m.

Conservation Status:— Not endangered.

Etymology:— From the Greek stachy (relating to a spike) and odes (resembling) in reference to the shape of the inflorescence ( McLeish et al. 1995).

Notes:— Prescottia stachyodes is a widespread species that is variable in shape and size. It has clearly petiolate leaves with lanceolate to elliptic blade, green flowers and an internally glabrous lip. The leaves are completely green or bear a central whitish or light-green stripe and sometimes white margins, the blade margins entire to lightly serrate. Leaves of P. stachyodes are variable in its shape and dimensions, apparently depending on the place where the plants grow. Plants growing on rocks differ from those in soil, which is probably the reason for the large number of synonyms for this species.

As currently circumscribed, P. stachyodes is highly variable in size, color, and shape. The delimitation proposed here for P. stachyodes is broader and includes some additional morphological variation (and additional species concepts). We could not find any geographic or ecological pattern or significant variation to recognize these proposed forms as separate species. There is continuous morphological variation, even in specimens within the same collection and population. On the other hand, seed morphology of this taxon ( Fig. 10G,H , 11 ) is unique and distinct from all other species of the genus (with fusiform intercellular spaces with beaded appearance; Fig. 10H , 11B,D,F,H ). All forms studied from different regions displayed the same seed morphology. In the future, population studies should be carried out to examine in detail populations and forms within this complex along their geographical range to assess whether it is possible to segregate any group(s) of populations as distinct species.

These plants appear to be self-pollinating since virtually all flowers set fruit. Singer & Sazima (2001) verified that P. stachyodes is self-compatible but pollinator-dependent. However, germinated pollen was observed in the anther cavity near the clinandrium ( Fig. 7E,F ) as reported for Epipactis microphylla (Ehrh.) Sw. ( Bonatti et al. 2006) , indicating variation in breeding system. The clinandrium turns back, and pollen comes into contact with the stigmatic surface. This is an efficient way to ensure seed set independent of pollinator presence. This process would explain somewhat distinct variants growing sympatrically.

Prescottia colorans was described as a unifoliate Prescottia with a purplish scape, probably the reason of the name. The holotype of P. colorans is annotated in Lindley’s handwriting with the information “ Brasil Loddiges”, in agreement with the information given in the protologue. This herbarium sheet has two collections, one being the type, a single inflorescence without leaves, and an illustration, and the other “ Wright 626, in Cuba Orientali”, which is not part of the type. The latter was collected in July 1856, after publication ( Lindley 1836) of P. colorans , but it was also determined and cited by Lindley (1858) as a P. colorans , which indicates that Lindley’s concept of P. colorans included big specimens not only from Brazil but also from Cuba.

When Lindley described Prescottia petiolaris ( Lindley 1836) , he did not say from where the type from. Later, Lindley (1840a) cited the type as: Hab. in Peruvia, Mathews, 1875 (exam. s. sp. in hb. Hooker). The specimen at K is stamped “Herbarium Hookerianum 1867” (1867 being the year that Kew bought Hooker’s herbarium), and this implies that it is the holotype of P. petiolaris . There is also a herbarium sheet in Lindley’s herbarium with a line drawing of the type he made from Hooker’s herbarium.

The type of Prescottia pellucida contains specimens with large flowers (lip 4.5–5.0 mm long) and short petioles (1.5–3.0 cm long). In his Flowering plants of Jamaica, Adams (1972) recognized P. pellucida as a good species (as did Ackerman 1989, 1995) but expressed some doubts and suggested that it may simply be an ecological variant of P. stachyodes . Prescottia pellucida has been regarded as synonym of P. stachyodes based on morphometric data taken from herbarium specimens collected throughout the Greater Antilles ( Ackerman 2000). Nir (2000) recognized P. pellucida as a good species and cited “ Cuba, Loma del Gato, Wright 626 (K-L)” as its type but without seeing the specimen. However, this number was cited by Lindley (1858) at the same time he published P. pellucida , exactly the line before it, as P. colorans . This specimen (Wright 626) is different from the type of P. pellucida .

In the protologue of Prescottia pellucida where Lindley (1858) listed orchids collected in eastern Cuba by Wright, he explicitly mentioned “Loma del Gato” (but no number). Ackerman (1989, 2000) cited “ Cuba, Loma del Gato, Wright s.n. (holotype K-L!, photograph of specimen at UPRRP!)” as the type of the species. The problem is that the material at K-L has three collections in the same sheet. One, Wright 1493, was from Monte Verde, but it was collected after the description in 1859, which excludes it as a possible type material. The other has no year and is labelled as “ Prescottia pellucida ?”, but it has only one leaf instead of the two leaves mentioned in the description. The other one (on the left) has two leaves but no number and was collected in 1856– 7 in “ Cuba Orientali”. This collection also has a line drawing by Lindley below, and therefore we selected it as the lectotype of P. pellucida .

When Schlechter (1920) described P. smithii he did not cite where the type was deposited. Four specimens of Smith 2277 were found at CM, GH, K, and NY. The original material of P. smithii was probably at B, but it could not be found and was probably destroyed during World War II. The original material of Smith was distributed by NY, and for this reason Azevedo & van den Berg (2007a) selected the collection at NY as the lectotype.

When describing P. longipetiolata, Schlechter (1921) characterized the new species as being similar to P. stachyodes , but with smaller flowers in a dense inflorescence, remarking that the type has only a notably long petiolate leaf, which was also emphasized by Lindley (1836) as a character of P. colorans . As Barbosa Rodrigues (1877) had already used the epithet longipetiolata, Hoehne (1945) chose P. schlechteri as a nomen novum for P. longipetiolata Schltr. Prescottia longipetiolata was considered as a synonym of P. cordifolia Rchb. f. by Hoehne (1945), Garay (1978), Hamer (1984), and Govaerts (2014), and here we consider it as a synonym of P. stachyodes .

Schlechter’s original material of P. longifolia and P. longipetiolata were probably destroyed along with the collections in B. Azevedo & van den Berg (2007a) lectotypified it. The latter we chose to lectotypify with the line illustration (tab. 291) printed in Mansfeld (1929), Figuren Atlas zu den Orchideenfloren der südamerikanischen Kordillerenstaaten.

When Carnevali & Vargas (1996) described Prescottia tepuyensis , they said that this taxon was closely related to P. stachyodes , but the plants and flowers are consistently smaller, and the floral bracts proportionally shorter. Despite these statements, this taxon is similar to the type of P. stachyodes in size. Carnevali & Vargas (1996) stated that small specimens of P. stachyodes have leaves of at least 10 cm long but are usually longer than 15 cm, and in P. tepuyensis the leaves do not exceed 8 cm in length (differing from the information given at the description: 8–11 cm long). The type of P. stachyodes has leaves 7–9 cm long. Prescottia stachyodes is extremely variable in shape and size, and the type of P. tepuyensis falls easily within the circumscription of P. stachyodes .

Figures

FIGURE 7. SEM micrographs of Prescottia pollen. Elastoviscine. A,B. Prescottia phleoides (Azevedo 344). Pollen exine. C. Prescottia phleoides (Azevedo 344). D. Prescottia stachyodes (Azevedo 288). Germinated pollen. E,F. Prescottia stachyodes (Azevedo 288).

FIGURE 10. SEM micrographs of Prescottia seed morphology.A,B. Prescottia phleoides (Azevedo 344). C,D. Prescottia plantaginifolia (Mori 10702). E,F. Prescottia spiranthophylla (Mulford 881). G,H. Prescottia stachyodes (Azevedo 318).

FIGURE 11. SEM micrographs of Prescottia stachyodes seed morphology. A,B. (Harris 10096). C,D (Kuhlmann 2742). E,F (Oliveira 842). G,H (Raven 19994).

FIGURE 31. Prescottia stachyodes. A. Habit. B,C. Flower. B. Front view. C. Lateral view. D. Floral bract. E. Perianth parts, clockwise from top: lip, dorsal sepal, petal, lateral sepal. F,G. Column F. Dorsal view. G. Ventral view. (Azevedo 318).

FIGURE 32. Distribution of Prescottia stachyodes.