Sarcorhachis sydowii Trel., Repert., 2012

Sarcorhachis sydowii Trel., Repert. View in CoL Spec. Nov. Regni Veg. 48: 16. 1940.—TYPE: ECUADOR. Pichincha: prope Mindo , 1,000–1,200 m, 6 Nov 1937, H. Sydow 317 (lectotype: US! sheet #1743685; isotype: US! sheet #1743684; designated by Arias et al. Novon 16: 207. 2006). Manekia sydowii (Trel.) Arias, Callejas & Bornstein, Novon View in CoL 16: 207. 2006.

Sarcorhachis sydowii Trel. var. hirsuta Yuncker, Ann. Missouri

Bot. Gard. 53: 380. 1966.— TYPE: ECUADOR. Zamora:

road from Loja to Zamora, km 45–51, 1,400 –1,600 m, 20

Nov 1961, C. H. Dodson & L. B. Thien 1423 (holotype:

MO!; isotype: NY !).

Sarcorhachis venezuelana Steyermark, Pittiera View in CoL 3: 33. 1971.—

TYPE: VENEZUELA. Carabobo: selva siempre verde a

lo largo del Río San Gían , al sur de Borburata, arriba de

la Planta Eléctrica, entre Los Tanques y la Toma, 750 m,

27–28 March 1966, J. A. Steyermark & C. S. Steyermark

95152 (holotype: VEN; isotypes: S!, US!). Manekia

venezuelana (Steyermark) Arias, Callejas & Bornstein ,

Novon 16: 207. 2006.

Stems with dark brown to black pellucid dots occasionally present, glabrous or rarely pubescent; internodes 0.9–18.7 (average 5.05) cm long. Prophylls 0.2–2.9(–0.73) cm long, drying dark brown to reddish brown. Leaves of monopodial axes with petioles 0.6–9.9(–3.35) cm long, vagination extending onto blade 0.1–1.7(–0.74) cm; laminae 4.4–18.1 + 2.0–19.5 (–8.88 + 7.3) cm, symmetrical or occasionally slightly asymmetrical, ovate to broadly ovate, rarely lanceolate, glabrous or rarely pubescent below on primary veins or near leaf base, base symmetrical, angle obtuse, shape rounded to truncate to cordate, apex straight to less often acuminate, dark, shiny green above, pale green below, chartaceous, drying subcoriaceous, pellucid dots reddish brown to black, visible above, below or on both surfaces, or apparently absent; venation suprabasal acrodromous, primary veins 7 or 9, often elevated on lower surface, three central veins united distal to petiole insertion 0.5–2.5 + cm. Spikes white to cream-colored in flower, solitary or paired with peduncles 0.2–6.3(–2.08) cm long, glabrous to pubescent near rachis attachment; rachis 0.9–22.7(–8.59) cm long; floral bracts rounded to triangular, ciliate to partly ciliate or with a few random hairs along margin; stamens with anthers 0.2–0.35(–0.26) mm long, the connective barely expanded at apex, 0.1–0.2 mm in diam, drying black; ovary with 2–6 stigma lobes. Fruits 1.70–4.25 + 0.50–1.75 (–2.53 + 0.99) mm.

Phenology— This species flowers and fruits throughout the year.

Distribution— This species is known from Dominica, Guadeloupe, Martinique, Costa Rica, Panama, Colombia, Ecuador, Peru, and Venezuela in montane woodlands, humid forests, cloud forests, tropical wet forests, primary forests, remnant woods, pastures, and river edges at 0–3,000 m. It has been reported by Callejas (2001) from Nicaragua on the basis of a single specimen, but not confirmed here. A cultivated specimen from Puerto Rico was also examined [Trujillo Plant Propagation Station, 27 Feb 1925, Britton & Boynton 8497 (NY)].

Notes— Arias et al. (2006) mentioned that M. incurva , M. naranjoana , M. sydowii , and M. venzuelana represent a difficult complex of taxa that share many floral similarities. As discussed above, our analysis (CDA), including vegetative features, does not allow for clear separation of these taxa. Therefore we prefer to lump these taxa together and recognize them as a single species, M. incurva , which represents the earliest name available. The simplest feature to distinguish this species from either M. urbanii or M. obtusa would be the supra-basal, acrodromous leaf venation pattern where

the three central veins are united for some distance distal to the petiole insertion.

The specimen of Hahn 1303 in the de Candolle herbarium was chosen as the lectotype of Piper guadeloupense since it was seen by C. de Candolle (based on information from the original publication), and is a representative specimen that displays important features of this taxon.

Additional Specimens Examined— (GenBank accession numbers are

1 2

indicated for collections that were amplified for rpl32-trnL, psbj-petA region1, 3 psbj-petA region 2, and 4 ITS).

COLOMBIA. Antioquia: Municipio Frontino, Corregimiento La Blanquita, 890–900 m, 6 45 0 N, 76 25 0 0 00 W, 13 Jul 1988, Callejas et al. 6720 ( MO). Choco: Río Chintado , 6 Feb 1967, Duke 9843 ( GH, MO, NY) GoogleMaps ; 37 km W of Las Animas , 180 m, 10 Jan 1979, Gentry & Renteria 23957 ( MO) ; Mecana, 14 Jan 1984, Juncosa 1926 ( MO, NY), 1 HQ731611 View Materials , 3 HQ731577 View Materials (clone 1), 3 HQ731578 View Materials (clone 2), 4 HQ731555 View Materials . Valle: Río Calima , 0–5 m, 11 Mar 1944, Cuatrecasas 16850 ( F + 2, US) ; trail from Las Cascadas to Alto de Yundo , Valley of Río Dagua, 500–600 m, 3 36 0 N, 76 44 0 W, 7 Jul 1984, Gentry & Monsalve 48247 ( MO, NY), 4 HQ731556 View Materials GoogleMaps ; 2–13 km N of Anchicaya , 300–490 m, 3 35 0 N, 76 42 0 W, 9 May 1984, Luteyn et al. 10372 ( G, MO, NY, US), 2 HQ731596 View Materials GoogleMaps .

COSTA RICA. Alajuela: 7 km E of Ciudad Quesada, 500 m, 10 22 0 N, 84 24 0 W, 17–18 May 1968, Burger & Stolze 4945 ( F); 850 m, 15 Aug 1977, Croat 43588 ( BM, MO); Reserva Forestal San Ramón , 800–900 m, 10 13 0 N, 84 37 0 W, 15 Nov 1988, Gomez-Laurito & Mora 11745 ( F); Reserva Forestal San Ramón , 10 12 0 53 00 N, 84 36 0 28 00 W, 2 Nov 1986, Herrera et al. 178 ( BM); Reserva Forestal de San Ramón , 10 12 0 53 00 N, 84 36 0 28 00 W, 3 May 1987, Herrera 619 ( BM); Upala, 350 m, 30 Mar 1988, Herrera 1644 ( BM, MO); Upala-Bijagua, 600–800 m, 10 42 0 45 00 N, 85 00 0 45 00 W, 17 Jul 1988, Herrera 2119 ( BM, F, MO); San Carlos-Fortuna , 700 m, 10 26 0 30 00 N, 84 42 0 35 00 W, 22 Nov 1992, Herrera 5569 ( MO); Río Toro, 1,550 m, 10 12 0 N, 84 19 0 W, 3 Sept 1972, Lent 2805 ( F); ridge top between Río Gorrión and Río Toro, 1,700 m, 10 12 0 N, 84 19 0 W, 7 Oct 1972, Lent 3003 ( F); Arenal Volcano, 600 m, 11 Apr 1974, Lent 3869 ( F); San Ramón , 900 m, 10 13 0 15 00 N, 84 33 0 25 00 W, 21 Apr 1994, Lépiz & Morales 307 ( NY); 13 km W of Fortuna on road to Arenal Dam, 500–550 m, 10 29 0 N, 84 43 0 W, 29 Apr 1983, Liesner et al. 15232 ( MO, WIS); San Ramón , 800–900 m, 10 13 0 10 00 N, 84 35 0 35 00 W, 28 Apr 1993, Morales et al. 1386 ( MO), 1 HQ731604 View Materials , 4 HQ731563 View Materials ; Along Río Zapote, 1 km from Zapote, 500 m, 10 45 0 N, 85 05 0 W, 5 Jan 1975, Taylor 18140 ( NY, US); 440 m, 11 Aug 1975, Utley & Utley 3258 ( F). Cartago: Refugio Nacional de Vida Silvestre, 1,300 –1,450 m, 14 Feb 1992, Almeda & Daniel 7006 ( MO); near Río Tuis along the road from Turrialba to Moravia, 730 m, 9 51 0 N, 83 34 0 W, 19–20 Dec 1966, Burger & Ramirez 3977 ( F); 10 km S of Tapantí on E slope above Río Grande de Orosi , 1,600 m, 9 42 0 N, 83 47 0 W, 10–24 Jun 1968, Burger & Stolze 6107 ( F); forest on ridge between Quebrada Casa Blanca and road to Transito, 1,400 m, 9 47 0 N, 83 47 0 W, 10 Aug 1984, Grayum & Jacobs 3729 ( BM, MO), 2 HQ731589 View Materials , 3 HQ731571 View Materials , 4 HQ731562 View Materials ; above Navarro del Socorro, 1,300 m, 9 47 0 N, 83 53 0 W, 30 Jan 1987, Greig & Berrocal 356 ( BM); Río Grande de Orosí, Tapantí , 1,200 m, 19 Jan 1964, Jimenez 1604 ( F, NY); 0.5 km N of Tapantí, 1,340 m, 3 Sept 1967, Lent 1248 ( F, MO, NY); 3 km E of Cachi, 20 Apr 1969, Lent 1585 ( F, MO, NY); Cerro Doan, 1,500 m, 17 Feb 1972, Lent 2350 ( F, G, GH, MO, NY, US); forest above Río Taus, Selva, 950 m, 9 47 0 N, 83 45 0 W, 2 Aug 1972, Lent 2751 ( F); woods beside Río Pejibaye, 780 m, 9 47 0 N, 83 44 0 W, 18 Nov 1972, Lent 3041 ( F, MO); 2 km E of Cachi, 1,450 m, 9 49 0 N, 83 47 0 W, 16 Dec 1972, Lent 3120 ( F); hills 4 km SW of Tobosi, 1,700 m, 9 50 0 N, 84 01 0 W, 11 Feb 1973, Lent 3167 ( F); beside Río Gato, 800 m, 9 47 0 N, 83 41 0 W, 29 Dec 1973, Lent 3729 ( F); woods along Río Naranjo, 2 km W of Orosi, 1,400 m, 9 48 0 N, 83 52 0 W, 16 Jan 1977, Lent 4052 ( F); 12 km S of Turrialba, 4 km SE of Pejibaye along Río Gato, 700 m, 9 48 0 N, 83 42 0 W, 16–17 Apr 1983, Liesner 14319 ( MO); Turrialba, 300–400 m, 9 58 0 20 00 N, 83 27 0 10 00 W, 22 Nov 2000, Mora & Rojas 1745 ( G + 3); forét de Juan Vinas, 1,135 m, 25 Jan 1890, Pittier & Durand 1871 ( G + 2); El Muneco, 1,400 –1,500 m, 6–7 Mar 1926, Standley & Valerio 50957 ( US). Guanacaste: W slope of Cerro Nubes- 2 km E of Silencio de Tilaran, 900 m, 10 28 0 N, 84 53 0 W, 26 Jan 1985, Grayum et al. 5010 ( MO), 2 HQ731591 View Materials ; 500 m, 3 Jun 1972, Lent 2567 ( F); Parque Nacional Guanacaste, 700–800 m, 10 59 0 26 00 N, 85 25 0 40 00 W, 18 Mar 1993, Morales et al. 1251 ( MO); Quebrada Serena, 700 m, 27 Jan 1926, Standley & Valerio 46172 ( US). Heredia: Parque Nacional Braulio Carrillo, 1,000 m, 10 16 0 38 00 N, 84 04 0 57 00 W, 22 Nov 1992, Boyle et al. 1186 ( MO); Parque Nacional Braulio Carrillo, Puesto El Ceibo , 520 m, 10 19 0 58 00 N, 84 04 0 44 00 W, 9 Dec 1992, Boyle 1435 ( MO), 1 HQ731603 View Materials , 3 HQ731570 View Materials , 4 HQ731560 View Materials ; plains of Santa Clara, 100 m, 8 May 1903, Cook & Doyle 616 ( US); Vara Blanca de Sarapiqui, 1,500 –1,750 m, Jul-Sept1937, Skutch3151 ( A, K, MO, NY, US); Cordillera Volcánica Central, 1,539 m, 10 13 0 45 00 N, 84 07 0 00 00 W, 8 Apr 2005, Soto & González 740 ( G). Limon: between Siquerres and the Río Pacuare, 50–100 m, 10 5 0 N, 83 29 0 W, 20–22 Dec 1969, Burger & Liesner 6975 ( F); Reserva Forestal Cordillera, 400 m, 10 11.5 0 N, 83 49 0 W, 17 Jun 1988, Hammel et al.17054 ( BM, MO); Cordillera de Talamanca, 100–200 m, 10 01 0 N, 83 24 0 W, 14 Oct 1988, Herrera 2185 ( BM, F, MO), 1 HQ731601 View Materials , 2 HQ731587 View Materials , 3 HQ731568 View Materials ; Canton de Guapiles, 1,100 m, 10 06 0 40 00 N, 83 59 0 40 00 W, 26 Feb 1990, Herrera & Schik 3834 ( BM, K, MO); 4 km NW of El Carmen, 10 m, 10 13 0 N, 83 28 0 W, 17 Mar 1973, Lent 3269 ( F); Cerro Coronel, 20–170 m, 10 41 0 N, 83 38 0 W, 16–23 Jan 1986, Stevens 23751 ( MO); Cerro Coronel, 80–120 m, 10 41 0 N, 83 38 0 W, 14 Mar 1987, Stevens et al. 24805 ( MO). Puntarenas: Finca las Alturas, 1,300 –1,400 m, 17 Feb 1991, Almeda et al. 6700 ( NY); Las Alturas Biological Station NE of San Vito , near Panama border, 1,530 m, 9 Jul 1993, Bornstein 180 ( SEMO) and Bornstein 181 ( SEMO); Monteverde, 1,450 –1,550 m, 10 47 0 N, 84 50 0 W, 21 Aug 1984, Gentry & Haber 48748 ( BM); Monteverde, 1,550 m, 10 48 0 N, 84 50 0 W, 12 Jul 1990, Gentry et al. 71575 ( MO), 1 HQ731602 View Materials , 2 HQ731588 View Materials , 3 HQ731569 View Materials ; Monteverde, 1,500 m, 10 20 0 N, 84 50 0 W, 24 Dec 1985, Haber 3910 ( BM, MO + 2, WIS); SW corner of Monteverde reserve, 1,100 –1,250 m, 10 16.5N, 84 47 0 W, 4 May 1986, Hammel et al. 15108 ( BM + 2, MO), 3 HQ731566 View Materials (clone 1), 3 HQ731567 View Materials (clone 2); La Pitahaya, 1,200 m, 10 15 0 10 00 N, 84 41 0 20 00 W, 27 May 1996, Herrera 9022 ( K); beside Río Bella Vista near Las Alturas, 1,300 m, 8 56 0 N, 82 51 0 W, 8 Jul 1972, Lent 2690 ( F, MO, NY); Monteverde, 1,550 m, 10 20 0 N, 84 50 0 W, 31 May 1988, Maffia 277 ( F); 6 km S of San Vito de Java , 1,220 m, 13 Aug 1967, Raven 21743 ( F). San José : Los Angeles de Siquirres, 3 km W and 1.9 km S from Guayacán, 1,000 m, 6 May 1983, Gómez et al. 20553 ( BM, WIS); Dulce Nombre Arriba-San Carlos , 650 m, 23 Jun 1966, Jimenez 4058 ( BM, F, WIS); between Río Para Blanca and Bajo la Hondura, 1,660 m, 10 03 0 N, 84 01 0 W, 22 Oct 1972, Lent 3027 ( F); vicinity of El General, 1,130 m, Feb 1936, Skutch 2593 ( A, GH, MICH, MO, NY, US); buissons á Las Vueltas, Tucurrique, 635 m, Nov 1898, Tonduz 12790 ( G + 5, GH, NY, P). Locality unknown: 1,520 –1,580 m, 15 Dec 1976, Dryer 1087 ( F); 1,400 m, Jan 1941, Valerio 1354 ( F).

DOMINICA. St. David : trail between Boeri Lake and Fresh Water Lake above Laudat , 760–915 m, 9 Mar 1993, Hill 24688 ( NY, US), 1 HQ731600 View Materials , 2 HQ731584 View Materials , 3 HQ731565 View Materials . St. George : south slope of Morne Macaque, Micotrin, 760–820 m, 28 May 1964, Ernst 1485 ( BM, US) ; south slope of Morne Macaque , Micotrin, 762–820 m, 25 Jun 1965, Ernst 1774 ( US) ; Fresh Water Lake area , south face Micotrin , 730–790 m, 1 Jun 1967, Wasshausen & Ayensu 338 (B-photo, US). St. Joseph: Point Lolo, 305 m, 26 Oct 1964, Nicolson 1852 ( A, US) ; En Haut Jean , 760–850 m, 24 Jun 1965, Webster 13507 ( US). St. Peter: Syndicate Estate, 26 Mar 1987, Whitefoord 5621 ( A, BM, MICH, US), 1 HQ731599 View Materials , 2 HQ731583 View Materials . Locality unknown: Apr 1882, Eggers 1006 ( K) ; 650 m, 2 Oct 1983, Whitefoord 3853 ( A) .

ECUADOR. Cotopaxi: Parque Nacional Cotopaxi, 3,000 m, 0 40 0 S, 78 30 0 W, 28 Sept 1983, Mantilla 54 ( NY). Esmeraldas: Quininde, Bilsa Biological Station, 400–600 m, 00 21 0 N, 079 44 0 W, 17 Nov 1994, Clark & Mora 264 ( MO). Imbabura: Lita, 600 m, 4 Oct 1980, Maas & Cobb 4686 ( NY); Collpi, 840 m, 4 Jun 1949, Solis 12782 ( F). Morona Santiago: Compamento La Playa, 1,050 m, 28 Jan 1981, Gentry et al. 30956 ( MO); Gualaquiza Cantón, Cordillera del Cóndor, 1,470 m, 3 29 0 S, 78 14 0 W, 19 Jul 1993, Gentry 80159 ( MO). Morona Santiago-Zamora: 21 Jul 1960, Mathias & Taylor 5210 ( F). Napo: Río Wai, 300 m, 0 15 0 S, 76 21 0 W, 8 Aug 1980, Brandbyge et al. 32648 ( MO), 2 HQ731594 View Materials , 3 HQ731576 View Materials , 4 HQ731558 View Materials ; San Pablo de los Secoyas, 300 m, 11 Aug 1981, Brandbyge et al. 33464 ( K); along road between Baeza and Tena, 45 km N of Archidona, 2,300 m, 21 Dec 1979, Croat 49545 ( MO, NY) and Croat 49594 ( MO, NY); vicinity of Baeza, 1,900 m, 27 Mar 1972, Dwyer & MacBryde 9624 ( MO). Napo- Pastaza: Borja-Virgilio Davila, 1,700 m, 15–26 Jan 1959, Harling 3910 ( NY). Pichincha: vicinity of Santo Domingo de los Colorados, near Hacienda Gloria Naria, 18 May 1955, Asplund 16400 (B-photo, F, NY); 20 km W of Santo Domingo de los Colorados, 305 m, 19 Oct 1961, Cazalet & Pennington 5081 ( US); 20 km W of Santo Domingo de los Colorados, 305 m, 20 Oct 1961, Cazalet & Pennington 5098 (B-photo, F, K, NY, US); 20 km W of Santo Domingo de los Colorados, 305 m, 29 Oct 1961, Cazalet & Pennington 5181 ( F, K, NY); 3 km S of Santo Domingo, 530 m, 8 Apr 1980, Dodson & Gentry 10366 ( F); 1 km E and 2 km inside of La Independencia at Km 12, La Concordia, Quito, 310–350 m, 29 Jul 1984, Dodson et al. 14604 ( MO), 1 HQ731612 View Materials , 3 HQ731579 View Materials , 4 HQ731561 View Materials . Zamora-Chinchipe: road from Loja to Zamora, km 54, 1,300 m, 4 02 0 S, 78 59 0 W, 18 Apr 1973, Holm-Nielsen et al. 3786 ( F, MO, NY). Locality unknown: 1,705 m, 25 Aug 1960, Grubb et al. 1349 ( K, NY).

GUADELOUPE. Bagatella , 510 m, 1892, Duss 2565 ( NY + 2, US) ; Basse Terre , 640–660 m, 13 Dec 1959, Proctor 20410 ( A) ; Cascade Vancheler , 18 Sept 1939, Questel 4139 ( P, US) ; forêt de Bains Jaunes, 800 m, 19 Jan 1936, Rodriguez 2457 ( NY) ; forêt de Bains Jaunes , 800 m, 6 Aug 1936, Stehlé & Stehlé 995 ( ILL, NY, P + 2) ; forêt de Bains Jaunes , 690 m, 8 Feb 1937, Stehlé & Stehlé 1328 ( ILL + 2, P) ; Chemin de Malanga , 650 m, 27 Jun 1937, Stehlé & Stehlé 1677 ( ILL, NY) ; Bains Jaunes , Bassin of Decantation, 600 m, 27 Jun 1937, Stehlé & Stehlé 2073 ( ILL, NY, P) ;. Locality unknown: 600 m, Collector unknown 728 ( P) ; 17–27 Apr 1979, Howard & Howard 19522 ( A, BM, NY, US) ; 1843, L’Herminier ( P + 2) ; Questel 1366 ( P + 2) ; 12 Jul 1941, Questel 4942 ( US) ; Richard s. n. ( P + 3) ; 680 m, 1 Mar 1937, Stehlé & Stehlé 1628 ( ILL, NY) ; 5 Sept 1944, Stehlé et al. 5632 ( US) .

MARTINIQUE. Pitons du Carbel, 1877/1880, Duss 1333 ( NY + 2, US) ; Montagne Pelee, 8 Jan 1979, Sastre 6517 ( A). Locality unknown: 1854, Belanger 460 ( G, K) ; 1861, Belanger 971 ( P, US) ; Sept 1900, Duss 4477 ( NY) ; Pleé 702 ( P) ; 1773, Ponthieu s. n. ( BM) ; 430 m, 30 Sept 1940, Stehlé & Stehlé 4628 ( US) ; 11 Dec 1942, Stehlé & Stehlé 6018 ( US) ; 580 m, 7 Mar 1946, Stehlé & Stehlé 6620 ( US) .

PANAMA. Bocas del Toro: Campamento la pata del Cedro, 1,525 m, 9 03 0 966 00 N, 82 43 0 931 00 W, 14 Mar 2004, Alfaro & Monro 5519. Chiriquí: cloud forest of Monte Rey , 21 Jul 1971, Croat 15706 ( GH, MO); Ojo de Agua, 1,520 m, 8 51 0 N, 82 45 0 W, 17 Jun 1987, Croat 66300 ( BM, MO); Las Lagunas area W of El Hato del Volcán, 1,400 m, 8 47 0 N, 82 40 0 W, 1,200 m, 23 Aug 1982, Hamilton et al. 938 ( BM); Bambito, 1.6 km SW Cerro Punta, 1,705 m, 26 Jun 1969, Tyson 5645 ( MO); Distrito Boquete, 1,200 m, 5 Feb 1985, Werff & Hardeveld 6527 ( MO). Coclé: vicinity of La Mesa beyond El Valle, 800–900 m, 11 Jul 1987, McPherson 11206 ( BM, MO), 2 HQ731585 View Materials , 4 HQ731559 View Materials . Darién: near Río Setigandi, 540–580 m, 18 Apr 1980, Gentry et al. 28547 ( F, MO, NY); Parque Nacional del Darién on ridge between Río Topalisa and Río Pucuro, 17 km E of Pucuro, 850 m, 8 03.5 0 N, 77 17 0 W, 17 Oct 1987, Hammel et al. 16274 ( MO), 2 HQ731586 View Materials ; along the Sambu River, Feb 1912, Pittier 5562 ( F, GH, NY, US). Panamá: Cerro Campana, 850 m, 11 Mar 1973, Busey 877 ( MO); Cerro Campana, 10 Sept 1970, Croat 12078 ( MO + 2); rainforest on summit of Cerro Campana, 31 Mar 1969, Porter et al. 4952 ( MO). San Blas : Nusgandi, 350 m, 9 19 0 N, 78 55 0 W, 6 Nov 1984, Nevers 4198 ( BM, MO + 2), 2 HQ731590 View Materials , 4 HQ731564 View Materials .

PERU. Huanuco: La Divisora between Tingo Maria and Aguaytia, Cordillera Azul , 1,500 –1,600 m, 3 Jun 1983, Gentry et al. 41465 ( MO, NY), 1 HQ731610 View Materials . Loreto: Explorama Inn , 1 km S of Indiana, 130 m, 73 1 0 W 3 30 0 S, 18 Jun 1986, Gentry et al. 54704 ( MO), 1 HQ731613 View Materials GoogleMaps ; Maynas Explorama Inn , 130 m, 3 30 0 S, 73 02 0 W, 12 Feb 1987, Gentry et al. 55636 ( MO), 4 HQ731554 View Materials GoogleMaps ; Maynas Sargento Lores , 95–110 m, 7 Jan 1983, McDaniel & Rimachi 26641 ( MO, NY), 1 HQ731609 View Materials , 2 HQ731595 View Materials , 4 HQ731553 View Materials ; Maynas Indiana , 120–140 m, 16 Aug 1988, McDaniel & Rimachi 30180 ( MO), 1 HQ731614 View Materials , 3 HQ731580 View Materials ; Maynas Region de Amazonas , 100–140 m, 3 20 0 S, 72 55 0 W, 21 Feb 1991, Pipoly et al. 13183 ( MO), 1 HQ731615 View Materials , 3 HQ731581 View Materials . San Martin : Mariscal Caceres, 10–12 m, 20 Jun 1978, Schunke 10272 ( F, MO, NY) GoogleMaps .

VENEZUELA. Anzoátegui: along Río Zumbador, 900 m, 1–2 Mar 1945, Steyermark 61301 ( F). Aragua: Parque Nacional Henri Pittier , 1,500 –1,900 m, 8–11 Mar 1972, Bunting 4712 ( NY); Parque Nacional Henri Pittier, Portachuelo trail, 1,000 –1,120 m, 10 20 0 N, 45 W, 13 Jan 1982, Luteyn et al. 8221 ( NY, VEN-photo); Parque Nacional Henri Pittier , 1,060 m, 9 Sept 1977, Steyermark & Huber 114218 ( NY, VEN-photo). Falcón: below hotel at Cerro Galicia, Sierra de San Luis , 1,420 m, 29 Mar 1984, Berry & Wingfield 4307 (VEN-photo); Sierra de San Luis , 1,450 m, 16 Jul 1967, Steyermark 98896 (B-photo, F, VEN-photo). Lara: 1500 m, Aug 1959, Aristeguieta 3949 ( NY); Jimenez, Parque Nacional Yacambu , 1,500 –1,800 m, 19 Apr 1972, Bunting 4988 ( NY); Distrito Palavecino, 1,100 –1,200 m, 4 Aug 1970, Steyermark et al. 103357 ( US) and 1300–1460 m, Steyermark et al. 103365 ( MO, US); Distrito Iribarren, 1,500 –1,550 m, 11 Aug 1970, Steyermark et al. 103686 ( F). Miranda: Distrito Federal, 1,300 –1,500 m, 12 Oct 1965, Steyermark 94398 ( US); Distrito Federal, 1300–1500 m, 24 Jun 1966, Steyermark & Nevling 95929 ( F). Monagas: 1,300 –1,400 m, 11 Apr 1945, Steyermark 61997 ( F, NY). Portuguesa: 15 km al este de Chabasquén, 67 km al NNO de Guanare, 1,450 –1,520 m, 9 26– 27 0 N, 69 54– 55 0 W, 29–31 Oct 1982, Steyermark et al. 126635 (VEN-photo). Sucre: Peninsula de Paria, Cerro de Humo, 900–1,060 m, 4 Mar 1966, Steyermark 95007 ( F, US). Tachira: vicinity of Mata Mula, N of Delicias on road to Bramon, 1,750 m, 9 11 0 N, 70 09 0 W, 26 Jul 1979, Steyermark & Liesner 118729 (VEN-photo). Trujillo: 13 km SE of Bocono, 1 km W of Guaramecal, 1,600 m, 9 11 0 N, 70 09 0 W, 16 Mar 1982, Liesner et al. 12904 ( MO, VEN-photo), 1 HQ731617 View Materials ; Grande entre Boconó y El Batatal, 1,800 m, 5 Sept 1966, Steyermark & Rabe 97369 ( US). Yaracuy: Distrito Nirgua-Distrito San Felipe border, Cerro la Chapa, 7 km N of Nirgua, 1,200 –1,300 m, 10 12 0 N, 68 35 0 W, 21 Oct 1982, Davidse et al. 20836 ( MO, VEN-photo), 1 HQ731618 View Materials , 2 HQ731597 View Materials ; Sierra de Aroa, 800–1,000 m, 10 26 0 N, 68 49 0 W, 30 Mar 1980, Liesner & Gonzalez 9683 ( MO, VEN-photo), 1 HQ731616 View Materials ; Cerro la Chapa, 1,200 –1,400 m, 9–10 Nov 1967, Steyermark et al. 100224 (B-photo, F, VEN-photo).

2. MANEKIA OBTUSA (Miquel) Arias, Callejas & Bornstein ,

Novon 16: 206. 2006. Artanthe obtusa Miq., Syst. Piperac.

416. 1844. Piper fluminense C. DC., Prodr. 16(1): 308.1869,

nom. superfl. for Artanthe obtusa . Sarcorhachis obtusa

(Miq.) Trel., Contr. U.S. Natl. Herb. 26: 118. 1929.—

TYPE: BRAZIL. F. Sellow s. n. (holotype: B, destroyed).

Piper convallariodorum C. DC., Bull. Herb. Boiss. , ser. 2, 1: 356.

1901. Sarcorhachis convallariodorum (C. DC.) Steyermark ,

Pittiera 3: 32. 1971.— TYPE: BRAZIL. Serra Ouro Preto, in

locis umbrosis ad rivulos, 1,050 m, 7 Jan 1894, C. A. W.

Schwacke 10227 ( holotype: G, photo at GH!, US!; possible

isotypes: R, RB).

Sarcorhachis obtusa (Miq.) Trel. var. cordata Yuncker, Bol. Inst.

Bot. (Sao Paulo) 3: 134, Fig. 117. 1966.— TYPE: BRAZIL.

Paraná: Jacarei, 14 August 1914, I. G. Jönsson 862a (holo-

type: S!, ILL! [fragment]; isotypes: LE, US ).

Stems with dark brown to black pellucid dots occasionally present, glabrous; internodes 1.5–15.2(–5.09) cm long. Prophylls 0.3–2.7(–0.81) cm long, drying dark brown. Leaves of monopodial axes with petioles 1.9–6.2(–3.63) cm long, vagination stopping 0.1–1.7(–0.32) cm before blade; laminae 3.9–11.7 + 3.4–10.9(–8.33 + 6.96) cm, symmetrical, ovate to broadly ovate, glabrous, base symmetrical, angle obtuse, shape truncate to cordate, apex angle acute, shape straight with slightly rounded tip (not especially sharp-pointed), dark green above, pale green below, chartaceous but drying coriaceous, pellucid dots reddish black to black, visible above and below, or above only; venation basal acrodromous with primary veins 7 or occasionally 5 or 9, often elevated on lower surface, all of the veins separate distal to petiole insertion. Spikes cream-colored in flower, usually solitary with peduncles 1.8–6.3(–3.73) cm long, mostly glabrous with few hairs near rachis attachment; rachis 2.2–11.3(–7.04) cm long; floral bracts rounded-triangular, ciliate to partly ciliate; stamens with anthers 0.35–0.60(–0.48) mm long, the connective noticeably expanded at apex to 0.5 mm in diam, drying black; ovary with 3–5 stigma lobes. Fruits 1.25–2.85 + 1.05–1.70 (–1.96 + 1.30) mm.

Phenology— Flowering specimens have been collected from October through January; fruiting material from February through June.

Distribution— This species is known from southeastern Brazil, rain forests at 50–1,500 m.

Notes— This species is the only member of Manekia where the petiole vagination stops before reaching the blade, an easy feature to examine. It is similar to Manekia urbanii in having the primary leaf veins separate distal to the petiole insertion, but can be distinguished based on the leaves being significantly larger in size as well as the obvious difference in distribution. If floral material is available, the large anthers with the noticeably expanded connective at the apex are also useful for distinguishing this species. According to notes on a few herbarium specimens, the fruiting spikes have a red coloration (when young?), which would also be distinct in the genus.

Additional Specimens Examined— (GenBank accession numbers are indicated for collections that were amplified for 1 rpl32-trnL, 2 psbj-petA region1, 3 psbj-petA region 2, and 4 ITS).

BRAZIL. Espirito Santo: Municipio de Santa Teresa, Vila nova, terreno do Gino , 640 m, 11 Mar 2001, Kollmann 3601 ( RB). Minas Gerais: Serra de Ouro Preto , 1962, Damazio 54528 ( RB). Paraná: Municipio de Cerro Azul , 25 15 0 S, 48 45 0 W, 6 Dec 1983, Callejas et al. 1856 ( NY), 1 HQ731606 View Materials , 3 HQ731574 View Materials , 4 HQ731557 View Materials GoogleMaps ; Municipio de Cerro Azul , 24 45 0 S, 48 45 0 W, 7 Dec 1983, Callejas et al. 1869 ( NY), 1 HQ731607 View Materials GoogleMaps ; Cerro Azul , 11 Feb 1960, Hatschbach 6743 ( NY) ; Río do Cedro, Municipio Guaraqueçaba , 20–50 m, 21 Nov 1968, Hatschbach 20367 ( C, F) ; Faisqueira, Municipio Antonia , 14 Nov 1980, Hatschbach 43277 ( C, NY, US) , 1 HQ731605 View Materials , 2 HQ731592 View Materials , 3 HQ731572 View Materials (clone 1), 3 HQ731573 View Materials (clone 2) ; Estrada da Graciosa, Ferradura, Municipio Morretes , 20 Jan 1987, Hatschbach & Silva 50877

( US), 1 HQ731608 View Materials , 3 HQ731575 View Materials ; Río Turvo , Municipio Dr. Ulysses , 2 Dec 1999, Hatschbach et al. 69805 ( G + 2). Rio de Janeiro : Paraty, estrada Paraty- Cunha, 13–15 km da entrada de Paraty, 11 Oct 2004, Carvalho-Silva et al.324 ( RB) ; Macaé , estrada Gliceri-Trajano de Morais, 19 May 2005, Carvalho-Silva et al. 372 ( RB) ; Resende , Visconde de Mauá, estrada Mauá-Resende, 22 20 0 7 00 S, 44 32 0 32 00 W, 1,055 m, 7 Dec 2006, Mansano & Marquete 06–354 ( RB) GoogleMaps ; Paraty , subindo o Rio Corisquinho, 240 m, 11 May 1994, Marquete 1782 ( RB) ; Petrópolis , vale das Videiras, morro da Cuca, 1,500 m, 17 Dec 1973, Martinelli 141 ( RB) ; Itatiaia , Parque Nacional de Itatiaia , lado direito da ponte do Rio Taquaral, em direção ao Centro de Visitantes, 18 Mar 2006, Monteiro et al. 156 ( RB) ; Itatiaia , Parque Nacional de Itatiaia, 2 Dec 2006, Monteiro & Gienerine 225 ( RB) ; Paraty , morro do Corisquinho , 480 m, 10 Mar 1994, Ribeiro et al. 17 ( RB). Santa Catarina : Municipio Blumenau, area virgem do Parque Nacional da Serra do Itajaí, 27 05 0 49 00, 26 00 S, 49 08 0 21.67 00 W, 660 m, 23 Nov 2009, Cadorin et al. 680 ( RB) ; Municipio Florianópolis, morro da Lagoa , 27 34 0 36.00 00 S, 48 28 0 32.02 00 W, 444 m, 23 Feb 2010, Cadorin et al. 1426 ( RB) ; Sáo Jaáo do Sul , 29 Jan 1976, Hagelund 10024D ( C) ; Sáo Jaáo do Sul , 21 Jan 1977, Hagelund 11072 ( C) ; Rodovia Indaial- Itaiópolis, 21 Jul 1994, Hatschbach & Poliquesi 60976 ( C) ; Municipio Pouso Redondo, Alto Pombinhas, 27 16 0 14.00 00 S, 50 02 0 30.00 00 W, 475 m, 7 May 2010, Korte & Kneiss 3151 ( RB) ; Novo Horizonte , Lauro Muller, 400 m, 24 Oct 1958, Reitz & Klein 7508 ( NY) ; Serra de Matador , Río do Sul, 500 m, 29 Dec 1958, Reitz 6076 ( NY) ; Sanga de Areia , Jacinto Machado, 200 m, 30 Oct 1959, Reitz & Klein 4141 (B-photo, F, G, K, NY, US) ; Municipio Taió, Fazenda Tarumã, 27 00 0 01.06 00 S, 50 07 0 46.01 00 W, 812 m, 24 Feb 2010, Schmitt et al. 1534 ( RB) ; Jacinto Machado Fundao do Tigre Preto , 3 Apr 1980, Waechter 47264 ( RB). Sao Paulo: 14 Nov 1947, Kuhlmann 1548 ( NY) ; Lorena Piquete , 2 Jun 1950, Kuhlmann 2371 ( NY, RB) ; Serra de Caracol , 5 Mar 1874, Mosen 1690 ( ILL, NY, P) ; Municipio de Ubatuba , estrada da Casa da Farinha, 9 Oct 1989, Ribeiro et al.753 ( RB) ; estrada para Caraguatuba , 750–820 m, 19 Sept 1973, Sucre 10226 ( RB + 3) ; Iporanga , núcleo Bulha D’água, trilha da caverna do Sr. Bento, 24 20 0 18.2 00 S, 48 29 0 58.2 00 W, 21 Apr 2003, Urbanetz et al. 126 ( RB). Locality unknown: Konus s. n. ( NY) ; 21 Feb 1955, Rambo 46015 ( NY) ; 1827, Riedel s. n. ( C, US) ; 1870, Riedel 44 ( K) ; Riedel 285 ( P) .

3. MANEKIA URBANII Trel., Repert. Spec. Nov. Regni Veg. 23:

313. 1927.— TYPE: HAITI. La Hotte, Morne Vandervelde,

700 m, 2 Dec 1925, E. L.Ekman 5242 (holotype: S; isotype: ILL!) .

Stems with black pellucid dots occasionally present, glabrous; internodes 1.2–8.5(–2.82) cm long. Prophylls 0.6 cm long, drying dark brown. Leaves of monopodial axes with petioles 0.8–2.6(–1.5) cm long, vagination extending onto blade 0.1–0.4(–0.2) cm; laminae 4.1–8.1 + 3–6.6(–5.79 + 4.71) cm, symmetrical, ovate, glabrous, base symmetrical, angle obtuse, shape rounded to slightly cordate, apex angle acute, shape straight to more commonly acuminate, dark shiny green above, pale green below, chartaceous but drying coriaceous, pellucid dots light brown to reddish brown, visible above and below; venation basal acrodromous with primary veins 5, usually elevated on lower surface, all of the veins separate distal to petiole insertion. Spikes most commonly paired with peduncles 0.4–2(–1.15) cm long, glabrous; rachis 0.6–2.7(–2.07) cm long; floral bracts rounded-triangular, usually ciliate; stamens with anthers 0.2–0.25(–0.24) mm long, the connective barely expanded at apex, ca. 0.1 mm in diam, drying black; ovary with 3–4 stigma lobes. Fruits 1.15–2.55 + 0.60–0.85(–1.61 + 0.73) mm.

Phenology— The few available herbarium specimens indicate that this species flowers from December to perhaps early March, with fruits evident from January to March.

Distribution— This species is endemic to Haiti on limestone cliffs at 700–1,200 m.

Notes— This species is most readily distinguished by the small leaves and short spikes, usually paired. Apparently it is different from other species of Manekia as it flowers while scrambling over exposed rock surfaces instead of climbing up tree trunks into the canopy. The current status of M. urbanii is unknown as it hasn’t been collected, to our knowledge, since 1983. A significant amount of deforestation has occurred in Haiti since then, but the area of Massif de la Hotte is relatively remote, so this taxon may still be extant.

Additional Specimens Examined— HAITI. Sud: Massif de la Hotte, 31 Mar 1927, Ekman 7941 ( ILL) ; Massif de la Hotte, Morne Descassiers, 1,200 m, 28 Jan 1928, Ekman 9525 ( ILL, G, GH, US + 2) ; Massif de la Hotte, 18 km N de Camp Perrin , en la carretera a Beaumont y Jérémie, 18 22 0 N, 73 53 0 W, 850 m, 11 Mar 1983, Zanoni et al. 25681 ( JBSD, MO) GoogleMaps .