Monstera tuberculata Lundell, 1939

51. Monstera tuberculata Lundell View in CoL , Lloydia 2: 78, t. 1. 1939. ( Figs. 88 View FIGURE 88 , 89 View FIGURE 89 )

Type: — BELIZE. El Cayo District, Valentin , June 1936, C.L. Lundell 6238 ( holotype MICH-MICH1115586!, isotype GH!) .

Nomadic vine, pendent habit. SEEDLINGS: filiform. JUVENILE PLANTS: root climbers; stems light-green to dark, smooth, flattened; internodes 3–6 cm long, 3–5 mm diam.; petiole not visible, dark green, smooth, 3–6 cm long; blades obovate or orbicular, subcordate to truncate at base, acuminate at apex, coriaceous, variegated, 5–10 × 4–9 cm, appressed to the phorophyte; fenestrations absent. ADULT PLANTS: root climbers; stems dark-green to light-green, smooth, cylindrical or dorsoventrally compressed and slightly sulcate on one side, with caducous epidermis; internodes 5–15 cm long, 1–2 cm diam., 5.0–7.5 times longer than wide; anchor roots dark-brown; feeder roots corky; petiole light or dark-green, smooth, 2–6 cm long, sheathed to base of the geniculum, the sheath apically prolonged in a free ligule, 3–6 cm long; petiole sheath dry-persistent for several days, leaving fibrous residues; geniculum smooth, flattened adaxially, convex abaxially, 5–10 mm long; blades ovate, cordulate to cordate at base, obtuse to short-acuminate at apex, coriaceous, drying blackish, yellowish or grayish, 5.5–20.0 × 4–12 cm, 1.4–1.9 times longer than wide, not decurrent on the geniculum; midrib convex on both surfaces, drying blackish abaxially, primary lateral veins 4–7 per side, obscure adaxially, prominent abaxially, drying yellowish or blackish; secondary veins reticulate and prominent, wavy throughout its length when dry; collective veins slightly visible; fenestrations absent or rarely present; margins entire. INFLORESCENCES on pendent stems, 1–10 simultaneously at flowering time, arranged in the leaf axils; peduncle smooth, 2–5 cm long, 2–6 mm diam.; spathe acuminate to long-acuminate, green externally during development, yellowish green or creamy white externally and white internally at anthesis, marcescent or deciduous after anthesis, 7–10 × 6–8 cm, up to 3 cm longer than the spadix; spadix white during development, yellow at anthesis, 4–7 cm long, 1.5–3.5 cm diam.; basal sterile flowers 4–6 mm long, with a yellowish stigmatic secretion; fertile flowers 5–9 mm long; stamens 1–5 mm long, with laminar filaments; anthers 1.5–2.0 mm long; ovary acute at base, ribbed, 3–5 × 2–3 mm; style pentagonal or hexagonal, 5–6 × 3.5–5.0 mm; stigmaphore thickly conical and recurved, 3–6 mm long; stigma linear, with a rusty red stigmatic secretion; berries with a yellowish-green stylar cap during development, mature stylar cap dark-green; pulp white; seeds brown with white dots, oblong, 6–9 mm long.

Distribution and ecology: —From México (Oaxaca) to West of Panamá, at 0‒1100 m. It lives in Tropical moist forest and Tropical wet forest life zones; primary and secondary forest.

Phenology: —Flowering has been recorded in February, April, June-July and November. Fruit in May-August.

Discussion: —The species, a member of sect. Echinospadix , is characterized by its silvery blotched juvenile leaves, appressed-climbing high growing adult plants with the flowering branches pendent, elongated internodes, short petioles with a ligule that may extend beyond base of the blade, the broadly ovate, usually subcordate acuminate blades with a short peduncle, spathe about as broad as long and white cylindroid spadix.

The two areas have two distinct varieties: Monstera tuberculata var. tuberculata in Mexico and Guatemala is known from elevations of 50–150 m and M. tuberculata var. brevinoda in Nicaragua to Panama ranges from 0– 150 m. Monstera tuberculata var. tuberculata in Belize ( Brewer 7635, MO), have adult leaf blades with abundant impressed glands on the underside, and in Mexico, in the region of Veracruz ( Ceja et al. 1881, XAL) they present leaves with a few fenestrations, a characteristic never observed in the populations of Costa Rica and Panama ( Cedeño-Fonseca et al. 2022).

Additional specimens examined: — MEXICO. Campeche: Calakmul. A 86 km al S de Ejido Constitución, 17°51’00”N 090°05’26”W, 218 m, 28 mayo 1998, E.M. Martínez et al. 30976 (MO!); Chiapas: Tropical rain forest remnant along Rio Pichucalco, richly hanging with lianas and epipytic bromeliads and orchids, 3 km NE of Pichucalco on road to Villahermosa, 17°32’N 093°04’W, 200 m, 2 Jul 1969, B. Marcks & C. Marcks 906 (MO!); Ocosingo. Límite entre la comunidad Lacandona y el monumento Bonampak, 16°43’32”N 091°04’38”W, 326 m, 10 agosto 2002, G. Águilar et al. 2036 (MEXU!, MO!); Zona urbana a 1.46 km del centro de Frontera Corozal al SO, 16°48’12”N 090°53’27”W, 123 m, 02 junio 2004, G. Águilar et al. 10390 (MEXU!, MO!); Entrada de Velasco Viejo a 4 km de la carretera, 16°58’12”N 091°13’48”W, 449–461 m, 30 marzo 2002, G. Águilar et al. 399 (MEXU!, MO!); Rapids on Río Santo Domingo at Santo Domingo. Forested river bank, 17°01’58”N 091°24’53”W, 490 m, 13 May 1982, G. Davidse et al. 20456 (MO!); Ostuacán. Two mi S of Chiapas border along highway 195. Disturbed area beside stream, 17°33’03”N 092°59’47”W, 80 m, 3 Jul 1977, T.B. Croat 40085 (MO!); Pichucalco. Along stream 10 km NE of Pichucalco, 17°32’56”N 093°03’39”W, 150 m, 10 May 1973, D.E. Breedlove 35047 (MO!); Along stream 11 km ENE of Pichucalco, 17°33’36”N 093°03’00”W, 100 m, 5 Oct 1980, D.E. Breedlove 45904 (CAS); Oaxaca: Mpio. Sta María Chimalapa: cafetales cerca de Sta. María; cefetales con Vochysia, Andira galeottiana, Termninalia, Enterolobium schomburgkii, Cassia moschata, Hymeneae,Apeiba, Guazuma, Didymopanax, Inga, Mosquitoxylum; suelos profundos, pendientes ligeras o fuertes, 16°55’N 094°40’W, 300 m, 6 May 1985, H. Hernández 1156 (MO!); Uxpanapa Region, along gravel road E of Sarabia on road to Uxpanapa, 4.2 mi NE of Río Corte. On and around karst limestone formations along road. Veg. type: ”selva alta perennifolia”, 17°09’N 095°49’W, 90 m, 18 Jan 1987, T.B. Croat & D.P. Hannon 63189 (MO!); Tabasco: Tacotalpa. Cerro del Madrigal, Km 7 ( 4.2 mi) de la estación Tacotalpa hacia Tapijulapa, 17°30’00”N 092°48’36”W, 90 m, 17 Jun 1983, C.P. Cowan et al. 3950 (MO!); Veracruz: Veracruz: San Andrés Tuxtla, EBT, camino a Laguna escondida, 18°35’02”N 095°04’38”W, 78 m, 06 March 2020, P. Díaz-Jiménez et al. 1456 (HEM!); Playa Escondida, 28 km NE of Catemaco, 18°34’12”N 095°03’36”W, 50 m, Aug 1972, M. Madison 631 (SEL!); 4 km NE of Minatitlan, 18°01’12”N 094°32’24”W, 50 m, Aug 1958, R. King 1114 (MICH, US); Estacion de biologia tropical Los Tuxtlas; selva alta perennifolia; bejuce Herbaceo, sobre Coussapoa purpusii. Inflorescencia verdosa. ”Telefono”, 18°34’N 095°44’W - 18°36’N 095°09’W, 450 m, 18 July 1985, S. Sinaca 143 (MO!, UMO); Entrance road to Playa Escondida near jct. with road between Santecomapan and Montepio, 18°33’36”N 095°03’36”W, 10–15 m, 25 Jul 1978, T. Cochrane et al. 8626 (MO!); Along road between Catemaco and Montepio, 15 km E of junction to Tebanca, Coyame and Nanciyaga, 18°29’50”N 095°02’31”W, 133 m, 6 Mar 2008, T.B. Croat & P. Díaz Jiménez 100323 (MO!); Catemaco. Vic. of Playa Escondida, 10 km N of Sontecomapan. Selva Alta Perennifolia. Evergreen tropical forest, some cleared for pastures, on slopes around and above Hotel Playa Escondida and the beach on the Gulf of Mexico, 18°35’24”N 095°03’00”W, 0–150 m, 30 Jun 1982, M. Nee 24740 (F!, MO!, XAL!); Minatitlán. 2 km N of Uxpanapa on road (Pob. 12) en camino al Pob. 13; arroyo pedregoso en selva de Dialium, Terminalia, Brosimum, Cynometra, Lonchocarpus spp , 17°12’36”N 094°12’36”W, 150 m, 7 Jul 1985, T. Wendt et al. 4886 (MO!); San Andrés Tuxtla. Estación de Biología Tropical ”Los Tuxtlas, Cerro Vigía 5, Lote 67. Selva alta perennifolia, 18°34’N 095°04’W, 450 m, 12Apr 2005, Thorsten Krömer 1921 (MO!); Balzapote, Parcela Escolar, Ejido Balzapote. Plantación de citricos, 18°36’N 095°04’W, 60 m, 4 Aug 2005, T. Krömer et al. 2461 (MO!); Soteapan. Mirador Pilapa, M. Vázquez T. 3448. BELIZE. Cayo: Valentin, 16°45’59”N 089°09’45”W, 2 Jul 1936, C.L. Lundell 6337 (LL!); Valentin, British Honduras, 16°45’59”N 089°09’45”W, June 1936 - August 1936, C.L. Lundell 6238 (MO!); Stann Creek: Watershed of the Cocoa Branch of the Sittee River. Sittee River Forest Reserve, 16°50’57”N 088°34’14”W, 300 m, 10 March 2016, S.W. Brewer & M. Paredes 7635 (BRH, MEXU, MO); Toledo: Vicinity of San Antonio, 16°14’N 088°59’W, 14 Jan 1946, P.H. Gentle 5484 (LL!). GUATEMALA. Alta Verapaz: Sebol, 15°48’22”N 089°56’44”W, 21 Apr 1964, E. Contreras 4453 (LL!); Izabal: Collected 2.5 mi. N. of Río Dulce on gravel road to Petén, 15°40’39”N 089°02’20”W, 100 ft, 08 August 1975, D.B. Dunn & D.G. LeDoux 22029 (MO!, UMO); El Estor. La Llorona, 15°39’18”N 089°10’37”W, 500 m, 23 Aug 1998, M. Véliz 98.6655 (MO!); Petén: Remate Road, 36 km from Tikal, 16°54’10”N 089°46’10”W, 25 Feb 1961, E. Contreras 1998 (LL!); Ca. 5 miles south of entrance to Tikal National Park, mature forest, 17°02’13”N 089°41’10”W, 19 Jun 1973, T.B. Croat 24747 (MO!); Dolores. Dolores, 16°30’45”N 089°24’57”W, 5 Sep 1961, E. Contreras 2855 (LL!); San Luis. San Luis, Km 51 of road, 16°11’59”N 089°26’31”W, 400 m, 10 Jul 1959, C.L. Lundell 16280 (LL!). NICARAGUA. Atlántico Sur: Municipio de Nueva Guinea, Reserva Indio-Maiz, Río Pijibaye entre el Caño Bijagua y el Cerro Chiripa, 11°22’N 084°01’W, 200–350 m, 15 enero 1999, R.M. Rueda et al. 10227 (MO!); Río San Juan: La Lupe, ca. 25 km ENE of Boca de Sabalo; primary forest lightly logged with silvicultural treatment, clay soils, hilly, 11°08’N 084°21’W, 80–120 m, 28 June 1997 – 2 July 1997, J. Salick & E. Stijfhoorn 8358 (MO!); La Palma, Río Santa Cruz, 11°02’N 084°25’W, 50 m, 21 marzo 1985, P.P. Moreno 25492 (MO!); Municipio de el Castillo, Reserva Indio-Maíz, zona de amortiguamiento, estación biológica La Lupe, 11°08’N 084°21’W, 100–200 m, 15 Marzo 1999, R.M. Rueda et al. 10371 (HULE, MO); Reserva Indio-Maíz, Municipio de el Castillo, en los alrededores de la desembocadura del Río Bartola, 10°58’N 084°20’W, 30–50 m, 13 Enero 1997, R.M. Rueda et al. 5667 (MO!, WU!); Municipio de San Carlos, Reserva Esperanza verde, 11°04’N 084°44’W, 60 m, 16 Septiembre 2001, R.M. Rueda et al. 16548 (HULE, MO); Between Río Santa Cruz and Caño Santa Crucita, La Palma; tall evergreen forest, 11°02’N 084°24’W - 11°04’N 084°26’W, 40–60 m, 30 Nov 1984 – 2 Dec 1984, W.D. Stevens et al. 23435 (MO!); Municipio El Castillo, Comunidad Las Maravillas 2 km al Oeste, 11°07’15”N 084°21’04”W, 100 m, 25 Abril 2005, W. Garrido 3917 (HULE, MO). COSTA RICA. Alajuela: San Carlos, Pocosol, Azucena, Concho de Cutris, 80 m, 4 April 1991, (Fl., Fr.), Q. Jiménez et al. 962 (CR!, MO!); Upala, Aguas Claras, P.N. Rincón de la Vieja, Cordillera de Guanacaste, Colonia Blanca, Finca Los Moras, junto a Quebrada Mora, 770 m, 13 June 1991, (Fr.), G. Rivera 1393 (CR!, MO!); Upala, San José o Pizote, San José, 4 km al Noroeste del pueblo, Laguna Las Camelias, 20 m, 19 April 1988, (Fl.), G. Herrera 1848 (CR!, MO!); Upala, Aguas Claras, Sector de Rio Aguas Verdes, 2 Km aguas arriba del puente, falda NE del volcan Santa Maria, 600 m, 12 February 1991, (Fl.), G. Rivera 1052 (CR!); Upala, Prov, Guanacaste-Alajuela, lower montane rainforest, 1500 m, 1 November 1982, (Infer.), L. Gómez 19175 (MO!); Cartago: Turrialba, Along road from Río Pacuare to Grano de Oro on road from Turrialba to Moravia, 600–1200 m, 30 June 1976, (Fr.), T.B. Croat 36574 (MO!); Heredia: Sarapiquí, Las Horquetas, Along Starkey Road”, 50 m, 7 August 1979, (Fr.), W. Stevens 13475 (CR!, MO!); Sarapiquí, La Virgen, S base of Cerros Sardinal, Chilamate de Sarapiquí (N side of Río Sarapiquí), 90 m, 4 July 1985, (Fr.), M.H. Grayum & B.E. Hammel 5544 (CR!, MO!); Sarapiquí, La Virgen, E.B. La Selva, Estación Biológico, La Selva, Along road leading to the reserve, 100 m, 16 August 1987, (Fr.), J.F. Smith 475 (CR!); Sarapiquí, Las Horquetas, La Esperanza, orillas de río San José, Finca propiedad de José Rufino Chaves, 168 m, 31 July 2010, (Fr.), A. Estrada 4624 (CR!); Sarapiquí, La Virgen, P.N. Braulio Carrillo, Estación Biológica Magsasay, Bosque primario y orillas de potreros, Sotobosque, 200 m, 23 June 1990, (Fr.), E. Alcázar 102 (CR!); Sarapiquí, La Virgen, Estacion Magsasay, Bosque primario y orillas de potreros, 200 m, 23 June 1990, (Fr.), E. Alcázar 118 (CR!, MO!); Sarapiquí, Puerto Viejo, Sendero Tres ríos y lindero el Peje, 0 m, 11 July 2003, (Fl., Fr.), R. Kriebel 3618 (CR!); Sarapiquí, Horquetas, Estación Biológica La Selva, 50 m, 24 Junio 2016, (Fr.), M. Cedeño & M. Blanco 907 (USJ!); Sarapiquí, La Virgen, Reserva Biológica La Tirimbina, 150 m, 16 April 2016, (Fr.), M. Cedeño & J. Ley 874 (USJ!); Limón: Talamanca, Cocles, 12 June 1981, (Infer.), R. Alfaro & R. Ocampo 27 (CR!); Pococí, Rita, Cariari de Pococí, Palmitas, El Triángulo, Las Brisas, finca de Guillemo Acosta , 30 m, 18 April 1997, (Fl.), A. Estrada et al. 743 (CR!); Talamanca, Cahuita, Punta de Riel, 4 June 1985, (Infer.), R. Soto 2128 (CR!); NW 0982, 5 km rd Cahuita Bribri, Sea level, flat land periodically flooded, pasture with shade tres, 1 February 1984, (Infer.), T. Pennington 11459 (CR!); Talamanca; R.V.S. Gandoca-Manzanillo, Trail between the town of Manzanillo and Punta Mona, Old secondary growth with old cocoa crops by the coast, 10 m, 3 August 1994, (Fr.), W. Alverson & V. Stilio 2580 (CR!); Siquirres, Pacuarito, Carretera a Siquirres, Entre Siquirres y río Pacuarito, 100 m, 17 November 1994, (Fl.), V. Nilsson et al. 572 (CR!); Talamanca, Cocles, 12 June 1981, (Fl.), R. Alfaro & R. Ocampo 26 (CR!); Limón, Río Blanco, Camino a Veragua Rainforest, entre Río Quito y las Brisas, Árboles en potreros, finca de Sr. Antonio Calderón, 118 m, 26 May 2011, (Fr.), A. Cascante et al. 2295 (CR!); Talamanca, Cahuita, R.V.S. Gandoca-Manzanillo, Sendero entre Manzanillo y Gandoca, Aprox. 1, 4 km de la entrada al sendero por el sector de Manzanillo, 10 m, 10 April 2012, (Fl.), A. Estrada et al. 5324 (CR!); Pococí, Rita, Finca La Suerte, 50 m, 10 July 1995, (Fl.), R. Aguilar 4206 (CR!, MO!); Talamanca, Sixaola, Cerro Manzanillo, sobre la fila, 100 m, 5 December 1992, (Infer.), A. Fernandez 477 (CR!); Talamanca, Sixaola, Punta Uva, peñón innominado adyacente a la Quebrada Ernesto, 0 m, 4 May 2001, (Fr.), J. Morales 7972 (CR!); Talamanca, Sixaola, Sendero a Punta Mona, 74 m, 4 April 2011, (Fr.), L. Vargas 4350 (CR!); Talamanca, Along the road S of Limón from the corner at the road to Bomba, 0–50 m, 26 February 1985, (Fr.), C. Taylor 4442 (MO!); Region between Hone Creek and ca. 4–5 km NW on a trail roughly paralleling the Río Carbon (Río Hone), Collection from Cacao and second growth, 60–100 m, 17 January 1974, (Fr.), J. Utley 659 (MO!); Talamanca, Road between Cahuita and Punto [Puerto] Vargas, 0–5 m, 7 July 1980, (Fr.), B.E. Hammel 9152 (MO!); Talamanca, Puerto Viejo, 500 m del cruce de Bribri, 40 m, 5 December 2012, (Fl., Fr.), J. Gómez et al. 15783 (USJ!); Talamanca, Cahuita, Camino detrás de la carretera, Finca La Montaña , 10 m, 11 July 1999, (Fl.), M. Blanco et al. 997 (USJ!); Talamanca, Cahuita, Bosques de Manzanillo, 50 m, 9 January 2017, (Fl.), M. Cedeño et al. 1104 (USJ!); Limón, Talamanca, Bribri, Proyecto ARA, 4 m, 30 September 2018, (Fl.), M. Cedeño et al. 1480 (USJ!). PANAMA. Bocas del Toro: 5 km S of Tiger Key on the mainland near Cacao Boquete just N of Enseñada de Bosquete, 09°10’12”N 082°10’24”W – 09°10’24”N 082°14’48”W, 5–110 m, 24 February 1989, Paul M. Peterson & Carol R. Annable 7064 (MO!); Distrito de Changuinola. File de la Sierpe, cerca de Quebrada Bonyic, 09°20’01”N 082°35’24”W, 111 m, 27 septiembre 2007, R. Carranza et al. 208 (MO, PMA, SCZ); Station Milla 7.5 on Changuinola-Almirante Railroad, along a slender ridge to World War II communications facility, ca. 2 km NW of ruins of U.S. Army Military barracks, 09°21’54”N 082°27’12”W, 0 – 100 m, 03 August 1976, T.B. Croat 38117 (MO!).