Monstera acuminata K.Koch, Index Seminum, 1855

Croat, Thomas B., Cedeño-Fonseca, Marco & Ortiz, Orlando O., 2024, Revision of Monstera (Araceae: Monsteroideae) of Central America, Phytotaxa 656 (1), pp. 1-197 : 16-20

2. Monstera acuminata K.Koch, Index Seminum [Berlin] 4. 1855. ( Figs. 3 , 4 )

Type: — Central America , live plant collected by Warszewicz cultivated at Berlin, K. Koch s.n. (holotype B) .

Monstera karwinskyi Schott, Oesterr. Bot. Z. 9: 99 (1859). TYPE:— Mexico: ¨a Colipa usque Papantla ubicumque frequens, scandens super arbores¨, [April 1841], W.F. Karwinsky [838] (holotype W destroyed, isotype LE-00000997 !, here designated as lectotype).

Monstera belizensis Lundell , Lloydia 2: 76 (1939). TYPE:— Belize, EI Cayo District, 23 June 1936, Lundell 6198 (holotype MICH-1115584 !).

Monstera grandifolia Standl. & Steyerm., Pub. Field Mus. Nat. Hist. Botany 23: 212 (1947). TYPE:— Guatemala, Peten, low forest between Finca Yalpemech and Chinaja, 28 March 1942, Steyermark 45423 (holotype F-1129317–9!, isotype US-1864956 !).

Monstera chiapensis Matuda, Madroño 10: 48 (1949). SYNTYPES:— Mexico, Chiapas, Finca California, Col. Turquia, Escuintla , 150 m., 29 August 1947, E. Matuda 17789 HEM [at Matuda Herbarium]; Finca Esperanza, 10 km. NE of Escuintla, 200 m., Matuda 17786 HEM (at Matuda Herbarium).

Monstera viridispatha Matuda, Rev. Soc. Mex. Hist. Nat. 11: 98 (1950). TYPE:— Mexico, Chiapas, Esperanza, Escuintla , 150–250 m., 7 Dec. 1949, Matuda 18717 (holotype HEM! [at Matuda Herbarium], isotypes NY!, UC-903869!, BH [photo]!).

Nomadic vine, appressed-climbing habit. SEEDLINGS: filiform. JUVENILE PLANTS: root climbers; stem dorsoventrally compressed, smooth, greenish; internodes 3–7 cm long, 3–5 mm diam.; petiole more or less distinct, greenish, smooth, 7–11 cm long, sheathed to base of the geniculum; petiole sheath persistent; blades obovate, cordate at base, acuminate at apex, coriaceous, 9–12 × 7–9 cm, slightly appressed to the phorophyte; fenestrations present or absent. ADULT PLANTS: root climbers; stems dark-green, smooth or verrucose, cylindrical or dorsiventrally compressed; internodes 1.0– 8.5 cm long, 1–3 cm diam., 0.3–2.8 times longer than wide; anchor roots brown or black; feeder roots corky, dark-brown; petiole dark green or greenish, smooth or verrucose at base, sheathed up to half the geniculum, 25–66 cm long; petiole sheath persistent; geniculum more or less evident, smooth, 2.0– 4.5 cm long; blades lanceolate or elliptical, widely truncate to obtuse at base, acuminate at apex, thinly coriaceous, drying blackish or greenish, 35–72 × 15–40 cm, 1.8–2.3 times longer than wide, slightly decurrent on the geniculum (decurrent part 1–2 mm wide); midrib sunken adaxially, raised abaxially, primary lateral veins 6–15 per side, slightly sunken adaxially, raised abaxially, departing midrib at 35–60°, secondary veins inconspicuous, collective veins absent; fenestrations present or absent, 1–6(9) oblong-elliptical perforations when present, that are located on both blade sides between the midrib and margin; margins entire or rarely pinnatilobed. INFLORESCENCES on ascending stems, 1–2 simultaneously at the flowering season, arranged in the leaf axils, rarely in deciduous cataphylls; peduncle smooth, 4–13 cm long; 1.5–2.5 cm diam.; spathe obtuse, length equal to the spadix, bright-green during development, light-green externally and white internally at anthesis, deciduous in fragments after anthesis; spadix white during development, cream at anthesis, 8–12 cm long, 2–3 cm diam., (3.1)4.5–7.0 times longer than wide; basal sterile flowers 5–6 mm long, with a yellowish stigmatic secretion; fertile flowers 5–7 mm long; stamens 2–7 mm long, with laminar filaments; anthers 1.0– 2.5 mm long; ovary quadrangular in longitudinal section, ribbed, 3–4 × 1.5–2.5 mm; style pentagonal or hexagonal, 1.5–2.5 × 1.5–2.5 mm; stigma linear, with a yellowish stigmatic secretion; berries with a stylar cap yellow, green during development; seeds not seen.

Distribution and ecology: — Monstera acuminata ranges from Mexico (Chiapas, Escuintla, Oaxaca, San Luis Potosí, Tabasco, Jalapa, Veracruz) to Guatemala (Escuintla, Izabal, San Marcos, Suchitepequez, Zacapa), Belize (Toledo), Honduras (Comayagua, Copn, Olancho, Santa Barbara, Santa Rita) and Nicaragua ( Granada, Río San Juan, Zelaya) at 50–1000(1524) m, in Tropical moist forest and Premontane wet forest life zones.

Phenology: —Flowering January-March, August-September. Fruiting in January-July, April-August, and November.

Discussion: —The species is a member of sect. Marcgraviopsis and is characterized by its appressed-climbing habit, minutely speckled petioles which are sheathed throughout with flaking or fibrous margins, ovate blades which are very asymmetric, unequal and truncate to acute at base and dry dark brown to grayish yellow-brown or blackened, usually with 1–9 perforations on both sides of the blade, 6–12 primary lateral veins as well as 1–2 inflorescences per axil with the peduncle typically shorter than the spadix, a spathe that is 3–5 cm longer than the spadix and a spadix about (3.1)4.5–7.0 times longer than broad and white to pale yellow-cream at anthesis with the fruiting spadix yellowish, pale yellow-green to white.

Monstera acuminata could be confused with M. spruceana but differs in having petiole 7–11 cm long (vs. 25–50 cm long), petiole sheath persistent (vs. petiolar sheath disintegrating as fibrous residues), spadix 8–12 × 2–3 cm (vs. spadix 15–25 × 3–6 cm), spadix 4.5–7.0 times longer than wide (vs. spadix 8–9 times longer than wide), and spathe 3–5 cm longer than the spadix (vs. spathe uniform or up to 0.5 cm longer than the spadix). Monstera acuminata is distributed from southern Mexico to Nicaragua, while M. sprucena is distributed from Costa Rica to Brazil.

Nomenclatural remarks: —In the protologue of Monstera karwinskyi, Schott (1859) indicated that the species was described using material from´ Mexico (Colipa)´and cited´Karwinsky´ as the collector.

Wilhelm Friedrich von Karwinsky von Karwin (1780–1855) was a German-Austrian naturalist who traveled in Brazil and Mexico. In this last country, Karwinsky made two expeditions. The first was carried out between 1826 and 1831 and was sponsored by the German-American Mining Society of Dusseldorf and the Bavarian government to make further collections of natural history objects. The second expedition, where he collected the type of M. karwinskyi , was carried out between 1841 and 1843 and was supported by five different sponsors in St. Petersburg, mainly to look for wildlife (plants and animals) and minerals. On this second trip to Mexico, Karwinsky met the Danish collector Frederik Liebmann (1813–1856) in Veracruz dunring February 1941, where they agreed to travel and explore together northward through the lowlands (Veracruz to Xicaltepec from February to May, 1841). Both naturalists arrived at Colipa, the type locality of M. karwinskyi , where they explored for 27 days (presumably until April 1941). As a result of the entire expedition, Karwinsky collected more than 2000 gatherings (in 6 major sets), which were partly named, and distributed c. 1851 by Ruprecht in St. Petersburg ( McVaugh 1980).

Engler & Krause (1908), in their revision of the Monsteroideae , recognized M. karwinskyi and cited three specimens, the presumed holotype from W (Karwinsky s.n.) and two other specimens, one from B (Kerber s.n.) and another from C (Liebmann s.n.). Both the specimen from W and B (photo at F) were destroyed in the second world war. However, we traced two extant specimens which also appear to match the diagnosis made by Schott (1859), one specimen from C (no collector information, barcode C 10021452) and another at LE (Karwinsky 838, barcode LE00000997). Both specimens were examined by M. Madison in 1975, where he noted ¨ holotype ¨ on the specimen from C, however on the specimen from LE he wrote ¨Karwinsky 838 = holotype ¨. Subsequently, Madison (1977) in his revision of Monstera , cited the specimen from C as ¨type¨ and further alleged that it was collected by Karwinsky; additionally, he designated the specimen from LE (Karwinsky 838) as ¨ paratype ¨.

The Material from C (barcode C10021452) bears handwritings on a few notes about habit, identifications, locality (¨Colipa¨) and date of collection (¨4/41¨), however it does not include collector information. This material bears annotations by two people: the label at the lower right with the information of the locality and identification (as M. adansonii Schott ) has a handwriting very similar to that of Liebmann; however, the other label (to the lower left) noted a different identification (as M. karwinskyi ) which corresponds to an annotation by Schott himself. It is possible that this specimen at C was actually originally collected by Liebmann during his trip with Karwinsky in May 1841; in fact, Engler & Krause (1908) cited Liebmann as the collector of this specimen. Considering this information, we believe that the material from C does not contain sufficient evidence to indicate that it represents a duplicate of the original material cited by Schott (1859) which was a collection made by Karwinsky. Additionally, Schott (1859) made no mention of Liebmann in the original protologue of M. karwinskyi , so this specimen should be excluded at a potential lectotype.

On the other hand, the specimen from LE (barcode LE00000997) seems to represent the only extant original material. This material bears a small handwritten label with an identification ( Caladium pertusum ), also noted ¨a Colipa usque Papantla ubicumque frequens, scandens super arbores¨; additionally, it has a typewritten label with the information on the collector and number (Karwinsky 838) and the locality (Iter Mexicanum, 1841, 1842). According to McVaugh (1980), much of the material collected by Karwinsky was not consistently numbered by the collector; however, the material was sorted into systematic order by family and genus, and numbers were arbitrarily assigned, presumably by Ruprecht. Additionally, it is known that Ruprecht, in circa 1851, distributed duplicates to other herbaria outside the former Soviet Union, but mostly without name or other data ( McVaugh 1980). Curiously, Schott (1859) included in the same work three new taxa of Araceae ( Anthurium karwinskyi Schott (1859: 101) , Monstera karwinskyi and Philodendron subincisum (1859: 99)), where the material examined was collected by Karwinsky in Colipa and Paplanta (during his second exploration done in 1841–1843). This information could support the notion that the original material examined by Schott was one of the duplicates sent by Ruprecht, this would explain why the original protologue of M. karwinskyi includes scarce information, as well as explain the fact that the material from LE (Karwinsky 838) has a numbering which was omitted in the original protologue in Schott (1859).

Considering the facts layed out above, we concluded that Madison’s citation of the specimen at C as (implicitly) the holotype was incorrect and given that the specimen is not demonstrably part of the original material on which the protologue was based, Madison’s citation of the “type” is not able to be interpreted as an implicit lectotypification. We therefore propose a corrected typification for M. karwinskyi , where we designate the specimen Karwinsky 838 (bardcode LE00000997) from LE, cited by Madison as a paratype, as the lectotype.

On the other hand, in the case of Monstera chiapensis , it was not possible to trace the syntypes cited by Matuda (1949), which are presumably deposited in the HEM Herbarium (at Matuda Herbarium); therefore, it was not possible to perform a proper typification.

Additional specimens examined: — BELIZE. Cultivated at Missouri Botanical Garden . Received from J. Dwyer s.n. ( MO!) ; Cayo: New Maria Camp. Forest , 16°49’38”N 089°01’W, 550 m, 3 May 1995, C. Whitefoord 9260 ( MO!) ; Resumadero Camp, at Chiquibul River. On fig tree, 16°37’41”N 089°04’05”W, 507 m, 14 Mar 1999, G. Reese, et al. 46 ( MO!) ; Forest along highway at mile 28.5 on Hummingbird Highway , south of Belmopan , 17°04’22”N 088°35’58”W, 200–300 ft, 14 June 1973 – 21 June 1973, T.B. Croat 24558 ( MO!) ; Stann Creek: Cockscomb Basin Wildlife Sanctuary , ca. 1 km west of visitors centre, ca. 30 m south of main trail before Wari Loop turnoff, plot 10, 16°46’50”N 088°27’55”W, 100 m, 06 March 2007, Z.A. Goodwin et al. 490 ( E!); Toledo: Jalacté , 16°11’18”N 089°10’54”W, 300 m, June 2008, B. Walshe-Roussel & F. Caal s.n. ( MO!) ; Southwestern Maya Mountains, Columbia River Forest Reserve. Trail between Union and Gloria Camps. On slightly undulating limestone hills, 16°23’22”N 089°08’10”W, 700–750 m, 13 April 1992, B.K. Holst 4382 ( MO!) ; Southwestern Maya Mountains , Columbia River Forest Reserve , Union Camp. On undulating limestone hills. Exposed limestone, common on hills and along streams. On dry hills E and N of camp, 16°23’N 089°09’W, 700–750 m, 06 April 1992, B.K. Holst 4121 ( MO!) ; Columbia River Forest Reserve : along SW base of Little Quartz Ridge, between Camp 1 and Union Camp. Moist, semievergreen forest on limestone soils with gently rolling hills and frequent limestone outcropping. MAP coordinates, 16°23’N 089°07’W, 700–750 m, 22 Feb 1997, B.K. Holst 5947 ( BRH!, CM!, COL!, MO!, NY!); Jimmy Cut, Salamanca , 16°18’N 089°03’W, 40 m, 28 May 1979, C. Whitefoord 1913 ( BM) ; Southern Maya Mountains , Bladen Nature Reserve , West Snake Creek. Along Snake Creek, 16°27’54”N 089°01’04”W, 580 m, 29 May 1997, D.L. Holland 15 ( BRH!, MO!); Southern Maya Mountains , Bladen Nature Reserve , central Snake Creek Camp, 16°27’35”N 088°58’52”W, 400 m, 28 May 1997, G. Davidse 36863 ( MO!, WU!); Southern Maya Mountains , Bladen Nature Reserve , Ek Xux Valley , stream side vegetation NW of EK Xux archeological site, 16°30’27”N 088°54’25”W, 280 m, 14 May 1996, G. Davidse 35953 ( BRH!, MO!); Southern Maya Mountains , Bladen Nature Reserve , Ek Xux Valley , stream side vegetation NW of EK Xux archeological site , 16°30’27”N 088°54’25”W, 280 m, 14 May 1996, G. Davidse 35910 ( BRH!, MO!); Southern Maya Mountains , Bladen Nature Reserve , streamside forest along river north of ”AC Camp”, ca. 2 airline km N of upper Bladen Branch, 16°30’15”N 088°55’07”W, 260 m, 11 May 1996, G. Davidse 35788 ( BRH!, MO!); Solomon Camp, vicinity of the junction of Richardson Creek and Bladen Branch, foothills of the Maya Mountains , 16°32’N 088°45’W – 16°33’N 088°46’W, 80–420 m, 05 March 1987 – 12 March 1987, G. Davidse & A.E. Brant 32195 ( MO!) ; Ca. 40 km SW of Punta Gorda, Dolores Estate. Growing along trail. Primary forest , 15°59’N 089°13’W, 1 Feb 1990, M.J. Balick et al. 2550 ( MO!) ; Between Condemned Branch Pine Ridge and Mafridyle [Mafredi] Lagoon, 16°09’56”N 088°57’16”W, 9 May 1952, P.H. Gentle 7687 ( LL!) ; Balsam Hill, Edwards Road , beyond Columbia , 16°16’17”N 088°57’15”W, 10 January 1948, P.H. Gentle 6360 ( LL!) ; Vic. San Antonio , 16°14’N 088°59’W, 27 Jan 1946, P.H. Gentle 5499 ( LL!) ; Vicinity of Condemn Branch Hills , 05 March 1945, P.H. Gentle 5254 ( LL!) ; Swasey Branch, Monkey River , 16°38’N 088°35’W, 9 Mar 1942, P.H. Gentle 3956 ( MO!) ; 1.5 miles S of Mayan village of San Jose, ca. 5 miles W of Columbia Forest Station , 16°15’N 089°04’W, 12 June 1973, T.B. Croat 24348 ( MO!) . EL SALVADOR. Santa Ana: m, E.M Martinez 174 ( TEFH!) . GUATEMALA. Alta Verapaz: 1 km W of Sebol , 15°48’07”N 089°57’09”W, 27 Apr 1964, E. Contreras 4518 ( LL!) ; Near Río Icvolay, near Hacienda Yaxcabnal , 5 mi NW of Cubilgüitz. Wooded slopes, 15°45’42”N 090°31’21”W, 250 – 300, 5 Mar 1942, J.A. Steyermark 44670 ( F!, MO!) ; Along road to El Estor (Lago Izabal), 2–4 miles E of Tamahú, 9–11 miles E of Highway CA-14 to Cobán, 15°18’44”N 090°16’34”W, 700–800 m, 18 Jul 1977, T.B. Croat 41496 ( MO!) ; Along road to El Estor (Lago Izabal), 5 miles W of Tucurú, 12 miles E of Highway CA-14 to Cobán, 15°18’36”N 090°15’10”W, 600 m, 18 July 1977, T.B. Croat 41513 ( MO!) ; 4 miles up road to Oxec, along road which turns north off Highway 7E (to El Estor), ca. 6 km NE of Panzos, 15°29’03”N 089°39’20”W, 500 m, 20 July 1977, T.B. Croat 41608 ( MO!) ; Escuintla: 6 miles north of Escuintla on road to Alotenango. Roadside near small river, 14°23’05”N 090°49’49”W, 650 m, 27 Jul 1977, T.B. Croat 42047 ( MO!) ; Izabal: Cerro San Gil , 15°37’38”N 088°47’48”W, 400 m, Oct 1996, C. Lopez s.n. ( MO!) ; Izabal, Cerro San Gil , 15°37’22”N 088°49’21”W, 300 m, 24 Oct 1996, C. Lopez 1996 ( MO!) ; Along dirt road which turns E into piña plantations from Highway CA-14, ca. 7 miles S of Puerto Barrios, 15°36’46”N 088°34’01”W, 50 m, 22 Jul 1977, T.B. Croat 41791A ( MO!) ; Along dirt road which turns E into piña plantations from Highway CA-14, ca. 7 miles S of Puerto Barrios, 15°36’46”N 088°34’01”W, 50 m, 22 Jul 1977, T.B. Croat 41814 ( MO!, TEX); Petén: Tikal National Park , [Tikal] , 17°13’30”N 089°36’47”W, 2 Mar 1961, C.L. Lundell 16834 ( LL!) ; Tikal National Park , [Tikal], 17°13’30”N 089°36’47”W, 2 Mar 1961, C.L. Lundell 16832 ( LL!) ; Tikal National Park, Tikal , 17°13’30”N 089°36’47”W, 8 Feb 1959, C.L. Lundell 15409 ( LL!) ; Río Pasion , Ceibal , above Sayaxche , 16°30’48”N 090°03’44”W, 3 Feb 1964, C.L. Lundell 17642 ( LL!) ; Tikal National Park, on Remate Road , S of Tikal, 17°13’30”N 089°36’47”W, 4 Mar 1961, C.L. Lundell 16858 ( LL!) ; Lacandon , 17°07’55”N 091°10’27”W, 3 Feb 1962, E. Contreras 3301 ( LL!) ; Ruins of Ceibal , 16°30’48”N 090°03’44”W, 240 m, 18 June 1971, W.E. Harmon & J.A. Fuentes 5764 ( MO!, UMO); Dolores. Dolores, on Río Mopan trail, about 3–5 km east of village, 16°32’22”N 089°22’44”W, 25 Apr 1961, E. Contreras 2185 ( LL!) ; San Andres . Parque Laguna del Tigre , 17°44’47”N 090°17’37”W, 110 m, 3 May 1996, M. Véliz 96.5505 ( MO!) ; Sayaxche. [Río Pasion], Sayaxche , about 3 km SSE of the village, 16°32’06”N 090°09’35”W, February 1964, C.L. Lundell 18032 ( LL!) ; Quetzaltenango: Along Highway CA- 2, 4 mi. NW of turnoff to Coloba, 14°41’56”N 091°52’41”W, 400 m, 22 Feb 1976, T.B. Croat 32781 ( MO!) ; Highway CA 2, 3 mi S of turnoff to Colomba, between Coatepeque and Retalhuleu. Coffee plantation, 14°39’30”N 091°46’36”W, 600 m, 22 Jan 1987, T.B. Croat & D.P. Hannon 63436 ( AAU, ENCB, HNMN, MEXU, MO!, SEL!, TEX); San Marcos: La Trinidad, ca. 2 km from Finca Armenia above San Rafael, 14°56’02”N 091°53’55”W, 1100–1250 m, 12 July 1977, T.B. Croat 40844 ( MO!) ; Suchitepéquez: Vicinity of Tiquisate. Near Santiago farm. Virgin forest , 14°13’46”N 091°19’50”W, 100 m, 17 Jun 1942, J.A. Steyermark 47651 ( MO!) ; 1 mi E of Mazatenango , 14°32’13”N 091°28’22”W, 300–500 m, 20 Aug 1977, T.B. Croat 43756 ( MO!) ; Zacapa: Along old road to Finca Agua Fría , 2 miles N from Route CA- 9, 41 miles S of turnoff to Petén (near Morales), 15°15’02”N 089°16’24”W, 150 m, 23 Jul 1977, T.B. Croat 41873 ( MO!) . HONDURAS. Atlántida: Jardin Botanico de Lancetilla , 15°41’59”N 087°28’00”W, 0-500 m, 29 Dec 1993, C.H. Nelson & R. Andino 17120 ( TEFH!) ; Jardin Botanico de Lancetilla, camino de los Pilis , 15°41’59”N 087°28’00”W, 20 m, 8 enero 1994, C.H. Nelson et al. 17192 ( TEFH!) ; Campamento Quebrada Grande ca. 10 km south west of La Ceiba. At base of north slope of Pico Bonito, from camp to 2 km south of camp. Río Bonito, forest above, and Cacao Plantation, 15°42’N 086°51’W, 80–140 m, 14 May 1993, R.L. Liesner 26317 ( MO!) ; Along trail to dam for municipal water supply of Tela, Lancetilla Botanical Gardens, on road ca. 2 mi WSW of Tela and S of main hwy. Printed label for specimens 64586–64646 reads 9/2/1987, but fieldbook states 10/2/1987, 15°44’N 087°27’W, 70–90 m, 10 February 1987, T.B. Croat & D.P. Hannon 64632A ( MO!) ; Tela. Lancetilla Valley ca. 10 miles southeast of Tela; in forest preserve along Río Lancetilla, on trail to water reservoir, 15°44’N 087°27’W, 10–150 m, 03 August 1977, T.B. Croat 42671 ( MO!) ; Comayagua: Southern end of Lago Yojoa , 14°46’40”N 087°59’08”W, 610 m, 11 May 1991, G. Davidse 34344 ( MO!) ; Copán: Santa Rita. 13 miles east of Copán along gravel road to La Entrada. disturbed roadsides, 14°52’N 089°04’W, 750 m, 1 Aug 1977, T.B. Croat 42530 ( MO!) ; Ca. 11 miles east of Copán on gravel road to La Entrada. Disturbed roadsides, 14°52’N 089°05’W, 750 m, 1 Aug 1977, T.B. Croat 42523 ( MO!) ; Cortés: Omoa. Ca. 1 mile northeast of Omoa on road to Puerto Cortes; sea level; pasture with a few trees bearing epiphytes, 15°47’06”N 088°01’26”W, 0 m, 2 Aug 1977, T.B. Croat 42552 ( MO!) ; Olancho: Along Río Olancho, on road between San Francisco de la Paz and Gualaco, 13.6 miles SW of Gualaco, on steep slope ca 0.5 miles E of main road, along gravel road into private property, 15°00’N 086°07’W, 1300 m, 06 February 1987, T.B. Croat & D.P.Hannon 64298 ( MO!) ; Catacamas.Catacamas. Montanas de Murmuyo [El Murmullo], Sierra de Agalto. Broadleaf forest, 14°52’57”N 085°56’10”W, 1000 m, 25 Feb 1982, S. Blackmore & G.L. Heath 1948 ( MO!) ; Santa Bárbara: Along Hwy. CA14 (National Hwy. 18), between Santa Rosa de Copán and San Pedro Sula, 51.3 mi WSW of junction of main San Pedro Sula–Tegucigalpa hwy. 39.2 mi SWS of frontier of Cortés Dept, 15°18’N 088°25’W, 500 m, 20 Jan 1987, T.B. Croat & D.P. Hannon 63870 ( CR!, MO!, P) . MEXICO. Cordoba , La Luz, E. Kerber s.n. ( MO!) ; 1841–1843, F.M. Liebmann s.n ( KEW 153) Chiapas: Berriozábal, La Pera , 16°51’43”N 093°18’53”W, 1140 m, 12 March 2020, P.D. Jiménez et al. 1486 ( HEM!) ; Road to Lacandona Reserva at the Nueva Palestina turnoff, 16°54’36”N 091°12’00”W, 300 m, 11 Oct 1986, B.E. Hammel et al. 15668 ( MO!) ; Lacondona Reserva ; Crucero frontera; Corozal , 16°54’36”N 091°04’48”W, 290 m, 11 Oct 1986, B.E. Hammel et al. 15674 ( MO!) ; Near ruins of Palenque . Mesophytic woods and roadside, 21 Jul 1971, D.L. Spellman et al. 161 ( MO!) ; Crucero Corozal, camino Palenque-Boca Lacantum. Veg. Selva alta subperennifolia, 16°45’36”N 091°00’00”W, 180 m, 8 Jan 1986, E.M. Martínez 15601 ( MO!) ; En reserva El Triunfo, 1900 m, 01 junio 1987, E.M. Martínez et al. 21501 ( MEXU!) ; Acacoyagua . m, A. López 55 ( HEM!) ; m, M. Martínez 918 ( HEM!) ; Angel Albino Corzo.Along slopes of Río Cuxtepec below Finca Cuxtepec, 15°43’48”N 092°57’36”W, 1080 m, 13 Dec 1980, D.E. Breedlove 48618 ( CAS) ; Ridge, just N of Finca Cuxtepec, 15°43’42”N 092°56’59”W, 1370 m, 11 January 1982, D.E. Breedlove & F. Almeda 56970 ( MO!) ; Escuintla . Along road from Escuintla to El Triunfo, ca. 1 mile N of Escuintla, 15°19’12”N 092°38’24”W, 100 m, 21 August 1977, T.B. Croat 43803 ( MO!) ; Along road between Escuintla and Monte Ovando, 2.8 km NW of Turquia, 15°21’N 092°39’W, 100 m, 13 February 1979, Thomas B. Croat 47509 ( MO!) ; About 12 miles NE of Escuintla, ca. 2 miles NE of El Triunfo, on gravel road to Permuta, 15°21’36”N 092°31’48”W, 300 m, 21 August 1977, T.B. Croat 43873 ( MO!) ; About 10 miles NE of Escuintla just above El Triunfo, 15°20’39”N 092°32’09”W, 300 m, 21 Aug 1977, T.B. Croat 43833 ( MO!) ; Along trail between Finca California (at base of S slope of Monte Ovando and ca. 4 km N of of Ovando Turquia) and summit of Cerro Ovando, 15°22’12”N 092°36’00”W, 450–850 m, 14 February 1979, T.B. Croat 47574 ( MO!) ; About 12 miles NE of Escuintla, ca. 2 miles NE of El Triunfo, on gravel road to Permuta, 15°21’36”N 092°31’48”W, 300 m, 21 August 1977, T.B. Croat 43874 ( MO!) ; La Concordia . m, C.A. Pérez 89 ( HEM!) ; La Trinitaria . m, A. Shilom Ton 2428 ( OAX) ; 10 km east northeast of Dos Lagos above Santa Elena, 16°06’36”N 091°34’18”W, 1170 m, 19 January 1982, D.E. Breedlove & F. Almeda 57561 ( MO!) ; Mapastepec. Sierra de Soconusco. New unfinished road to Tuxtla Gutiérrez from Highway 200 (5.5 mi NW of turnoff to Mapastepec), 6.5–8.5 mi up road, 15°39’26”N 093°02’39”W, 350–450 m, 20 January 1987, T.B. Croat & D.P. Hannon 63338 ( MO!) ; Montecristo de Guerrero. m, 147 m, R. Reinoso 500 ( HEM!) ; Ocosingo . m, E.M. Martínez 15601 ( MEXU!) ; 5 km al S de Crucero Corozal. Selva alta perennifolia, 250 m, 21 Apr 1985, E.M. Martínez 12247 ( MO!) ; Nuevo Guerrero sobre el camino Palenque-Boca Lacantum, 16°59’24”N 091°17’24”W, 340 m, 14 abril 1986, E.M. Martínez 18140 ( MO!) ; A 1.6 km al SE de San Javier, 16°47’28”N 091°05’43”W, 604 m, 25 abril 2003, G. Águilar & C. Chancayun 6598 ( MEXU!, MO!); A 1.63 km de Frontera Corozal al SE , 16°48’06”N 090°52’33”W, 144 m, 04 junio 2004, G. Águilar & M. Méndez 10437 ( MEXU!, MO!); Al S de Frontera Corozal, 16°47’35”N 090°52’09”W, 132 m, 19 abril 2004, G. Águilar & M. Méndez 9827 ( MEXU!, MO!); A 1.07 km al SO del crucero San Javier, 16°47’25”N 091°04’34”W, 361 m, 09 diciembre 2003, G. Águilar et al. 8776 ( MEXU!, MO!); A 1.25 km al SE de San Javier. Selva alta perennifolia, 16°47’32”N 091°05’48”W, 411 m, 27 febrero 2003, G. Águilar et al. 5904 ( MEXU!, MO!); 5 km SW of Santo Domingo (120 km SE of Palenque on road to Bonampak), 16°59’24”N 091°24’36”W, 600 m, 13 May 1982, G. Davidse et al. 20400 ( MO!) ; Ocozocoautla de Espinosa. m, D. Torres-Arguello 58 ( CHIP!) ; 20 mi N of Ocozocouatla along gravel road to Apitpac, near km 31 marker, 16°58’15”N 093°30’10”W, 700 m, 09 July 1977, T.B. Croat 40655 ( MO!) ; 13 miles north of Ocozocoautla on gravel road to Apitpac, near Km 21 marker, 16°54’N 093°27’W, 900 m, 9 Jul 1977, T.B. Croat 40631 ( 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 40078 ( MO!) ; Palenque. Along gravel road from Palenque to Bonampak, 60 miles SE of Palenque, 17°00’03”N 091°19’11”W, 400 m, 05 July 1977, T.B. Croat 40196 ( MO!) ; Along gravel road between Palenque and Bonampak, 88–90 miles SE of Palenque, 16°52’30”N 091°12’03”W - 16°51’41”N 091°09’49”W, 350–370 m, 05 July 1977, T.B. Croat 40253 ( MO!) ; 5 miles SE of Palenque, on road to Chancala, Ocosingo and San Cristóbal de las Casas, 17°28’12”N 091°57’36”W, 200 m, 4 Jul 1977, T.B. Croat 40138 ( MO!) ; Siltepec . m, R. Reinoso 582 ( HEM!) ; Tuxtla Gutiérrez . m, E. Palacios 879 ( CHIP!) ; m, E. Palacios 2316 ( CHIP!) ; Oaxaca: Tuxtepec to Oaxaca; 5 km S of Tuxtepec; on forested hill at paper mill, 18°04’N 096°10’W, 130 m, 26 Sep 1986, B.E. Hammel & M. Merello 15461 ( MO!) ; Uxpanapa Region, along gravel road between Esmeralda (17 km E of Sarabia) and Río Manea (tributary of Río Verde), 11.5–13.5 mi S of Esmeralda. Roadside vegetation with primary forest elements. Veg. type: ”selva alta perennifolia” , 17°03’36”N 094°45’W, 90–110 m, 19 Jan 1987, T.B. Croat & D.P. Hannon 63276 ( F!, GUAT!, MO!, PMA!); Uxpanapa Region, along gravel road from Esmeralda (17 km E of Sarabia) to Río Verde, 1.1 mi S of Esmeralda. Edge of pasture and in forest on karst limestone formation on steep slopes with cliffs. Veg. type: ”selva alta perennifolia”, 17°09’36”N 094°45’W, 100 m, 18 Jan 1987, T.B. Croat & D.P. Hannon 63240 ( MO!) ; Uxpanapa region, along gravel road between Esmeralda (17 km E of Sarabia) and Río Verde, 1.1 mi S of Esmeralda; edge of pasture and forest on karst limestone. Veg. type: ”selva alta perennifolia”, 17°10’N 094°45’W, 100 m, 19 Jan 1987, T.B. Croat & D.P. Hannon 63306 ( MO!) ; Along Rte. 175 between Valle Nacional and Oaxaca, 7–10 km S of bridge over Río San Juan Bautista at Valle Nacional. Steep roadside cliffs, 17°43’33”N 096°19’39”W - 17°43’26”N 096°19’18”W, 597–666 m, 3 Mar 2008, T.B. Croat & P.D Jiménez 100163 ( MO!) ; Tuxtepec. Mun. San Juan Bautista Valle Nacional. Sierra de Juárez, along Highway 175, between Valle Nacional and Oaxaca, 14 miles SW [above/west] of Valle Nacional, 17°39’36”N 096°19’12”W, 1210 m, 30 June 1977, T.B. Croat 39784 ( MO!) ; [Mun. San Juan Bautista Valle Nacional].About 14 miles S of Valle Nacional, along Highway 175 to Oaxaca, 17°39’36”N 096°19’12”W, 1220 m, 22 August 1977, T.B. Croat 43904 ( MO!) ; Puebla: San Miguel Tzinacapan. Vicinity: Partiendo hacia el suroeste de Tzinacapan hacia el poniente rumbo a Masacowatah pero no llegando hasta allá. Justamente por el arroyo de Atekokomol, 20°01’03”N 097°33’01”W, 870 m, 12 October 2015, J.D. Amith 2673 ( MO!) ; Cuetzalan del Progreso. Taxipchual. Bejando por el lado oriente de un potrero por donde termina la brecha a la cascada y llegando a un arroyo y después volviendo a subir hacia la brecha entre Taxipehual y las cascadas, 20°01’38”N 097°27’19”W, 534 m, 11 October 2015, J.D. Amith 2667 ( MO!) ; m, J.L. Contreras 6136 ( HUAP!) ; Querétaro: 2 km al suroeste de Neblinas, Río Tancuilin, municipio de Landa. Bosque de Bursera, Cedrela, Higerón y Oshite; ladera de cerro; orilla de arroyo; cañada, 610 m, 12 Sep 1990, H. Rubio 1959 ( MO!) ; Landa de Matamoros. 2 km al SO de Neblinas, Río Tancuilin, 21°15’00”N 099°03’00”W, 610 m, 12 Sep 1990, H. Rubio 1959 ( IEB!) ; San Luis Potosí: Creek bottom, north of Tamazunchale, 4 July 1935, O.M. Clark 7407 ( MO!) ; Tamazunchale. Hills above Highway 85, 6 mi NW of Tamazunchale, 21°17’24”N 098°47’24”W, 250 m, 25 June 1977, T.B. Croat 39269 ( MO!) ; Tabasco: Cárdenas. A 2 km del Poblado C-29, 18°03’49”N 093°27’24”W, 14 m, 10 Sep 2008, P.D Jiménez 613 ( MO!) ; Jalpa de Médez. Km 2 (1.2 miles) de la carretera Jalapa-Tacotalpa. Km 33.9 (21.2 miles) de la desviación de la carretera hacia Lomas Alegres y Castañal, 17°34’12”N 092°36’36”W, 26 m, 16 June 1983, C.P. Cowan 3929 ( MO!) ; Tacotalpa. 0.2 km abajo (NW) de, y antes de entrar a Tapijulapa. Hasta 0.5 km arriba del camino por el arroyo a pie, 17°28’00”N 092°46’27”W, 43–47 m, 30 mayo 1982, C.P. Cowan et al. 3533 ( MO!) ; Veracruz: Laguna Escondida, 5 km de la Estacion de Biologia de Los Tuxtlas. Orilla de la selva alta perennifolia, primaria, 300 m, 17 Sep 1974, M. Soto & C.C. Horvitz 27 (F); 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.D. Jiménez 100324 ( MO!) ; Along secondary road between Hidalgotitlán and Jaltipan, along Río Coatzacoalcos, vicinity Buena Vista, 4 km S of Buena Vista, 17°49’17”N 096°38’17”W, 150 m, 5 Mar 2008, T.B. Croat & P.D. Jiménez 100281 ( MO!) ; Alrededores de la Estación de Biologia Tropical ”Los Tuxtlas” , carr. Catermaco-Montepio, Mpio. de San Andrés Tuxtla, 18°35’N 095°04’W, 120 m, 11 May 2005, T. Krömer 2172 ( MO!); Estación de Biologia Tropical ”Los Tuxtlas, Cerro Vigia 5, Lote 67, Mpio. de San Andrés Tuxtla , 18°34’N 095°04’W, 450 m, 12 April 2005, T. Krömer 1914 ( MO!) ; Catemaco . 19 km E of Catemaco, 18°26’24”N 094°58’12”W, Mar 1965, L.G. Quintero 2243 ( MICH, US); Vicinity 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, 21 Apr. 1983, M.H. Nee 26708 ( F!, MO!, NY!, XAL!); Coatzacoalcos. April 1895, C.L. Smith 1069 ( US!); Fortín. Near Fortín along walkway from hydroelectric plant to railroad tracks. Open, mostly cleared area, 18°53’24”N 096°59’24”W, 1000–1150 m, 27 June 1977, T.B. Croat 39468 ( MO!) ; Huatusco. ”El Mirador”, 21 km E of Huatusco at Km 45 along Highway to Puente National, 19°12’36”N 096°52’12”W, 1200 m, 23 August 1977, T.B. Croat 43998 ( MO!) ; ”El Mirador”, 21 km E of Huatusco at Km 45 along Highway to Puente National, 19°12’36”N 096°52’12”W, 1200 m, 23 August 1977, T.B. Croat 43992 ( MO!) ; ”El Mirador”, 21 km E of Huatusco at Km 45 along Highway to Puente National, 19°12’36”N 096°52’12”W, 1200 m, 23 August 1977, T.B. Croat 44012 ( MO!) ; Minatitlán . Minatitlán, 17°58’45”N 094°33’45”W, 14 m, 30 January 1892, J.G. Smith 240 ( MO!) ; 13.7 km al E de La Laguna sobre terracería a Uxpanapa, luego 6.7 km al N sobre camino nuevo (no completo) a Belisario Domínguez (brecha 93); zona kárstica con selva perennifolia de Dialium, Brosimum, Spondias, Bursera, Robinsonella mirandae, etc, con afloramientos grandes de piedra, 17°19’48”N 094°22’48”W, 130 m, 26 Nov 1981, T.L. Wendt et al. 3482 ( MO!) ; Papantla. 100 m, Feb 1968, L. Gutiérrez R. 75 (US!); San Andrés Tuxtla . m, G.I. Manríquez & et al. 2138 ( FEZA) ; Alrededores de la Estación de Biologia Tropical ”Los Tuxtlas”, carr. Catemaco-Montepio. Selva alta perennifolia, 18°35’N 095°04’W, 120 m, 11 May 2005, T. Krömer & A. Acebey 2178 ( MO!) ; Alrededores de la Estación de Biologia Tropical ”Los Tuxtlas” , 18°35’N 095°04’W, 120 m, 05 noviembre 2005, T. Krömer & A. Acebey 2178 ( MO!) ; Ejido 1º de Mayo, faldas del Volcán San Martín. Selva alta perennifolia, 18°33’N 095°13’W, 920 m, 2 May 2005, T. Krömer & A. Acebey 2060 ( MO!) ; Santiago Tuxtla. E of Santiago Tuxtla, 18°27’36”N 095°17’24”W, 25 Jan. 1965, M. Sousa S. 2156 ( MEXU!) ; Totutla. Km 45 on highway between Conejo (near Puente Nacional) and Huatusco, Hacienda ‘El Mirador’, slopes of barranca de Santa Maria, 19°12’00”N 096°45’36”W, Sept. 1961, H.E. Moore & G.S. Bunting 8859 ( BH) ; Zongolica. Wooded area on the north side of airdrome. Campo Experimental de Hule, El Palmar, 19 Nov 1944 – 20 Nov 1944, J.V. Santos 3635 ( MICH) . NICARAGUA. Atlántico Norte: Cerro Baká , 6.5 km E of Río Coperna; lower mountain slopes and secondary areas, 13°40’N 084°30’W, 200–320 m, 19 Mar 1979, J.J. Pipoly, III 4932 ( MO!) ; Caño Hormiguero , SE of Cerro La Pimienta No. 1, 13°46’N 084°59’W, 750 m, 10 April 1979, J.J. Pipoly, III 5084 ( MO!) ; Reserva Bosawas , Municipio de Siuna, sendero rumbo a la cara oeste del cerro Saslaya, 13°42’N 084°56’W, 100–250 m, 9 Abril 1996, R.M. Rueda et al. 4404 ( MO!) ; Along new road fron Siuna to Matagalpa, ca. 31.4 km beyond Río Ulí (near Waní), ca. 8.9 km beyond Rosa Grande La Balsama and near major highway construction camp, 13°36’N 085°05’W, 100–200 m, 18 March 1978, W.D. Stevens 7515 ( MO!) ; Estelí: Municipio de Esteli, ásentamiento Puertas Azules , 13°16’N 086°16’W, 1300 m, 17 Abril 1999, R.M. Rueda & W. Velásquez 10928 ( MO!) ; Granada: Volcán Mombacho, Hacienda Las Delicias , ca 10 km al SE de Ciudad Granada; bosque húmedo tropical, 11°50’50”N 085°57’50”W, 600–800 m, 24 junio 1984, D. Soza et al. 104 ( MO!) ; Volcán Mombacho, near Finca Cutirre , 11°49’N 085°56’W, 250–300 m, 01 July 1975, J.T. Atwood & D.A. Neill AN66 ( MO!) ; Volcán Mombacho, entre las fincas Maria Auxiliadora y el Cacao , 11°51’N 085°56’W, 400–500 m, 24 mayo 1980, P.P. Moreno 443 ( MO!) ; NE del Volcán Mombacho, en el empalme de los caminos a Santa Isabel y Cutirre , 11°51’N 085°57’W, 340–360 m, 16 septiembre 1980, P.P. Moreno 2624 ( MO!) ; Volcán Mombacho, 2 km al SE de la Hacienda Cutirre, El Chipote , 11°48’N 085°54’W, 190–195 m, 11 junio 1982, P. P. Moreno 16453 ( MO!) ; Volcán Mombacho al Sur de la Hacienda Cutirre , 11°49’N 085°55’W, 300–400 m, 11 junio 1982, P.P. Moreno 16449 ( MO!) ; Volcán Mombacho, arriba y sureste de la Hacienda Cutirre , 11°49’N 085°56’W, 290–300 m, 29 enero 1981, P.P.Moreno 6311 ( MO!) ; Municipio de Granada. Reserva Natural Volcán Mombacho , 11°50’27”N 085°58’59”W, 1150–1350 m, 2 Diciembre 2003, R.M. Rueda & L.D. Paguaga 17621 ( HULE) ; Upper slopes of Volcán Mombacho, along W shore of Lake Nicaragua , ca. 15 km S of Granada, 11°50’N 085°58’W, 1100–1200 m, 19 March 1977, T.B. Croat 39104 ( MO!) ; Jinotega: Along rock road 3.8 miles SE of Yalí, between Condega on Route 1 and Jinotega on Route 3. Roadside and disturbed primary forest on steep slopes, 13°15’N 086°09’W, 1350 m, 6 Aug 1977, T.B. Croat 42933 ( MO!) ; Managua: Ca. 5.4 km NE of El Crucero, ridge of Sierra de Managua near Hda. Palmira , 12°01’N 086°16’W, 800–900 m, 25 August 1977, W.D. Stevens 3503 ( MO!) ; Matagalpa: Ridge along road between La Danta and La Luna , E of Esquipulas, 12°40’N 085°43’W, 960–1000 m, 25 January 1979, W.D. Stevens 11841 ( MO!) ; Río San Juan: Near Caño Chontaleño , 20 km NE of El Castillo, 11°08’N 084°12’W, 200 m, 18 April 1978 – 21 April 1978, D.A. Neill & P.C. Vincelli 3625 ( MO!) .

Figures

FIGURE 3. Monstera acuminata from Tuxtla Gutiérrez, Mexico. (A) Developing inflorescence. (B) Inflorescence in male anthesis with fragments of spathe. (C) Spathe fragments in back view. (D) Fertile flower; in lateral view (left), and longitudinal section (right). (E) Stylar plate with stigma (left) and one stamen (right). (F) Spathe fragments in frontal view. (G) Sterile flower; in lateral view (left), and longitudinal section (right). (H) Developing infructescence. P. Diaz-Jiménez et al. 1486 (HEM). Photos by M. Cedeño-Fonseca.

FIGURE 4. Monstera acuminata. Adult plant growing in Acacoyagua, Mexico. Photo by M. Cedeño-Fonseca. (Not collected).