Tynanthus guatemalensis Smith (1893: 6)

Medeiros, Maria Cláudia Melo Pacheco De & Lohmann, Lúcia G., 2015, Taxonomic Revision of Tynanthus (Bignonieae, Bignoniaceae), Phytotaxa 216 (1), pp. 448-450 : 448-450

6. Tynanthus guatemalensis Smith (1893: 6) . Lectotype (designated here):— GUATEMALA. Quezaltenango : Banks of Rio Ocosito, April 1892, J. Donnell Smith 1488 (US! (barcode 125785); isolectotypes GH! (barcode 93265) photo, K! (barcode 449544), M!, NY! (barcode 328979), P! (barcode 468599), US! (barcode 125786)).

Fig. 8 : G–J

Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, lenticeled to densely lenticeled, pubescent to glabrescent, with simple and peltate trichomes; interpetiolar ridge present; interpetiolar patelliform glands absent; prophylls of the axillary buds 3.5–20 mm long, 2.1–14 mm wide, foliaceous, ovate to elliptic, pubescent or puberulent to glabrescent throughout, with simple, peltate and patelliform trichomes. Leaves 2–3 foliolated (more commonly 3); terminal leaflets often modified into simple tendrils, without adhesive-disks on tip; petioles and petiolules pubescent throughout surface, with simple and peltate trichomes; petioles (0.9–) 1.5–5.6 cm long; petiolules (0.4–) 0.6–3.4 cm long; leaflets (2.5–) 3.5–12.2 cm long, (1.2–) 1.9–7.9 cm wide, membranous to chartaceous (sometimes subcoriaceous), discolor or concolor, elliptic; apex caudate, mucronate; base cuneate or obtuse, symmetrical; margin entire; the abaxial surface pubescent to glabrescent on and near the veins, with simple, peltate and patelliform trichomes; the adaxial surface pubescent to glabrescent on and near the veins, with simple, peltate and patelliform trichomes; glandular trichomes distributed especially on the abaxial surface; second venation weak brochidodromous; pocket domatia without (sometimes with) trichomes. Inflorescence axilar or terminal, a thyrse or a compound thyrse, lax, with conical aspect, first order (1.5–) 2.6–10.5 cm long, second order 0.5–2.1 cm long; axis pubescent, with simple and peltate (sometimes also with patelliform) trichomes; bracts of the

22 • Phytotaxa 216 (1) © 2015 Magnolia Press

MEDEIROS & LOHMANN inflorescence caducous or persistent, pubescent throughout (sometimes only at margins), 0.4–3.3 mm long; floral bracts 0.3–0.5 mm long; floral pedicels 1–4 mm long. Calyx green, 1.5–2.5 mm long, 1.3–2.5 mm wide, with transversal aperture, truncate or minutely 5-denticulate, pubescent to puberulent (sometimes glabrescent) throughout outside, with patelliform glands; lobes 0.1–0.4(–0.5) mm long. Corolla white, cream or pale yellow, 0.5–0.9 cm long, 1.9–3.5 mm wide at the tube opening; tube 2–5 mm long, internally pubescent at the base, with simple and long and short stipitate trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent throughout lower ones and at the margin of (sometimes throughout) upper ones; upper ones 0.3–1.8 mm long, 0.6–1.9 mm wide, acute to obtuse; lower ones 1.4–3.1 mm long, 1.6–3.2 mm wide, obtuse to rounded. Androecium with fertile stamens inserted ca. 1 mm from the base of the corolla; shorter ones 2–4 mm long; longer ones 3–5 mm long; anthers thecae 0.6–1 mm long, obovate to elliptic, subexserted; connective extending 0.1–0.3 mm beyond anther attachment; staminode 1.7–2 mm long, glabrous. Gynoecium 4–6.5 mm long; ovary 1–1.4 mm long, 0.6–0.8 mm wide, conical, densely pubescent; style 3–5 mm long, tomentose to pubescent at the base. Fruit a linear flattened capsule, (12.7–) 14–38 cm long, 0.8–2.4 cm wide, woody, smooth or granular throughout, without lenticels to densely lenticeled, glabrescent (sometimes pubescent at extremities), with peltate and patelliform trichomes (sometimes also with simple); central ridge single, slightly or not prominent; margins prominently raised (winged), 0.3–0.9 cm wide. Seeds body 0.8–2 cm long, 0.5–0.9 cm wide; wings 0.6–1.8 cm long.

Phenology: —Flowers from April to July and produces fruits from August to April.

Distribution and habitat: —Occurs in moist broadleaf forests from Belize ( Belize, Cayo, Orange Walk, Stann Creek and Toledo), Guatemala (Alta Verapaz, El Petén, Izabal and Quetzaltenango) and Mexico (Campeche, Chiapas, Quintana Roo, Tabasco and Veracruz-Llave) ( Fig. 10 ).

Additional specimens examined: — BELIZE. Belize: Gracy Rock Bank (“ Gracie Rock ”), Sibun River , 27 May 1935, P. H. Gentle 1652 ( K, MO, NY, P). Maskall, 25 May 1934, P. H. Gentle 1324 ( F, K, MO, NY, US). South of Yalbac Hills , Terra Nova Medicinal Plant Reserve , 40 m, 11 June 1993, J. B. Walker & L. Romero 138 ( NY). Cayo: Cayo village , May 1907, E. Campbell 112 ( K). Arenal – Valentin road, June–August 1936, C. L. Lundell 6168 ( F, NY, US). 53 Miles Section, Humming Bird Highway, 18 June 1956, P. H. Gentle 9132 ( F, K, MO). Maccal River , bridge between Augustine and Cuevas, 7 October 1972, J. D. Dwyer & R. Pippin 10189 ( MO). Vicinity of

TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)

Phytotaxa 216 (1) © 2015 Magnolia Press • 23 Cuevas south of Millionario , 29–30 May 1973, T. B . Croat 23599 ( F, MO, NY). South of Millionario , 1900 ft, 29 May 1973, A. H . Gentry 7661 ( F, INPA, MO, NY, US). Vicinity of Millionario , 1800 ft, 30 May 1973, A. H . Gentry 7689 ( MO, NY); 30 May 1973, A. H . Gentry 7730 ( F, MO, NY). Between Cuevas & Millionario, 30 May 1973, J . D. Dwyer 10814 ( MO, US). Vicinity of Millionario between the McCal River and Cuevas , 1900 ft, 30 May–3 June 1973, T. B . Croat 23715 ( F, K, MO, NY, US). 3 miles S of Grano de Oro , 1700 ft, 2 June 1973, A. H . Gentry 7755 ( F, MO, NY). Vicinity of Grano de Oro lumber camp south of Millionario , 1700 ft, 2 June 1973, A. H . Gentry 7768 ( F, MO, NY). 4 miles south of Grano de Oro on road to La Flor, 2 June 1973, J . D. Dwyer 10923 ( MO, US). Vicinity of La Flor at Rio de La Flor, 6 mi. south of Grano de Oro , 1700–2000 ft, 3 June 1973, T. B . Croat 23812 ( F, MO). Western Highway, near Belmopan , 3 June 1981, C . Whitefoord 3115 ( BM, MO). Monkey Tail track, 450 m, 29 May 1995, C . Whitefoord 9499 ( BM, F, MO). Chiquibul Forest Reserve, San Pastor track, c. 1 km from Las Cuevas , 560 m, 10 June 1996, A. I . Garcia A68 ( BM, MO). Augustine, Rio Frio Caves road, Mountain Pine Ridge Forest Reserve , 460 m, 3 July 1989, J . Meave & A . Howe 1433 ( BM, MO). Valentin , June–July 1936, C. L . Lundell 6354 ( F, NY, US). Orange Walk: Mile 58 on Northern Highway , ca. 4 miles south of Tower Hill , 23 June 1973, A. H . Gentry 8489 ( MO). 1.5 km S of Program for Belize Camp , 13 May 1991, R . Arvigo et al. 502 ( NY). Programme for Belize Rio Bravo Research Station, 150 m, 28 May 1997, D. Lentz et al. 2744 ( NY). Stann Creek: Stann Creek Valley , Mountain Cow Ridge , 25 March 1940, P. H . Gentle 3275 ( MO, NY, US). Cockscomb Mountains, tributary of Cocoa Branch of Sittee River, 2 km due north of Victoria Peak , 300–500 ft, 5–6 June 1973, A. H . Gentry 8014 ( MO). Toledo: Temash River , 200 ft, 7 May 1935, W. A . Schipp 8910 ( F photo); 5 June 1944, P. H . Gentle 4654 ( MO, NY). Near Agua Caliente, near San Antonio , 18 April 1945, P. H . Gentle 5335 ( F, K, MO). Beyond Central Camp, Edwards Road beyond Columbia , 21 May 1951, P. H . Gentle 7339 ( F, K, MO, NY). Broken ridge, Joe Taylor creek , 2 October 1952, P. H . Gentle 7775 ( F, K, MO). 1.5 miles from Maya village of San Jose on road to Columbia Forestry Station , 12 June 1973, A. H . Gentry 8119 ( MO). Mile 13 on road west from Punta Gorda, near junction of Southern Highway and road to San Antonio , 100 ft, 14 June 1973, A. H . Gentry 8222 ( MO). Southwestern Maya Mountains, Columbia River Forest Reserve, Trail between Gloria Camp and Edwards Camp to the South , 15 April 1992, B. K . Holst 4468 ( MO). GUATEMALA. Alta Verapaz: Along Río Icvolay between Río Apia and Río Soctelá, 8–10 miles northwest of Cubilgüitz , 200–210 m, 14 March 1942, J. A . Steyermark 45058 ( F). Panzós, Finca Mercedes, Teleman , 70 m, 3 September 1988, E . Martínez S . et al. 23472 ( MO). El Petén: Fallabón–Yaxha Road , 22 March 1933, C. L . Lundell 2193 ( F). Carmelita , 1 m south of village, on trail to Flores, 24 June 1942, F. E . Egler 42-227 ( F). Tikal National Park, Tikal, in ramonal on Remate road, 19 May 1959, C. L . Lundell 15999 ( US); Ibid., in zapotal on Remate road, 3 July 1959, C. L . Lundell 16142 ( F, G, NY, US); Ibid. , in ramonal covering the ruines, 27 August 1960, E . Contreras 1467 ( F, K, MO, NY). Remate, on Tikal Road, in zapotal, about 12 km NE of the village, 14 May 1960, E . Contreras 942 ( US). Dos lagunas, Ixcanrio, on Aguas Turbias Road, 12 km E , in zapotal, 11 May 1969, E . Contreras 8517 ( F). Laguna Yaxja, Banks of Laguna Yaxja and dry wooded hills of limestone to north and south, 160–250 m, 28 June 1970, W. E . Harmon & J . D. Dwyer 2772 ( MO). Dolores, Rio Mopan , 21 February 1971, E . Contreras 10583 ( MO, NY). Santa Elena, en la orilla del camino para San Andrés , a km 17, lado noreste, 25 May 1971, R. T . Ortíz 1782 ( F, MO, NY, US). Ca. 5 miles S of Tikal, logging road 0–4 miles W of road to Tikal , 19 June 1973, A. H . Gentry 8346 ( MO, US). Izabal: Trail from Los Amates to Izabal , 31 May 1919, S. F . Blake 7802 ( US). Vicinity of Quiriguá , 75–225 m, 15–31 May 1922, P. C . Standley 24535 ( MO, NY, US). MEXICO. Yucatán, September 1935, R. S . Flores s.n. ( F 782747 ). Campeche: Calakmul, A 500 m al NW de Dos Naciones , 190 m, 23 October 1997, E . Martínez S . et al. 29324 (NY); A 5 km al E de La Mancolona , 200 m, 17 November 1997, E . Martínez S . et al. 29720 (NY); 6.5 km al O de Flores Magón , 153 m, 12 March 2002, J. C . Soto et al. 22687 ( MBM); A 6.7 km al E de La Nueva Vida , 302 m, 13 May 2003, D. Álvarez 5026 ( MO, SPF); A 4.3 km al O del poblado Flores Magón , 182 m, 3 June 2003, D. Álvarez 5219 ( MBM, MO); A 7.6 km al E del poblado "Unión 20 de Junio" (antes la Mancolona ), 160 m, 2 August 2003, D. Álvarez & C . Jiménez J . 6140 (MO). Hopelchén, A 4 km al O del ejido Santa Rosa, sobre el camino al ejido Carlos A . Madrazo , 100 m, 25 September 1996, P . Alvaro M . 605 (BM, MO); A 2.7 km al S de Chan Chen , 12 June 2004, D. Álvarez et al. 8957 ( MO); A 6.64 km al S de Xmejia, 164 m, 15 June 2004, D. Álvarez et al. 9199 ( MBM, MO). Chiapas: Ocosingo, Ruins of Bonampak , 14 March 1975, W. S . Hoover 244 ( MO, US); En el viejo poblado de Jalisco ( Selva Lacandona ), 350 m, 1 December 1976, J. I . Calzada et al. 2907 ( MO); South of Santo Domingo on road to Bonampak and Echeverria from Chancala , 455 m, 25 January 1982, D. E . Breedlove & F . Almeida 57938 ( MO); Siria, a 62 Km al SE de Palenque, 400 m, 12 May 1982, M . Sousa et al. 12397 ( BM, MO); A 9 km al NW de Boca Lacantum, camino a Crucero Corozal–Palenque , 160 m, 16 October 1984, E . Martínez S . 8301 (MO); A 5 km al

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MEDEIROS & LOHMANN

NO de Crucero Corozal, 170 m, 14 June 1985, E . Martínez S . & G . Aguilar 15000bis ( MO); Camino a las ruinas de Bonampak, reserva Montes Azules, 10 km NE del poblado de Lacanjá , 200 m, 26 October 1985, S . S . Colín & G . I . Manriquez 319 ( MO); En Ejido Roberto Bárrios a 60 km al S de Boca Lacantum, camino a Chajul , 200 m, 18 April 1986, E . Martínez S . 18357 (F, MO); A 15 km al NW de Boca Lacantum, camino a Palenque , 220 m, 12 June 1986, E . Martínez S . & M .A. Soto A . 18582 ( F photo, MO); En Nuevo Chihuahua a 70 km al S de Boca Lacantum , 200 m, 23 June 1986, E . Martínez S . 18966 ( F, MO); Borde del río Lacantún, Estación Biológica de Chajul , 150 m, 2 August 1996, G . I . Marínquez et al. 4035 ( MO); Nuevo Guerrero, 200 m, 11 April 2002, G . Aguilar M . 472 (MBM, MO); A 0.2 km de Nuevo Jerusalem camino a Nuevo Francisco León , 12 m, 4 May 2002, G . Aguilar M . et al. 630 ( MBM, MO); 4 May 2002, G . Aguilar M . et al. 685 (MO); A 6.5 km al S de Nuevo Guerrero sobre el camino a Santo Domingo, 400 m, 6 May 2002, D. Álvarez et al. 961 ( M, RB); 0.5 km al S de Nuevo Francisco León ( Restaurante El Paraíso ), 177 m, 10 May 2002, J . C . Soto et al. 23490 ( M, MO); A 1 km al S de El Paraíso , 325 m, 22 May 2002, G . Aguilar M . 992 ( MO); A 600 m al SO de Nuevo Jerusalém , 340 m, 31 May 2002, G . Aguilar M . et al. 1286 (MO); A 200 m de Nuevo Francisco León, restaurante El Paraíso , 174 m, 16 June 2002, G . Aguilar M . et al. 1399 (M, MBM, MO); A 2 km al N de Nuevo Guerrero, 190 m, 22 June 2002, G . Aguilar M . et al. 1561 (M, MO, NY); A 1 km de Nuevo México, al Norte del Poblado , 176 m, 9 July 2002, G . Aguilar M . & F . Aguilar 1805 ( M, MO); A 3.8 km al SE del paraíso, 385 m, 14 October 2002, G . Aguilar M . & D. Álvarez 3449 ( MBM, MO, NY); A 4 km al S de San Javier , 350 m, 26 January 2003, G . Aguilar M . et al. 5206 (MBM, MO, NY, SPF); A 2.1 km al SE de la comunidad Lacanjá Chansayab , 320 m, 17 June 2003, D. Álvarez 5378 ( MO); A 1 km de al NO del poblado " El limonar", 521 m, 30 August 2003, D. Álvarez & A . Chambor 6297 ( MO); A 0.57 km al SO del crucero de Bonampak , 340 m, 15 October 2003, G . Aguilar M . et al. 8178 (MO); A 0.64 km al O del crucero de Bonampak , 313 m, 13 December 2003, G . Aguilar M . et al. 8908 ( MBM). Palenque, Ruinas de Palenque , 160 m, 11 May 1982, M . Sousa et al. 12382 ( BM, MBM, MO). Quintana Roo: A 15 km al Sur de Ejido Laguna OM , 8 June 1980, O . Téllez & E . Cabrera 2423 ( BM, MO, NY). A 8 km al Norte de La Unión , 9 June 1980, O . Téllez & E . Cabrera 2456 ( MO, NY). A 12 km al Sur del Ejido Laguna OM , sobre el camino a Tomas Garrido , 7 August 1980, O . Téllez & E . Cabrera 3054 ( MO, NY). A 7 km al norte de Bacalar , 21 December 1982, E . Cabrera & H . de Cabrera 4139 ( MO). 4 km al NO de Estero Franco, sobre el camino en construccion a Tomas Garrido, lugar llamado El Danto , 17 May 1985, E . Cabrera et al. 8349 ( BM, MO). Adolfo de la Huerta, A 4.2 km al ESE de San Carlos, camino a Pozo Pirata , 102 m, 20 June 2004, D. Álvarez et al. 9517 ( MBM, MO). Othón P . Blanco, A 1.5 km al N de Dos Aguadas , 66 m, 21 June 2004, D. Álvarez et al. 9560 ( MO). A 1.8 km al O de El Martirio, camino a Pozo Pirata, 97 m, 9 September 2004, D. Álvarez et al. 10585 ( BM). Tabasco: Balancán, La Palma , 1–6 June 1939, E . Matuda 3277 ( F, K, MO, NY). Huimanguillo, El 32 rumbo a Díaz Ordaz , 150 m, 15 May 1985, F . Ventura A . 21496 ( MO, NY). Tacotalpa, Disperso a la orilla de la selva Milpa em el cerro cerca del Ejido Lázaro Cárdenas al este del ejido, 50 m, 7 June 1979, A . Orozco et al. 2208 ( MO). Veracruz-Llave: Hidalgotitlán, Hermanos Cedillo , La escuadra por el Rio Soloxuchil, 150 m, 17 August 1974, M . Vasquez 927 ( M). Minatitlán , 2.5 km al N de Poblado 10 por el camino a Pob. 9, 180 m, 9 May 1986, T . Wendt et al. 5280 ( MO) .

Taxonomic notes: — Tynanthus guatemalensis is morphologically similar to the Atlantic Forest species T. micranthus , with which it shares elliptic leaflets with caudate-mucronate apices, lax and conical inflorescences, a similar corolla length (0.5–0.9 mm in both species), and winged fruits. However, T. guatemalensis presents simple tendrils (versus trifid in T. micranthus ), foliaceous prophylls of the axillary buds (versus minute in T. micranthus ) and a pubescent inflorescence axis (versus glabrescent in T. micranthus ). Despite the morphological similarities between those two species, T. guatemalensis is more closely related to T. croatianus , another Central American species but morphologically very distinct from T. guatemalensis (see notes under T. croatianus ).

Nomenclatural notes: —In the protologue of T. guatemalensis, Donnell Smith (1893) did not clearly indicate which of the materials examined was the holotype. Two duplicates of the J. Donnell Smith 1488 collection were located at US, where Donnell Smith worked (Stafleu & Cowan 1985: 698), and the best quality material is here selected as the lectotype .

Figures

FIGURE 8. A–F. Tynanthus fasciculatus: A. Flowering branch; B. Detail of tomentose indumentum in branchlets; C. Calyx with laciniate apex; D. Open corolla showing the androecium; E. Gynoecium (G. Edwall s.n., F 896072); F. Fruit showing the subtetragonal cross section (M. Barreto 1824, F). G–J. T. guatemalensis: G. Flowering branch; H. Interpetiolar region showing the foliaceous prophylls of the axillary buds; I. Flower (E. Contreras 8517, F); J. Fruit (P.H. Gentle 7775, F). Illustration by Klei Sousa.

FIGURE 10. Distribution of Tynanthus guatemalensis.