Celtis triflora (Ruiz ex Klotzsch) Miquel (1853: 181)

Zamengo, Henrique Borges, Chamorro, Débora C., Houtepen, Erika. T., Gaglioti, André Luiz, Pederneiras, Leandro Cardoso, Prado, Darién E. & Oakley, Luis J., 2025, Taxonomic revision of the Celtis iguanaea complex (Cannabaceae), Phytotaxa 689 (1), pp. 53-98 : 88-89

13. Celtis triflora (Ruiz ex Klotzsch) Miquel (1853: 181) ( Figures 3G–G3 , 4L , 5Y–Z , 6A 1–A 2 , 7 ).

≡ Momisia triflora Ruiz ex Klotzsch Linnaea 20: 537 (1847).

Lectotype (designated by Zamengo et al. 2024b):— PERU. Pasco: Oxapampa, Pozuzo, s.d., fr., H. Ruiz-López & J.A. Pavón y Jiménez s.n. (lectotype: MA [811136] image!; isolectotypes: G [00412625 sheet 1/2, 00412625 sheet 2/2] images!, HAL [0110269] image!, MA [811135, 818705] images!, P [00089388] image!).

= Celtis asperula Miquel (1853: 176) .

Lectotype (designated by Zamengo et al. 2023b):— BRAZIL. Pará: Without a specific location, s.d., fr., C.F.P. von Martius s.n. (lectotype: U [0007899, fragment] image!) . Epitype (designated by Zamengo et al. 2023b):— BRAZIL. Pará: Itaituba, Rio Tapajós, 22 September 1922, fr., A. Ducke s.n. (RB [00439255] image!).

= Celtis williamsii Rusby (1901: 497) .

Lectotype (designated by Zamengo et al. 2023c):— BOLIVIA. La Paz : San Buenaventura, 30 November 1901, fl., R.S. Williams 603 (lectotype: NY [00133652] image!; isolectotypes: K [000575984] image!, NY [00133651] image!, US [00089699] image!).

Scrambling shrubs, 2–15 m tall; secondary and tertiary branches chestnut-brown, sinuous or straight, terete or sulcate, glabrous to pilose, the trichomes ivory-white; thorns 1–15 mm long, in pairs or solitary, curved, semi-curved or straight, maroon-red or stramineous-yellow, glabrous to subglabrous, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns or scarce throughout. Leaf: petiole 2–10 mm long, subglabrous to pilose, the trichomes ivory-white, leaf blades elliptic, widely-elliptic, widely-ovate or ovate, 4.5–13 × 2–6.5 cm, concolorous (buff-yellow, cinnamon-brown, coppery-brown, emerald-green, olive-green or stramineous-yellow), chartaceous, the apex acuminated or cuspidate, the base asymmetric or symmetric, rounded or subcordate, the margins crenate-serrate, serrate or serrulate, teeth congested emerging from the proximal third to the distal (immature leaves), from the middle to the distal third (mature leaves) or restricted to the distal third (mature leaves), adaxial surface smooth, lustrous or opaque, glabrous to subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, abaxial surface smooth, subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, veins protruding, chestnut-brown, pocket domatia, inconspicuous, glabrous to subglabrous throughout. Cymes paniculiform, peduncles 4–6 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow, bracts present. Staminate flowers: pedicels 1–2 mm long, subglabrous to pilose, the trichomes ivory-white, sepals abaxially glabrous, the trichomes ivory-white, margins ciliate. Pistillate flowers: pedicels 1–2.5 mm long, pilose, the trichomes ivory-white or lemon-yellow; ovary 2–3 × 1.5–2 mm, subglabrous to pilose, the trichomes ivory-white, throughout or concentrated at the base and scarce over the rest of the surface, scabrous, the style conspicuous (0.6–1 mm long), inconspicuous (0.1–0.5 mm long) or null, the stigmatic branches 4.5–8 mm long, bifid, the lobes 4–6 mm long. Drupe: globose or ovate, 10–17 × 5–8 mm, epicarp primrose-yellow, smooth, subglabrous to pilose, the trichomes ivory-white; mesocarp viscous, membranous, not ornamented; pyrene elliptic or ovate, 6–10 × 5–7.5 mm, ivory-white, cristate-verrucose surface with proeminent crests and randomly distributed acicular or rounded warts, monoapiculate, the apiculum aciculate, 1–2.5 mm long, linear apex apiculum, scar absent.

Etymology: —Unknown.

Vernacular names: —Azedinho, cipó farinha seca, esporão de galo, farinha seca, jatobá do brejo, laranjinha cipó, limãozinho ( Brazil), cagalero, chustustis, escobo, fariña seca, guayaba, maíz tostado, noquichapirr, ojoso negro, perro isma, pomarroso, quechua, sartajchi, satajchi, satajchi amarillo, tsachik, uña de gato, uñe-gato, vara, wayja maek and yahuatáo ( Bolivia, Colombia, Ecuador, Peru and Venezuela).

Distribution, habitat and ecology: —Distributed in northern South America and southern Central America ( Figure 7 ). It inhabits the Amazon rainforest and is found mainly on river banks and has an affinity for sandy and clayey soils. Heliophilous or sciophyte, growing under high or low light incidence.

Phenology: —Flowering and fruiting year-round.

Taxonomic notes: — Celtis triflora was synonymized under C. iguanaea by Berg & Dahlberg (2001). These species should be considered distinct because C. iguanaea occurs in the southern United States, on the Caribbean islands and the Caribbean coast of South America, and northern South America ( Figure 7 ); has branches cinereous-gray, terete; stigmatic branches 1.5–2 mm long ( Figure 5N ) with lobes 0.6–1 mm long ( Figure 5N ); mature drupe fulvous-orange ( Figure 6M ); mesocarp non-viscous; and verrucose pyrene surface ( Figure 6N ), whereas C. triflora occurs in the Amazon region of northern South America ( Figure 7 ); has branches chestnut-brown, sulcate; the stigmatic branches 4.5–8 mm long ( Figure 5Z ) with lobes 4–6 mm long ( Figure 5Z ), mature drupe primrose-yellow ( Figure 6A 1 ), mesocarp viscous, and cristed-verrucose pyrene surface ( Figure 6A 2 ).

Additional material examined: — BOLIVIA. Pando: Without locality, Rio Madre de Díos, 2 October 1923, fr., J.G. Kuhlmann 563 ( RB). BRAZIL. Acre: Acrelândia, Rio Abuna, projeto de assentamento extrativista ( PAE) Porto Dias, km 108 of BR-364 (Rio Branco-Porto Velho), then 30 km S on side road, colocação Palhau, 9°58’46” S, 66°48’5” W, 4 October 2003, fl., D.C. Daly et al. 12156 ( RB). Xapuri, Projeto de assentamento Agroextrativista Cachoeira, BR-117, ramal da cachoeira, arredores da pousada, 10°49’32” S, 68°25’2” W, 2 November 2011, fr., H. Medeiros et al. 892 ( RB). COLOMBIA. Santander: Girón, sector Cruces, Muro Presa, Antigua vía Barranca, Bucaramanga, 7°5’31” N, 73°23’19” W, 25 November 2017, fr., H. David et al. 5563 ( HUA). ECUADOR. Napo: Ahuano, Reserva Biológica Jatun Sacha, Río Napo, 8 km abajo de Misahualli, 1°4’ S, 77°36’ W, 6 February 1987, fr., C. Cerón 730 ( MO, US). FRENCH GUYANA. Cayenne Cedex: Remire-Montjoly, Rorota, 23 March 1984, fr., D. Sabatier 826 ( CAY, P, U). GUYANA. Without a specific location: Western extremity of Kanuku Montains, in drainage of Takutu River, March 1938, fl., A.C. Smith 3237 (K, P). PERU. Junin: Chanchamayo, San Ramón. Sector Nueva Italia, 10°59’20” S, 75°25’15” W, 18 March 2017, fr., L. Valenzuela et al. 31791 ( HOXA, MO, USM). Pasco: Oxapampa, bosque de proteccion San Matias-San Carlos, 10°43’54” S, 74°54’23” W, 6 July 2003, fr., A.M. Monteagudo & K. Meza 5746 ( HOXA, MO). SURINAME. Brokopondo: Albufeira de Brokopondo, NW Brokopondo Stuwmeer lake, SE of Brownsberg Nature Reserve, near mouth of Withey Creek, 4°35’ N, 55°7’ W, 25 February 1998, fr., B. Hoffman & F. van Troon 5276 (U, US). VENEZUELA. Aragua: Girardot, orilla de la carretera vía Punta Palmita, 6 December 1979, fr., H. Rodríguez 965 ( MY).

Figures

FIGURE 1. Lectotype of Rhamnus iguanaea Jacquin. Illustration made by Maria Moninckx (1686-1690) according to Wijnands 1983.

FIGURE 2. Leaves, indumentum and domatia.—A. Celtis alnifolia, scale bar 6 cm, A1. Adaxial surface, with detail of indument; A2. Abaxial surface, with detail of indument; A3. Domatia. B. C. brasiliensis, scale bar 3.5 cm; B1. Adaxial surface, with detail of indument; B2. Abaxial surface, with detail of indument; B3. Domatia. C. C. dichotoma, scale bar 4.5 cm, C1. Adaxial surface, with detail of indument; C2. Abaxial surface with detail of indument; C3. Domatia. D. C. diffusa, scale bar 6 cm, D1. Adaxial surface, with detail of indument; D2. Abaxial surface with detail of indument; D3. Domatia. E. C. gardneri, scale bar 3 cm, E1. Adaxial surface, with detail of indument; E2. Abaxial surface with detail of indument; E3. Domatia. F. C. hilariana, scale bar 2 cm F1. Adaxial surface, with detail of indument; F2.Abaxial surface with detail of indument; F3. Domatia. [Photos: H.B. Zamengo].

FIGURE 3. Leaves, indumentum and domatia.—A. Celtis iguanaea, scale bar 6 cm, A1. Adaxial surface, with detail of indument, A2. Abaxial surface, with detail of indument, A3. Domatia; B. C. orthacanthos, scale bar 4 cm, B1. Adaxial surface, with detail of indument, B2.Abaxial surface, with detail of indument, B3. Domatia; C. C. pubescens, scale bar 4 cm, C1. Adaxial surface, with detail of indument, C2. Abaxial surface, with detail of indument, C3. Domatia; D. C. spinosa, scale bar 2.5 cm, D1. Adaxial surface, with detail of indument, D2. Abaxial surface, with detail of indument, D3. Domatia; E. C. spinosissima, scale bar 6 cm, E1. Adaxial surface, with detail of indument, E2. Abaxial surface, with detail of indument, E3. Domatia; F. C. tarijensis, scale bar 4 cm, F1. Adaxial surface, with detail of indument, F2. Abaxial surface, with detail of indument, F3. Domatia; G. C. triflora, scale bar 6 cm, G1. Adaxial surface, with detail of indument, G2. Abaxial surface, with detail of indument, G3. Domatia. [Photos: H.B. Zamengo].

FIGURE 4. Type of cymes.—A. Celtis alnifolia, paniculiform cyme, scale bar 2 mm; B. C. brasiliensis, glomerulate cyme, scale bar 1 mm; C. C. dichotoma, paniculiform cyme, scale bar 3 mm; D. C. diffusa, glomerulate cyme, scale bar 2 mm; E. C. gardneri, glomerulate cyme, scale bar 2 mm; F. C. iguanaea, paniculiform cyme, scale bar 1 mm; G. C. orthacanthos, paniculiform cyme, scale bar 2 mm; H. C. pubescens, glomerulate cyme, scale bar 2 mm; I. C. spinosa, glomerulate cyme, scale bar 1 mm; J. C. spinosissima, glomerulate cyme, scale bar 1 mm; K. C. tarijensis, glomerulate cyme, scale bar 1 mm; L. C. triflora, paniculiform cyme, scale bar 3 mm. [Photos: A, B, C, D, E, G, J, K, H.B. Zamengo; F, E.A.T. Houtepen; H, R. Fernandez-Hilario; I, A. Gonzáles; L. H. Medeiros].

FIGURE 5. Staminate and pistillate flowers.—A–B. Celtis alnifolia, A. Staminate flower, scale bar 0.5 mm, B. Pistillate flower, scale bar 3 mm; C–D. C. brasiliensis, C. Staminate flower, scale bar 1 mm, D. Pistillate flower, scale bar 3 mm; E–F. C. dichotoma, E. Staminate flower, scale bar 0.5 mm, F. Pistillate flower, scale bar 6 mm; G–H. C. diffusa, G. Staminate flower, scale bar 1 mm, H. Pistillate flower, scale bar 6 mm; I–J. C. gardneri, I. Staminate flower, scale bar 1 mm, J. Pistillate flower, scale bar 1.5 mm; K–L. C. hilariana, K. Staminate flower, scale bar 1 mm, L. Pistillate flower, scale bar 2.5 mm; M–N. C. iguanaea, M. Staminate flower, scale bar 1 mm, N. Pistillate flower, scale bar 4 mm; O–P. C. orthacanthos, O. Staminate flower, scale bar 1 mm, P. Pistillate flower, scale bar 3 mm; Q–R. C. pubescens, Q. Staminate flower, scale bar 1 mm, R. Pistillate flower, scale bar 1 mm; S–T. C. spinosa, S. Staminate flower, scale bar 1 mm, T. Pistillate flower, scale bar 3 mm; U–V. C. spinosissima, U. Staminate flower, scale bar 1 mm, V. Pistillate flower, scale bar 4 mm; W–X. C. tarijensis, W. Staminate flower, scale bar 1 mm, X. Pistillate flower, scale bar 5 mm; Y–Z. C. triflora, Y. Staminate flower, scale bar 1 mm, Z. Pistillate flower, scale bar 3 mm. [Photos: H.B. Zamengo].

FIGURE 6. Mature drupes and pyrenes.—A–B. Celtis alnifolia, A. Mature drupe, scale bar 9 mm, B. Pyrene, scale bar 2.75 mm; C–D. C. brasiliensis, C. Mature drupe, scale bar 10.5 mm, D. Pyrene, scale bar 3.5 mm; E–F. C. dichotoma, E. Mature drupe (type), scale bar 6 mm, F. Pyrene (type), scale bar 3.75; G–H. C. diffusa, G. Mature drupe, scale bar 6 mm, H. Pyrene, scale bar 5 mm; I–J2. C. gardneri, I. Mature drupe, scale bar 3.75 mm, J1. Pyrene (type), scale bar 2.5 mm, J2. Pyrenes, scale bar 2.5 mm; K–L. C. hilariana, K. Mature drupe, scale bar 3.25 mm, L. Pyrene, scale bar 2 mm; M–N. C. iguanaea, M. Mature drupe, scale bar 5 mm, N. Pyrene, scale bar 4 mm; O–P. C. orthacanthos, O. Mature drupe, scale bar 6.5 mm, P. Pyrene, scale bar 4.5 mm; Q–T. C. pubescens, Q. Mature drupe, scale bar 5 mm, R. Mature drupe (type), scale bar 3 mm, S. Indumentum lemon, scale bar 1 µm, T. Pyrene, scale bar 3.25 mm; U–V. C. spinosa, U. Mature drupe, scale bar 5 mm, V. Pyrene, scale bar 3 mm; W–X. C. spinosissima, W. Mature drupe, scale bar 7 mm, X. Pyrene, scale bar 5 mm; Y–Z. C. tarijensis, Y. Mature drupe, scale bar 6 mm, Z. Pyrene, scale bar 5 mm; A1–A2. C. triflora, A1. Mature drupe, scale bar 8.5 mm, A2. Pyrene, scale bar 5 mm. [Photos: H.B. Zamengo].

FIGURE 7. Distribution map of Celtis alnifolia, C. brasiliensis, C. dichotoma, C. diffusa, C. gardneri, C. hilariana, C. iguanaea, C. orthacanthos, C. pubescens, C. spinosa, C. spinosissima, C. tarijensis and C. triflora.