Celtis pubescens Sprengel (1824: 931)

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 : 81-83

9. Celtis pubescens Sprengel (1824: 931) ( Figures 3C–C3 , 4H , 5Q–R , 6Q–T , 7 ).

≡ Mertensia pubescens Kunth (1817: 26) nom. illeg. non Mertensia pubescens Humboldt & Bonpland ex Willdenow (1810: 73) . Momisia pubescens (Kunth) F. Dietrich (1819: 123) nom. illeg.

Lectotype (designated [as type] by Baehni 1936, second step of lectotypification designated by Zamengo et al. 2024b):— ECUADOR. Guayas: Guayaquil, February 1802, fr., A.J.A.G. Bonpland & F.W.H.A. von Humboldt 3792 (lectotype, P [00307271] image!; isolectotypes: B [-W 04666 -01 0] image!, P [06815234, 00307270] images!).

= Momisia brevifolia Klotzsch (1847: 538) . Celtis brevifolia (Klotzsch) Miquel (1853: 180) .

Type:— ECUADOR. Guayas: Guayaquil, s.d., fl., H. Ruiz López s.n. (holotype: B [10 0247965] image!).

= Celtis goudotii Planchon (1848: 312) .

Type:— COLOMBIA. Tolima: “ between Ibagué and Rusaguasuga ”, s.d., J. Goudot s.n. (holotype: K [000575977] image!; isotype: P [06781653] image!) .

= Celtis velutina Planchon (1848: 313) .

Lectotype (designated by Zamengo et al. 2024b):— PERU. Junin: Huancayo, Quebrada de Pariahuanca , s.d., fl., Matthews 826 (lectotype: K [000575981] image!; isolectotypes: E [00641649] image!, K [000575982] image!).

Scrambling shrubs, 4–40 m tall; secondary and tertiary branches fuscous-brown, sinuous or straight, terete, pubescent to velutinous, the trichomes chestnut-brown or lemon-yellow; thorns 2–8 mm long, in pairs or solitary, curved, semi-curved or straight, fuscous-brown, pubescent to velutinous throughout, the trichomes chestnut-brown, concentrated both at the base and on the entire surface of the thorns. Leaf: petiole 2–5 mm long, pubescent to velutinous, the trichomes chestnut-brown or lemon-yellow, leaf blades elliptic, ovate or ovate-elliptic, 6–8.5 × 2.5–4 cm, concolorous (olive-green), chartaceous, the apex acuminated, the base symmetrical, acute, rounded, obtuse or subcordate, margins entire or serrulate, teeth congested emerging from the middle to the distal third (immature and mature leaves) or restricted to the distal third (mature leaves), adaxial surface scabrous, opaque, pilose to pubescent throughout, the trichomes chestnut-brown, ivory-white or lemon-yellow, abaxial surface velvety, pubescent to velutinous throughout, the trichomes chestnut-brown or lemon-yellow, veins protruding, chestnut-brown, tuft domatia, conspicuous, velutinous throughout, the trichomes lemon-yellow. Cymes glomerulate, peduncles 3–6 mm long, velutinous, the trichomes chestnut-brown or lemon-yellow, bracts absent. Staminate flowers: pedicels 0.5–1 mm long, velutinous, the trichomes chestnut-brown or lemon-yellow, sepals abaxially velutinous, the trichomes chestnut-brown or lemon-yellow, margins ciliate. Pistillate flowers: pedicels 0.5–1 mm long, velutinous, the trichomes chestnut-brown or lemon-yellow; ovary 0.5–1 × 0.5–1 mm, pubescent, the trichomes lemon-yellow, concentrated at the base and scarce over the rest of the ovary, velvety, the style conspicuous (0.6–1 mm long), the stigmatic branches 1–2 mm long, bifid, the lobes 1–1.5 mm long. Drupe: globose or ovate, 8–10 × 4–6 mm, epicarp lemon-yellow, velvety or scabrous, pilose to pubescent, the trichomes lemon-yellow; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 5–6.5 × 4–4.5 mm, ivory-white, surface alveolate-crateriform, monoapiculate, the apiculum aciculate, 1–2 mm long, linear apex apiculum, scar present.

Etymology: —The epithet “ pubescens ” refers to the indumentum of branches and leaves.

Vernacular names: —Cipó farinha seca, esporão de galo, farinha seca and laranjinha cipó ( Brazil).

Distribution, habitat and ecology: —Endemic to the Amazon region ( Bolivia, Brazil [Acre state], Colombia, Ecuador, Peru and Venezuela, Figure 7 ), growing in primary and secondary forests, on roadsides and riverbanks with and affinity for clayey soils. Heliophilous or sciophilous, growing under high or low incidence of light.

Phenology: —Flowers in September and bear fruits from October to January.

Taxonomic notes: — Baehni (1936) highlighted the yellow color of the trichomes present at the bifurcations of the abaxial surface of the leaf blades, but was unable to differentiate C. pubescens from other taxa that also have abaxial leaf surfaces velvety. As a result, several taxa were synonymized under C. pubescens . Baehni (1936) indicated a large distribution range of C. pubescens including Argentina, Bolivia, Brazil, Colombia, Ecuador, Paraguay, and Peru, and presented a great variability of habits, which would justify the different taxa described over the years. This author mentioned that C. pubescens presents transitory characters such as entire or serrate leaf margins and the type of indumentum (pilose to velutinous) varying on the same individual according to the age and height of the collected branch.

Baehni (1936) also mentioned that the fruits are useless for the differentiation of species, which by other authors ( Hunziker & Dottori 1976, Sattarian & Van Der Maesen 2006, Zarafshar et al. 2010, Zamengo et al. 2020, 2021, Chamorro et al. 2021, Chamorro 2022) is not supported, because fruits and pyrenes are indeed crucial for the differentiation of species. The circumscription of C. pubescens by Baehni (1936) should be disregarded because the lack of field observations and the neglect of characters related to fruits led him to synonymize most taxa that showed any level of indumentum. Some of these (e.g., C. brasiliensis and C. clausseniana ) should be considered independent, while others should be synonymized into distinct taxa.

Baehni’s circumscription was adopted by different authors ( Baehni 1937, Romanczuk & Martínez 1978, Dottori & Hunziker 1994, Marchioretto 1988, Rocha et al. 2000, Torres & Luca 2005). As a result, C. pubescens became frequent in herbarium collections.

Berg & Dahlberg (2001) synonymized C. pubescens under C. iguanaea . Torres & Luca (2005) re-established C. pubescens . After comparing the type of C. pubescens , other specimens from the Amazon region, and the specimens mentioned by Torres & Luca (2005), we conclude that the specimens mentioned by these authors should be identified as C. clausseniana . Celtis clausseniana and C. pubescens have abaxial leaf surfaces velvety, which is one of the main characters used by Torres & Luca (2005) for species identification (see Torres & Luca 2005). In addition to this character, the same authors mentioned that “ C. pubescens ” has domatia in pockets, Baehni (1936), however, mentioned tufts of yellowish trichomes along the bifurcations of the leaf veins, which matches our own observations. Based on these observations, we suggest to update the characterization of “ C. pubescens ” senso Torres & Luca (2005) to C. clausseniana .

The synonymization of C. pubescens with C. iguanaea is easily refuted, because different authors ( Jacquin 1763, Lamarck 1789, Planchon 1848, Miquel 1853, Planchon 1873, Boldingh 1913) who worked with the Caribbean populations of C. iguanaea highlighted that the species has a glabrous to subglabrous abaxial leaf surface ( Figure 3A 2 ), whereas C. pubescens has a pubescent to velutinous ( Figure 3C 2 ) abaxial leaf surface ( Kunth 1817, Dietrich 1819, Planchon 1848, Miquel 1853, Planchon 1873, Baehni 1936). In addition to this character, we highlight that C. pubescens has a scabrous adaxial leaf surface, tuft domatia ( Figure 3C3 ); glomerulate cymes ( Figure 4H ); lemon-yellow mature drupes ( Figure 6Q ), pilose to pubescent epicarp ( Figure 6Q, R ) covered with lemon-yellow trichomes ( Figure 6S ), and alveolate-crateriform pyrene surface ( Figure 6T ), whereas C. iguanaea has smooth adaxial leaf surface, pocket domatia ( Figure 3A3 ); paniculiform cyme ( Figure 4F ); fulvous-orange mature drupes ( Figure 6M ), glabrous to subglabrous epicarp with ivory-white trichomes ( Figure 6M ), and verrucose pyrene surface ( Figure 6N ).

Additional material examined: — BOLIVIA. Pando: Manuripi, 35 Km al norte de Puerto América entrando bosque alto cerca Alianza, 11º44’S, 67º59’W, 2 May 1994, fr., A. Jardim 639 ( MO, SI). Nicolás Suárez, en la zona de Campoana, junto a la barraca San José, 16 January 1983, fr., Fernández Casas & Susanna 8308 ( HUA, TEFH). BRAZIL. Acre, Acrelândia, Basin of Rio Madeira, Rio Abunã, Porto Dias, Km 130 of BR 130 of BR-364, then 30 km on Ramal do Pelé, then Ramal do Gordo, colocação Bom Jardim I, 9°58’38” S 66°47’35” W, 16 May 2005, fr., D.C. Daly et al. 13709 ( NY, RB, UFACPZ). Sena Madureira, Bacia do rio Purus, fazenda Nova Olinda, margem direita do rio Iaco, carreador São Bento I, ca. 15 km da sede, 10°7’ S, 69°13’ W, 26 October 1993, fr., M. Silveira et al. 656 ( NY, UFACPZ), Rio Macauã, reserva extrativista de Macauã, Seringal Capital, colocação extrema, 9°24’ S, 68°54’ W, 3 April 1994, fr., L. de Lima et al. 583 ( NY, UFACPZ). Senador Guiomard, fazenda experimental Catuaba, BR 364, Km 35, 10°4’ S, 67°37’ W, 1 November 2008, fr., H. Medeiros et al. 126 ( NY, RB, SP, U, UFACPZ), February 2022, fr., D.F. Silva et al. 362 ( RB). Xapuri, Rio Acre 3 hours by boat downstream from Xapuri and 1 hr walking inland from left bank, 10°45’S 68°20’W, 4 November 1991, fr., D.C. Daly et al. 7124 ( NY, UFACPZ). COLOMBIA. Tocaima: December 1932, fr., E.P.A. Perez 2132 ( COL, US). ECUADOR. El Oro: near junction of Rio Luis & Rio Ambocas, 10 km due south of Portovelo, 6 October 1944, fl., I.L. Wiggins 10897 ( US). PERU. Pasco: Oxapampa, Distrito de Pozuzo, carretera Yanahuanca a Tingo Mal Paso, 10°2’20” S 75°34’57” W, 26 May 2009, fr., R. Vásquez et al. 35779 ( HOXA, HUT, MO, USM). Huancavelica: Taycaja, Colcabamba, cerca de la línea de transmisión en parches de bosque seco, 12°23’55” S, 74°42’9” W, 27 September 2013, fl., R. Fernandez-Hilário 450 ( RB). VENEZUELA. Bolívar: Santa Elena, Mata Cutia, as margens do igarapé, 11 September 1979, fr., N.A. Rosa & O.C. Nascimento 3384 ( INPA, MG, NY).

Figures

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.