Celtis alnifolia (Weddell) Miquel (1853: 176)

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 : 65-66

1. Celtis alnifolia (Weddell) Miquel (1853: 176) ( Figures 2A–A 3 , 4A , 5A–B , 6A–B , 7 ).

≡ Momisia alnifolia Weddell (1852: 193) .

Lectotype (designated by Zamengo et al. 2023b):— BRAZIL. Mato Grosso: Cuiabá , “ flumen Cuxipo ” [Coxipó river], November to December 1844, fr., H.A. Weddell 3028 (lectotype: P [00089360] image!; isolectotypes: P [00089361, 00089362] images!).

= Celtis morifolia Planchon (1848: 311) nom. illeg. non Celtis morifolia Rafinesque (1838: 34) . Celtis flavovenarum Zamengo (2023: 212) as a new name for C. morifolia Planchon. Lectotype (designated by Zamengo et al. 2023b):— BRAZIL. Tocantins: Natividade, December 1839, fr., G. Gardner 3426 (lectotype: K [000512922] image!; isolectotypes: BR [0000013534707] image!, K [000512923] image!, U [0007900 fragment] image!).

Scrambling shrubs or scrambling trees, 2–9 m tall; secondary and tertiary branches fawn- brown, sinuous, terete or sulcate, pilose to pubescent, the trichomes ivory-white; thorns 2–7 mm long, in pairs or solitary, semi-curved to straight, buff-yellow or stramineous-yellow, subglabrous or pilose, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns. Leaf: petiole 4–12 mm long, pilose or pubescent, the trichomes ivory-white; leaf blades widely-elliptic, widely-ovate or ovate, 3.5–12 × 2–7 cm, concolorous (emerald-green or olive-green), chartaceous, the apex cuspidate, the base symmetrical, rounded or obtuse, the margins crenate-serrate, serrate or serrulate, teeth congested emerging from the proximal third to the distal third (immature leaves) or from the middle to the distal third (mature leaves), adaxial surface scabrous, opaque, subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, abaxial surface, scabrous, subglabrous or pilose, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, veins protruding, buff-yellow or lemon-yellow, pocket domatia, conspicuous or inconspicuous, subglabrous to pilose, the trichomes ivory-white, dispersed over the entire surface. Cymes paniculiform, peduncles 3–5 mm long, pilose, the trichomes ivory-white or lemon-yellow, bracts absent. Staminate flowers: pedicels 1–2 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow; sepals abaxially pilose or pubescent, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 2–7 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow; ovary 2–4 × 1–3 mm, subglabrous to pubescent, the trichomes ivory-white, throughout or sometimes concentrated only at the base and scarce over the rest of surface, scabrous, the style conspicuous (0.6–2 mm long), the stigmatic branches 2–3 mm long, bifid, the lobes 1–1.5 mm long. Drupe: ovate, 7.5–9 × 5–6 mm, epicarp lemon-yellow, scabrous, glabrous or subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 4.5–5.5 × 3.5–4.5 mm, ivory-white, surface alveolate-crateriform, monoapiculate, the apiculum aciculate, 0.25–0.5 mm long, linear apex apiculum, scar present.

Etymology: —The epithet “ alnifolia ” refers to the leaves similar to those in the genus Alnus Miller (1754: 51) ( Betulaceae ).

Vernacular names: —Canjiquinha, crista de galo, esporão de galo, grão de galo, joá, juá espinheira, juá mirim, rouba tempo, sarã and taleira ( Brazil).

Distribution, habitat and ecology: —Endemic to Brazil, occurring in cerrado and caatinga biomes of the Midwest, Northeast and Southeast ( Figure 7 ). Celtis alnifolia is an heliophilous and inhabits dry forests, gallery forests, anthropic areas with an affinity for sandy, clayey and calcareous soils.

Phenology: —Flowering from September to November and fruiting from January to October.

Taxonomic notes: — Planchon (1873) synonymized C. alnifolia under C. gardneri Planchon (1848: 311) . We consider these species to be distinct, since C. alnifolia has leaf blades with trichomes concentrated on the veins and scarce on the blade surface ( Figure 2A 1 , A 2 ), abaxial surface scabrous; mature drupe lemon-yellow ( Figure 6A ), epicarp scabrous and pyrene with alveolate-crateriform surface ( Figure 6B ), whereas C. gardneri has leaf blades with trichomes on both the veins and the blade surface ( Figure 2E 1 , E 2 ), abaxial surface velvety or smooth; mature drupe fulvous-orange ( Figure 6I ), epicarp smooth and pyrene with alveolate-crateriform-verrucose surface ( Figure 6J ).

Baehni (1936) and Berg & Dahlberg (2001) synonymized C. alnifolia under C. iguanaea . Celtis alnifolia can be differentiated by having pilose to pubescent thorns; mature leaves with margins serrate from the proximal third to the distal third ( Figure 2A ) or emerging from the middle to the distal third, scabrous leaf surfaces; pistillate flowers with conspicuous style ( Figure 5B ); lemon-yellow mature drupes, 7.1–7.5 × 7.9–8.6 mm ( Figure 6A ), pyrenes 4.2–5 × 3.5–4.2 mm with alveolate-crateriform surface ( Figure 6B ), whereas C. iguanaea has glabrous to subglabrous thorns; mature leaves with entire margins ( Figure 3A ) or serrate on distal third ( Figure 9D ), leaf surfaces smooth; pistillate flowers with inconspicuous ( Figure 5N ) or no style; fulvous-orange mature drupes, 8.1–14 × 7.2–13 mm ( Figure 6M ), and pyrenes 4–8 × 4–6 mm with verrucose surface ( Figure 6N ).

Nomenclatural notes: — Celtis flavovenarum was proposed as a new name for C. morifolia Planch. ( Zamengo et al. 2023b) and later determined to be conspecific with C. alnifolia . C. alnifolia is the oldest validly published name and has therefore nomenclatural priority over C. flavovenarum (≡ C. morifolia Planch., ICN Art. 11.3, Turland et al. 2018).

Additional material examined: — BRAZIL. Goiás: Alvorada do Norte, 3 December 2003, fr., G. Pereira-Silva et al. 8053 ( CEN). Luziânia, AHE Corumbá IV, local desmatado para a construção da casa de força, margem esquerda do rio Corumbá, 16°19’52” S, 48°11’26” W, 23 November 2001, fr., M. Carvalho-Silva 94 ( CEN), Fazenda Elias Pena, 9 November 2002, fr., G. Pereira-Silva et al. 6982 ( CEN). Minaçu, Estrada Minaçu canteiro da obra Km 7, 13°26’30” S, 48°12’34” W, 20 September 2001, fl., G. Pereira-Silva et al. 5397 ( CEN). São Domingos, Fazenda São Domingos, 13°36’10” S, 46°45’37” W, 11 October 1999, fl., A.C. Sevilha & S.C.S. Xavier 1859 ( CEN), Fragmento intacto da fazenda São Domingos, 13°37’47” S, 46°44’31” W, 31 October 2000, fl. & fr., A.C. Sevilha 2006 ( CEN), Fazenda São Domingos, 13°37’45” S, 46°44’4” W, 11 March 2004, fr., A.A. Santos et al. 2315 ( CEN). Mato Grosso: Coxipó da Ponte, Rio Coxipó, Avenida Fernando Correa da Costa, em baixo da ponte, rio Coxipó, margem esquerda, 15º37’33” S 56º3’32” W, 24 April 2022, H.B.Z. Souza et al. 220 ( RB). Tocantins: Natividade, Bloco praia, próximo ao córrego prainha, chácara do casal Albany Nunes Cerqueira & Denise Viana Camila, 31 July 2019, fr., H.B.Z. Souza 130 ( PMSP, RB). Palmeirópolis, Linha de transmissão São Salvador – Canabrava, entre as torres 77 e 79, fazenda do Sr. Joaquim, fazenda Esplanada, 13°9’8” S, 48°13’18” W, 6 November 2009, fr., B.M.T. Walter et al. 5918 ( CEN).

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.

FIGURE 9. Main characteristics of Celtis iguanaea (Jacquin) Sargent.Photographs taken in Curaçao.—A. Habit, scale bar 2 m; B. Branch and thorns, scale bar 1 cm; C. Immature leaves, scale bar 2 cm; D. Mature leaves with teeth restricted to the upper third, scale bar 4 cm; E. Mature leaf, margin entire, scale bar 2 cm; F. Imature paniculiform cyme, scale bar 1 mm; G. Mature paniculiform cyme, scale bar 2 mm; H. Pistillate flowers, scale bar 4 mm; I. Staminate flowers, scale bar 1 mm; J. Yellow drupe in the ripening process, scale bar 4 mm; K. Fully ripe orange drupe, scale bar 6 mm; L. Pyrene, scale bar 4 mm. [Photos: A–K, E.A.T. Houtepen; L, H.B. Zamengo]