Celtis spinosa 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 : 83-84

10. Celtis spinosa Sprengel (1824: 931) ( Figures 3D–D3 , 4I , 5S–T , 6U–V , 7 ).

Lecotype (designated by Baehni 1936):— BRAZIL. Rio Grande do Sul: Without a specific location, s.d. F. Sellow 536 (lectotype: B [10 0247968] image!; isolectotypes: K [000964296] image!, P [00089381] image!) .

= Celtis bonplandiana Planchon (1873: 190) .

Type:— ARGENTINA. Corrientes: Without a specific location, s.d., fr., A.J.A.G. Bonpland 943 (holotype: P [00089351] image!).

= Celtis tala Gillies ex Planchon var. gaudichaudiana Planchon (1873: 191) .

Type:— BRAZIL. Rio Grande do Sul: Without a specific location, 1833, fr., C. Gaudichaud-Beaupré 1732 (holotype: P [00089382] image!).

Scrambling shrubs, 1–7 m tall; secondary and tertiary branches cinereous-gray, sinuous, terete, glabrous to subglabrous, the trichomes ivory-white; thorns 5–17 mm long, in pairs or solitary, curved, semi-curved or straight, buff-yellow, cinereous-gray or stramineous-yellow, glabrous to subglabrous, the trichomes ivory-white, scarce throughout. Leaf: petiole 3–4 mm long, pilose, the trichomes ivory-white, leaf blades oblong, 2–5 × 1.5–2.5 cm, concolorous (emerald-green or olive-green), chartaceous, the apex acuminated or cuspidate, the base symmetrical, cuneate, obtuse or subcordate, the margins crenate-serrate or serrate, teeth congested emerging from the middle to the distal third (immature and mature leaves) or restricted to the distal third (mature leaves), adaxial surface smooth, lustrous, 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, buff-yellow or chestnut-brown, pocket domatia, conspicuous, subglabrous, the trichomes ivory-white, ciliate. Cymes glomerulate, peduncles 1–2 mm long, subglabrous to pilose, the trichomes ivory-white, bracts absent. Staminate flowers: pedicels 0.25–0.5 mm long, subglabrous to pilose, the trichomes ivory-white, sepals abaxially subglabrous, the trichomes ivory-white, the margins entire. Pistillate flowers: pedicels 1–2 mm long, subglabrous to pilose, the trichomes ivory-white; ovary 2.5–3 × 1.5–2 mm, pilose throughout or concentrated at the base and scarce over the rest of the surface, the trichomes ivory-white, scabrous, the style conspicuous (0.6–1 mm long), the stigmatic branches 1.5–2 mm long, bifid, the lobes 0.6–2 mm long. Drupe: globose or ovate, 8.4–10 × 8.4–10 mm, epicarp fulvous-orange, smooth, glabrous or subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 4.5–6 × 4–4.5 mm, ivory-white, verrucose surface with proeminent, rounded, randomly distributed warts, monoapiculate, the apiculum attenuate, 0.1–0.3 mm long, linear apex apiculum, scar absent.

Etymology: —The epithet “ spinosa ” refers to the thorns found on the branches of the species.

Vernacular names: —Espora de galo, esporão de galo, grão de galo, grapiá, grapiá branco, grupiá, tala, taleira, tobeira, unha de gato ( Brazil), tala gateadora and tala trepadora ( Argentina).

Distribution, habitat and ecology: —Endemic to southern South America, with a restricted occurrence in southern Brazil (Rio Grande do Sul and Santa Catarina states), northeastern Argentina, southern Paraguay, and Uruguay ( Figure 7 ). It inhabits roadsides, forest edges, savannas and riparian forests, and has an affinity for sandy and clayey soils. In Rio Grande do Sul, the species also was collected in a restinga area. Heliophilous or sciophyte, growing under high or low light incidence.

Phenology: —Flowering from September to December and fruiting from November to April.

Taxonomic notes: — Baehni (1936) synonymized C. tala under C. spinosa . Romanczuk (1976) reestablished C. tala , indicating that C. spinosa is a scrambling shrub, whereas C. tala is a non-scrambling tree. Dottori & Hunziker (1994) synonymized C. spinosa under C. iguanaea , which was corroborated by Berg & Dahlberg (2001).

Celtis spinosa was re-established by Torres & Luca (2005) and recognized for the state of Rio de Janeiro by Pederneiras et al. (2011). After comparing the specimens cited by both Torres & Luca (2005) and Pederneiras et al. (2011), we concluded that none correspond to C. spinosa , but rather to C. alnifolia and C. clausseniana ( Torres & Luca 2005) and C. brasiliensis ( Pederneiras et al. 2011) . We therefore suggest disregarding the categorization of C. spinosa by these authors.

Berg & Dahlberg (2001) synonymized C. spinosa under C. iguanaea . These species should be considered distinct because C. iguanaea has mature leaf blades up to 12 cm long, elliptical or wide-elliptical ( Figures 3A ; 9D, E ); paniculiform cymes ( Figures 4F ; 9G ); mature drupes up to 14 mm long ( Figure 6M ), and pyrenes up to 8 mm long ( Figure 6N ), whereas C. spinosa has mature leaf blades up to 5 cm long, oblong ( Figure 3D ); glomerulate cymes ( Figure 4I ); mature drupes up to 10 mm long ( Figure 6U ), and pyrenes up to 6 mm long ( Figure 6V ).

Additional material examined: — ARGENTINA. Buenos Aires: Parks gardens and environs of the city of Buenos Aires, 1913, fr., H.M. Curran s.n. ( US 03414966). La Plata, Punta Lara, 3 December 1971, fr., J.P. Carauta 1418 ( RB). Isla Martín García, reserva Natural y sitio histórico Isla Martin García, 30 October 2004, fl., S.T. Robles et al. 2024 ( LP, US). Corrientes: Capital, Puente Pexoa, 16 January 1974, fr., C. Romanczuk 852 ( SI, US). Entre Ríos: Arroyo Martínez, Delta del Paraná, 14 October 1944, fl., O. Boelcke 1030 ( LIL, US). BRAZIL. Rio Grande do Sul: Arambaré, 27 September 2003, fl., G. Schlindwein s.n. ( ICN 128232). Arroio Grande, Pontal, próximo ao farol Ponta Alegre, 4 October 1997, fl., J.A. Jarenkow & E.N. Garcia 3566 ( ESA, MBM, PEL). Barra do Guaraí, Espinilho, 17 November 1984, fr., M. Sobral 3391 ( ICN, PAMG, RB, SP). Osório, Estrada Pontal dos Diehl , nº860, 29º49’55” S 50º07’58” W, 12 April 2022, fl. & fr., H.B.Z. Souza et al. 196, 197, 198, 199, 200, 201 ( PMSP, RB). Santa Catarina: Anita Garibaldi, passo do rio Canoas, 12 April 1963, fr., R. Reitz & R.M. Klein 14753 (R). Araranguá, lagoa da Serra, 13 June 1995, V. Boff 13 ( CRI), Morro dos Conventos , 28°56’4” S, 49°21’49” W, 26 December 2016, fr., L.A. Funez 5823 ( FURB). PARAGUAY. Central: Granja Ysapy, 29 January 1966, fr., A. Krapovickas et al. 12197 ( CTES). Neembucú, Curupayty, Humaitá, 9 November 1978, fl., Bernardi 18423 (U). URUGUAY. Rocha: Fuerte San Miguel, 21 February 1960, fr., Boelcke 8303 ( BAA).

Figures

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]