Celtis spinosissima (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 : 84-86

11. Celtis spinosissima (Weddell) Miquel (1853: 176) ( Figures 3E–E3 , 4J , 5U–V , 6W–X , 7 ).

≡ Momisia spinosissima Weddell (1852: 195) .

Type:— BRAZIL. Rio de Janeiro, Without a specific location, August 1828, fl., C. Gay Mouret s.n. (holotype: P [00089383] image!).

Scrambling shrubs, 1–15 m tall; secondary and tertiary branches chestnut-brown, fawn-brown or maroon-red, straight or sinuous, sulcate, pilose, the trichomes ivory-white; thorns 2–10 mm long, in pairs or solitary, curved, semi-curved or straight, buff-yellow or fuscous-brown, glabrous to pilose throughout or concentrated at the base and sparse over the rest of the surface, the trichomes ivory-white. Leaf: petiole 5–15 mm long, subglabrous to pilose, the trichomes ivory-white, leaf blades widely-ovate, ovate or oval-lanceolate, 3–15 × 1.5–7.5 cm, concolorous (emerald-green or olive-green), chartaceous or membranous, the apex acuminated, caudate or falcate, the base asymmetric or symmetric, obtuse, rounded or subcordate, the margins crenate-serrate, teeth congested emerging from the proximal third to the distal (immature and mature leaves), adaxial surface smooth, opaque, glabrous to pilose throughout or concentrated on the veins and scarce on the blade surface, the trichomes ivory-white, abaxial surface smooth, subglabrous to pilose, 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 to pilose, the trichomes ivory-white, ciliate. Cymes glomerulate, peduncles 1–1.5 mm long, pilose to pubescent, the trichomes ivory-white, bracts absent. Staminate flowers: pedicels 0.5–1 mm long, pilose to pubescent, the trichomes ivory-white, sepals abaxially subglabrous to pilose, the trichomes ivory-white, margins ciliate. Pistillate flowers: pedicels 1.5–2 mm long, pilose to pubescent, the trichomes ivory-white; ovary 3–4 × 1.5–2 mm, pilose to pubescent throughout,, the trichomes ivory-white, velvety or scabrous, the style conspicuous (0.75–1 mm long), the stigmatic branches 2–6 mm long, bifid, the lobes 0.6–2 mm long. Drupe: globose or ovate, 10–14.5 × 6.5–13 mm, epicarp primrose-yellow, smooth, subglabrous to pilose, the trichomes ivory-white; mesocarp viscous, chartaceous, alveolate; pyrene ovate, 8.5–10 × 5–5.7 mm, ivory-white, verrucose surface with proeminent, rounded, randomly distributed warts, monoapiculate, the apiculum aciculate, 0.5–1 mm long, linear apex apiculum, scar present.

Etymology: —The epithet “ spinosissima ” refers to the abundance of thorns.

Vernacular names: —Anzol de lontra, barra de ferrão, bugro de bode, canjiquinha, carupiá, cipó de juá, coatindiba, corrupiá, corubá, corupiá, crista de galo, espora de galo, espora de pinto, esporão, esporão de gato, fruta de galo, galinha choca, gamelinha, grão de galo, grapiá, guajissara, juá, nhapindá, rompe gibão, rouba tempo, salta-martim, unha de gato, vurupiá ( Brazil), tala amarillo, tala brava, tala gateadora, tala guiadora and tala trepadora ( Argentina).

Distribution, habitat and ecology: —Endemic to South America, with records from Argentina, Brazil, and Paraguay ( Figure 7 ). It inhabits Atlantic forests, dry forests, gallery forests, restingas, savannas, forest edges and secondary forests, and has affinity for sandy soils. Heliophilous or sciophyte, growing under high or low light incidence.

Phenology: —Flowers from August to November, and fruits from November to June.

Taxonomic notes: —From Weddell (1852) to Baehni (1936) C. spinosissima was accepted, being synonymized under C. iguanaea by Berg & Dahlberg (2001). Celtis spinosissima was re-established by Zamengo et al. (2020), because C. iguanaea has mature leaves rounded ( Figures 3A ; 9D, E ), margins entire ( Figures 3A ; 9E ) or with teeth restricted to the distal third (mature leaves, Figure 9D ), abaxial surface glabrous; mature drupes fulvous-orange ( Figure 6M ) with glabrous epicarp, whereas C. spinosissima has mature leaves oval-lanceolate ( Figure 3E ), margins crenateserrated from the proximal third to the apex ( Figure 3E ), abaxial surface pilose; mature drupes primrose-yellow ( Figure 6W ) with pilose epicarp.

To support this re-establishment, we add the following characters that also differentiate these species. Celtis iguanaea has branches cinereous-gray; paniculiform cymes ( Figures 4F ; 9G ); ovary subglabrous to pilose ( Figures 5N ; 9I ), style inconspicuous ( Figures 5N ; 9I ) or null; and mesocarp non-ornamented, whereas C. spinosissima has branches chestnut-brown, fawn-brown or maroon-red; glomerulate cymes ( Figure 4J ); ovary pilose to pubescent ( Figure 5V ), style conspicuous ( Figure 5V ); and mesocarp alveolate (see Figure 8B in Chamorro et al. 2021).

Additional material examined: — ARGENTINA. Jujuy: Ledesma, camino a Valle Grande, Três Cruces, 15 October 1964, fl., A.L. Cabrera & H.A. Fabris 16016 ( CTES). Misiones: 25 de Mayo, Ruta 2, cerca del Río Uruguay, - 27,5451 S, - 54,8263 W, 18 December 2016, fr., D.C. Chamorro et al. 82 ( UNR). Salta, La Caldera, ruta 9 camino de cornisa, - 24, 5135 S, - 65, 3436 W, 2 December 2016, D.C. Chamorro et al. 33 ( UNR). BRAZIL. Alagoas: Quebrangulo, Reserva Biológica Federal da Pedra Talhada, 9° 13’ 40.71” S, 36° 25’ 37.58” O, 26 November 2015, fr., L. Nusbaumer 4832 ( MAC). Bahia: Ilhéus, Área do CEPEC (Centro de Pesquisas do Cacau), Km 22 da rodovia Ilhéus/Itabuna (BR 415) 7 December 1988, fr., T. S. dos Santos 4443 ( ALCB, CEPEC, MBM). Espírito Santo: Cariacica, Reserva Biológica Duas Bocas, Trilha do pescador, margem da represa, 20°16’22” S, 40°28’36” W, 14 February 2008, fr., A.M.A. Amorim et al. 7084 ( CEPEC, MBML, RB). Minas Gerais: Aiuruoca, Serra do Papagaio, 20 June 1943, fr., M. Magalhães 5599 ( IAN). Paraíba:Alagoa Nova, 20 January 1959, fl. & fr., J.C. Moraes 2019 (K, NY, P, US). Paraná:Adrianópolis, Barra Grande, 19 April 1995, fr., J. Cordeiro & J.M. Silva 1213 ( NY, US). Rio de Janeiro: Rio de Janeiro, Alto da Boa Vista, Estrada Dona Castorina, - 22.965444 S, - 43.245957 W, 13 August 2021, fl., H.B.Z. Souza 150 ( RB). Rio Grande do Sul: Caxias do Sul, Galópolis, 24 September 1997, fl., S. Diesel 996 ( ULBRA, US). Santa Catarina: Apiúna, Morro Dom Bosco, 27°03’1” S, 49°22’00” W, 1 April 2017, fr., L.A. Funez 6401 ( FURB). São Paulo: São Paulo, Parque Estadual do Jaraguá, trilha do Mauro, 27 July 2019, fl., H.B.Z. Souza & R.B.Z. Souza 129 ( PMSP), 27 October 2019, fl. & fr.,

H.B.Z. Souza & D.C. Chamorro 134 ( PMSP). Sergipe: Estância, APA Sul, - 11.2546136 S, - 37.4675338 W, 5 March 2012, fr., D.A. Campos et al. 61 ( ASE). PARAGUAY. Alto Paraná: Reserva biológica Itabó, 15 September 1988, fr., G.C. Marmori 1629 ( CTES).

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 8. Epitype of Celtis iguanaea (Jacquin) Sargent [Photo: RB herbarium].

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]