Celtis orthacanthos Planchon (1848: 309)

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 : 80-81

8. Celtis orthacanthos Planchon (1848: 309) ( Figures 3B–B3 , 4G , 5O–P , 6O–P , 7 ).

Lectotype (designated by Zamengo et al. 2023b):— BRAZIL. Bahia: “ in collibus ”, 1831, fl. & fr., P.Salzmann s.n. (lectotype:K [000512940] image!; isolectotypes: E [01080234] image!, G [00354635] image!, K [000964265 lower branch] image!, MO [1406850] image!, MPU [017540, 017541, 017542, 017543, 017544] images!, P [00089365, 00089366, 00089367] images!, R [010027399] image!).

= Celtis glycycarpa Martius ex Miquel (1853: 174) .

Lectotype (designated by Zamengo et al. 2023b):— BRAZIL. Minas Gerais: “ Near Itambé and Duas Pontes ”, 1839, fl., J.B.E. Pohl s.n. (lectotype: BR [0000013540890] image!; isolectotypes: U [0238802, fragment] image!) . Syntype:— BRAZIL. Minas Gerais: “ Praesidium S. Joan Baptistae ”, April, fr., C.F.P. von Martius 1092 (M [destroyed, photos in F [IRN number 246150, image!] and MO [1409400] image!, U [0007901 fragment] image!).

Scrambling shrubs, 3–12 m tall; secondary and tertiary branches chestnut-brown or cinereous-gray, sinuous, terete, glabrous to pilose, the trichomes ivory-white; thorns 3–20 mm long, in pairs or solitary, curved, semi-curved or straight, buff-yellow or maroon-red, glabrous to pilose, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns or scarce throughout. Leaf: petiole 4–10 mm long, subglabrous to pilose, the trichomes ivory-white; leaf blades elliptic or widely-elliptic, 4–12.5 × 1.5–5.5 cm, concolorous (buff-yellow, emerald-green, cinnamon-brown, coppery-brown, olive-green or stramineous-yellow), chartaceous or membranous, the apex acuminated, the base symmetrical, obtuse, rounded or subcordate, the margins entire, serrate or serrulate, congested teeth emerging from the proximal third to the distal third (immature and mature leaves), from the middle to the third (mature leaves) or restricted to the distal third (mature leaves), adaxial surface smooth, lustrous, glabrous to subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, 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 or inconspicuous, glabrous to pilose throughout. Cymes paniculiform, peduncles 5–11 mm long, pilose, the trichomes ivory-white, bracts present. Staminate flowers: pedicels 0.5–1 mm long, pilose, the trichomes ivory-white, sepals abaxially glabrous to subglabrous, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 1–3 mm long, pilose, the trichomes ivory-white; ovary 1.5–3 × 1–2 mm, subglabrous to pilose throughout, the trichomes ivory-white, velvety or scabrous, the style conspicuous (0.6–1 mm long) or null, the stigmatic branches 5–6 mm long, bifid, the lobes 3–4 mm long. Drupe: globose or ovate, 11.5–13 × 9.5–10.5 mm, epicarp primrose-yellow, smooth, glabrous or subglabrous, the trichomes ivory-white; mesocarp viscous, membranous, not ornamented; pyrene ovate, 7–9 × 5–6 mm, ivory-white, surface alveolate-crateriform, monoapiculate, the apiculum aciculate, 1–1.5 mm long, linear apex apiculum, scar present.

Etymology: —The epithet “ orthacanthos ” refers to the straight thorns present on the branches of the species.

Vernacular names: —Coatindiba, corindiba, corindiuba, corubá, cotindiba, curubá, esporão, esporão de galo, grão de galo, juá, juazeiro, tala, talã and taleiro ( Brazil).

Distribution, habitat and ecology: —Endemic to Brazil with records from the Atlantic Forest ( Figure 7 ). It occurs in high-altitude grasslands, “capoeiras”, on hillsides, in dry forests, secondary forests, on riverbanks, “matas de tabuleiro” and sandbanks. Celtis orthacanthos has an affinity for clayey and sandy soils. Heliophilous or sciophilous, growing in environments with very high and very low light.

Phenology: —Flowers from August to July and fruits all year round.

Taxonomic notes: —Based on the brown color of the leaves, Planchon (1873) synonymized C. orthacanthos under C. aculeata var. laevigata . After analyzing the protologue and specimens from the type localities ( C. orthacanthos from Bahia and C. aculeata var. laevigata from Venezuela), we concluded that C. aculeata var. laevigata and C. orthacanthos should be considered distinct. Celtis aculeata var. laevigata should be synonymized under C. iguanaea , and C. orthacanthos should be recognized as a distinct species.

Celtis iguanaea and C. orthacanthos have similar leaf and flower morphologies, but can be distinguished based on fruits and pyrenes. Celtis iguanaea has fulvous-orange mature drupes ( Figure 6M ) and verrucose pyrene surface ( Figure 6N ), whereas C. orthacanthos has primrose-yellow mature drupes ( Figure 6O ) and alveolate-crateriform pyrene surface ( Figure 6P ).

Berg & Dahlberg (2001) recognized C. orthacanthos , however their characterization should be disregarded, because all the specimens mentioned and their respective characteres refer to C. atlantica ( Zamengo et al. 2024a) . With regard to C. glycycarpa, Baehni (1936) synonymized it under C. triflora and Berg & Dahlberg (2001) synonymized it under C. iguanaea . Celtis iguanaea has fulvous-orange mature drupes ( Figure 6M ), whereas C. orthacanthos (= C. glycycarpa , Figure 6O ) and C. triflora ( Figure 6A 1 ) have primrose-yellow mature drupes. After comparing specimens from Bahia and Minas Gerais (localities of the types of C. orthacanthos and C. glycycarpa respectively), we conclude that C. glycycarpa should be synonymized under C. orthacanthos . Celtis orthacanthos (= C. glycycarpa ) differs from C. triflora because it has 5–6 mm long stigmatic branches ( Figure 5P ), 3–4 mm long stigmatic lobes ( Figure 5P ); pyrenes with scar, 1–1.5 mm long distal apiculum, and alveolate-crateriform surface ( Figure 6P ), whereas C. triflora has the 4.5–8 mm long stigmatic branches ( Figure 5Z ), 4–6 mm long stigmatic lobes ( Figure 5Z ); pyrene without scar, 1–2.5 mm long distal apiculum, and cristed-verrucose surface ( Figure 6A 2 ).

Additional material examined: — BRAZIL. Bahia: Alcobaça, km 5-8 da rodovia BA 001, trecho Alcobaça- Caravelas, 17°33’ S, 39°12’ W, 4 July 1970, fr., A.J. Ribeiro et al. 20 ( CEPEC, RB). Almadina , rodovia Almadina- Ibitupã , entrada ca. 5km W da sede do município, fazenda Cruzeiro do Sul , Serra dos Sete-Paus , ca. 8km da entrada, 14°44’6” S, 39°41’46” W, 15 January 1998, fr., J.G. Jardim et al. 1246 ( CEPEC, NY, RB), Serra Corcovado , 9,8km SW of Coarci on road to Almadina , then N into fazenda São José , 14°42’21” S, 39°36’12” W, 19 September 2004, fr., W.W. Thomas et al. 14162 ( CEPEC, NY, RB). Amargosa , Recôncavo Sul , entrada da porteira fazenda Acaju , 13°1’ S, 39°36’ W, 27 May 2005, fr., M.A.A. Costa & M.L. Guedes 17 ( ALCB). Amélia Rodrigues , estrada Feira-Salvador , 30-35km de Feira de Santana , 23 March 1994, fl. & fr., F. França & E. Melo 955 ( HUEFS). Ilhéus , CEPEC, Km 22 da BR415, 14º47’5” S, 39º13’48” W, 22 April 2022, fr., H.B.Z. Souza 216 ( PMSP, RB). Espírito Santo: Conceição da Barra , Flona do Rio Preto , trilha do Canavial (sede abandonada, lado direito), 25 July 2019, fr., A. Nepomuceno et al. 870 ( VIES). Minas Gerais: Bela Vista de Minas , duplicação da BR381 Norte , Rio Santa Bárbara , 14 April 2005, fr., E. Tameirão-Neto 3853 ( BHCB, RB). Carangola , rio Carangola , 20°43’ S, 42°1’ W, 28 October 1988, fl., L.S. Leoni s.n. ( GFJP489 , RB736645 , SP276479 ) ; fazenda Santa Clara, 1 June 2008, fr., L.S. Leoni 7129 ( GFJP, RB, VIES). Catas Altas, fazenda do Engenho , 23 January 2004, fr., J.R. Stehmann et al. 3524 ( BHCB, RB). Rio de Janeiro: Teresópolis , Providência , Estrada Rio Bahia, Fazenda do senhor Isaias , estrada de terra após as plantações, borda direita do rio Paquequer , borda de mata, - 22.2664125 S, - 42.9251813 W, 23 May 2021, fr., H.B.Z. Souza & M. Pfister 148 ( RB). São Paulo: Arapeí, fragmento da Baleia, 22°37’23” S, 44°26’36” W, 11 May 2011, fr., G.R. Souza et al. 1399 ( RB, VOLRE) .

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.