taxonID	type	description	language	source
03A0BD67312A9D6354D5FA6378BAA13C.taxon	materials_examined	Type: — Celtis australis Linnaeus (1753: 1043).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67312A9D6354D5FA6378BAA13C.taxon	materials_examined	Type: — Mertensia laevigata Kunth (1817: 31) nom. illeg., non Mertensia laevigata Willdenow (1810: 75).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67312A9D6354D5FA6378BAA13C.taxon	materials_examined	Type: — Momisia laevigata (Kunth) F. Dietrich (1819: 123), designated by Britton & Wilson (1924). Trees, shrubs, or lianas, monoecious; bark lenticellate, branches sometimes with brachyblasts, thorns if present, then straight, curved, or semi-curved in various colors. Leaves alternate and distichous, chartaceous to membranous, petiole adaxially canaliculate, leaf blade usually widely-elliptic, widely-ovate, oblong or ovate, the apex acuminate, acute, attenuate, cuspidate, or obtuse, the base obtuse, rounded, or subcordate, the margins entire, crenulate, crenate, serrulate or serrate, leaf blade trinerved from the base, adaxially opaque or lustrous, smooth or scabrous, abaxial surface velvety, scabrous or smooth, leaf blades concolorous or discolorous, pocket or tuft domatia if present, then mostly in the abaxial axils of secondary, sometimes also os higher level veins. Cymes glomerulate or paniculiform, peduncle sometimes with bract. Staminate flowers light green, pedicellate, glomerulate, (4 –) 5 (– 6) sepals and stamens. Pistillate flowers light green, ovary ovate, style conspicuous, inconspicuous or null, stigmatic lobes entire, bifid, bilobed or trifid. Drupe globose to ovate, pedicellate, epicarp yellow, orange, black or purple; pyrene globose or ovate, ivory-white, base sometimes with basal apiculum, apex sometimes with a scar in various shape and a distal apiculum, if basal and distal apiculum present (= biapiculate), if only distal apiculum present (= monoapiculate), ornamentation surface varied at the specific level. All species in this treatment belong to the C. subg. Mertensia. The main characters of this subgenus are the thorns and the divided stigmatic lobes (bifid or bilobed).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731239D6854D5FF297EF9A2E0.taxon	etymology	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731239D6854D5FF297EF9A2E0.taxon	distribution	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 2 A 1, A 2), abaxial surface scabrous; mature drupe lemon-yellow (Figure 6 A), epicarp scabrous and pyrene with alveolate-crateriform surface (Figure 6 B), whereas C. gardneri has leaf blades with trichomes on both the veins and the blade surface (Figure 2 E 1, E 2), abaxial surface velvety or smooth; mature drupe fulvous-orange (Figure 6 I), epicarp smooth and pyrene with alveolate-crateriform-verrucose surface (Figure 6 J). 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 2 A) or emerging from the middle to the distal third, scabrous leaf surfaces; pistillate flowers with conspicuous style (Figure 5 B); lemon-yellow mature drupes, 7.1 – 7.5 × 7.9 – 8.6 mm (Figure 6 A), pyrenes 4.2 – 5 × 3.5 – 4.2 mm with alveolate-crateriform surface (Figure 6 B), whereas C. iguanaea has glabrous to subglabrous thorns; mature leaves with entire margins (Figure 3 A) or serrate on distal third (Figure 9 D), leaf surfaces smooth; pistillate flowers with inconspicuous (Figure 5 N) or no style; fulvous-orange mature drupes, 8.1 – 14 × 7.2 – 13 mm (Figure 6 M), and pyrenes 4 – 8 × 4 – 6 mm with verrucose surface (Figure 6 N). Nomenclatural notes: — Celtis flavovenarum was proposed as a new name for C. morifolia Planch. (Zamengo et al. 2023 b) 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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731239D6854D5FF297EF9A2E0.taxon	materials_examined	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, 15 º 37 ’ 33 ” S 56 º 3 ’ 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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731229D6B54D5FD697BAFA7F6.taxon	etymology	Etymology: — The epithet “ brasiliensis ” refers to the country of Brazil where the type was collected. Vernacular names: — Corrupiá, espora de galo, espora de pinto, esporão de galo, grão de galo, joá branco, joá cipó, joá de espinho, juá and joazeiro (Brazil).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731229D6B54D5FD697BAFA7F6.taxon	distribution	Distribution, habitat and ecology: — Endemic to Brazil, all records from the Atlantic Forest in the states of Espírito Santo and Rio de Janeiro (Figure 7). Except for the types of synonymized taxa, Berg & Dahlberg (2001) do not mention any other specimens analyzed, making it impossible to understand their morphological criteria and correct the identifications of taxa erroneously identified as C. brasiliensis. We noted that the majority of specimens identified as C. brasiliensis by Berg & Dahlberg (2001) belong to three species: C. clausseniana (occurring mainly in the dry areas of Argentina, Bolivia, Brazil, and Paraguay), Celtis serratissima Zamengo, Torres, Gaglioti & Romaniuc (2020: 953), which occurs in the dry areas of Bolivia, Brazil, and Paraguay), and C. pubescens (occurring in the Amazonian areas of Bolivia, Brazil, and Peru). Celtis brasiliensis occurs in Atlantic Rainforest, restingas and anthropic areas and has an affinity for sandy soils. Heliophilous, all records associated with open environments with high light incidence.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731229D6B54D5FD697BAFA7F6.taxon	description	Phenology: — Flowering from July to March and fruiting from June to May of the following year. Taxonomic notes: — Baehni (1936) synonymized C. brasiliensis under C. pubescens. These species are distinct. Celtis brasiliensis is endemic to the Brazilian Atlantic Forest (states of Espírito Santo and Rio de Janeiro states, Figure 7), it has secondary and tertiary branches glabrous to pilose with ivory-white trichomes; thorns glabrous to subglabrous, the trichomes ivory-white; petiole subglabrous to pilose, the trichomes ivory-white or lemon-yellow, leaf blades adaxially surface subglabrous with trichomes concentrated on the veins and scarce on the blade surface (Figure 2 B 1), abaxially surface subglabrous to pubescent (Figure 2 B 2), with pocket domatia (Figure 2 B 3); ovary pilose; mature drupe primrose-yellow (Figure 6 C), 10.3 – 10.6 × 8.5 – 9.3 mm (Figure 6 C), epicarp glabrous (Figure 6 C) or subglabrous with trichomes ivory-white; epicarp surface scabrous to smooth, pyrene 6.5 – 7 × 5.5 – 5.7 mm (Figure 6 D), without scar (Figure 6 D), surface alveolate-crateriform-verrucose (Figure 6 D), whereas C. pubescens it is endemic to the Amazonian regions of Bolivia, Brazil (Acre state), Colombia, Ecuador and Venezuela (Figure 7), has secondary and tertiary branches pubescent to velutinous, the trichomes chestnut-brown or lemon-yellow; thorns pubescent to velutinous, the trichomes chestnut-brown; petiole pubescent to velutinous, the trichomes chestnut-brown or lemon-yellow, leaf blade adaxially surface pilose to pubescent with trichomes both on the veins and along the blade surface (Figure 3 C 1), abaxially surface pubescent to velutinous (Figure 3 C 2), with tuft domatia (Figure 3 C 3); ovary pubescent; mature drupe lemon-yellow (Figure 6 Q, R), 8 – 10 × 4 – 6 mm (Figure 6 Q, R), epicarp pilose to pubescent with lemon-yellow trichomes concentrated both at the base and along the surface of the epicarp (Figure 6 Q, R), scabrous surface, pyrene 5 – 6.5 × 4 – 4.5 mm (Figure 6 T), with scar (Figure 6 T), and alveolate-crateriform pyrene surface (Figure 6 T). After comparing the type of C. brasiliensis and other specimens from Rio de Janeiro, we concluded that C. membranacea should be synonymized under C. brasiliensis. In this sense, C. brasiliensis can be differentiated from C. iguanaea by the following characters: C. brasiliensis has mature leaves with serration from the middle to the distal third (Figure 2 B), adaxial surface pilose to velutinous, scabrous (Figure 2 B 1), abaxial surface with trichomes both on the veins and on blade surface (Figure 2 B 2); glomerulate cymes (Figure 4 B); conspicuous style (Figure 5 D); mature drupe primrose-yellow (Figure 6 C), mesocarp viscous, alveolate, pyrene without scar, surface alveolate-crateriform-verrucose with non-prominent warts (Figure 6 D), whereas C. iguanaea has mature leaves with entire leaf margins (Figure 3 A) or with teeth restricted to the upper third (Figure 9 D, E), adaxial surface glabrous to subglabrous, smooth, abaxial surface with trichomes concentrated on the veins and scarce on the blade surface (Figure 3 A 2); paniculiform cymes (Figure 4 F); inconspicuous (Figure 5 N) or no style; mature drupe fulvous-orange (Figure 6 M), mesocarp non-viscous, not ornamented, pyrene with scar, verrucose pyrene surface, and prominent warts (Figure 6 N).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731229D6B54D5FD697BAFA7F6.taxon	materials_examined	Additional material examined: — BRAZIL. Espírito Santo: Aracruz, PORTOCEL, na restinga, 19 February 1992, fr., O. J. Pereira 2723 (SP, VIES). Cariacica, PNM do Mochuara, 20 º 17 ’ 42 ” S 40 º 25 ’ 49 ” W, 19 May 2018, fr., A. M. Assis et al. 4464 (VIES). Linhares, Pontal do Ipiranga, 10 March 1996, fr., A. M. Assis et al. 103 (SP, VIES), São Rafael, Rodovia Antônio Armani, 19 º 27 ’ 43 ” S, 40 º 12 ’ 32 ” W, 5 April 2022, fr., D. F. Silva et al. 391 (VIES). Marataízes, na restinga, 3 November 1972, fr. P. L. Krieger 11865 (CESJ, SP). Serra, Jacareípe, restinga, 20 º 8 ’ 60 ” S 40 º 11 ’ 00 ” W, 7 December 1996, fl., A. R. Azevedo & F. Passamani 105 (VIES). Vila Velha, Interlagos, restinga, 10 November 1995, fl., O. Zambom & M. Fernandes 161 (SP, VIES), Alagados do vale, Vale Encantado, 25 January 2015, fr., R. T. Valadares 1282 (VIES). Vitória, Praia Comprida, 5 May 1946, fr., A. C. Brade et al. 18101 (RB); Reserva Ecológica Restinga de Camburi, 23 August 1997, fr., A. M. de Assis & I. W. Júnior 316 (VIES). Rio de Janeiro: Araruama, Praia Seca, restinga, 8 March 2008, fr., A. C. S. Cavalcanti et al. 63 (RB). Armação dos Búzios, estrada para Búzios, 22 January 1967, fr., D. Sucre 1447 (RB); praia rasa, restinga, 12 November 1998, fr., A. Q. Lobão et al. 390 (RB); restinga de Manguinhos, 12 November 1999, fl., D. Fernandes & J. Caruzo 289, 297 (RB); mata residual na margem da lagoa de Geribá, 21 October 2000, fr., C. Farney et al. 4241 (RB); Rasa, 22 ° 44 ’ 28 ” S, 41 ° 57 ’ 15 ” W, 3 April 2004, fr., H. G. Dantas & R. D. Ribeiro 247 (RB); Tauá, 7 March 2004, fr., A. F. P. Machado et al. 95 (RB); Rasa 22 ° 45 ’ 18 ” S, 41 ° 59 ’ 04 ” W, 27 August 2004, fl., H. G. Dantas 393 (RB). Cabo Frio, Restinga do Peró, 17 September 1968, fl., D. Sucre 1447 (RB). São Pedro da Aldeia, Avenida Wilson Mendes 478, - 22.8679219 S, - 42.0288895 W, 27 March 2021, fr., H. B. Z. Souza & F. M. Bastos 145 (RB).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D6A54D5FF297F52A728.taxon	materials_examined	Type (designated by Zamengo et al. 2024 b): — PERU. Without a specific location: s. d., fl., H. Ruiz s. n. (lectotype: K [000575978] image!; isolectotypes: G [00354630, 00354632] images!, HAL [0110267] image!, P [00089349] image!). Scrambling shrubs, height unknown, secondary and tertiary branches buff-yellow, fawn-brown, stramineous-yellow or tawny-brown, straight or sinuous, terete, pubescent to velutinous, the trichomes lemon-yellow; thorns 3 – 9 mm long, solitary, curved, buff-yellow, pilose to pubescent throughout, the trichomes lemon-yellow, concentrated both at the base and on the surface of the thorns. Leaf: petiole 2 – 3 mm long, pubescent to velutinous, the trichomes lemon-yellow; leaf blades elliptic, ovate or ovate-lanceolate, 3.9 – 9 × 1.6 – 3.7 cm, concolorous (emerald-green or olive-green), chartaceous, the apex acuminate, the base symmetrical, rounded, the margins entire or laxely serrate from the middle to distal third (mature leaves), adaxial surface scabrous, opaque, pubescent throughout, the trichomes ivory-white or lemon-yellow, abaxial surface velvety, pubescent to velutinous throughout, the trichomes lemon-yellow, veins protruding veins, buff-yellow or chestnut-brown, pocket domatia inconspicuous, pubescent to velutinous, the trichomes lemon-yellow dispersed over the entire surface. Cymes paniculiform, peduncles 20 – 25 mm long, pubescent to velutinous, the trichomes lemon-yellow, bracts absent. Staminate flowers: pedicels 2 – 4 mm long, pubescent to velutinous, the trichomes lemon-yellow, sepals abaxially pubescent to velutinous, the trichomes lemon-yellow, the margins ciliate. Pistillate flowers: pedicels 2 – 3 mm long, pubescent to velutinous, the trichomes lemon-yellow; ovary 4 – 6 × 3 – 4 mm, pubescent to velutinous throughout, the trichomes lemon-yellow, velvety, style conspicuous (0.6 – 1 mm long), inconspicuous (0.1 – 0.5 mm long) or null, the stigmatic branches 7 – 9 mm long, bifid, the lobes 4 – 5 mm long. Drupe: ovate, 10 – 12 × 8 – 9 mm, epicarp lemon-yellow or primrose-yellow (see Ruiz 1777 – 1788: 178 – 179), scabrous, subglabrous or pilose, the trichomes lemon-yellow; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 7.5 × 6.5 mm, ivory-white, surface alveolate-crateriform, monoapiculate, the apiculum truncate, 0.5 mm long, truncate apex apiculum, scar absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D6A54D5FF297F52A728.taxon	etymology	Etymology: — The epithet “ dichotoma ” refers to the apparently dichotomous cymes. Vernacular names: — Unknown.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D6A54D5FF297F52A728.taxon	distribution	Distribution, habit and ecology: — Endemic to Peru (Figure 7), know from only two records in the Departamento of Pasco. In addition to the type, a new specimen was located, collected in secondary forests of Pozuzo, at about 1250 m. Phenology: — Unknown. Taxonomic notes: — Berg & Dahlberg (2001) synonymized C. dichotoma under C. iguanaea. These species must be considered distinct because C. dichotoma has adaxial leaf blade pubescent (Figure 2 C 1), scabrous, abaxial leaf surface pubescent to velutinous on veins and blade surface, trichomes lemon-yellow (Figure 2 C 2); ovary pubescent to velutinous (Figure 5 F), stigmatic branches 7 – 9 mm long, stigmatic lobes 4 – 5 mm long (Figure 5 F); mature drupe lemon-yellow or primrose-yellow (see Ruiz 1777 - 1788: 178 – 179), pyrene with alveolate-crateriform surface (Figure 6 F), whereas C. iguanaea has adaxial leaf surface glabrous to subglabrous (Figure 3 A 1), smooth, leaf abaxially subglabrous mostly on the veins and scarcely on the blade surface, trichomes ivory-white, (Figure 3 A 2); ovary subglabrous to pilose (Figure 5 N), stigmatic branches 1.5 – 2 mm long, stigmatic lobes 0.6 – 1 mm long (Figure 5 N), mature drupe fulvous-orange (Figure 6 M), pyrene with verrucose surface (Figure 6 N). We located a new specimen of C. dichotoma, which is sterile, and thus lacking information on phenology. The label of this specimen indicates that it is a shrub without information on the height, which justifies the absence of this information in the description of the species.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D6A54D5FF297F52A728.taxon	materials_examined	Additional material examined: — PERU. Pasco: Oxapampa, Chacahuassi, Pozuzo, Puesto de vigilancia Huampal, bosque secundário, 10 ° 11 ’ S, 75 ° 34 ’ W, 1250 m alt., 23 September 2002, A. Monteagudo et al. 3977 (HOXA).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D7454D5F8117E60A2DC.taxon	description	Lectotype (designated by Zamengo et al. 2023 b): — BRAZIL. Minas Gerais: Mercês, 1841, fl., G. Gardner 5184 (lectotype: K [000512921, left branch] image!; isolectotype: K [000512920] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D7454D5F8117E60A2DC.taxon	description	Scrambling shrubs or scrambling trees, 3 – 20 m tall; secondary and tertiary branches cinereous-gray or chestnut-brown, sinuous or straight, terete or sulcate, pilose to velutinous, the trichomes ivory-white or chestnut-brow; thorns 2 – 8 mm long, in pairs or solitary, curved, semi-curved or straight, chestnut-brown or maroon-red, pilose to velutinous throughout, the trichomes ivory-white or chestnut-brown. Leaf: petiole 3 – 10 mm long, pubescent to velutinous, the trichomes ivory-white or chestnut-brown, leaf blades widely-ovate or ovate-lanceolate, 3.8 – 12.2 × 2.1 – 7.5 cm, concolorous (olive-green or chestnut-brown) or discolorous (adaxial surface olive-green and abaxial chestnut-brown), chartaceous, the apex acuminated, the base symmetrical, cordate or sub-cordate, the margins entire, serrate or serrulate, congested teeth emerging from the proximal third to the distal (immature leaves), from the middle to the distal third (mature leaves) or restricted to the distal third (mature leaves), adaxial surface scabrous, opaque, pilose to pubescent throughout, the trichomes ivory-white or chestnut-brown, abaxial surface velvety, pubescent to velutinous throughout, the trichomes ivory-white or chestnut-brow, veins sometimes protruding, chestnut-brown, pocket domatia, inconspicuous, pubescent to velutinous throughout, the trichomes ivory-white or chestnut-brown. Cymes glomerulate, peduncles 4 – 10 mm long, pubescent to velutinous, the trichomes ivory-white or chestnut-brown, bracts absent. Staminate flowers: pedicels 1 – 2 mm long, pubescent to velutinous, the trichomes ivory-white or chestnut-brown, sepals abaxially pubescent to velutinous, the trichomes ivory-white or chestnut-brown, the margins ciliate. Pistillate flowers: pedicels 2 – 4 mm long, pubescent to velutinous, the trichomes ivory-white or chestnut-brown; ovary 4 – 6 × 2 – 4.5 mm, velutinous throughout, the trichomes ivory-white or chestnut-brown, velvety, the style conspicuous (0.6 – 1 mm long), the stigmatic branches 3 – 4 mm long, bifid, the lobes 1 – 2 mm long. Drupe: globose or ovate, 7.5 – 12 × 8 – 9 mm, epicarp primrose-yellow, velvery, velutinous, the trichomes ivory-white or chestnut-brown; mesocarp viscous, membranous, not ornamented; pyrene ovate, 8 – 10 × 6 – 6.7 mm, ivory-white, surface striated-verrucose, nonproeminent warts, rounded warts randomly distributed, biapiculate, distal apiculum aciculate, 0.5 – 1 mm long, linear apex apiculum, scar absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D7454D5F8117E60A2DC.taxon	etymology	Etymology: — The epithet “ diffusa ” refers to the growth pattern. Vernacular names: — Coatindiba, corindiba, corindiuba, corrupiá, corubá, cotindiba, curubá, espora de galo, espora de pinto, esporão, esporão de galo, galinha choca, grão de galo, guajissara, gurrupiá, gurupiá, joá branco, joá cipó, joá de espinho, juá, juá preto, juazeiro, limoeiro, nhapindá, salta-martim, tala and taleiro (Brazil).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D7454D5F8117E60A2DC.taxon	distribution	Distribution, habit and ecology: — Most records from the coast of Brazil with only one record from Paraguay (Figure 7). Common in areas of the Atlantic Forest, scarce in the Brazilian Cerrado. In Brazil, the species occurs at the edges of dense ombrophilous forests, “ capoeirões ” and dry forests. Celtis diffusa has an affinity for sandy, sandyclayey and clayey soils. Heliophilous, as all its records are associated with environments with high incidence of light. Phenology: — Flowers and fruits occur from March to December. Taxonomic notes: — Baehni (1936) synonymized C. diffusa under C. pubescens. These species must be considered distinct because C. diffusa occurs in the southeast region of Brazil with only one record for Paraguay (Figure 7); has pocket domatia (Figure 2 D 3); ovary 4 – 6 × 2 – 4.5 mm (Figure 5 H), stigmatic branches 3 – 4 mm long (Figure 5 H); epicarp with chestnut-brown trichomes (see Figure 8 F in Chamorro et al. 2021), biapiculate pyrene 8 – 10 × 6 – 6.7 mm with striated-verrucose surface (Figure 6 H), whereas C. pubescens occurs in the Amazon regions of Bolivia, Brazil (Acre), Colombia, Ecuador, Peru, and Venezuela (Figure 7); has tuft domatia (Figure 3 C 3); ovary 0.5 – 1 × 0.5 – 1 mm (Figure 5 R), stigmatic branches 1 – 2 mm (Figure 5 R); epicarp with lemon-yellow trichomes (Figure 6 S), monoapiculate pyrene 5 – 6.5 × 4 – 4.5 mm with alveolate-crateriform surface (Figure 6 T). Berg & Dahlberg (2001) synonymized C. diffusa under C. iguanaea. These species are distinct, because C. diffusa has branches, thorns (see Figure 2 F in Chamorro et al. 2021), abaxial leaf surface (Figure 2 D 2), and ovary pilose to velutinous (Figure 5 H); glomerulate cymes (Figure 4 D); style conspicuous (Figure 5 H), stigmatic branches 3 – 3.5 mm long (Figure 5 H); drupe primrose-yellow (Figure 6 G), epicarp covered by chestnut-brown trichomes (see Figure 8 F in Chamorro et al. 2021), biapiculate pyrene with striated-verrucose surface (Figure 6 H), whereas C. iguanaea has branches, thorns (Figure 9 B), abaxial leaf surface (Figure 3 A 2) and ovary glabrous to pilose (Figure 5 N); dichotomous cymes (Figure 4 F); style inconspicous (Figure 5 N) to none, stigmatic branches 1.5 – 2 mm long (Figure 5 N); mature drupe fulvous-orange (Figure 6 M), epicarp subglabrous with ivory-white trichomes (Figure 6 M), monoapiculate pyrene with verrucose surface (Figure 6 N).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731209D7454D5F8117E60A2DC.taxon	materials_examined	Additional material examined: — BRAZIL. Bahia: Itagibá, Litoral sul, mata do laterítico, 14 ° 10 ’ 36 ” S, 39 ° 44 ’ W, 13 July 2009, fr., M. L. Guedes et al. 16355 (ALCB, NY, US). Medeiros Neto, comunidade Agrovila Panorama, núcleo Jacarandá, matriz 5, 17 ° 25 ’ 7 ” S, 46 ° 09 ’ 35 ” W, 16 November 2013, fl., Grupo de coletores do núcleo Jacarandá 5 (RB). Espírito Santo: Barra de São Francisco, Boa Sorte, propriedade do Sr. Vitorino Cortelette, 10 July 1984, fr., R. M. Pizziolo 179 (MBML, SP). Bairro Alvorada, Fazenda Santa Catarina, 13 May 1988, fr., M. F. dos Santos s. n. (MBML 5003, SP 293364). Minas Gerais: Caratinga, Estação Biológica de Caratinga, 11 October 1987, fl., I. R. Andrade et al. 241 (NY); 30 May 2003, fr., J. P. Boubli s. n. (ESA 85798, BHCB 82204). Descoberto, reserva biológica da represa do Grama, 3 November 2002, fl., L. C. S. Assis et al. 614 (CESJ, RB). Juiz de Fora, Caetés, 18 May 1982, fr., L. Krieger s. n. (CESJ 19925, SP 304097). Mercês, Estrada Espírito Santo, borda da estrada, 21 ° 10 ’ 53.8 ” S 43 ° 20 ’ 03.4 ” W, 1 August 2022, fr., H. B. Z. Souza 227 (RB). Rio de Janeiro: Alto da Boa Vista, Vista Chinesa, 29 November 1945, fr., J. G. Kuhlmann s. n. (NY barcode 00441119!, P barcode 06781629!, RB barcode 00438837!), Estrada Dona Castorina, 22 º 58 ’ 14 ” S, 43 º 15 ’ 19 ” W, 11 May 2022, fr., H. B. Z. Souza 223 (RB). São Paulo: São José dos Campos, Rua B ao lado da Rodovia Governador Carvalho Pinto, - 23.3001336 S, - 45.8849443 W, 22 November 2020, fr., H. B. Z. Souza 142, 143, 144 (PMSP). PARAGUAY. Caaguazú: 19 November 1876, fr., B. Balansa 2423 (P).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313E9D7754D5FD0D7AF6A2DC.taxon	description	Lectotype (designated by Zamengo et al. 2023 b): — BRAZIL. Sergipe: Porto da Folha, São Pedro Island, São Francisco River, February – April 1838, fl. & fr., G. Gardner 1406 (lectotype: K [000575973] image!; isolectotypes: G [00354615, 00354624] images!, GH [00034260] image!, K [000964240 left branch] image!, NY [00441092] image!, P [00089353, P 00089354] images!, S [8976] image!, US [01117635] image!). Scrambling shrubs, 2 – 6 m tall; secondary and tertiary branches chestnut-brown or cinereous-gray, sinuous, sulcate, pilose, the trichomes ivory-white; thorns 3 – 6 mm long, in pairs or solitary, curved, semi-curved to straight, cinereous-gray or fuscous-brown, pilose to pubescent, the trichomes ivory-white, concentrated at the base and scarce over the rest of the surface. Leaf: petiole 5 – 7 mm long, pilose to pubescent, the trichomes ivory-white, leaf blades elliptic, 5 – 6 × 2.5 – 3 cm, concolorous (emerald-green, cinnamon-brown or olive-green), chartaceous, the apex acuminated, the base symmetrical, rounded or obtuse, the margins serrate, congested teeth emerging from the proximal third to the distal (immature leaves) or from the middle to the distal third (mature leaves), adaxial surface scabrous or smooth, lustrous, subglabrous to pilose throughout or sometimes concentrated on the veins and scarce on the blade surface, the trichomes ivory-white, abaxial surface velvety or smooth, subglabrous to pilose throughout, the trichomes ivory-white, veins non-protruding, chestnut-brown or cinnamon-brown, pockets domatia conspicuous, pilose, the trichomes ivory-white, ciliate. Cymes glomerulate, peduncles 2 – 4 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 pilose, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 2 – 4 mm long, pilose to pubescent, the trichomes ivory-white; ovary 2.5 – 3 × 2 – 2.5 mm, subglabrous to pubescent throughout or concentrated at the base and scarce over the rest of the surface, the trichomes ivory-white, velvety or smooth, style conspicuous (0.6 – 1 mm long), the stigmatic branches 2 – 2.5 mm long, bifid, the lobes 0.6 – 1 mm long. Drupe: globose or ovate, 7.1 – 7.5 × 7.9 – 8.6 mm, epicarp fulvous-orange, smooth, glabrous or subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 4.2 – 5 × 3.5 – 4.2 mm, ivory-white with alveolate-crateriform-verrucose surface and proeminent, rounded, randomly distributed warts, monoapiculate, the apiculum aciculate, 0.75 – 1 mm long, linear apex apiculum, scar absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313E9D7754D5FD0D7AF6A2DC.taxon	etymology	Etymology: — The epithet “ gardneri ” honors George Gardner, the collector of the type. Vernacular names: — Espora de galo, espora de pinto, esporão, esporão de galo, grão de galo, joá branco, joá cipó, joá de espinho, juá and juazeiro (Brazil).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313E9D7754D5FD0D7AF6A2DC.taxon	distribution	Distribution, habitat and ecology: — Endemic to Brazil, most records from the Caatinga (Figure 7). The species occurs in gallery forests, dry forests and steppe savannas. It has an affinity for sandy soils and litholic neosols. Heliophilous, all records from open environments with high incidence of light. Phenology: — Flowering from January to October and fruiting all year. Taxonomic notes: — Baehni (1936) and Berg & Dahlberg (2001) synonymized C. gardneri under C. iguanaea. These species should be recognized independently, because C. gardneri has sulcate secondary and tertiary branches; pilose to pubescent thorns; mature leaves with margins serrated from the middle to the distal third (Figure 2 E), abaxial surface with trichomes throughout, conspicuous domatia (Figure 2 E 3); glomerulate cyme (Figure 4 E); conspicuous style (Figure 5 J); and alveolate-crateriform-verrucose pyrene surface (Figure 6 J), whereas C. iguanaea has terete secondary and tertiary branches; glabrous to subglabrous thorns; mature leaves with entire margins (Figure 3 A) or with teeth restricted to the distal third (Figure 9 D), abaxial surface with trichomes concentrated on the veins and scarce on the blade surface, inconspicous domatia (Figure 3 A 3); paniculiform cyme (Figure 4 F); inconspicous or no style (Figure 5 N), and verrucose pyrene surface (Figure 6 N).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313E9D7754D5FD0D7AF6A2DC.taxon	materials_examined	Additional material examined: — BRAZIL. Alagoas: Maravilha, subida da Serra da Caiçara em direção a torre de telefonia, 23 March 2006, fr., R. P. Lyra-Lemos et al. 9267 (MAC). Palmeira dos Índios, Fazenda Boa Sorte, 9 º 32 ’ 38 ” S, 35 º 49 ’ 58 ” W, 5 April 2008, fl., R. P. Lyra-Lemos et al. 11055 (MAC). Pão de Açúcar, próximo ao Rio São Francisco, 9 º 42 ’ 65 ” S, 37 º 30 ’ 13 ” W, 23 June 2002, fr., R. P. Lyra-Lemos et al. 6911 (MAC). Pilar, Fazenda Lamarão, 20 June 2006, fr., R. P. Lyra-Lemos et al. 9616 (MAC). Santana do Ipanema, Serra do Gugi, 31 January 2010, fl. & fr., Chagas-Mota 7466 (MAC). Viçosa, Serra Dois Irmãos, margem do Rio Paraíba, 28 February 2009, fr., Chagas-Mota 2238 (MAC). Ceará: Aracoiaba, 4 º 30 ’ 00 ” S, 38 º 42 ’ 00 ” W, 19 February 2014, fl., W. Batista 263 (EAC, RB). Barbalha, 24 March 2023, fr., A. C. Sampaio 1 (RB). Fortaleza, beira alta do riacho da 1 º ponte da estrada de Maracanaú, 7 October 1955, fl., A. Ducke 2492 (RB). Sergipe: Porto da Folha, povoado dos índios Xocó, divisa entre Alagoas e Sergipe, nas margens do rio São Francisco, 9 º 47 ’ 18 ” S, 37 º 21 ’ 51 ” W, 24 March 2022, fl., & fr., H. B. Z. Souza & I. M. dos Santos 224 (RB).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313D9D7754D5FD0D7F1FA7B4.taxon	description	Lectotype (designated by Zamengo et al. 2023 b): — BRAZIL. Minas Gerais: Without a specific location, 1816 – 1821, fl. & fr., A. F. C. P. de Saint-Hilaire B 1 1877 (lectotype: P [00089358] image!; isolectotype: P [00089359] image!). Shrubs or trees, 3 – 12 m tall; secondary and tertiary branches fawn-brown, straight, terete, subglabrous to pilose, the trichomes ivory-white; thorns 1 – 2 mm long, solitary, straight, buff-yellow, subglabrous, the trichomes ivory-white, concentrated at the base and scarce over the rest of the surface. Leaf: petiole 5 – 8 mm long, subglabrous to pilose, the trichomes ivory-white; leaf blades elliptic, widely-elliptic, widely-ovate or ovate, 2.3 – 4 × 1.5 – 2 cm, concolorous (emerald-green or olive-green), chartaceous, the apex acute or attenuate, the base symmetrical, obtuse, the margins serrate, congested teeth emerging from the middle to the distal third (mature leaves), adaxial surface smooth, lustrous, subglabrous to pilose mostly on the veins and scarce on the blade surface, the trichomes ivory-white, abaxial surface smooth, subglabrous to pilose mostly on the veins and scarce on the blade surface, the trichomes ivory-white, veins protruding, chestnut-brown, pockets domatia, conspicuous, pilose throughout, the trichomes ivory-white. Cymes glomerulate, peduncles 2 – 4 mm long, subglabrous to pilose, the trichomes ivory-white, bracts absent. Staminate flowers: pedicels 1 – 2 mm long, subglabrous, the trichomes ivory-white, sepals abaxially pilose, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 1 – 3 mm long, subglabrous to pilose, the trichomes ivory-white; ovary 2 – 2.5 × 1 – 1.5 mm, subglabrous to pilose throughout, the trichomes ivory-white, concentrated both at the base and on the surface of the ovary, smooth, the style conspicuous (0.6 – 1 mm long), stigmatic branches 1 – 1.5 mm long, bifid, the lobes 0.1 – 0.2 mm long. Drupe: globose or ovate, 6 – 6.5 × 3.5 – 4.5 mm, epicarp fulvous-orange, smooth, subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamentede; pyrene ovate, 3.5 – 4 × 3 – 3.7 mm, ivory-white, verrucose surface with proeminent, rounded, randomly distributed warts, apiculum absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313D9D7754D5FD0D7F1FA7B4.taxon	etymology	Etymology: — The epithet “ hilariana ” honors Auguste Saint-Hilaire, the collector of the type. Vernacular names: — Espora de galo, espora de pinto, esporão, esporão de galo and juá (Brazil).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313D9D7754D5FD0D7F1FA7B4.taxon	distribution	Distribution, habitat and ecology: — Endemic to Brazil, with only few records for the states of Bahia (in the Caatinga biome) and Minas Gerais (in the Cerrado biome, Figure 7). The species inhabits riverbanks and roadsides. Phenology: — Flowering in November, fruiting from November to January. Taxonomic notes — Baehni (1936) and Berg & Dahlberg (2001) synonymized C. hilariana under C. iguanaea. These species should be recognized independently, because C. hilariana has arboreal habit (see the specimens Flores et al. 419 [ESA] and Pereira et al. 1660 [UB]); mature leaves 4 cm long (Figure 2 F); glomerulate cymes; and pyrene without apiculum (Figure 6 L), whereas C. iguanaea is a scrambling shrub; mature leaves greater than 10 cm long (Figure 3 A); paniculiform cymes; and pyrene apiculate (Figure 6 N). We recommend that new expeditions be carried out in order to locate new specimens of C. hilariana, because the current quantity is insufficient to provide ecological and conservation data.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313D9D7754D5FD0D7F1FA7B4.taxon	materials_examined	Additional material examined: — BRAZIL. Bahia: Muquém de São Francisco, Margem esquerda do rio São Francisco, 13 March 1991, fr., B. A. S. Pereira et al. 1660 (BG, GUA, IBGE, UB). Minas Gerais: Jaíba, Beira da estrada para pegar a balsa, 15 ° 12 ’ 26 ” S, 43 ° 51 ’ 8 ” W, 31 January 2010, fr., T. B. Flores et al. 419 (ESA, RB). Januária, Vazante do rio São Francisco, 4 km de Januária, 29 November 1953, fl. & fr., M. Magalhães 6086 (BHZB, RB).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	materials_examined	Type: — VENEZUELA. Sucre: Cumaná, Bordones, s. d., fl. & fr., A. J. A. G. Bonpland & F. W. H. A. von Humboldt 352 (holotype: P [00669755] image!; isotypes: B [10 0247963] image!, P [00669756] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	materials_examined	Type: — MEXICO. Querétaro: Maconí, Near Moctezuma, January 1840, fr., C. Ehrenberg 1114 (holotype: B [10 0003604] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	materials_examined	Type: — MEXICO. Morelos: Near Yantepec, 5 July 1901, fl. & fr., C. Pringle 8535 (holotype: GH [00267593] image!; isotypes: A [00267593] image!, B [10 0247962] image!, CM [0480] image!, F [143619] image!, G [00357312, 00357313] images!, GH [00034252] image!, GOET [011255] image!, HBG [512849] image!, LL [00370450] image!, MEXU [00010647, 01231732] images!, MIN [1000579] image!, MO [291620] image!, NY [00133648] image!, P [00722004, 00722005] images!, PH [00003989] image!, UC [141980] image!, US [00089707, 00089708, 01013908], VT [031079] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	description	Scrambling shrubs, 1 – 8 m tall; secondary and tertiary branches cinereous-gray, sinuous, terete, subglabrous to pilose, the trichomes ivory-white; thorns 3 – 17 mm long, in pairs or solitary, curved, semi-curved or straight, cinereous-gray or fuscous-brown, glabrous to subglabrous, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns. Leaf: petiole 4 – 5 mm long, subglabrous to pilose, the trichomes ivory-white, leaf blades elliptic or widely-elliptic, 2.5 – 12 × 1.5 – 6 cm, concolorous (emerald-green or olive-green), chartaceous or membranous, the apex acuminated or cuspidate, the base symmetrical, rounded or subcordate, the margins entire or serrate, congested teeth emerging from the middle to the distal third or restricted to the distal third, adaxial surface smooth, lustrous or opaque, glabrous to subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, adaxial surface smooth, subglabrous, the trichomes ivory-white, concentrated on the veins and scarce on the blade surface, veins protruding, chestnut-brown or lemon-yellow, pocket domatia, inconspicuous, subglabrous to pilose, the trichomes ivory-white, ciliate. Cymes paniculiform, peduncles 2 – 4 mm long, pilose, the trichomes ivory-white, bracts absent. Staminate flowers: pedicels 0.5 – 1 mm long, pilose, the trichomes ivory-white, sepals abaxially pilose, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 1.5 – 2 mm long, pilose, the trichomes ivory-white; ovary 2.5 – 4 × 2 – 3.5 mm, pilose throughout, the trichomes ivory-white, scabrous, the style inconspicuous (0.25 – 0.5 mm long) or null, the stigmatic branches 1.5 – 2 mm long, bifid, the lobes 0.6 – 1 mm long. Drupe: globose or ovate, 8.1 – 14 × 7.2 – 13 mm, epicarp fulvous-orange, scabrous or smooth, subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 4 – 8 × 4 – 6 mm, ivory-white, verrucose surface with proeminent, rounded, randomly distributed warts, monoapiculate, the apiculum aciculate, 0.15 – 0.25 mm long, linear apex apiculum, scar present.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	etymology	Etymology: — The epithet “ iguanaea ” refers to the iguanas (Reptilia: Iguanidae) that Commelijn (1697) observed in Curaçao using the trees as perches and feeding on the fruits (Commelijn 1697). Vernacular names: — Beishi di yuana, beshi di juana, rambèshi, rombèshi, palu di djuku, yerba di Juana and yerba di yuana (Curaçao), cock pai (Antigua & Barbuda), beishi di yuana, rombèshi (Aruba), wild cherry (Belize), beishi di yuana, palu di djuku (Bonaire), escobo, guayaba, maíz tostado, pomarroso, uña de gato, vara (Colombia), gallinasa, guanasa, guasiriano, hueso, ramón de costa, ramón de sierra (Cuba), cagalero (El Salvador), biscucuy, capulin sylvestre (Guatemala), palo de achiote (Honduras), bainoro, bejuco, bimora, callado, carboncillo, cumbro, garabato, grangeno, granjero, guichegueda, gumbro, ixi’im che, jarambullo, k’anbe’eb, mora, muk, naranjita, palo blanco, queretano, raspa sombrero, roupe-capa, safrancillo, sits’muuk, uchika, uña de gato, xmuuy-xmuk, yàg-quê-gád (Mexico), azufaifo (Puerto Rico, Acevedo-Rodríguez 2005), iguana hackberry (USA), fruta de gracheraca, and lladoners (Venezuela).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	distribution	Distribution, habitat and ecology: — Growing naturally from the southern United States throughout Central America, on the Caribbean islands, and in northern South America (Colombia, Guianas, and Venezuela) (Figure 7). In Curaçao, C. iguanaea is mostly found in the northwestern part of the island (locally known as Bandabou) in different habitats, mostly in seasonal gullies (locally known as “ rooien ” (Spanish: arroyo) with running water only during the rainy season). The species has also been observed on steeply inclined hill slopes, directly adjacent to the sea (close to where a seasonal gully enters the sea), below limestone cliffs, and on road sides. Most of these locations seem to share a seasonal availability of water, except for steep slopes and road sides, which could potentially explain the smaller size and reduced growth of the specimens at these locations. Historical locations of C. iguanaea also include the developed urban southeast part of the island of Curaçao (locally known as Bandariba). In the rest of the distribution range, this species appears to grow in scrub (Spanish: matorrales) in the semi-arid states of Mexico, in dry seasonal forests on the Pacific slope of Mexico and Central America, in dry seasonal and semi-deciduous forests of the Yucatan peninsula, in coastal thickets of numerous Caribbean islands, and in equivalent environments in northern South America (Figure 7). Phenology: — Flowering from October to December and fruiting from November to April. Taxonomic notes: — For more than 250 years, the name C. iguanaea has been attributed throughout the Neotropics to different populations of Celtis species with noticeable morphological differences between populations. As a result of these interpretations, C. iguanaea is frequently identified in herbarium collections and during floristic surveys throughout the Neotropics and is considered an ecological generalist (e. g., DRYFLOR 2016, Acevedo-Rodrigues 2020). The main reasons why this taxon has been interpreted very broadly include (1) scant descriptive information in the protologue, (2) scarcity of original material from the island of Curaçao, (3) different interpretations of taxonomic characters, and (4) few informative elements of the lectotype (Figure 1). C. iguanaea has been the only species of the genus known to occur in Curaçao or the Lesser Antilles, and therefore there is no doubt regarding its identity and its morphological features. In general, C. iguanaea is characterized as: small trees or shrubs, often scrambling (Figure 9 A), cinereous-gray bark (Figure 9 B), branches with stipulate or solitary thorns, curved, semi-curved to straight (Figure 9 B); leaf blade elliptic to widely elliptic, apex acuminate to cuspidate, base rounded to subcordate (Figures 3 A, 9 D, E), margins entire (in mature leaves, Figure 9 D, E) or serrate (in immature, Figure 9 C, or mature leaves, Figure 9 D), leaf surfaces concolorous (emerald-green, Figure 3 A); stigmatic branches 1.5 – 2 mm long (Figure 5 N), bifid, lobes 0.5 – 1 mm long (Figure 5 N); mature drupe 8.1 – 14 × 7.2 – 13 mm (Figure 6 M), fulvous-orange (Figure 6 M), epicarp subglabrous, trichomes ivory-white, pyrene ivory, ovoid, monoapiculate, and with verrucose surface (Figure 6 N). The analyzed individuals from the type locality of C. iguanaea (Curaçao) have thorns 3 – 17 mm long and straight, curved or semi-curved, mostly solitary or geminate (Figure 9 B); leaves 2.5 – 12 × 1.5 – 6 cm (Figures 3 A; 9 D, E), glabrous or subglabrous, smooth on both surfaces, margin entire (Figure 9 E) or with teeth restricted to the distal third (Figure 9 D); pistillate flowers with subglabrous ovary (Figures 5 N; 9 I), drupes fulvous-orange, up to 14 mm long (Figure 6 M). Most of these characters were cited by Jacquin (1760, 1763, 1788) during his studies, which proves that the specimens analyzed match Jacquin’s description. Most of these characters are different from those detailed in the description of Berg & Dahlberg (2001). For example, C. iguanaea s. str. has no pubescent leaf surface, neither scabrous adaxial surface, margins with teeth emerging from the base (in mature leaves), yellow drupes and epicarp glabrous to velutinous. In C. iguanaea s. str. the fruits turn lemon-yellow (Figure 9 J) during the ripening process, but when they are fully ripe, they are fulvous-orange (Figure 9 K). We analyzed several additional names (Appendix 1), of which the majority has been traditionally incorporated in the synonymy of C. iguanaea (Sargent 1895, Baehni 1936, Berg & Dahlberg 2001). Based on the results of the analysis of the protologue and type materials, we confirmed that nine of them are taxonomic synonyms of C. iguanaea. In addition, we synonymize C. punctata (≡ S. punctatus) under C. iguanaea because the native specimens from Haiti (type locality of S. puncatatus) have the same characters as the individuals from Curaçao (type locality of C. iguanaea). For example, both populations have subglabrous leaf surfaces, smooth (see the protologue Urban & Ekman 1926), paniculiform cymes, fulvous-orange ripe drupes, and verrucose pyrene surface. After comparing the main morphological characters of the Neotropical species of Celtis (Chamorro et al. 2021) with C. iguanaea s. str. and with the types, protologue, and other publications (Kunth 1817, Klotzsch 1847, Planchon 1848, Miquel 1853, Planchon 1873, Baehni 1936, Dottori 1976, Hunziker & Dottori 1976, Sattarian 2006, Sattarian & Van Der Maesen 2006, Zarafshar et al. 2010, Zamengo et al. 2020, Chamorro et al. 2021, Chamorro 2022) we were able to update the circumscription of C. iguanaea and the species reestablished here (Table 2). Nomenclatural notes: — 1. One factor contributing to the incorrect interpretation of C. iguanaea is its lectotype (Figure 1). The figure illustrates a immature specimen with armed branches, oval-lanceolate leaves with serrated margins and four (apparently dried) fruits. All types of species in Celtis subg. Mertensia have these same characters, meaning that the lectotype (Figure 1) is not diagnostic and cannot be used to accurately name the taxon. We have therefore designated an epitype (Figure 8, E. A. T. Houtepen 1 - RB 01443256) for the correct interpretation of the species, following Art. 9.9 of the International Code of Nomenclature (Turland et al. 2018). The epitype (Figure 8) was collected on the island of Curaçao, the type locality of the species, and has grayish branches, straight and curved thorns, elliptic to widely-elliptic leaves, glabrous and subglabrous leaf surfaces, staminate and pistillate flowers, and orange (fulvous) mature drupes with verrucose pyrenes. 2. When Roemer & Schultes (1819) described Ziziphus commutata, C. aculeata was cited as a synonym in its protologue but with the explicit exclusion of Rhamnus iguanaeus (as ‘ iguaneae ’); thus, Ziziphus commutata is a validly published and legitimate name. However, these authors did not mention any specimen in the protologue, but they indicated the type locality: “ In fruticetis montosis Jamaica and Hispaniola ” and refer to the following: “ eadem cum planta Swartzii, nostra commutatis, videtur ”. This is a hint that they worked with material collected by Swartz. In the Caribbean collection of Swartz in the S herbarium, we found a specimen from Jamaica, which had apparently been reviewed by Roemer & Schultes because the handwritten label (Bürdet 1978 a, 1978 b) was checked where the name Z. commutata appears. We designated this specimen (S 08 - 978) as the lectotype because it agrees with the characters indicated in the protologue: branches, thorns, leaves, and flowers. 3. About C. anfractuosa, Liebmann (1851) cited the locality “ México-Jalcomulco ” but did not mention any specimens. Berg & Dahlberg (2001) recognized as holotype a specimen from herbarium C (F. M. Liebmann 5894), collected at the same locality. A duplicate of this material was located in F (614098). Considering that the proposal of Berg & Dahlberg (2001) cannot be considered a lectotypification, we designate the specimen C (10019649) as lectotype (Art. 9.3, ICN, Turland et al. 2018). 4. In the descriptions of the varieties of C. aculeata, Grisebach (1864) quoted collections by A. Prior and Macfayden, but did not mention the herbarium. Berg & Dahlberg (2001) recognize GOET materials as holotypes (A. Prior s. n. or 291 for C. aculeata var. pubescens and Macfayden s. n. for C. aculeata var. serrata). Duplicates of these specimens have been found in K. All specimens are considered to be syntypes, because Grisebach (1864) did not specify the holotype (Art. 9.6, ICN, Turland et al. 2018). The GOET materials should not be considered holotypes because Grisebach (1864) did not specify the type and it is not known whether he analyzed only GOET materials. In this sense, we have designated lectotypes for both varieties (Art. 9.3, ICN, Turland et al. 2018). For C. aculeata var. pubescens, we chose K (000575970) because it has a flower and a fruit, which were not observed in the GOET material. For C. aculeata var. serrata, we choose K (000575971) because it has flowers, which was not observed in the GOET material. 5. For Sarcomphalus punctatus, Urban & Ekman (1926) cited the specimen “ H- 1041 ” without indicating the herbarium. Berg & Dahlberg (2001) recognized the material H. Ekman H- 1041 of B as a holotype, and mentioned an isotype deposited in S. In this case, a lectotypification is necessary because it is not known which specimen was used to describe the species, since Urban worked at B and Ekman at S. With this in mind, we have selected specimen B (10 0247969) as the lectotype. 6. In the case of Mertensia zizyphoides, Kunth (1817) cited the locality as “ Crescit locis humidis prope Mompox, Minchiqueo et Peñones de Roso ad ripam Magdalenae ” but did not mention any material. Baehni (1936) cited “ Bonpland & Humboldt 1520, hb. Paris ”. Four samples were located in P. Because Baehni (1936) did not specify the material, we propose a second step of lectotypification and elect P (00669757) according to ICN Art. 9.17 (Turland et al. 2018).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67313C9D7E54D5FF2978CEA3D9.taxon	materials_examined	Additional material examined: — ANTIGUA. Freetown Rd. at upper waldrons, 28 August 1937, fl., H. E. Box 634 (US). ARUBA. 1909 - 1910, fl., I. Boldingh 6432 (U); fr., I. Boldingh 7055 (NY, US). BARBUDA. Near Tarafoot Bay, 15 May 1937, H. E. Box 634 (US). BELIZE. Toledo, Chavarria’s Road, Resemederes, across Columbia River, 4 October 1947, P. H. Gentle 6268 (US). BONAIRE. Along the road north of BOPEC, 27 February 1999, fr., A. S. J. van Proosdij et al. 858 (U). Columbia, 4 November 1952, fl., A. L. Stoffers 527 (U). Playa Foentchi, 14 December 1952, fl., A. L. Stoffers 997 (U). Put Bronswinkel, 14 September 1952, fl., A. L. Stoffers 676 (U). COLOMBIA. Atlántico: espinares de Sabanilla em lomas bajas arenosas cerca de la orilla del mar, 26 July 1959, fr., A. Dugand 5165 (US). Magdalena: Comisaria Goajira, 16 August 1944, fl., O. Haugth 4306 (P, US). Meta, Granada, 1 January 1844, fr., J. Goudot s. n. (P barcode 06885259). COSTA RICA. Puntarenas: Osa, Sierpe, Reserva Forestal Golfo Dulce, Rincón, cerca de Banegas Los Charcos, 1 km al Este del centro del Pueblo Banegas, Estación Biológica Los Charcos de Osa, 8 º 40 ’ 18 ” N 83 º 30 ’ 17 ” W, 23 June 2009, fr., R. Aguilar 12151 (U). San José: El General, April 1939, fl., A. F. Skutch 4292 (US). CUBA. Chickens not far from Playa de Marianao, November 1984, fl. & fr., Dr. León 4685 (NY). Las Villas: Farallones de Guajimico, on coast East of Cienfuegos, 24 February 1956, fl. & fr., C. V. Morton 10467, 10472 (US). Havana: 11 October 1904, fr., H. A. Van Hermann 195 (NY). Matanzas bay, 1849, fl., F. Rugel 623 (FLAS). Pinar Del Rio: San Diego de los Baños, 3 September 1910, N. L. Britton et al. 6707 (US). CURAÇAO. Hojje Abrau: 27 February 1917, H. M. Curran & M. Haman 185 (US). In type C, 17 February 1917, H. M. Curran & M. Haman 55 (US). Near Piedra Mulina, between Mt. Christoffel and Mt. Garcia, 31 March 1997, A. S. J. van Proosdij 398 (U). Rooi Beru: 5 October 1952, fl., A. L. Stoffers 208 (U), fl., A. L. Stoffers 210 (U). Sta. Barbara: 18 October 1952, fl., A. L. Stoffers 400 (U). Without locality: s. d., fr., I. Boldingh 4970 (U), s. d., fl., I. Boldingh 5113 (U), March 1913, N. L. Britton & J. A. Shafer 3104 (U, US). Westpoint: Bandabou, Bandabou, 12 ° 21 ’ 36 ” N, 69 ° 8 ’ 34 ” W, 27 March 2021, fl. & fr., E. A. T. Houtepen 2 (RB). DOMINIC. North of Goodwill, Roseau, 6 July 1965, fl. & fr., W. R. Ernst 1840 (US); North side of the east Cabrit, 4 June 1977, fl., D. H. Nicolson 4200 (US). Petit Coulibri: GR 788 828, 8 August 1989, fl., C. Pendry 80805 (E). Point Michel: 21 July 1983, Akeassi & Porticop 16515 (P). DOMINIC REPUBLIC. Altagracia: 1 - 1,5 km norte del poblado de Boca de Yuma en el camino viejo a El Caracol, paralelo ao río Duey (= Río Yuma), 18 º 24 ’ N, 68 º 37 ’ W, 22 November 1983, fr., T. Zanoni et al. 27989 (U). Reserva del Este, 25 July 2001, fr., P. Acevedo-Rodríguez 11785 (US). Azua: March 1913, fl., J. N. Rose et al. 4096 (US). Barahoana: Pedernales, 7 km al sur de los Tres Charcos (de Oviedo) y aprox. 7 - 8 km más en el camino a La Playa Blanca, 17 º 45 ’ N, 71 º 25 ’ O, 9 February 1986, fr., T. Zanoni et al. 36115 (U), Sierra de Bahoruco, Enriquillo N del poblado Higüero, en el lugar llamado El Maniel Viejo, 18 º 3 ’ 5 ” N 71 º 19 ’ 5 ” W, 9 June 1993, fr., R. García et al. 4900 (FLAS), May 1901, fl., M. Fuertes 241 (L); Bay road at base of Morne Daniel, s. d., W. H. Hodge 3889 (US). Cordillera Central: Azua, aprox. 2 km al E del pueblo de Padre Las Casas, 2 October 1987, fr., R. García & J. Pimentel 2527 (US). La Canela, 30 January 1947, fr., J. J. S. Jiménez 1315 (US). Loma Bandera: from Cutupu to La Vega, Limestone hill, 22 May 1968, fl., B. A. H. Liogier 11325 (US). San Juan, Hillslopes, vicinity of Rio Arriba del Norte, north of San Juan, September 1946, R. A. & E. S. Howard 8950 (US). San Pedro de Macoris: March 1913, fl., J. N. Rose et al. 4276 (US). Santo Domingo, 1 May 1911, fl., P. Fuertes 241 (E, P, U, US). EL SALVADOR. San Ignacio: Chalatenango, 14 October 1958, fr., P. H. Allen 7044 (US), La Union, February 1922, fr., P. C. Standley 20647 (US). GUADALUPE ISLAND. Morne la Vierge: Northeast of Capesterre, 2 June 1960, fl., R. Proctor 21149 (US), Grande Terre. 200 m de la pointe des Chateaux. Guadalupe, 19 April 1974, fl., C. Sastre & F. Sastre 2579 (P). GUATEMALA. Dpto. Peten: Tikal, on El Remate road about 8 Km, in high forest, 26 September 1959, fr., E. Contreras 182 (XAL), bordering lake Petén Itzá, in zapotal, about 2 km from San José, 23 September 1976, fr., C. L. Lundell & E. Contreras 20393 (IBUG). HAITI. Without locality, s. d., fl., Poiteau s. n. (P barcode 06884775). Artibonite: 5, 6 km al oeste de Gonaives en la carretera costera a Anse Rouge, 19 º 28 ’ N 72 º 43 ’ W, 11 June 1985, fl., T. Zanoni et al. 35164 (U), Hard limestone foothills at Case-à-Roche, 4 April 1926, fl., E. L. Ekman 5850 (US). Île de Tortue: West side of La Vallée Valley, 29 December 1929, fr., E. C. Leonard & G. M. Leonard 11276 (US). Massif des Cahos: Dessalines, at Case-à-Roche, 13 March 1925, fl., E. L. Ekman 3498 (US). Massif du Nord: St- Michel, road to La Lome, 3 June 1927, fl., E. L. Ekman 8344 (US), Ile La Ganave, 3 August 1927, E. L. Ekman 8805 (U). Nord-Óuest: Île de la Tortue; Ravine on north side of Tortue, northeast of Basse Terre, 26 March 1929, fr., E. C. Leonard & G. M. Leonard 12462 (US). Vicinity of Bassin Bleu: Road to Gros Morne, 15 April 1929, fr., E. C. Leonard & G. M. Leonard 14685 (US). Vicinity of Gros Morne: river flat west of Trois Rivières, 18 February 1926, fr., E. C. Leonard 9900 (US). Vicinity of Jean Rabel: Road toward Port de Pai, 13 March 1929, E. C. Leonard & G. M. Leonard 13830 (US). Vicinity of St. Michel de l’ Atalaye: 17 November 1925, E. C. Leonard 7100 (US); 20 November 1925, E. C. Leonard 7256 (US). Vicinity of St. Raphael: Departament du Nord, 3 December 1925, fr., E. C. Leonard 7715 (US). HONDURAS. Dpto. Lamaní: Comayagua, 20 September 1975, fr., E. N. Vargas 208 (TEFH). Dpto. Copán: La Vegona, Thickets La Vegona, Copán river 2 km east of Copán ruins, 23 November 1969, fr., A. R. Molina & A. R. Molina 24784 (US). Dpto. Morazán: Villa Nueva, 13 June 1947, fl., R. A. Molina 91 (US). Dpto. Olancho: Orilla del río Catacamas, 1 May 1988, fl., C. Nelson & R. Andino 10210 (TEFH). JAMAICA. Srove Place near Mile Sully: 23 September 1908, N. L. Britton 10619 (P). Manchester: 22 October 1975, fl., G. R. Proctor 35378 (US). Grove Place near Mile Gully: 23 September 1908, W. H. Harris & L. Britton 10619 (US). Without locality: s. d. fl., P. O. Swartz s. n. (S 08 - 978). MARTINIQUE. Sarmenteux: Carbet, Prêcheur, 1881, fl., P. A. Duss 1401 (US). Without locality: 1 January 1870, Hahn 1301 (P); 1 January 1857, fl., M. Belanger 279 (P). MEXICO. Campeche: Calakmul, 1 km al N de Rancho El Sacrificio, camino a Nuevo Centro de Población Ejidal Ley de Fomento Agropecuario, 17 º 59 ’ 23 ” N, 89 º 23 ’ 40 ” W, 5 August 1997, fr., E. S. Martínez et al. 28118 (MBM, MEXU); a 4 km al N de Xpujil, camino a Dzibalchén, 18 º 32 ’ 21 ” N, 89 º 24 ’ 7 ” W, 11 August 1997, fr., E. S. Martínez et al. 28363 (MBM, MEXU), 2, 4 km al NE de Bel-Há, 18 º 56 ’ 42 ” N, 89 º 15 ’ 29 ” W, 12 May 2002, fl., J. Calónico Soto & D. Álvarez 23598 (MBM, MEXU), a 4 Km al E de Nuevo Becal, 18 ° 35 ’ 59 ” N, 89 ° 15 ’ 56 ” W, 13 May 2002, fl., D. Alvárez & J. Calónico Soto 1174 (MBM, MEXU). Hopelchén: Laguna Noh, 18 ° 37 ’ N, 90 ° 18 ’ W, 20 October 1981, fr., C. Chan & E. Ucán 966 (XAL). Chiapas: Ocosingo, 5 km al S de Crucero Corozal, 21 April 1985, fr., E. S. Martinez 12223, 12280 (MBM, MEXU), restaurante la escondida, en la 11 de Julio, camino a Palenque, 17 ° 10 ’ 38 ” N, 91 ° 29 ’ 17 ” W, 12 June 2002, fr., G. M. Aguilar & J. Aguilar 1377 (MBM, MEXU, XAL), 1 km al SO del crucero Bonampak, 16 º 45 ’ 34 ” N, 42 º 91 ’ 6 ” W, 22 May 2003, fl., G. M. Aguilar et al. 6822 (MBM, MEXU), 5 km al NE del crucero San Javier, rumbo a San Jacinto, 16 º 48 ’ 50 ” N, 91 º 3 ’ 39 ” W, 16 June 2003, fr., D. Álvarez 5318 (MBM, MEXU), 3, 43 km al E de San Javier, 16 º 45 ’ 39 ” N, 91 º 4 ’ 33 ” W, 27 July 2003, fr., G. M. Aguilar et al. 7535 (MBM, MEXU). Chihuahua: Batopilas, 6 October 1898, fr., E. A. Goldman 219 (US). Colima: Manzanillo, 5 a 10 Km al N de Santiago, brecha Santiago a Chandiablo, 23 June 1984, fl. & fr., F. J. M. Santana & N. A. Cervantes 323 (IBUG). Durango: Pueblo Nuevo, Agua Caliente, 23 ° 17 ” N, 105 ° 20 ” W, 12 October 1983, fr., M. González 1315 (IBUG). Toyaltita: 92 km al SW de San Míguel de Cruces, 24 º 9 ’ N, 105 º 57 ’ W, 7 July 1984, fl., P. L. Tenorio et al. 6290 (MBM, MEXU). Guanajuato: El Charco, 12 Km al sureste de Atarjea, 10 November 1988, fr., E. Ventura & E. López 6373 (XAL), Ocampo, 12 Km al N de Flores Magón, s. d., fl., A. Baliente et al. 41 (MBM, MEXU). Guerrero: La Unión, en El Cedral, a 19 km de Las Lagunas, 12 km al SO de la desviación a La Unión, 21 July 1985, fr., J. C. Soto-Núñez 9583 (MBM, MEXU), Zumpango del Rio, 7 Km al N de Zumpango del Rio, 21 May 1986, fl., J. C. Soto Nuñes 12668 (MBM, MEXU). Hidalgo: Pacula, cañon del Río Moctezuma, camino de Pacula-El Fraile-Mina La Negra-Río Moctezuma, cerca del campamento de la CFE, 22 February 2007, A. Castro-Castro 646 (IBUG). Jalisco: Selva de Cuitzmala, 19 ° 31 ’ 0 ” N, 105 ° 2 ’ 28 ” W, 29 September 1997, fr., A. M. Dominguez 695 (IBUG). Mexico: Temascaltepec, Ixtapan, 13 May 1933, fl., G. B. Hinton 3924 (GBH). Michoacan: Aguililla, 2 km al No de Aguililla, carretera Dos Aguas, 11 July 1985, fl., J. C. Soto-Núñez 9212 (MBM, MEXU), Tiquicheo, Tzentzénnguaro, 26 Km al N de Tiquicheo, 10 October 1981, fr., J. C. Soto Nuñez & G. R. Silva 3282 (XAL). Morelos: Yautepec, July 1905, fl., J. N. Rose et al. 8582 (US), Nayarit, San Blas, 2 Ocotober 1925, fr., R. S. Ferris 5315 (US). Oaxaca: Chahuites, 1 Km al N por el Alto, 28 December 1987, fr., M. A. García-Bielma 147 (XAL), Tehuantepec, San Pedro Huamelula, a 5 km al N de Ayuta, 15 º 56 ’ 23 ” N, 95 º 50 ’ 55 ” W, 15 May 1999, fl., E. S. Martínez et al. 32156 (MBM, MEXU), Guadalupe Victoria, 11 km N de la desviación em la carretera costera, 15 º 59 ’ 10 ” N, 95 º 51 ’ 46 ” W, 30 September 1998, fr., C. Perret et al. 44 (MBM), Yautepec, Santa Maria Ecatepec, río Otate, al S de Santa Maria Ecatepec, 16 º 14 ’ N 95 º 55 ’ W, 9 May 1993, fr., P. L. Tenorio 18761 (MBM, MEXU). Querétaro: Jalpan, 1 Km de El Puerto de Animas, camino a Ahuacatlán, 22 May 1989, fl., E. G. Carranza 1725 (IBUG, XAL), Las adjuntas confluencia entre los ríos Moctezuma y Estórax, 14 May 1993, fl., S. Zamudio et al. 9078 - b (XAL), Landa de Matamorros, Matzacintla, 29 km al noroeste Jalpan, 28 April 1982, fl., R. M. Hernández & P. L. Tenório 7179 (HUA). Quintana Roo: Adolfo de la Huerta, a 3, 3 km al NE de la Carolina, 19 º 37 ’ 17 ” N, 89 º 1 ’ 17 ” W, 19 June 2004, fr., D. Álvarez et al. 9491 (MBM, MEXU). Isla de Cozumel, sobre el camino al Cedral a aproximadamente 18 km al S de San Miguel de Cozumel, 18 June 1987, fr., E. Cabrera & H. de Cabrera 13618 (MBM, MEXU), Solidaridad, Carretera Tulum-Cobá, Km 10 - 15, 20 ° 17 ’ 20 ” N, 87 ° 30 ’ 15 ” W, 8 November 2002, fr., F. May et al. 2280 (XAL). San Luis Potosí: Valles, 19 May 1981, fr., P. A. Fryxell & W. R. Anderson 3503 (US). Sinaloa: Mocorito, 21 September 1923, fr., G. N. Collins & J. H. Kempton 55 (US). Sonora: Alamo, In the vicinity of Alamos. Arroyo near Tocan, 17 March 1910, J. N. Rose et al. 13009 (US), Tabasco, Parque Nacional de Agua Blanca, Macuspana, 17 ° 38 ” N, 92 ° 30 ’ W, 28 November 1987, fr., L. Ruiz Pavón et al. 20616 (GUADA). Tamaulipas: Without locality, Nacimiento del río Sabinas, 4 km al Norte del Ejido la Libertad, 18 May 1982, fr., A. B. Valiente et al. 140 (MBM, MEXU); Victoria, June 1907, fr., E. Palmer 584 (US). Veracruz: Coatepec, 3 km antes de Jalcomulco sobre la carretera Tuzamapan-Jalcomulco, 19 º 21 ’ N 96 º 46 ’ W, 28 July 1979, fr., G. C. Castillo & L. Tapia 797 (MBM, XAL); Naolinco, mal país a orillas de la carretera actopan km 9, 25 June 1976, fr., R. Ortega 294 (MBM, MEXU), Teocelo, 19 ° 22 ’ 28 ” N, 96 ° 51 ’ 29 ” W, 8 September 2016, fr., B. V. Bonilla 1 (XAL). Yucatan: Valladolid, San Miguel hacia el oriente, 20 ° 42 ’ N, 88 ° 14 ” W, 18 September 1981, fr., E. Ucán 1494 (XAL). MONTSERRAT. Isles Bay: 3 April 1979, P. G. Howard 18946 (US). NICARAGUA. Esteli: ca. 7 km from Hwy 1 (at ca. km 193) on road to Pueblo Nuevo, from Quebrada Jamaili to near summit of Cerro El Pedrero, 13 º 24 ’ N, 86 º 27 ’ W, 3 July 1977, fr., W. D. Stevens & B. A. Krukoff 2619 (MBM, MO), ca. 1, 1 km from Hwy 1 (at ca. km 193) on road to Pueblo Nuevo, 13 º 25 ’ N, 86 º 25 ’ W, 29 December 1977, fl., W. D. Stevens & B. A. Krukoff 5739 (MBM, MO). Granada: s. d. fl., P. Levy 1109 (P). Leon: slope and ridge immediately W Quebrada of Las Ruedas, N of road, NW of El Transito, 12 º 5 ’ N 86 º 43 ’ W, 13 May 1981, fl., W. D. Stevens et al. 20140 (MBM, MO). Managua: ca. 4, 5 km NNW of Hwy 12 along road on ridge of Sierra de Mateare, 12 º 7 ’ N, 86 º 23 ’ W, 8 July 1978, fr., W. D. Stevens & B. A. Krukoff 9246 (MBM, MO), 2, 3 km from Hwy 12 on road along ridge of Sierra de Managua from Hwy 12 (Carretera vieja a Léon) at km 17 to Hwy 2 (Carretera Sur), 12 º 4 ’ N, 86 º 22 ’ W, 29 July 1977, fr., W. D. Stevens & B. A. Krukoff 2928 (MBM, MO), ca. 1, 9 km W of Montelimar, 11 º 49 ’ N, 86 º 32 ’ W, 31 August 1977, fr., W. D. Stevens & B. A. Krukoff 3620 (MBM, MO), along highway 8 ca. 2, 4 km SW of intersection with highway 2, Km 28, 11 ° 57 ’ N, 86 ° 20 ’ W, 18 September 1977, fr., W. D. Stevens et al. 3967 (MBM, MO). PANAMA. Bocas del Toro: 8 ° 45 ’ N, 82 ° 15 ” W, 10 August 1986, fr., G. McPherson 9930 (MO, US). Província de Panamá: Subida para Cerro Jefe, 13 March 1969, fl., D. Sucre et al. 4750 (RB). PERU. Tumbes: Contralmirante Villar, 2 May 1990, fr., C. Díaz & E. Peña 4035 (MO, MOL). PUERTO RICO. Along Quebrada Marunguey: 5 km NE of Camp García, 4 June 1978, F. R. Fosberg 57591 (US). Arecibo: Bo. Dominguito, 12 January 1995, P. Acevedo-Rodriguez & A. Siaca 7237 (US). Coamo Springs: Riverbanks, March 1913, E. G. Britton & D. W. Marble 2319 (US), in saxosis, 16 December 1885, fr., P. E. E. Sintensis 3085 (US). Culebra Island: March 1906, N. L. Britton & W. M. Wheeler 105, 107 (US). Fajardo, Along trail from Cabeza Chiquita to playa Convento, 18 º 22 ’ 77 ” N 65 º 38 ’ 59 ” W, 19 December 1994, P. Acevedo-Rodriguez & A. Siaca 6980 (US). San Ildefonzo, Coamo Springs Rd, 22 November 1899, G. P. Goll 708 (US). Juncos: in monte Florida, 25 August 1985, fr., P. Sintensis 2234 (P, US). Tarjavo: in monte Emajaqua, 4 May 1885, fl., P. E. E. Sintensis 1318 (US). Vieques Island: U. S. Fish & Wildlife Refuge in western Vieques, along road to summit of Monte Pirata, 18 º 5 ’ 37 ” N, 65 º 33 ’ 10 ” W, 22 October 2002, fl., G. L. Breckon et al. 6657 (US), Cerra Encantada, 28 January 1914, J. A. Shafer 2561 (US). SABA. Morypoint: 1906, fl., I. Boldingh 1476, 2116 (U). Windwardside: In Village and outskirts of village on road leading to the Bottom, 16 August 2006, fr., S. A. Mori et al. 26413 (NY). Without locality: 24 August 1947, fl., F. Arnoldo 828 (U), 18 July 1906, fl., I. Boldingh 1950 (U). SAINT BATH. Without locality: 29 May 1938, fr., A. Questal 706 (US). SAINT CHRISTOPHER. Without locality: Lower NE slope of Sir Timothy Hill., 17 16 ’ 56 ” N 62 41 ’ 03 ” W, 19 July 2014, fl., S. Carrington 2348 (US). SAINT EUSTATIUS. Quill National Park, Coucher Mountain Trail and small portion of the around the mountain, 17 º 28 ’ 59 ” N 62 º 57 ’ 55 ” W, 2 February 2008, fr., B. M. Boom et al. 11212 (NY). Signaehuel: 1906, fl., I. Boldingh 1029 (U). SAINT MARTIN. Guana Bay: 6 August 1906, fl., I. Boldingh 2389 (U), fl., I. Boldingh 2399 (U). Paradise: 12 September 1906, fl., I. Boldingh 3293 (U). Without locality: June 1945, P. Bena 2174 (P), 1906, fl., I. Boldingh 2572 (NY, U), 21 August 1906, fl., I. Boldingh 2680 (U), 30 August 1906, fl., I. Boldingh 3079 (U). US TRINIDAD AND TOBAGO. Erin seashore: 6 December 1933, W. E. Broadway 9334 (P, U). UNITED STATES OF AMERICA. Florida: Collier County, 3 September 1970, G. N. Avery 838 (FLAS), Horr’s Island near Cape Romano, 10 May 1922, fl., J. K. Small 10479 (NY), Lee County, Sanibel Island, 14 April 1954, fl., Cooley et al. s. n. (FLAS barcode 66617), 15 December 1971, W. C. Brumbach 7772 (FLAS), Upper Captiva, 14 October 1972, W. C. Brumbach 8060 (FLAS, NY), Upper Captiva, 30 March 1978, W. C. Brumbach 9348 (NY), Manatee County, 23 November 1891, J. H. Simpson 405 (NY). VENEZUELA. Anzoategui: 29 October 1949, fr., F. D. Smith 105 (US). Aragua: alrededores de Maracay prolongación de Calle Principal de San Vicente (Tapatapa), aprox. 4 Km hacia el lago, al lado de la carretera, 4 October 1969, fr., G. S. Bunting 3417 (MY). Carababe: environs de Puerto Cabello, 1 January 1843, fl., J. Linden 1532 (P). Margarita Island: 17 July 1901, fl., O. O. Miller 18 (P), 14 August 1903, fl., J. R. Jhonston 116 (US). Falcon: Dabajuro, Buchivacca, 13 July 1976, fl., A. Colma et al. 146 (MY). Guárico: a unos 15 km de Altagracia de Orituco, June 1966, fl., L. Aristeguieta 6131 (US). Chaguaramas: Infante, 19 October 1980, fr., B. Trujillo et al. 17201 (MY). Lara: Barquisimeto, November 1923, fl., J. Saer 81 (US). Miranda: Suroeste del Valle de Caracas, Colinas de Bello Monte, 27 March 1992, fr., N. Ramirez & M. López 3175 (MY). Zulia: Perijá, 1917, E. Tejera 24 (US). US VIRGIN ISLANDS. St. Croix Island. Christiansted: Cane Bay Estat, 22 February 1913, J. N. Rose et al. 3603 (US). St. John Island. Cinnamon Bay: ruins, 24 August 1992, fr., P. Acevedo-Rodriguez et al. 5150 (US). Cruz Bay: Route 104, 100 m east of great Cruz Bay, 18 January 1988, fr., P. Acevedo-Rodriguez 2361 (US). Green Cay: 17 January 1980, F. R. Fosberg 59290 (US). Lameshur Bay: 19 January 1989, fl., P. Acevedo-Rodriguez 2691 (US). Reff Bay Quarter: Along road from little Lameshur to Great Lameshur, 25 August 1987, fr., P. Acevedo-Rodriguez 2021 (CTES, US), Lang’s Peak (Jakobsberg) Eliza’s Retreat, above Altena, 13 January 1979, fr., F. R. Fosberg 58924 (US). Maho Bay Quarter: 100 m from Center Line Road (Road 10), km 4,5, 7 January 1991, fr., P. Acevedo-Rodriguez & A. Siaca 3815 (US), Potwood pond to West End, February 1913, J. A. Shafer 1171 (US). St. Thomas Island. Without locality: 1 August 1881, fl., B. Eggers & A. Toepffer 323 (P). BRITISH VIRGIN ISLANDS. Tortola Island. Without locality, 6 May 1919, fl. & fr., W. C. Fishlock 409 (US), June 1881, fl., B. Eggers s. n. (US barcode 00704115), 17 January 1990, fr., P. Acevedo-Rodriguez 3237 (US). WEST INDIES. Without locality: s. d., fl., W. Bredemeyer s. n. (B - W 18996 - 01 0). s. d., Jacquin s. n. (S-R 11088), s. c., s. n. (S 08 - 980), fr., P. O. Swartz s. n. (S 08 - 975).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	description	Lectotype (designated by Zamengo et al. 2023 b): — 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!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	description	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.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	etymology	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	distribution	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.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	description	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 6 M) and verrucose pyrene surface (Figure 6 N), whereas C. orthacanthos has primrose-yellow mature drupes (Figure 6 O) and alveolate-crateriform pyrene surface (Figure 6 P). 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. 2024 a). 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 6 M), whereas C. orthacanthos (= C. glycycarpa, Figure 6 O) and C. triflora (Figure 6 A 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 5 P), 3 – 4 mm long stigmatic lobes (Figure 5 P); pyrenes with scar, 1 – 1.5 mm long distal apiculum, and alveolate-crateriform surface (Figure 6 P), whereas C. triflora has the 4.5 – 8 mm long stigmatic branches (Figure 5 Z), 4 – 6 mm long stigmatic lobes (Figure 5 Z); pyrene without scar, 1 – 2.5 mm long distal apiculum, and cristed-verrucose surface (Figure 6 A 2).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731349D7954D5FC017AF8A6B8.taxon	materials_examined	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. 5 km W da sede do município, fazenda Cruzeiro do Sul, Serra dos Sete-Paus, ca. 8 km 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, 9,8 km 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 - 35 km de Feira de Santana, 23 March 1994, fl. & fr., F. França & E. Melo 955 (HUEFS). Ilhéus, CEPEC, Km 22 da BR 415, 14 º 47 ’ 5 ” S, 39 º 13 ’ 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 BR 381 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. (GFJP 489, RB 736645, SP 276479); 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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	materials_examined	Type: — ECUADOR. Guayas: Guayaquil, s. d., fl., H. Ruiz López s. n. (holotype: B [10 0247965] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	materials_examined	Type: — COLOMBIA. Tolima: “ between Ibagué and Rusaguasuga ”, s. d., J. Goudot s. n. (holotype: K [000575977] image!; isotype: P [06781653] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	description	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.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	etymology	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	distribution	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 3 A 2), whereas C. pubescens has a pubescent to velutinous (Figure 3 C 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 3 C 3); glomerulate cymes (Figure 4 H); lemon-yellow mature drupes (Figure 6 Q), pilose to pubescent epicarp (Figure 6 Q, R) covered with lemon-yellow trichomes (Figure 6 S), and alveolate-crateriform pyrene surface (Figure 6 T), whereas C. iguanaea has smooth adaxial leaf surface, pocket domatia (Figure 3 A 3); paniculiform cyme (Figure 4 F); fulvous-orange mature drupes (Figure 6 M), glabrous to subglabrous epicarp with ivory-white trichomes (Figure 6 M), and verrucose pyrene surface (Figure 6 N).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731339D7B54D5F8A17AF2A5D0.taxon	materials_examined	Additional material examined: — BOLIVIA. Pando: Manuripi, 35 Km al norte de Puerto América entrando bosque alto cerca Alianza, 11 º 44 ’ S, 67 º 59 ’ 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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	description	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!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	materials_examined	Type: — ARGENTINA. Corrientes: Without a specific location, s. d., fr., A. J. A. G. Bonpland 943 (holotype: P [00089351] image!).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	materials_examined	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.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	etymology	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	distribution	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731319D7A54D5FA197BE6A75C.taxon	materials_examined	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, nº 860, 29 º 49 ’ 55 ” S 50 º 07 ’ 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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731309D4454D5F88D7BC0A150.taxon	materials_examined	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.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731309D4454D5F88D7BC0A150.taxon	etymology	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731309D4454D5F88D7BC0A150.taxon	distribution	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 3 A; 9 D, E), margins entire (Figures 3 A; 9 E) or with teeth restricted to the distal third (mature leaves, Figure 9 D), abaxial surface glabrous; mature drupes fulvous-orange (Figure 6 M) with glabrous epicarp, whereas C. spinosissima has mature leaves oval-lanceolate (Figure 3 E), margins crenateserrated from the proximal third to the apex (Figure 3 E), abaxial surface pilose; mature drupes primrose-yellow (Figure 6 W) 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 4 F; 9 G); ovary subglabrous to pilose (Figures 5 N; 9 I), style inconspicuous (Figures 5 N; 9 I) or null; and mesocarp non-ornamented, whereas C. spinosissima has branches chestnut-brown, fawn-brown or maroon-red; glomerulate cymes (Figure 4 J); ovary pilose to pubescent (Figure 5 V), style conspicuous (Figure 5 V); and mesocarp alveolate (see Figure 8 B in Chamorro et al. 2021).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD6731309D4454D5F88D7BC0A150.taxon	materials_examined	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).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	description	10 A – D).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	materials_examined	Type: — BOLIVIA. Tarija: “ In sylvis ”, August 1846, fr., H. A. Weddell 4816 (holotype: P [00089387] image!, Figure 10 A). Epitype (here designated): BOLIVIA. Tarija: Provincia Arce, 5 km S of comunidad Guayvillas (28.3 km S of Padcaya) on road to Bermejo, 22 º 01 ” S, 64 º 39 ” W, 5 May 1983, fr., J. C. Solomon 10496 (SI [094759] image! Figure 10 B; isoepitypes: LPB [no registration number, image!, Figure 10 C], MO [1429929] image!, Figure 10 D). Scrambling shrubs or scrambling trees, 5 – 15 m tall; secondary and tertiary branches cinereous-gray, sinuous or straight, terete, glabrous to pilose, the trichomes ivory-white; thorns 1 – 10 mm long, in pairs or solitary, semi-curved to straight, buff-yellow or stramineous-yellow, glabrous to subglabrous, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns or scarce throughout. Leaf: petiole 10 – 20 mm long, glabrous to pilose, the trichomes ivory-white, leaf blades widely-ovate, oblong, ovate or ovate-lanceolate, 1.5 – 8.3 × 1 – 4 cm, concolorous (emerald-green or olive-green), chartaceous or membranous, the apex acuminate, the base symmetrical, rounded or obtuse, the margins serrate, teeth congested emerging from the proximal third to the distal (immature leaves) or from the middle 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, the trichomes ivory-white or lemon-yellow, concentrated on the veins and scarce on the blade surface, veins protruding, buff-yellow or lemon-yellow, the domatia in pocket, conspicuous, subglabrous to pilose, the trichomes ivory-white or lemon-yellow, ciliate. Cymes glomerulate, peduncles 3 – 5 mm long, pilose, the trichomes ivory-white or lemon-yellow, bracts present. Staminate flowers: pedicels 1 – 2 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow, sepals abaxially subglabrous, the trichomes ivory-white, the margins ciliate. Pistillate flowers: pedicels 3 – 8 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow; ovary 3 – 7 × 2 – 5 mm, subglabrous to pilose, the trichomes ivory-white or lemon-yellow, concentrated at the base and scarce over the rest of the surface, scabrous or smooth, the style conspicuous (0.6 – 1 mm long), inconspicuous (0.1 – 0.5 mm long) or null, the stigmatic branches 2 – 4 mm long, bifid, the lobes 0.6 – 2 mm long. Drupe: Drupe: globose or ovate, 5 – 12 × 4 – 11 mm, epicarp fulvous-orange, smooth, glabrous or subglabrous, the trichomes ivory-white; mesocarp not viscous, membranous, not ornamented; pyrene ovate, 5 – 10 × 4 – 7 mm, ivory-white, verrucose surface with proeminent, rounded, randomly distributed warts, monoapiculate, the apiculum aciculate, 0.5 – 2.5 mm long, linear apex apiculum, scar absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	etymology	Etymology: — The epithet “ tarijensis ” refers to Tarija, a city in Bolivia and locality where the type was collected. Vernacular names: — Anzol de lontra, coruoia, corupiá, crista de galo, espora de galo, esporão de galo, grão de galo, grapiá, grapiá branco, grupiá, gurupiá, tobeira, unha de gato (Brazil), tala, tala-gateadora, taleira and taleiro (Argentina).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	distribution	Distribution, habitat and ecology: — Endemic to southern South America with records from Argentina, Bolivia, Brazil, Paraguay, and Uruguay (Figure 7). Celtis tarijensis prefers the Atlantic rainforest and can be found on forest edges, in clearings, in dry forests, araucaria forests, on roadsides and riverbanks. Heliophilous or sciophyte, growing under high or low light incidence. Phenology: — Flowers from August to November and fruits from November to July. Taxonomic notes: — When Weddell (1852) described Momisia tarijensis, he mentioned the red color (“ Baccae rubrae ”) of the fruits. The South American species of Celtis do not have red fruits, so we are interpreting the color “ rubrae ” as fulvous-orange. Based on this character, we identified other species native to southern South America, close to the type locality of Momisia tarijensis, that have fulvous-orange fruits. These are C. chichape (see Figure 7 E in Zamengo et al. 2024 a), C. pallida var. discolor (see Figure 7 Q in Zamengo et al. 2024 a) C. pallida var. pallida (see Figure 7 S in Zamengo et al. 2024 a) and C. spinosa (see Figure 7 W in Zamengo et al. 2024 a). After comparing the characters of the types, protologue, and other specimens, we identified differences between Momisia tarijensis and these taxa. Among the specimens examined we highlight the type (Figure 10 A) of Momisia tarijensis (P 00089387) and the others with the same identification (H. A. Weddell 4051, P 06885598, 06885599, 06885600, and 06885601). We analyzed these other specimens, because the type of Momisia tarijensis has deteriorated and consists only of leaf fragments and five fruits (Figure 10 A). After comparisons, we conclude that Momisia tarijensis is different from the four species mentioned above.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	description	Celtis chichape has rounded leaves; inconspicuous style; and crateriform pyrene surface (see Figure 7 F in Zamengo et al. 2024 a), whereas Momisia tarijensis has oval-lanceolate leaves; conspicuous style; and verrucose pyrene surface (Figure 6 Z). Celtis pallida has foliated brachyblasts, whereas C. tarijensis does not present this character. Celtis spinosa is similar to Momisia tarijensis. Both species occur in similar habitats (southern South America, Figure 7); have oval lanceolate leaves (Figure 3 D, F); the style conspicuous (Figure 5 T, 5 X); and verrucose pyrene surface (Figure 6 V, Z). Despite these similarities, the species can be differentiated by the following characters: Celtis spinosa are scrambling shrubs, up to 5 m high; with petioles 3 – 4 mm long, leaf blades up to 5 cm long (Figure 3 D); mature drupes up to 10 mm long (Figure 6 U), pyrenes up to 6 mm long with an apiculum 0.1 – 0.3 mm long (Figure 6 V); C. tarijensis are trees up to 15 m tall; with petioles 10 – 20 mm long, leaf blades more than 5 cm long (Figure 3 F); mature drupes up to 12 mm long (Figure 6 Y), and pyrenes up to 10 mm long with an apiculum that is 0.5 – 2.5 mm long (Figure 6 Z). Berg & Dahlberg (2001) synonymized Momisia tarijensis under C. iguanaea. We found characters allowing to distinguish these species. Celtis iguanaea has mature leaf blades elliptic to widely-elliptic (Figures 3 A; 9 D, E), up to 12 cm long (Figures 3 A; 9 D, E), margins entire (Figure 9 E) or with teeth restricted to the distal third (Figure 9 D); paniculiform cymes (Figure 4 F); and pyrenes with apiculum scar (Figure 6 N), whereas Momisia tarijensis has mature leaf blades widely-ovate, oblong, ovate to ovate-lanceolate (Figure 3 F), up to 8.5 mm long (Figure 3 F), the margins with teeth emerging from the proximal third to the distal third or from the middle to the distal third (Figure 3 F); glomerulate cymes (Figure 4 K); and pyrenes without apiculum scar (Figure 6 Z). After comparing these structures, we concluded that C. iguanaea and Momisia tarijensis are distinct taxa; therefore, we propose a new combination: Celtis tarijensis (Weddell) Zamengo, Chamorro & Oakley comb. nov. Nomenclatural note: — We designated an epitype (Figure 10 B) for Momisia tarijensis because the holotype (Figure 10 A) is damaged and makes it impossible to correctly identify the taxon. Because the holotype shows only few morphological characters, the identity of Momisia tarijensis was already considered doubtful; e. g., Miquel (1853) suggested that the species was a synonym of C. glycycarpa. Among the common features, Miquel (1853) mentioned that both taxa have glabrous mature leaves and pubescent immature leaves. Planchon (1873) suspected that the material Regnell 251 (P 06885588) is Momisia tarijensis, but we identified this specimen as C. spinosissima. These two species are easy to confuse as they share the same habitats, have oval-lanceolate leaves with trichomes concentrated on the veins, domatia in pockets, pistillate flowers with bifid stigma lobes, and pyrenes with a verrucose surface. Both species can be differentiated based on the color of the fruits. In C. spinosissima the drupes are yellow (primrose, Figure 6 W) and in C. tarijensis they are orange (fulvous, Figure 6 Y), as evident in the other material used by Weddell (1852) for the protologue. Like the original material, the proposed epitype (Figure 10 B) was collected in Tarija (Bolivia) and has branches with thorns, leaves, ripe fruits orange (fulvous) and pyrenes. Duplicates of the epitype are in LPB (Figure 10 C, without registration number) and MO (1429929, Figure 10 D) herbaria.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310E9D4754D5FE9078D3A87C.taxon	materials_examined	Additional material examined: — ARGENTINA. Jujuy: El Carmen, Ruta 9, a orillas del arroyo Las Lansas, - 24.4542 S, - 65.2978 W, 2 December 2016, fr., D. C. Chamorro et al. 35 (UNR). BOLIVIA. Burnet O’Connor: 2,8 Km SE of Narvaez on road to Entre Rios, 21 ° 25 ’ S, 64 ° 16 ’ W, 4 October 1983, fl., J. C. Solomon 11026 (CTES, LPB, MBM, MO, SI, U), Valley of the Río Chillaguatas, below Ranco Nogalar on trail between Sidaras and Tariquia, 22 º 5 ’ S 64 º 25 ’ W, 16 October 1983, fl., J. C. Solomon 11244 (MBM, MO, U), Entre Rios 8 Kms hacia Villamontes, 23 October 1983, fl., S. G. Beck & M. Liberman 9674 (CTES), Salinas, El Mesón área alambrada, 21 º 45 ’ S 64 º 12 ’ W, 17 October 2005, fl., S. G. Beck 31618 (L). Tarija: Aniceto Arce, 5 Km S of comunidad Guayavillas (28,3 Km S of Padcaya) on road to Bermejo, 5 May 1983, fr., J. C. Solomon 10496 (MO, SI), 7,4 Km E of Emboroza on road to Bermejo, 22 º 16 ’ S 64 º 30 ’ W, 24 April 1983, fr., J. C. Solomon 10061 (MO, SI), Upper Río Cambarí, ca. 5 Km from the first pass, on trail from Sidras To Tariquia, 22 º 10 ’ S 64 º 26 ’ W, 13 October 1983, fl., J. C. Solomon 1174 (CTES, MO), Hillsides Vicinity of Sidras, 15,5 Km N of Emborozú, 22 ° 12 ’ S, 64 ° 32 ’ W, October 1983, fl., J. C. Solomon 11126 (INPA, MO, SI, US). BRAZIL. Mato Grosso do Sul: Caarapó, fazenda Campanário, 22 ° 51 ’ 32 ” S 55 ° 3 ’ 9 ” W, 4 February 2008, fr., G. A. Damasceno-Júnior et al. 4997 (CGMS). Paraná: Curitiba, Jardim Botânico Municipal, - 25.44331 S - 49.2644707 W, 28 March 2022, fr., H. B. Z. Souza & D. F. Silva 188, 189, 190, 191, 192 (PMSP, RB). Rio Grande do Sul, Canela, 22 October 1998, fl., S. Diesel 1691 (ULBRA, US). Caxias do Sul, São Pedro, 21 October 1999, fl., A. Kegler 283 (UCS, US). Santo Antônio da Patrulha, BR 290, Rodovia Osvaldo Aranha, Km 20, 29 º 53 ’ 10 ” S 50 º 38 ’ 22 ” W, 12 April 2022, fr., H. B. Z. Souza & M. Z. Souza 202, 203, 204, 205, 206, 207 (RB). Santa Catarina: Abelardo Luz, Comunidade da Grama, assentamento José Maria, 26 ° 33 ’ 1 ” S, 52 ° 6 ’ 32 ” W, 14 April 2009, fr., A. Stival-Santos & S. Silveira 585 (FURB, IBGE). São Paulo: Bom Sucesso do Itararé, Estrada de Bom Sucesso de Itararé, a 2 Km da Mineração São Judas, 24 ° 20 ’ 07 ” S, 49 ° 04 ’ 37 ” W, 15 December 1997, fr., S. I. Elias et al. 153 (ESA, FUEL, RB). PARAGUAY. Caazapá: Yuty, 15 km S de Capitindy, 10 September 1987, fl., M. M. Arbo et al. 2794 (CTES). URUGUAY. Artigas, 4 April 1986, fr., A. I. Del Puerto et al. 18190 (MVFA).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310C9D4054D5F9477BCDA13C.taxon	description	Scrambling shrubs, 2 – 15 m tall; secondary and tertiary branches chestnut-brown, sinuous or straight, terete or sulcate, glabrous to pilose, the trichomes ivory-white; thorns 1 – 15 mm long, in pairs or solitary, curved, semi-curved or straight, maroon-red or stramineous-yellow, glabrous to subglabrous, the trichomes ivory-white, concentrated at the base and scarce along the surface of the thorns or scarce throughout. Leaf: petiole 2 – 10 mm long, subglabrous to pilose, the trichomes ivory-white, leaf blades elliptic, widely-elliptic, widely-ovate or ovate, 4.5 – 13 × 2 – 6.5 cm, concolorous (buff-yellow, cinnamon-brown, coppery-brown, emerald-green, olive-green or stramineous-yellow), chartaceous, the apex acuminated or cuspidate, the base asymmetric or symmetric, rounded or subcordate, the margins crenate-serrate, serrate or serrulate, teeth congested emerging from the proximal third to the distal (immature leaves), from the middle to the distal third (mature leaves) or restricted to the distal third (mature leaves), adaxial surface smooth, lustrous or opaque, glabrous to 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, chestnut-brown, pocket domatia, inconspicuous, glabrous to subglabrous throughout. Cymes paniculiform, peduncles 4 – 6 mm long, subglabrous to pilose, the trichomes ivory-white or lemon-yellow, bracts present. Staminate flowers: pedicels 1 – 2 mm long, subglabrous to pilose, the trichomes ivory-white, sepals abaxially glabrous, the trichomes ivory-white, margins ciliate. Pistillate flowers: pedicels 1 – 2.5 mm long, pilose, the trichomes ivory-white or lemon-yellow; ovary 2 – 3 × 1.5 – 2 mm, subglabrous to pilose, the trichomes ivory-white, throughout or concentrated at the base and scarce over the rest of the surface, scabrous, the style conspicuous (0.6 – 1 mm long), inconspicuous (0.1 – 0.5 mm long) or null, the stigmatic branches 4.5 – 8 mm long, bifid, the lobes 4 – 6 mm long. Drupe: globose or ovate, 10 – 17 × 5 – 8 mm, epicarp primrose-yellow, smooth, subglabrous to pilose, the trichomes ivory-white; mesocarp viscous, membranous, not ornamented; pyrene elliptic or ovate, 6 – 10 × 5 – 7.5 mm, ivory-white, cristate-verrucose surface with proeminent crests and randomly distributed acicular or rounded warts, monoapiculate, the apiculum aciculate, 1 – 2.5 mm long, linear apex apiculum, scar absent.	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310C9D4054D5F9477BCDA13C.taxon	etymology	Etymology: — Unknown. Vernacular names: — Azedinho, cipó farinha seca, esporão de galo, farinha seca, jatobá do brejo, laranjinha cipó, limãozinho (Brazil), cagalero, chustustis, escobo, fariña seca, guayaba, maíz tostado, noquichapirr, ojoso negro, perro isma, pomarroso, quechua, sartajchi, satajchi, satajchi amarillo, tsachik, uña de gato, uñe-gato, vara, wayja maek and yahuatáo (Bolivia, Colombia, Ecuador, Peru and Venezuela).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310C9D4054D5F9477BCDA13C.taxon	distribution	Distribution, habitat and ecology: — Distributed in northern South America and southern Central America (Figure 7). It inhabits the Amazon rainforest and is found mainly on river banks and has an affinity for sandy and clayey soils. Heliophilous or sciophyte, growing under high or low light incidence. Phenology: — Flowering and fruiting year-round. Taxonomic notes: — Celtis triflora was synonymized under C. iguanaea by Berg & Dahlberg (2001). These species should be considered distinct because C. iguanaea occurs in the southern United States, on the Caribbean islands and the Caribbean coast of South America, and northern South America (Figure 7); has branches cinereous-gray, terete; stigmatic branches 1.5 – 2 mm long (Figure 5 N) with lobes 0.6 – 1 mm long (Figure 5 N); mature drupe fulvous-orange (Figure 6 M); mesocarp non-viscous; and verrucose pyrene surface (Figure 6 N), whereas C. triflora occurs in the Amazon region of northern South America (Figure 7); has branches chestnut-brown, sulcate; the stigmatic branches 4.5 – 8 mm long (Figure 5 Z) with lobes 4 – 6 mm long (Figure 5 Z), mature drupe primrose-yellow (Figure 6 A 1), mesocarp viscous, and cristed-verrucose pyrene surface (Figure 6 A 2).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
03A0BD67310C9D4054D5F9477BCDA13C.taxon	materials_examined	Additional material examined: — BOLIVIA. Pando: Without locality, Rio Madre de Díos, 2 October 1923, fr., J. G. Kuhlmann 563 (RB). BRAZIL. Acre: Acrelândia, Rio Abuna, projeto de assentamento extrativista (PAE) Porto Dias, km 108 of BR- 364 (Rio Branco-Porto Velho), then 30 km S on side road, colocação Palhau, 9 ° 58 ’ 46 ” S, 66 ° 48 ’ 5 ” W, 4 October 2003, fl., D. C. Daly et al. 12156 (RB). Xapuri, Projeto de assentamento Agroextrativista Cachoeira, BR- 117, ramal da cachoeira, arredores da pousada, 10 ° 49 ’ 32 ” S, 68 ° 25 ’ 2 ” W, 2 November 2011, fr., H. Medeiros et al. 892 (RB). COLOMBIA. Santander: Girón, sector Cruces, Muro Presa, Antigua vía Barranca, Bucaramanga, 7 ° 5 ’ 31 ” N, 73 ° 23 ’ 19 ” W, 25 November 2017, fr., H. David et al. 5563 (HUA). ECUADOR. Napo: Ahuano, Reserva Biológica Jatun Sacha, Río Napo, 8 km abajo de Misahualli, 1 ° 4 ’ S, 77 ° 36 ’ W, 6 February 1987, fr., C. Cerón 730 (MO, US). FRENCH GUYANA. Cayenne Cedex: Remire-Montjoly, Rorota, 23 March 1984, fr., D. Sabatier 826 (CAY, P, U). GUYANA. Without a specific location: Western extremity of Kanuku Montains, in drainage of Takutu River, March 1938, fl., A. C. Smith 3237 (K, P). PERU. Junin: Chanchamayo, San Ramón. Sector Nueva Italia, 10 ° 59 ’ 20 ” S, 75 ° 25 ’ 15 ” W, 18 March 2017, fr., L. Valenzuela et al. 31791 (HOXA, MO, USM). Pasco: Oxapampa, bosque de proteccion San Matias-San Carlos, 10 ° 43 ’ 54 ” S, 74 ° 54 ’ 23 ” W, 6 July 2003, fr., A. M. Monteagudo & K. Meza 5746 (HOXA, MO). SURINAME. Brokopondo: Albufeira de Brokopondo, NW Brokopondo Stuwmeer lake, SE of Brownsberg Nature Reserve, near mouth of Withey Creek, 4 ° 35 ’ N, 55 ° 7 ’ W, 25 February 1998, fr., B. Hoffman & F. van Troon 5276 (U, US). VENEZUELA. Aragua: Girardot, orilla de la carretera vía Punta Palmita, 6 December 1979, fr., H. Rodríguez 965 (MY).	en	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): 53-98, DOI: 10.11646/phytotaxa.689.1.5, URL: https://doi.org/10.11646/phytotaxa.689.1.5
