Celtis tarijensis (Weddell) Zamengo, Chamorro & Oakley, 2025

Zamengo, Henrique Borges, Chamorro, Débora C., Houtepen, Erika. T., Gaglioti, André Luiz, Pederneiras, Leandro Cardoso, Prado, Darién E. & Oakley, Luis J., 2025, Taxonomic revision of the Celtis iguanaea complex (Cannabaceae), Phytotaxa 689 (1), pp. 53-98 : 86-87

12. Celtis tarijensis (Weddell) Zamengo, Chamorro & Oakley comb. nov. ( Figures 3F–F3 , 4K , 5W–X , 6Y–Z , 7 ,

10A–D ).

≡ Momisia tarijensis Weddell (1852: 194) .

Type :— BOLIVIA. Tarija: “ In sylvis ”, August 1846, fr., H.A. Weddell 4816 (holotype: P [00089387] image!, Figure 10A ). Epitype (here designated): BOLIVIA. Tarija: Provincia Arce, 5 km S of comunidad Guayvillas (28.3 km S of Padcaya) on road to Bermejo, 22º01” S, 64º39” W, 5 May 1983, fr., J.C. Solomon 10496 ( SI [094759] image! Figure 10B ; isoepitypes: LPB [no registration number, image!, Figure 10C ], MO [1429929] image!, Figure 10D ) .

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.

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).

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 7E in Zamengo et al. 2024a), C. pallida var. discolor (see Figure 7Q in Zamengo et al. 2024a) C. pallida var. pallida (see Figure 7S in Zamengo et al. 2024a) and C. spinosa (see Figure 7W in Zamengo et al. 2024a). 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 10A ) 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 10A ). After comparisons, we conclude that Momisia tarijensis is different from the four species mentioned above.

Celtis chichape has rounded leaves; inconspicuous style; and crateriform pyrene surface (see Figure 7F in Zamengo et al. 2024a), whereas Momisia tarijensis has oval-lanceolate leaves; conspicuous style; and verrucose pyrene surface ( Figure 6Z ). 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 3D, F ); the style conspicuous ( Figure 5T, 5X ); and verrucose pyrene surface ( Figure 6V, 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 3D ); mature drupes up to 10 mm long ( Figure 6U ), pyrenes up to 6 mm long with an apiculum 0.1–0.3 mm long ( Figure 6V ); C. tarijensis are trees up to 15 m tall; with petioles 10–20 mm long, leaf blades more than 5 cm long ( Figure 3F ); mature drupes up to 12 mm long ( Figure 6Y ), and pyrenes up to 10 mm long with an apiculum that is 0.5–2.5 mm long ( Figure 6Z ).

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 3A ; 9D, E ), up to 12 cm long ( Figures 3A ; 9D, E ), margins entire ( Figure 9E ) or with teeth restricted to the distal third ( Figure 9D ); paniculiform cymes ( Figure 4F ); and pyrenes with apiculum scar ( Figure 6N ), whereas Momisia tarijensis has mature leaf blades widely-ovate, oblong, ovate to ovate-lanceolate ( Figure 3F ), up to 8.5 mm long ( Figure 3F ), the margins with teeth emerging from the proximal third to the distal third or from the middle to the distal third ( Figure 3F ); glomerulate cymes ( Figure 4K ); and pyrenes without apiculum scar ( Figure 6Z ). 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 10B ) for Momisia tarijensis because the holotype ( Figure 10A ) 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 6W ) and in C. tarijensis they are orange (fulvous, Figure 6Y ), as evident in the other material used by Weddell (1852) for the protologue. Like the original material, the proposed epitype ( Figure 10B ) 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 10C , without registration number) and MO (1429929, Figure 10D ) herbaria.

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, 22º5’S 64º25’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, 21º45’S 64º12’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, 22º16’S 64º30’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, 22º10’S 64º26’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, 29º53’10” S 50º38’22” 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).

Figures

FIGURE 3. Leaves, indumentum and domatia.—A. Celtis iguanaea, scale bar 6 cm, A1. Adaxial surface, with detail of indument, A2. Abaxial surface, with detail of indument, A3. Domatia; B. C. orthacanthos, scale bar 4 cm, B1. Adaxial surface, with detail of indument, B2.Abaxial surface, with detail of indument, B3. Domatia; C. C. pubescens, scale bar 4 cm, C1. Adaxial surface, with detail of indument, C2. Abaxial surface, with detail of indument, C3. Domatia; D. C. spinosa, scale bar 2.5 cm, D1. Adaxial surface, with detail of indument, D2. Abaxial surface, with detail of indument, D3. Domatia; E. C. spinosissima, scale bar 6 cm, E1. Adaxial surface, with detail of indument, E2. Abaxial surface, with detail of indument, E3. Domatia; F. C. tarijensis, scale bar 4 cm, F1. Adaxial surface, with detail of indument, F2. Abaxial surface, with detail of indument, F3. Domatia; G. C. triflora, scale bar 6 cm, G1. Adaxial surface, with detail of indument, G2. Abaxial surface, with detail of indument, G3. Domatia. [Photos: H.B. Zamengo].

FIGURE 4. Type of cymes.—A. Celtis alnifolia, paniculiform cyme, scale bar 2 mm; B. C. brasiliensis, glomerulate cyme, scale bar 1 mm; C. C. dichotoma, paniculiform cyme, scale bar 3 mm; D. C. diffusa, glomerulate cyme, scale bar 2 mm; E. C. gardneri, glomerulate cyme, scale bar 2 mm; F. C. iguanaea, paniculiform cyme, scale bar 1 mm; G. C. orthacanthos, paniculiform cyme, scale bar 2 mm; H. C. pubescens, glomerulate cyme, scale bar 2 mm; I. C. spinosa, glomerulate cyme, scale bar 1 mm; J. C. spinosissima, glomerulate cyme, scale bar 1 mm; K. C. tarijensis, glomerulate cyme, scale bar 1 mm; L. C. triflora, paniculiform cyme, scale bar 3 mm. [Photos: A, B, C, D, E, G, J, K, H.B. Zamengo; F, E.A.T. Houtepen; H, R. Fernandez-Hilario; I, A. Gonzáles; L. H. Medeiros].

FIGURE 5. Staminate and pistillate flowers.—A–B. Celtis alnifolia, A. Staminate flower, scale bar 0.5 mm, B. Pistillate flower, scale bar 3 mm; C–D. C. brasiliensis, C. Staminate flower, scale bar 1 mm, D. Pistillate flower, scale bar 3 mm; E–F. C. dichotoma, E. Staminate flower, scale bar 0.5 mm, F. Pistillate flower, scale bar 6 mm; G–H. C. diffusa, G. Staminate flower, scale bar 1 mm, H. Pistillate flower, scale bar 6 mm; I–J. C. gardneri, I. Staminate flower, scale bar 1 mm, J. Pistillate flower, scale bar 1.5 mm; K–L. C. hilariana, K. Staminate flower, scale bar 1 mm, L. Pistillate flower, scale bar 2.5 mm; M–N. C. iguanaea, M. Staminate flower, scale bar 1 mm, N. Pistillate flower, scale bar 4 mm; O–P. C. orthacanthos, O. Staminate flower, scale bar 1 mm, P. Pistillate flower, scale bar 3 mm; Q–R. C. pubescens, Q. Staminate flower, scale bar 1 mm, R. Pistillate flower, scale bar 1 mm; S–T. C. spinosa, S. Staminate flower, scale bar 1 mm, T. Pistillate flower, scale bar 3 mm; U–V. C. spinosissima, U. Staminate flower, scale bar 1 mm, V. Pistillate flower, scale bar 4 mm; W–X. C. tarijensis, W. Staminate flower, scale bar 1 mm, X. Pistillate flower, scale bar 5 mm; Y–Z. C. triflora, Y. Staminate flower, scale bar 1 mm, Z. Pistillate flower, scale bar 3 mm. [Photos: H.B. Zamengo].

FIGURE 6. Mature drupes and pyrenes.—A–B. Celtis alnifolia, A. Mature drupe, scale bar 9 mm, B. Pyrene, scale bar 2.75 mm; C–D. C. brasiliensis, C. Mature drupe, scale bar 10.5 mm, D. Pyrene, scale bar 3.5 mm; E–F. C. dichotoma, E. Mature drupe (type), scale bar 6 mm, F. Pyrene (type), scale bar 3.75; G–H. C. diffusa, G. Mature drupe, scale bar 6 mm, H. Pyrene, scale bar 5 mm; I–J2. C. gardneri, I. Mature drupe, scale bar 3.75 mm, J1. Pyrene (type), scale bar 2.5 mm, J2. Pyrenes, scale bar 2.5 mm; K–L. C. hilariana, K. Mature drupe, scale bar 3.25 mm, L. Pyrene, scale bar 2 mm; M–N. C. iguanaea, M. Mature drupe, scale bar 5 mm, N. Pyrene, scale bar 4 mm; O–P. C. orthacanthos, O. Mature drupe, scale bar 6.5 mm, P. Pyrene, scale bar 4.5 mm; Q–T. C. pubescens, Q. Mature drupe, scale bar 5 mm, R. Mature drupe (type), scale bar 3 mm, S. Indumentum lemon, scale bar 1 µm, T. Pyrene, scale bar 3.25 mm; U–V. C. spinosa, U. Mature drupe, scale bar 5 mm, V. Pyrene, scale bar 3 mm; W–X. C. spinosissima, W. Mature drupe, scale bar 7 mm, X. Pyrene, scale bar 5 mm; Y–Z. C. tarijensis, Y. Mature drupe, scale bar 6 mm, Z. Pyrene, scale bar 5 mm; A1–A2. C. triflora, A1. Mature drupe, scale bar 8.5 mm, A2. Pyrene, scale bar 5 mm. [Photos: H.B. Zamengo].

FIGURE 7. Distribution map of Celtis alnifolia, C. brasiliensis, C. dichotoma, C. diffusa, C. gardneri, C. hilariana, C. iguanaea, C. orthacanthos, C. pubescens, C. spinosa, C. spinosissima, C. tarijensis and C. triflora.

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

FIGURE 10. Holotype and epitypes of Celtis tarijensis (≡Momisia tarijensis).—A. Holotype of Momisia tarijensis, herbarium P; B. Epitype of C. tarijensis, herbarium SI; C. Isoepitype of C. tarijensis, herbarium LPB; D. Isoepitype of C. tarijensis, herbarium MO.