Celtis iguanaea (Jacquin) Sargent (1895: 64)

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 : 72-77

7. Celtis iguanaea (Jacquin) Sargent (1895: 64) ( Figures 1 , 3A–A3 , 4F , 5M–N , 6M–N , 7 , 8 , 9 ).

≡ Rhamnus iguanaea Jacquin (1760: 16) as “ iguanaeus ”. Ziziphus iguanaea (Jacquin) Lamarck (1789: 318) . Mertensia iguanaea (Jacquin) Schultes (1820: 312) . Momisia iguanaea (Jacquin) Rose & Standley (1912: 8) . Celtis aculeata Swartz (1788: 53) nom. illeg. superfl. Momisia aculeata (Swartz) Klotzsch (1847: 539) . Celtis epiphylladena Ortega (1798: 79) nom. illeg. superfl. Celtis rhamnoides Willdenow (1806: 998) nom. illeg. superfl.

Lectotype: (designated by Wijnands 1983):—Tab. 73, in J. Commelijn, Horti Med. Amstelod. 1: 141. 1697 ( Figure 1 ). Epitype (designated here):— CURAÇAO. Westpoint: Bandabou, 12°21’36” N, 69°8’34” W, 27 March 2021, fl. & fr., E.A.T. Houtepen 1 (RB [01443256] image!) ( Figure 8 ).

= Ziziphus commutata Roemer & Schultes (1819: 336) . Mertensia rhamnoides Schultes (1820: 313) nom. illeg. superfl. Mertensia commutata (Roemer & Schultes) Hemsley (1883: 138) .

Lectotype (designated here):— JAMAICA. Without a specific location, s.d., fl., P. O. Swartz s.n. (lectotype: S [08-978] image!).

= Mertensia laevigata Kunth (1817: 31) nom. illeg. non Mertensia laevigata Willdenow. Momisia laevigata (Kunth) F. Dietrich (1819: 123) . Celtis laevigata (Kunth) Sprengel (1824: 932) nom. illeg. non Celtis laevigata Willdenow. Celtis glabrata Sprengel (1828: 150) nom. illeg. non Celtis glabrata Steven ex Planchon. Celtis aculeata var. laevigata (Kunth) Planchon (1873: 187) .

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!) .

= Mertensia zizyphoides Kunth (1817: 26) . Momisia zizyphoides (Kunth) F. Dietrich (1819: 124) . Celtis zizyphoides (Kunth) Sprengel (1824: 932) .

Lectotype (designated [as type] by Baehni 1936):— COLOMBIA. Bolívar: Río Magdalena, “ Mompox ” (currently: Santa Cruz de Mompox ), Minchiqueo et Peñones de Roso ad ripam Magdalenae , May 1831, fl., A.J.A.G. Bonpland & F.W.H.A. von Humboldt 1520 (second step lectotypification, designated here: P [00669757] image!; isolectotypes: B [10 0247964 fragment] image!, P [00089363, 00089364, 00089380] images!).

= Momisia ehrenbergiana Klotzsch (1847: 538) . Celtis ehrenbergiana (Klotzsch) Liebmann (1851: 339) .

Type:— MEXICO. Querétaro: Maconí, Near Moctezuma, January 1840, fr., C. Ehrenberg 1114 (holotype: B [10 0003604] image!).

= Celtis anfractuosa Liebmann (1851: 338) . Momisia anfractuosa (Liebmann) Rose & Standley (1912: 8) .

Lectotype (designated here):— MEXICO. Jalcomulco : July 1841, fr., F. Liebmann 5894 (lectotype: C [10019649] image!; isolectotype: F [614098] image!).

= Celtis aculeata var. pubescens Grisebach (1859: 149) .

Lectotype (designated here):— JAMAICA. Without a specific location, 25 July 1850, fl., A. Prior 291 (lectotype: K [000575970] image!; isolectotype: GOET [011254] image!).

= Celtis aculeata var. serrata Grisebach, (1859: 149) .

Lectotype (designated here):— JAMAICA. Without a specific location: March 1856 fl., Macfayden s.n. (lectotype: K [000575971] image!; isolectotype: GOET [011253] image!).

= Celtis platycaulis Greenman (1903: 78) . Momisia platycaulis (Greenman) Rose & Standley (1912: 8) .

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!) .

= Sarcomphalus punctatus Urban & Ekman (1926: 19) Celtis punctata (Urban & Ekman) Urban & Ekman (1928: 14) .

Lectotype (designated here):— HAITI. “ Montagnes du Trou d’Eau , near Glore ”, 22 July 1924, E.L. Ekman H-1041 (lectotype: B [10 0247969] image!; isolectotype: S [07-8525] image!).

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.

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).

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 9A ), cinereous-gray bark ( Figure 9B ), branches with stipulate or solitary thorns, curved, semi-curved to straight ( Figure 9B ); leaf blade elliptic to widely elliptic, apex acuminate to cuspidate, base rounded to subcordate ( Figures 3A , 9D, E ), margins entire (in mature leaves, Figure 9D, E ) or serrate (in immature, Figure 9C , or mature leaves, Figure 9D ), leaf surfaces concolorous (emerald-green, Figure 3A ); stigmatic branches 1.5–2 mm long ( Figure 5N ), bifid, lobes 0.5–1 mm long ( Figure 5N ); mature drupe 8.1–14 × 7.2–13 mm ( Figure 6M ), fulvous-orange ( Figure 6M ), epicarp subglabrous, trichomes ivory-white, pyrene ivory, ovoid, monoapiculate, and with verrucose surface ( Figure 6N ).

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 9B ); leaves 2.5–12 × 1.5–6 cm ( Figures 3A ; 9D, E ), glabrous or subglabrous, smooth on both surfaces, margin entire ( Figure 9E ) or with teeth restricted to the distal third ( Figure 9D ); pistillate flowers with subglabrous ovary ( Figures 5N ; 9I ), drupes fulvous-orange, up to 14 mm long ( Figure 6M ).

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 9J ) during the ripening process, but when they are fully ripe, they are fulvous-orange ( Figure 9K ).

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 1978a, 1978b) 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).

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, 8º40’18” N 83º30’17” 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), 18º24’ N, 68º37’ 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, 17º45’ N, 71º25’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, 18º3’5” N 71º19’5” 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. 200m 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, 19º28’ N 72º43’ 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, 17º59’23” N, 89º23’40” W, 5 August 1997, fr., E.S. Martínez et al. 28118 ( MBM, MEXU); a 4 km al N de Xpujil, camino a Dzibalchén, 18º32’21” N, 89º24’7” W, 11 August 1997, fr., E.S. Martínez et al. 28363 ( MBM, MEXU), 2, 4 km al NE de Bel-Há, 18º56’42” N, 89º15’29” 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, 16º45’34” N, 42º91’6” 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, 16º48’50” N, 91º3’39” W, 16 June 2003, fr., D. Álvarez 5318 ( MBM, MEXU), 3, 43 km al E de San Javier, 16º45’39” N, 91º4’33” 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, 24º9’N, 105º57’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, 15º56’23” N, 95º50’55” 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, 15º59’10” N, 95º51’46” W, 30 September 1998, fr., C. Perret et al. 44 ( MBM), Yautepec, Santa Maria Ecatepec, río Otate, al S de Santa Maria Ecatepec, 16º14’N 95º55’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, 19º37’17” N, 89º1’17” 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, 19º21’N 96º46’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, 13º24’N, 86º27’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, 13º25’N, 86º25’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, 12º5’N 86º43’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, 12º7’N, 86º23’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), 12º4’N, 86º22’W, 29 July 1977, fr., W.D. Stevens & B.A. Krukoff 2928 ( MBM, MO), ca. 1, 9 km W of Montelimar, 11º49’N, 86º32’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, 18º22’77” N 65º38’59” 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, 18º5’37” N, 65º33’10” 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, 17º28’59” N 62º57’55” 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-R11088), s.c., s.n. (S08-980), fr., P.O. Swartz s.n. (S08-975).

Figures

FIGURE 1. Lectotype of Rhamnus iguanaea Jacquin. Illustration made by Maria Moninckx (1686-1690) according to Wijnands 1983.

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

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

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

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

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

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

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