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Coccoloba P. Browne is the richest genus of tropical trees and shrubs within the Polygonaceae (Howard 1959a, b, 1961, Melo 2004, Burke & Sanchez 2011, Ortiz-Díaz 2023). The presence of an ochrea, small flowers with 5 tepals, 10 stamens and a globose to ovoid achene embedded in the tepals, and/or the expanded hypanthium are the diagnostic characteristics of Coccoloba (Melo 2004, Burke et al. 2010, Ortiz-Díaz 2023). The genus is native and widely distributed in the Neotropics, recognizing four regions with high species richness and endemisms (Koenemann & Burke 2020): Mesoamerica, The Antilles, Amazonia and Southeast Brazil.
The only monographic study of Coccoloba was carried out by Lindau in 1891, who recognized 125 species, but more recent studies estimate the number of species from 120 (Hernández-Ledesma et al. 2015) to 150 (Howard 1961, Brandbyge 1990, Melo 2004, Acevedo-Rodríguez & Strong 2012, Koenemann & Burke 2020). In Lindau´s treatment of Coccoloba, the genus was classified in four sections: Coccoloba sect. Rhigia, C. sect. Paniculatae, C. sect. Campderia and C. sect. Coccoloba, estimating 80 % of all species are including in the last one. Coccoloba sect. Coccoloba comprises the species with racemiform and spiciform inflorescences; achene entirely or partially embedded in between the tepals, and/or the expanded hypanthium obtuse or crowned, at apex. Recent phylogenetic analyzes of the subfamily Eriogonoideae have shown the monophyly of the Coccoloba and Neomillspaughia Blake placed as a sister to the genus, which together with Podopterus Bonpl. form the Coccolobeae tribe (Burke et al. 2010, Burke & Sanchez 2011, Koenemann & Burke 2020). Infrageneric classification of the sections remain untested.
As part of the taxonomic study of the Coccoloba species for Mexico, some specimens were observed that do not correspond to the species previously described within the section Coccoloba and here they are proposed as two new species. Therefore, the objective of this work is to describe, illustrate and map the distribution of two new species of Coccoloba for Mexico. The morphological characters of these new species are compared with similar morphological species to delimit them.
Materials and methods
Field and herbarium work. The study was conducted at Herbarium UADY of the Universidad Autónoma de Yucatán. During the review of herbarium specimens for the taxonomic treatment of the genus Coccoloba for Mexico, more than 150 specimens deposited in the BM, CICY, MEXU, MO, NY, UADY and XAL herbaria (acronyms follow Thiers 2023) determined as Coccoloba ascendens, C. barbadensis, C. escuintlensis, C. ibarrae, C. hondurensis and C. spicata, were examined. All type specimens of the previously listed species were consulted, as well as the general collections housed in virtual herbaria, including those maintained by JSTOR Global Plants (plants.jstor.org), Red de Herbarios del Noreste de México (herbanwmex.net), speciesLink (specieslink.net) and National Autonomous University of Mexico (MEXU; datosabiertos.unam.mx/biodiversidad). Specialized taxonomic literature on Coccoloba was consulted; in particular, Howard (1959a, b), Lindau (1891), Melo (2004) and Ortiz-Díaz (2023). Additionally, the International Plant Names Index (www.ipni.org), and Tropicos (tropicos.org) were consulted to update the current nomenclature and geographical information.
Botanical exploration was also carried out in the field to locate living populations. In November 2022, botanical exploration was carried out in the Catemaco region to collect specimens of Coccoloba burkeae sp. nov. In December 2022 the authors contacted the biologist Francisco Hernández Najarro to plan the expedition in the state of Chiapas to collect the species C. najarroi sp. nov, but unfortunately it was not possible to access living populations due to the insecurity of the region. The distribution map was created in SimpleMappr (Shorthouse 2010) using the geographic coordinates of the herbarium labels.
Morphological analysis and conservation assessment. Morphological descriptions were made from herbarium specimens. The Coccoloba burkeae specimens collected in Catemaco (holotype and isotypes) were herborized before the measurements in order to homogenize the results. In Coccoloba the size, shape and indumentation of the fruit and leaves are taxonomically informative characters and allow the delimitation of the species. These morphological characters were measured using a Mitutoyo Absolute Digimatic digital caliper (Japan). The conservation status of the new species was assessed using the IUCN Red List Criteria (IUCN 2023). We relied on criterion B, geographical distribution assessed both as B1 (extent of occurrence) or B2 (area of occupancy), as implemented in GeoCAT (Bachman et al. 2011).
Results
Coccoloba burkeae J.J. Ancona, J.J. Ortiz-Díaz & J. Tun sp. nov. (Figures 1 and 2).
Figure 1 Coccoloba burkeae J.J. Ancona, Ortiz-Díaz & J. Tun. A. Branch with leaf and inflorescence; B. Petiole and ochrea; C. Details of the inflorescence and ochreoles; D. Achene embedded in the expanded hypanthium and tepals; E. Achene.
Figure 2 Coccoloba burkeae in the locus classicus: A. Stem; B. Tree; C. Branches and fruits; D. Mature inflorescence and fruits; E. Base of the leaf-blade; F. Apex acuminate of leaf-blade; G. Fruit, from top to bottom: fleshy fruit showing the appressed perianth lobes; dry fruit showing a slightly pointed apex and tepals covering the crowned apex of the achene; achene.
Type. México. Veracruz: municipio Catemaco, Carretera Catemaco-Coyame, alrededor del Lago Catemaco, 18°26′29″ N 95°01ʹ33ʺ W, Elev. 363 m., 29 octubre 2022, J.J. Ancona, J. Tun, R. Ramos & R. Peraza 369 (holotype UADY!, isotype MEXU!, XAL!).
Diagnosis. Coccoloba burkeae is morphologically similar to C. barbadensis, C. spicata and C. hondurensis, but it is distinguished by: the leaf blade lanceolate, with long acuminate apex and obtuse base; fruit globose, of 6-6.5 (7) mm long, 5-6 mm broad, tepals appressed at the achene apex when fresh and slightly acuminate or crowned when dry, rounded base. Figure 3 shows the diagnostic characters and their comparison with morphologically more similar species.
Figure 3 Comparison of diagnostic characters between Coccoloba burkeae and related species; from left to right: C. spicata, C. barbadensis, C. burkeae and C. hondurensis. A. Fruits (achene embedded in the hypanthium and perianth lobes); B. Achenes (naked). C. Inflorescence showing the fruits and bracteoles D. Leaf-blades shape and size.
Description. Trees 10-25 m tall, bark scaly with rectangular pieces. Branches internodes solid, glabrous, striated. Ochrea 0.8-1 cm long, papery, cylindrical, glabrous. Leaves simple, alternate; petiole 1.2-1.5 cm long, 2-3 mm thick, striated, glabrous, inserted at base of the ochrea; leaf-blade lanceolate to lanceolate elliptic, 9.5-12.5(-15) × 4.5-6.5 (-7.5) cm, coriaceous to chartaceous, glabrous on both surfaces, apex acuminate to long acuminate, base obtuse; venation anastomosed, 5-7 pairs of secondary veins, primary and secondary veins prominent, glabrous. Inflorescence terminal, racemiform, 10-21cm long, raquis striated, glabrous to sparsely scabrid; bracteole 0.6-0.8 mm long, coriaceous, glabrous, cymbiform rounded in the apex; ochreolae 0.7-1 mm long, tubular, membranaceous, glabrous, lacerated; pedicel 0.7-1 mm long, glabrous. Flowers on the rachis of the inflorescence, solitary or two per fascicle, tepals 1.5-2 mm long, 0.5-1.0 mm long; hypanthium 0.8-1 mm long, green to whitish; tepals 5, 1-1.2 mm long, whitish. Fruit 6-6.5 (-7) mm long, 5-6 mm diameter, globose, glabrous; achene embedded in the succulent hypanthium and, in the tepals, this appressed at apex when fresh to slightly acuminate when dry, 6-6.5 mm long, 5-6 mm diameter, light brown-green, apex warty, base rounded.
Distribution and habitat. Coccoloba burkeae is endemic to Mexico. This species is distributed in the state of Veracruz (Figure 4). Populations are known from the high evergreen forests of the Tuxtlas region and around Lake of Catemaco at elevations of 300 to 400 m asl.
Figure 4 Geographical distribution of Coccoloba burkeae (circles) and Coccoloba najarroi (triangles).
Etymology: The specific name is dedicated to Dr. Janelle M. Burke, who has studied the phylogenetic relationships of the Polygonaceae family with emphasis on the subfamily Eriogonoideae and the genus Coccoloba.
Conservation status. The GeoCAT tool (Bachman et al. 2011) estimated the Extent of Occurrence (EOO) of C. burkeae as 3,064.323 km2, and its Area of Occupancy (AOO) as 52 km2, based on cells of 2 × 2 km. Following the IUCN (2023) criteria, the EOO and AOO results place C. burkeae in the Endangered (EN) category.
Additional specimens examined. Mexico, Veracruz, municipality Angel R. Cabada, Laguna El Majahual, 18° 39’ 14” N, 95° 18’ 17” W, 29 noviembre 1975, M. Sousa 3345 (MEXU). Municipality Catemaco, lado N de Lago Catemaco en cerros al E de Coyame, 18° 26’ 04” N, 95° 01’ 09” W, 501 m, 27 octubre 1971, J. H. Beaman 5182 (MEXU); Pipiapan, 18° 27’ 37” N, 95° 02’ 33” W, 610 m, 29 noviembre 1975, F. Ventura A. 12186 (MEXU); Laguna de Sontecomapan, 18° 31’ 03” N, 95° 02’ 04” W, 11 m, 14 diciembre 1972, R. Cedillo-Trigos & J. I. Calzada 62 (MEXU); Laguna de Nixtamalapan al N de Catemaco, 18° 25’ 54” N, 95° 06’ 00” W, 390 m, 31 mayo 1985, J. I. Calzada 11868 (MEXU). Municipality Ixhuatlán del Sureste, 5.05 km al NO de Ixhuatlán del Sureste, 18° 01’ 42” N, 94° 25’ 04” W, 17 septiembre 2011, J. Calónico Soto & B. Gómez Chagala 27839 (MEXU). Municipio Jáltipan de Morelos, Jaltipa-Tilapa, cerca del rancho Mirador Santa Isabel, al N de Ahuatepec, 17° 45’ 99” N, 94° 42’ 99” W, 27 mayo 2000, A. Rincón, G. F. Lorea, H. C. Durán E 1693 (MEXU). Municipality San Andrés Tuxtla, 6, 3 km al N de la Est. de Biol. Los Tuxtlas, yendo a Montepío, 28 mayo 2011, A. Campos V. 7401 (MEXU); Brecha de la Est. de Biol. Los Tuxtlas -Montepío, 14 julio 2004, A. Campos V. & Y. Ramírez Amezcua 6013 (MEXU); Est. de Biol. Los Tuxtlas, lote 67, 18 mayo 2004, A. Campos V. 5965 (MEXU); Est. de Biol. Los Tuxtlas, área de enseñanza, A. Campos V. 5752 (MEXU); Ejido Laguna Escondida, 32 km de Catemaco a Montepío. Dentro de 1 km de la latitud, 01 agosto 2005, E. Velasco Sinaca 718 (MEXU); Lote 71 Est. de Biol. Los Tuxtlas, 30 km de Catemaco a Montepío, 08 diciembre 2004, E. Velasco Sinaca 121 (MEXU); El poblado de Laguna escondida a 5 km de la Est. de Biol. Los Tuxtlas, 18° 35’ 27” N, 95° 05’ 16” W, 148 m, 26 julio 1974, J. I. Calzada 1392 (MEXU); Camino Laguna Escondida 1.5 km NO. Est. de Biol. Los Tuxtlas, 18° 35’ 27” N, 95° 05’ 16” W, 148 m, 20 febrero 1984, G. Ibarra Manríquez, G. Gómez V. & Santiago Sinaca Colín 1309 (MEXU); Laguna Escondida, 3 km. NO de la Est. de Biol. Los Tuxtlas, 29 diciembre 1984, G. Ibarra Manríquez & S. Sinaca Colín 2218 (MEXU); Est. de Biol. Los Tuxtlas, Laguna Zacatal o Laguna "Seca”, 18° 34’ N, 95° 04’ W, 02 febrero 1987, G. Ibarra Manríquez, L. González García & D. Van Dorp 3024 (MEXU). Municipality Santiago Tuxtla, 5.2 km W de Santiago Tuxtla, 18° 27’ 30” N, 95° 20’ 37” W, 10 octubre 1965, M. Sousa 2370 (MEXU); Cerro El Vigía Santiago Tuxtla Veracruz, 08 septiembre 1968, V. M. Toledo 157 (MEXU). Municipality Uxpanapa, 4-6 km del Campamento Hermanos Cedillo, camino a Río Alegre, 140 m, 15 junio 1974, J. Dorantes 3220 (MEXU, XAL).
Coccoloba najarroi Ortiz-Díaz & J.J. Ancona sp. nov. Figure 5.
Figure 5 Coccoloba najarroi Ortiz-Díaz & J.J. Ancona: A. Leaf-blade; B. Petiole and ochrea; C. Branch with leaf and fruits; D. Details of the pedicels and ochreoles; E. Achene embedded in the expanded hypanthium and tepals; F. Achene.
Type. Mexico, Chiapas, Municipio La Independencia, 45-50 km east of Lagos de Montebello National Park on road to Ixcan from Santa Elena. 760 m, 22 January 1982, D.E. Breedlove & F. Almeda 57722 (holotype: MO!, isotype: MEXU!).
Diagnosis. By the morphology of the fruit Coccoloba najarroi is similar to C. ascendens Duss ex Lindau and C. ibarrae J.J. Ancona & J.J. Ortiz-Díaz, but differs by its leaf blades chartaceous, glabrous, with obtuse to rounded apex and, cordate asymmetric base; inflorescence 15-30 cm; fruit oval, apex and base narrowed, 15-17.5 mm long, 10-10.5 (11) mm wide; achene included, light brown, apex helmet-shaped or semicircular, base truncated. Figure 6 exhibits of C. najarroi morphology with similar species.
Figure 6 Coccoloba najarroi morphology and related species; from left to right: Coccoloba ibarrae, C. ascendens, C. najarroi. A. Fruits (achene included in the hypanthium and tepals); B. Achenes (naked); C-E. Shape and size of the leaf blade: C. C. najarroi; D. C. ibarrae; E. C. ascendens.
Description. Tree 10-20 m tall. Branches internodes solid, glabrous, striated. Ochrea 8-10 mm long, glabrous, coriaceous. Leaves simple, alternate; petiole 2.5-4 cm long, 3-5 mm thick, striated, glabrous, inserted at base of the ochrea; leaf-blade elliptical 25-32 cm long, 14-16.5 cm wide, coriaceous, glabrous, apex slightly obtuse to rounded, base cordate; venation brochidodromous, 9-11 pairs of secondary veins, primary and secondary veins prominent, glabrous. Inflorescence racemiforme, terminal, 20-30 (-35) cm long, 3-5 mm thick, glabrous; rachis striate, glabrous; bracteole when mature 1.5-1.7 mm long, coriaceous, glabrous, cymbiform, ochreolae 1.2-1.5 mm long, tubular, coriaceous, glabrous, lacerated margin; pedicel 2.5-3 mm long, glabrous. Flowers, not seen. Fruit 15-17.5 mm long, 10-11.5 mm diameter, oval, apex and base narrowed, glabrous, ridged; achene almost completely covered by succulent hypanthium, tepals free at apex, ovoid, 7.5 mm long, 8 mm diameter, light brown, apex helmet-shaped or semicircular, base rounded to truncated.
Distribution and habitat. Coccoloba najarroi is known only from the high evergreen forests of the municipalities of La Independencia and La Trinitaria, Chiapas, around Lagos de Montebello, at elevations of 700-1,300 m asl (Figure 4).
Etymology. The specific epithet is dedicated to our friend Francisco “Paco” Hernández Najarro, botanist, collector and great connoisseur of Chiapas flora.
Phenology. Fruits in the months of November to January; bloom unknown.
Conservation status. The GeoCAT tool (Bachman et al. 2011) estimated the Extent of Occurrence (EOO) of C. najarroi as 0 km2, and its Area of Occupancy (AOO) as 16 km2, based on cells of 2 × 2 km. Following the IUCN (2023) criteria, the EOO and AOO results place C. najarroi in the Critically Endangered (CR) and Endangered (EN) categories, respectively.
Additional specimens examined. México, Chiapas, municipality La Independencia, Lower Montane Rain Forest in valley of Santa Elena along road to Ixcan, 800 m, 29 November 1976, D. E. Breedlove 41954 (MEXU, MO). Municipality La Trinitaria, Montane Rain Forest 10 km east northeast of Dos Lagos above Santa Elena. 1,170 m, 19 December 1980, D.E. Breedlove 48814 (MO). Municipality Tila, Chewpaj, 17° 17’ 24” N, 92° 24’ 50” W, 1000 m, 10 diciembre 1982, A. Méndez-Tun 5236 (MEXU). Municipality Ocosingo, la comunidad Lacandoná de Nahá, se localiza en 27.0 km. Al sureste de Palenque, por la carretera fronteriza hasta el crucero Chacalá, después 55.6 km. Por el camino de terracería hacia Monte Líbano, 16° 59’ N, 91° 36’ W, 950 m, 22 abril 1995, A. Durán F. & S. Levy T. 386 (MEXU).
Discussion
Currently, there is no revision of the genus Coccoloba for Mexico. However, Villaseñor (2016) reports 30 species of Coccoloba for the Mexican flora, of which seven are endemic. Ancona et al. (2023) described a new endemic species from Mexico plus the two new species described here, the richness of the genus increases to 33 species, of which 10 are endemic. Coccoloba burkeae is widely represented in herbarium collections. However, the specimens had been misidentified under the names of C. barbadensis, C. hondurensis, and C. escuintlensis. The first two species and C. burkeae belong to the section Coccoloba, while C. escuintlensis belongs to the section Campderia (Lindau 1891). The Coccoloba sect. Coccoloba (Lindau 1891) is distinguished by the presence of racemiform or spiciform inflorescences, the achene embedded entirely or partially by the hypanthium, and the tepals are fleshy, appressed, or free, covering the achene apex. While the Coccoloba sect. Campderia also the inflorescences are racemiform or spiciform; however, the hypanthium expands below the middle of the achene and the tepals are papery, free and completely enveloping the rest of the achene.
Due to the morphological similarity of Coccoloba burkeae with the aforementioned species, they can be considered as part of the C. barbadensis complex, which includes species that are characterized by the presence of chartaceous, ovate-elliptic leaf-blades with cordate to subcordate base, the inflorescences may vary from spiciform to racemiform, with short pedicels less than 1.5 mm long, succulent fruits, ovoid or spherical, apex prominent or crowned. They are distributed from Mexico to Honduras. Throughout its taxonomic history, 16 names have been published in this complex, but only five of them are accepted (Howard 1992), including the species Coccoloba floresii (Ortiz-Díaz et al. 2015) that were determined in the herbarium collections such as C. barbadensis or C. hondurensis. A taxonomic revision in this complex could elucidate and define the limits between the species of this complex.
Regarding their distribution, Coccoloba burkeae and C. barbadensis apparently are distributed in the same geographical areas of Veracruz. However, in ecological terms they are very different. C. burkeae inhabits more humid areas, in high evergreen forests, riverbanks and lagoons with very humid or swampy soils. While C. barbadensis occupies drier areas, in low deciduous forest vegetation, dry and stony soils. On the other hand, the species C. hondurensis and C. spicata are distributed in disjunct geographical areas and far from C. burkeae. Coccoloba spicata is distributed in the Yucatan Peninsula biotic province and C. hondurensis in high evergreen forests of Chiapas and Central America (Belize, Guatemala, El Salvador and Honduras).
The species Coccoloba najarroi seems to be a very different species from those known in Mexico and Central America. However, in morphology of the fruit it is very similar to C. ascendens and C. ibarrae, a recently described species of section Paniculata (Ancona et al. 2023). While C. ascendens and C. najarroi belong to the section Coccoloba. In the few herbarium collections, C. najarroi were identified as C. spicata. However, they are morphologically very different, C. najarroi presents racemose inflorescences and C. spicata, as its name indicates, presents spiked inflorescences.
In terms of its distribution, C. najarroi is only known from the high evergreen forests of the municipalities of La Independencia and La Trinitaria in the state of Chiapas. While C. ibarrae is known from the cloud forests of Oaxaca and Veracruz and C. ascendens is known from the humid forests of Panama and northern South America.
