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Casearia emarginata C. Wright ex Griseb. (Salicaceae), was proposed as a member of the Flacourtiaceae by A.H.R. Grisebach in 1866 (Figure 1A) based on a plant collected by Charles Wright (Wright 1894, MO!!) in Cuba. The plant was described as having obovate, subrotund, emarginate leaves. In the same page, Grisebach also described a second species, C. ramosissima, based on another of Wright’s specimens (Wright 1893, MO!!), which was said to be related to the precedent but differing in being a shrub 3-6 feet high with white flowers. These two characters were not provided for C. emarginata. However, C. ramosissima was further described as bearing divergent branches, which are rigid and subspinescent. Both species were said to have been collected in eastern Cuba at the locality of Quemado (Grisebach 1866).
Figure 1 Types of Casearia ehrenbergiana Urban and C. emarginata C. Wright ex Griseb. A) Casearia ehrenbergiana (Ekman 9278, NY). B) Casearia emarginata (Bro. Hiram 4195, NY).
In 1899 Ignatz Urban, then working in his Symbolae Antillanae, proposed a third species, Casearia ehrenbergiana (Figure 1A), based upon a collection of Carl A. Ehrenberg from Haiti (Ehrenberg 9, HAL!). This taxon featured lanceolate or obovate leaves and a different vegetative architecture. In its comments on the new species, Urban stated that it was fairly similar to C. emarginata but indicated that C. emarginata featured shorter, non spinescent branches, and the spinescent branches are conspicuous in the type plant of this name (Urban 1899).
In 1941 Cyrus L. Lundell, in one of its many contributions on the Flora of Yucatan, described a fourth entity, Casearia randioides (Figure 2), based upon a collection by himself and his wife Amelia Lundell (Lundell & Lundell) along the Mérida-Progreso road in northern Yucatán, Mexico. The protologue included an illustration of a relatively common shrub or subshrub found in many places of the Yucatán Peninsula Biotic Province (YPBP sensu Carnevali Fernández-Concha et al. 2021). In the original description, Lundell stressed “ … branchlets stout, erect or suberect … internodes short, unarmed.” (Lundell 1941).
Figure 2 Casearia randioides Lundell (Salicaceae). A) Branch with short brachyblasts. B) Branch without brachyblasts. C) Flowers blooming from the branch. E) Details of the flowers. D Herbarium specimen (G. Carnevali et al. 10361, CICY). (Photographs A-E. Germán Carnevali. F. Mayte R. Aguilar Canché ).
These four entities are unusual among Neotropical Casearia Jacq. species because of their general habit and very small flowers and they have been variously synonymized in the literature (see below). The leaves are fasciculate along shortened branchlets or brachyblasts, and range from suborbicular to obovate or obovate elliptic or even lanceolate for C. ehrenbergiana, apically rounded to emarginate, sessile or almost, the base cuneate, 10-24 (-27) mm long, with a broadly crenate margin, the abaxial face glabrous to laxly puberulent along the midnerve; the flowers are minute with narrowly oblong, truncate green tepals ca. 3.8-4.2 × 0.8-0.9 mm; stamens are 10, whitish or very pale green, with the stamens glabrescent whereas the corona-forming staminodes are densely pubescent. Taxa referrable to this group of entities are hereinafter referred to as the Casearia emarginata-complex. Populations belonging to these four, apparently closely related entities, are distributed in the Mexican Yucatán Peninsula, in Cuba, and in the Hispaniola (Sleumer 1980, Gutiérrez Amaro 2000). We will henceforth refer to this aggregate of taxa as the Casearia emarginata complex.
Populations of this complex belong to Casearia sect. Decandrae (sensuSleumer 1980), probably a polyphyletic aggregation of taxa (de Mestier et al. 2022) that are characterized mainly by their flowers with 10 subequal stamens, and subglobose, narrowly ovoid or oblongoid flowers in overall shape, their base rounded or attenuate just before anthesis, their apex attenuated in bud. However phenetically similar, these series of populations seem to be easily discernible into several taxa based on a few striking characters. The most distinct are the plants for the Mexican portion of the Yucatán Peninsula (MYP). These plants have fairly straight, few-branched stems bearing very short brachyblasts; these, as well as the branches, lack a spinescent apex. Given this main difference between the populations referred as Casearia emarginata from the Yucatán peninsula, and those from the Antilles, we analyzed relevant local populations with the comparative method. The main question is whether the populations currently assigned to Casearia emarginata in the MYP are conspecific with the West Indian species and, if not, whether they should be referred to any other described taxon or constitute an undescribed taxon. More specifically, we address the following question: is the species from the MYP different from Casearia emarginata or other similar species of Casearia sect. Decandrae?
The latest updates of the Angiosperm phylogenetic classification (APG IV 2016), particularly of the Malpighiales, expand the family Salicaceae to include a large part of Flacourtiaceae, whereas another portion of the family was assigned to the family Achariaceae. These molecular analyses also affected Casearia (Alford 2005), because members of the genera Hecatostemon S. F. Blake, Laetia sect. Laetia, Laetia sect. Casinga (Griseb.) Warburg, Samyda Standl., and Zuelania A. Rich. (Samarakoon & Alford 2019) as well as Euceraea Mart. and Neoptychocarpus Buchheim (de Mestier et al. 2022), are now generally included within Casearia.
According to their ecological preferences in the Mexican Yucatán Peninsula, Casearia species could be arranged in two main groups: those that occur preferentially in the dry northwest whereas others occur in the more humid areas of the south of the península. The first group, which includes fewer species, occurs mainly as deciduous understory shrubs or subshrubs in dry forests on thin, skeletal, rocky calcareous soils and usually flower along with a flush of new leaves. The second group includes taxa from tropical subhumid forests and are medium to large-sized trees (4-10 m) in the canopy or upper strata of the forests.
Materials and methods
The study was conducted at the Centro de Investigación Científica de Yucatán, AC. (CICY), Yucatán, Mexico, at the Oakes Ames Orchid Herbarium, and the Missouri Botanical Garden during 2023-2025. Herbaria in tropical America and the USA were visited during 2024. Plants and materials in the field in Mexico were obtained under scientific permits (SGPA/DGVS/008421/18 and SGPA/DGGFS/712/2913/17) issued by the Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT) to researchers at CICY. The study covers the area constituted by the Yucatán Peninsula Biotic Province, henceforth YPBP, particularly, the three Mexican states of the peninsula (MYP), as well as Cuba and Hispaniola.
Herbaria specimens relevant for Casearia and the biogeographical area relevant to this study were examined, either physically or through digital access. These include A, CAS, CICY, DUKE, F, GH, K, L, LL, MEXU, MICH, MO, NY, TEX, UADY, UCAM, and US (acronyms according to Thiers, 2025). Specimens revised where cited as follows: those physically studied by us are indicated with !!, those that were digitally seen as well as in photographs are indicated as !, whereas those cited from the literature but otherwise not seen by us are not marked in any way. In the main text, we cite only a handful of specimens for each Mexican state of the Yucatán peninsula, as well as West Indian material. Other specimens are found in Supplementary Material. Collected specimens were deposited at CICY and duplicates sent to the herbaria indicated after the citations. A further step consisted in analyzing the protologues of morphologically similar species of Sect. Decandrae (sensuSleumer (1980) as well as relevant floristic treatments featuring descriptions and keys (Nee 1999, Liogier 1994, Gutiérrez Amaro 2000), and Pool 2001). Nomenclature decisions follow the Madrid code (Turland et al. 2025). Species treated here were defined according to the Unified Species Concept (de Queiroz 2007) where species are considered as separately evolving metapopulations where the secondary species criteria (morphology, biogeography, molecular data, etc.) are treated as different lines of evidence (operational criteria) relevant to assessing lineage separation.
The description of the Casearia emarginata-like populations from the Mexican Yucatán Peninsula was prepared from live, pickled, and herborized material, complemented with available iconography. Measurements were taken with a ruler or with a Metromex Vernier caliper. Photographs by the authors of both live plants and specimens of Casearia from the Mexican Yucatán Peninsula were taken with a Nikon SD330 camera with an AF-S DX NIKKOR 18-55 mm lens (mainly photographed by G. Carnevali) or with a Nikon D800 and D850 cameras with a Nikon Micro 105 f/2.8G IF-ED AF-S VR lens (most of the photographs taken by Alfredo Dorantes). Cartography was created with QGIS Development Team 2025 (qgis.osgeo.org) and edited with Inkscape Project 2025 (inkscape.org). The composite plates were created and edited with GIMP Development Team (2025) and with Adobe Photoshop (Adobe Inc. 2019).
Herbarium material was rehydrated by boiling and then soaked in a soapy solution or in a concentrated ammonium hydroxide solution for about one minute, then rinsing it in water. The flowers were returned to the corresponding herbarium specimens; no additional material received such treatment. The study of the MYP’s Casearia was relatively easy based upon actual specimens and digital images of herbarium material, and after studying a large amount material we became familiar with the taxa of the genus in the Mexican Yucatán Peninsula because of our knowledge of overall morphology, geographical origin, and ecological preferences. This allowed us to confidently refer particular herbarium specimens of Casearia to the species that occur in the area.
The conservation status of Casearia taxa of the Mexican Yucatán Peninsula is provided only for taxa restricted to the region (three species) and was assessed according with the IUCN Red List Criteria (IUCN 2012) and the IUCN Red List Categories and Criteria Version 3.1 (IUCN 2016). Because population data were not available, we relied on the B criteria, geographical distribution, assessed both as B1 (EOO, extent of occurrence) or B2 (AOO, area of occupancy) as implemented in GeoCAT (Bachman et al. 2011). We complemented these assessments with our own field experience, information and opinions provided by experts, published data and iconography, whenever available. Distribution was documented through herbarium vouchers and online resources, like iNaturalist (www.inaturalist.org) and Symbiota (https://sample.symbiota.org/collections/list.php?usethes=1&taxa=15678).
Results
Populations of the Mexican Yucatán Peninsula currently assigned to Casearia emarginata are morphologically distinct from populations of C. emarginata from Cuba and Hispaniola (Table 1) because they lack conspicuous, short branches transformed into spines found in their West Indian counterparts. The elongated branches with spinescent tips of these West Indian plants often lack leaves but can produce flowers. In the type specimens of Casearia emarginata and C. ehrenberghii these spinescent branches are thin and more or less perpendicular to the axis of the main branches and most conspicuous when leafless. Some other specimens feature thick, short and thick, spinescent branches. Relevant differences between the two sets of populations are summarized in Table 1. Information about the color of the flowers of the West Indian plants is scant but at least in one case (Eyerdams 58, A, GH!!, MO!!) the flowers are described as pinkish as opposed to the Mexican Yucatán Peninsula plants where flowers are white and green. The name Casearia randioides Lundell, based upon a Yucatán specimen is available for the Mexican Yucatán Peninsula populations, which are also a perfect match to Lundell’s species circumscription. We herein provide an amended, amplified description based only upon material from the Mexican Yucatán Peninsula.
Table 1 Relevant morphological character states distinguishing between Casearia emarginata s.l. (including C. ehrenbergiana) and C. randioides.
| Casearia emarginata (Cuba)* | Casearia randioides (Mexican Yucatán Peninsula) | |
|---|---|---|
| Branches * | apically spinescent | Apically not spiny |
| Leaf blade, shape * | oblong-elliptic, obovate-elliptic or elliptic | obovate, obovate-elliptic, obovate-circular |
| Leaf margin * | revolute, slightly serrate, laxly toothed | with 1-3 teeth on the superior portion or entire |
| Secondary nerves, number * | 4-6 | 2-4 |
| Inflorescence, flower number | 3-5 (rarely more) | 1-2 |
| Calyx lobes, shape * | suborbicular to elliptic | oblong-elliptic |
| Calyx lobes, size (mm) * | 1.5-2.0 | 2.0-3.5 |
| Filament length (mm) * | ± 1.0 | 2.4-2.5 |
| Filament, pubescence | Only on the distal section | pilose or subglabrous |
(*) The data and measurements of Casearia emarginata are based on Gutiérrez Amaro (2000) and our study of herbarium material.
Casearia randioides Lundell, Contributions from the University of Michigan Herbarium 6: 48, f. 3. 1941. Type: México. Yucatán, km 20 Mérida-Progreso road, fl., May-Aug, 1938, C.L. Lundell & A.A. Lundell 7982, (holotype MICH!, isotypes CAS!, DUKE!, F!, K!, L, LL!, NY!, TEX!, US!!). Figures 2A-D; 5C, D
Casearia emarginata Auct. non Wright, sensuSleumer, 1980, Sosa et al. 1985, Durán et al. 2000, Carnevali Fernández-Concha et al. 2010.
Suffrutescent shrub to rarely a small tree, 1-3(-4) m tall. Branches sericeous, erect or suberect, glabrous or slightly puberulent, lacking a terminal spine; branches where leaves arise forming brachyblasts, about 1 mm long, stipules present, minute, drooping, petioles subsessile or short up to 1.5 mm long. Leaves simple, alternate, arranged more or less in a single plane, chartaceous, obovate, obovate-elliptic, obovate-circular, (0.9-)1.2-1.8(-2.2) × (0.6-)0.8-1.4(1.8) cm, apex rounded, retuse, base cuneate, margins with 1 to 3 teeth on the upper portion of the lamina to completely entire, abaxial face glabrous or sparsely puberulent along midrib, adaxial face glabrous, 2-4 secondary nerves ascending. Flowers minute, white and greenish, 1-2 per node; 1.0-2.0 mm long, puberulent. Calyx pentamerous, 2.0-3.5 mm long, lobes oblong-elliptic, dull green, glabrous, apex rounded, margins sparsely ciliate. Corolla absent. Stamens 10, white, united at base for 0.4-2.5 mm, pilose or sparsely pilose, alternating with 10 smaller staminodes together conforming a shallow corona, hairy on its outer surface and on the margins; anthers yellow, basifixed, about 0.5 mm long, filaments white. Ovary about 1 mm high, ellipsoidal, style 1.5 mm long, glabrous, attenuate toward apex, stigma yellow, capitate, entire. Fruit a spherical-ovoid capsule, 0.6-0.7 × 0.5-0.6 cm, rugulose, 3-valvate, shortly apiculate, first green, purple upon maturity, drying black; seeds 6-8, obovate, spherical, 4-5 × 1-1.1 mm, glabrous, shiny, enveloped by a red aril.
Our revision of the Casearia herbarium specimens collected in the MYP reveals that beyond C. randioides, there are seven additional species in the area, namely C. aculeata Jacq., C. corymbosa Kunth, C. laetioides (A. Rich.) Northr., C. subsessiliflora Lundell (reinstated from the synonymy of C. silvestris Sw. by Carnevali Fernández-Concha et al. 2010), C. thamnia (L.) T. Samar. & M.H. Alford, C. tremula (Griseb.) Griseb. ex C. Wright, and C. yucatanensis (Standl.) T. Samar. & M.H. Alford. These are synoptically summarized below after the discussion section.
Discussion
Sleumer (1980) accepted a broad concept for the Casearia species with small leaves, reduced vegetative size, and minute flowers with ten stamens. He applied the name Casearia emarginata to them and three names were included under synonymy: C. ehrenbergiana, C. ramosissima (but see below; Gutierrez-Amaro synonimyzed this taxon with C. littoralis T.Samar. & M.H.Alford, a movement we do not agree with), and C. randioides. The reasons for this synonymization were not explained, but it is possible that the general appearance (e.g., reduced vegetative size, dimensions and shape of the leaves) and the branches transformed into spines were contributing factors, but this would not apply to C. randioides, which is not armed. Here, we follow authors dealing with the West Indian members of this complex (e.g.,Gutiérrez Amaro 2000) and treat them as a single species under the name C. emarginata although acknowledging that C. ramosissima may be distinct. There is, however, apparently much variation within these plants, particularly in leaf shape and the thickness of the spinescent branchlets as well as their angle relative to the main stem, suggesting that more than one taxon might be involved. Even very broadly defined, at least two sets of populations are readily distinct based on vegetative architecture and floral structures. The first set of populations are restricted to the Mexican Yucatán Peninsula; the second, to Cuba and Hispaniola.
As stated above, the Mexican Yucatán Peninsula populations correspond to the taxon proposed by Lundell, Casearia randioides as they match perfectly the type specimens and the description provided in the protologue. Thus, we reinstate this species from the synonymy of C. emarginata and supply the information (morphology, distribution, and aspects of its ecology) not originally provided by Lundell (1941). The specific epithet suggests a similarity to species of Randia L. (Rubiaceae); in this case with R. obcordata S. Watson, and to a lesser degree, to the MYP endemic R. truncata Greenm. & C.H. Thomps., both of which are sympatric with C. randioides, but have opposite leaves and paired spines at the internodes and endings of branches. Furthermore, Casearia randioides superficially recalls the endemic, also sympatric, Acalypha gaumeri Pax & K.Hoffm. (Euphorbiaceae) that is a smaller plant with strikingly different flowers. The fact that several totally unrelated, vegetativelly similar taxa, coexist in these habitats suggests similar selective pressures leading to convergent evolution.
On the other hand, as compared to the West Indian populations of the Casearia emarginata complex, C. randioides is much more homogeneous and most plants bear leaves that are obovate, obovate-elliptic, or obovate-circular, and only rarely elliptic. Furthermore, the West Indian plants are apparently much more floriferous and the flowers are borne on brachyblasts on the spinescent branchlets, whereas in C. randioides, the flowers arise directly from the main stem. Finally, the flowers in C. emarginata are clearly pedicellate whereas in C. randioides they are almost sessile. Both species seem to prefer calcareous soils or rocky outcrops. In the case of Casearia ramosissima, it is important to mention that it is a clearly distinct species bearing flowers lacking staminodes. Because of this, it has been placed in Samyda (Gutiérrez Amaro 2000), although here we follow Samarakoon & Alford (2019) in treating it as a member of Casearia. Thus, in the light of all this evidence, and because we do not know these plants firsthand, we have refrained from making any taxonomic decisions on the West Indian members of the complex.
Selected material examined (see Supplementary Material for additional records). Mexico. Campeche. Mun. Calakmul, 2 km al NE, 18° 07’ 24” N, 89° 47’ 25” W, 8 September 1997 (fr.), E.M. Martínez, D. Álvarez, S. Ramírez & G. Bacab 28209 (CICY!!, MEXU!); Mun. Campeche, 1 km al S de Samulá, 19° 48’ 30” N, 90° 33’ 00” W, 28 June 2003, (fl., fr.), C. Gutiérrez 7810 (CICY!!, MEXU!); Mun. Hopelchén, 3.1-3.3 km en línea recta de la salida de Dzibalchén por la carretera hacia Xpujil, 1.1-1.3 km a lo largo de una terracería que sale al lado NNE de la carretera DzibalchénXpujil y que llega a campos de cultivo y milpas, 19° 27’ 21” N, 89° 41’ 51” W, 27 November 2023 (flowering), G. Carnevali et al. 10361 (CICY!!); Quintana Roo. Mun. José María Morelos, 3 km al E de la desviación de Presumida, por el camino a Santo Tomás, cruzando la Laguna Esmeralda, por la carretera Ticul, Yucatán a José María Morelos, 178 m snm, 19° 48’ 40” N, 88° 43’ 00” W, 20 September 2002, (fl., fr.), J.L. Tapia et al. 1313 (CICY!!, MEXU!!); Mun. Solidaridad, camino de Rancho Viejo a Balcheil, km 12, Tulum- Cobá, 20° 17’ 50” N, 87° 31’ 10” W, 10 March 1974, (fl.), R.M. López-Franco et al.132 (CICY!!, UCAM!). Yucatán: Mun. Kinchil, Área de Conservación de Kinchil (ACK), propiedad de Kekén, unos 8-9 km al WWS del desvío a Kinchil por la carretera Mérida-Celestún, unos 7.8 km al W de Bella Flor, unos 20-21 km al E del puente a la entrada de la Ría Celestún. Colecciones en los alrededores y márgenes del helipuerto en el extremo SW del área de los biodigestores de Kekén, unos 6-6.3 km al S de la carretera Mérida-Celestún, 90 m snm, 20° 48’ 37.45” N, 90° 08’ 20.5” W, 17 March 2022, (fl.), G. Carnevali et al. 9073 (CICY!!, MEXU!!); Mun. Mérida, carretera Mérida-Progreso, a 1 km del desvío hacia Dzityá, 21° 03’ 31” N, 89° 38’ 26” W, 18 November 2003, (fl., fr.), F. May et al. 2430 (CICY!!, MEXU!!); Mun. Muna, 2.5 km al NE de la entrada a las granjas Kekén y 1.5 km al W de la carretera Mérida-Muna, 11 km antes de llegar a Muna, 167 m snm, mosaico de vegetación muy perturbada con remanentes relativamente bien conservados, 20° 35’ 51” N, 89° 43’ 18” W, 15 September 2009, (fl.), J.L. Tapia et al. 2063 (CICY!!, GH!!).
Phenology. Casearia randioides flowers from November to September. It fruits from March to November.
Distribution and habitat. Casearia randioides is endemic to the Mexican Yucatán Peninsula, in the states of Campeche, Quintana Roo, and Yucatán, where it grows mainly in tropical dry forest of low to medium height, as well as in the secondary vegetation associated with these forests. It is usually found in the understory, growing on rocky, calcareous soils. It has been very rarely collected in flower or fruits, presumably because these are inconspicuous, and, at least the flowers, fugacious.
IUCN Conservation assessment. LC (Least Concern). Casearia randioides meets criterion B2a for an Endangered species (EN) but it does not meet at least two of the three additional conditions necessary for the assignment of this category (IUCN 2016). It is known from many localities (62) in the three states of the MYP, its Extent of Occurrence (EOA) is of 93.632.609 km2 (far exceeding the 20,000 km2 limit) and the Area of Occupancy (AAO) is of 176,000 km2. Because of this last figure, it could have been assigned to the category of an Endangered species (EN) but it only meets one (B Criteria) of the three additional conditions. The habitat of the species is moderately fragmented under natural conditions but is being severely threatened by anthropogenic activities, particularly touristic and agricultural developments. However, the species is relatively common, apparently resilient, and locally abundant. In fact, as stated before, the number of herbarium collections most likely does not accurately reflect the abundance of the species as many collectors fail to gather it because the flowers and fruits are inconspicuous and only very occasionally detected. Furthermore, several populations (about 10) are known to occur in the Biosphere Reserve of Calakmul (Campeche), and others in a few state of private reserves. This is the first conservation assessment of this species, and for all reasons mentioned before, the species is best regarded as LC.
Casearia emarginata s.l. additional material examined. CUBA. Occidente. Prope Quemado, 1860, C. Wright 1894 (type of C. emarginata, holotype, isotype, MO!! (MO-279838/A:2063163), photograph, NY!); prope Quemado (Quermado), February 2, 1861, Charles Wright 1893 (type of C. ramosissima, holotype, B!, isotypes, BM!, G!, GH!!, GOET!, K!, L!, MO!!, S!, YU!, S!). Habana, Vedado, Coastal thickets, Renté, Santiago Bay, Bro. León 9777 (GH!); Vedado, Río Macambo, Vía Azul, 28 December 1954, Bro. Alain & M. López Figueiras 4193 (as C. emarginata, GH!!). Centro: Santa Clara, Río San Juan, “rocky hillside … tree 8 m”, 24-25 May 1910, N.L. Britton, F.S. Earle & P. Wilson 5882 (as C. emarginata, NY!). Oriente. Santiago de Cuba, Renté, April 1973, Bro. Clemente 2623 (LS, NY); Santiago de Cuba, “Low hills ... shrub, 2 m tall”, 9 August 1913, Bro. Leon 3936 (NY); Vicinity of Guantánamo,
“Dry, poor soil; shrub 8 feet”, 23 February 1911, J.A. Shafer 8910 (as Casearia ramosissima, NY!); Guantánamo, Finca San Francisco, 21 October 1920, Bro. Hioram 4195 (as C. emarginata, GH!! NY!); Novaliches, Guantanamo, shrub, coastal thickets, 5 January 1939, Bros. León & M. Victorin 18648 (as C. emarginata, GH!!, NY!). HAITI. Hispaniola, no date, C.A. Ehrenberg s.n., (type of C. ehrenbergiana, holotype B; isotypes, F, GH!!). Massif de la Selle, 25 August 1926, E.L. Ekman 6737 (as C. ehrenbergiana, A, GH!!); Gonave Island, Pte. A Raquette “tree in mountain slope, flowers pinkish”, 200 m, 24 July 1927, W.J. Eyerdam 58 (as C. ehrenbergiana, A, GH!!, MO!!, NY!); Plaine Cul-de-Sac, Port-au-Prince at Fort-National, 11 November 1927, E. L. Ekman 9278 (as C. ehrenbergiana, GH!!, NY!); Plaine Cul-de-Sac, Port-au-Prince, road to Pétionville, Limestone, 13 April 1926, E.L. Ekman 5852 (as C. emarginata, MO!!); Gonave Island, Pte-ã-Raquettes, “light forest, 2-3 m high, very common, 24 November 1927, E. L. Ekman. H8650 (MO!!); vicinity of Mole St. Nicolas, “Shrub 10 feet high, branches numerous, flowers green; road through mole gorge”, 13-19 February 1929, E.C. Leonard & G. M. Leonard 13133 (as C. ehrenbergiana, NY!).
Synopsis of Casearia in the Mexican Yucatán Peninsula. Casearia species are a common and widespread floristic element in the MYP, yet are rarely found in fertile condition. As in many other tree groups, fruiting material is usually more commonly represented in herbaria than flowering material. As a genus, they can be recognized vegetatively by alternate, entire, pinnately nerved leaves, that are usually pellucid-punctate or pellucid-lineolate and usually bear serrate margins (Gentry 1993). The flowers lack petals and usually have a series of staminodes aranged in a coronalike manner. We offer the following key for the Casearia species native of the three Mexican states of the Yucatán peninsula.
Key to Casearia in the Mexican Yucatán peninsula
1. Leaves obovate, broadly obovate to spatulate obovate, the apex rounded to obtuse, truncate to emarginated; flowers solitary or geminate (rarely 3 flowers), axillary ................................................................................2
1. Leaves ovate to narrowly ovate, elliptic, oblong to lanceolate-oblong, apex acute, acuminate, shortly obtuse to subacute; flowers in axillary or terminal fascicles or corymbs .......................................................................3
2. Leaves obovate to obovate-cuneate, in short branches (brachyblasts), 2.0-5.0(-6.5) cm × 1.5-3.5 cm, pubescent on both sides, the pubescence denser and yellowish on the underside; flowers solitary, 1.3-2.2 cm long, white to pale yellow, very fragrant; stamens (10-)12, almost entirely fused to form a tube, adnate to calyx tube; disc without lobes or appendages; stigma depressed-globose, inconspicuously 3-4 lobed; fruit capsular, ellipsoid to subglobose, 1.5-2.0 cm diameter ........................................................................ C. yucatanensis
3(1). Leaves ovate to narrowly ovate; flowers sessile or subsessile, in dense axillary fascicles, wholly covered with scarious bracts; style trifid; stigma capitate ................................................................. C. subsessiliflora
3 Leaves oblong, oblong-elliptic to lanceolate-elliptic; flowers pedicellate in dense axillary or terminal fascicles, or in distinctly pedunculate corymbose inflorescences, not covered by scarious bracts; style terete, short or absent; stigma capitate, depressed-globose or subsessile, peltate, flattened ........................................................... 4
4(3). Flowers grouped in dense axillary or terminal fascicles ..................................................................... 5
4. Flowers clustered in distinctly pedunculate corymbose inflorescences ....................................................... 6
5(4). Shrubs or trees often with strong thorns on trunk and larger branches (not common in herbarium material); leaves elliptic to oblong-elliptic, 6.0-10.5 × 2.0-4.5 cm, apex acute or shortly acuminate; base cuneate to obtuse; flowers in axillary fascicles, 3.0-4.0 mm long, 1.5-2.0 mm diameter; stamens (7-)8(-9); style conspicuous, short; stigma capitate; fruit capsular, subglobose, 5-10 mm diameter, glabrescent........ C. aculeata
5. Trees without thorns on trunk or branches; leaves oblong to oblong-elliptic, (6-) 10-25 × (2.5-) 3.0-6.0 (-9.0) cm, apex subacute, rounded-obtuse or acuminate, base obtuse to rounded, subcordate, asymmetrical; flowers in dense terminal fascicles sometimes axillary, 6. 0-7.0 mm long, 3.5-5.0 mm diameter; stamens 20-40; style absent; stigma inconspicuous, subsessile, peltate, flattened, thick; fruit baccate, depressed-globose, fleshy and succulent, 2.5-7.5 cm diameter, puberulent ............................................................................. C. laetioides
6(4). Leaves chartaceous, glabrous on both sides; inflorescences corymbose, terminal or axillary, pedicels not acrescent in fruit; flowers fragrant; stamens 60-80; disc without lobes or appendages; stigma slightly 4-5 lobed; fruit ferruginous-tomentulate................................................................................................... C. thamnia
6. Leaves subcoriaceous, glabrescent or hairy along lower half of midrib on underside; inflorescences corymbose to corymbiform or racemes, always axillary, pedicels acrescent in fruit; flowers not fragrant; stamens 6-24; disc lobed or appendaged; stigma capitate, unlobed; fruit not ferruginous-tomentose.......................................... 7
7(6). Leaves oblong-elliptic; apex acute, rarely acuminate, base acute, rounded, to obtuse-truncate, sometimes slightly cordate, inconspicuously asymmetrical, glabrescent, opaque on both sides; lateral veins rather irregular, conspicuous on both sides, mainly on underside, reticulation of nervation inconspicuous; petiole straight, 4-6 mm long; inflorescences corymbose to corymbiform, axillary; flowers 3-4 mm long, ca 2 mm diameter; stamens (6-)8; disc with linear-spatulate lobes; fruit globose to ellipsoid, 1.-1.2 cm diameter, not fleshy, opening always in 3 valves............................................................................................................................... C. corymbosa
7. Leaves ovate-oblong or ovate-elliptic, sometimes elliptic, rarely even lanceolate on the same plant, apex shortly obtuse, attenuate or subacuminate, sometimes rounded or even slightly emarginate, base cuneate to rounded, rarely subcordate, conspicuously asymmetric, glabrous or hairy along the lower half of the midrib on the underside, shiny on the upper side; lateral nerves arcuate, the upper ones closer together, conspicuous only on the underside, reticulation of dense and finely conspicuous veins on both sides; petiole curved, often deflexed, (5-)10-18(-24) mm long; inflorescences in much abbreviated racemes or corymbs; flowers 4-6(-8) mm long, 2-3 mm diameter; stamens 15-24; disc with rounded lobes; fruit subglobose, 3-4(-5) cm diameter, fleshy, opens in 3 or 4 valves............................................................................................................................. C. tremula
Synopsis of species of Casearia in the Mexican Yucatán Peninsula (excluding C. randioides).
Casearia aculeata Jacq., Enumeratio Systematica Plantarum, quas in insulis Caribaeis 21. 1760. Type: Hispaniola.
Haiti: (no known extant authentic specimen), Jacquin anno 1757-58. Figure 3A, B.
Figure 3 Casearia in the Mexican Yucatán Peninsula. A-B) Casearia aculeata Jacq. (not vouchered). C-D) Casearia corymbosa Kunth (C. C. Ramírez D. s/n, CICY; D. J. Muñoz 61, CICY). E-F) Casearia laetioides (A. Rich.) Warb (E. G. Carnevali et al. 9281, CICY; F, not vouchered). G-H) Casearia subsessiliflora Lundell (G. Carnevali et al. 10329, CICY). (Photographs A-B, F. Alfredo Dorantes. C-D, F-H. Germán Carnevali).
Casearia sylvestris auct. non Sw. [sensuSosa et al. 1985, pro parte].
General Distribution. Mexico, Central America, the Antilles, and South America. Widespread in México; in the Mexican Yucatán Peninsula, only in Campeche and Quintana Roo (Figure 4). It has not been reported in the northwesternmost, driest part of the peninsula, within the limits of the state of Yucatán, and most likely it does not grow there.
Figure 4 Distribution map of Casearia in the Mexican Yucatán Peninsula. A) Casearia aculeata Jacq. B) Casearia corymbosa Kunth. C) Casearia l aetioides (A. Rich.) Warb. D) Casearia subsessiliflora Lundell. E) Casearia thamnia (L.) T. Samar. & M.H. Alford. E) Casearia tremula (Griseb.) Griseb. ex C. Wright. F) Casearia yucatanensis (Standl.). T. Samar. & M.H. Alford (Mayte R. Aguilar Canché).
Selected Specimens from the MYP. Mexico. Campeche. Mun. Escárcega, 58 km al SE de la desviación a Palizada camino a Escárcega, 18° 14’ 30” N, 91° 20’ 00” W, selva mediana perturbada, 4 August 1993, P. Simá et al. 1662 (CICY!!, MEXU!, MO!, NY). Quintana Roo: Mun. [Tulum], Ruinas de Cobá, 20° 29’ 30” N, 89° 47’ 55” W, [20 m snm], [selva mediana subperennifolia], 24 November 1980, C. Chan et al. 66 (CICY!!, XAL).
Ecology and vegetation. Tropical semideciduous forests.
Notes (morphology). Casearia aculeata is a shrub with spines in trunk and branches (not always present in herbarium specimens). This is a unique morphological character among the Casearia of the Mexican Yucatán Peninsula. (Figure 3A, B). It is often confused with the unarmed C. corymbosa, but in this later species the flowers are arranged on a peduncled corymb.
Common names. ts’iu che’, pak’aal che’ (Maya).
Casearia corymbosa Kunth, Nova Genera et Species Plantarum (quarto ed.) 5: 366-367. 1821[1823]. TYPE: Colombia: Crescit prope Mompox et Hondam, in ripa fluminis Magdalenae, floret aprili, Humboldt & Bonpland s.n.‒ Lectotype: Humboldt & Bonpland 1705, Colombia: Tolimá: Honda (P-Bonpl.!), designated by Nee, Fl. Veracruz 111: 15 (1999). Figure 3C, D.
Casearia nitida auct. non. Jacq. [sensuSosa et al. 1985]
General Distribution. Mexico, Central America, and South America (Colombia and Venezuela). Widespread in México. In the Mexican Yucatan Peninsula it has been found in the three states: Campeche, Quintana Roo, and Yucatán (Figure 4).
Selected specimens from the Mexican Yucatán Peninsula. Mexico. Campeche: Mun. Carmen, 35 km al SO de la desviación a Sabancuy, sobre la carretera Campeche-Ciudad del Carmen, 18° 51’ 15” N, 91° 25’ 40” W, duna costera, 21 September 1987, E. Cabrera & H. de Cabrera 14524 (CICY!!, MEXU!). Quintana Roo: [Mun. Tulum], entrance to Cobá, QR, 26 June 2002, P. Acevedo-Rodríguez & F. May Pat 12199 (CICY!!, MO!!, US!!). Yucatán: Mun. Valladolid, 7 km al SO de Valladolid, sobre el camino a Dzitnup, 20° 39’ 00” N, 88° 14’ 24” W, 24 m, acahual de selva mediana, 29 June 1985, Edgar Fco. Cabrera C. & H. de Cabrera 8752 (CICY!!, MEXU!, MO).
Ecology and vegetation. tropical dry and tropical semideciduous forests.
Notes (morphology). Shrubs or trees up to 20 meters tall with long (up to 10 cm long), oblanceolate leaves, on long branches, with medium sized flowers. Probably the most common of the Casearia with a tree habit in the Mexican Yucatán Peninsula.
Uses. Timber, ritual-religious, melliferous, medicinal.
Common names. ix iim che’ (Maya).
Casearia laetioides (A. Rich.) Northr., Mem. Torrey Bot. Club 12: 55. 1902.-Zuelania laetioides A. Rich. In: Sagra, Hist. Phys. Cuba, Pl. Vasc. 10: 89-90. 1845. TYPE: Cuba. R. de la Sagra s.n. Lectotype, designated by Samarakoon & Alford, Novon 27(1): 69. (P [bc] P00789605; isotypes, BM, K, P [bc] P00789606). Figure 3E, F.
=Laetia guidonia Sw., Prodr. [O. P. Swartz] 83. 1788.-Zuelania guidonia (Sw.) Britton & Millsp., Bahama Fl. (Britton & Millspaugh) 285. 1920.-Casearia guidonia (Sw.) Lundell, Wrightia 5: 41-42. 1974, nom. illeg., non C. guidonia Benth., J. Linn. Soc., Bot. 5(Suppl. 2): 89. 1861. TYPE: Jamaica. O. Swartz s.n.-Lectotype, designated by Kiger ex Sleumer [Sleumer, 1980: 280], S; isolectotypes, BM, SBT)
General Distribution. Mexico (Chiapas, Oaxaca, Querétaro, San Luis Potosí, Tamaulipas, Veracruz), Central America, northern South America (Colombia, Venezuela), and the West Indies. In the Mexican Yucatán Peninsula it is found in the three states. (Figure 4).
Selected specimens from the MYP. Mexico. Campeche: Mun. Candelaria, 2 km al N del poblado de Luna, a lo largo de la carretera Candelaria-Reforma Agraria, unos 24 km al N de Candelaria, 18° 18’ 47” N, 90° 53’ 56” W, selva mediana subperennifolia, 19 May 2000, G. Carnevali, J. L. Tapia M., F. May, L. Carrillo & G. Mondragón 6172 (ASU!!, CICY!!, MEXU!!, MO!!, UCAM!!). Quintana Roo: Mun. José María Morelos, Laguna de Chinchankanab, 19° 50’ 17” N, 88° 45’ 29” W, selva mediana, 5 m, 8 May 1982, C. Chan V. & M. Burgos C. 1375 (CICY!!, XAL). Yucatán: Mun. Kinchil, Área de Conservación de Kinchil (ACK), propiedad de Kekén , unos 8-9 km al WWS del desvío a Kinchil por la carretera Mérida-Celestún, unos 7.8 km al W de Bella Flor, unos 20-21 km al E del puente a la entrada de la Ría de Celestún, 20° 51’ 27.60” N, 90° 8’ 20.10” W, 7 m, selva mediana subperennifolia, 26 May 2022, G. Carnevali, J.L. Tapia, N. Raigoza & J. Muñoz 9281 (CICY!!, G!!, GH!!, MO!!, SEL!!, UADY!!).
Ecology and vegetation. Tropical semideciduous and evergreen forest.
Notes (morphology). Trees up to 30 m high, leaves long (up to 25 cm long), oblong, elliptic to oblong-lanceolate, along elongated branches, and medium sized flowers.
Common names. volador (Spanish), ta’may, sabak che’ (Maya).
Casearia subsessiliflora Lundell, Contributions from the University of Michigan Herbarium 6: 50. 1941. Type: Mexico: Quintana Roo, Cobá, in advanced forest east of the ruins, 5 July 1938, C.L. Lundell & Amelia A. Lundell 7824 (holotype: MICH!; isotypes: A, F, LL, US!!). Figure 3G, H.
Casearia sylvestris auct. non Sw. [sensuSosa et al. 1985, Villaseñor 2016].
General Distribution: Only in the Mexican Yucatán Peninsula; Quintana Roo and Yucatán. Endemic. (Figure 4)
Selected Specimens from the YPBP. Mexico. Quintana Roo: Mun. Lázaro Cárdenas, Rancho Santa María, approximately 10 km SE of El Eden Ecological Reserve, 21° 12’ 51” N, 87° 11’ 50” W, 6 m, 7 August 1999, G.P. Schultz 1304 (MO!!, UCR). Yucatán: Zona Arqueológica de Sayil, 20° 10’ 37” N, 89° 39’ 90” W, ca. 150 m, 21 November 2006, G. Carnevali et al. 7186 (CICY!, FLAS!!, GH!!, MEXU!!, MO!!, NY!!, TEX!!, UAMIZ!!, UCAM!!, XAL!!).
Ecology and vegetation. Tropical semideciduous forest.
Notes (morphology). This species was reinstated from the synonymy of Casearia sylvestris Sw. (Carnevali et al. 2008) from which it is amply distinct. Casearia subsessiliflora features medium (up to 8 cm long) ovate to ovateelliptic leaves, along elongated branches, and small-sized flowers.
Common names. unknown.
IUCN Conservation assessment. The IUCN Conservation assessment for this species was assigned by Carnevali Fernández-Concha et al. (2021) as VU (Vulnerable). A new assessment is possible now, including 20+ additional occurrences. Casearia subsessiliflora meets criterion B2a for an Endangered species (EN) but it does not meet at least two of the three additional conditions necessary for the assignment of this category (IUCN 2016). It is known from many localities (30) two states of the MYP, its Extent of Occurrence (EOA) is 35.451.282 km2 (exceeding the 20,000 km2 limit), the Area of Occupancy (AAO) is 96.000 km2. The habitat of the species is extremely fragmented under natural conditions due to anthropogenic activities, particularly agricultural developments, and human settlements. Few populations (less than 5) are known to occur at private or public protected areas and for this reasons we believe the appropriate status is Near Threatened (NT).
Casearia thamnia (L.) T. Samar. & M.H. Alford, Novon 27(1): 69. 2019. (15 Mar 2019) ‒Laetia thamnia L., Pl. Jamaic. Pug. 31. 1759. TYPE: Jamaica. Red hills above the Angels, P. Browne s.n. (holotype: LINN Cat. 680.1). Figure 5A, B.
Figure 5 Casearia (Salicaceae) in the Mexican Yucatán Peninsula. A-B) Casearia thamnia (L.) T. Samar. & M.H. Alford. (non vouchered). C-D) Casearia randioides Lundell. (G. Carnevali et al. 10361, CICY). E-F) Casearia tremula (Griseb.) Griseb. ex C. Wright. (vouchered). G-H) Casearia yucatanensis (Standl.) T. Samar. & M.H. Alford. (non vouchered). (photographs A-B. Alfredo Dorantes. C-D, G-H. Germán Carnevali. E-F. Alexis Valdez).
General Distribution. México, Central America, South America (Colombia), and the Antilles. In Mexico in Chiapas, Oaxaca, and Tabasco; in the Mexican Yucatán Peninsula in the three states. (Figure 4).
Selected Specimens from the YPBP. Mexico. Campeche: Mun. Champotón, Aguada de Chuiná, alrededor, km 40 carretera Escárcega a Champotón, 18° 57’ 47” N, 90° 42’ 50” W, selva mediana subperennifolia, 9 May 1983, E. Góngora 420 (CICY!!, XAL). Quintana Roo: Mun. Othón P. Blanco, 2 km al NE de Lázaro Cárdenas, vía corta Chetumal-Mérida, 4.5 km al W de la desviación a José María Morelos, por la carretera Chetumal-Carrillo Puerto, 18° 59’ 16” N, 88° 11’ 39” W, 12 m, selva baja caducifolia, 2 August 2006, J. L. Tapia-Muñoz, G. Carnevali, R. Duno & R. Balam Narváez 1916 (CICY!!, MEXU!!, MO!!, XAL!!). Yucatán: Mun. Tizimín, 5.5. km al E de la población de Moctezuma, 13 km antes de El Cuyo, por la carretera Colonia Yucatán-El Cuyo, 21° 26’ 19” N, 87° 41’ 21” W, 10 m, selva baja caducifolia, 19 May 2006, J. L. Tapia-Muñoz, C. Espadas M. & J.A. González I. 1847 (CICY!!, MEXU!!, MO!!, XAL!!).
Ecology and vegetation. This species ocurs in semi-evergreen forests, and several types of seasonally dry forests.
Notes (morphology). Small tree 3-7 m tall, with long (up to 11 cm long) elliptic to elliptic-lanceolate or obovate leaves, and medium size flowers.
Common names. zapote amarillo (Spanish), ch’aw che’, ix iim che’ (Maya).
Casearia tremula (Griseb.) Griseb. ex C. Wright, Anales de la Academia de Ciencias Médicas, Físicas y Naturales de la Habana, Revista Cientifica 5: 201. 1868.‒Zuelania tremula Griseb., Catalogus plantarum cubensium . . . 9. 1866. Type: Cuba or. et occ., Wright 2640 (GOET; isotypes: GH!, MO (BC: MO-279837/A:1916993), NY (fragm.).
General Distribution. Mexico, Central America, South America (Colombia, Venezuela), and the Antilles. In Mexico it ocurrs in the Pacific coast from Baja California, and Sinaloa to Chiapas, and in the Atlantic coast, only in the Mexican Yucatán Peninsula (Figure 4). This constitutes a new floristic record for the region (the specimen looks very much like the type specimen). Casearia tremula was not mentioned in any of the previous checklists of the region (e.g., Durán et al. 2000, Martínez et al. 2001, Carnevali Fernández-Concha et al. 2010). A second specimen in Quintana Roo, J.J. Ortíz 924 (MEXU!), also determined as C. tremula, is Casearia thamnia. Villaseñor (2016) cites the species from Campeche and Quintana Roo but no vouchers as support are provided.
Selected Specimens from the YPBP. Mexico. Campeche: Mun. Calakmul, Centenario, Laguna Silvituc, 18° 38’ 49” N, 90° 17’ 20” W, 30 m, 10 January 1999, E. Martínez S. 31907 (MEXU!).
Ecology and vegetation. Casearia tremula is only known from one specimen from the Laguna of Silvituc, the biggest lagoon in the state of Campeche, which is surrounded by medium to high subperennifolius forest. The label of the lone specimen describes an aquatic vegetation, and the plant as a herb one-meter-high with orange flower (clearly a mistake). The occurrence of this species in the region must be more accurately confirmed.
Notes (morphology). Small tree 3-7 m tall with long (up to 11 cm long), elliptic, elliptic-lanceolate or obovate blade, and medium sized flower.
Common names. unknown.
Casearia yucatanensis (Standl.) T. Samar. & M.H. Alford, Novon 27(1): 68. 2019 (15 Mar 2019) -Samyda yucatanensis Standl., Contr. U.S. Natl. Herb. 23: 842. 1923. TYPE: Mexico. Yucatán: Mérida, A. C. V. Schott 603 (holotype: US!!; isotypes: BM, F [3], MO!! [2]). Figure 5G, H.
General Distribution: Endemic to Mexico, only in the three states of the Mexican Yucatán Peninsula: Campeche, Quintana Roo, and Yucatán. (Figure 4).
Selected Specimens from the MYP. Mexico. Campeche: Mun. Campeche, 2 km al S de Samulá, 19° 48’ 30” N, 90° 32’ 30” W, 20 m, selva baja caducifolia, 4 February 2002, C. Gutiérrez B. 7313 (CICY!!, MEXU!, UCAM), same locality and date, (fl.), C. Gutiérrez B. 7315 (UCAM!). Quintana Roo: Mun. Solidaridad, Punta Laguna, 12.5 km al norte de Cobá, 20° 38’ 45” N, 87° 37’ 53” W, 20 m, selva mediana subperennifolia, 7 May 2004, R. Duno & M. Fierros 1963 (CICY!!, MEXU!!, XAL). Yucatán: Mun. Kinchil, Área de Conservación de Kinchil (ACK), propiedad de Kekén, unos 8-9 km al WWS del desvío a Kinchil por la carretera Mérida-Celestún, unos 7.8 km al W de Bella Flor, unos 20-21 km al E del puente a la entrada de la Ría de Celestún, Colecciones en los alrededores y márgenes de la carretera cerca de la caseta interior, unos 250-260 m al S del puente interior, unos 1,000-1,250 m al SSE de la entrada al Centro de Demostración de la ACK, unos 1,700-1,800 m al norte de la entrada al helipuerto de la ACK, 20° 49’ 46.74” N, 90° 8’ 26.58” W, 5-6 m, selva baja caducifolia, 7 April 2022, G. Carnevali, J.L. Tapia, D. Angulo & J. Muñoz 9136 (CICY!!, GH!!).
Ecology and vegetation. Casearia yucatanensis grows in seasonally dry low or medium statured forests, and secondary vegetation associated with these two types of vegetation. It is a common species and ocurrs between 5 to 300 m.
Notes (morphology). Casearia yucatanensis is a remarkable species. It is a shrub or small tree up to 5(-12) meters high with medium (up to 6 cm long), obovate to obovate-cuneate leaves in short branches (brachyblasts); the flowers are generally solitary, with many produced simultaneously in different branches of the tree, making for a very showy (yet fugaciously flowered) tree. Flowers are extremely fragrant, about 2 cm diameter, with white sepals.
Uses. It is often used as an ornamental plant for the beauty of its fowers and their fragrance, but more work is needed to improve on the lasting qualities of the flowers to make it become an important horticultural plant.
Common names. aguja de tórtola, jazmín de monte (Spanish), puuts’ mukuy, kaba pak’aal che’, xikin juj (Maya).
IUCN Conservation assessment. LC (Least concern) [DDATA: EOO = 88,788.035, AOO = 192.000, L = 48, L PNA= 5]. (Carnevali Fernández-Concha et al. 2021).
Casearia yucatanensis is known from many localities (48) in the three states of the MYP, its Extent of Occurrence (EOA) is of 88,788.035 km2 (far exceeding the 20,000 km2 limit) and the Area of Occupancy (AAO) is of 192.000 km2. Because of this last figure, it could have been assigned to the category of an Endangered species (EN) but it only meets one (B Criteria) of the three additional conditions. The habitat of the species is moderately fragmented under natural conditions but is being severely threatened by anthropogenic activities, particularly touristic and agricultural developments. However, the species is relatively common, apparently resilient, and locally abundant. Furthermore, several populations (more than 20) are known to occur in the Biosphere Reserve of Calakmul (Campeche) and in the Biosphere Reserves of Celestún and Ría Lagartos (Yucatán), as well as other state and private reserves or protected areas; for all reasons mentioned before, the species is best regarded as LC.
Supplementary material
Supplemental data for this article can be accessed here: https://doi.org/10.17129/botsci.3731
