# Synopsis and typification of New World members of the tribe Chiococceae (Rubiaceae, Dialypetalanthoideae)

Piero G Delprete

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## To cite this version:

Piero G Delprete. Synopsis and typification of New World members of the tribe Chiococceae (Rubiaceae, Dialypetalanthoideae). Journal of the Botanical Research Institute of Texas, 2025, 19 (4), pp.501-605. 10.17348/jbrit.v19.i4.1443. hal-05506741

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## SYNOPSIS AND TYPIFICATION OF NEW WORLD MEMBERS OF THE TRIBE CHIOCOCCEAE (RUBIACEAE, DIALYPETALANTHOIDEAE)

Piero G. Delprete

AMAP Lab, IRD, CNRS, CIRAD, INRA, Université de Montpellier, 34398 Montpellier, FRANCE

AMAP Lab, IRD, Herbier de Guyane, B.P. 90165, 97323 Cayenne, French Guiana, FRANCE

ORCID: http://orcid.org/0000-0001-5844-3945

Email: piero.delprete@ird.fr

ORCID. PGD: 0000-0001-5844-3945

### ABSTRACT

The tribe Chiococceae sensu Paudyal et al. (2018) is a monophyletic group of 30 genera and ca. 210 species. The tribe has amphi-Pacific geographical disjunction between the Neotropics and the West Pacific, with ca. 160 species present in the Greater Antilles, which represents its center of diversity, ca. 25 species present in Central and South America, and 28–30 species in the South Pacific (Philippines, New Caledonia, Marianas to Melanesia and Tonga Islands). The tribe has been shown to be monophyletic by numerous molecular phylogenetic studies. Members of the Chiococceae are highly variable in habit, ranging from subshrubs, erect or scandent shrubs, vines, treelets to tall trees. Their flowers are also highly variable in shape and dimensions, with corolla tubes ranging from ca. 3 mm long (e.g., *Erithalis*) to 27 cm long (e.g., *Osa*), with stamens inserted near or at the base of corolla tube or on the disc, and with spinulose pollen. Fruits of this tribe are also variable, as they can be capsular, drupaceous, or baccate, and the seeds can be flat, polygonal or globose, included in pyrenes or not. A synopsis of the Neotropical taxa of the tribe is here presented. The generic delimitations adopted in this synopsis follow those of Paudyal et al. (2018). The objective of this synopsis is to present a complete synonymy and typification of all the names involved, and to provide the identity of all the names of Neotropical taxa. A total of 83 lectotypifications, 20 neotypifications, and 1 epitypification, are here presented. The new combinations *Coutaportla campanilla* (DC.) Delprete, *Scolosanthus crucifer* ssp. *microphyllus* (Borhidi) Delprete, *Solenandra coriacea* (Poir.) Delprete, *Solenandra parviflora* (Bonpl.) Delprete, and the new species *Scolosanthus nipensis* Borhidi ex Delprete, are here published. In the Neotropics, 27 genera, 183 species, 8 subspecies, and 2 varieties, are here recognized in the tribe, of which 16 genera, 83 species, 5 subspecies, and 1 variety occur in Cuba.

**KEY WORDS:** Neotropics, Condamineae, Catesbaeeae, nomenclature, Rubiaceae, typification

### RESUMEN

La tribu Chiococceae sensu Paudyal et al. (2018) es un grupo monofilético de 30 géneros y ca. 210 especies. La tribu tiene una disyunción geográfica anfipacífica entre el Neotrópico y el Pacífico Occidental, con ca. 160 especies presentes en las Antillas Mayores, que representa su centro de diversidad, ca. 25 especies presentes en América Central y del Sur, y 28–30 especies en el Pacífico Sur (Filipinas, Nueva Caledonia, Marianas a Melanesia y las Islas Tonga). Se ha demostrado que la tribu es monofilética mediante numerosos estudios filogenéticos moleculares. Los miembros de Chiococceae son muy variables en hábito, desde subarbustos, arbustos erectos o escandentes, enredaderas, arbolitos hasta árboles altos. Sus flores también son muy variables en forma y dimensiones, con tubos de corola que van desde ca. 3 mm de largo (p. ej., *Erithalis*) a 27 cm de largo (p. ej., *Osa*), con estambres insertados cerca o en la base del tubo de la corola o en el disco, y con polen espinuloso. Los frutos de esta tribu también son variables, ya que pueden ser capsulares, drupáceos o bacados, y las semillas pueden ser planas, poligonales o gúbosas, incluidas en pirenos o no. Aquí se presenta una sinopsis de los taxones neotropicales de la tribu. Las delimitaciones genéricas adoptadas en esta sinopsis siguen las de Paudyal et al. (2018). El objetivo de esta sinopsis es presentar una sinonimia y tipificación completa de todos los nombres involucrados, y proporcionar la identidad de todos los nombres de los taxones neotropicales. Aquí se presenta un total de 83 lectotipificaciones, 20 neotipificaciones y 1 epitipificación. Se publican aquellas nuevas combinaciones *Coutaportla campanilla* (DC.) Delprete, *Scolosanthus crucifer* ssp. *microphyllus* (Borhidi) Delprete, *Solenandra coriacea* (Poir.) Delprete, *Solenandra parviflora* (Bonpl.) Delprete y la nueva especie *Scolosanthus nipensis* Borhidi ex Delprete. En el Neotrópico, se reconocen 27 géneros, 183 especies, 8 subespecies y 2 variedades de la tribu, de las cuales 16 géneros, 83 especies, 5 subespecies y 1 variedad se encuentran en Cuba.

**PALABRAS CLAVES:** Neotrópico, Condamineae, Catesbaeeae, nomenclatura, Rubiaceae, tipificación

The tribe Chiococceae sensu Paudyal et al. (2014, 2018) is a monophyletic group of 30 genera and about 210 species (Delprete & Motley 2003; Delprete 2004; Motley et al. 2005; Negron-Ortiz 2005; Borhidi 2007, 2008; Borhidi et al. 2009; Manns & Bremer 2010; Barrabé et al. 2011; Alejandro et al. 2014; Jardim et al. 2015; Franck

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et al. 2017; Paudyal et al. 2014, 2018). The tribe has amphi-Pacific geographical disjunction between the Neotropics and the West Pacific. Members of the Chiococceae are primarily distributed in the Neotropics, with a center of diversity in the Greater Antilles, where ca. 160 species are found; and ca. 25 species are distributed in Central and South America. Several species are widespread in the Neotropics and subtropical areas (e.g., *Chiococca alba* (L.) Hitchc., *Coutarea hexandra* (Jacq.) K. Schum.).

In the West Pacific, the Chiococceae are represented by *Bikkia* Reinward (1828:8), *Thiollierea* Montrouzier (1860:217), and *Badusa* A. Gray (1859:308). These three genera include a total of ca. 28–30 species, ranging from the Philippines, New Caledonia, Marianas to Melanesia and Tonga Islands (Motley et al. 2005; Manns et al. 2012; Govaerts et al. 2017; Barrabé et al. 2011; Paudyal et al. 2014, 2018; Barrabé & Fleurot 2021). The Pacific genera are not treated in the present synoptic treatment.

Motley et al. (2005) performed a molecular phylogenetic analysis of the *Catesbaeeae-Chiococceae* complex (CCC) using ITS and *trnL-F* sequences. They sampled 61 species, representing 24 of the 28 genera that they recognized in the CCC. The phylogenies obtained could not fully determine many relationships owing to poor branch support and polytomies. Their phylogenies confirmed the monophyly of *Catesbaea*, *Isidorea*, *Portlandia*, and *Scolosanthus*, and retrieved *Exostema*, *Solenandra*, *Bikkia*, and *Chiococca* as polyphyletic.

Manns and Bremer (2010), according to their molecular phylogenies, delimited the Chiococceae including *Strumpfia* with 26 other genera, supporting the conclusions of Bremer and Eriksson (2009). In that study, they tentatively included *Ceuthocarpus* and *Thogsennia* in the tribe, but did not include *Shaferocharis* Urb., although these three taxa were previously included in the group by Motley et al. (2005). They included *Eosanthe* in the Chiococceae, which was not included in the tribe by Motley et al. (2005).

Paudyal et al. (2014) studied the genetic divergences among the tribes of subfamily Dialpetalanthoideae (sensu Razafimandimbison & Rydin 2024, as “Cinchonoideae”), using *trnL-F* data, analyzed using maximum likelihood and Bayesian inference methods, and provided additional evidence for excluding *Strumpfia* from the Chiococceae, along with the striking morphological peculiarities, and included it in the monotypic tribe Strumpfieae. In *Strumpfia* the flowers are protogynous, buzz-pollinated, the anthers united into a synandrum, opening by a common apical pore, and the pollen grains have a verrucose exine. In the Chiococceae the flowers are not buzz-pollinated, the anthers are free from each other, and the pollen grains have a spinulose exine. In Paudyal et al. (2014), the tribe Chiococceae was re-delimited to include 29 genera, *Badusa*, *Bikkia*, *Catesbaea* (including *Phyllacanthus*), *Ceratopyxis*, *Ceuthocarpus*, *Chiococca* (including *Asemnantha*), *Coutaportla*, *Coutarea*, *Cubanola*, *Eosanthe*, *Erithalis*, *Exostema*, *Hintonia*, *Isidorea*, *Lorencea*, *Morierina*, *Nernstia*, *Osa*, *Phialanthus*, *Portlandia*, *Salzmannia*, *Schmidtottia*, *Scolosanthus*, *Shaferocharis*, *Siemensia*, *Solenandra*, *Thogsennia*, and *Thiollierea*.

Paudyal et al. (2018) performed the most comprehensive phylogenetic study of the Chiococceae using two nuclear (ETS, ITS) and two plastid (petD, *trnL-F*) datasets analyzed with Bayesian and Maximum Likelihood (ML) methods. The phylogenetic trees generated from the analyses of the combined dataset (all markers) resulted in the most fully resolved phylogeny. The majority rule consensus tree was used by Paudyal et al. (2018:fig. 3) for taxonomic decisions. That tree has four well-supported main clades, designated A to D. Each clade is discussed in detail below.

In Paudyal et al.'s (2018:fig. 3) analysis, clade A was strongly supported (BPP = 1.0, BS = 100) as sister to the remainder of Chiococceae. In this clade are positioned *Coutaportla* and *Lorencea*, genera endemic to Mexico and northern Central America. It should be noted that in the analyses using the combined dataset they were placed on a strongly supported clade, while in the analyses using only plastid data, these two genera were positioned on separate clades.

Clade B of Paudyal et al. (2018:fig. 3) included all the genera with capsular fruits and wind-dispersed seeds of the Chiococceae sensu Paudyal et al. (2014, 2018), namely *Coutarea*, *Exostema*, *Hintonia* and *Solenandra*. The monophyly of clade B was well supported (BPP = 0.93, BS = 62), and the monophyly of *Hintonia* was strongly supported, along with its close relationship with *Exostema sensu* McDowell (1996) and *Coutarea*. In the phylogenetic study of Paudyal et al. (2018:fig. 3), *Coutarea* and *Exostema* were not resolved as
monophyletic. Following the monophyletic groups detected within Clade B, Paudyal et al. (2018) recircum-
scribed three of these four genera (except *Hintonia*, which was maintained as traditionally delimited) and
named three additional small genera, *Coutareopsis* Paudyal & Delprete, *Motleyothamnus* Paudyal & Delprete,
and *Adolphoduckea* Paudyal & Delprete. In clade B, *Exostema*, as traditionally delimited, was confirmed to be
paraphyletic, and required most taxonomic changes. In the well-supported (BPP = 0.86, BS = 63) subclade B1
were retrieved the species of *Exostema* with axillary inflorescences. This result agrees with the morphology-
based delimitation of *E. sect.* *Exostema* of McDowell (1996). Because *E. caribaeum* (Jacq.) Roem. & Schult. is
the type of the genus, Paudyal et al. (2018) treated the species of this clade as *Exostema* s.str., a genus of 8
species that occur in Cuba and Hispaniola, with the exception of *E. caribaeum*, which is also present in other
Antillean islands, southern Florida, Mexico and Central America. The presence of axillary vs. terminal inflo-
rescences is a significant character that has been used to define many genera in the Rubiaceae, and axillary
inflorescence represents a strong morphological synapomorphy for *Exostema* s.str.

Still within clade B of Paudyal et al. (2018), a subset of the species traditionally positioned in *Exostema*
were retrieved on the strongly supported (BPP = 1) subclade B4, which are all the species with terminal inflo-
rescences, many short flowers per inflorescence, and (except for one species) represent the species of the “*E.
sect. Pitonia*” of McDowell (1996). Paudyal et al. (2018) merged all the species present on subclades B4a and
B4b into the expanded genus *Solenandra* s.l., and published the necessary new combinations. The monophyly of
*Solenandra* sensu Borhidi (2002) was retrieved in previous molecular studies (McDowell & Bremer 1998;
McDowell et al. 2003; Manns & Bremer 2010; Manns et al. 2012). *Solenandra*, as delimited by Paudyal et al.
(2018), is a genus of 22 species, characterized by terminal inflorescences, infundibular, white corollas, capsular
fruits basipetally septicidal, placentas linear, narrowly ellipsoid to lanceolate, and winged seeds acropetally,
centripetally or basipetally aligned. Paudyal et al. (2018) divided *Solenandra* s.l. into two sections: *S. sect.
*Solenandra* (12 spp.), with corollas 0.6–2.2(–3.0) cm long, turning pale yellow after anthesis, and *S. sect.
*Pitonia* (DC.) Paudyal & Delprete (10 spp.), with corollas 4–21 cm long, turning pink to maroon after
anthesis.

Greuter and Rankin-Rodríguez (2021, 2022a) interpreted the clades within Clade B of Paudyal et al. (2018: fig. 3) differently than Paudyal et al. (2018), and instead of recognizing the phylogenetically and morphologically well supported genera, as delimited by Paudyal et al. (2018), opted to treat the entire Clade B as the broadly expanded, morphologically diverse genus *Exostema*. Their wide circumscription submerged the long-established genera *Coutarea* and *Hintonia*, as well as Paudyal et al.’s (2018) recently recognized segregated genera into a very broadly delimited *Exostema*. Because the genus *Coutarea* is positioned within Clade B, Greuter and Rankin-Rodríguez (2021) published a formal proposal (2831) to conserve the name *Exostema* against *Coutarea*, as the latter has nomenclatural priority.

Delprete and Paudyal (2023) in a rebuttal to Greuter and Rankin-Rodríguez's (2021, 2022a) proposal expressed their strong disagreement with naming the entire Clade B of Paudyal et al. (2018) as a widely expanded, highly heteromorphic *Exostema*, and their preference to recognize of the phylogenetically and morphologically well-supported genera of clade B. Greuter and Rankin-Rodríguez (2021), in their proposal, argued that the use of the name *Exostema* for all the species of clade B would require fewer new combinations than using the name *Coutarea* for the whole clade. Several additional arguments presented by Greuter and Rankin-Rodríguez (2021, 2022a) to recognize their expanded delimitation of *Exostema* are supported by erroneous information, and are amply discussed and contrasted in Delprete and Paudyal (2023). Greuter and Rankin-Rodríguez (2021, 2022a) supposed that the wide morphological variation present in their broadly delimited *Exostema* is mainly due to shift in pollination syndromes, without taking into account the strong set of additional morphological features that characterize each subclade within Paudyal et al.'s Clade B: inflorescence position, flower merosity, corolla symmetry, ovary placentation, style morphology, among many other characters. The rebuttal of Delprete and Paudyal (2023) to Greuter and Rankin-Rodríguez's proposal, was summarized as follows: "(1) The wide expansion of *Exostema* and the proposal to reject *Coutarea* vs. *Exostema* proposed by Greuter and Rankin-Rodríguez (2021, 2022a) is causing and will continue to produce disruption
in nomenclatural stability of traditional and current usage of generic and specific names within the
Chiococcaceae; (2) The broad synonymization under *Exostema* s.l. proposed by Greuter and Rankin-Rodríguez
(2021, 2022a) entails the lumping of six genera (*Adolphoduckea*, *Coutarea*, *Coutareopsis*, *Hintonia*, *Motleyothamnus*,
*Solenandra*) under a broadly distributed, widely polymorphic genus, decreasing the value of the diagnostic
information of each monophyletic taxon present in Paudyal et al.'s (2018) Clade B and reduces the systematic and
morphological information of the species, as members of smaller monophyletic genera; (3) All the internal
clades of clade B retrieved in Paudyal & al.'s (2018) phylogenies are well- to strongly supported, and each of
them corresponds to a genus characterized by a unique set of morphological characters; (4) Paudyal & al.'s
(2018) genera have been accepted by numerous Rubiaceae specialists and managers of specialized websites;
(5) The broad circumscription of *Exostema* proposed by Greuter and Rankin-Rodríguez (2021, 2022a) does
not fulfill the principle of maximizing the ease of identification of the ca. 40 species included in such a highly
polymorphic genus, and therefore increases the difficulty of species identification by the botanical commu-
nity." Following the above considerations, which are in agreement with the basic principles of classification
adopted by the Angiosperm Phylogeny Group (1998), Delprete and Paudyal (2023) advised the Nomenclature
Committee to reject Greuter and Rankin-Rodríguez's (2021) proposal to conserve the generic name *Exostema*
against *Coutarea*. Whether the proposal to reject *Coutarea* vs. *Exostema* published by Greuter and Rankin-
Rodríguez (2021) is recommended or not, and whether those authors will eventually proceed in publishing
new combinations in either *Coutarea* or *Exostema*, to our knowledge, most Rubiaceae specialists will continue
to recognize the genera of Clade B as delimited by Paudyal et al. (2018), with the obvious disruption in nomen-
latural stability and in the traditional usage of generic names in this group.

Several examples of acceptance of the genera of Clade B as delimited by Paudyal et al. (2018) have been
recently published. Borhidi et al. (2018) in an article focusing on additions and corrections to their floristic
treatment of Cuban Rubiaceae (Borhidi et al. 2017), followed all generic delimitations proposed by Paudyal et al.
(2018), recognizing the genera *Solenandra sensu* Paudyal & Delprete, *Exostema sensu* Paudyal & Delprete,
*Adolphoduckea*, and *Motleyothamnus*. Torres-Montúfar et al. (2022) in a synopsis of the Chiococcaceae occurring
in Mexico, recognized *Coutarea*, *Exostema*, *Hintonia*, and *Solenandra* as delimited by Paudyal et al. (2018). Torres-
Montúfar et al. (2023) recognized the South-American genera *Adolphoduckea*, *Coutareopsis*, and *Motleyothamnus*
as delimited by Paudyal et al. (2018). The multi-authored internet site *Flora e Funga do Brasil* (2024) recognized
*Adolphoduckea* and *Coutarea* occurring in Brazil as delimited by Paudyal et al. (2018). The multi-authored
website *Plants of the World Online* (POWO, 2024) accepted all the genera of the Chiococcaceae as delimited by
Paudyal et al. (2018). The generic delimitations of Paudyal et al. (2018) in clade B, are followed in the present
treatment.

Clade C of Paudyal et al. (2018:fig. 3), also called the *Catesbaea-Portlandia* clade, was strongly supported (BPP = 1.0, BS = 100) as monophyletic and comprised *Catesbaea*, *Cubanola*, *Isidorea*, *Nernstia*, *Osa*, *Phyllacanthus*, and *Portlandia*. Based on a phylogenetic analysis using morphological characters, Delprete (1996) included these seven genera into an expanded *Catesbaeeae*, with eight other genera of Chiococcaceae, i.e., *Bikkia*, *Ceuthocarpus*, *Coutaportla*, *Coutarea*, *Hintonia*, *Schmidtottia*, *Siemensia* and *Thogsennia*. However, *Thogsennia* has not been included in any molecular phylogenetic studies, including that of Paudyal et al. (2018), because it is an extremely rare, monotypic genus from eastern Cuba, only known by a few old collections. The genera of clade C were already resolved as one clade in several previous studies (Rova et al. 2002; Motley et al. 2005; Robbrecht & Manen 2006; Manns & Bremer 2010; Manns et al. 2012). Aiello (1979) segregated *Osa* from *Hintonia*, and *Cubanola* and *Nernstia* (as *Cigarrilla*) from *Portlandia* based mainly on placenta and seed characters. She distinguished *Osa* from *Hintonia* as having large wingless seeds (vs. small winged seeds in *Osa*) with tuberculate testa (vs. reticulate) and persistent funicle (vs. non-persistent), long trumpet-shaped corollas (vs. funnelform) and large leaves with attenuate apex (vs. medium-sized leaves with acute to acuminate apex). Similarly, Aiello (1979) distinguished *Cubanola* from *Portlandia* in having reticulate to foveate seeds (vs. tuberculate in *Portlandia*) and non-persistent funicle (vs. persistent), loculicidally and septicidally dehiscent capsule (vs. basipetally loculicidally dehiscent), placenta circular in cross section (vs.
linear and adnate to septum), and chartaceous to subcoriaceous leaves (vs. coriaceous). *Nernstia* is distinguished from *Portlandia* in having colliculate seeds (vs. tuberculate) with acropetally imbricate arrangement and non-persistent funicle (vs. persistent) and large, spongy placenta (vs. linear, adnate to the septum). All Aiello's segregations are supported by the molecular phylogenies of Paudyal et al. (2018). The monotypic *Nernstia*, endemic to Mexico, and *Osa*, endemic to Central America, are resolved on a strongly supported clade (subclade C2), sister to the *Catesbaea-Portlandia-Isidorea* clade. *Cubanola*, a genus of two species endemic to Cuba and Hispaniola, was placed as sister to all the other genera of the clade. These relationships are similar to those retrieved in the molecular phylogenetic trees of Motley et al. (2005) and Manns & Bremer (2010).

*Catesbaea*, a genus of 17 species occurring in the Florida Keys, Bahamas, and the Antilles, in Paudyal et al. (2018) formed a strongly supported clade (BPP = 1.0, BS = 100) with *Phyllacanthus* Hook.f. nested in it, corroborating several previous molecular phylogenetic studies (Rova et al. 2002; Motley et al. 2005; Robbrecht & Manen 2006; Manns & Bremer 2010; Manns et al. 2012). *Phyllacanthus* was segregated from *Catesbaea* by Hooker (1871) because of its large, laterally flattened, triangular thorns and uniserial ovules. Aside from these characters, the two genera are morphologically similar. Although the analysis of Paudyal et al. (2018) using combined data resolved a different relationship, based on their *trnL-F* results and morphological similarity with *C. flaviflora* Urb., which has flowers almost identical to those of *P. grisebachianus* Hook.f., *Phyllacanthus* is here treated as a synonym of *Catesbaea*.

*Portlandia*, positioned in subclade C4, is a genus of six species endemic to Jamaica (Delprete & Motley, 2003). In Paudyal et al.'s (2018) study it formed a strongly supported monophyletic clade (BPP = 1.0, BS = 100), sister to the *Isidorea* clade. The sister relationship of *Portlandia* and *Isidorea* was already shown in previous phylogenetic studies (Delprete 1996; Rova et al. 2002; Delprete & Motley 2003; Motley et al. 2005; Robbrecht & Manen 2006; Manns & Bremer 2010).

*Isidorea*, retrieved in subclade C5 of Paudyal et al.'s (2018) study, is a genus of 13 species endemic to Cuba and Hispaniola. It was strongly supported to be monophyletic (BPP = 1.0, BS = 100). It differs from *Portlandia* in having stiff, pungent, coriaceous leaves and stipules divided at the base into two units, looking like four, apically pungent, stipules per node (Aiello, 1979). Paudyal et al. (2018) included ten *Isidorea* species in their study, six of which were not included in previous phylogenetic analyses. The *Isidorea* clade was divided into two subclades, one with the species from Cuba (subclade C5a), and the other with the species from Dominican Republic (subclade C5b).

Clade D of Paudyal et al. (2018) was strongly supported (BPP = 1.0, BS = 100) as monophyletic, and included 14 genera, namely *Badusa*, *Bikkia*, *Ceratopyxis*, *Ceuthocarpus*, *Chiococca*, *Eosanthe*, *Erithalis*, *Morierina*, *Phialanthus*, *Salzmannia*, *Schmidtottia*, *Scolosanthus*, *Siemensia* and *Thiollierea*. These genera were retrieved in five strongly supported subclades (subclades D1-D5). Several genera of this clade are endemic to Cuba (*Ceratopyxis*, *Ceuthocarpus*, *Eosanthe*, *Schmidtottia*, *Siemensia*), some others occur in Cuba and other Antilles (*Erithalis*, *Phialanthus*, *Scolosanthus*), two are widespread in the Neotropics (*Chiococca*, *Salzmannia*), and four occur in the western Pacific region (*Badusa*, *Bikkia*, *Morierina*, *Thiollierea*). Corolla shapes, sizes and fruit types vary greatly among members of this clade, showing no distinct pattern in the evolution of these characters and supporting the evolutionary plastic nature of them. Although clade D is strongly supported as monophyletic (BPP = 1.0, BS = 100), the relationships between the five subclades were not well supported in Maximum Likelihood analysis (ML; BS < 70), and these subclades formed an unresolved grade. The genera mentioned above are also grouped as a monophyletic alliance in most of the previous molecular studies (Rova et al. 2002; Motley et al. 2005; Robbrecht & Manen 2006; Manns & Bremer 2010; Manns et al. 2012); however, they did not form a monophyletic group in Bremer & Eriksson (2009).

In Paudyal et al.'s (2018) molecular analysis, the genera previously included in the *Chiococcae* by Hooker (1873a) are found on clade D, which divided into two separate clades: clade D5, with *Chiococca* (including *Asemnantha*), *Erithalis*, *Salzmannia* and *Scolosanthus*; and clade D2, with *Phialanthus*, *Ceratopyxis*, *Ceuthocarpus*, *Eosanthe* and *Schmidtottia* (the last four endemic to Cuba). The four genera from the western Pacific islands, *Badusa*, *Bikkia*, *Morierina* and *Thiollierea*, were found in two separate clades (subclades D1 and
D4). *Morierina* and *Thiollierea*, endemic to New Caledonia, formed a strongly supported monophyletic clade (BPP = 1.0, BS = 100; subclade D1) with the morphologically distinct *Morierina* nested in *Thiollierea*. *Morierina montana* Vieill. is a large tree with narrowly tubular corollas, found in forested area, whereas *Thiollierea* species are shrubs with large, colorful, campanulate flowers, growing in scrub coastal vegetation on ultrabasic soils (Motley et al. 2005). Bremer (1992) and Delprete (1996), due to the overall morphological similarity of *Morierina* and *Exostema*, e.g., corollas with long-narrow tube and reflexed lobes, in their phylogenetic analyses using morphological data retrieved the two genera as sister taxa. Motley et al. (2005) were the first to include *Morierina* in a molecular phylogenetic analysis and found it to be nested in the *Thiollierea* clade. *Thiollierea* was resurrected by Barrabé et al. (2011) to include ten species endemic to New Caledonia, which were previously placed in *Bikkia*. They amended *Thiollierea* mostly following the molecular analysis of Motley et al. (2005), further supported by the morphological data presented in Barrabé et al. (2011). The characters that they used to distinguish *Thiollierea* from *Bikkia* s.s. are: anthers that twist at anthesis (vs. not twisting in *Bikkia*), sheathing, truncate stipules (vs. free, acuminate), flat ovules (vs. globose), drooping inflorescence (vs. erect) and flat seeds (vs. angular or diamond-shaped). Despite acknowledging the fact that *Morierina* in molecular phylogenetic trees is nested in *Thiollierea*, Barrabé et al. (2011) refrained from adequately addressing its systematic position within *Thiollierea*, as shown by Motley et al. (2005). Paudyal et al. (2018) were unable to include in their study the second species of *Morierina*, *M. propinqua* Brongn. & Gris, which is probably extinct. However, they felt that *Morierina* is another example of extreme morphological variation in a single genus present in this tribe. Although only one species was included in their analysis, Paudyal et al. (2018) transferred both *Morierina* species to *Thiollierea*, and published the necessary new combinations.

The molecular phylogenies of Paudyal et al. (2018) resolved the four Cuban endemics *Ceuthocarpus*, *Ceratopyxis*, *Eosanthe* and *Schmidtottia* and the West Indian genus *Phialanthus* on the strongly supported subclade D2 (BPP = 1.0, BS = 92). This result supported the same relationships retrieved in the molecular studies of Rova et al. (2002), Motley et al. (2005), Bremer and Eriksson (2009), and Manns and Bremer (2010), although it contradicted the phylogenetic trees produced by Robbrecht and Manen (2006), in which *Eosanthe* was placed in a different position.

Urban (1923a) segregated *Schmidtottia* from *Portlandia* because of its terminal inflorescence (vs. lateral in *Portlandia*), sheathing, truncate stipules (vs. interpetiolar, triangular), septicidal capsules (vs. loculicidal) and ovate to obovate placenta (vs. linear). Paudyal et al. (2018) included seven *Schmidtottia* species in their phylogenetic study. Those species were found on a strongly supported (BPP = 1.0, BS = 100) clade along with *Ceuthocarpus involucratus* (Wernham) Aiello. Because *Ceuthocarpus* was nested within the *Schmidtottia* clade, Paudyal et al. (2018) returned *Ceuthocarpus* to *Schmidtottia*, in agreement with Alain (1959).

Paudyal et al. (2018) confirmed the placement of *Ceratopyxis* within the *Chiococeae*, another Cuban endemic monotypic genus, as sister to the *Phialanthus-Eosanthe* clade. *Ceratopyxis* has been positioned as sister to *Phialanthus* in most previous studies, except those produced by Manns & Bremer (2010), where *Schmidtottia* and *Phialanthus* were retrieved as sister genera, with no support value stated. *Phialanthus*, a genus of 22 species occurring in the Bahamas and the Greater and Lesser Antilles, was already positioned in the *Chiococeae* s.s. by Hooker (1873a). Bremer (1992) excluded it from the amended *Chiococeae* citing the presence of free filaments, but later Rova et al. (2002) showed that it is closely related to other members of *Chiococeae*. In Motley et al. (2005), *Phialanthus* was supported to be monophyletic.

Paudyal et al. (2018) included nine species of *Phialanthus* in their phylogenetic study, and in both combined and separate analyses using plastid and nuclear datasets, this genus was retrieved as non-monophyletic. Eight species of *Phialanthus* were found on one clade forming a trichotomy with *Eosanthe* and *Phialanthus hispaniolae* Alain & R.G.García as sister taxa in a strongly supported Subclade D2c (BPP = 1.0, BS = 100). Delprete (1999a), based on morphological observations, pointed out that *Eosanthe* is similar to *Phialanthus* because of the sheathing stipules, axillary inflorescence, persistent four-lobed calyx, filaments not connate to the corolla tube and the two-seeded indehiscent fruits. Delprete (1999a) also indicated that due to the resinous branches, thick coriaceous leaves, foliose calyx lobes, ridged corolla tube, and linear-oblong anthers,
already suggested by Motley et al. (2005). *Scolosanthus* is a genus of about 27 species occurring in the Bahamas and Greater and Lesser Antilles.

*Chiococca cubensis*, a species endemic to Cuba, in Paudyal et al.'s (2018) analysis was retrieved on the strongly supported (BPP = 1.0) subclade D5d, as a sister taxon to *Scolosanthus*, in the combined and nuclear phylogenies, while in those using the plastid dataset *C. cubensis* was nested within the *Scolosanthus* clade. In their analyses, the *Scolosanthus-C. cubensis* clade is sister to the *Salzmannia* clade. *Chiococca cubensis* is similar to *Salzmannia*, as delimited by Paudyal et al. (2018), by the tubular-subcylindrical corollas, from which it differs by the branches without resinous exudate (vs. with resinous exudate), cymose inflorescence (vs. subcapitate or small racemes), corollas deep purple-brown outside and yellow inside (vs. white, pale yellow, yellow orangish-yellow to greenish) and by being endemic to Cuba (vs. occurring in coastal cordillera of Venezuela and coastal Brazil). *Chiococca cubensis* differs from *Scolosanthus* by being a scandent or climbing shrub lacking thorns [vs. erect shrubs with bifurcate or trifurcate thorns (rarely unarmed) in *Scolosanthus*]. In addition, *C. cubensis* can be distinguished from most *Chiococca* species in having corollas tubular-subcylindrical, purple-brown outside and yellow inside (vs. campanulate to funnelform, white, cream-white to pale yellow throughout in *Chiococca*, with the exception of *C. rubriflora* Lundell, from southern Mexico and Guatemala, which has corollas red outside and orange inside). Hence, Paudyal et al. (2018) transferred *C. cubensis* to the new genus *Ramonadoxa* Paudyal & Delprete, along with the necessary new combination, and treated it as a sister genus of *Scolosanthus*.

In clade D of Paudyal et al. (2018), *Erithalis* was retrieved as monophyletic on the strongly supported subclade D5b (BPP = 1.0, BS = 98), corroborating previous phylogenetic analyses (Negron-Ortiz & Watson 2002, 2003; Motley et al. 2005; Manns & Bremer 2010).

In conclusion, Paudyal et al. (2018), according to the results of the molecular phylogenetic analyses, recognized the following Neotropical genera in the *Chiococceae*: *Adolphoduckea* Paudyal & Delprete (1 sp.), *Catesbaea* L. (including *Phyllacanthus* Hook.f., 17 spp.), *Ceratopyxis* Hook.f. (1 sp.), *Chiococca* P. Browne (22 spp.), *Coutaportla* Urb. (5 spp.), *Coutarea* Aubl. (2 spp.), *Coutareopsis* Paudyal & Delprete (3 spp.), *Cubanola* Aiello (2 spp.), *Eosanthe* Urb. (1 sp.), *Erithalis* (9 spp.), *Exostema* (Pers.) Rich. (8 spp.), *Hintonia* Bullock (3 spp.), *Isidorea* A. Rich. ex DC. (13 species), *Lorencea* Borhidi (1 sp.), *Motleyothamnus* Paudyal & Delprete (1 sp.), *Nernstia* Urb. (1 sp.), *Osa Aiello* (1 sp.), *Phialanthus* Griseb. (22 spp.), *Portlandia* P. Browne (6 spp.), *Ramonadoxa* Paudyal & Delprete (1 sp.), *Salzmannia* DC. (4 spp.), *Schmidtottia* Urb. (including *Ceuthocarpus* Aiello, 11 spp.), *Scolosanthus* Vahl (27 spp.), *Shaferocharis* Urb. (3 spp.), *Siemensia* Urb. (1 sp.), *Solenandra* Hook.f. s.l. (14 spp.), and *Thogsennia* Aiello 1 sp.). The Paleotropical genera included in the *Chiococceae* by Paudyal et al. (2018) are *Badusa* A. Gray (3 spp.), *Bikkia* Reinw. (11 spp.), and *Thiollierea* Montrouz. (including *Morierina* Vieill., 12 spp.), and are not addressed in the present synopsis.

Members of the *Chiococceae sensu* Paudyal et al. (2014, 2018) are highly variable in habit, ranging from subshrubs, erect or scandent shrubs, vines, treelets to tall trees. This group is also highly variable in flower shape and dimensions, having corolla tubes ranging from ca. 3 mm long (e.g., *Erithalis* P. Browne) to 27 cm long (e.g., *Osa Aiello*). This wide corolla variation is sometimes present even within the genus *Catesbaea* L. (including *Phyllacanthus* Hook.f.), as some of its species have large, funnel-shaped corollas with tubes 13–20 cm long (e.g., *C. spinosa* L.) and, at the other extreme, some species have small, campanulate corollas with tubes 5–7 mm long (e.g., *C. parviflora* Sw.). Fruits in this tribe are also highly variable, as they can be capsular, drupaceous, or baccate. Seeds can be flat, polygonal or globose, included in pyrenes or not, and winged or unwinged.

There is no single morphological synapomorphy to distinguish the *Chiococceae sensu* Paudyal et al. (2014, 2018), and only a combination of two homoplasious characters can be used to define this group: (1) stamens inserted near or at the base of corolla tube or on the disc and (2) presence of spinulose pollen (Huysmans et al. 2000; Motley et al. 2005). However, neither of these characters is peculiar to this tribe. Basal stamen insertion, although a rare character state in Rubiaceae, is also found in other groups in subfamily Dialypetalanthoideae (sensu Razafmandimbison & Rydin 2024), e.g., in the tribe Hamelieae, in *Chione* DC.
(tribe Chioneae; Razafimandimbison & Rydin 2024), and in *Neobertiera* Wernham (tribe Sipaneeae; Delprete 2015a, 2022). Other Rubiaceae taxa also have spinulose pollen, e.g., some species in the tribe Spermacoceae (Dessein et al. 2002), in subfamily Rubioideae (sensu Razafimandimbison & Rydin 2024).

*Placocarpa* J.D. Hook. is a monospecific genus endemic to Mexico, which remains with dubious position within the Rubiaceae, as it was included or excluded from the Chiococceae by different authors. Joseph Dalton Hooker (1873a:107-108) published *Placocarpa* Hook.f. with a sole species, *P. mexicana* Hook.f., and cited the material studied as "Mexici incola, prope Orizaba, a Broterio detecta." The collector of the material cited is the naturalist Matteo Botteri (1808-1877). Hooker positioned *Placocarpa* in the Chiococceae, where it was maintained by Robbbrecht (1988, 1994). Delprete (1996) based on a phylogenetic analysis using morphological characters, positioned it in the *Chiococceae-Catesbaeae-Exostema* complex, as a genus related to *Scolosanthus*. Huysmans et al. (1999), based on palynological characters, excluded *Placocarpa* from the *Chiococceae-Catesbaeae-Exostema* complex because of the reticulate-columellate exine, while the genera of the Chiococceae have microechinate pollen grains. The sheet of Botteri 913 at K, barcode K000432683, has a several pencil sketches by J.D. Hooker, depicting the foliaceous calyx lobes, corollas with cylindrical tube and four round lobes, stamens inserted near the corolla mouth, and ovary with two pendulous ovules. Borhidi (2006, 2012) treated *Placocarpa* in his two editions of *Rubiáceas de México*. In the first edition, Borhidi (2006) included it in the *Chiococceae*. In the second edition (Borhidi 2012), he did not suggest any systematic position, and described its fruits as splitting into two indehiscent cocci. *Placocarpa* has never been included in any molecular phylogenetic analysis. Because of pollen grains with reticulate-columellate exine (i.e., not microechinate), and the fruit splitting into two indehiscent cocci, I exclude it from the *Chiococceae* sensu Paudyal et al. (2014, 2018), and its systematic position within the Rubiaceae remains to be tested.

## MATERIAL AND METHODS

The generic delimitations this synopsis follow those of Paudyal et al. (2018). This study is based on examination of herbarium specimens, material preserved in 60% ethanol, and field observations. Herbarium specimens were studied either on site or through loans from the B, B-W, BBS, BM, BR, BRB, C, CAY, COL, F, FI, G, G-DC, HAC, HAJB, HAL, IAN, INPA, K, L, LINN-SM, M, MBM, MG, MO, MPU, NY, P, P-AD, P-BONPL, P-JJR, P-JU, P-LA, R, RB, U, UMP, US, and VEN herbaria. Digital images of herbarium specimens were analyzed through the following websites:

- Jstor Global Plants (https://plants.jstor.org/),

- Reflora - Brazilian Plants: Historic Rescue and Virtual Herbarium for Knowledge and Conservation of the Brazilian Flora (http://reflora.jbrj.gov.br/reflora/PrincipalUC/PrincipalUC.do?lingua=en),

- INCT - Herbario Virtual da Flora e Fungos (http://inct.splink.org.br/),

- JACQ (https://www.jacq.org/#),

- Smithsonian National Museum of Natural History (https://collections.nmnh.si.edu/search/botany/),

- NYBG C.V. Starr Virtual Herbarium (https://sweetgum.nybg.org/science/vh/),

- CJBG - Conservatoire et Jardin de la Ville de Genève (https://www.ville-ge.ch/musinfo/bd/cjb/chg/advanced.php?lang=fr),

- BoGART - The Inventory Database of the Botanical Garden of Berlin-Dahlem (https://ww2.bgbm.org/bogartdb/BogartPublic.asp),

- BR virtual herbarium Plantentuin Meise (https://www botanica collections.be/#/en/search/specimen),

- virtual herbarium of the Muséum National d'Histoire Naturelle of Paris (https://science.mnhn.fr/institution/mnhn/collection/p/item/search/form?lang=en_US),

- Type Database of the Herbarium of Göttingen (https://gwdu64.gwdg.de/pls/herbar/typen$.startup),

- Sweden's Virtual Herbarium (http://herbarium.emg.umu.se/index.html).

Barcode numbers of herbarium specimens, when available, are cited in square brackets after the herbarium code; when the barcode number is not available, the accession number, is cited preceded by "No." in square brackets.

Important information about the history of Cuban herbaria and their codes, collection numbers, accession numbers, and series numbers of specimens preserved in them is available in Regalado Gabancho et al. (2008). The HAC herbarium specimens do not have barcode numbers or accession numbers. Hence the HAC type specimens are here described in detail, in order to distinguish the type specimens of the same gathering
in HAC. The LS and SV herbaria have been integrated in HAC, with some duplicates in HAJB. The original herbaria of LS and SV is specified when citing type specimens, to unequivocally indicate a given specimen. Many botanical collectors who worked in Cuba did not use collection numbers. Specimens without collection number that were included in HAJB, were assigned HAJB accession numbers and barcode numbers.

Information about collecting localities in Cuba was obtained from the *Diccionario Geográfico de Cuba* (Comisión Nacional de Nombres Geográficos, 2000).

All specimens cited have been examined, unless indicated by "n.v." (not seen) after the herbarium code.

Information about collection dates and localities of specimens gathered by Martius in Brazil were obtained from the text and the maps available in Spix and Martius (1976). Pohl's collection dates and localities in Brazil were obtained by Pohl's (1976) travel diary.

## SYSTEMATIC TREATMENT

# ADOLPHODUCKEA

Poeppig (1841:31–32) published *Exostema maynense* Poepp. and provided a detailed description of the species. Among the numerous features described, he wrote that this species is a beautiful tree, to 60 feet tall (ca. 18 m tall), with very hard, reddish wood, persistent, ovate stipules, terminal, corymbose inflorescences, corolla tube 4.5 inches long (ca. 11.5 cm long), with tube distally infundibuliform and linear lobes, exserted stamens and style, obovate capsules with persistent calyx, and oblong, compressed, winged seeds.

In the phylogenetic study of Paudyal et al. (2018:fig. 3), *Exostema maynense* was retrieved on a strongly supported clade with *Coutarea hexandra*. These two species are similar in having terminal inflorescences and 6-merous flowers; the former differs from the latter by having cymose inflorescences (vs. simple or compound dichasia in *C. hexandra*), actinomorphic flowers (vs. zygomorphic), oblong corolla lobes (vs. ovate-deltoid), equal stamens (vs. unequal, sigmoidal), and capsules weakly laterally compressed (vs. strongly laterally compressed), occurring at 120–500(–1100) m from Ecuador to Bolivia (vs. from near sea level to 500 (–850) m, distributed throughout the Neotropics, from Mexico to Argentina). Based on these differences, Paudyal and Delprete transferred *Exostema maynense* to the monospecific genus *Adolphoduckea* Paudyal & Delprete, a name dedicated to Adolpho Ducke (1876–1959), the famous naturalist who worked in the Amazon Basin for most of his life, and collected and described hundreds of insects and plants.

*Adolphoduckea* can be distinguished from *Exostema sensu* Paudyal et al. (2018) by its terminal inflorescences (vs. axillary in *Exostema*), 6-merous flowers (vs. 4- or 5-merous), calyx lobes broad, long, with obtuse tip (vs. short, with acute tips), strongly flattened capsules (vs. slightly flattened), and by being a tree 10–30 tall, found in the lowland forests of the western Amazon basin (vs. shrubs or small trees, 1–6(–10) m tall, occurring in Cuba and Hispaniola, and *E. caribaeum*, distributed in other Antilles, Mexico, and Central America). For a complete description of the genus, see Paudyal et al. (2018:385).

*Adolphoduckea* is similar to *Solenandra sensu* Paudyal et al. (2018) in having terminal inflorescence and narrowly cylindrical corolla tubes; the former differs from the latter in having 6-merous flowers (vs. 5-merous), laterally compressed capsules (vs. not laterally compressed, terete), and seeds acrobasipetally arranged (vs. acropetally, centripetally or basipetally arranged), and by being a large tree growing in the western Amazon Basin (vs. subshrubs, shrubs or trees ranging from Mexico and Central America, to the Greater and Lesser Antilles).

*Adolphoduckea* is also similar to *Motleyothamnus* Paudyal & Delprete in having terminal inflorescence, laterally compressed capsules and acrobasipetal seed arrangement; the former differs from the latter in having 6-merous flowers (vs. 5-merous), corollas that turn pink to maroon with age (vs. turning pale yellow), and by being a tall tree that grows in the forests in lowland or at low elevations of the western Amazon Basin (vs. shrub growing in open places and shrublands of the Andes, at 1000–2800 m elevation).

*Adolphoduckea* is a monospecific genus, endemic to Amazonian lowlands of Ecuador, Peru, and Bolivia, occurring at 120–500(–1100) m elevation
**Adolphoduckea** Paudyal & Delprete, Bot. J. Linn. Soc. 187:384. 2018. **TYPE:** *Adolphoduckea maynense* (Poepp.) Paudyal & Delprete

1. **Adolphoduckea maynense** (Poepp.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:385. 2018. *Exostema maynense* Poepp. (as 'Exostemma'), Nov. Gen. Sp. Pl. 3(3-4):31, tab. 237. 1841; Delprete in Andersson, Fl. Ecuador 62:51-53, f. 17. 1999 (as *Exostema maynense*). **TYPE:** PERU. Loreto: "crescit in sylvis primaevis provinciae Maynas et Yurimaguas," 1831, E.F. *Poeppig* 2392 (LECTOTYPE) (Delprete 1999b:51); NY [00077350]; ISOLECTOTYPES: W-Rchb. [No. 1189-0294658], W [No. W0049010]).

**Distribution.**—Ecuador, Peru, and Bolivia.

## CATESBAEA

Linnaeus (1753:109) described *Catesbaea spinosa* L., and dedicated the genus name to Mark Catesby (1683-1749), the famous English naturalist, who studied plants and animals (mostly birds) in North America and the West Indies. *Catesbaea* was historically treated as a member of the tribes Gardenieae (e.g., Candolle, 1830; Baillon, 1880, 1881) or *Catesbaeae* (e.g., Hooker, 1873a; Verdcourt, 1958), due to its multi-seeded, succulent fruits. Robbbrecht and Puff (1986) excluded it from Gardenieae, and Robbbrecht (1988) listed it among the "genera incerte sedis." Delprete (1996) included it in the *Portlandia* group, in which it is an eccentric genus. Huysmans et al. (1999) confirmed its position within the *Catesbaeae-Chiococcaceae-Exostema* complex because of its spinulose pollen. Its position in the Chiococcaceae and its monophyly (including *Phyllacanthus*) was shown by Paudyal et al. (2014, 2018). *Catesbaea* has never been the subject of a monographic treatment. It displays a wide variation in shape and size of the corollas, the smallest are found on *C. parviflora* Sw., which has corolla tubes 5-7 mm long, and the largest ones in *C. spinosa* L., which has corollas tubes 13-20 cm long. Liogier (1962, 1995) published keys to the nine species occurring in Cuba (1962), and to the seven species occurring in Hispaniola (1995). Borhidi (2017) in *Rubiáceas de Cuba*, stated that *Catesbaea* is genus of about 30 species, 9 of them occurring in Cuba. In the present study, it is recognized as a genus of 17 species, 10 of which occurring in Cuba, of shrubs, with acicular thorns at nodes, and baccate, multi-seeded fruits, ranging from southern Florida, the Bahama Archipelago, to the Greater and Lesser Antilles.

**Catesbaea** L., Sp. Pl. 109. 1 May 1753; Standley, N. Amer. Fl. 32(3):209-214. 1934; Liogier, Fl. Cuba 5:68-70. 1962; Correll & Correll, Fl. Bahama Arch.: 1380-1383. 1982; Howard, Fl. Lesser Antill. 6(3):396-397. 1989; Liogier, Fl. España 7:225-230. 1995; Borhidi et al., *Rubiáceas Cuba* 59-64. 2017; Borhidi et al., *Acta Bot. Hung.* 60(3-4):292-294. 2018. **TYPE:** *Catesbaea spinosa* L.

*Catesbya Cothenius, orth. var.*, Disp. 6. Jan-May 1790.

*Phyllacanthus Hook. f.*, Hook. Icon. Pl. 11:77. 1871. [*Phyllacantha Hook. f.* in Bentham & J.C. Hooker, Gen. Pl. 2:78. 1873]. **TYPE:** *Phyllacanthus grisebachianus* Hook. f.

*Echinodendrum A. Rich.* in R. de la Sagra, Hist. Fis. Cuba 11:18. 1850. **TYPE:** *Echinodendrum campanulatum* (Sagra ex DC.) A. Rich. [= *Catesbaea campanulata* Sagra ex DC., Prodr. 4:401. 1830]

1. **Catesbaea ekmaniana** Urb., Ark. Bot. 20A(5):54. 1926. **TYPE:** HAITI: Massif de la Selle, Monte Cabaio, Roberjot, 1700 m, 23 Aug 1924, E.L. *Ekman H-1587* (LECTOTYPE, here designated: S [No. S10-12160]; ISOLECTOTYPES: F [No. 710735 (frag. ex US)], S [No. S-R-7815], US [00138219]).

**Distribution.**—Haiti (Massif de la Selle).

**Notes.**—Urban (1926:54-55), in the protologue of *Catesbaea ekmaniana* Urb., cited the material studied as "Massif de la Selle in faucibus Morne Cabaio prope Roberjot rarissima et parce 1,700 m alt., m. Aug flor.: [Ekman] n. H-1587," without citing the herbarium of deposit. The original material at B was destroyed during WWII. In S there is a specimen, with accession number S10-12160, that has a label with the annotation "Catesbaea Ekmaniana Urb." handwritten by Urban. The specimen consists of a ramified branch with numerous leaves and a small branch with a flower in anthesis, and is here designated as the lectotype of this name.

2. *Catesbaea flaviflora* Urb., Symb. Antill. 9:157. 1923. **TYPE:** CUBA: Santiago de Cuba, in declivibus calcareis, 25 Sep 1916, E.L. *Ekman 7731* (LECTOTYPE, here designated: S [No. S-R-7824]; ISOLECTOTYPE: K [K000173530]).

**Distribution.**—Cuba (Santiago de Cuba, Holguín).

**Notes.**—Urban (1923a:157), in the protologue of *Catesbaea flaviflora* Urb., cited the gathering *Ekman 7731*, without citing the herbarium of deposit. The original material at B was destroyed during WWII. Borhidi et al. (2017:60; 2018:293) cited the type of this name as "[...] *Ekman 7731*; Holotipo: B†; lectotipo: S!" According
to the *International Code of Nomenclature for algae, fungi, and plants* (Turland et al. 2018; hereafter the *Code*), starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al.’s (2017, 2018) lectotype citations are not valid.

In S there is a specimen of *Ekman* 7731, with accession number S-R-7824, which has a label with the annotation “Catesbaea flaviflora Urb.” handwritten by Urban. This specimen consists of a ramified branch with numerous long thorns subtended by minute leaves, without flowers. This specimen is here designated as the lectotype of *Catesbaea flaviflora*.

In K there is a specimen of *Ekman* 7731, with barcode K000173530, which has a label with annotation “Catesbaea flaviflora Urb. [...] , det. I. Urban [...]” handwritten by an unknown author. It consists of a branch with pairs of long thorns at each node, subtended by minute leaves, with numerous flowers in anthesis. This specimen is an isotype.

**3. Catesbaea foliosa** Millsp., Publ. Field Mus., Bot. 2:312. 1909. **TYPE:** BAHAMAS: Caicos Islands, West Caicos, 20 Dec 1907, P. Wilson 7761 (HOLOTYPE: F [No. 221880]).

*Distribution*.—Bahamas, Cayman Islands.

**4. Catesbaea fuertesii** Urb., Symb. Antill. 7:402. 1912. **TYPE:** DOMINICAN REPUBLIC. BARAHONA: “in sylvis apertis,” Nov. 1910, M. *Fuertes* 641 (*NEOTYPE*, **here designated**: GH [00058995]; *ISONEOTYPES*: A [00105428], BM n.v., F [No. 768291], M [M-0187158], MO [No. 707104, barcode MO-03687], NY [00115054], US [No. 658445]).

*Distribution*.—Dominican Republic.

*Notes*.—Urban (1912:402–403), in the protologues of *Catesbaea fuertesii* Urb., cited the gathering *Fuertes 641*, without indicating the herbarium of deposit. The original material at B was destroyed during WWII. Four specimens of this gathering were found in A, GH, M, and NY, all of them without proof that they were studied by Urban. Hence a neotype needs to be designated. The GH specimen, with barcode 00058995, consists of a much-ramified branch, with numerous leaves subtended by acicular thorns, and numerous flowers in anthesis. This specimen is here designated as the neotype of this name.

**5. Catesbaea gamboana** Urb., Symb. Antill. 9:524. 1928. **TYPE:** CUBA. LAS TUNAS: Gamboa, in moist forest, 26 Aug 1922, E.L. *Ekman* 14950 (*LECTOTYPE*, **here designated**: S [No. S-R-7823]; *ISOLECTOTYPES*: F [No. 604767], G [G00439777], NY [00115055]).

*Distribution*.—Cuba (Las Tunas, Granma).

*Notes*.—Urban (1928:524), in the protologue of *Catesbaea gamboana* Urb., cited the gathering *Ekman* 14950, without indicating the herbarium of deposit. The original material at B was destroyed during WWII. Borhidi et al. (2017:60; 2018:293) cited the type of this name as “*Ekman* 14950; Holotipo: B†; lectotipo: S!” According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al. (2017:60; 2018:293) lectotype designations are not valid.

A specimen in S, with accession number S-R-7823, has a label with the annotation “Catesbaea gamboana Urb. (typus)” handwritten by Urban. The specimen consists of two ramified branches with numerous acicular thorns subtended by pairs of minute leaves. This specimen is here designated as the lectotype of this name.

**6. Catesbaea glabra** Urb., Symb. Antill. 7:401. 1912. **TYPE:** DOMINICAN REPUBLIC. SANTIAGO: Llanos de Rafael, 200 m, 12 May 1887, H.F.A. *Eggers* 1912 (*NEOTYPE*, **here designated**: BM [000081474]; *ISONEOTYPES*: F [No. 776379 (frag.)], G [3 sheets, G00423003, G00423004, G00423005]).

*Distribution*.—Haiti, Dominican Republic.

*Notes*.—Urban (1912:401), in the protologue of *Catesbaea glabra* Urb., cited several syntypes from Haiti (*Picarda* 80; *Ehrenberg* 97; *Picarda* 1341b; *Picarda* 335; *Picarda* 853), and Dominican Republic (*Rob. Schomburgk* 22; *Rob. Schomburgk* 118; *Eggers* 1912). The original material at B was destroyed during WWII. A specimen in BM with barcode 000081474, has a label with the heading “Eggers: Flora Indiae Occid. Exs.” and the annotations “[Nr.] 1912, Rubiaceae sp. [Santo Domingo] in fruticetis. Llanos de Rafael, 200 m, 12.V.1887” handwritten by an unknown author. The specimen consists of a densely ramose branch with three acicular thorns at each node, and is here designated as the neotype of *C. glabra*.
**7. Catesbaea grayi** Griseb., Pl. Wright. 2:503. 1862 [preprint of Mem. Amer. Acad. Arts, n.s. 8(2):503. 1863].
**TYPE:** CUBA. HOLGUÍN: Monte Verde, "frutex orgyalis, baccis albis, in sylvis at Monte Verde," s.d. [1860-1864], C. Wright 371 (LECTOTYPE, here designated: GOET [GOET010202]; ISOLETOTYPES: BR [000000557709], G [4 sheets, G00389798, G00389799, G00389800, G00389801], GH n.v., GOET [GOET010203], HAC [ex SV No. 951], HAC, K [2 sheets, K000173531, K000173532], MO [No. 2091978], P [P00559101], PH [0001648], S [No. S-R-7822, S10-12237], UC n.v., YU [YU.066110])].

*Distribution.*—Cuba (Las Tunas, Granma, Guantánamo, Holguín, Santa Clara).

*Notes.*—Grisebach (1862:503) in the protologue of *Catesbaea grayi* Griseb., cited the gathering Wright 371, without citing the herbarium of deposit. Borhidi et al. (2017:61; 2018:293) cited the type of *Catesbaea grayi* as "Tipo: [...] Wright 371; Holotipo: GOET; isotipo: GH!" According to the *Code* (Turland et al. 2018), starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:61; 2018:293) type citations cannot be treated as an inadvertent lectotype designation.

According to Stafleu and Cowan (1979:1007) most of Grisebach's types are in GOET. In GOET there are two specimens of Wright 371 annotated with this name. The specimen with barcode GOET010202, has a label with the annotation "C. Grayi m." handwritten by Grisebach. The specimen consists of two branches with numerous leaves, and one of them with a few flowers in anthesis, and is here designated as the lectotype of *Catesbaea grayi*.

The GOET specimen with barcode GOET010203, has a label with the annotation "C. Grayi Gr." handwritten by Grisebach. The specimen consists of two sterile branches, and is an isotype.

**8. Catesbaea holacantha** C. Wright ex Griseb., Cat. Pl. Cub. 122. 1866. **TYPE:** CUBA: "Cuba Occ., pr. Mayari abajo," s.d. [1860-1864], C. Wright 2657 (HOLOTYPE: GOET [GOET010204]; ISOTYPES: BM [000081495 (specimen on right side of the sheet)], G [2 sheets, G00389796, G00389797], GH [00058989], HAC [ex LS], HAC, K [K000173533], MO [No. 2091979], NY [00115056], P [P00559100], S [No. S-R-7821], UC n.v., YU [YU.001726]).

*Distribution.*—Cuba (Las Tunas, Holguín, Granma, Guantánamo, Camagüey, Santi Spiritus, Villa Clara).

*Notes.*—Grisebach (1866:122) in the protologue of *Catesbaea holacantha* Wright ex Griseb., cited the gathering Wright 2657, without citing the herbarium of deposit. According to Stafleu and Cowan (1979:1007), the GOET herbarium contains most of his types. Borhidi et al. (2017:61; 2018:293) stated that the holotype of this name is at GOET. In GOET there is a single specimen of Wright 2657, barcode GOET010204, which has a label with the annotation "Catebaea holocantha" handwritten by Grisebach. The specimen consists of a much ramified branch, with dense sturdy thorns, subtended by minute leaves, and with numerous flowers in anthesis. This specimen is the holotype of this name.

In BM there is a sheet, with barcode 000081495, which has two different gatherings affixed on it. This barcode is assigned to the specimen on the left side of the sheet. On the right side of the sheet there is a specimen that has a label with the heading "Plantae Cubenses Wrightianae," the number "2657" handwritten by an unknown author, and the printed annotation "Coll. C. Wright, 1860-1864." The specimen on the right side of the sheet, to which it should be assigned a different barcode, is an isotype of *Catesbaea holacantha*.

Borhidi et al. (2017:61; 2018:293) spelled the name of this species "*Catesbaea holoacantha* C. Wr. ex Griseb.,” but the specific epithet was published as “*holacantha*” in the original publication, and it should be kept as such.

**9. Catesbaea longispina** A. Rich., in Sagra, Hist. Fis. Cuba, Bot. 11:12. 1850. **TYPE:** CUBA: ["Jagua, *De la Ossa* s.n." but the specimen label says "legit Ramon de la Sagra, Cienfuegos, 873"] Cienfuegos, s.d., *R. de la Sagra* 873 (HOLOTYPE: P [P00559104]; ISOTYPES: F [No. 972513 (frag, ex P), No. 630955 (frag, ex P)]).

*Distribution.*—Cuba (Cienfuegos, Ciego del Avila, Camagüey, Las Tunas, Holguín, Granma, Santiago de Cuba).

*Notes.*—Achille Richard (in Sagra 1850:12) in the protologue of *Catesbaea longispina* A. Rich., cited the material studied as "Crescit prope *Jagua*, ubi detexit clar. *De la Ossa*." The attribution of the gathering to De la Ossa is certainly an error because on the label of the specimen in P it is written that it was collected by Ramon de la Sagra. The holotype specimen of *C. longispina* has a label with the heading "Herbarium Richard" handwritten in red ink, the annotation "Catesbaea longispina Nob. Sp. nova" handwritten by Achille Richard, and the annotations "Cuba, legit Ramon de la Sagra" handwritten in red ink. Above that label is affixed another
label with the annotation "Cienfuegos. 873" handwritten by Ramon de la Sagra. The specimen consists of three branches with long, narrow thorns subtended by minute leaves, with several ovaries with the corollas fallen off. On the sheet is also affixed a single corolla in anthesis.

10. *Catesbaea macrantha* A. Rich., in R. de la Sagra, Hist. Fis. Cuba, Bot. 11:12, tab. 47. 1850. *TYPE:* CUBA. La Habana: Bacuranao, "Crescit in insula Cuba, prope Potrero de Barreto, Bacuranao," s.d., *R. de la Sagra s.n.* (HOLOTYPE: P [P00559106]).

*Non Catesbaea macrantha* C. Wright, Anales Acad. Ci. Méd. Fis. Nat. Habana 6:99. 1869 [= *Catesbaea spinosa* L.].

**Distribution.**—Cuba (La Habana, Cienfuegos, Sancti Spiritus, Las Tunas).

*Notes*.—Achille Richard (in Sagra 1850:12), in the protologue of *Catesbaea macrantha* A. Rich., cited the material studied as "*Protrero de Barreto* [sic!], i.e. Potrero], *Bacuranao* (*Ramon de la Sagra*)." In P there is one specimen, with barcode P00559106, which has a label with the heading "Herbarium Richard" and the bottom annotation "Scripsit A. Richard," both handwritten in red ink. The label also has the annotations "Catesbaea macrantha Nob. C. latifolia" handwritten by Achille Richard. The specimen consists of two branches with pairs of acicular spines at each nodes, numerous small leaves, and five large flowers in anthesis. This specimen is the holotype of *C. macrantha* A. Rich.

11. *Catesbaea melanocarpa* Krug. & Urb., in Urban, Symb. Antill. 1:427. 1899. *TYPE:* PUERTO RICO: Guanica, La Plata, 17 Jan 1886, P. *Sintenis* 3786 (*LECTOTYPE*, here designated: BM [000081494]; *ISONEOTYPES*: US [No. 091360, barcode 00476949], US [No. 958600, without barcode]).

**Distribution.**—Puerto Rico, Lesser Antilles.

*Notes*.—Krug and Urban (in Urban, 1899:427-428) in the protologue of *Catesbaea melanocarpa* Krug. & Urb., cited several gatherings from Puerto Rico (*Sintenis* 3362, *Sintenis* 3542), St. Croix (*Eggers s.n*.), and Antigua (*Barber & Tillson* 70, *Wullschlägel* 262). The original material studied by Krug and Urban in B was destroyed during WWII. I was able to trace several specimens of the gatherings cited by Krug and Urban, although none of them are annotated by the authors. A specimen in BM, with barcode 000081494, has a label with heading "P. SINTENIS: PLANTAE PORTORICENSIS," the annotation "3786. Catesbaea parviflora Sw. Guanica, ad La Plata, 17.1.1886" handwritten by an unknown author, and the printed annotation "det. I. Urban." This specimen consists of three branches, all with long, acicular thorns. Two branches have a few leaves and several fruits. The third branch has leaves at all nodes and several fruits. This specimen is here designated as the lectotype of *C. melanocarpa*.

Another specimen of *Sintenis* 3786 is in US, with barcode 00476949. It has the same label and the same annotations as the BM specimen. It consists of three branches, all of them with long, acicular thorns, a few leaves, and a few fruits. This specimen is an isolectotype.

Howard (1989:396-397) cited *Catesbaea melanocarpa* as present in Puerto Rico and St. Croix, and reported that various annotations on herbarium specimens suggested that it might be a synonym of *Catesbaea parviflora* Sw. However, I differ from his opinion, and I prefer to recognize it as a distinct species.

12. *Catesbaea microcarpa* Urb., Symb. Antill. 5:509. 1908. *TYPE:* HAITI: "Haiti in sylvis apricis ad Petite Rivière de Bajonnais, 100 m," Oct 1905, W. *Buch 1031* (*NEOTYPE*, here designated: GH [00058991]).

**Distribution.**—Haiti (Gonave Island).

*Notes*.—Urban (1908:509), in the protologue of *Catesbaea microcarpa* Urb., cited the gathering *Buch 1031*, without citing the herbarium of deposit. The original material at B was destroyed during WWII. A search in numerous herbaria retrieved a sole specimen of *Buch 1031* in GH, with barcode 00058991. There is no evidence on the sheet that it was studied by Urban. It consists of a small branch with pairs of small acicular thorns subtended by pairs of small leaves. No flowers or fruits are present. This specimen is here designated as the neotype of *C. microcarpa*.

13. *Catesbaea nana* Greenm., in R. Combs, Trans. Acad. Sci. St. Louis 7:428. 1897. *TYPE:* CUBA. Santa Clara: Distr. Cienfuegos, Cieneguita, on sandy arid savannah, 10 Aug 1895, *R. Combs 406* (HOLOTYPE: GH [00058992]; *ISOTYPES*: F [2 sheets, No. 358209, No. 17029], G [G00389797], K [K000173541], MO [No. 2091980], NY [00115058], P [P00559102], US [00138221] (frag. ex NY]).

**Distribution.**—Cuba (Cienfuegos, Villa Clara, Ciego del Ávila, Camagüey).
**Notes.**—Greenman (in Combs 1897:428–429), in the protologue of *Catesbaea nana* Greenm., cited the material studied as “Collected by Robert Combs in dry poor soil at Cieneguita, Cuba, August 10, 1895, no. 406. A species related to *C. parviflora* Sw.—Plate XXXV,” without citing the herbarium of deposit. According to Stafleu and Cowan (1979:364–365), Greenman’s herbarium and types are at “GH, duplicates A, B, DS, E, F, L, MO, MONT, NY, US, W.” In GH there is a specimen of *Combs 406*, barcode 00058992, with a label that has the annotations “Catesbaea nana, Greenman, n. sp.” handwritten by Greenman. This specimen is the holotype of this name.

Borhidi et al. (2017:62; 2018:293) erroneously cited the type of *Catesbaea nana* as “Cuba, Distr. Cienfuegos, Cieneguita, *Combs* 271 (B†, F).”

**13. Catesbaea parviflora** Sw., Prodr. 30. 1788. *Echinodendrum parviflorum* (Sw.) A. Rich., in R. de la Sagra, Hist. Fis. Cuba, Bot. 11:18. 1855. TYPE: JAMAICA: without locality, s.d., *Swartz s.n.* (LECTOTYPE, here designated: S [No. S-R-950]; ISOLECTOTYPES: B-W [B-W 02800 -01 0], LD [No. 1250417], SBT [SBT13328]).

**Distribution.**—USA (Southern Florida), Bahamas, Cuba (La Habana, Mayabeque, Artemisa, Matanzas), Jamaica, Puerto Rico, Lesser Antilles.

**Notes.**—Swartz (1788:30), along with the succinct description of *Catesbaea parviflora* Sw., cited the collection locality as “Jamaica” without citing the herbarium of deposit. According to Stafleu and Cowan (1986: 117) “The Swartz herbarium constitutes, with the Alströlmer herbarium, the basis of [...] S. [...] The West-Indian collections (1784–1786) at S, however, are not complete.” Swartz specimens are present in many herbaria, and, according to Stafleu and Cowan (1986:117), “Swartz was very liberal with his specimens and Swartz types will be found for instance in BM and LD.”

Borhidi et al. (2017:62; 2018:293) cited the type of *Catesbaea parviflora* as “Tipo: Jamaica. leg.: Swartz s.n. Holotipo: S!” According to the *Code*, starting from 1 January 2001, the designation of a lectotype, or a neotype, must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al.’s (2017, 2018) type designations are not valid.

In S there is a specimen, with accession number S-R-950, which, on the verso of the sheet, has the annotations “Catesbaea parviflora Swartz. D. Swartz” and “E Jamaica: Swartz” handwritten by two different authors. The specimen consists of a small branch with pairs of acicular thorns and numerous pairs of small leaves. By the side of the branch are affixed the flowering branches of a *Spermacoce* species, and two medium-sized leaves that do not belonging to any *Catesbaea* species. The branch with acicular thorns is here designated as the lectotype of *Catesbaea parviflora*.

*Catesbaea campanulata* Sagra ex DC., Prodr. 4:401. 1830. *Echinodendrum campanulatum* (Sagra ex. DC.) A. Rich., in Sagra, Hist. Fis. Cuba, Bot. 11:18, tab. 47bis. 1850. TYPE: CUBA: Havana, s.d., *R. de la Sagra s.n.* (HOLOTYPE: G-DC [G00666482]).

**Notes.**—Candolle (1830:401) cited the material studied of *Catesbaea campanulata* Sagra ex DC. as “in Cuba propè Havanam legit cl. Ramon de la Sagra. An à C. parviflorà satis differt? (v.s.).” With this citation, he expressed doubts of whether this taxon might differ from *C. parviflora* Sw., and indicated that he saw a herbarium specimen, without citing the herbarium of deposit. In G-DC there is a single specimen, with barcode G00666482, annotated as “Catesbaea campanulata” by Candolle. On the specimen is pinned a label with the annotation “66. Catesbaea campanulata spec. nov. La Havanne Mr. Ramon de la Sagra, fevrier 1829” probably handwritten by Ramón de la Sagra. The specimens consists of a small branch with a few lateral branches, several pairs of though thorns, and congested minute leaves, and is the holotype of this name.

*Catesbaea parviflora* var. *septentrionalis* Krug & Urb., in Urban, Symb. Antill. 1:429. 1899. TYPE: USA. FLORIDA: Southern Florida, Bahia Honda Key, s.d., A.H. *Curtiss 1130* (LECTOTYPE (Gillis 1974:162): GH [00058994]; ISOLECTOTYPE: A [00105426]).

**Notes.**—Krug and Urban (in Urban 1899:429), in the protologue of *Catesbaea parviflora* var. *septentrionalis* Krug & Urb., cited several gatherings from Florida, The Bahamas, and Cuba. The original material at B was destroyed during WWII. Gillis (1974:162) cited the type of *C. parviflora* var. *septentrionalis* as “Lectotype: Florida, Bahia Honda Key, *Curtiss 1130* (GH); isotype: GH.” There are two specimen of *Curtiss 1130*, one in GH and the other in A. These two herbarium codes belong to specimens present in the Harvard University Herbarium, and Gillis cited both of them as present in GH.

In GH there is a single specimen of *Curtiss 1130*, on which is affixed a label with the typewritten annotation “Lectotype: Catesbaea parviflora var. septentrionalis Krug & Urb., Symb. Antill. 1: 429. 1899. William T. Gillis August 1974.” This specimen, with barcode 00058994, has a label with printed heading “Curtiss, North American Plants” and the printed annotations “Shrub 4–6 feet high; fruit white. Bahia Honda Key, South Florida.” It consists of two branches densely covered with small leaves, with pairs of acicular thorns at every node, and numerous immature fruits. This is the specimen that Gillis designated as the lectotype of *Catesbaea parviflora* var. *septentrionalis*.
The specimen of *Curtiss 1130* in A, with barcode 00105426, has the same labels of the GH specimen. It is the specimen that Gillis (1974:162) cited as "isotype: GH." The specimen consists of a ramified branch densely covered with small leaves, with aciculat thorns at every node, and several immature fruits, and is an isotype.

*Catesbaea fasciculata* Northr., Mem. Torrey Bot. Club 12:66, tab. 16. 1902. **TYPE:** BAHAMAS: ANDROS: Fresh Creek, 6 Jun 1890, J.I. Northrop & A.R. Northrop 627 (LECTOTYPE (Gillis 1974:162): F [No. 130711]; ISOLECTOTYPES: A [00058993], G [G00389802], GH [00105427], K [K000173529], NY [00007353]).

**Notes.**—Alice Northrop (1902:66–67, tab. 16) in the protologue of *Catesbaea fasciculata* Northr., cited the material studied as “Collected in Fresh Creek, Andros, June 6. The same as *Eggers 4508* from Hog Island (627).” The number “627” corresponds to the gathering *Northrop & Northrop 627*. Hence, she cited two gatherings, *Northrop & Northrop 627*, from Andros Insland, and *Eggers 4508*, from Hog Island, without citing the herbarium of deposit. According to Stafleu and Cowan (1981:775), Northrop's herbarium and types are at “F, other material in A, B, G, GH, K, NY, US.” Northrop (1902:10) in the chapter entitled “Analysis of the Collections” stated “The type specimens are in the herbarium of Columbia University [now integrated in NY]. Sets are also at Kew, The Royal Botanical Garden at Berlin, the Gray herbarium, and Geneva.” Article 9.1 Note 1 of the *Code* (Turland et al. 2018), states, “If the author used only one specimen or illustration, either cited or uncited, when preparing the account of the new taxon, it must be accepted as the holotype, but the possibility that the author used additional, uncited specimens or illustrations (which may have been lost or destroyed) must always be considered.” John and Alice Northrop produced various duplicates of their gatherings, which are now at B, F, G, GH, K and NY. Alice Northrop did not prepare the protologue from a single specimen now at NY. The duplicates of *Northrop & Northrop 627* and *Eggers 4508* are syntypes, one of which should be designated as lectotype. Hence, I disagree with Kass (2005) that the specimen *Northrop & Northrop 627* in NY, barcode 00007353, is the holotype of *C. fasciculata*.

Gillis (1974:162) treated *Catesbaea fasciculata* as a synonym of *C. parviflora* Sw. and cited its type as, “Type: Fresh Creek, Andros, *Northrop and Northrop 627* (Lectotype: F-130711; Isotypes: A, GH, NY).” Therefore, Gillis designated the F specimen with accession number 130711 as the lectotype of *C. fasciculata*, which is here followed.

**14. Catesbaea parvifolia** DC., Prodr. 4:401. **TYPE:** HISPANIOLA: without locality, s.d. [1819–1820], *C. Bertero s.n.* (HOLOTYPE: G-DC [G00666416]).

**Distribution.**—USA (Southern Florida), Jamaica, Puerto Rico, Haiti, Dominican Republic, Antigua.

**Notes.**—Candolle (1830:401) in the protologue of *Catesbaea parvifolia* DC., cited the material studied and previously published names as “in Santo-Domingo legit cl. Bertero. Catesbaea parviflora var. Domingensis Spreng! syst. 1. p. 416. Gardenia parvifolia Lam. dict. 3. p. 441. ex Roem. et Schult. syst. 5. p. 247.—Randia parvifolia Lam. dict. 3. p. 25 (excl. Sloan. syn.) ill. t. 156. f. 2. huc refer. videtur, etiamsi flos 5-fidus depictus sit. (v.s.).” Just below, Candolle (1830:401) wrote: “*Species exclusae*—[...] C. [Catesbaea] parviflora Lam. = Scolosanthus versicolor.” In G-DC, there is a single original specimen, with barcode G00666416, with two labels. One label has the annotation “Randia parvifolia Lam. Catesbaea _________ [meaning *Catesbaea parvifolia*] DC. handwritten by Candolle. The other label has the annotations “Catesbaea parvifolia W. ex Spreng. H. Balbis” handwritten by Candolle, and “S. Doming. Bertero, M. Balbis 1821” handwritten by another author. The specimen consists of a branch with numerous lateral branches, minute leaves, and pairs of acicular thorns at each node. This specimen is the holotype of this name.

**15. Catesbaea phyllacantha** Griseb., Cat. Pl. Cub. 122. 1866. *Phyllacanthus grisebachianus* Hook. f., Hook. Icon. Pl. 11:77, tab. 1095. 1871. **TYPE:** CUBA: [Pinar del Río] “Cultivated at Retiro but indigenous on north coast,” 1863, C. Wright 2655 (HOLOTYPE: GOET [GOET010428]; ISOTYPES: GH [00058984], K [K000173528], MO [MO-038765], NY [00115059], S [No. S-R-7820], YU [YU.001746]).

**Distribution.**—Cuba (Pinar del Río, Toscano, in forests bordering the mangroves towards Inojal). Probably extinct.

**16. Catesbaea sphaerocarpa** Urb., Symb. Antill. 7:401. 1912. **TYPE:** HAITI: “Prope André,” Apr 1899, *W. Buch* 196 (NEOTYPE, here designated: GH [00058987]).

**Distribution.**—Haiti.

**Notes.**—Urban (1912:401–402) in the protologue of *Catesbaea sphaerocarpa* Urb., cited the material studied as “Hab. in Haiti prope Gonaïves in Plaine, m. Dec. fruct.: Picarda n. 1558, ? prope André m. April. flor. [...]: Buch n. 196 [...].” The original material at B was destroyed during WWII. As a result of a search in numerous herbaria, a single specimen of *Buch 196* was found in GH, with barcode 00058987. There is no evidence on the sheet that it was studied by Urban. On the sheet are affixed five ramified branches with pairs of acicular thorns, some of them subtended by pairs of minute leaves, and a few flower buds. This specimen is here designated as the neotype of *C. sphaerocarpa*.
17. *Catesbaea spinosa* L., Sp. Pl. 109. 1 May 1753. TYPE: [icon.]: "Frutex Spinosus Buxi foliis, plurimis simul nascentibus; flore tetrapetaloide, pendulo, sordide flavo, tubo longissimo; fructu ovali croceo, semina parva continentem" in Catesby, Nat. Hist. Carolina p. 100, tab. 100. 1743. (lectotype (Dandy Sloane Herbarium: 112. 1958). Voucher: Herb. Sloane 232:21 (BM-SL; collected by Catesby near Nassau, New Providence, The Bahamas).

**Distribution.**—Cultivated in southern Florida, Bahamas, Cuba (the whole island).

**Notes.**—Linnaeus (1753:109–110), in the protologue of *Catesbaea spinosa* L., cited "*Catesb. carol. 2. p. 100. t. 100. Habitat in Providencia*." With the expression "*Catesb. carol.*," he meant Catesby's publication *Natural History of Carolina*, and specifically referred to volume 2, page 100, table 100. Dandy (1958:112) in his publication *Sloane Herbarium*, cited the same reference provided by Linnaeus, which is associated with the specimen in the Sloane Herbarium No. 232: 21 at BM-SL.

*Catesbaea latifolia* Lindl., Bot. Reg. 10:Tab. 858. 1825 ["1824"). TYPE: [icon.]: Tab. 858 of Lindl., Bot. Reg. 10:1825 ["1824"] (LECTOTYPE, here designated).

**Notes.**—Lindley (1825:Tab. 858) cited the material studied of *Catesbaea latifolia* Lindl., as "Received from the West Indies by Mr. Colville, in whose Nursery our drawing was made this summer." In Lindley's plate 858 in the *Botanical Register* is depicted a branch with pairs of long acicular thorns, subtended by pairs of medium-sized leaves. On the branch are inserted two flowers in anthesis with large corollas, exerted stamens and style. All the elements depicted in this plate are sufficient for the application of the name. Hence, in absence of original specimens associated with this name, Table 858 is here designated as the lectotype of this name.

*Catesbaea longiflora* Sw., Prodr. 30. 1788, nom. superfl.

**Notes.**—Swartz (1788:30) below *Catesbaea longiflora* Sw. cited *C. spinosa* L. (as "*ipinosa*"). Hence *C. longiflora* is a superfluous name.

*Catesbaea macracantha* C. Wright, Anales Acad. Ci. Méd. Habana 6:99. 1869. TYPE: CUBA. Sancti Spiritus: "En el Potrero Manatí jurisdiccion de Trinidad cerca de los manglares," s.d., C. Wright 3575 (LECTOTYPE, here designated: GH [00058990]; ISOLECTOTYPES: NY [00115057], US [0138220]).

**Notes.**—Wright (1869:99) in the protologue of *Catesbaea macracantha* C. Wright, cited the collection locality and his own collection number, without citing the herbarium of deposit.

Borhidi et al. (2017:61–62; 2018:293) cited the type of *C. macracantha* as "Cuba. Oriente, Potrero Manatí, Wright 3575. Holotype: GH; isotype, HAC!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al. (2017:61–62; 2018:293) lectotype designations (as "holotype") are not valid.

As a result of a general search in numerous herbaria, three specimens of Wright 3575 were retrieved. The specimen in US, barcode 00138220, has a label with the annotation "Catesbaea macracantha Wr." handwritten by Wright. The specimen consists of two branches. The branch on the left side of the sheet is densely ramified, densely spinose, with several minute leaves, without fruits or flowers. The branch on the right side is sparsely spinose, with longer spines, subtended by minute leaves, and has a ripe fruit.

The NY specimen of Wright 3575, with barcode 00115057, has a label with the annotation "Catesbaea macracantha Wr." handwritten by Wright. The specimen consists of four branches. All of them with sparse long spines, often subtended by minute leaves, and a ripe fruit.

The GH specimen of Wright 3575, with barcode 00058990, has a label with the annotation "Catesbaea macracantha Wr." handwritten by an unknown author. On the left side of the sheet is affixed a label with the annotation "Catesbaea, Mar 15, Fruticosis 10-15 f. Fr. pallido. Low lands near Mangrove swamps, Potrero Manatí. Trinidad." handwritten by Wright. The specimen consists of two branches, one unbranched, with sparse, long spines and numerous minute leaves. The other branch is densely ramified, with sparse, long spines, and has one ripe fruit. The material on this sheet is here designated as the lectotype of *Catesbaea macracantha*.

Borhidi et al. (2017:61–62; 2018:293), treated *Catesbaea macracantha* C. Wr. as a distinct species. This name is a synonym of *C. spinosa*.

## CATESBAEA EXCLUDED TAXA

*Catesbaea elliptica* Spreng. ex DC., Prodr. 4:401. 1830. TYPE: Pro syn. *Catesbaea tavassorii*. = *Exostema spinosum* (Le Vavass) Krug & Urb. (See below).

*Catesbaea erecta* Sesse & Mociño ex DC., Prodr. 4:401. 1830. TYPE: Based on unpublished illustration for Fl. Mex. = *Bouvardia erecta* (Sesse & Mociño ex DC.) Standl., in N.L. Britton et al. (eds.), N. Amer. Fl. 32:110. 1921.

*Catesbaea inermis* Spreng., Syst. Veg., ed. 16, 1:416. 1825 ["1824"). TYPE: Not traced. = *Rondeletia inermis* (Spreng.) Krug & Urb., in Urban, Symb. Antill. 1:416. 1899.

**Notes.**—Sprengel (1825:416), in the protologue of *Catesbaea inermis* Spreng., cited the locality and collector as "*Portorico*. Bertero" without citing the herbarium of deposit. He described *C. inermis* as "C. foliis oblongis obtusis mucronatis venoso-lineatis, floribus
axillaribus solitariis." The original material studied by Sprengel at B was destroyed during WWII. As a result of a general search in TO, G, G-DC, M, W, and the Jstor Global Plants and Jacq virtual herbaria, no original material was found. Hence, a neotype for this name needs to be designated. Krug and Urban (1899:416-417) transferred *C. inermis* to *Rondeletia*, as *R. inermis* (Spreng.) Krug. & Urb., under which they recognized four varieties.

*Catesbaea parahybensis* Vell., Fl. Flum. 54. 1829 [1825]. [fl. Flum. Icon. 1:Tab 145. 1831 ["1827"] [Protologue] "*Florentem vidi Sept. ad ripas fluvii vulgó dieti Parahyba prope locum 15 Ilhas denominatum.*"
= *Faramea* sp.

**Notes.**—Vellozo (1829:54) in the protologue of *Catesbaea parahybensis* Vell., described the collection locality near the Rio Parahiba, southern Brazil. Vellozo's herbarium has been lost, and the plates published in *Flora Fluminensis Icones* were published two years after the description (Vellozo, 1831), hence they are not original material. The original drawings incommended by Conceicao Vellozo and included in the *Florae Fluminensis Icones* are preserved in two institutions: 1) the Manuscript Section of the National Library in Rio de Janeiro (https://bndigital.bn.gov.br/acervodigital), which is a complete set of the original plates, bound in 11 volumes, with each plate identified by a unique catalogue number; and 2) the Archives of the Torre do Tombo, Lisbon (https://digitarq.arquivos.pt/results?t=florae+fluminensis), which is an incomplete set, with only volumes 6, 7, 8, 9, and 11, and with the drawings slightly different from those in the

Manuscript Section of the Brazilian National Library. The typification of *C. parahybensis* Vell. is not attempted in this treatment.

*Catesbaea triacantha* Spreng., Neue Entdeck. Pflanzenk. 3:47. 1822. *Chomelia triacantha* (Spreng.) Griseb., Cat. Pl. Cub.:133. 1866. *Guettarda triacantha* (Spreng.) F. Maza, Anales Soc. Esp. Hist. Nat. 23:290. 1894. = *Scolosanthus triacanthus* (Spreng.) DC. (See below).

*Cinchona spinosa* Vavass., Observ. Phys. 37(2):243, tab. 2. 1790. *Catesbaea vavassorii* Spreng., Syst. Veg. 1:416. 1825, nom. nov. = *Exostema spinosum* (Le Vavass.) Kr. & Urb. (See below under *Exostema spinosum*). https://gallica.bnf.fr/ark:/12148/bpt6k9603516s/f257.item (Just after page 320)

## CERATOPYXIS

Joseph Dalton Hooker (1872:24, tab. 1125) described the genus *Ceratopyxis* Hook.f., with a sole species, *C. verbenacea* (Griseb.) Hook.f., which is based on *Rondeletia verbenacea* Griseb. He did not explain the etymology of the genus name. From the Greek, *cerato-* (κέρατο, horn) and pixys (*πήεης*, capsule), could mean either horny capsule or capsule with horns. *Ceratopyxis* has traditionally been included in the Chiococcceae (Bremer 1992; Delprete 1996; Robbrecht 1988, 1994). It is distinguished from other genera within the tribe by the composition of the following characters: erect shrub, resinous at nodes, stipules large, ovate, intrapetiolar, persistent, inflorescence terminal, frondose, thyrsiform, cylindrical, densiflorous, ovaries 2-locular, with one pendulous ovule per locule, fruits capsular, subcoriaceous, laterally compressed, septicidal, seeds pendulous, oblong-subreniform, and laterally compressed. Its position in the Chiococcceae was confirmed by the molecular phylogenies of Rova et al. (2002) and Paudyal et al. (2018). *Ceratopyxis* is a monotypic genus endemic to the Cuban province of Pinar del Río.

**Ceratopyxis** Hook.f., Hook. Icon. Pl. 12:24, tab. 1125. 1872; Liogier, Fl. Cuba 5:89. 1962; Vales & Babos, Acta Bot. Acad. Sci. Hung. 23(1-2):275-283. 1977; Borhidi et al., Rubiáceas Cuba 64-65, f. 12. 2017. **TYPE:** *Ceratopyxis verbenacea* (Griseb.) Hook. f.

1. **Ceratopyxis verbenacea** (Griseb.) Hook. f., Hook. Icon. Pl. 12(1):24, tab. 1125. 1872. *Rondeletia verbenacea* Griseb., Cat. Pl. Cub. 130. 1866. **TYPE:** CUBA: "Cuba occ.," Pinar del Río, s.d., *C. Wright* 2695 (HOLOTYPE: GOET [GOET010206]; ISOTYPES: G [2 sheets, G00389794, G00389738], GH [00095875 (without collection number)], K [1 sheet, K000432658, K000432659], MO [No. 2091124], NY [02200784], S [No. S05-990], YU [YU.001727]).

**Distribution.**—Cuba (Pinar del Río: Sierra de los Órganos).

**Notes.**—Grisebach (1866:130) in the protologue of *Rondeletia verbenacea* Griseb., wrote "Habitu accedit Lercheam et probabiliter novum genus formabit.—Cuba occ. (Wr. [Wright] 2695). E." In GOET, where Grisebach worked, there is a sole specimen, with barcode GOET010206, which has two labels with the annotation "Rondeletia verbenacea m." followed by a detailed description handwritten by Grisebach. The specimen consists of a small branch with several leaves and two inflorescences with numerous dehisced capsules. This specimen is the holotype of this name.
## CHIOCOCCA

Patrick Browne (1756:164) described the genus *Chiococca*, without mentioning a species, and cited the vernacular name as “The climbing Snow-berry.” Accordingly, he derived the genus name from the Greek, chion-(κιών, snow) and -kokkos (κόκκος, berry), referring to the white fruits. Paudyal et al.’s (2018) included 17 species of *Chiococca* in their molecular phylogenetic study. In the phylogenies obtained, 14 species were resolved on one clade, while two species, *C. naiguatensis* Steyerm. (Venezuela) and *C. plowmanii* (coastal dunes of Brazil), were found on a strongly supported separate clade, along with two entries of *Salzmannia nitida* (coastal dunes of Brazil). In Paudyal et al.’s (2018) phylogenies, another species of *Chiococca*, *C. cubensis* Urb., was found on yet another strongly supported clade, as sister taxon to the *Scolosanthus* clade. Hence, they transferred *C. cubensis* to the monospecific genus *Ramonadoxa* (See below). *Chiococca* is the type genus of the tribe Chiococceae, and has variable habit, ranging from small shrub to vine to lianas, small, deeply lobed flowers, usually white or yellow corollas (rarely red outside), and 2-seeded (rarely 1- or 3-seeded) spongy drupes. *Chiococca* (including *Asemnantha*) is here recognized as genus of 22 species occurring throughout the Neotropics, and in subtropical regions, with the center of diversity in Mexico and Central America, with 2-3 species in the Greater Antilles, and a few species in South America.

**Chiococca** P. Browne, Civ. Nat. Hist. Jamaica 164. 1756; Standley, N. Amer. Fl. 32(4):285-290. 1934; Liogier, Fl. Cuba 5:93-94. 1962; Steyermark, Mem. New York Bot. Gard. 23:373-384, f. 63. 1972; Steyermark, in Lasser & Steyermark, Fl. Venezuela 2:870-888, f. 140-142. 1974; Correll & Correll, Fl. Bahama Arch. 1383-1386, f. 603-604. 1982; Liogier, Fl. Española 7:231-233. 1995; Lorence, Fl. Mesoamericana 4.2:47-51. 2012; Borhidi et al., Rubiáceas Cuba 68-73, f. 14-15. 2017. **TYPE:** *Chiococca alba* (L.) Hitchc.

*Asemnantha Hook.f.*, in Bentham & Hooker, Gen. Pl. 2(1):106. 7-9 April 1873 [Hook.f., in Hooker's Icon. Pl. 12:40, pl. 1145. April 1873 (as "Asemnantha")] **TYPE:** *Asemnantha pubescens* Hook.f., in Bentham & Hooker, Gen. Pl. 2(1):107. 7-9 April 1873 [Hook.f. in Hooker's Icon. Pl. 12:40, pl. 1145. April 1873 (as "Asemnantha pubescens")] [= *Chiococca motleyana* Borhidi, Acta Bot. Hung. 53:268. 2011].

**Notes.**—The genus *Asemnantha* Hook.f. was published by Joseph Dalton Hooker in 1873, in both Bentham and Hooker's *Genera Plantarum* (as *Asemnantha*) and *Hooker's Icones Plantarum* (as *Asemnantha*). Stafleu and Cowan (1970:294-296), for the entry of *Hooker's Icones plantarum*, on page 294, supplied the date of publication for volume 12 as 1876, which is the date reported on the title page of the volume. However, on page 296, they gave different dates for the four parts of volume 12. Stafleu and Cowan (1970:296) reported that page 40 and plate 1145 of *Asemnantha pubescens* Hook.f. (as *Asemnantha pubescens*) are included in the second part of volume 12, which was published in April 1873 (Hooker, 1873b). This was determined from notes in the copies in BM and K. Article 31.1 of the *Code* (Turland et al. 2018) makes the exception that proof establishing the date of publication supersedes the date on the title page. Because both publications were published on April 1873, it is impossible to establish which of them has priority. As the publication of *Genera Plantarum* vol. 2, part 1, has the date of 7-9 April 1873 (Hooker, 1873a), it is here treated as the first publication of those names, and name *Asemnantha* is retained for this genus.

**1. Chiococca alba** (L.) Hitchc., Report Missouri Bot. Gard. 4:94. 1893. *Lonicera alba* L., Sp. Pl. 175. 1753. **TYPE:** [proto-logue] "Jamaica, St. Catherine Parish, savanna, towards Two-mile-Wood," [icon.] "Jasminum forte," Sloane, Voy. Jamaica 2, tab. 188, fig. 3. 1725 (LECTOTYPE (Franck et al. 2021:45); EPITYPE (Franck et al. 2021:45): "Jamaica," *H. Sloane s.n.* (BM [000594058]).

**Distribution.**—From Southern USA (Texas & Florida), Mexico to Argentina.

**Notes.**—This is the most widespread species in the genus, ranging throughout the Neotropics, and extending in subtropical regions of southern USA and northern Argentina. As here delimited, it is a very variable species, in terms of vegetative and reproductive characters. It has a variable habit ranging from shrub, vine, or liana, climbing on trees up to 15 m from the ground. A detailed study of specimens from throughout the range of this species might reveal that some of the names here listed in synonymy might deserve to be recognized as distinct species or infraspecific taxa.

*Chiococca racemosa* L., Syst., Nat., ed. 10, 2:917. 1759, *nom. superfl.* (*Lonicera alba* L. (1753) is cited in synonymy).

*Chiococca brachiata* Ruiz & Pav., Fl. Peru. 2:67, pl. 219b. 1799. **TYPE:** PERU: Chinchao, ["in nemoribus Chinchao Quebrada ad Macora praedium"], s.d., *H. Ruiz López & J.A. Pavón Jiménez s.n.* (LECTOTYPE (Franck et al. 2021:45): MA [MA815658]; ISOLECTOTYPE: MA [MA815659]).

*Chiococca pubescens* Humb. & Bonpl. ex Schult., in Roemer & Schultes, Syst. Veg., ed. 15[bis], 5:202. 1819. *Chiococca anguifuga* Mart. var. β *pubescens* (Humb. & Bonpl. ex Schult.) DC., Prodr. 4:483 (1830). **TYPE:** VENEZUELA: Cumaná, ["Habitat in America meridionali"], s.d., A.J.A. Bonpland & F.W. Humboldt *s.n.* (LECTOTYPE, here designated: B-W [B-W 04141-01 0]).
**Notes.**—Josef August Schultes (in Roemer & Schultes, 1819:202) attributed the name *Chiococca pubescens* to Humboldt and Bonpland, and in the protologue he also cited "Reliqu. Willd. MS." meaning that the specimen was previously studied by Willdenow. In B-W there is a folder [Willdenow folder B-W 04140] with the label "Pentandria Monogynia, Chiococca parviflora [...] Habitat in Cumana" handwritten by Willdenow. Inside the folder, there are two sheets, both of them with the annotation "Ch. parviflora" handwritten on the upper right corner. Specimen with barcode B-W 04141-01 0, kept inside that folder, is here designated as the lectotype of *Chiococca pubescens* Humb. & Bonpl. ex Schult.

*Chiococca parviflora* Humb. & Bonpl. ex Schult., in Roemer & Schultes, Syst. Veg., ed. 15 [bis] 5:202. 1819. TYPE: VENEZUELA: Cumana, s.d., A.J.A. Bonpland & F.W. Humboldt s.n. (LECTOTYPE (Franck et al. 2021:45): B-W [B-W 04140 01 0]; POSSIBLE ISOLECTOTYPES: B-W [B-W 04140 02 0], HAL [0113824]).

**Notes.**—Schultes (in Roemer & Schultes, 1819:202) attributed the name *Chiococca parviflora* to Humboldt and Bonpland, and in the protologue he also cited "Reliqu. Willd. MS." meaning that the specimen was previously studied by Willdenow. In B-W there is a folder [Willdenow folder B-W 04140] with the label "Pentandria Monogynia, Chiococca parviflora [...] Habitat in Cumana" handwritten by Willdenow. Inside the folder, there are two sheets, both of them with the annotation "Ch. parviflora" handwritten on the upper right corner. Specimen with barcode B-W 04140 01 0, kept inside that folder, was designated as the lectotype of *C. parviflora* Humb. & Bonpl. ex Schult. by Franck et al. (2021:45).

*Chiococca paniculata* Hoffm. ex Schult., in Roemer & Schultes, Syst. Veg., ed. 15[bis]. 5:203. 1819, not validly published.

*Chiococca anguifuga* Mart., Denkschr. Königl. Akad. Wiss. München 1824:93, tab. 9, figs. 20–21. 1824. TYPE: BRAZIL. MINAS GERAIS: "Crescit in sylvis primaevis ad Praesium S. Joannis Baptistae et alibi in parte orientali Provinciae Minas Gerais," s.d., C.F.P. Martius s.n. [Martius Obs. 1088] (LECTOTYPE, here designated: M [M-0187149]).

**Notes.**—Martius (1824:93) in the protologue of *Chiococca anguifuga* Mart., cited the collection locality of a gathering collected by himself, without citing his collection number or the herbarium of deposit. Müller Argoviensis (1881:51) synonymized *C. anguifuga* Mart. under *C. brachiata* var. β genuina Müller. Arg. [= C. alba]. In M there is a specimen, barcode M-0187149, which has a label with the heading "Dr. Martius Iter Brasil." and the handwritten annotations "Chiococca anguifuga Mart., Raiz preta Bras. Habitat in sylvis aboriginibus ad Presid. S. Joannis Babtistae, Provinciae Minarum, Observ. 1088." The specimen consists of a branch with several leaves and several inflorescences with filiform rachis and sparse, secundiflorous secondary branches. On the inflorescences are present several fruits. This specimen, collected and described by Martius, is here designated as the lectotype of *C. anguifuga*.

*Chiococca densifolia* Mart., Denkschr. Königl. Akad. Wiss. München 1824:93, tab. 6. 1824. *Chiococca brachiata* var. κ densifolia (Mart.) Müll. Arg., in Martius et al., Fl. Bras. 6(5):53. 1881. TYPE: BRAZIL. BAHIA: "Habitat in sylvis primaevis ad Almada et Ferradas nec non alibi in saltu ille vacatissimo, montium maritimum tractum in Provincia Bahiensi laeta et aeterna fronde tegente," s.d., C.F.P. Martius Herb. Bras. 603 (LECTOTYPE, here designated: M [M-0187142]; ISOLECTOTYPES: GH [00257765], HAL [HAL0113133], W [Reicherbach fil. No.W1889-0287615]).

**Notes.**—Martius (1824:93), in the protologue of *Chiococca densifolia* Mart., cited the collection locality of a gathering collected by himself, without mentioning the collection number or the herbarium of deposit. Müller Argoviensis (1881:51) published the new combination *C. brachiata* var. κ densifolia (Mart.) Müll. Arg. [= C. alba], under which he cited, among other gatherings, "Martius Hb. Flor. Bras. n. 602, 603." In M there is a specimen, barcode M-0187142, which has a label with the handwritten annotation "Chiococca densiflora [sic] Mart. var. (Ch. anguifuga in herb. Mart.)," the printed annotation "Martii Herbar. Florae Brasil. No.,” and the handwritten number "603." This specimen consists of two branches with numerous leaves and several axillary inflorescences. Because this specimen was collected and described by Martius, it is here designated as the lectotype of *C. densifolia*.

*Chiococca racemosa* Jacq. var. β scandens DC., Prodr. 4:482. 1830. TYPE: PORTO RICO: without locality, 1827, H. Wydler 368 (LECTOTYPE, here designated: G-DC [G00666532]).

**Notes.**—Candolle (1830:482), in the protologue of *Chiococca racemosa* var. β scandens DC., cited "In Jamaicâ, ins. Stae-Crucis. Brown. jam. 164. n.2. Swartz obs. var. β (v.s.)". In G-DC there is a single specimen associated with this name, which is annotated as "Chiococca racemosa β scandens Sw. Pers." by Candolle. This specimen, with barcode G00666532, has a label with the annotation "H. Wydler. Portorico. 1827. N° 368, Chiococca racemosa." This specimen is here designated as the lectotype of *C. racemosa* var. scandens.

*Chiococca racemosa* Jacq. var. γ laxiflora DC., Prodr. 4:482. 1830. TYPE: cultivated: ["v.s. cult. in hort. Calc."], 1819, N. Wallich s.n. (LECTOTYPE (Franck et al. 2021:45): G-DC [G00666534]).

*Chiococca racemosa* Jacq. var. δ longifolia DC., Prodr. 4:482. 1830. TYPE: GUADALOUPE: ["In Guadalupâ legit cl. Badier. (v.s.)"], without locality, s.d. [1780–1790], Badier s.n. (LECTOTYPE, (Franck et al. 2021:45): G-DC [G00666535]; POSSIBLE ISOLECTOTYPE: P-LA [P00308513]).

**Notes.**—Candolle (1830:482), in the protologue of *Chiococca racemosa* var. γ laxiflora DC., cited the material studied as "In Guadalupâ legit cl. Badier. (v.s.)". In G-DC there is a sheets with two specimens affixed on it. On the bottom right corner is pinned a label with the annotation "Chiococca racemosa γ laxiflora DC." handwritten by Candolle. The specimen on the right side of the sheet, with barcode G00666535, was designated as the lectotype of *C. racemosa* var. longifolia by Franck et al. (2021:45).

In P-LA, there is a specimen with barcode P00308513, which has the two labels affixed on the bottom right corner. The bottom label has the annotation "Chiococca racemosa L." handwritten by an unknown author, and the label just above has the annotation "de la Guadeloupe coll. Badier. This specimen is a possible isolectotype of *C. racemosa* var. longifolia."
*Chiococca racemosa* var. jacquiniana Griseb., Fl. Brit. W. I.:337. 1861.

**Notes.**—Grisebach (1864:337) intended *Chiococca racemosa* var. jacquiniana Griseb. as the typical variety, but the typical variety of this species is var. *racemosa*.

*Chiococca latifolia* Raf., Alsogr. Amer. 75. 1838. TYPE: CUBA: without locality, s.d., *Jalambic s.n.* (LECTOTYPE: (Franck et al. 2021:46); DWC n.v.).

*Chiococca macrocarpa* M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11(1):230. 1844. TYPE: MEXICO. VERA CRUZ: Mirador, Jun.—Oct 1840, H. Galeotti 7064 (HOLOTYPE, BR [000000530537]).

**Notes.**—Martens and Galeotti (1844:230–231), in the protologue of *Chiococca macrocarpa* M. Martens & Galeotti, cited the gathering Galeotti 7064, and the locality “Mirador,” without citing the herbarium of deposit. In BR there is a sole specimen, with barcode 000000530537, annotated with this name, which is the holotype.

*Chiococca* var. *a. grandifolia* Müll. Arg. in Martius et al., Fl. Bras. 6(5):50. 1881. TYPE: PERU: Tarapoto, 1855–1856, *R. Spruce* 3933 (LECTOTYPE, here designated: G [G00389863]; ISOLECTOTYPES: BR [2 sheets, 000000557803, 000000557738]).

*Chiococca brachiata* var. *β. genuina* Müll. Arg., in Martius et al., Fl. Bras. 6(5):51. 1881. TYPE: BRAZIL. SÃO PAULO: without locality, s.d., W.J. Burchell 3214 (LECTOTYPE, here designated: BR [000000558529]; ISOLECTOTYPE: GH [00257763]).

*Chiococca brachiata* var. *γ. valida* Müll. Arg. in Martius et al., Fl. Bras. 6(5):51. 1881. TYPE: BRAZIL. BAHIA: without locality, 1834, J.S. Blanchet 1156 (LECTOTYPE, here designated: G [G00389835]).

*Chiococca brachiata* var. *δ. conjengens* Müll. Arg., in Martius et al., Fl. Bras. 6(5):51. 1881. TYPE: BRAZIL. RIO DE JANEIRO: Without locality, s.d., L. Riedel 368 (LECTOTYPE, here designated: BR [000000530739, specimen on the right side of the sheet]).

*Chiococca brachiata* var. *η. intermedia* Müll. Arg., in Martius et al., Fl. Bras. 6(5):52. 1881. TYPE: BRAZIL. MINAS GERAIS: Lagoa Santa, s.d., E. Warming s.n. (HOLOTYPE: G [G00389862]).

*Chiococca brachiata* var. *ζ. biformis* Müll. Arg., in Martius et al., Fl. Bras. 6(5):52. 1881. TYPE: BRAZIL. RIO DE JANEIRO: without locality, Sep 1836, Ildephonso Gomez [A.I. Gomes de Freitas] 23 (LECTOTYPE, here designated: G [G00389876]).

*Chiococca brachiata* var. *η. intermedia* Müll. Arg., in Martius et al., Fl. Bras. 6(5):52. 1881. TYPE: BRAZIL. MINAS GERAIS: “Prope Fachada,” Nov 1824, L. Riedel 854 (LECTOTYPE, here designated: BR [000000530741]).

*Chiococca brachiata* var. *θ. diplomorpha* Müll. Arg., in Martius et al., Fl. Bras. 6(5):52. 1881. TYPE: BRAZIL. GOIAS: “prope urbem Goyaz” [town of Goiás], s.d., W.J. Burchell 6524 (LECTOTYPE, here designated: BR [000000530674]; ISOLECTOTYPE: GH [00257762]).

*Chiococca brachiata* var. *t. rigida* Müll. Arg., in Martius et al., Fl. Bras. 6(5):53. 1881. TYPE: BRAZIL. BAHIA: “parte meridionali prov. Bahia,” 1840, J.S. Blanchet 3108A (LECTOTYPE, here designated: G [G00389843]; ISOLECTOTYPES: G [G00389885], W [No. W0027947]).

*Chiococca brachiata* var. *λ. acutifolia* Müll. Arg., in Martius et al., Fl. Bras. 6(5):53, tab. 7. 1881. TYPE: BRAZIL. RIO DE JANEIRO: Without locality, s.d., A.F.M. Glaziou 3034 (LECTOTYPE, here designated: BR [000000530506]).

*Chiococca brachiata* var. *μ. microphylla* Müll. Arg., in Martius et al., Fl. Bras. 6(5):53. 1881. TYPE: BRAZIL. BAHIA: without locality, s.d., J.S. Blanchet 3276 (LECTOTYPE, here designated: BR [000000530678]).

*Chiococca brachiata* var. *v. acuminata* Müll. Arg., in Martius et al., Fl. Bras. 6(5):53. 1881. TYPE: BRAZIL. MATO GROSSO: Cuiabá, “ubi ab incolis Caringa nuncupatur,” 1832, M. da Silva Manso 24 (LECTOTYPE, here designated: G [G00389881]).

*Chiococca brachiata* var. *ξ. subrhombea* Müll. Arg., in Martius et al., Fl. Bras. 6(5):54. 1881. TYPE: BRAZIL. RIO DE JANEIRO: “prope Rio de Janeiro,” s.d., W.J. Burchell 2774 (LECTOTYPE, here designated: BR [000000530677]).

*Chiococca brachiata* var. *o. lanceolata* Müll. Arg., in Martius et al., Fl. Bras. 6(5):54. 1881. TYPE: BRAZIL. MINAS GERAIS: Caldas, “opp. Caldas in cafoes,” Dec 1854, A. Lindberg 97b (LECTOTYPE, here designated: BR [000000530708]).

*Chiococca brachiata* var. *π. petiolaris* Müll. Arg., in Martius et al., Fl. Bras. 6(5):54. 1881. TYPE: BRAZIL. BAHIA: “ad Igreja Velha,” 1841, J.S. Blanchet 3276 (LECTOTYPE, here designated: G [G00389853]; ISOLECTOTYPE: BR [000000530711]).

*Chiococca brachiata* var. *tenuifolia* Müll. Arg., in Martius et al., Fl. Bras. 6(5):458. 1881. TYPE: BRAZIL. Rio de Janeiro: without locality, s.d., A.F.M. Glaziou 10941 (LECTOTYPE, here designated: G [G00389837]).

*Chiococca micrantha* J.R. Johnston, Proc. Amer. Acad. Arts 40:696. 1905; Proc. Bot. Soc. Nat. Hist. 34:t, 29, F. 3a-c. 1909. *Chiococca alba* (L.) Hitchc. var. *micrantha* (J.R. Johnston.) Steyerm., Acta Bot. Venez. 6:139. 1971. TYPE: VENEZUELA. NUEVA ESPARTA: Island of Margarita, San Juan Mountains, woods above South Hill, 27 Jul 1903, J.R. Johnston 115 (LECTOTYPE: Howard 1899:399, as “holotype”): GH [00057576]; ISOLECTOTYPES: F [No. 1745174], G [2 sheets, G00389832], K [000432648], NY [00099439], US [00138518]). **Notes.**—Johnston (1905:696–697), in an article entitled “New plants from the Islands of Margarita and Coche, Venezuela,” described *Chiococca micrantha* J.R. Johnston. In the protologue, he cited the material studied as “San Juan Mts., in woods above South hill, alt. 400 m, Johnston, no. 115, July 27, 1903,” without citing the herbarium of deposit. Howard (1899:399) cited the type of this name as “Type: Johnston 115 (holotype GH!). Although Johnston’s label on the holotype indicates a collection from the Island of Margarita, Steyermark (Fl. Venezuela 9:886. 1974) attributes the collection to Cerro de San Juan, Edo. Nueva Esparta, Venezuela.” Howard’s correction is accurate, as the label of the GH specimen has the heading “Plants of the Island of Margarita, Venezuela.”

*Chiococca bermudiana* S. Br., Proc. Acad. Nat. Sci. Philadelphia 61:493. 1910. TYPE: BERMUDA: north shore, near Flatts, Mt. Langdon, 31 Aug.–20 Sep 1905, S. Brown & N.L. Britton 181 (HOLOTYPE: PH [00005273]; ISOTYPES: A [00057578], F [No. 203773], GH [00057577], PH [00005272]).

*Chiococca racemosa* var. yucatana Loes., Repert. Spec. Nov. Regni Veg. 18(524/530):361. 1922. TYPE: MEXICO. YUCATAN: “im distrikt Ticul, auf der Hacienda Yaxché im Walde,” E. Seler & C. Seler 5591 (B, destroyed). MEXICO: YUCATAN: 4 km W of Buctzotz, sobre la carrera Tizimin–Merida, 21°11′N, 88°49′W, 24 Jul 1987, E.F. Cabrera C. & H. Cabrera 13715 (NEOTYPE, here designated: MO [No. 3927696, barcode MO-508961]).
**Notes.**—Loesener (1922:361) in the protologue of *Chiococca racemosa* var. *yucatana* Loes., cited the gathering *Seler 5591*, without citing the herbarium of deposit. According to Stafleu and Cowan (1981:141), Loesener's original material was at B. That material was destroyed during WWII. A general search in numerous herbaria did not retrieve any original specimen. Hence, a neotype needs to be designated for this name. The specimen *Cabrera & Cabrera 13715* at MO, accession number 3927696, barcode MO-508961, is here designated as the neotype of this name.

*Chiococca alba* ssp. *parvifolia* var. *micrantha* f. *pilosa* Steyerm., Acta Bot. Venez. 6:141. 1972 ["1971"]. TYPE: VENEZUELA. LARA: Cercanías de Barquisimeto, near la Ruesga, Aug 1923, *J. Saer 4* (HOLOTYPE: VEN [No. 15153]; ISOTYPE: US [00138510]).

*Chiococca parviflora* R.O. Williams & Cheesman, in R.O. Williams, Fl. Trinidad & Tobago 2:29. 1928, *orth. var.* of *Chiococca parvifolia*

*Chiococca petenensis* Lundell, Wrightia 5(1):7. 1972. *Chiococca petensis* Lundell, Wrightia 5(1):7. 1972, *orth. var.* TYPE: GUATEMALA. PETEN: Dolores, forest on Santo Toribio, 7 Jun 1961, *E. Contreras* 2439 (HOLOTYPE: LL [00000270]; ISOTYPES: LL [00373120], S [2 sheets, Nos. S10-24110, S05-995]).

*Chiococca trisperma* Hook. f., Trans. Linn. Soc. London 20:219. 1847. TYPE: ECUADOR. GALAPAGOS ISLANDS: Chatam Island, Sep 1835, C.R. *Darwin 109* (LECTOTYPE (Porter 1980): CGE n.v.; ISOLECTOTYPES: K [one sheet, K000432644, K000432646]).

**Notes.**—Joseph Dalton Hooker (1847:219), in the protologue of *Chiococca trisperma* Hook. f., cited the material studied as "*Hab. Chatam Island, Charles Darwin, Esq.*," without citing the herbarium of deposit. Porter (1980) designated as lectotype of this name a specimen in CGE. In K there is one sheet, on which are affixed two specimens. The specimen on the upper portion of the sheet, with barcode K000432646, has a label with the handwritten annotation "109. Chiococca trisperma Nob. Galapagos." The specimen on the left side of the sheet, barcode K000432644, has the stamp "Herbarium Hookerianum" and the handwritten annotation "Chatam Island, Galapagos, C. Darwin" handwritten by Hooker. These two K specimens, affixed on the same sheet, are isolectotypes of *C. trisperma*.

*Chiococca trisperma* f. *angustifolia* Andersson, Kongl. Vetensk. Acad. Handl. 1853:193. 1855 ["1853"). TYPE: ECUADOR. GALAPAGOS ISLANDS: Albermarle Island, "insula A bermarle a Dr Macroe lectam," s.d., *Macroe s.n.* (not found).

**2. Chiococca auyantepuiensis** Steyerm., Mem. New York Bot. Gard. 23:374. 1972. TYPE: VENEZUELA. BOLIVAR: Auyán-tepui, near Río Lomita Camp, 1800 m, 10 May 1964, *J.A. Steyermark* 93652 (HOLOTYPE: VEN [No. 15495]).

**Distribution.**—Venezuela (Auyán-Tepui).

**3. Chiococca belizensis** Lundell, Amer. Midl. Naturalist 29:492. 1943. TYPE: BELIZE. ["British Honduras"). TOLEDO DISTR.: Cow Pen, near Monkey River, 2 Sep 1942, *P.H. Gentle* 4115 (HOLOTYPE: MICH [1210532]; ISOTYPES: A [00057582], F [No. 1258120], LL [00000266], MO [No. 1253527], NY [00099446], US [00138511]).

**Distribution.**—From southern Mexico to Colombia and Peru.

*Chiococca durifolia* Dwyer, Ann. Missouri Bot. Gard. 67:87. 1980. TYPE: PANAMA: Cocle: La Mesa, above El Valle, 3 Jan 1974, *J.D. Dwyer* 11865 (HOLOTYPE: MO [No. 2188155]; ISOTYPES: GH [00257767])

**4. Chiococca capitata** Wernham, J. Bot. 51:323. 1913. TYPE: JAMAICA: without locality, s.d., *W. Wright s.n.* (HOLOTYPE: BM [000551619]). https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.bm00055169

**Distribution.**—Jamaica.

**Notes.**—Wernham (1913:323) distinguished *Chiococca capitata* Wernham by the congested subcapitate cymes and the tubulose-infundibular corolla with minute lobes reflexed at anthesis. He described the congested inflorescences as "Heads barely 1 cm in diameter, even including the corollas." Adams (1972), in *Flowering Plants of Jamaica*, did not mention this name. The combination of congested inflorescences, which is unique within the genus, and the narrowly tubular corollas, is diagnostic in recognizing it as distinct species. To my knowledge, this species is only known by the holotype specimen in BM, with barcode 000551619.

**5. Chiococca caputensis** D.H.Lorence & C.M.Taylor, Novon 5:201. 1995. TYPE: PANAMA. PANAMA: Cerro Jefe, NE of Panama City, 850–900 m, 12 Jul 1986, *G. McPherson* 9744 (HOLOTYPE: MO [No. 3624428]; ISOTYPE: PMA [No. 035797]).

**Distribution.**—Panama (Cerro Jefe).

**6. Chiococca coriacea** M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11(1):231. 1844. *Chiococca racemosa* var. *coriacea* (M. Martens & Galeotti) Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjøbenhavn 1852 (2-4):40. 1853. TYPE: MEXICO. VERA CRUZ: environs de Vera Cruz, s.d., *J.J. Linden* 423 (HOLOTYPE: BR [000000530743]; ISOTYPE: BR [000000530570]).

**Distribution.**—Mexico (Tamaulipas, Veracruz).

**7. Chiococca filipes** Lundell, Contr. Univ. Michigan Herb. 7:53. 1942. TYPE: MEXICO. CHIAPAS: Nuevo Amatenango, 1300 m, 17 Jul 1941, *E. Matuda* 4783 (HOLOTYPE: MICH [1210533]; ISOTYPES: A [00057583], LL [00000267], MEXU [MEXU00084035], MO [No. 1219325], NY [00099445], US [00138516]).

**Distribution.**—Mexico (Chiapas, Oaxaca).
8. **Chiococca grandiflora** Lorence & T.Van Devender, in D.H. Lorence, T.R. Van Deventer & G.M. Ferguson, Phytokeys 98:74, figs. 1–3. 2018. **TYPE:** MEXICO. **SONORA:** Mun. Alamos, near Tepopa, NNW of Chirobo, 1100–1400 m, 22 Aug 1992, P.S. Martin, P. Comtois, C. Lindquist, S.A. Meyer, B. Risner, & D.A. Yetman s.n. sub P. Jenkins 92-135 (HOLOTYPE: ARIZ [309922]; ISOTYPES: ARIZ [383348], PTBG [105887]).

**Distribution.**—Mexico (Sonora).

9. **Chiococca henricksonii** M.C.Johnst., Madroño 28:30. 1981. **TYPE:** MEXICO. **COAHUILA:** 1–2 km N of Puerto Colorado, near crest at SW end of Sierra de La Fragua, above 1750 m, 2 Sep 1941, I.M. Johnston 8738 (HOLOTYPE: LL [00000268]; ISOTYPES: F [No. 1723223], GH [00057580], MICH n.v.).

**Distribution.**—Mexico (Coahuila).

10. **Chiococca lucens** Standl. & Steyerm., Fieldiana, Bot. 28:571, fig. 122. 1953. **TYPE:** VENEZUELA. **BOLIVAR:** Sororopán-tepui, between quebrada and summit, 2130–2250 m, 15 Nov 1944, J.A. Steyermark 60184 (HOLOTYPE: F [No. 1182124]; ISOTYPES: F [No. 118191], US [00138517], VEN [No. 25401]).

**Distribution.**—Venezuela (Bolívar).

11. **Chiococca motleyana** Borhidi, Acta Bot. Hung. 53:268. 2011. *Asemnantha pubescens* Hook.f., in Hook. Icon. Pl. 12:40. Apr 1873 (as "Asemnanthe pubescens"). *Asemnantha pubescens* Hook.f., in Bentham & J.D. Hooker, Gen. Pl. 107. 7–9 Apr 1873. **TYPE:** MEXICO. **YUCATÁN:** "in sylvis humidis Yucatan ad Chiapas lecta," s.d., J.J. Linden 1264bis (HOLOTYPE: K [K000432633]).

**Distribution.**—Mexico (Campeche, Quintana Roo, Yucatán), Guatemala, Belize.

12. **Chiococca nitida** Benth., Hooker's J. Bot. 3:236. 1841. [Delprete, Rev. Biol. Neotrop. 1:4–10. 2005 ["2004"]; Jardim et al., Phytotaxa 202(1):15–25. 2015]. **TYPE:** GUYANA: near Mt. Roraima, 1839, Rob.H. Schomburgk ser. I, 1055 (LECTOTYPE, here designated: K [K000173187]; ISOLECTOTYPES: BM [000551614], F [No. 767823], G [G00389943], L [L0057946], NY [00099436], photo-K at NY).

**Distribution.**—Venezuela, Guyana, French Guiana, Brazil (Amazonas, Pará, Rondônia, Bahia), and Peru.

*Chiococca nitida* Benth. var. *amazonica* Muell. Arg. in Martius et al., Fl. Bras. 6(5):50. 1881. **TYPE:** BRAZIL. **PARA:** Caripi, juxta Pará [now city of Belém], Aug 1849, R. Spruce s.n. (HOLOTYPE: G [G00389944]; ISOTYPES: GH [00257766], M [M-0187135], TCD [TCD0005809]; photo-G at NY).

*Chiococca erubescens* Wernham, J. Bot. 51:322 (1913). **TYPE:** FRENCH GUIANA: without locality, s.d., J. Martin s.n. (LECTOTYPE (Steyermark 1972:376): K [K000432651]; ISOLECTOTYPE: K [K000432650]).

*Chiococca cordata* Cowan, Brittonia 7:411. 1952. *Chiococca nitida* Benth. var. *amazonica* Muell. Arg. f. cordata (Cowan) Steyerm., Mem. New York Bot. Gard. 23:373 (1972). **TYPE:** GUYANA: Kanuku Mts., Soirntau, 725 m, Sep 1948, G. Wilson-Browne 230 (Forest Dept. 5707) (HOLOTYPE: NY [00099441]; ISOTYPE: K [K000432649])

**Notes.**—Cowan (1952:411–412), in the protologue of *Chiococca cordata* Cowan, cited the type as "Type: Wilson-Browne 230 (F. D. 5707) (NY)." The NY specimen with barcode 00099441 has an isotype label, and "holotype" handwritten in pencil. It is the only specimen of *Wilson-Browne* 230 at NY and is the holotype of this taxon.

*Chiococca multipedunculata* Steyerm., Mem. New York Bot. Gard. 23:373. 1972. **TYPE:** GUYANA: Potaro Distr., Mt. Kanaima, 485 m, 20 Aug 1959, B.A. Whitton 120 (HOLOTYPE: NY [00099438]; ISOTYPE: U [U0099834]).

*Chiococca nitida* var. *chimantensis* Steyerm., Mem. New York Bot. Gard. 23:376. 1972. **TYPE:** VENEZUELA. **BOLIVAR:** Chimantá Massif, SW facing portion of Chimantá-tepui, 1410 m, 15 May 1953, J.A. Steyermark 75406 (HOLOTYPE: VEN [No. 71471]; ISOTYPES: BM [000551618], F n.v., MO [2 sheets, Nos. 2012043, 2011333], NY [00099434], WIS [v 0262263 WIS]).

13. **Chiococca oaxacana** Standl, Publ. Field Mus. Nat. Hist., Bot. Ser. 22:383. 1940. **TYPE:** MEXICO. **OAXACA:** Cerro San Antonio, 1650 m, 26 Jun 1906, C. Conzatti 1418 (HOLOTYPE: F [No. 579006]; ISOTYPES: GH [00244641 (frag.)], MEXU [2 sheets, MEXU00033025, MEXU00534928], US [00289016]).

**Distribution.**—Mexico (Morelos, Oaxaca, Puebla, Tamaulipas).

14. **Chiococca pachyphylla** Wernham, J. Bot. 51:323. 1913. **TYPE:** MEXICO. **VERACRUZ:** Between Acatlán and Chiconquiaco, Sep 1829, C.J.W. Schiede & F. Deppe s.n. (HOLOTYPE: BM [000551613]).

**Distribution.**—Mexico, Belize, Guatemala, Honduras, El Salvador, and Nicaragua).

*Chiococca mexicana* Lundell, Wrightia 5:71. 1974. **TYPE:** MEXICO. **PUEBLA:** Tespilco, Mun. Zacapoaxtla, 1800 m, 23 Apr 1970, F. Ventura A. 955 (HOLOTYPE: LL [00000269]; ISOTYPES: ENCB [ENCB003697], F [No. 1706520], MICH [1210521]).

15. **Chiococca petrina** Wiggins, Contr. Dudley Herb. 3:76, tab. 20, fig. 5–13. 1940. **TYPE:** MEXICO. **Sonora:** gorge of andesitic rock 23 mi S of Divisadero, 26 Sep 1934, I.L. Wiggins 7493 (HOLOTYPE: DS [DS No. 263330, CAS barcode 0026742]; ISOTYPES: A [00257760], ARIZ [ARIZ-BOT-0004895], F [No. 11108857], LL [00000271], UC [UC 665454], US [00138519]).

**Distribution.**—Mexico (Chihuahua, Coahuila, Sinaloa, Sonora).
A specimen in K, barcode K000432657, has a label with the annotations "Chiococca phaenostemon Schltdl. Linnaea 9. 594. Haz. de la Laguna. Jul. 29. C. racemosa viz differt." handwritten by several authors.

The GOET specimen, barcode GOET010209, has a label with the annotation "1. Chiococca phaenostemon n. sp. Prov. Jalapa. Comm. Cl. dr. Schlechtendal 1839." probably handwritten by Schlechtendal.

A specimen in GH, barcode 00057579, has a label with the annotation "Chiococca phaenostemon n. sp. Prov. Jalapa" handwritten by Schlechtendal. The collector or the collection number is not reported on the label. The specimen consists of a small branch with four leaves, and a branch with several leaf pairs and two pairs of axillary inflorescences, with flower buds and flowers in anthesis.

A specimen in BM, barcode 000551612, has a label with the annotation "Chiococca, Haz. de la Laguna, Jul. 29" handwritten by an unknown author (Schiede?). There is no evidence on the sheet that it was studied by Schlechtendal. The specimen consists of a branch with a pair of axillary infructescences with mature fruits.

*Chiococca staminea* M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11(1):231. 1844. **TYPE:** MEXICO. **VERACRUZ:** Jalapa ["environs de Xalapa"], s.d., *J. Linden* 48 (**HOLOTYPE:** BR [000000530710]).

**17. Chiococca pinetorum** Britton, Publ. Field Columb. Mus., Bot. Ser. 2(3):171. 1906. **TYPE:** BAHAMAS: New Providence, near Nassau, 1 Sep 1904, N.L. Britton & L.J.K. Brace 430 (LECTOTYPE (Franck et al. 2021: 53): NY [00099443]; ISOLECTOTYPES: F [No. 171854], K [K000432652], US [00138520]).

**Distribution.**—Southern USA (Florida), and Bahama Archipelago.

**Notes.**—Britton (1906: 171) in the protologue of *Chiococca pinetorum* Britton, cited as type the gathering *Britton & Brace 430* without indicating the herbarium of deposit. At the time he published this name, he was the Director of the New York Botanical Garden. In NY there is a specimen, barcode 00099443, which has a label with the heading "The New York Botanical Garden, Exploration of the Bahamas, New Province" and the annotation "Type. 430. Chiococca pinetorum Britton, Pinelands, Harold Road" handwritten by Britton. Above that label is affixed a field label with the annotation "430. C. pinetorum. Cor. white, veined with purple" handwritten by Britton. The specimens consists of a long branch with numerous lateral branches, numerous small leaves, and several infructescences with mature fruits. This specimen was designated as the lectotype of *C. pinetorum* by Franck et al. (2021:53).

The specimen of *Britton & Brace 430* in F, accession number 171854, has a "holotype" label made by W.T. Gillis in July 1974. It is unknown to me the reasoning of labelling this specimen as "holotype." It consists of a long branch with numerous lateral branches, numerous small leaves, and several infructescences with mature fruits. This specimen is an isotype.

**18. Chiococca rubriflora** Lundell, Wrightia 5:7. 1972. **TYPE:** GUATEMALA. **PETEN:** in *corazoal*, W of km 165 of Poptum road, woody vine, flowers dark red outside, orange inside, 10 Sep 1969, E. *Contreras 9110* (**HOLOTYPE:** LL [00000272]; **ISOTYPES:** DUKE [10000635], F [No. 1968054], K [K000432654], LL [00373121], S [2 sheets, Nos. S05-997, S10-24087]).

**Distribution.**—Mexico (Chiapas) and Guatemala.

**19. Chiococca semipilosa** Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 22:279. 1940. **TYPE:** GUATEMALA. **CHIQUIMULA:** Caracol Mountain, 1.5 mi N of Quezaltepeque, 1200–1400 m, 7 Nov. 1939, *J.A. Steyermark 31406* (**HOLOTYPE:** F [No. 1039351]).

**Distribution.**—Mexico, Guatemala, and Honduras.

*Chiococca pubescens* var. *peninsularis* Wiggins, Contr. Dudley Herb. 4:24. 1950. **TYPE:** MEXICO. **BAJA CALIFORNIA:** Sierra de San Franciscoquito, 18 Oct 1890, T.S. *Brandegee 265* (**HOLOTYPE:** DS [No. 20437], barcode CAS0026743); **ISOTYPE:** UC [UC102610]).

*Chiococca pueblensis* Lundell, Wrightia 5:7. 1972. *Chiococca pubescens* Standl., Contr. U.S. Natl. Herb. 20:209. 1919, *nom illeg. superfl.* Non Humb. & Bonpl. ex Schult. (Syst. Veg. 5:202. 1819). **TYPE:** MEXICO. **PUEBLA:** San Luis Tamaulipas, 320 m, Jul 1908, C.A. *Purpus* 3334 (**HOLOTYPE:** US [No. 810975]; **ISOTYPES:** E [E00285117], F [No. 244045], G [G0038939]).

*Chiococca vestita* Lundell, Wrightia 5:8. 1972. **TYPE:** GUATEMALA. **PETEN:** Poptun, 2 km E, in hammock in pineland, 13 Jul 1959, *C.L. Lundell 16425* (**HOLOTYPE:** LL [00000273]; **ISOTYPES:** S [No. S10-24103]).

*Chiococca vestita* var. *glaberrima* Lundell, Wrightia 5:9. 1972. **TYPE:** GUATEMALA. **PETEN:** Poptun, 2 km E, in hammock in pineland, 13 Jul 1959, *C.L. Lundell 16425A* (**HOLOTYPE:** LL [00000274]; **ISOTYPES:** S [2 sheets, Nos. S05-998, S10-24114]).

*Chiococca gracilis* Borhidi, Acta Bot. Hung. 49:63. 2007. **TYPE:** GUATEMALA. **IZABAL:** El Estor, margin of open nickel mine, 410 m, 30 Aug 1988, W.D. Stevens & E. *Martinez* S. 2527 (**HOLOTYPE:** MEXU [MEXU 00490722]; **ISOTYPES:** MEXU [MEXU 01054938], MO [No. 4293149], PTBG [PTBG 1000000148]).
*Portlandia*, indicating that it is somewhat intermediate between the two genera. Aiello (1979) maintained *Coutaportla* as a monotypic genus. She discussed the identity of *Portlandia guatemalensis* Standl. and positioned it close to *Coutaportla*, without publishing the necessary new combination because of lack of mature fruits. Lorence (1986) transferred *P. guatemalensis* to *Coutaportla*, and Borhidi (2003) transferred it to the monospecific *Locencea* Borhidi, which is here followed (See below). *Coutaportla* is characterized by the interpetiolar stipules, triangular or deltoid, inflorescences terminal, uni- or pauci-florous, 4(5)-merous flowers, laterally compressed hypanthium, white, pink or purple corollas, 2-locular ovary, with (1–)2–5 ovules per locules, placenta subapical, medial or basal, capsules laterally compressed, loculicidal (secondarily septicidal), and seeds dorso-ventrally compressed, wingless (Torres-Montúfar, 2024; Torres-Montúfar et al. 2023). In this genus are here included five species of shrubs or small trees (0.1–)0.5–3 m tall, four of them occurring in Mexico, and one in Venezuela.

*Coutaportla* Urb., Symb. Antill. 9:146. 1923; Lorence, Syst. Bot. 11(1):209–213. 1986; Villareal, Sida 12(1):223–225, f. 1. 1987; Ochterena-Booth, Fl. Mesoamericana 4.2:69. 2012; Borhidi, Acta Bot. Hung. 60(1–2):31–45. 2018; Torres-Montúfar et al., Pl. Ecol. Evol. 156(1):5. 2023; Pío-León et al., Acta Bot. Mex. 130:e2167, f. 1–2. 2023; Torres-Montúfar, Acta Bot. Mex. 131:35–52. 2024. TYPE: *Coutaportla ghiesbreghtiana* Urb.

1. **Coutaportla ghiesbreghtiana** (Baill.) Urb., Symb. Antill. 9:147. 1923. *Portlandia ghiesbreghtiana* Baill., Adansonia 12:300. 1879. TYPE: MEXICO. OAXACA: Hda. Huijastla, 1842–43, A. Ghiesbrecht 27 (FIRST-STEP LECTOTYPE (Lorence 1999: 44), SECOND-STEP LECTOTYPE, here designated: P [P002273491]; ISOLECTOTYPES: F [No. 972609], G [G00436041, without collection number], P [P002273492], US [2 sheets, 00997912, 00137327, without collection number]).

**Distribution.**—Mexico (Oaxaca).

*Notes*.—Baillon (1879:300) described and discussed *Portlandia ghiesbreghtiana* Baill., but did not cite any collection or herbarium of deposit. Lorence (1999:44) cited the type of *Portlandia ghiesbreghtiana* as “[...] *Ghiesbrecht* 27 (Holotype P (photos MEXU, PTBG); Isotypes P, 3 sheets (photos MEXU), US (photos MEXU, PTBG)).” Because he did not specify which specimen in P is the “holotype,” Lorence’s citation is a first-step lectotypification. In P, where Baillon worked, there are four specimens of *Ghiesbrecht* 27. The specimen with barcode P002273491, has a label with the annotation “Portlandia Ghiesbreghtiana H. B. Sect. Coutaportla, Voy. Adansonia XII” handwritten by Baillon. This specimen is here designated as the second-step lectotype of this name.

The P specimen of *Ghiesbrecht* 27 with barcode P002273492, has a label with the annotation “Coutarea” handwritten by an unknown author, and “Ghiesbreghtiana H. B.” handwritten by Baillon; this specimen is an isotype. The P specimens of *Ghiesbrecht* 27 with barcodes P002273493 and P002273494, have labels with the annotation “Coutarea Ghiesbreghtiana H. B.” handwritten by Baillon; these specimens are isotypes.

2. **Coutaportla helgae** Pío-León, Torr.-Montúfar & H. Ávila, Acta Bot. Mex. 130:e2167, p. 3, figs. 1–2. 2023. TYPE: MEXICO: SINALOA: Mun. Cosalá, 600 m E of community Las Mimbres del Padre, 24°17′N, 106°46′W, 300 m, 17 Jan 2022, J.F. Pío-León & J. Beltran 289 (HOLOTYPE: CIIDIR n.v.; ISOTYPES: FESC n.v., HCIAD n.v., MEXU n.v., USON n.v.).

**Distribution.**—Mexico.

3. **Coutaportla campanilla** (DC.) Delprete, **comb. nov.** *Coutarea campanilla* DC., Prodr. 4:350. 1830. *Coutarea hexandra* var. *campanilla* (DC.) Steyerm., Mem. New York Bot. Gard. 23:297–299. 1972. TYPE: VENEZUELA: near Caracas, s.d., Vargas s.n. (HOLOTYPE: G-DC [2 sheets, G00665778 (one flower), G00665780 (two branches))].

**Distribution.**—Venezuela (between Caracas and Mérida).

*Notes*.—In G-DC there are two sheets associated with *Coutarea campanilla* DC., which are kept together. The sheet with barcode G00665778, with a single flower in an envelope, has a label with the annotation “Coutarea campanilla DC” handwritten by Candolle. The sheet with barcode G00665780 does not have a label handwritten by Candolle, and consists of two branches with numerous leaves, one of them with an inflorescence with flower buds, and the other with an infructescence with several dehisced capsules. On top of those branches is affixed a label with the annotation “Vulgó-campanilla-fortaspe [? Illegible] ad gen. cinch. pertinet Barbacoan ad May 26 se halla junto con la pubescens-palida” handwritten by Vargas. According to Art. 8.3 of the Code (Turland et al. 2018), because the two sheets are kept together, with Candolle's annotation
on the first sheet, they are here treated as a single specimen with multiple preparations, which is the holotype of *C. campanilla* DC.

*Coutaportla campanilla* (DC.) Delprete is similar to *Coutarea hexandra* because of its terminal inflorescences and the lenticellate stems; from which it differs by the actinomorphic, 4-merous flowers; while in *C. hexandra* the flowers are zygomorphic, curved, bilaterally symmetrical, and commonly 6-merous. *Coutaportla campanilla* differs from *Coutarea alba* by having terminal inflorescences, while in *C. alba* they are usually axillary or terminal on lateral short shoots. *Coutarea campanilla* is here transferred to *Coutaportla* based on floral characters. Ovaries and fruits are needed, to confirm if they have the same kind of placentation found in the other species of *Coutaportla*, which have 2-5 ovules per locule, basally inserted. The addition of this species to *Coutaportla* extends the geographical range of the genus to Venezuela, while the other species occur in Mexico.

*Coutarea lindeniana* Baill., Adansonia 12:300. 1879. TYPE: VENEZUELA. Entre Caracas et Merida, s.d. [1842], J.J. Linden 376 (HOLOTYPE: P [P00559103]; ISOTYPE: G [G00436038]).

**Notes.**—Baillon (1879:300) in the protologue of *Coutarea lindeniana* Baill., cited the gathering *Linden* 376 collected in Venezuela. In P there is a specimen that has a label with the annotations “Coutarea” handwritten by an unknown author, and “Lindeniana H. B.” handwritten by Baillon. The P specimen is the holotype of this name. The specimens of *Linden* 376 present in P and G have small, 4-merous, actinomorphic flowers.

4. **Coutaportla lorenceana** Torr.-Montúfar, Ochot.-Booth & Art. Castro, Pl. Ecol. Evol. 156(1):5. 2023. TYPE: MEXICO. SINALOA: Mun. Concordia, El Palmito, alrededores del acceso principal al Santuario Chara Pinta; 23.56444°N, 105.848882°W, 1980 m, 10 Sep 2019, A. Castro-Castro, Avila-González & Zavala-Pérez 4532 (HOLOTYPE: MEXU n.v.; ISOTYPES: CIIDIR n.v., FCME n.v., FESC n.v., IEB n.v., IBUG n.v., MO n.v., PTBG n.v., SLPM n.v.).

**Distribution.**—Mexico.

*Chiococca grandiflora* Lorence & T.Van Devender *pro parte*, in D.H. Lorence, T.R. Van Deventer & G.M. Ferguson, Phytokeys 98:74, figs. 1-3. 2018. *PARATYPES* cited in Lorence et al. (2018:76): *T. Walker s.n.* (ARIZ [212520]), *S. Walker s.n.* (UTC [00263027], ARIZ [181630]); *S. Walker* 70043 (K).

5. **Coutaportla pailensis** Villareal, Sida 12:223, fig. 1. 1987. TYPE: MEXICO. COAHUILA: Cerro de Paila, Cañon de Loma Prieta, NW de Hipólito, 25°51'N, 101°30'W, 6 Aug 1986, J.A Villareal, M.A. Carranza & J. Whebe 3305 (HOLOTYPE: MEXU [MEXU 00424882]; ISOTYPES: ANSM n.v., ENCB [ENCB003687], TEX [00000020]).

**Distribution.**—Mexico.

## COUTAPORTLA EXCLUDED TAXA

*Coutaportla guatemalensis* (Standl.) Lorence, Syst. Bot. 11:210. 1986. *Portlandia guatemalensis* Standl., J. Wash. Acad. Sci. 18:162. 1928.
= *Lorencea guatemalensis* (Standl.) Borhidi, Acta Bot. Hung. 45:17. 2003 (See below under *Lorencea guatemalensis*).

## COUTAREA

Aublet (1775:314) published the genus *Coutarea* Aubl., with the sole species *C. speciosa* Aubl. [= *C. hexandra* (Jacq.) K. Schum.]. He supplied the French vernacular name “Coutar de la Guiane,” suggesting that the generic name was of indigenous origin, which was probably “coutar,” “kutar” or something similar. The position and delimitation of *Coutarea* has a complex history. Ochterena-Booth (1994), completed a master thesis on the revision of *Coutarea*, which remained unpublished. A generic and specific description, with extensive synonymy, was published by Delprete (1999b, 2010, 2024). The genus is unique within the Chiococcaceae by the following combination of characters: shrubs or trees to 15 m tall (to 30 m tall in the Amazon Basin), young branches commonly lenticellate, inflorescences terminal (terminal on lateral short-shoots in *C. alba*), in simple or compound dichasia, flowers 6-merous, zygomorphic, stamens well-exserted, unequal, stigmatic portion in two lines along the style, capsules strongly laterally compressed, seeds large, winged, vertically imbricate. In the present treatment, two species are recognized in this genus, ranging from Mexico to Argentina.

*Coutarea* Aubl., Hist. Pl. Gui. Franc. 1:314, pl. 122. 1775; Standley, N. Amer. Fl. 32(2):126-128. 1921; Steyermark, Mem. New York Bot. Gard. 23:297-299. 1972; Steyermark, in Lasser & Steyermark, Fl. Venezuela 9(1):203-209, f. 29. 1974; Delprete in Andersson, Fl. Ecuador 62:44-50, f. 15-16. 1999; Borhidi, Rubiáceas México 145, 147, f. 29. 2006; Borhidi, Rubiáceas México, 2nd Ed 158, 160, f. 33. 2012;
Ochoterena-Booth, Fl. Mesoamericana 4.2:69-70. 2012; Aguirre Arbo & Salas in Zuloaga & Zanotti, Fl. Argentina 19(3):248-250, 2 figs. 2022. TYPE: *Coutarea speciosa* Aubl. [= *Coutarea hexandra* (Jacq.) K. Schum.].

1. **Coutarea alba** Griseb., Abh. Königl. Ges. Wiss. Göttingen 24:153. 1879. TYPE: ARGENTINA. JUJUY: [Depto. Ledesma], San Lorenzo, Nov. 1873, P.G. Lorenz & G.H.W. Hieronymus 217 (LECTOTYPE (Aguirre Arbo & R.M. Salas in F.O. Zuloaga & C.A. Zanotti, Fl. Argentina 19(3):248. 2022): CORD [CORD00006240 & CORD00006241]; ISOLECTOTYPES: E [00514966], G [G00436040]).

**Distribution.**—Venezuela, Bolivia, Brazil, Paraguay, Argentina (Salta, Jujuy).

2. **Coutarea hexandra** (Jacq.) K. Schum, Fl. Bras. 6(6):196. 1889. *Portlandia hexandra* Jacq., Enum. Syst. Pl. 16. 1760 [Jacq., Select. Stirp. Amer. Hist. 182, f. 20. 1763]. TYPE: COLOMBIA. BOLIVAR: vicinity of Cartagena, s.d., Jacquin s.n. (LECTOTYPE (Delprete 1999b:47): BM [000081660, one flower]).

**Distribution.**—From Mexico to Argentina (absent in the Greater Antilles).

*Coutarea speciosa* Aubl., Hist. Pl. Guiane 1:314, t. 122. 1775. *Portlandia speciosa* (Aubl.) Baillon, Hist. Pl. 7:381. 1880. *Coutarea hexandra* var. *β speciosa* K. Schum. in Martius et al., Fl. Bras. 6(6):197. 1889. TYPE: FRENCH GUIANA: "Caienne & in sylvis Guianae," s.d., Aublet s.n. (NEOTYPE (Delprete 2015b:603): P-JU [No. 9927a, branch with flowers]; ISONEOTYPE: P-JU [No. 9927b, fruits and seeds]).

*Portlandia acuminata* Willd., in Roemer & Schultes, Syst. Veg., ed. 15[bis], 5:23. 1819. TYPE: VENEZUELA: Caracas, s.d., F. Bredemeyer s.n. (HOLOTYPE: B-W [B -W 03932 -01 0]).

*Exostoma souzanum* Mart., in Martius & Spix, Reise Bras. 2:789. 1828. *Cinchona souzana* (Mart.) Brign., Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis., ser. 2, 1:63. 1862. TYPE: Not found.

*Coutarea flavescens* DC., Prodr. 4:350. 1830. TYPE: [icon.]: [MEXICO]. Torner Collection Plate No. 0734, in the Hunt Institute for Botanical Documentation (LECTOTYPE, McVaugh 2000: 463).

**Notes.**—Augustinus Pyramus de Candolle (1830:350) in the protologue of *Coutarea flavescens* DC., stated "C. latiflora (fl. mex. ic. ined.) pediculis pluribus sub floribus ebracteatis, floris diametro longitudine dimido ferè minore.— In Mexico. Folia superiora interdum terna, ovalia basi attenuata. Pediculi ternato-verticillati bis bifidi ad ramificationes tautum bracteati. Flores flavescentes minores et praesertim angustiores quam priorum." Therefore, the authority of this name should be attributed solely to him.

There is no original specimen of *Coutarea flavescens* in G, G-DC, or MA. McVaugh (1969, 1972, 1977, 1980, 1982, 1987, 1990, 1998, 2000), discussed in great detail the history, itinerary, herbarium, and illustrations of the Sessé & Mociño Expedition (1781-1803), which is below summarized under *Coutarea latiflora* DC. [= *Hintonia latiflora* (DC.) Bullock].

McVaugh (2000:463) cited the type of *Coutarea flavescens* as follows, "Lectotype: No. 0734 of the Torner Collection, annotated "Coutarea flava" by Alphonse de Candolle. DC. plate 459, as cited in Calques des Dessigns (Field Mus. neg. 30688), is an incomplete copy of Torner 0734. It is annotated "Coutarea flava" and, by Alphonse de Candolle, "Sans doute C. flavescens DC."

Ochoterena-Booth (2012:119) cited the type of "*Coutarea latiflora* Sessé et Moc. ex DC." as "Holotipo: México, Michoacán, Sessé y Mociño s.n. (G-DC!)." However, as explained above, the authority of this name should be attributed solely to Candolle, and there is no original specimen associated with this name in G-DC.

*Coutarea pubescens* Pohl, Pl. Bras. Icon. Descr. 2:148, tab. 200. 1833. *Coutarea hexandra* var. *e pubescens* (Pohl) K. Schum. in Mart., Fl. Bras. 6(6):198. 1889. *Coutarea hexandra* f. *pubescens* (Pohl) Steyerm., Mem. New York Bot. Gard. 23:298. 1972. TYPE: BRAZIL: without locality, s.d., J.B.E. Pohl s.n. (LECTOTYPE, here designated: M [M-0187237]).

*Coutarea mollis* Cham., Linnaea 9:259. 1834. TYPE: BRAZIL: without locality, s.d., C.F.P. Martius Herb. Fl. Brasil. No. 995 (NEOTYPE, here designated: M [M-0187239]).

**Notes.**—Chamisso (1834:259), in the protologue of *Coutarea mollis* Cham., cited the material studied as "E Brasilia aequinotialis misit Sellow" without citing the herbarium of deposit. A general search in numerous herbaria did not retrieve any original specimen associated with this name. Therefore, a neotype needs to be designated. A specimen in M, with barcode M-0187239, has the handwritten annotations "Coutarea pubescens Pohl, Martii Herbar. Florae Brasil. N° 995." The specimen consists of a ramified branch with lenticellate stems, numerous leaves, and terminal inflorescences with flowers in anthesis. This specimen is here designated as the neotype of *C. mollis*.

### Notes about *Bignonia triflora* Vell. (*Bignoniaceae*)

*Bignonia triflora* Vell., Fl. Flum. 249. 1829 ["1825"). TYPE: [icon. ined.] "Didyn. Angiosp. BIGNONIA trifolia [triflora] Tab. 38," Divisão de Manuscritos, Biblioteca Nacional, Rio de Janeiro, No. mss1198655_042 (LECTOTYPE, Nascimento et al. 2024).

**Notes.**—Delprete (1999b:47), treated "*Bignonia triflora* Pav. ex DC." (1830:148) as a synonym of *Coutarea hexandra*, and cited the type as "Type: Pavón s.n., Peru, Dept. Loreto (MA, not seen)." Alphonse de Candolle (1845:148), in the protologue of *Bignonia triflora*, wrote: "22. B. TRIFLORA (Vell. fl. flum. 6. t. 38), scandens, ramis teretibus; foliis conjugatis, petiolo simpliciter cirrhoso, foliolis ovatis acuminatis basi obtusis, petiolulis petiolo duplo brevioribus, pedunculis axillaribus brevissimis pedicellos 3 elongatos gerentibus, calyce campanulato-oblongo 5-dentato, corolle tubo obconico, lobis subrotundis patentibus, stamimum filamentis villosis. in Brasilis prov. Fluminensi. Caet. ign. B. triflora Pav! herb. est Rubiacea quaedam." Hence, he treated *B. triflora* as a species of *Bignonia*, in the family Bignoniaceae, and attributed the species name to Vellozo. The phrases following the description can be broken into the following: 1. "in Brasilis prov. Fluminensi."—in the Brazilian province of Rio de Janeiro; 2. "Caet. ign."—
*Distribution*.—Ecuador (Azuay).

**3. Coutareopsis fuchsioides** (C.M. Taylor) Paudyal & Delprete, Bot. J. Linn. Soc. 187:386. 2018. *Coutarea fuchsioides*
C.M. Taylor, Novon 20:101. 2010. TYPE: PERU. LA LIBERTAD: Prov. Pataz, rd. (Huamachuco to) Chagagua-Pataz, after Chagual on
steep slope up the Cordillera, 7°48'S, 77°37'W, 1799 m, 21 Apr 2004, M. Weigand & C. Schwarzer 7917 (HOLOTYPE: USM n.v.; ISOTYPES:
BSB n.v., M [M-0210026], MO [No. 6145369]).

*Distribution*.—Peru (La Libertad).

CUBANOLA

Aiello (1979) transferred two species of *Portlandia*, one endemic to Cuba, and the other to Dominican Republic, to the newly founded *Cubanola*. She stated that "The name of the genus derives from the fact that one of the two species is endemic to Cuba and the other to Hispaniola." She distinguished *Cubanola* from *Portlandia* by its reticulate-foveate seeds (vs. tuberculate in *Portlandia*), non-persistent funicle (vs. persistent), loculicidal and septicidal capsule (vs. loculicidal), placenta circular in cross section (vs. linear and adnate to septum), and chartaceous to subcoriaceous leaves (vs. coriaceous). The separation of *Cubanola* as a distinct genus was supported by the molecular phylogenetic trees of Motley et al. (2005), Manns and Bremer (2010), and Paudyal et al. (2018).

**Cubanola** Aiello, J. Arnold Arbor. 60:111. 1979; Standley, N. Amer. Fl. 32(1):13. 1918; Liogier, Fl. Cuba 5:27. 1962; Liogier, Fl. Española 7:241, 243, f. 198-9. 1995; Borhidi, Rubiáceas Cuba 87, f. 22. 2017. TYPE: *Cubanola daphnoides* (R. Graham) Aiello

*Gonianthes* A. Rich., in Sagra, *nom. superfl.*, Hist. Cuba 11:10. 1850. TYPE: *Gonianthes sagrana* A. Rich. [= *Cubanola daphnoides* (R. Graham) Aiello]

Non *Gonianthes* Blume (1823) (Burmanniaceae).

1. **Cubanola daphnoides** (R. Graham) Aiello, J. Arnold Arbor. 60:112. 1979. *Portlandia daphnoides* R. Graham, Edinb. New Philos. J. 30:206. 1841. TYPE: CUBA. MATANZAS: Yumury River Valley, near Matanzas, 1849, F.I.X. *Rugel* 374 (FIRST-STEP NEOTYPE (Aiello 1979: 112), SECOND-STEP NEOTYPE, here designated: NY [00126755]; ISONEOTYPES: BM [2 sheets, 000081662], GH [00058978], K [K000173488], L [2 sheets, L 0000272, L 0000273], NY [3 sheets, 00126752, 00126753, 00126754]).

*Distribution*.—Cuba (Guantánamo, Holguín, Villa Clara, Matanzas, Habana, Pinar del Río).

*Notes*.—Graham (1841:206–207), in the protologue of *Portlandia daphnoides* R. Graham, wrote that the plant is native of Cuba, but did not cite any specimen or herbarium of deposit. Aiello (1979:112) cited the type of *P. daphnoides* as "NEOTYPE: Cuba, Matanzas, Yumury Valley, near Matanzas, *Rugel* 374 (NY!)." There are four sheets of *Rugel* 374 at NY. Aiello did not indicate which of those sheets is the neotype. Hence, Aiello's (1979:112) citation is a first-step neotypification. The NY sheet with barcode 00126755 has a label with the annotations "*Cubanola daphnoides* (R. Grah.) Aiello. NEOTYPE: *Portlandia daphnoides* R. Grah. Edinb. N. Phil. Journ. 30. 206. 1840. LECTOTYPE: *Portlandia longiflora* Meiss. ex Griseb. Pl. Cuba 126. 1866" type-written by Aiello [underlines by Aiello]. According to the *Code* (Turland et al. 2018; e.g., Art. 30.1), an annotation affixed on a specimen does not count as effective publication. The NY specimen with barcode 00126755 is here designated as the second-step neotype of *P. daphnoides*.

*Gonianthes sagrana* A. Rich., in Sagra, Hist. Fis. Cuba, Bot. 11:10. 1850 (as "*sagreana*"). TYPE: CUBA: without locality [Havana], s.d., *Ramon de la Sagra* s.n. (FIRST-STEP LECTOTYPE (Aiello 1979:112), SECOND-STEP LECTOTYPE, here designated: P [P02273495]; ISOLECTOTYPES: P [2 sheets, P00582018, P02273496]).

*Notes*.—Achille Richard (in Sagra, 1850:10) in the protologue of *Gonianthes sagrana* A. Rich. (as "*sagreana*"), cited two gatherings as "Crescit in ruderais circa *Havanam*, mense maio florens (*Ramon de la Sagra*), et prope Santiago (*Linden*)." Aiello (1979:112) cited the type of *G. sagrana* as "TYPE: Cuba, without further locality, Sagra s.n. (holotype, P!)." In P there are three sheets collected by de la Sagra, annotated with this name, hence Aiello's citation is a first-step lectotypification. The P specimen with barcode P02273495 has a label with the heading "Herbarium Richard" and the footnote "Legit Ramon de la Sagra," both handwritten in red ink, and "Gonianthes Sagraeana Nob." handwritten by Achille Richard in black ink. This specimen has another label with the typewritten annotation "Cubanola daphnoides (R. Grah.) Aiello. Holotype: Gonianthes sagraeana A. Rich. Annette Aiello 1977." The specimen consists of three branches, one with a pair of flower buds, one with a pair of flowers in anthesis, and a small branch with a pair of very young flower buds. This specimen is here designated as the second-step lectotype of *G. sagrana*.

The P specimen with barcode P00582018 has a label the printed heading "HERBIER E. DRAKE" and the annotation "Cuba, Legit Ramon de la Sagra" handwritten by an unknown author. This specimen is an isolectotype.
The P specimen with barcode P02273496 has a label with the heading "Herbarium Richard" and the footnote "Legit Ramon de la Sagra," both handwritten in red ink, and "Gonianthes sagraeana Rich" handwritten in black ink by an unknown author (not Achille Richard). This specimen is an isolectotype.

*Portlandia longiflora* Meisner ex Griseb., Cat. Pl. Cub:126. 1866. **TYPE:** CUBA. MATANZAS: Yumury River Valley, near Matanzas, 1849, F.I.X. Rugel 374 (FIRST-STEP LECTotype (Aiello 1979:112), SECOND-STEP LECTotype, here designated: NY [00126755]; ISOLECTOTYPES: BM [2 sheets, 000081662], GH [00058978], K [K000173488], L [2 sheets, L 0000272, L 0000273], NY [3 sheets, 00126752, 00126753, 00126754]).

**Notes.**—Grisebach (1866:126) in the protologue of *Portlandia longiflora* Meisner ex Griseb., cited two gatherings as "Cuba occ., in praeruptis as Yumury pr. Matanzas (Rug. [Rugel] 374); Cuba or. et occ. (Wr. [Wright] 2678)." Aiello (1979:112) cited the type of *P. longiflora* as "TYPE: Cuba, Matanzas, Yumury Valley, near Matanzas, Rugel 374 (lectotype, NY!)." As explained above (under *Gonianthes sagrana*), there are four sheets of Rugel 374 at NY. Because Aiello did not indicate which of those sheets is the lectotype, her citation is a first-step lectotypification of *P. longiflora*. The NY specimen with barcode 00126755 is here designated as the second-step lectotype of this name.

**2. Cubanola domingensis** (Britton) Aiello, J. Arnold Arbor. 60:113. 1979. *Portlandia domingensis* Britton, Bull. Torrey Bot. Club 41:24. 1914. **TYPE:** DOMINICAN REPUBLIC. SAN PEDRO DE MACORÍS: San Pedro de Macorís, 26 Mar 1913, J.N. Rose, W.R. Fitch & P.G. Russell 4176 (LECTotype, here designated: NY [00126745]; ISOLECTOTYPES: GH [00058982], NY [00126744], US [00137325]).

**Distribution.**—Dominican Republic (La Altagracia, San Pedro de Macorís, La Romana).

**Notes.**—Britton (1914:24), in the protologue of *Portlandia domingensis* Britton, cited the material studied as "Near San Pedro de Macorís, Santo Domingo, March 26, 1913 (Rose, Fitch & Russell 4176)" without citing the herbarium of deposit. Aiello (1979:113) cited the type of *P. domingensis*, as "TYPE: Dominican Republic, [...], Rose et al. 4176 (holotype, NY! isotypes, GH!, NY!, US!)." As stated by Aiello, there are two specimens of Rose et al. 4176 in NY. Because she did not specify which of the two NY specimens is the holotype, her citation cannot be treated as a lectotypification.

The NY sheet with barcode 00126745, has a label with the annotation "Portlandia dominguensis Britton, n.s." handwritten by Britton. It has a second label with the annotations "Cubanola dominguensis (Brit.) Aiello. Holotype: Portlandia dominguensis Britt. Bull. Torr. Bot. Club 41: 24 (1914)" typewritten by Aiello in 1977. The specimen consists of two branches with several undehisced capsules. This specimen is here designated as the lectotype of *Portlandia dominguensis*.

The NY sheet with barcode 00126744, has a label with the annotation "Portlandia dominguensis Britton" handwritten by Britton. The specimen consists of two branches with a few undehisced capsules. This specimen is an isolectotype.

## EOSANTHE

Urban (1923a:162–163) described *Eosanthe* Urb., as a genus belonging to the Rubiaceae, but did not position it in any tribe. He derived the generic name from the Greek, *Eos* (ήώς), the Greek goddess of the dawn, and anthers (female form of anthos, ἀνθος), flower, referring to the orange calyx lobes of this genus. Robbrecht (1988, 1994) treated it as a genus *incertae sedis*. Delprete (1999a), based on morphological observations, indicated that *Eosanthe* is similar to *Phialanthus* because of the sheathing stipules, axillary inflorescence, persistent four-lobed calyx, filaments not connate to the corolla tube. Delprete (1999a:220) also wrote "*Eosanthe* resembles some species of *Schmidtottia* (Catesbaeeae sensu Delprete 1996) in having sheathing stipules, ovate coriaceous leaves, resinous branches, foliose calyx lobes, and linear-oblong anthers. The species of *Schmidtottia* with sheathing stipules and ovate thick-coriaceous leaves that resemble those of *Eosanthe* are *S. sessilifolia* (Britton) Urb. and *S. elliptica* (Britton) Urb. *Schmidtottia* differs from *Eosanthe* in having terminal few-flowered inflorescences (vs. solitary axillary flowers) and many-seeded capsules (vs. two-seeded pseudosamaras). [...] *Eosanthe* is tentatively included in the *Catesbaeeae-Chiococceae* complex." Manns and Bremer (2010), following their molecular phylogenies, included *Eosanthe* in the *Chiococceae*. The molecular phylogenies of Paudyal et al. (2018) are in agreement with Delprete's morphological observations, and Manns and Bremer's (2010) molecular phylogenies, and confirmed its position within the *Chiococceae*. *Eosanthe* is an extremely rare, monotypic genus, endemic to eastern Cuba (Sierra de Cristal, 1200–1300 m), with foliaceous, bright
orange-red calyx lobes, yellow corolla, and fruits that are narrowly winged pseudo-samaras (due to the persistent calyx lobes), which is a unique fruit type in the tribe. A full description of genus and species is available in Delprete (1999a).

**Eosanthe** Urb., Symb. Antill. 9:162. 1923; Delprete, Brittonia 51:217-230. 1999; Standley, N. Amer. Fl. 32(4):298. 1934; Liogier, Fl. Cuba 5:98. 1962; Borhidi, Rubiáceas Cuba 97-98. 2017. **TYPE:** *Eosanthe cubensis* Urb.

**1. Eosanthe cubensis** Urb., Symb. Antill. 9:162. 1923. **TYPE:** CUBA. HOLGUÍN: Sierra del Cristal, prope acumen, 1200-1300 m, 8 Mar 1916, E.L. Ekman 6816 (LECTOTYPE (Delprete 1999a:229): S [No. S-R-7947]).

**Distribution.**—Cuba (Holguín: Sierra del Cristal).

**Notes.**—Urban (1923a:162-163), in the protologue of *Eosanthe cubensis* Urb., cited the gathering *Ekman 6816*, without indicating the herbarium of deposit. The original material at B studied by Urban was destroyed during WWII. Delprete (1999a:229) designated the lectotype of this name as "Type. CUBA. Holguín: Sierra del Cristal, [...], Ekman 6816 (HOLOTYPE: B-destroyed; LECTOTYPE, here selected: S)." In S there is a specimen of *Ekman 6816*, with accession number S-R-7947, that has a label with the annotation "Eosanthe cubensis Urb (gen. nov., spec. nov.)" handwritten by Urban. The specimen consists of two branches with several coriaceous leaves, and two separate branch tips, with terminal inflorescences, and conspicuous, orange calyx lobes. This is the specimen that was designated by Delprete (1999a:229) as the lectotype of *E. cubensis*.

*Eosanthe cubensis* is an extremely rare species. To my knowledge, aside from the type gathering, it is known only by one additional collection from the same locality: CUBA. Sierra del Cristal, 1100-1325 m, 15 Dec 1922 (young fr), E.L. Ekman 15990 (HAC [2 sheets]).

## ERITHALIS

Patrick Browne (1756:165, t. 17, fig. 3) described and illustrated the genus *Erithalis* P. Browne, in *The Civil and Natural History of Jamaica*, without citing a species. The generic name is derived from the Greek "erithales" (ἐριθαλές), which means "herb that grows on walls." Linnaeus (1759:930) published the species *Erithalis fruticosa* L., citing Browne's description and illustration. The molecular phylogenies of Negrón-Ortiz and Watson (2002, 2003), Motley et al. (2005), and Manns and Bremer (2010) proved that this genus belongs to the Chiococcaceae. Paudyal et al.'s (2018) molecular phylogenies confirmed its position in the tribe, and resolved *Erithalis* as monophyletic, corroborating previous results. *Erithalis* is characterized by being shrubs or small trees, with interpetiolar stipules, connate at base, inflorescences axillary, paniculate, corymbose, racemose, or uniflorous, corollas actinomorphic, hypocrateriform, narrowly infundibular or rotate, stamens inserted at the base of corolla tube, style branches 2 or 5-8, ovaries with 2-5 locules, 5-20-celled, and drupaceous fruit with woody pirenes. The genus was revised by Negrón-Ortiz (2005), and her species delimitations are here adopted, except for *E. insularis* (Ridl.) D. Zappi & T.S. Nunes, which is here recognized as a distinct species. Another species, *E. orbiculata* (Proctor) A.R. Franck, P.A. Lewis & Oberli, from Jamaica, was recently transferred to this genus (Franck et al. 2017). *Erithalis*, as here delimited, is a genus of 10 species, exceptional in the Chiococcaceae by having 5-20-celled drupes, widespread in the Caribbean region (including Caribbean coast of Venezuela), and one species endemic to Fernando de Noronha Island (northeastern Brazil).

**Erithalis** P. Browne, Civ. Nat. Hist. Jamaica 165, t. 17, fig. 3. 1756; Standley, North Amer. Flora 32(4):279-281. 1934; Liogier, Fl. Cuba 5:89-90. 1962; Correll & Correll, Fl. Bahama Arch. 1386-1388, f. 605. 1982; Liogier, Fl. Española 7:250, 252-253, f. 198-213. 1995; Negrón-Ortiz & Watson, Pl. Syst. Evol. 234:71-83. 2002; Negrón-Ortiz & Watson, Syst. Bot. 28(2):442-451. 2003; Negrón-Ortiz, Sida 21:1565-1598. 2005; Lorence, Fl. Mesoamericana 4.2:85-86. 2012; Borhidi et al., Rubiáceas Cuba 98-102, f. 26. 2017. **TYPE:** *Erithalis fruticosa* L.

**1. Erithalis angustifolia** DC., Prodr. 4:465. 1830. **TYPE:** [LESSER ANTILLES (See notes below)]: Collector unknown s.n. (HOLOTYPE: G-DC [G00666855]).

**Distribution.**—Lesser Antilles (Martinique, St. Vincent and the Granadines), and Trinidad.

**Notes.**—In G-DC there is a sheet, with barcode G00666855, which has the annotation "Erithalis angustifolia DC." handwritten by Candolle. The specimen consists of a small branch with numerous leaves, flowers
in anthesis, and mature fruits. At the base of the branch is pinned a label with the annotations "Rubiaceae. La Havane. Mr de la Ossa 1825" handwritten by an unknown author. The latter annotation is incorrect, as this species only occurs in the Lesser Antilles and Trinidad. This specimen is the holotype of this name.

*Erithalis acuminata* Krug & Urb., Notizbl. Bot. Gart. Berlin-Dahlem 1:319. 1897. **TYPE:** MARTINIQUE: Guidling, 1877, *P. Duss* 206 (LECTOTYPE; Negrón-Ortiz 2005:1582); NY [00743699]).

**Notes.**—Krug and Urban (1897:319), in the protologue of *Erithalis acuminata* Krug & Urb., cited the material studied as "Habitat in Martinique, solummodo in Montagne-Pelée 900–1000 m alt., sed ibidem satis frequens: Duss n. 206, 937, 1724; St. Vincent: Guidling."

Negrón-Ortiz (2005:1582), for *Erithalis acuminata*, cited the type as "Martinique, Guidling, 1877, *Duss* 206 (holotype: B, destroyed; Lectotype: NY!; here designated)." In NY there is a sheet of *Duss* 206, with barcode 00743699, which has a label with the printed annotations "Lectotype. Erithalis acuminata Kr. and Urb. Notizbl. Königl. Bot. Gart. Berlin. 1:319–320. 1897 [...] Det.: V. Negrón-Ortiz in 2004." The specimen consists of a small branch with numerous leaves, and several infructescences with mature fruits. This is specimen that was designated as the lectotype of *E. acuminata* by Negrón-Ortiz.

In NY there is another specimen, with barcode 00115111, which has a label with the annotation "No. 206, 937, 1724, Erythalis angustifolia Griseb. (non DC), "acuminata Kr. et Urb. n. sp. [...] Montagne Pelée/versant du fond Coré, [...] 900–1000, 1877, 1879, 1883." It is apparent that the annotations on this label simply reproduced the information of the material cited Krug and Urban (1897:319). Hence, this specimen is dubious original material.

*Chiococca pulcherrima* Wernham, J. Bot. 51:322. 1913. **TYPE:** ST. VINCENT: without locality, s.d., *Anderson* 308 (HOLOTYPE: BM [000551620]; ISOTYPE: K n.v.).

2. *Erithalis diffusa* Correll, J. Arnold Arbor. 58:47. 1977. **TYPE:** BAHAMAS: San Salvador, N of airstrip and Teachers College, near Riding Rock, 21 Nov 1974, *D.S. Correll* 43840 (HOLOTYPE: A [00092883]; ISOTYPES: F [No. 1784095], FTG n.v., NY [00115113]).

**Distribution.**—Bahamas (San Salvador and Crooked Islands).

3. *Erithalis fruticosa* L., Syst. Nat., Ed. 10, 2:930. 1759. **TYPE:** [icon.]; Tab. 17, Fig. 3 in P. Browne, Civ. Nat. Hist. Jamaica, vol. 1. 1756 (LECTOTYPE, here designated).

**Distribution.**—Southern USA (Florida), Bahamas, Greater and Lesser Antilles, Trinidad, Mexico (Veracruz, Campeche, Quintana Roo), Venezuela (Caribbean coast and Caribbean Islands), Colombia (Santa Catalina Island, Caribbean region).

*Notes.*—Jarvis (2007:504) regarding the type of *Erithalis fruticosa* L., wrote "Negrón-Ortiz (in Sida 21:1584) wrongly refers to the cited Browne plate as the holotype (there is no original material in S). Although this could be corrected to lectotype under Art. 9.8, the absence of "designated here" with the type statement (Art. 7.11) means that this cannot be accepted as a formal typification (See Art. 9.8, Note 4)." Following Jarvis's observations, Tab. 17, Fig. 3 of P. Browne's *Civil and Natural History of Jamaica* (1756), is here designated as the lectotype of this name.

*Erithalis odorata* Pers., Syn. Pl. 1:200. 1805, nom. alt.

*Erithalis inodora* Jacq., Select. Stirp. Amer. Hist.:73. 1763. *Erithalis fruticosa* var. inodora (Jacq.) Persoon, Syn. Pl. 1:200. 1805. **TYPE:** LEEWARD ISLANDS. Dominica: Forest bluff into sea near Anse Du Me on the northern coast, ca. 7 mi E of Portsmouth, 11 Aug 1964, R.L. *Wilbur*, E.L. *Dunn*, H.A. *Hespenheide* & D.R. *Wiseman* 8304A (*NEOTYPE*, here designated: MO [No. 4626927, barcode MO-1432073]).

*Notes.*—Jacquin (1763:73), in the protologue of *Erithalis inodora* Jacq., cited the material studied as "Habitat passim in Curação" without citing the herbarium of deposit. As already reported by Negrón-Ortiz (2005:1585), I was also unable to locate any original specimen. Therefore, a phenotype needs to be designated. The specimen *Wilbur et al.* 8304A in MO, with accession number 4626927 (barcode MO-1432073), is here designated as the neotype of *E. inodora*.

*Erithalis elliptica* Raf., Sylva Tell. 123. 1838. **TYPE:** Based on Swartz's description of *E. fruticosa*.

*Erithalis parviflora* Griseb., Cat. Pl. Cub. 134. 1866. **TYPE:** CUBA: without locality, 1860–1864, Wright 2721 (HOLOTYPE: GOET [GOET010231]; ISOTYPES: K [K000173942], MO [No. 2092104], NY [00743698], S [No. S05-983], US [00138504], YU [YU.065584]).

*Erithalis harrisii* var. angusta S. Moore ex Rendle, J. Bot. 73:279. 1935. **TYPE:** JAMAICA: Luana Point, collection date unknown, *Harris* 9821 (HOLOTYPE: BM n.v.).

*Erithalis revoluta* Urb., Symb. Antill. 3:379. 1903. **Type:** PUERTO RICO: Guánica, Caño Gordo, s.d. [13 Feb 1886], *P. Sintenis* 3796 (LECTOTYPE, here designated: F [No. 189202]).

*Notes.*—Urban (1903:379) in the protologue of *Erithalis revoluta* Urb., cited the material studied as "Portorico in sylvis litoralibus ad Caño Gordo, m. Febr. de flor.: Sintenis n. 3796." The original material at B studied by Urban was destroyed during WWII. There is a specimen in F, accession number 189202, which has a label with the heading "P. SINTENIS: PLANTAE PORTORICENSES," the annotations "3796. Erithalis revoluta Urb. (typus). Prope Guanica in sylvis litoralibus ad Caño Gordo, 13.II. 1886," and the
watermark “Herb. Krug et Urban.” At the bottom of the label is printed “det. I. Urban.” This specimen is here designated as the lectotype of *E. revoluta*.

4. **Erithalis harrisii** Urb., Symb. Antill. 5:514. 1908. **TYPE:** JAMAICA: near Troy, 30 Jun 1904, W.H. Harris 8743 (LECTOTYPE (Lebrón-Ortiz 2005:1588): NY [00115115]; ISOLECTOTYPES: E [E00285119], NY [00115114], US [00138503]).

**Distribution.**—Jamaica.

5. **Erithalis insularis** (Ridl.) D. Zappi & T.S. Nunes, Kew Bull., 55(3):655. 2000. *Palicourea insularis* Ridl., J. Linn. Soc. 27:41. 1890. *Chiococca insularis* (Ridl.) C.M. Taylor & M.R.V. Barbosa, Phytotaxa 202:18. 2015. **TYPE:** BRAZIL. PERNAMBUCO: Island of Fernando de Noronha, 26 Aug 1887, H.N. Ridley, T.S. Lea & G.A. Ramage 86 (HOLOTYPE: BM [BM000949953]; ISOTYPES: GH [00094899], K [2 sheets, K000016495, K000016496]).

**Distribution.**—Brazil, Island of San Fernado de Noronha.

**Notes.**—Ridley (1890:41), in the protologue of *Palicourea insularis* Ridl., cited the material studied as “Main Island, in the Sapate woods, only a few bushes at one spot,” without citing the herbarium of deposit. Most importantly, he described the fruit as “Drupa viridis, 5-angulata, 5-loculis.” Stafleu and Cowan (1983:786) regarding Ridley’s collections specified that his gatherings from Fernando de Noronha are in BM. Negrón-Ortiz (2005:1585) regarding the type of *P. insularis*, wrote “Type: BRAZIL. Pernambuco: Fernando de Noronha Island, 1887, *Ridley, Lea & Ramage* 86 (holotype BM-n.v., not at BM per BM staff); lectotype: K! here designated; isotype B-destroyed, photo NY!).” The holotype specimen was later found at BM, barcode BM000949953, as reported by Figueira et al. (2020).

The generic position of this species has a complex history. Rydley (1890) initially described it as a species of *Palicourea*. Zappi and Nunes (2000) transferred it to *Erithalis*, proposing the new combination *E. insularis* (Ridl.) D. Zappi & T.S. Nunes, and stated that it “differs from all other species of the genus by its poorly branched, few-flowered inflorescences, which remain hidden by the surrounding leaves. The leaves are distributed along the new branches, while most *Erithalis* tend to have leaves grouped distally on the branches, resembling species of *Terminalia* (Combretaceae).” The characters used by those authors to transfer this species to *Erithalis* are not really distinctive from other species of the genus, for example, in many *Erithalis* species the leaves are distributed along the branches, e.g., *E. harrisii*, *E. angustifolia*. Also, the type specimens (*Ridley et al.* 86) show that the inflorescences vary in length and architecture, as they are short and few-flowered in the GH and K specimens, and paniculate, many-flowered, and almost as long as the leaves in the BM holotype.

Negrón-Ortiz (2005:1585) treated *Erithalis insularis* as a synonym of *E. fruticosa*, with no further comment, showing that the specialist who revised the genus opted to include this species in *Erithalis*.

Taylor and Barbosa (in Jardim et al. 2015:18) transferred this species to *Chiococca*, with the new combination *C. insularis* (Ridl.) C.M. Taylor & M.R.V. Barbosa. Those authors stated that *Palicourea insularis* was originally described by Ridley (1890) as a shrub or small tree, with five-merous corollas with apparently valvate aestivation, and fruits with four pyrenes. However, Ridley (1890:41) described the fruits of this species as 5-locular. Taylor and Barbosa continued their discussion by stating that “However several of its characters do not agree with *Palicourea* (Palicoureeae), in particular its apparent lack of raphides, triangular stipules, axillary inflorescences, quincuncial corolla aestivation, and fusiform stigmas that are positioned together with the anthers in open flower; *Palicourea* is characterized by raphides in the tissues, bilobed stipules, terminal inflorescences, valvate corolla aestivation, and bifid stigmas that are positioned above or below the anthers in the open flowers.” They complemented those statements with a detailed comparison between *Erithalis* and *Chiococca*, and concluded that this species is better positioned in *Chiococca*.

Figueira et al. (2020) presented a long discussion on the ecology and conservation of *Chiococca insularis*, mostly stimulated by the fact that this rare species, previously thought to be extinct, was rediscovered in Fernando de Noronha. They also added a plate illustrating the features of this species. The characters shown in figure 3 of Figueira et al. (2020), in my opinion, provide significant evidence that this species should be maintained in *Erithalis*. In figure 3E is shown an ovary in cross section, with 4 locules and one ovule each, while most species of *Chiococca* have 2-locular ovaries, which commonly develop into 2-seeded fruits, or in some species 1-seeded fruits by abortion of one ovule (e.g., *C. nitida*). In *C. alba*, the ovaries are 2-locular, developing into 2-seeded fruits. In some rare populations of *C. alba*, the ovaries have been reported to be
exceptionally 3-locular (Müller Argoviensis, 1881; Robinson, 1902). In figure 3F is shown a paniculate inflorescence, similar to those of several species of *Erithalis*, e.g., *E. harrisii* and *E. angustifolia*. In figure 3F are also shown several flowers in anthesis, with deeply divided corollas and narrowly oblong lobes, as in most species of *Erithalis*. Finally, in figure 3G are shown smooth, black fruits, which are also typical of many *Erithalis* species, while the fruits of most species of *Chiococca* are commonly white. In conclusion, this taxon should be treated as a distinct, extremely rare species of *Erithalis*, endemic to the Fernando de Noronha Island.

6. **Erithalis odorifera** Jacq., Sel. Stirp. Amer. 72. 1763. *Erithalis odorata* Raf., Sylva Tell. 123. 1838 (intended to replace *E. fruticosa* in the sense of Jacquin and Plumier; no publication cited). *Erithalis fruticosa* var. *odorifera* (Jacq.) Griseb., Fl. Brit. W. I. 336. 1861. *Erithalis fruticosa* ssp. *odorifera* (Jacq.) Steyerm., Fl. Venez. 9:869. 1974. TYPE: [Icon.]: Plumier, Icon 249, f. 2. (LECTOTYPE (Lebrón-Ortiz 2005:1589)).

*Distribution*.—Bahamas, Greater and Lesser Antilles, and Caribbean coast of Venezuela.

*Erithalis rotundata* Griseb., Pl. Wright. 2:507. 1862. TYPE: CUBA. HOLGUIN: Monte Verde, Jan–Jul 1859, C. Wright 1268 (LECTOTYPE (Lebrón-Ortiz 2005:1589): GOET [GOET008938]; ISOLECTOTYPES: GH [2 sheets, 00092684, 00092685], MO [2 sheets, Nos. 2092105, 2092106], NY [00743697], PH [00012880 (“Sep 1859–Jan 1860”)], S [No. S05-988], YU [2 sheets, YU.065585, YU.065586]).

7. *Erithalis orbiculata* (Proctor) A.R. Franck, P.A. Lewis & Oberli, Phytotaxa 297(3):278. 2017. *Exostema orbiculatum* Proctor, J. Arnold Arbor. 63:303. 1982. TYPE: JAMAICA: Trelawny, Crown Lands area, ca. 5 mi NW of Troy, 1750–2000 ft. [530–610 m], 10 Jun 1975, *G. Proctor* 35236 (HOLOTYPE: IJ [No. 61431]; ISOTYPES: A [00046005], US [00130605]).

*Distribution*.—Jamaica.

8. *Erithalis quadrangularis* Krug & Urb., Notizbl. Bot. Gart. Berlin-Dahlem 1:320. 1897. TYPE: JAMAICA: New Green, near Manchester, 29 Apr 1896, *W.H. Harris* 6318 (LECTOTYPE (Lebrón-Ortiz 2005:1591): NY [000115116]).

*Distribution*.—Jamaica, St. Vincent and the Granadines.

*Notes*.—Krug and Urban (1897:320), in the protologue of *Erithalis quadrangularis* Krug & Urb., cited the material studied as “Habitat in Jamaica in New Green 700 m alt., m. April. florif.: Bot. Dep. Herb. (W. Harris) n. 6318.” The original material at B studied by Krug and Urban was destroyed during WWII. Lebrón-Ortiz (2005:1591) designated specimen *Harris* 6318 in NY as the lectotype.

9. *Erithalis salmeoides* Correll, J. Arnold Arbor. 58:49, fig. 8. 1977. TYPE: BAHAMAS: Great Inagua, between Conch Shell Point and Lantern Head, 3 Aug 1975, *D.S. Correll* 45897 (HOLOTYPE: A [00092686]; ISOTYPE: FTG n.v.).

*Distribution*.—Bahamas (Great Inagua, Little Inagua, Mayaguana), Turks and Caicos, Jamaica, and Cuba (Santiago de Cuba).

10. *Erithalis vacciniifolia* (Griseb.) Wright, Anal. Acad. Ci. Habana 6:126. 1869. *Chione vacciniifolia* Griseb., Cat. Pl. Cub. 133. 1866. *Erithalis fruticosa* ssp. *vacciniifolia* (Griseb.) Borhidi, Bot. Közlem. 58:177. 1971. TYPE: CUBA. Baracoa, 1860–1864, C. Wright 2719 (HOLOTYPE: GOET n.v.; ISOTYPES: GH [00092520], K [K000173941]).

*Distribution*.—Cuba and Dominican Republic.

EXOSTEMA

Bonpland (in Humboldt et Bonpland, Pl. Aequin. 1:131. Apr 1807 [“1808”]) published the genus *Exostema* (Pers.) Bonpl. by raising *Cinchona* subg. *Exostema* Pers. to genus rank, hence, the type of *Exostema* is the type of *C. subg. Exostema*, *C. caribaea* Jacq. [= *Exostema caribaeum* (Jacq.) Schult.] Bonpland derived the genus name by referring to the exserted stamens of the genus. The taxonomic relationships among *Coutarea*, *Hintonia* and *Exostema* have been variously interpreted by different authors. For example, Hooker (1873a), Schumann (1891), and Robbrecht (1988), placed *Coutarea* and *Exostema* in *Cinchoneae*, because of their capsular fruits and winged seeds. McDowell (1995, 1996) delimited *Exostema*, by having terminal inflorescences and corolla tubes 13–21 cm long (e.g., *E. longiflorum* Roem. & Schult.) or axillary inflorescences and corolla tubes 1–4 cm long (e.g., *E. nitens* Urb.). Delprete (1996), with a phylogenetic analysis of the *Catesbaeae-Chiococceae-Exostema* complex using morphological data, retrieved three clades, which he called *Catesbaeae* (*Portlandia*-group, *Catesbae*-group), *Exostema*-group, and *Chiococceae*. *Exostema*, as traditionally delimited (e.g., McDowell, 1995, 1996), is a highly polymorphic taxon, which was later shown to be paraphyletic, and
this reflected on the difficulties in coding the polymorphic morphological characters used in the phylogenetic
analysis by Delprete (1996). McDowell (1996) and McDowell and Bremer (1998) retrieved *Exostema sensu*
McDowell (1996) as monophyletic; however, this result is misleading because in their analysis they did not
include any additional ingroup genera, and they used *Coutarea hexandra* as the only outgroup, which is closely
related to the South American species of *Exostema sensu* McDowell (McDowell et al. 2003; Motley et al. 2005;
Paudyal et al. 2018). Paudyal et al.'s (2018) molecular phylogenetic analysis reiterated that *Exostema*, as tradi-
tionally circumscribed, is not monophyletic, corroborating most of the previous molecular phylogenetic
studies (McDowell et al. 2003; Motley et al. 2005; Robbrecht & Manen 2006; Bremer & Eriksson 2009; Manns
& Bremer 2010; Manns et al. 2012). Paudyal et al.'s (2018) subclade B1 comprised only the eight *Exostema* spe-
cies with axillary inflorescences. Their results agreed with the morphology-based delimitation of *Exostema*
section *Exostema* of McDowell (1996). Accordingly, Paudyal et al. (2018) restricted the delimitation of
*Exostema* by including only the species with axillary inflorescences, which are distributed in Cuba and
Hispaniola, and *E. caribaeum*, which is also present in other Antilles, Mexico, and Central America.

*Exostema* sensu Paudyal et al. (2018) is a genus of eight species of shrubs or small trees 1–6(–10) m tall, with axillary inflorescences, 4–5-merous flowers, fragrant during the day, actinomorphic corollas, stamens exserted or partially exserted, style clavate or subcapitate, capsules obovate in outline, slightly laterally compressed, and seed basipetally, acropetally or centrally inserted.

**Exostema** (Pers.) Bonpl. in Humboldt et Bonpland, Pl. Aequin. 1:131. Apr 1807 ["1808"). *Cinchona* subg. *Exostema* Pers., Syn. Pl. 1:196. 1 Apr–15 Jun 1805; W. Wright, Phil. Trans. 62(2):504–506, f. 10. 1778; Standley, N. Amer. Fl. 32(2):117–126. 1921; Liogier, Fl. Cuba 5:20–25, f. 2. 1962; Correll & Correll, Fl. Bahama Arch. 1394, f. 607. 1982; Borhidi & Fernández Zequeira, Acta Bot. Hung. 35(1–4):287–307. 1989; Liogier, Fl. Española 7:254, 256–265, f. 198–15. 1995; McDowell. Monogr. *Exostema* (Rubiaceae), Ph.D. Thesis, Duke University, 1995; McDowell, Opera Bot. Belg. 7:277–295. 1996; McDowell & Bremer, Pl. Syst. Evol. 212:215–246. 1998; Darók et al., Acta Bot. Hung. 42(1–2):85–96. 2000; McDowell et al., Syst. Bot. 28:431–441. 2003; Lorence, Fl. Mesoamericana 4.2:86–87. 2012; Borhidi et al., Rubiáceas Cuba 103–119, f. 27–28. 2017; Paudyal et al., Bot. J. Linn. Soc. 187(3):365–396. 2018; Borhidi et al., Acta Bot. Hung. 60(3–4):302–311. 2018; Greuter & Rankin-Rodríguez, Taxon 70:906. 2021; Greuter & Rankin-Rodríguez, Taxon 71:210–215. 2022; Delprete & Paudyal, Taxon 72(5):1098–1108. 2023. LECTotype (Rogers 1987:165): *Exostema caribaeum* (Jacq.) Schult.

1. **Exostema acuminatum** Urb., Symb. Antill. 3:378. 1903. TYPE: DOMINICAN REPUBLIC. SANTO DOMINGO: Santo Domingo, near the city, s.d., *Rob. H. Schomburgk 42* (NEOTYPE, **here designated**: K [K000173622]).

**Distribution.**—Haiti and Dominican Republic.

**Notes.**—Urban (1903:378–379) in the protologue of *Exostema acuminatum* Urb., cited the following gatherings: *Buch* 753 from Haiti; *Rob. Schomburgk 42*, *Rob. Schomburgk 85*, and *Eggers 1798* from Dominican Republic; without citing the herbarium of deposit. The original material at B studied by Urban was destroyed during WWII. Three corresponding specimens are present in separate herbaria: *Schomburgk 42* in K, *Schomburgk 85* in F, and *Buch* 753 in GH. None of those specimens shows proof that they were examined by Urban. The K specimen of *Schomburgk 42*, barcode K000173622, has a label with the annotation “No. 42” followed by a detailed description handwritten by Robert Schomburgk. On the sheet are affixed two branches with numerous leaves and numerous flowers in anthesis. This specimen is here designated as the neotype of *E. acuminatum*.

2. **Exostema caribaeum** (Jacq.) Schult. in Roem. & Schult., Syst. Veg., ed. 15[bis], 5:19. 1819. *Cinchona caribaea* Jacq., Enum. Pl. Carib. 16. 1760. TYPE: DOMINICA: Prince Rupert Head, 8 Jun 1792, *J. Findlay s.n.* (NEOTYPE (McDowell 1996:283, 289): BM [000028058]).

**Distribution.**—USA (southern Florida), Bahamas, Greater and Lesser Antilles, Mexico, Guatemala, Honduras, Costa Rica, Panama.

**Notes.**—Jacquin (1760:16) published *Cinchona caribaea* Jacq. with the succinct description “pedunculis unifloris” without citing any specimen. Three years later, Jacquin (1763:61–62, tab. 179, fig. 95) published a detailed description of *C. caribaea*, followed by a statement regarding the collection localities and phenology, “Habitat in fruticetis circa Havanam; & in Domingo ad finum Bayaha. Floret Septembri & Octobri. Fructus maturos collegi Decembri.” In Jacquin’s table 179, figure 95 is depicted a dehisced capsule and one half of a
dehisced capsule, which could belong to a number of Rubiaceae genera. This figure is not original material, and does not represent reliable material for the application of this name. Article 9.4 of the *Code* (Turland et al. 2018) states, in part, “For the purposes of this *Code*, original material comprises the following elements: (a) those specimens and illustrations (both unpublished and published prior to publication of the protologue) that the author associated with the taxon, and that were available to the author prior to, or at the time of, preparation of the description, diagnosis, or illustration with analysis (Art. 38.7 and 38.8) validating the name; [...].”

Howard (1973), regarding the material gathered by Jacquin in the Caribbean Region, stated, “The consignments of living plants had been considerable; herbarium material was scanty, but many drawings and descriptions had been made on the spot.” According to Art. 9.4 (Turland et al. 2018), if Jacquin’s original drawing of *Cinchona caribaea* still exists, it would be original material. In the Botany Library of the Natural History Museum of London, is preserved a volume with Jacquin’s original notes and drawings, with the description “Von Jacquin botanical notes. 18th-century botanical drawings and notes made by French scientist and botanist Nicolaus Joseph von Jacquin (1727–1817).” Rosie Jones, Special Collection Librarian of the Natural History Museum, carefully searched the volume for a possible drawing of *Cinchona caribaea*, and confirmed that no such drawing is present in the volume.

McDowell (1996) searched BM and LINN and did not find any original specimen of *Cinchona caribaea*. A recent search in the W virtual herbarium (https://jacq.org) also did not find any original specimen. McDowell's (1996:283, 289) type citation of *C. caribaea* is somewhat confusing. On page 283, he designated a specimen of *De Ponthieu s.n.* in BM, collected in Dominica in 1792, as the neotype, and on page 289, he designated it as the lectotype. In BM there is a specimen of *De Ponthieu s.n.*, barcode 000028156, which McDowell identified in 1994 as *Exostema sanctae-luciae* (Kentish) Britten. Since McDowell identified the *De Ponthieu s.n.* specimen as *E. sanctae-luicae*, it cannot be the neotype of *E. caribaeum* (Jacq.) Schult. Hence, McDowell cited the wrong specimen as neotype. According to article 9 of the *Code* (Turland et al. 2018), errors of a type citation can be corrected. In BM there is a specimen, barcode 000028058, which was collected by Captain Findlay on Prince Rupert Head in Dominica, on 8 June 1792. Hence, specimen 000028058 is assumed to be the one that McDowell really meant to cite in his typification of *C. caribaea*.

*Cinchona jamaicensis* W. Wright, Philos. Trans. 67:506, f. 10. 1778, nom. *superfl.* of *Cinchona caribaea*.

*Cinchona myrtifolia* Stokes, Bot. Mat. Med. 1:359. 1812. Type: Not traced.

**Notes.**—Jonathan Stokes (1812:359), in the protologue of *Cinchona myrtifolia* Stokes, cited the material studied as “*Obs*. 6211. Specimen gathered by Broughton in Jamaica.”

*Cinchona racemosa* Schrank ex Steud., Nom. Bot. ed. 2, 1:363. 1840, nom. *illeg. superfl.*

**Notes.**—*Exostema caribaeum* (as “*Exostemma*”) is cited in synonymy under this name, which corresponds to *Cinchona caribaea* Jacq, a valid name cited on the same page.

*Exostema longicuspe* Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1852:48. 1853. Type: COSTA RICA: Puntarenas, Island of San Lucas, s.d., A.S. *Oersted 11403* (*LECTOTYPE*, here designated: C [C10018137]; isolectotypes: C [C10018138], K n.v.).

**Notes.**—Oersted (1853:48) in the protologue of *Exostema longicuspe* Oerst., cited his own collection as “Paa den lille Ø San Lucas ved af Costa-Rica i Naerheden af Puntarenas.” There are two specimens in C associated with this name and collected by Ørsted, both of them with the collection number 11403 and the handwritten annotation “Exostemma longicuspis Ørsted. In insula San Lucas.” Specimen with barcode C10018137 is here designated as the lectotype of *E. longicuspe*.

*Exostema caribaeum* var. *velutinum* Urb., Symb. Antill. 8:665. 1921. TYPE: HAITI: Plaine in hills, s.d., C.G. *Ehrenberg* 339 (not traced).

*Exostema caribaeum* var. *pubescens* Borhidi & Muñiz, Acta Bot. Acad. Sci. Hung. 18:48. 1973. TYPE: CUBA. *SANTIAGO DE CUBA*: near Versailles, s.d., A. *Borhidi*, Muñiz & S. *Vazquez s.n.* (HOLOTYPE: BP n.v.).

*Exostema veraensis* Kitanov, Fitologya 11:51. 1979. TYPE: CUBA. *SANTIAGO DE CUBA*: Between Santiago de Cuba airport and Punta Gorda, in shrubs on limestone, s.d., B.P. *Kitanov SV No. 18660* (“1860”) (HOLOTYPE: HAC [ex SV]).

**3. Exostema glaberrimum** Borhidi & M. Fernández, Acta Bot. Hung. 35:302. 1989. TYPE: CUBA. *GUANTÁNAMO*: Mun. Baracoa, parte del Yunque de Baracoa, bosque siempre verde mesófilo, 17 Apr 1986, I. *Arias*, M. *Díaz*, E. *Genes*, J. *Cutíerrez*, R. *Oviedo*, R. *Rankin* & G. *Stohr HAJB 58878* (HOLOTYPE: HAJB [HAJB G 000452]; ISOTYPE: B [B 10 0280881]).

**Distribution.**—Cuba (Guantánamo: Yunque de Baracoa).
**4. Exostema lancifolium** Borhidi & Acuña, in Borhidi & Muñiz, Acta Bot. Acad. Sci. Hung. 17:28, fig. 12. 1972 [“1971”]. **TYPE:** CUBA. HOLGUÍN: Moa, Mina Potosi, above Yamanigüey, 1968, *V. Samek No. 26830* (*LECTOTYPE*, **here designated**: HAC [ex SV, specimen A (Fig. 1)]; *ISOLECTOTYPE*: HAC [ex SV, specimen B]).

**Distribution.**—Cuba (Holguín: Moa; Guantánamo: Baracoa).

**Notes.**—Borhidi and Acuña (in Borhidi & Muñiz, 1972:28–29) cited the type of *Exostema lancifolium* Borhidi & Acuña as “Typus: Prov. Oriente; Region de Moa, in fruticetis serpentinosis ad Minas Potosi, supra Yamanigüey. Leg. V. Samek V. 1968, No. 26830 SV.” The code “SV” refers to the Herbarium of the Estación Experimental Agronómica of Santiago de las Vegas, which has been integrated into HAC. Borhidi et al. (2017:110) cited the type of *Exostema lancifolium* as “Tipo: Cuba, Prov. Holguín, Region de Moa, [...], supra Yamanigüey. Leg. V. Samek, 1968. Holotipo: SV 26830 (HAC).”

In HAC there are two specimens of *Samek No. 26830*. Both of them are annotated as “Exostema lancifolium Borhidi & Acuña” by Borhidi. To distinguish them, I wrote the letters “A” and “B” in pencil directly on the sheets. These two letters mean that the two sheets should be treated as two different specimens. Specimen A (Fig. 1) consists of a branch with numerous leaves, two dehisced capsules, and several flower buds, and is here designated the lectotype of *Exostema lancifolium*. Specimen B consists of a branch with numerous leaves, half of a dehisced capsule, and two flower buds, and is an isotype.

*Exostema lucidum* Borhidi & M. Fernández, Acta Bot. Hung. 35:302. 1989. *Exostema lancifolium* var. *lucidum* (Borhidi & M. Fernández) Borhidi, Rubiáceas de Cuba 110. 2017. Type: CUBA. Holguín: Moa, near Minas Jaragua, barranco del Arroyo Jaragua cerca la Mina Jaragua, 200 m, 3 May 1980, A. Álvarez de Zayas, J. Bisse, J. Gutiérrez & F.K. Meyer HAJB 42659 (*LECTOTYPE*, **here designated**: HAC [ex HAJB]; isotype: B [B 10 0273062]).

**Notes.**—Borhidi & Fernández Zequeira (1989:302), in the protologue of *Exostema lucidum* Borhidi & M. Fernández, did not specify which HAJB specimen is the holotype. Borhidi et al. (2017:110) cited the type of *Exostema lucidum* as “Tipo: Cuba Oriental, Prov. Holguín; Moa; Barranco del arroyo Jaragua [...] Col. Alvarez et al. 3.05.1980. Holotipo: HAJB 42659, isotype: HAJB.”

As a result of a detailed search in HAJB, no original material *Exostema lucidum* was found. The HAJB curator, Eldis R. Béquer (pers. comm.), is not aware of the possible location of the original specimens of this name. All the HAJB specimens of *Exostema* previously on loan to JPU, for study by Borhidi, have been returned.

In HAC there is a specimen of *Alvarez et al. HAJB 42659*, which was donated by HAJB to HAC. On the upper right corner of the sheet is the stamp “Herbario del Jardín Botánico Nacional, Universidad de La Habana. HAJB.” On the specimen is affixed a label with the annotation “ISOTYPUS! *Exostema lucidum* Borhidi et Fernández by Borhidi.” On the specimen label the collectors are cited as “J. Bisse, F. Meyer, A. Alvarez, J. Gutierrez.” The specimen consists of three branches with numerous leaves and numerous flowers in anthesis, and is here designated the lectotype of this name.

**5. Exostema nitens** Urb., Rep. Sp. Nov. 17:7. 1921. **TYPE:** DOMINICAN REPUBLIC. PEREVIA: Cordillera Central, San José de Ocoa, en el lugar llamado Amacey Gordo, 18°33'N, 70°35'W, 810 m, 8 Mar 2000, A. Veloz & M. De La Cruz 2066 (*NEOTYPE*, **here designated**: FLAS [No. 209895]; *ISOTYPES*: MO [No. 5677546], NY n.v.).

**Distribution.**—Dominican Republic (Cordillera Central).

**Notes.**—Urban (1921b:7), in the protologue of *Exostema nitens* Urb., cited the gathering *Bertero* 325 without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. A search in Jstor Global Plants and Jacq virtual herbaria, and in G, G-DC, and TO herbaria, did not find any original specimen associated with this name. Hence it is necessary to designate a neotype for this name. The gathering *Veloz & De La Cruz* 2066 has flowers in anthesis and mature fruits, and has duplicates distributed in several herbaria. The specimen in FLAS, with accession number 209895, is here designated as the neotype of *E. nitens*.

**6. Exostema purpureum** Griseb., Cat. Pl. Cub. 125. 1866. **TYPE:** CUBA. HOLGUÍN: Cuchillas de Baracoa, 1860–1864, C. Wright 2671 (*LECTOTYPE*, **here designated**: GOET [GOET010241]; *ISOLECTOTYPES*: BM [BM001008864], G [G00436068], GH [00046006], MO [No. 2091689], YU [YU.001737]).

**Distribution.**—Eastern Cuba (Guantánamo: Baracoa; Holguín: Sierra Cristal, Sierra de Nipe, Sierra de Moa y Toa; Santiago de Cuba).

**Notes.**—Grisebach (1866:125), in the protologue of *Exostema purpureum* Griseb., cited the two Cuban gatherings, Wright 2671 from the Baracoa Province, and Wright 2680 from the Majari Province, without citing the herbarium of deposit. Grisebach’s original material is in GOET, where he worked.
<figure><img src="image_4.png" /><figcaption>Fig. 1. Lectotype of *Exostema lancifolium* Borhidi & Acuña (Samek No. 26830, HAC [ex SV]). Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
Borhidi et al. (2017:113; 2018:305) cited the type of *Exostema purpureum* as "Tipo: C. Wright 2761, Baracoa, holotipo: GOET, isotipos: GH, HAC, NY, BM, S, US." According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:113; 2018:305) "holotype" citations cannot be treated as valid lectotype designations.

The GOET specimen of Wright 2671, with barcode GOET010241, has a label with the annotation "E. glabrum, foliis ovati acuti floris axillaris superant. [...]" handwritten by Grisebach. This specimen consists of a branch with numerous leaves and a pair of old flowers with the corolla fallen off. In the pocket affixed on the sheet there is a corolla in anthesis and a corolla in bud. This specimen is here designated as the lectotype of *E. purpureum*.

The GOET specimen of Wright 2680, with barcode GOET010240, has a label with the annotation "Exostema an purpureum? n. sp. E. caribeam aff.? [...]" handwritten by Grisebach. This specimen consists of a branch with numerous leaves and a few capsules. In the pocket affixed on the sheet are included parts of a capsule and several seeds.

**7. Exostema salicifolium** Griseb., Cat. Pl. Cub. 125. 1866. TYPE: WESTERN CUBA: without locality, 1860-1864, C. Wright 2676 (FIRST-STEP LECTotype (Borhidi & Fernández Zequeira 1989:292), SECOND-STEP LECTotype, here designated: GOET [GOET010242]; ISOLeTYPeS: BM [00028153], G [G00436065], GH [00046010], HAC, K, MO [No. 2091692], NY [00077385], US [00130606], YU [YU.001739]).

**Distribution.**—Cuba (Artemisa: Bahia Honda, Toscano; La Habana; Pinar del Río).

**Notes.**—Grisebach (1866:125), in the protologue of *Exostema salicifolium* Griseb., cited the material examined as "Frutex arborescens, floribus albis odoratis" (Schomb.).—Cuba occ. (Wr. 2676); Haiti pr. S. Domingo (Schomb. 42)" without citing the herbarium of deposit.

Borhidi and Fernández Zequeira (1989:292) cited the type of *Exostema salicifolium* as "Tipo: Wr. 2676 [Wright 2676]" without citing the herbarium of deposit. Hence, their citation is a first-step lectotype designation.

Borhidi et al. (2017:115; 2018:306) cited the type of *Exostema salicifolium* as "Tipo: C. Wright 2676 Cuba Occidental. Holotipo: GOET, isotipos: GH, HAC!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:115; 2018:306) type citations cannot be treated as inadvertent lectotype designations.

The specimen Wright 2676 in GOET, with barcode GOET010242, is here designated as the second-step lectotype of *Exostema salicifolium*.

**8. Exostema spinosum** (Le Vavass.) Krug & Urb., Ber. Deutsch. Bot. Ges. 15:262. 1897.

**8a. Exostema spinosum** (Le Vavass.) Krug & Urb. ssp. **spinosum**, Ber. Deutsch. Bot. Ges. 15:262. 1897.
*Cinchona spinosa* Le Vavass., Observ. Phys. 37:243. 1790. TYPE: [icon]; Table 2 of Vavasseur, Observ. Phys. 37(2). 1790 (LECTotype, here designated).

**Distribution.**—Cuba (the whole island), Haiti, and Dominican Republic.

**Notes.**—Le Vavasseur (1790:243–244, tab. 2) described *Cinchona spinosa* Vavass. with ample details, without citing any specimen. After a general search in many herbaria, no original specimen associated with name was found. In Le Vavasseur’s Plate 2 is depicted a branch with numerous lateral branchlets, each of them with numerous minute leaves. Each branchlet terminates with an acicular thorn, which has extremely reduced internodes. On the branch are also depicted several flower buds at different stages of development,

<footnote>*Exostema purpureum* ssp. *mensurensis* Kitanov, Ann. Univ. Sofia Fac. Biol. 64:275. 1972. TYPE: CUBA. HOLGUIN: Sierra de Nipe, Mayarí, Loma de la Mensura, 21 Apr 1960, Alain [Liogier], Acuña [J. Acuña Gale] & Ramos 7985 (HOLOTYPE: HAC [ex SV]). (Fig. 2).</footnote>

<footnote>*Exostema purpureum* ssp. *avenium* Borhidi & M. Fernández, Acta Bot. Hung. 35:302. 1989. TYPE: CUBA. HOLGUIN: Majari Province, 1860-1864, C. Wright 2680 (HOLOTYPE: HAC; ISOTYPES: BM [000028157], F [No. 776427 (frag.)], GH [00061490], GOET [GOET010240], K [K000173621], NY n.v., MO [2091688], YU [YU.065588]).</footnote>

<footnote>*Exostema revolutum* Borhidi & M. Fernández, Acta Bot. Hung. 35:301. 1989. TYPE: CUBA. GUANTÁNAMO: Laderas al NE de la confluencia del Río Baracoa con Arroyo del Cayo Peladeros de Jauco, 20 Feb 1979, R. Berazain, L. Catasús, M. Duharte, R. Capote & A. López HAJB No. 39753 (HOLOTYPE: HAJB [HAJB G 000461]; ISOTYPES: B [B 10 0009700], HAJB [2 sheets, HAJB G 000462, HAJB G 000463]).</footnote>
<figure><img src="image_5.png" /><figcaption>Fig. 2. Holotype of *Exostema purpureum* ssp. *mensurensis* Kitanov (Alain et al. 7985, HAC [ex SV]). = *Exostema purpureum* Griseb. Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
and two flowers in anthesis with four exerted stamens, and the exerted style. On the plate are also depicted a flower bud (a), a flower in anthesis (b), a separate ovary and style (c), dehisced capsules (d), details of placentation (e), and seeds (f and g). The details on this plate are sufficient for the unequivocal application of the name, and this plate is here designated as the lectotype of *Cinchona spinosa*. https://gallica.bnf.fr/ark:/12148/bpt6k9603516s/f257.item (plate after page 320).

*Catesbaea vavassorii* Spreng., Syst. Veg., ed. 16, 1:416. 1825. *Catesbaea elliptica* Spreng. ex DC., Prodr. 4:401. 1830, *nom. illeg. pro syn.* (under *Catesbaea vavassorii* Spreng.). *Exostema vavassorii* (Spreng.) Griseb., Cat. Pl. Cub. 126. 1866. **TYPE:** [icon]: Table 2 of Vavasseur, Observ. Phys. 37(2). 1790 (*LECTOTYPE*, **here designated**).

*Notes*.—Sprengel (1825:416), in the protologue of *Catesbaea vavassorii* Spreng., cited the material studied as “*Hispaniola*. (Cinchona spinosa Vavass. in Rozier. obs. phys. tom. 37).” Hence he cited table 2 of Vavasseur, Observ. Phys. 37(2). 1790, which is here designated as the lectotype of this name.

**8b. Exostema spinosum** ssp. **tortuense** (Urb.) Borhidi, Bot. Koz. 62:27. 1975. *Exostema spinosum* var. *tortuense* Urb., Arkiv. Bot. 21A (5):72. 1928 [“1927”]. **TYPE:** HAITI: Île de Tortue, coastal cliffs at Boucan-Guêpes, 22 May 1925, E.L. Ekman 4087 (*LECTOTYPE*, **here designated**: S [S07-14968]; *ISOLECTOTYPE*: S [S07-14962]).

**Distribution.**—Haiti (Île de Tortue).

*Notes*.—Urban (1928a [“1927”]:72) in the protologue of *Exostema spinosum* var. *tortuense* Urb., cited the material studied as “Insula la Tortue in scopulosis littoralibus ad Boucan-Guêpe, m. Majo fl. et fr: [Ekman] n. H 4087,” without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. In S there are two specimens of *Ekman 4087*. The specimen with accession number S07-14968 has a label with the annotation “Exostema spinosum (Vavass.) Krug et Urb. var. tortuense (typus)” handwritten by Urban. The specimen consists of three branches with numerous small leaves, and several capsules, and is here designated as the lectotype of *E. spinosum* var. *tortuense*.

The S specimen with accession number S07-14962 has a label with the same annotations of the other *Ekman 4087* at S, but is not annotated by Urban. This specimen also consists of three branches with numerous small leaves, and several capsules, and is an isotype.

## EXOSTEMA EXCLUDED TAXA

*Exostema australe* A. St. Hil., Pl. Usuel. Bras. 1(3):2–4, tab. 3. 1824. **TYPE:** [icon.]; A. Saint-Hilaire, Plantes Usuelles des Brasiliens 1(3):2–4, tab. 3B. 1824 (*LECTOTYPE* (Germano Filho 1999:59)). BRAZI: [São Paulo], Mogy das Cruzes, s.d., A. *Saint-Hilaire Cat. D, N° 653* (*EPITYPE*, **here designated**: P [P03921477]; *ISOEPITYPE*: P [P03921483]).

= *Bathysa australis* (A. St. Hil.) K. Schum., in Martius et al., Fl. Bras. 6(6):239. 1889.

*Notes*.—Saint-Hilaire (1824:1(3):2–4), in the protologue of *Exostema australe* A. St. Hil., provided a detailed description, and cited the range of the species as growing in the forests of southern Brazil, southwards until the state of São Paulo. Germano Filho (1999:59) cited as type of this name table 3, fig. B of Saint-Hilaire's first volume of *Plantes Usuelles du Brasil*, associated with name *Exostema*. In figure B of table 3, are only depicted a capsule in side view, and a flower in anthesis in side view with exerted stamens. These characters are present in several species of *Exostema*, *Bathysa*, and numerous other Rubiaceae genera with capsular fruits. Hence, figure B is not sufficient to undoubtedly identify this species. To remove any doubt about the application of the name, an epitype needs to be designated. In P there are several specimens associated with this name, collected by A. Saint-Hilaire in Brazil. Specimen with barcode P03921477 has a label affixed on the bottom-left corner of the sheet, with the annotation “Exostema australe Aug. de S. Hil., Plan. us. Bras. Bois vierges prov. Mogy das Cruzes (A. de Saint-Hilaire Scr.)” handwritten by A. Saint-Hilaire. On the bottom-right corner of the sheet is affixed another label with the annotation “BRÉSIL.—Province de Saint-Paul. Voyage de Saint-Hilaire, de 1816 à 1821. Cat. D, N° 653.” The specimen consists of a distal portion of a branch, with large stipules, a large leaf, and infructescences with numerous capsules. This specimen is here designated as the epitype of *E. australe*.

*Exostema bicolor* Poepp, in Poeppig & Endlicher, Nov. Gen. Sp. Pl. 3:32. 1841.

= *Motleyothamnus corymbosum* (Ruiz & Pav.) Paudyal & Delprete (See below).

*Exostema canescens* Bartl. ex DC., Prodr. 359. 1830 (as “*Exostemma*”). *Cinchona canescens* (Bartl. ex DC.) Brign., Mem.

Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. II, 1:63. 1862.

= *Hintonia latiflora* (DC.) Bullock (See below).

*Exostema capitatum* Spreng., Neue Entd. 2:143. 1821. **TYPE:** Not traced.

= *Phragmanthera capitata* (Spreng.) Balle, Adansonia, n.s., 1:251. 1962. (Loranthaceae).
*Exostema coriaceum* (Poir.) Schult., in Roemer & Schultes, Syst. Veg. ed. 15 [bis], 5:20. 1819. [Cinchona coriacea
Poir. in Lamarck, Encycl. 6:38. 1804]

= *Solenandra coriacea* (Poir.) Delprete, **comb. nov.** (See below)

*Exostema corymbosum* (Ruiz & Pav.) Spreng., Syst. Veg. 1:706. 1824 [“1825”. Portlandia corymbosa Ruiz & Pav., Fl. Peruv. 2:49. 1799.

= *Motleyothamnus corymbosum* (Ruiz & Pav.) Paudyal & Delprete (See below).

*Exostema coulteri* Hook. f., in Hemsley, Diagn. Pl. Nov. Mexic. 32. 1879. TYPE: MEXICO. HIDALGO: Zimapán, s.d., J. Coulter 209 (HOLOTYPE: K [K000173537]; ISOTYPE: MO [No. 3751087]).

= *Syringantha coulteri* (Hook. f.) T. McDowell, Novon 6(3):278. 1996.

*Exostema cuspidatum* A. St. Hil., Pl. Usuel. Bras., tab. 3. 1824. *Schoenleinia cuspidata* (A. St. Hil.) Klotzsch, Getreue Darstell. Gew. 14(2):tab. 15. 1846. *Cinchona cuspidata* (A. St. Hil.) Brign., Mem. Mat. Fis. Ital. Sci. Modena, Pt. Mem. Fis., ser. 2, 1:63. 1862. *Bathysa cuspidata* (A. St. Hil.) Hook. f. ex K. Schum. in Martius et al., Fl. Bras. 6(6):237. 1889. TYPE: BRAZIL: Minas Gerais, “Cueilli dans les bois près Itajuru, Cap de Moines,” 1816–1821, A. *Saint-Hilaire Catal. BI No. 970* (LECTOTYPE (Taylor et al. 2011:503): P [P00752481]; ISOLECTOTYPE: P [2 sheets, P00752482, P00752483]; POSSIBLE ISOLECTOTYPE: F [No. 970756, barcode F0069068F (ex P, without collection number)]).

= *Schizocalyx cuspidatus* (A. St. Hil.) Kainul. & B. Bremer, Amer. J. Bot. 97:1976. 2010.

*Exostema dissimiliflorum* (Mutis ex Humb.) Schult., in Roemer & Schultes, Syst. Veg., ed. 15[bis], 5:17. 1819. *Cinchona dissimiliflora* Mutis ex Humb., Mag. Neuesten Entdeck. Gesammemten Naturk. Feunde Berlin 1:120. 1807. *Ladenbergia dissimiliflora* (Mutis ex Humb.) Klotzsch, Getreue Darstell. Gew. 14(2):tab. 15. 1846. *Macrocnemum dissimiliflorum* (Mutis ex Humb.) Triana, Revista Acad. Colomb. Ci. Exact. 2:416. 1938. TYPE: COLOMBIA: without locality (not found).

= *Ferdinandusa dissimiliflora* (Mutis ex Humb.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 7:22. 1930.

*Exostema formosum* Cham. & Schltdl. ex DC., Prodr. 4(4):361. 1830. *Exostema formosum* Cham. & Schltdl., nom. inval., Linnaea 4:179. 1829. *Rustia formosa* (Cham. & Schltdl. ex DC.) Klotzsch, in Hayne, Getr. Darstell. Gew. 14:tab. 15. 1846. TYPE: BRAZIL: without locality, s.d., *Collector Unknown s.n.* (LECTOTYPE (Delprete 2024:144): G-DC [G00665732]).

= *Rustia formosa* (Cham. & Schltdl. ex DC.) Klotzsch, in Hayne, Getr. Dartell. Gew. 14:tab. 15. 1846.

*Exostema formosum* var. *α leprosum* DC., Prodr. 4(4):361. 1830. *Exostema formosum* forma *β laeve* Cham. & Schltdl., nom. inval., Linnaea 4: 179. 1829. TYPE: BRAZIL: without locality, s.d., *F. Sellow s.n.* [1814] (LECTOTYPE (Delprete 2024:144): HAL [HAL097786]).

= *Rustia formosa* (Cham. & Schltdl. ex DC.) Klotzsch, in Hayne, Getr. Dartell. Gew. 14:tab. 15. 1846.

*Exostema formosum* var. *β laeve* DC., Prodr. 4(4):361. 1830. *Exostema formosum* forma *α leprosum* Cham. & Schltdl., nom. inval., Linnaea 4:179. 1829. TYPE: BRAZIL: without locality, s.d., *Collector Unknown [Sellow] s.n.* (LECTOTYPE (Delprete 2024:144): G-DC [G00665732]; ISOLECTOTYPE: HAL [HAL097787 (collected by Sellow)]).

= *Rustia formosa* (Cham. & Schltdl. ex DC.) Klotzsch, in Hayne, Getr. Dartell. Gew. 14:tab. 15. 1846.

*Exostema leonis* Standl., Publ. Field. Mus. Nat. Hist., Bot. Ser. 8:338. 1931. TYPE: CUBA. PINAR DEL RÍO: Loma Pelada, Cayajabos, 9 Aug 1928, Bro. León & J.T. Roig 13531 (HOLOTYPE: NY [00077374]).

= *Acunaeanthus tinifolius* (Griseb.) Borhidi, Acta Bot. Acad. Sci. Hung. 26:286. 1981 [“1980”].

*Exostema macrocnemia* (Mart.) G. Don, Gen. Hist. 3:482. 1834. *Cinchona macrocnemia* Mart., Ann. Sci. Nat., Bot., sér. 3, 10:13. 1848. TYPE: COLOMBIA. [CAQUETA]: “Mount Aracaroa” [Aracaruara], s.d. [Jan 1820], C.F.P. Martius *s.n.* (M? not found).

= *Remijia macrocnemia* (Mart.) Wedd., Ann. Sci. Nat., Bot., sér. 3, 10:13. 1848.

*Exostema nerifolium* A. Rich., in R. de la Sagra, Hist. Fis. Cuba, Bot. 11:7. 1850. TYPE: CUBA: “Vuelta de abajo,” s.d., J. Maria Valenzuela *s.n.* (P? [not there]).

= *Suberanthus nerifolium* (A. Rich.) Borhidi & M. Fernández, Acta Bot. Acad. Sci. Hung. 27:316. 1982 [“1981”].

*Exostema occidentale* Benth., Bot. Voy. Sulphur 104. 1845 [“1844”. TYPE: COLOMBIA: NARINO: Isla Gorgona, 1841, R.B. Hinds 354 (LECTOTYPE (Delprete 1999b:7): K [K000470115]; ISOLECTOTYPE: BM [barcode unknown]).

= *Rustia occidentalis* (Benth.) Hemsl., Biol. Centr.-Amer., Bot. 2:14. 1881.

*Exostema orbiculatum* Proctor, J. Arnold Arbor. 63:303. 1982. TYPE: See under *Erithalis orbiculata* (above).

= *Erithalis orbiculata* (Proctor) A.R. Franck, P.A. Lewis & Oberli (See above).

*Exostema souzanum* Mart., in Martius & Spix, Reise Bras. 2:789. 1828. *Cinchona souzana* (Mart.) Brign., Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis., ser. 2, 1:63. 1862.

= *Coutarea hexandra* (Jacq.) K. Schum. (See above).