Domingo Bello y Espinosa (1817–1884; Fig. 1) was a natu-ralist from the Canary Islands whose modest yet important contributions to the botany of Puerto Rico during the late 19th century have been consigned to obscurity. Indeed, for the 400 years that the island was under the dominion of the Spanish Crown, Bello y Espinosa (hereinafter Bello) was the only Spanish-born resident of Puerto Rico who published taxonomic work on the local flora. Prior to Bello’s botanical accounts, official Spanish plant exploration in Puerto Rico was limited to a single expedition that took place between 1796 and 1797, which was conducted by the Spanish botanist Martín de
Sessé (1751–1808) and the Cuban botanist José Estévez Cantal (1771–1841; Blanco & al., 2000). Bello’s work, entitled Apuntes para la flora de Puerto-Rico (hereinafter Apuntes; Bello, 1881, 1883; Fig. 2), made him the only Canary Island native included in Taxonomic literature II (Stafleu & Cowan, 1976). Although biographical literature pertinent to personalities from the Canary Islands singles him out as a relevant historical figure (e.g., Pinto, 1884; Padrón Acosta, 1968; Rodríguez Delgado, 1995; Izquierdo, 2005), and despite the fact that his Apuntes are cited in the botanical literature related to Puerto Rico (e.g., Ackerman, 1995; Liogier, 1996; Acevedo-Rodríguez & Strong, 2005), there has not been a single work focusing on his botani-cal contributions. This article aims to address that neglect by
Domingo Bello y Espinosa (1817–1884) and the new taxa published in his Apuntes para la flora de Puerto-RicoEugenio Santiago-Valentín,1,2 Lázaro Sánchez-Pinto3 & Javier Francisco-Ortega4,5
1 Departamento de Biología, Universidad de Puerto Rico, Recinto de Río Piedras, Apartado Postal 23360, San Juan, Puerto Rico 00931-3360, U.S.A.
2 Herbario, Universidad de Puerto Rico, Jardín Botánico Sur, 1187 Calle Flamboyán, San Juan, Puerto Rico 00926-1117, U.S.A.3 Museo de la Naturaleza y el Hombre, Calle Fuente Morales No. 2, Santa Cruz de Tenerife, 38003 Tenerife, Spain4 International Center for Tropical Botany, Department of Biological Sciences, Florida International University, Miami,
Abstract Domingo Bello y Espinosa (1817–1884) was an attorney and naturalist from the Canary Islands who lived in Mayagüez (Puerto Rico) between 1848 and 1878. He is the author of a two-part publication dated 1881 and 1883, which represents one of the first contributions on plant taxonomy made by an island resident. These studies include a total of 88 new taxa. Eighty-four of them are new taxonomic descriptions (3 genera, 59 species, 22 varieties) and 4 of them are new combinations. Eight of these species are still accepted either as published originally by Bello (3 species) or as basionyms of subsequent taxonomic combi-nations (5 names). Stahlia Bello is also currently accepted. Twelve of Bello’s names are not legitimate (11 species, 1 variety). Bello did not leave a herbarium and published illustrations for only 3 of the new species that he described. Therefore, Bello’s names need to be interpreted on the basis of his descriptions and indirect accounts primarily provided by Ignatz Urban. This German botanist had access to illustrations (made by Leopold Krug) and plant collections (from Krug and Agustín Stahl) that were originally identified with Bello’s names. Here we assign neotypes for 42 of the 59 species described by Bello. The remaining 17 species not typified include 3 names typified in previous works, 9 illegitimate names, 3 names that we could not assign to any infrageneric taxon, and 2 names that are both illegitimate and can not be interpreted taxonomically. In addition, we lectotypify Caesalpinia monosperma (synonym of Stahlia maritima). We present a summary of the life of Domingo Bello that includes new data from archival research in the Canary Islands and Puerto Rico, as well as the only known portrait of him.
Keywords Antilles; botanical history; botanical nomenclature; Canary Islands; Caribbean Island Biodiversity Hotspot; neotypification; Puerto Rico
Supplementary Material The Electronic Supplement (Table S1) is available in the Supplementary Data section of the online version of this article at http://www.ingentaconnect.com/content/iapt/tax
B OTAN I C AL H IS TO RY / N O M EN CL AT U R E
324
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
presenting a taxonomic review of the plants he described for Puerto Rico. Given the scarce information published on the life of this naturalist, we also present a biography that incorporates data from primary sources, including archives in the Canary Islands (La Laguna, Tenerife: Archivo Municipal and Archivo Diocesano; Santa Cruz de Tenerife: Biblioteca Municipal) and Puerto Rico (Mayagüez: Archivo Histórico Municipal and Archivo Parroquial Catedral Nuestra Señora de la Candelaria).
Bello was born in La Laguna on 31 July 1817, and was baptized Domingo Francisco Ignacio Antonio del Salvamento in the parish church of Nuestra Señora de los Remedios on 2 August 1817. His parents were Domingo Bello Lenard, a uni-versity professor of mathematics, and Ana Espinosa y Carta, both native to La Laguna (see: “Fondo Santo Domingo de Guzmán de La Laguna, libro 36, signatura 36”; Folios 244 [verso]–245 [recto] from Archivo Diocesano, La Laguna). We are not certain of the specific location where Bello was born, but by 1823, at the age of six, he lived with his parents, a brother and a sister at Calle del Laurel (see: “Padrón de habitantes
varios municipios 1836–1841, P–I, S–II, 13” from Archivo Municipal, La Laguna). This street is currently known as Calle de Anchieta. He obtained his law degree at the Universidad de San Fernando, at La Laguna, in 1842 (Izquierdo, 2005). At that time he was still living at Calle del Laurel with his mother (then a widow), a brother, and three sisters (see: “Padrón de habitantes La Laguna 1841–1847, P–V [3], S–II” from Archivo Municipal, La Laguna).
In 1848, Bello migrated to Puerto Rico and settled in the western port city of Mayagüez, a common destination for Canarian immigrants during the 19th century (Cifre de Loubriel, 1995). In Mayagüez, he married Leocadia Raldiris Fernández, a member of a distinguished Puerto Rican family of that city (see: “Libro de matrimonios 1851, Vol 9, folio 105” from Archivo Parroquial, Catedral Nuestra Señora de la Candelaria, May-agüez), with whom he had a daughter (Isabel) and a son (José). While in Mayagüez, Bello practiced his profession as an attor-ney for a commercial company in the city (Wiley & al., 2014), and was the director of a school (see: “Documentos Históricos
Fig. 1. Undated portrait of Domingo Bello y Espinosa (1817–1884) wearing his lawyer’s toga. This is, at present, the only known portrait of him. Courtesy of Carlos Gaviño de Franchy.
Fig. 2. Title page of the first part of the Apuntes para la flora de Puerto-Rico (Bello, 1881).
325
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
de Mayagüez 1848, V–I” from Archivo Histórico Municipal de Mayagüez). After 30 years in Puerto Rico, Bello returned with his family to Tenerife in 1878 (Gundlach, 1880; Urban, 1898). By 1883 he was again living at Calle del Laurel in La Laguna (see: “Padrón de habitantes La Laguna 1883–1884, P–XXII [1], S–II, 1” from Archivo Municipal, La Laguna). He died a few years after returning to the Canaries on 21 January 1884, in a house located at Calle de la Candelaria No. 9 in Santa Cruz de Tenerife (see: “Fondo Santo Domingo de Guzmán de La Laguna, libro 20, signatura 105”; Folios 112 [recto]–112 [verso] from Archivo Diocesano, La Laguna). Bello is buried in the old Cementerio de San Rafael y San Roque, in Santa Cruz de Tenerife (García Pulido, 2000; Santiago-Valentín & al., 2014).
Bello was a cultured, well-known public figure in the Canary Islands. Between 1845 and 1847, he was the secretary of the influential Colegio de Abogados de Santa Cruz de Tenerife (Bar Association of Santa Cruz de Tenerife) (Izquierdo, 2005). He became Mayor of La Laguna twice; first in 1842, and then shortly after his return from Puerto Rico, between 1881 and 1883 (Padrón Acosta, 1968; Izquierdo, 2005; Santiago-Valentín & al., 2010, 2014). His formal obligations did not divert him in any way from cultivating a deep interest in natural history, and he was instrumental in the publication of a catalog of plants from the Canary Islands written in 1808 by the most distinguished Canary Islands naturalist of the 18th and 19th centuries, José de Viera y Clavijo (1731–1813; Viera y Clavijo, 1882a, b, c). In addition, between 1879 and 1880 Bello published a novel in ten parts that was centered on the plants of an imaginary botanic garden located in Shanghai that was devoted to the cultiva-tion of Canarian plants (Bello, 1879a–f, 1880a–d). Further-more, shortly after his arrival to Tenerife, in 1878, he wrote an unpublished manuscript titled “Apuntes botánicos de las Islas Canarias” (Botanical notes on the Canary Islands) (Steffen, 1948; Francisco-Ortega & al., in prep.).
Bello was part of the first generation of residents of Puerto Rico devoted to the study of its natural history. With the col-laboration of scientists from abroad, these naturalists fostered a scientific “golden age” especially for botany (Santiago-Valentín & al., 2010, 2014). The group included the wealthy German busi-nessman and naturalist enthusiast Leopold Krug (1833–1898), who also lived in Mayagüez, and Agustín Stahl (1842–1917), a physician native to Puerto Rico who resided in Bayamón (a town near the island capital, San Juan). Knowledge culled from their plant collections, illustrations, and publications was employed by Ignatz Urban (1848–1931), a professor at the Botanic Garden of Berlin and a leading taxonomist of Antillean plants, in the invaluable Symbolae antillanae (Howard, 1996; Liogier, 1996). Bello developed a friendship with Leopold Krug and together they studied the flora and fauna of Mayagüez and its environs (Howard, 1996). Krug was a patron of natural his-tory studies. For instance, he provided support to the German naturalist (then settled in Cuba) Johannes Gundlach (1810–1896) during his expeditions to collect plants and animals of Puerto Rico in 1873, and 1875–1876, as well as to the German botanist Paul Sintenis (1847–1907), who collected an impressive num-ber of plant specimens in Puerto Rico between 1884 and 1887 (Howard, 1996; Santiago-Valentín & González López, 2002).
Shortly after leaving Puerto Rico, Bello published his Apuntes as a two-part series in the journal of the Sociedad Española de Historia Natural (Bello, 1881, 1883). The first part was published in 1881, making the Apuntes one of the first taxonomic publications entirely devoted to the plants of Puerto Rico. Bello, however, acknowledged that the work was not a complete flora of the island: “Estos apuntes están muy léjos de formar un catálogo siquiera aproximado de la Flora de Puerto-Rico” (These notes are very far from constituting even an approximate catalog of the Flora of Puerto Rico). The Apuntes comprise a catalog of the 963 taxa of phanerogams and pteridophytes studied by him during his 30-year residence, focusing primarily on material from the floristically diverse western region of the island. Agustín Stahl’s Estudios sobre la flora de Puerto Rico (Stahl, 1883), and Estudios para la flora de Puerto Rico (Stahl, 1884, 1885, 1886, 1887, 1888) are the first attempt at a published Flora for Puerto Rico.
METHODS
Bello (1881) mentioned that insects destroyed his herbar-ium. In addition, Urban (1902) indicated that Bello did not make a herbarium and that the original materials for his work became part of the Krug herbarium, housed at B. Unfortunately, Krug’s collections and illustrations were destroyed during the Second World War, when most of the Berlin herbarium was bombed (Hiepko, 1996). The first author visited this institution in 2010 and did not find any of Krug’s collections or illustrations relevant to the Apuntes, nor specimens collected by Bello. Therefore, we faced the challenge of not having original plant material to interpret his names. Bello’s Apuntes included line engravings, but for only four species (also shown by Santiago-Valentín & al., 2014); two of them (Bello, 1883) illustrating the endemic orchids Epidendrum kraenzlinii Bello (accepted name Psychilis kraenzlinii (Bello) Sauleda; Figs. 3, 4D) and E. krugii Bello (accepted name P. krugii (Bello) Sauleda; Figs. 3, 4C). The other two illustrations were for Tarenaya spinosa (Jacq.) Raf. (Cleomaceae, published by Bello, 1881 as Cleome pun-gens Willd.) and Meliosma herbertii Rolfe (Sabiaceae, pub-lished by Bello, 1881 as Atelandra laurina Bello). It is not cer-tain who authored the artwork to produce these line engravings, but we infer that they were made by Bello himself. A separate study of correspondence sent by Gundlach from Puerto Rico to Cuban colleagues confirms that by 1876 Bello made sev-eral illustrations of Puerto Rican plants, birds and butterflies, and that these were compiled into a pictorial album (Santiago-Valentín & González López, 2002). In addition, Bello (1881) also referred to his plant drawings in his accounts for Doyerea emetocathartica Grosourdy (Cucurbitaceae, accepted name Anguria glomerata Eggers; Acevedo-Rodríguez & Strong, 2012) and Stellaria ovata Willd. ex Schltdl. (Caryophyll aceae, accepted name Stellaria antillana Urb. var. antillana; Acevedo- Rodríguez & Strong, 2012). While the current work was in press, we located the original illustrations of plants from Puerto Rico, deposited at the Museo Municipal de Bellas Artes de Santa Cruz de Tenerife, in the Canary Islands. This material
326
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
is being assessed for a future publication (Santiago-Valentín & al., in prep.).
Our assessment of the new taxa in the Apuntes relied on our interpretations of the morphological descriptions provided by Bello. Crucial to our work was Ignatz Urban’s Symbolae antillanae (Urban, 1903, 1905, 1910, 1911). Urban (1911) con-sidered that original materials for the Apuntes were Krug’s plants and illustrations, because these plants had been named by Bello and because Krug’s illustrations were, in a way, pre-pared under Bello’s supervision/guidance. In the fourth volume of Symbolae antillanae, Urban (1911) identified as “Bello!” 36 taxa for which a specimen with Bello’s name was available to him in the Krug herbarium (Electr. Suppl.: Table S1). In addi-tion, Urban included localities for seven taxa that were based
on information provided in the Apuntes, and were annotated as “ex Bello” (“according to Bello”).
Urban must have rigorously analyzed Bello’s descriptions even for those entities for which he had no material that came under Bello’s names. Urban was an astute and meticulous tax-onomist, and many of his taxonomic contributions are still widely accepted today. He had the great advantage of com-paring specimens from different collectors and from different areas in the Antilles. Urban’s studies not only included Krug’s collections from Puerto Rico, but also drawings made by the latter of plants from that island. Urban (1903, 1905, 1910, 1911) cites 66 of the new names (3 genera, 42 species, 21 variet-ies) published by Bello as related to Krug’s drawings, indicat-ing that these illustrations were originally accessioned under Bello’s names (Electr. Suppl.: Table S1). In addition, Urban (1903, 1910, 1911) reported specimens for nine species (Atelandra laurina Bello, A. obtusifolia Bello, Bignonia odorata Bello, Crescentia microcarpa Bello, Heteropterys pubiflora (DC.) Bello, Magnolia portoricensis Bello, Psychotria pseudopa-vetta Bello, Spathodea portoricensis Bello, Tephrosia aniloides Bello) and seven varieties (five within Mangifera indica L. and two within Anacardium occidentalis L.) from Stahl’s collec-tions that were originally labeled using Bello’s names (Electr. Suppl.: Table S1).
Among the plant specimens from Puerto Rico evaluated by Urban were those collected by Paul Sintenis between 1884 and 1887 (Liogier, 1996), right after the publication of the sec-ond part of the Apuntes (Bello, 1883). The collection, which became part of the “Krug & Urban herbarium” (located at B) was studied by Urban and associates (e.g., Karl M. Schumann, 1851–1904, a German botanist, determined specimens of Cereus quadricostatus Bello and Opuntia repens Bello, both Cact-aceae) with duplicates sold to several U.S. and European her-baria (Howard, 1996). Therefore, Sintenis’s specimens are linked to names published by Bello by means of the thorough taxonomic work of Urban published in his fourth volume of Symbolae antillanae. Although most of the original set of Sin-tenis (located at B) was lost, his many duplicates represent the largest number of Puerto Rico specimens surviving from that period, thus serving as an invaluable material for typification.
In his work, Stahl included Bello’s names listed in the Apuntes, and indicated that they were difficult to interpret and that the identification of many of them were incorrect and taxonomically questionable. However, Stahl acknowledged that he was unfamiliar with several of the plants listed by Bello. From this we concluded that Stahl did not study many of the original plant collections used by Bello for his new descrip-tions. However, based on Bello’s account for Stellaria ovata (see accepted name above), we infer that he was familiar with some of the illustrations of Puerto Rican plants made by Stahl (see Acevedo-Rodríguez, 2007 for a review of Stahl’s illustra-tions and collections).
In this work we assign neotypes to 42 of the 59 new species described by Bello. The remaining 17 species include 3 names typified in previous works. The orchids Epidendrum kraenzli-nii Bello and E. krugii Bello (Bello, 1883), were lectotypified by Sauleda (1988) using Bello’s published illustration. In addition,
Fig. 3. Original engravings of the orchids Epidendrum kraenzlinii Bello and E. krugii Bello from the Apuntes para la flora de Puerto-Rico (Bello, 1883). These illustrations were assigned as their lecto-types by Sauleda (1988).
327
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
Fig. 4. Species that are widely accepted by taxonomists, either as they were originally published by Bello or as basionyms of other taxa. A, Vriesea macrostachya (Bello) Mez; B, Metastelma lineare Bello; C, Psychilis krugii (Bello) Sauleda; D, Psychilis kraenzlinii (Bello) Sauleda; E, Meliosma obtusifolia (Bello) Krug & Urb. — Photo credit: A & E, E. Santiago-Valentín; B, R. Joglar; C, A. Cuevas-Pradó; D, J. Ackerman.
328
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Grant (1997) assigned neotype and isoneotypes to Caraguata macrostachya Bello (Bromeliaceae, accepted name Vriesea macrostachya (Bello) Mez; Fig. 4A). The limited descriptions of 3 of these 17 species did not allow unequivocal determina-tions and thus they were not typified. Furthermore, nine of the species that were taxonomically determined correspond to illegitimate names. Finally, two of Bello’s species names are both illegitimate and can not be interpreted taxonomically. In this study we lectotipify the name Caesalpinia monosperma Tul., which is synonym of Stahlia maritima Bello (see entry below for Stahlia maritima).
Bello (1881, 1883) made use of codes to identify taxa not found in the botanical literature consulted by him, and for which he gave a name and a description. New species were marked with one asterisk and new genera with two asterisks. These notations helped us to interpret Bello’s names since five of them (e.g., “Galactia filiformis”, “Jacquemontia tam-nifolia”, “Miconia pyramidalis”, “Pimenta vulgaris”, “Rumex berlandieri ”) were not marked with asterisks but were given full Latin/Spanish descriptions although they lacked author-ships. We believe that authorities for these names were not included because of typographical errors. Therefore, we did not consider these to be new names and assume that they refer to taxa previously described by other authors (see Discussion below with details on the interpretation of these five names and their authorities). Bello also described 22 new infraspecific taxa, although did not use codes to identify them in the way he did for his new species and genera. Only two of them (Emilia sonchifolia (L.) DC. var. rosea Bello, Rivinia humilis L. var. canescens Bello) were clearly labelled as varieties, but the rest were not assigned to any infraspecific rank. We have regarded all of these names as varieties, and their taxonomic interpreta-tion is also included in our study.
Our research was also based on other bibliographic/taxo-nomic database resources, including the Kew Bibliographic Database (Royal Botanic Gardens, Kew, 2013), the Interna-tional Plant Name Index (IPNI, 2013), the Plant List (The Plant List, 2013), the World Checklist of Selected Plant Families (Govaerts & al., 2000, 2008, 2013a, b), and the Malpighiaceae Nomenclature Website (Anderson & al., 2008). Besides the aforementioned Puerto Rican Floras by Stahl (1883, 1884, 1885, 1886, 1887, 1888) and Urban (1903, 1905, 1910, 1911) we also con-sulted subsequent floristic studies made by Britton & Wilson (1923, 1924a–c, 1925a–b), Liogier (1985, 1988, 1994, 1995, 1997), Acevedo-Rodríguez (2005a), Acevedo-Rodríguez & Strong (2005), and Axelrod (2011). The recent catalog of seed plants of the West Indies (Acevedo-Rodríguez & Strong, 2012) was use-ful to compile lists of synonyms to Bello’s names. Furthermore, we consulted taxonomy specialists (see the Acknowledgments section below) to guide us in the taxonomic interpretation of these names. We studied plant collections from the Caribbean Islands deposited at B, GH, NY, SJ, UPR, UPRRP, as well as digital specimens from B, BM, G, GH, K, LD, MO, NY, P, S, US, and W.
The new taxonomic descriptions and taxonomic combina-tions published by Bello (1881, 1883) are presented below in the same sequence as originally found in his works. We have only
focused on those names that Bello considered as novelties (see above). For each of these names we provide a list of synonyms. Currently accepted names appear in bold face italics type. In addition, we provide a discussion of each of these names with details pertinent to: (1) interpretation of these names in taxonomic/floristic works and (2) references to Bello’s names originally assigned to Stahl’s specimens (as Stahl!) and Krug’s illustrations/specimens as indicated by Urban (1903, 1905, 1910, 1911). Table S1 (Electr. Suppl.) provides a list of names published by Bello (1881, 1883) with their current taxonomic interpreta-tion and assigned types. This table also provides details on those names for which there was material and/or illustrations from Krug or Stahl. As neotypes we have mostly chosen her-barium specimens collected by Sintenis that Urban associated to Bello’s names. We assigned neotypes from collections by A.A. Heller (three specimens) or A. Stahl (two specimens) that were also evaluated by Urban, in those instances when we were unable to select Sintenis’s material. Fifteen taxa could not be typified with specimens cited in Urban’s work.
NAMES PUBLISHED BY DOMINGO BELLO Y ESPINOSA IN 1881
Magnolia portoricensis Bello in Anales Soc. Esp. Hist. Nat. 10: 233. 1881 (Magnoliaceae) – Neotype (designated here): PUERTO RICO. Prope Jayuya in sylv. mont. ad San Patricio, 14 Jun 1886, P.E.E. Sintenis 4581 (B barcode B 10 0383085; isoneotype: LD barcode 1694661).Urban (1905) considered this a good species based on
plant material collected by Stahl and one illustration made by Krug that originally were identified with Bello’s name (Electr. Suppl.: Table S1). Stahl (1884) accepted this name, although he indicated that plants of this species were unknown to him. Subsequent taxonomists working in the region (e.g., Britton & Wilson, 1924a; Liogier, 1985; Acevedo-Rodríguez & Strong, 2012) have followed Urban’s (1905) taxonomic interpretation. Magnoliaceae specialists accept Bello’s name (Magnolia Soci-ety International, 2007) and we concur with this placement. This species is endemic to Puerto Rico.
Capparis cynophallophora L. var. longifolia Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 (Capparaceae) – Type: not designated.This is an undetermined taxon within Capparaceae.Urban (1905) mentioned one plate from Krug to refer to
this taxon and considered this variety a synonym of “Capparis cynophallophora L. var. β angustifolia Eichl. in Mart.” From the brief description provided by Bello it is not certain if this taxon belongs to Cynophalla (DC.) J.Presl (as C. flexuosa (L.) J.Presl) or to Quadrella (DC.) J.Presl (as Q. cynophallophora (L.) Hutch., Capparaceae; X. Cornejo, pers. comm.). There-fore, we leave Bello’s variety as an unplaced taxon within the Capparaceae.
Capparis cynophallophora L. var. biflora Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
329
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
= Cynophalla flexuosa (L.) J.Presl. in Berchtold & Presl, Prir. Rostlin 2: 275. 1825 ≡ Morisonia flexuosa L., Pl. Jamaic. Pug.: 14. 1759 (Capparaceae) ≡ Capparis flexuosa (L.) L., Sp. Pl., ed. 2: 722. 1762.Urban (1905) examined one plate from Krug that referred
to this variety and merged this taxon within “Capparis cyno-phallophora L. var. α normalis Eichl. in Mart.” Bello’s descrip-tion highlights the presence of axillary glands, a feature of Cynophalla that does not occur in Quadrella (X. Cornejo, pers. comm.). From Bello’s description this taxon is tentatively assigned to Cynophalla flexuosa (X. Cornejo, pers. comm.). See Iltis & Cornejo (2010b) for a synopsis of Quadrella.
Capparis cynophallophora L. var. triflora Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Cynophalla flexuosa (L.) J.Presl. in Berchtold & Presl, Prir. Rostlin 2: 275. 1825 ≡ Morisonia flexuosa L., Pl. Jamaic. Pug.: 14. 1759 (Capparaceae) ≡ Capparis flexuosa (L.) L., Sp. Pl., ed. 2: 722. 1762.Urban (1905) examined one plate from Krug that referred
to this variety and merged this taxon within “Capparis cyno-phallophora L. var. α normalis Eichl. in Mart.” From Bello’s description this taxon is tentatively assigned to Cynophalla flexuosa (X. Cornejo, pers. comm.).
Capparis cynophallophora L. var. elliptica Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Cynophalla flexuosa (L.) J.Presl. in Berchtold & Presl, Prir. Rostlin 2: 275. 1825 ≡ Morisonia flexuosa L., Pl. Jamaic. Pug.: 14. 1759 (Capparaceae) ≡ Capparis flexuosa (L.) L., Sp. Pl., ed. 2: 722. 1762.Urban (1905) examined one plate from Krug that referred
to this variety and merged this taxon within “Capparis cyno-phallophora L. var. α normalis Eichl. in Mart.” From Bello’s description this taxon is tentatively assigned to Cynophalla flexuosa (X. Cornejo, pers. comm.).
Capparis cynophallophora L. var. acutifolia Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Cynophalla amplissima (Lam.) Iltis & Cornejo in Rodrigué-sia 61: 154. 2010 ≡ Capparis amplissima Lam., Encycl. 1: 607. 1785.Urban (1905) examined one plate from Krug that referred
to this variety and considered this taxon a synonym of Cap-paris portoricensis Urb. (accepted name Cynophalla amplis-sima (Lam.) Iltis & Cornejo; X. Cornejo, pers. comm.). Based on Urban’s account this name has been tentatively assigned to Cynophalla amplissima (X. Cornejo, pers. comm.).
Capparis cynophallophora L. var. mollis Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Quadrella cynophallophora (L.) Hutch., Gen. Fl. Pl. 2: 309. 1967 ≡ Capparis cynophallophora L., Sp. Pl.: 504. 1753.Urban (1905) studied one plate from Krug that referred to
this variety and considered Bello’s name as Capparis cynophal-lophora L. “Var. α normalis Eichl in Mart.”
Capparis breynia L., nom. illeg., var. atropurpurea Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Quadrella cynophallophora (L.) Hutch., Gen. Fl. Pl. 2: 309. 1967 ≡ Capparis cynophallophora L., Sp. Pl.: 504. 1753.Urban (1905) examined one plate from Krug that referred
to this variety and merged this taxon within Capparis jamai-censis Jacq. (accepted name Quadrella cynophallophora; X. Cornejo, pers. comm.). Based on Urban’s account this name has been tentatively assigned to Q. cynophallophora (X. Cornejo, pers. comm.). Iltis & Cornejo (2010a) provided a taxonomic revision for the Q. cynophallophora complex. In here we followed X. Cornejo (pers. comm.) and do not recog-nize Quadrella jamaicensis (Jacq.) J.Presl. as a distinct spe-cies but as part of Q. cynophallophora s.l. (X. Cornejo, pers. comm.).
Capparis breynia L., nom. illeg., var. rosea Bello in Anales Soc. Esp. Hist. Nat. 10: 237. 1881 – Type: not designated.
= Quadrella cynophallophora (L.) Hutch., Gen. Fl. Pl. 2: 309. 1967 ≡ Capparis cynophallophora L., Sp. Pl.: 504. 1753.See entry for Capparis breynia L. var. atropurpurea Bello
for a taxonomic discussion of this name.
Sida purpurea Bello in Anales Soc. Esp. Hist. Nat. 10: 239. 1881 (Malvaceae).This is an undetermined taxon within Malvaceae.Urban (1910) cited this taxon as a “Species dubia”, although
he examined plant material and one illustration from Krug’s collections (Electr. Suppl.: Table S1) that originally were labeled with Bello’s name. Stahl (1884) accepted the name, but he did not study any plant material belonging to this species. Britton & Wilson (1924c) also accepted Bello’s name yet indicating that it was only known from the original description and that they were not certain about its generic placement. Blanchard (O. Blanchard, pers. comm.) suggested that Bello’s original description suitably matches Melochia pyramidata L. (Malv-aceae). However, the taxon described by Bello has its flowers on terminating axillary branchlets. In contrast, M. pyrami-data has inflorescenses opposite the leaves (P.A. Fryxell, pers. comm.). Dorr (2012) suggests that the name corresponds to a species of Kosteletzkya C.Presl (Malvaceae). Because of these taxonomic uncertainties, we have decided to leave this name as an unplaced taxon within the family.
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
For interpreting Bombycella betulina and B. phoenicea (see next entry) it is necessary to examine: (1) the meaning of the question marks (“ ? ”) as quoted by Bello, and (2) the actual account provided by Bello to propose these two names. Bello used the question marks when he identified a taxon based on a name reported in the literature that he consulted, but doubted that the name corresponded to the plant he studied. In these cases he highlighted characters that differ between the taxon described in the literature and the samples he stud-ied. Bello listed these names (Hibiscus betulinus ?, H. ban-croftianus ?, H. phoeniceus ?, H. unilateralis ?) but was not sure that they were taxonomically correct nor that they were part of Hibiscus. We believe that Bello created new combi-nations; support for this comes from the clarification pro-vided by Bello himself at the end of the entries for Bomby-cella betulina and B. phoenicea. There, he stated that there are four characters robust enough to segregate Hibiscus sect. Bombycella as a distinct genus. The actual text is as follows: “me parecen caractéres suficientes para tomar como género la seccion Bombicella [sic] de los Hibiscos” (I think they are good enough characters to place section Bombicella [sic] of Hibiscus as a genus).
Bello incorrectly cited Grisebach as the species author-ity for Hibiscus bancroftianus, instead of H. bancroftianus Macfad. (accepted name H. poeppigii (Spreng.) Garcke; Dorr 2012). Urban (1910) studied two illustrations made by Krug that originally were assigned to Bombycella betulina (Electr. Suppl.: Table S1). However Urban (1910) considered Bello’s name a synonym of H. brasiliensis. Subsequent taxonomists have merged B. betulina within H. brasiliensis (Britton & Wilson, 1924c) or H. phoeniceus (Acevedo-Rodríguez & Strong, 2012). The names H. brasiliensis L. and H. phoeniceus Jacq. have been considered by some authors to refer to the same taxon. However, as indicated by Fryxell & Berazaín (2007), the former name cannot be typified as from the original description it is not clear to what plants Linnaeus (1763) referred. Therefore, Fryxell & Berazaín (2007) proposed to reject this name and instead to accept the later name H. phoeniceus for nomen-clatural stability. The Nomenclature Committee for Vascular Plants accepted the proposal (Brummitt, 2009).
Bombycella phoenicea (Jacq.) Bello in Anales Soc. Esp. Hist. Nat. 10: 241. 1881 (“Bombicella”) ≡ Hibiscus phoeniceus Jacq., Hort. Bot. Vindob. 3: 11–12. 1776.See entry for Bombycella betulina (DC.) Bello for a discus-
sion concerning the use of Hibiscus brasiliensis vs. H. phoeni-ceus. Bello (1881) recognized two morphs of B. phoenicea. The first one has “petalis splendidè purpureis”, the second form has “petalis roseis”. He was not certain if this entity should still be considered part of Hibiscus (either as H. phoeniceus or as H. unilateralis, see entry for Bombycella betulina (DC.) Bello for an explanation on the segregation of Hibiscus sect. Bombycella as a distinct genus). None of the taxonomists work-ing in the region have followed Bello’s name and they have placed B. phoenicea within Hibiscus: either as H. brasiliensis (Urban, 1910; Britton & Wilson, 1924c) or as H. phoeniceus Jacq (Stahl, 1884; Dorr, 2012). Urban (1910) studied one illustration
made by Krug that was originally identified as belonging to B. phoenicea (Electr. Suppl.: Table S1).
Gossypium janiphifolium Bello in Anales Soc. Esp. Hist. Nat. 10: 242. 1881 (Malvaceae) – Neotype (designated here): PUERTO RICO. Without location, 1889, A. Stahl 775b (US barcode 00698357).
= Gossypium hirsutum L., Sp. Pl., ed. 2: 975. 1763 ≡ Gossy-pium barbadense L. var. hirsutum (L.) Triana & Planch. in Ann. Sci. Nat., Bot., ser. 4, 17: 171. 1862 ≡ Gossypium herbaceum L. var. hirsutum (L.) Mast. in Hooker, Fl. Brit. India 1: 347. 1874.
331. 1927.= Gossypium birkinshawii G.Watt in Bull. Misc. Inform. Kew
1927: 330. 1927.= Gossypium marie-galante G.Watt in Bull. Misc. Inform.
Kew 1927: 344. 1927 ≡ Gossypium barbadense L. var. marie-galante (G.Watt) A.Chev. in Rev. Int. Bot. Appl. Agric. Trop. 18: 118. 1938 ≡ Gossypium hirsutum var. marie-galante (G.Watt) J.B.Hutch., Evol. Gossypium Different. Cult. Cottons: 43. 1947.
= Gossypium ekmanianum Wittm., Bot. Kult. Baumwolle 4(1): 174. 1928.Urban (1910) examined plant material sent to him by Krug
and Stahl that originally was labeled as Gossypium janiphi-folium (Electr. Suppl.: Table S1). Stahl (1884), Urban (1910) and Britton & Wilson (1924c) recognized G. janiphifolium as an endemic species for Puerto Rico. Nevertheless, Stahl (1884) indicated that this species was unknown to him and Britton & Wilson (1924c) stated that they were never able to find the plant in the field. Fryxell (1979) and Dorr (2012) treated this species as a synonym of G. lanceolatum Tod.; however, the latter is currently considered to be a synonym of the wide-spread G. hirsutum (P.A. Fryxell, pers. comm.). Liogier (1994) merged Bello’s species with G. hirsutum var. marie-galante (G.Watt) J.B.Hutch.
Banisteria chrysophylla Bello in Anales Soc. Esp. Hist. Nat. 10: 245. 1881 (Malpighiaceae) nom. illeg., non Lam., Encycl. 1: 368. 1785.
= Heteropterys wydleriana Juss. in Ann. Sci. Nat., Bot., sér. 2, 13: 275. 1840 ≡ Banisteria wydleriana (Juss.) C.B.Rob. in Britton, N. Amer. Fl. 25(2): 137. 1910.
= Heteropterys nitida Kunth var. obtusifolia Kuntze, Revis. Gen. Pl. 1: 88. 1891.
= Heteropterys bellonis Urb., Symb. Antill. 1: 330. 1899.Bello (1881) based this species description on sterile
331
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
material, and was doubtful about its generic status. Urban (1899) studied plant material from Krug’s collections (Electr. Suppl.: Table S1) and was aware that Bello’s name was a hom-onym of what was previously published by Lamarck. Urban described Heteropterys bellonis as a new species that was dedi-cated to Bello, to accommodate this taxon. Later, Urban (1905) considered both Banisteria chrysophylla Bello and H. bellonis as synonyms of H. wydleriana. Stahl (1884) accepted Baniste-ria chrysophylla Bello but was not familiar with the plant. He identified this species as “Stigmaphyllon (Banisteria) chryso-phylla Bello”. We are not certain if this was a new combination that Stahl proposed to include Bello’s name within Stigmaphyl-lon A.Juss. (Malpighiaceae), or if he wanted to point out that he was not sure about the generic placement of this taxon. Britton & Wilson (1924b) considered Bello’s name a synonym of B. wydleriana. Other taxonomists working in the region (e.g., Liogier, 1988; Acevedo-Rodríguez & Strong, 2012) fol-lowed Urban’s (1905) taxonomic assignment. The most recent nomenclature treatment for the family (Anderson & al., 2008) also agreed with this taxonomic interpretation for Bello’s name. This species is endemic to Puerto Rico.
= Heteropterys laurifolia (L.) Juss. in Ann. Sci. Nat., Bot., ser. 2, 13: 276. 1840 ≡ Banisteria laurifolia L., Sp. Pl., ed. 2: 611. 1762.Urban (1905) examined one illustration made by Krug
and plant material from Stahl’s and Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Stahl (1884) recognized Bello’s species. However, Urban (1905) assigned this species to synonymy under Heteropterys laurifolia as did Britton & Wilson (1924b). Other taxono-mists working in the region (e.g., Liogier, 1988; and Acevedo-Rodríguez & Strong, 2012) have followed Urban’s taxonomic interpretation. Anderson & al. (2008) considered H. laurifolia an accepted name.
Tetrapterys paniculata Bello in Anales Soc. Esp. Hist. Nat. 10: 245. 1881 (“Tetrapteris”) (Malpighiaceae) – Neotype (designated here): PUERTO RICO. Prope Bayamon, 1885, A. Stahl 338 (S No. S14-48530).
gested that it might refer to Stigmaphyllon fulgens Juss. (Malpighiaceae, accepted name S. emarginatum (Cav.) Juss.; Anderson & al., 2008; listed as “S. fulgens L.” by Stahl, 1884). Stahl (1884) accepted Bello’s name, but indicated that he was not familiar with this species. Urban (1905) did not study any plant material or illustrations referring to Bello’s name but merged Tetrapterys paniculata with T. citrifolia (Sw.) Pers., a species that he considered also to include T. inaequalis. Other taxonomists working in the region (e.g., Britton & Wilson, 1924b; Liogier, 1988; Acevedo-Rodríguez, 2005a) have fol-lowed Urban’s taxonomic interpretation. The most recent nomenclature treatment for the family (Anderson & al., 2008)
considered Bello’s name simply as Tetrapterys sp. However, we believe that T. paniculata corresponds to the Caribbean endemic T. inaequalis as it is the only species of the genus found in Puerto Rico and T. citrifolia is a species endemic to Jamaica (Acevedo-Rodríguez & Strong, 2012).
Turpinia glandulosa Bello in Anales Soc. Esp. Hist. Nat. 10: 250. 1881 (Staphyleaceae, placed in Celastraceae by Bello).This is an undetermined taxon.Urban (1910) did not study any plant material or illustra-
tions referring to Bello’s name and wrote the following state-ment for this taxon: “in dubio haeret; an hujus familiae?” (there is a doubt as to the family?). Stahl (1886) accepted Turpinia glandulosa, indicating that he was not familiar with the plant. Britton & Wilson (1924c) and Acevedo-Rodríguez & Strong (2012) also treated Bello’s species as an uncertain species. Croat (1976) included Bello’s taxon as a synonym of Turpinia occi-dentalis subsp. occidentalis and incorrectly listed two speci-mens collected by L. Picarda (790, 833) from Haiti as types.
Ilex exandra Bello in Anales Soc. Esp. Hist. Nat. 10: 251. 1881 (Aquifoliaceae, placed in Celastraceae by Bello) – Neotype (designated here): PUERTO RICO. Guajataca Gorge, Quebradillas, Apr 1985, A.H. Liogier 35506 (UPR barcode UPR 04142).
= Goetzea elegans Wydler in Linnaea 5: 423. 1830 (Solanaceae).This species is endemic to Puerto Rico, and we are not
certain why it was assigned to the Aquifoliaceae by Bello (1881). Clearly, he was not aware that this species had previ-ously been described by Wydler (1830) within the Antillean endemic genus Goetzea Wydler. Urban (1911) did not study original plant material or illustrations with Bello’s name, and yet he considered this species referrable to G. elegans. Stahl (1886) accepted Bello’s name but indicated that he was not familiar with the plant. All subsequent taxonomists working in the region (e.g., Britton & Wilson, 1925b; Liogier, 1995; Knapp, 2012) have followed Urban’s taxonomic interpretation and we concur with this view.
Anacardium occidentale L. var. rubrum Bello in Anales Soc. Esp. Hist. Nat. 10: 252. 1881 (Anacardiaceae) – Type: not designated.
= Anacardium occidentale L., Sp. Pl.: 383. 1753.Bello (1881) described two varieties within the cultivated
cashew. We place Bello’s varieties without any taxonomic rank in Anacardium occidentale.
Anacardium occidentale L. var. luteum Bello in Anales Soc. Esp. Hist. Nat. 10: 252. 1881 (Anacardiaceae) – Type: not designated.
= Anacardium occidentale L., Sp. Pl.: 383. 1753.See entry for Anacardium occidentale L. var. rubrum Bello
for a taxonomic discussion of this name.
Mangifera indica L. var. macrocarpa Bello in Anales Soc. Esp. Hist. Nat. 10: 253. 1881 (Anacardiaceae) – Type: not designated.
332
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
= Mangifera indica L., Sp. Pl.: 200. 1753.Bello (1881) described five varieties within the cultivated
mango tree, and this particular variety was also identified with the Spanish name “Mangó de puerco”. We place Bello’s variet-ies without any taxonomic rank in Mangifera indica.
Mangifera indica L. var. leiosperma Bello in Anales Soc. Esp. Hist. Nat. 10: 253. 1881 (Anacardiaceae) – Type: not designated.
= Mangifera indica L., Sp. Pl.: 200. 1753.See entry for Mangifera indica L. var. macrocarpa Bello
for a taxonomic discussion of this name. This particular vari-ety was also identified with the Spanish name “Mangó de Filipinas”.
Mangifera indica L. var. viridis Bello in Anales Soc. Esp. Hist. Nat. 10: 253. 1881 (Anacardiaceae) – Type: not designated.
= Mangifera indica L., Sp. Pl.: 200. 1753.See entry for Mangifera indica L. var. macrocarpa Bello
for a taxonomic discussion of this name.
Mangifera indica L. var. intermedia Bello in Anales Soc. Esp. Hist. Nat. 10: 253. 1881 (Anacardiaceae) – Type: not designated.
= Mangifera indica L., Sp. Pl.: 200. 1753.See entry for Mangifera indica L. var. macrocarpa Bello
for a taxonomic discussion of this name.
Mangifera indica L. var. armeniaca Bello in Anales Soc. Esp. Hist. Nat. 10: 253. 1881 (Anacardiaceae) – Type: not designated.
= Mangifera indica L., Sp. Pl.: 200. 1753.See entry for Mangifera indica L. var. macrocarpa Bello
for a taxonomic discussion of this name. This particular variety was also identified with the Spanish name “Mangó de rosa.”
Stahlia Bello in Anales Soc. Esp. Hist. Nat. 10: 255. 1881 (Fab-aceae) – Type: Stahlia maritima Bello.
Stahlia maritima Bello in Anales Soc. Esp. Hist. Nat. 10: 255. 1881 (Fabaceae) – Neotype (designated here): PUERTO RICO. Guánica, in sylvis inter Barina et la Boca, 2 Mar 1886, P.E.E. Sintenis 3876 (NY barcode 00993999; iso-neotypes: BM barcodes BM001124488 & BM001124489, G barcodes G00375840, G00375842 & G00375843, GH barcode 00460917, NY barcode 00993992, P barcodes P02770284, P02770285 & P02770286, W Herb. Mus. Palat. Vindob. Aequ. 1899 No. 6828).
= Stahlia monosperma (Tul.) Urb., Symb. Antill. 2: 285. 1900 ≡ Caesalpinia monosperma Tul. in Arch. Mus. Hist. Nat. 4: 148. 1844 – Lectotype (designated here): PUERTO RICO. Without location or date, A. Plée 713 (P barcode P03090076).Bello (1881) described the species within a new unispecific
genus dedicated to Agustín Stahl, but in doubt that a tree fairly well known for its excellent wood was not described until the publication of his Apuntes. Stahl saw living individuals of the
tree (Stahl, 1885) and accepted Bello’s name, but was doubtful of its validity. Bello was not aware that this plant had previously been described by Tulasne (1844) within Caesalpinia L. Urban (1900) subsequently proposed the combination S. monosperma. The name S. monosperma is widely accepted (e.g., Liogier, 1988; Axelrod, 2011; Acevedo-Rodríguez & Lewis, 2012). Urban (1905) studied one illustration made by Krug that was originally associated with Bello’s name (Electr. Suppl.: Table S1).
The original description of Caesalpinia monosperma refers to three specimens (Louis Claude Richard s.n. from St. Lucia, A. Plée 713, A. Plée 971 from Puerto Rico), all of which are deposited in P. We designated Plée 713 as the lecto-type because this collector is repeatedly cited throughout the original morphological description, and because it is the only of Plée’s specimens annotated by Tuslane. The genus Stahlia is considered as endemic to Puerto Rico and the Dominican Republic, although there is a syntype specimen of Caesalpinia monosperma collected by L.C. Richard (P, P03090065) attrib-uted to the island of St. Lucia.
Tephrosia aniloides Bello in Anales Soc. Esp. Hist. Nat. 10: 258. 1881 (Fabaceae) ≡ Cracca aniloides (Bello) O.F.Cook & G.N.Collins in Contr. U.S. Natl. Herb. 8: 128. 1903 (Fabacaeae) – Neotype (designated here): PUERTO RICO. Coamo, in sylvis, 28 Dec 1885, P.E.E. Sintenis 3265 (NY barcode 00988693).
= Coursetia caribaea (Jacq.) Lavin var. caribaea, Adv. Legume Syst. 3: 63. 1987 (Fabaceae) ≡ Galega caribaea Jacq., Select. Stirp. Amer. Hist.: 212. 1763 (Fabaceae) ≡ Tephrosia caribaea (Jacq.) DC., Prodr. 2: 251. 1825 ≡ Cracca carib-aea (Jacq.) Benth. ex Oerst. in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1853(1–2): 9. 1854 ≡ Brit-tonamra caribaea (Jacq.) Kuntze, Revis. Gen. Pl. 1: 164. 1891 (Fabaceae) ≡ Benthamantha caribaea (Jacq.) Kuntze, Revis. Gen. Pl. 3(3): 53. 1898 (Fabaceae).Urban (1905) examined plant material from Stahl’s col-
lections (Electr. Suppl.: Table S1) that originally was identi-fied as Tephrosia aniloides, but Urban did not accept Bello’s name, merging it with Cracca caribaea. Stahl (1885) suggested that T. aniloides should be placed in another genus, but with-out giving an alternative name. Taxonomists working in the region have assigned Bello’s name to Benthamantha caribaea (Britton & Wilson, 1924b), Cracca caribaea (Liogier, 1988), or Coursetia caribaea (Acevedo-Rodríguez & Lewis, 2012). We accept the nomenclature proposed by Lavin (1988) in his monograph of Coursetia DC.
Aeschynomene fistulosa Bello in Anales Soc. Esp. Hist. Nat. 10: 259. 1881 (Fabaceae) – Neotype (designated here): PUERTO RICO. Mayagüez, ad margines fossarum, 22 Oct 1884, P.E.E. Sintenis 94 (US barcode 00711161).
= Aeschynomene sensitiva Sw. var. sensitiva, Prod.: 107. 1788.Urban (1905) examined two illustrations made by Krug that
originally were labeled with Bello’s name (Electr. Suppl.: Table S1). However, Urban (1905) assigned this to Aeschynomene sensitiva. Stahl (1885) gave little credit to this species since it was described on the basis of sterile material that easily could
333
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
be confused with other members of the genus. Subsequent tax-onomists working in the region (e.g., Britton & Wilson, 1924b; Acevedo-Rodríguez & Lewis, 2012) and legume specialists (Rudd, 1955) have followed Urban’s taxonomic interpretation for Bello’s name.
Cajanus luteus Bello in Anales Soc. Esp. Hist. Nat. 10: 260. 1881 (Fabaceae) – Neotype (designated here): PUERTO RICO. Lares, ad Perchas cult., 21 Jan 1887, P.E.E. Sintenis 6015 (US barcode 00712515).
Bello (1881) was uncertain about the placement of this spe-cies and even suggested that it might be a mere variety of Caja-nus indicus. He also indicated that this was a cultivated species. Indeed, C. cajan is the only species of the genus reported for Puerto Rico (Acevedo-Rodríguez & Strong, 2012). Urban (1905) did not study plant material or illustration referring to Bello’s collection, but merged this species with C. indicus. Stahl (1885) also considered Bello’s taxon to be a synonym of C. indicus. Subsequent taxonomists working in the region (e.g., Britton & Wilson, 1924b; Liogier, 1988; Acevedo-Rodríguez & Lewis, 2012) have regarded Bello’s species as a synonym of C. cajan, an opinion agreed with by L. Rico (pers. comm.).
Phaseolus lanceolatus Bello in Anales Soc. Esp. Hist. Nat. 10: 262. 1881 (Fabaceae) – Neotype (designated here): PUERTO RICO. Pueblo Viejo, 19 Jul 1914, J.A. Stevenson 2097 (NY barcode 01007063).
= Vigna trichocarpa (C.Wright) A.Delgado in Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1254. 1993 (Fabaceae) ≡ Phaseolus trichocarpus C.Wright in Anales Acad. Ci. Med. Habana 5: 337. 1869.Urban (1905) examined one illustration and plant mate-
rial from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1) and assigned this spe-cies to Phaseolus ovatus Benth. Stahl (1885) accepted Bello’s name, although he indicated that he did not study plant material. Britton & Wilson (1924b) assigned the species to P. trichocar-pus. Other taxonomists working in the region (e.g., Liogier, 1988; Acevedo-Rodríguez, 2005a; Acevedo-Rodríguez & Lewis, 2012) have considered Bello’s name a synonym of Vigna longifolia (Benth.) Verdc. However, Delgado-Salinas & al. (in prep.) indicated that the lanceolate and peltate stipules reported in Bello’s description clearly suggest that this species should be assigned to Vigna trichocarpa.
Phaseolus cochleatus Bello in Anales Soc. Esp. Hist. Nat. 10: 262. 1881, nom. illeg., non Vell., Fl. Flumin.: 312. 1829.
Maréchal, Mascherpa & Stainier in Taxon 27: 202. 1978.Urban (1905) examined one illustration and plant mate-
rial from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). He considered this spe-cies to be a synonym of Phaseolus adenanthus. This place-ment was accepted by Britton & Wilson (1924b). Stahl (1885) included Bello’s name in his work but did not study material of this species. Acevedo-Rodríguez & Lewis (2012) followed Urban’s taxonomic interpretation, placing it within Vigna Savi as V. adenantha.
Phaseolus cochleatus Bello var. pallidus Bello in Anales Soc. Esp. Hist. Nat. 10: 262. 1881 (“pallida”) – Type: not designated.
= Leptospron adenanthum (G.Mey.) A.Delgado in Amer. J. Bot. 98: 1710. 2011 ≡ Phaseolus adenanthus G.Mey., Prim Fl. Esseq.: 239. 1818 ≡ Vigna adenantha (G.Mey.) Maréchal, Mascherpa & Stainier in Taxon 27: 202. 1978.Bello (1881) distinguished two varieties within Phaseolus
cochleatus (“α violacea”, “β pallida”). Urban (1905) only rec-ognized var. violacea as a synonym of P. adenanthus. For var. pallida, Urban stated: “mihi ignotus” (it is unknown to me). We consider these two morphs as mere variants within Leptospron adenanthum and we do not give them any taxonomic recogni-tion. We have followed the latest taxonomic arrangements sug-gested for this group by Delgado-Salinas & al. (2011).
Phaseolus cochleatus Bello var. violaceus Bello in Anales Soc. Esp. Hist. Nat. 10: 262. 1881 (“violacea”) – Type: not designated.
= Leptospron adenanthum (G.Mey.) A.Delgado in Amer. J. Bot. 98: 1710. 2011 ≡ Phaseolus adenanthus G.Mey., Prim Fl. Esseq.: 239. 1818 ≡ Vigna adenantha (G.Mey.) Maréchal, Mascherpa & Stainier in Taxon 27: 202. 1978.See our discussion under Phaseolus cochleatus Bello var.
pallidus Bello.
Acacia leptosperma Bello in Anales Soc. Esp. Hist. Nat. 10: 265. 1881 (Fabaceae) – Neotype (designated here): PUERTO RICO. Inter Algarrobo et Cari-blanco, 2 Dec 1885, P.E.E. Sintenis 2931 (US barcode 00710660).
= Acuan bahamense Britton & Rose in Britton, N. Amer. Fl. 23(2): 132. 1928 (Fabaceae).
= Desmanthus strictus Bertol. in Giorn. Arcadico Sci. 21: 190. 1824 ≡ Desmanthus virgatus (L.) Willd. var. strictus (Bertol.) Griseb., Fl. Brit. W. I.: 218. 1860.Urban (1905) examined one illustration and plant material
from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1), concluding that this species is a synonym of Desmanthus virgatus var. strictus. Stahl (1885) admitted that he was not familiar with the plant, and he mistak-enly rendered the name as “Albizzia leptosperma Bello” (Faba-ceae). We are not certain if this was a typographical error and, therefore, Stahl (1885) instead referred to Acacia leptosperma
334
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Bello. Another possibility is that he intended to transfer the taxon to Albizia Durazz, thus making a new combination under this genus. Britton & Wilson (1924b) considered Bello’s name a synonym of Acuan virgatum (L.) Medik. (accepted name D. vir-gatus (L.) Willd.; Luckow, 1993). Both Liogier (1988) and Ace-vedo-Rodríguez & Lewis (2012) merged Acacia leptosperma with D. virgatus. In her taxonomic monograph of Desmanthus Willd., Luckow (1993) placed Bello’s taxon under the category of “doubtful and excluded names” and tentatively considered this species a synonym of D. virgatus. However, in our study we follow Urban’s taxonomic interpretation of this species since he examined material labeled with Bello’s name.
Jussiaea plumeriana (DC.) Bello in Anales Soc. Esp. Hist. Nat. 10: 267. 1881 (Onagraceae) ≡ Jussiaea erecta L. var. plumeriana DC., Prodr. 3: 55. 1828.
= Jussiaea acuminata Sw., Fl. Ind. Occid. 2(1): 745. 1798.Bello (1881) listed two names for this taxon: “Jussiaea
erecta, DC. var. plumeriana” (without any description and without an author for the varietal rank; p. 266) and “J. plume-riana” (with a Latin description but without an author or aster-isk). It is unclear why Bello used two different names for this taxon. We believe that there was a typographic error, and that it is likely that with this variety Bello intended to refer to Jussiaea erecta L. var. plumeriana DC., as the name “Jussiaea erecta DC.” was not published by Augustin de Candolle. We inter-pret Bello’s name, J. plumeriana, as a new combination (basi-onym: J. erecta L. var. plumeriana DC.). Urban (1910) exam-ined one illustration made by Krug that was labeled J. erecta var. plumeriana (Electr. Suppl.: Table S1) and considered this variety as well as J. plumeriana to be synonyms of J. erecta. Britton & Wilson (1925a) followed Urban’s taxonomic inter-pretation. Stahl (1886) erroneously assigned J. plumeriana to Augustin de Candolle, but merged this taxon within J. acu-minata. Subsequent taxonomists and Onagraceae specialists (e.g., Raven, 1963; Liogier, 1995; Acevedo-Rodríguez & Strong, 2012; P. Hoch, pers. comm.) considered Bello’s combination a synonym of Ludwigia erecta.
Homonoma Bello in Anales Soc. Esp. Hist. Nat. 10: 269. 1881 (Melastomataceae) – Type: H. aridum Bello.
= Nepsera Naudin in Ann. Sci. Nat., Bot., sér. 3, 13: 28. 1850 (Melastomataceae).
Homonoma aridum Bello in Anales Soc. Esp. Hist. Nat. 10: 269. 1881 – Neotype (designated here): PUERTO RICO. Sa. [Sierra] de Luquillo, in monte Jimenes, Jun 1885, P.E.E. Sintenis 1487 (NY barcode 01312866).
= Spennera sieberi Steud. in Flora 27: 720. 1844 ≡ Aciotis sie-beri (Steud.) Triana in Trans. Linn. Soc. London 28(1): 52. 1871.Urban (1905) examined one illustration and plant material
from Krug’s collections that originally were labeled with Bel-lo’s name (Electr. Suppl.: Table S1). He considered the species to be a synonym of Nepsera aquatica. Subsequent taxonomists working in the region and with Melastomataceae (e.g., Stahl, 1886; Britton & Wilson, 1925a; Liogier, 1995; Michelangeli & Bécquer-Granados, 2012; W. Judd, pers. comm.) have fol-lowed Urban’s taxonomic interpretation of Bello’s name.
Pimenta acuminata Bello in Anales Soc. Esp. Hist. Nat. 10: 270. 1881 (Myrtaceae) – Neotype (designated here): PUERTO RICO. Maunabo, ad Emajagua versus, 18 Sep 1886, P.E.E. Sintenis 5061 (NY barcode 00819380).
pimento (O.Berg) Griseb., Fl. Brit. W. I.: 241. 1860.Urban (1910) examined plant material of this species sent to
him by Krug (Electr. Suppl.: Table S1) that originally was iden-tified with Bello’s name, and considered this to be a synonym of Amomis caryophyllata. Britton & Wilson (1925a) followed Urban’s taxonomic treatment. Stahl (1886) recognized Pimenta acuminata although he indicated that he was not familiar with the plant. Other subsequent taxonomists (e.g., Liogier, 1994; Acevedo-Rodríguez & Strong, 2012) have considered Bello’s name a synonym of P. racemosa var. racemosa. Landrum (1986), in his monograph of Pimenta Lindl., used a question mark for the placement of P. acuminata. In our study we fol-low Urban’s treatment, as he studied plant material originally identified with Bello’s name.
Eugenia calyculata Bello in Anales Soc. Esp. Hist. Nat. 10: 271. 1881 (Myrtaceae) – Neotype (designated here): PUERTO RICO. Luquillo, highway to Bella Vista, 10 Aug 1950, E.L. Little Jr. 13596 (UPR barcode UPR 08568).
= Eugenia domingensis O.Berg in Linnaea 27: 296. 1856.Urban (1910) examined one illustration and plant material
from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). He considered this species to be a synonym of Eugenia aeruginea DC. However, Urban (1910) misapplied E. aeruginea to the taxon known as E. domingensis (Sandwith, 1934). Britton & Wilson (1925a) also assigned this species to E. aeruginea sensu Urban (1910); Stahl (1886) sug-gested that this species may correspond to E. portoricensis DC. (accepted name E. pseudopsidium Jacq.; Acevedo-Rodrí-guez & Strong, 2012), indicating that original material was not studied by him. Liogier (1994), Govaerts & al. (2008), Acevedo-Rodríguez & Strong (2012), and F. Barrie (pers. comm.) sug-gested that Bello’s species is a synonym of E. domingensis.
335
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
Eugenia tetrasperma Bello in Anales Soc. Esp. Hist. Nat. 10: 271. 1881 – Neotype (designated here): PUERTO RICO. On a ravine on the north side of road no. 3, km 109.6, between Maunabo and Arroyo, 26 Jul 1972, J.L. Vivaldi 72-186 (UPR barcode UPR 08048).
= Aulacocarpus quadrangularis Griseb., Fl. Brit. W. I.: 239. 1864 (Melastomataceae).Urban (1910) examined one illustration and plant material
from Krug’s collections that originally were identified with Bello’s name (Electr. Suppl.: Table S1). He regarded this spe-cies to be a synonym of Mouriri domingensis (as “Mouriria”). Stahl (1886) considered Eugenia tetrasperma to be a distinct species. Subsequent taxonomists (e.g., Britton & Wilson, 1925a; Liogier, 1995; Michelangeli & Bécquer-Granados, 2012) have followed Urban’s taxonomic interpretation of Bello’s name. Morley (1976) in his monograph of tribe Memecyleae (Melas-tomataceae) also agreed with Urban’s taxonomic placement although indicating, based on Bello’s description, that some of the morphological traits of this species are of dubious applica-tion to Mouriri Aubl.
Eugenia paniculata Bello in Anales Soc. Esp. Hist. Nat. 10: 271. 1881, nom. illeg., non Jacq., Collectanea 2: 108. 1789.This is an undetermined taxon within Eugenia.Urban (1910) examined one illustration from Krug’s col-
lections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). He was uncertain of the identity of this material and reported it as “Species dubia”. Previously, Urban (1895a) suggested that this taxon might be a synonym of Euge-nia eggersii Kiaersk. Stahl (1886) recognized E. paniculata Bello, although he did not study any plant material. Britton & Wilson (1925a) stated that the species “has not been identified by recent botanists”. We are certain that Bello’s name does not refer to E. paniculata Jacq. (accepted name Myrcia guianensis (Aubl.) DC.; Acevedo-Rodríguez & Strong, 2012), a species that occurs in Puerto Rico (Acevedo-Rodríguez & Strong, 2012), because Myrcia DC. ex Guill. has 5-merous flowers and an embryo with well-defined, leafy cotyledons. Bello’s description of his E. paniculata referred to a species with 4-merous flow-ers and undifferentiated embryos, suggesting that it belongs to Eugenia L. From the brief description provided by Bello (1881) it is not obvious to what species of Eugenia he was referring. Therefore, we have considered this name as an undetermined taxon within this genus.
Eugenia costata Bello in Anales Soc. Esp. Hist. Nat. 10: 272. 1881, nom. illeg., non Cambess. in Saint-Hilaire, Fl. Bras. Merid. 2: 359. 1833 ≡ Eugenia serrasuela Krug & Urb. in Bot. Jahrb. Syst. 19: 652. 1895.This is an undetermined taxon within Eugenia.Urban (1910) considered this species to be a synonym of
Eugenia serrasuela, but with no mention to plant material
nor illustrations. The only locality cited (“Prope Anones”) is based on what is reported by Bello in the Apuntes. Previously, Krug and Urban realized that Bello’s name was a homonym of E. costata and proposed the new name E. serrasuela for this poorly known Puerto Rican endemic without making reference to any herbarium collections (Urban, 1895a). Stahl (1886) rec-ognized E. costata Bello (although he was not familiar with the species) and suggested that it belonged to “Autocarpus Berg.”, a genus that we have not found in the taxonomic literature. Subse-quent taxonomists (e.g., Britton & Wilson, 1925a; Liogier, 1994) followed Urban’s taxonomic interpretation of Bello’s name, but indicating that E. sarrasuela was only known from the species descriptions provided by Bello (1881) and Urban (1895a, 1910). As noticed by Liogier (1994) and F. Barrie (pers. comm.), the description provided by Bello (1881) is confusing because he described plants with 5-merous flowers (only found in anoma-lous flowers of Eugenia), but with a solid and undifferentiated embryo (as typical in Eugenia). Therefore, we are not certain to which taxon Bello referred. Because we cannot rule out that Bello observed plants of Eugenia with abnormal flowers, we have chosen to assign this name to an undetermined species within this genus.
Psidiastrum Bello in Anales Soc. Esp. Hist. Nat. 10: 272. 1881 (Myrtaceae) – Type: Psidiastrum dubium Bello.
= Eugenia L., Sp. Pl.: 470. 1753.
Psidiastrum dubium Bello in Anales Soc. Esp. Hist. Nat. 10: 272. 1881 – Neotype (designated here): PUERTO RICO. Guánica, Cóbanas to Barinas, 11 Oct 1940, L.E. Gregory 208 (UPR barcode UPR 08569).
= Eugenia yumuryensis O.Berg in Linnaea 27: 234. 1856.= Eugenia matanzensis O.Berg in Linnaea 27: 235. 1856.
Urban (1910) examined one illustration made by Krug that was labeled with Bello’s name (Electr. Suppl.: Table S1), and assigned this species to Eugenia axillaris. Stahl (1886) accepted Psidiastrum dubium, but indicated that Bello’s morphological observations were erroneous, and what Bello (1881) consid-ered to be many seeds were in reality fruit debris that resulted from insect damage. Still, Stahl (1886) stated that he was not familiar with this species, but indicated that Bello’s description matched that of E. flavovirens O.Berg. According to Acevedo-Rodríguez & Strong (2012) Stahl’s concept of E. flavovirens is referrable to E. axillaris. Subsequent taxonomists and Euge-nia specialists (e.g., Britton & Wilson, 1925a; Liogier, 1994; Acevedo-Rodríguez & Strong, 2012; F. Barrie, pers. comm.) followed Urban’s taxonomic interpretation of Bello’s name.
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
(designated here): PUERTO RICO. Dorado, 11 Feb 1959, R.O. Woodbury s.n. (UPR barcode UPR 08580)
= Cayaponia americana (Lam.) Cogn. in Candolle & Candolle, Monogr. Phan. 3: 785. 1881 ≡ Bryonia americana Lam., Encycl. 1: 498. 1785 (Cucurbitaceae).
= Trichosanthes tamnifolia Poir. in Lamarck, Encycl., Suppl. 1: 386. 1810 (Cucurbitaceae).Urban (1911) studied one illustration made by Krug that
originally was labeled with Bello’s name (Electr. Suppl.: Table S1) and considered this species to be a synonym of Cayaponia americana (Lam.) Cogn. var. angustiloba Cogn. (accepted name Cayaponia americana; Acevedo-Rodríguez & Strong, 2012). The original description of var. angustiloba did not refer to Bello. Stahl (1886) accepted Cionandra angustiloba as a distinct species but indicated that he was not familiar with this plant. Acevedo-Rodríguez & Strong (2012) and Nee (unpub.) considered Bello’s name a synonym of Cayaponia americana.
Turnera ovata Bello in Anales Soc. Esp. Hist. Nat. 10: 275. 1881 (Turneraceae) ≡ Piriqueta ovata (Bello) Urb. in Jahrb. Königl. Bot. Gart. Berlin 2: 76. 1883 (Turneraceae) – Neo-type (designated here): PUERTO RICO. In thickets, near seashore, Pitahaya, Cabo Rojo, 21 Feb 1984, A.H. Liogier & al. 34929 (UPR barcode UPR 08579).
that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Previously, Urban (1883) accepted Bello’s name as the basionym of Piriqueta ovata. Stahl (1886) followed Urban’s interpretation for this taxon, although indicating that he was not familiar with the species. Both Britton & Wilson (1924c) and Liogier (1994) also considered P. ovata a distinct species. We have followed the taxonomic conclusions by Arbo (1995) who monographed Piriqueta Aubl. and suggested that Bello’s name is a synonym of P. racemosa. Acevedo-Rodríguez & Strong (2012) also agreed with this taxonomic placement.
Cereus quadricostatus Bello in Anales Soc. Esp. Hist. Nat. 10: 276. 1881 (Cactaceae) ≡ Leptocereus quadricostatus (Bello) Britton & Rose in Contr. U.S. Natl. Herb. 16: 242. 1913– Neotype (designated here): PUERTO RICO. Prope Guayanilla in montibus calcareis ad Los Indios, 29 Aug 1886, P.E.E. Sintenis 4919 (B barcode B 10 0383086; isoneo-types: BM barcode BM001124486, G barcode G00375848, GH barcode 00359288, K barcode K000251796, W Herb. Mus. Palat. Vindob. Aequ. No. 132).Urban (1910) studied one illustration made by Krug that
originally was labeled with Bello’s name (Electr. Suppl.: Table S1) and accepted Cereus quadricostatus as a distinct species. Stahl (1886) also cited the name, although he stated that he was not familiar with the species. Subsequent taxono-mists and Cact aceae specialists (e.g., Britton & Rose, 1913; Britton & Wilson, 1924c; Liogier, 1994; Hunt, 2006; Acevedo- Rodríguez & Strong, 2012) have considered Leptocereus quad-ricostatus to be the correct name for this species.
Cereus leiocarpus Bello in Anales Soc. Esp. Hist. Nat. 10: 276. 1881 (Cactaceae) – Neotype (designated here): PUERTO RICO. In coastal thickets, El Tuque, Ponce, 25 Nov 1980, A.H. Liogier & al. 31238 (UPR barcode UPR 08578).
= Pilosocereus royenii (L.) Byles & G.D.Rowley in Cact. Succ. J. Gr. Brit. 19(3): 67. 1957 (Cactaceae) ≡ Cactus royenii L., Sp. Pl.: 467. 1753 ≡ Cephalocereus royenii (L.) Britton & Rose in Contr. U.S. Natl. Herb. 12: 419. 1909.
= Cereus nobilis Haw., Syn. Pl. Succ.: 179. 1812 ≡ Cephalo-cereus nobilis (Haw.) Britton & Rose in Contr. U.S. Natl. Herb. 12: 418. 1909 ≡ Pilosocereus nobilis (Haw.) Byles & G.D.Rowley in Cact. Succ. J. Gr. Brit. 19: 67. 1957.
= Cereus monoclonos DC., Prodr. 3: 464. 1828 ≡ Cephalocer-eus monoclonos (DC.) Britton & Rose in Contr. U.S. Natl. Herb. 12: 418. 1909.
= Cereus swartzii Griseb., Fl. Brit. W. I.: 301. 1860 ≡ Cepha-lo cereus swartzii (Griseb.) Britton & Rose in Contr. U.S. Natl. Herb. 12: 420. 1909.
= Pilocereus urbanianus K.Schum., Gesamtbeschr. Kakt.: 193. 1897 ≡ Pilosocereus urbanianus (K.Schum.) Byles & G.D.Rowley in Cact. Succ. J. Gr. Brit. 19: 69. 1957.
= Cephalocereus millspaughii Britton in Contr. U.S. Natl. Herb. 12: 417. 1909 ≡ Pilosocereus millspaughii (Britton) Byles & G.D.Rowley in Cact. Succ. J. Gr. Brit. 19: 67. 1957.
= Cephalocereus barbadensis Britton & Rose, Cact. 2: 44. 1920 ≡ Pilosocereus barbadensis (Britton & Rose) Byles & G.D.Rowley in Cact. Succ. J. Gr. Brit. 19: 66. 1957.Urban (1910) studied one illustration from Krug’s collec-
tions that originally were labeled with Bello’s name (Electr. Suppl.: Table S1), and considered this species to be a syn-onym of Pilocereus royenii (as “royeni”). Earlier, Stahl (1886) regarded Bello’s species as a synonym of Cereus swartzii. Britton & Wilson (1924c) placed Cereus leiocarpus in syn-onymy under Cephalocereus royenii. Other taxonomic special-ists followed Urban’s interpretation for this species (Liogier, 1994; Acevedo-Rodríguez & Strong, 2012). Bello’s description indicated that his species has smooth fruits, and P. royenii is the only cereoid cactus found in Puerto Rico with that diagnostic character (C.M. Taylor, pers. comm.).
Opuntia repens Bello in Anales Soc. Esp. Hist. Nat. 10: 277. 1881 (Cactaceae) – Neotype (designated here): PUERTO RICO. Guánica, 23 Feb 1886, P.E.E. Sintenis 4019 (G bar-code G00375847; isoneotypes: BM barcode BM001124487, GH barcode 00303867, K barcode K000035295, MO bar-code MO-133051, NY barcode 00873852, W Herb. Mus. Palat. Vindob. Aequ. 1900 No. 454).The name “Opuntia repens Karw.” (in Salm-Dyck, Hort.
Dyck.: 361. 1834) is a nomen nudum as it was published without a description (Crook & Mottram, 2002). In addition, this name referred to plants from Mexico (Salm-Reifferscheid-Dyck, 1834) that do not occur in Puerto Rico. Opuntia repens is a spe-cies endemic to Puerto Rico and the Virgin Islands (Acevedo-Rodríguez & Strong, 2012). Bello (1881) suggested that this
337
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
species might be a variety of O. spinosissima Mill. (accepted name Consolea spinosissima (Mill.) Lem., Cact aceae; Hunt, 2006). Urban (1910) examined one illustration made by Krug that originally was labeled with Bello’s name (Electr. Suppl.: Table S1). Opuntia repens has been widely accepted by other taxonomists working in the region and by Cactaceae special-ists (e.g., Stahl, 1886 [indicating that he was not familiar with flowers/fruits of the species]; Britton & Wilson, 1924c; Liogier, 1994; Hunt, 2006; Acevedo-Rodríguez & Strong, 2012).
Viscum randiae Bello in Anales Soc. Esp. Hist. Nat. 10: 278. 1881 (Santalaceae, placed by Bello in Loranthaceae) ≡ Phoradendron randiae (Bello) Britton in Sci. Surv. Porto Rico Virgin Islands 5: 257. 1924 (Santalaceae) – Neotype (designated here): PUERTO RICO. Cabo-Rojo, in “Monte Grande”, 16 Jan 1885, P.E.E. Sintenis 885 (US barcode 00706638).
= Phoradendron quadrangulare (Kunth) Griseb., Fl. Brit. W. I.: 711. 1864 ≡ Loranthus quadrangularis Kunth in Humboldt & al., Nov. Gen. Sp. (quarto ed.) 3: 444. 1820.
= Phoradendron quadrangulare (Kunth) Griseb. var. gracile Krug & Urb. in Bot. Jahrb. Syst. 24: 37. 1897 ≡ Phoraden-dron gracile (Krug & Urb.) Trel., Phoradendron: 112. 1916
= Phoradendron antillarum Trel. var. antillarum, Phoraden-dron: 111. 1916.
= Phoradendron antillarum Trel. var. orientale Trel., Phora-dendron: 112. 1916 (“orientalis”) ≡ Phoradendron randiae (Bello) Britton var. orientale (Trel.) Stehlé in Bull. Soc. Bot. France, Mem. 1953–1954: 25. 1954.
= Phoradendron moringicola Trel. in Repert. Spec. Nov. Regni Veg. 26: 343. 1929.
= Phoradendron rehderianum Urb. in Ark. Bot. 23A(5): 62. 1930.Urban (1905) examined one illustration made by Krug that
originally was labeled with Bello’s name (Electr. Suppl.: Table S1). However, he considered this name to be a synonym of Pho-radendron quadrangulare. Britton & Wilson (1924a) accepted Bello’s name as the basionym of P. randiae, and Liogier (1985) also followed this taxonomic arrangement. Subsequent taxono-mists working in the region agreed with Urban’s taxonomic inter-pretation for this species (e.g., Liogier, 1997; Kuijt, 2003, 2012).
Coffea arabica L. var. stenophyllea Bello in Anales Soc. Esp. Hist. Nat. 10: 280. 1881 (Rubiaceae) – Type: not designated.
= Coffea arabica L., Sp. Pl.: 200. 1753.Bello (1881) described this variety for the cultivated coffee
tree, and we place this taxon without any taxonomic rank in Coffea arabica.
Psychotria pseudopavetta Bello in Anales. Soc. Esp. Hist. Nat. 10: 281. 1881 (“pseudo-pavetta”) (Rubiaceae) ≡ Uragoga pseudopavetta (Bello) M.Gómez, Dicc. Bot. Nom. Vulg. Cub. Puerto-Riqu.: 85. 1889 (Rubiaceae) – Neotype (des-ignated here): PUERTO RICO. Juncos, in monte Goyo, 24 Aug 1885, P.E.E. Sintenis 2810 (NY barcode 01328094).
Urban (1911) examined one illustration made by Krug and plant material collected by Stahl that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Urban considered the species to be a synonym of Palicourea domingensis. Britton & Wilson (1925a) agreed with Urban’s taxonomic placement, while Stahl (1887) regarded Bello’s name as a synonym of Palicourea pavetta. Subsequent taxonomists working in the region (e.g., Liogier, 1997; Acevedo-Rodríguez & Strong, 2012) interpreted Bello’s name as a synonym of Psychotria domin-gensis. Taylor & al. (2010) proposed that the correct name for Psychotria domingensis is Palicourea domingensis, and in our study we follow their taxonomic conclusions.
Geophila cordata Bello in Anales Soc. Esp. Hist. Nat. 10: 282. 1881 (Rubiaceae), nom. illeg., non Miq. in Linnaea 17: 72. 1843.
Urban (1911) examined one illustration made by Krug that originally was labeled with Bello’s name (Electr. Suppl.: Table S1), and considered this species to be a synonym of Geophila herbacea (accepted name G. repens; Johnston, 1949). In addition, Urban (1911) indicated that Bello’s name did not refer to G. cor-data Miq. The latter was described from Suriname and has gen-erally been treated as a synonym of G. repens (Steyermark, 1972; C.M. Taylor, pers. comm.). Britton & Wilson (1925a) agreed with Urban’s placement of this species. Acevedo-Rodríguez & Strong (2012) regarded Bello’s species as a synonym of G. repens. Taylor (C.M. Taylor, pers. comm.) agreed with this taxonomic place-ment, as it is the only species of the genus present in Puerto Rico and because of its distinct cordate leaves.
Cephaelis triplocephala Bello in Anales Soc. Esp. Hist. Nat. 10: 282. 1881 (Rubiaceae) ≡ Uragoga triplocephala (Bello) M.Gómez, Dicc. Bot. Nom. Vulg. Cub. Puerto-Riqu.: 102. 1889 (Rubiaceae) – Neotype (designated here): PUERTO RICO. N. side Luquillo Mts., wet woods, 13 Apr 1899, A.A. Heller 1087 (NY barcode 01327633).
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Urban (1911) examined one illustration made by Krug that originally was labeled with Bello’s name (Electr. Suppl.: Table S1), and considered this species a synonym of Psychotria uliginosa. Stahl (1887) accepted Cephaelis triplocephala, but was not familiar with this plant. Britton & Wilson (1925b) and Liogier (1997) agreed with Urban’s taxonomic placement for this species. Taylor (2001) proposed that P. uliginosa should be transferred to Notopleura uliginosa, and Acevedo-Rodríguez & Strong (2012) placed Bello’s name as a synonym of the latter.
Diodia nitens Bello in Anales Soc. Esp. Hist. Nat. 10: 283. 1881 (Rubiaceae) – Neotype (designated here): PUERTO RICO. Mayagüez, along the beach, 2 Feb 1900, A.A. Heller 4508 (NY barcode 00875529).
= Diodia radicans Cham. & Schltr. in Linnaea 3: 350. 1828.Like Stahl (1887), Urban (1911) did not study any plant
material or illustrations that could be referred to Bello’s species. However, Urban considered the species a synonym of Diodia maritima. Based on the species description, Stahl (1887) also indicated that Diodia nitens was morphologically similar to Diodia sarmentosa Sw. (accepted name Diodella sarmentosa (Sw.) Bacigalupo & Cabral ex Borhidi; Borhidi, 2006) sug-gesting that Bello’s species was not distinct. Britton & Wilson (1925) agreed with Urban’s placement for this species. Liogier (1997) and Acevedo-Rodríguez & Strong (2012) followed the nomenclatural treatment published by Exell (1944) and adopted Diodia maritima as a synonym of Diodia serrulata. The latest taxonomic treatment was made by Borhidi (2006) who consid-ered Diodia serrulata to be a synonym of Diodella serrulata.
Borreria alternans Bello in Anales Soc. Esp. Hist. Nat. 10: 283. 1881 (Rubiaceae) ≡ Spermacoce alternans (Bello) Kuntze, Revis. Gen. Pl. 3(2): 123. 1898 – Neotype (desig-nated here): PUERTO RICO. Sierra de Luquillo in monte Jimenes, 28 May 1885, P.E.E. Sintenis 1647 (BM barcode BM000832499).
= Spermacoce ocymifolia Willd. ex Roem. & Schult., Syst. Veg. 3: 530. 1818 ≡ Hemidiodia ocymifolia (Willd. ex Roem. & Schult.) K.Schum. in Martius, Fl. Bras. 6(6): 29. 1888 (Rubiaceae) ≡ Diodia ocymifolia (Willd. ex Roem. & Schult.) Bremek. in Recueil Trav. Bot. Neerl. 31: 305. 1934.
= Spermacoce portoricensis Balb. ex DC., Prodr. 4: 552. 1830.Urban (1911) did not study any plant material or illustrations
that could be referred to Bello’s species, and yet he considered the species to be a synonym of Hemidiodia ocymifolia. Britton & Wilson (1925b) agreed with Urban’s placement while Stahl (1887) assigned Bello’s name to synonymy under Spermacoce
portoricensis. Liogier (1997) assigned the name to synonymy under Diodia ocymifolia. More recently Acevedo-Rodríguez & Strong (2012) and C.M. Taylor (pers. comm.) followed the recommendations of Govaerts & al. (2013) and considered Bello’s species a synonym of S. ocymifolia.
Emilia sonchifolia (L.) DC. var. rosea Bello in Anales Soc. Esp. Hist. Nat. 10: 286. 1881 (Asteraceae) – Type: not designated.
= Emilia fosbergii Nicolson in Phytologia 32: 34. 1975.Urban (1911) did not recognize this variety and assigned
this species to synonymy under Emilia sonchifolia. Here we fol-low the latest published account for West Indian Asteraceae by Robinson & Funk (2012), who did not recognize Bello’s name and merged this variety within Emilia fosbergii.
Conradia pedunculosa DC. var. pallida Bello in Anales Soc. Esp. Hist. Nat. 10: 288. 1881 (Gesneriaceae) – Type: not designated.
= Gesneria pedunculosa (DC.) Fritsch in Engler & Prantl, Nat. Pflanzenfam. 4(3b): 184. 1894 ≡ Conradia pedunculosa DC., Prodr. 7: 525. 1839.Bello (1881) described two varieties within Conradia
pedunculosa that were assigned to synonymy under Gesneria albiflora (Decne.) Kuntze (accepted name G. pedunculosa; Skog, 2012) by Urban (1911). None of Bello’s varieties are cur-rently recognized as distinct taxa and they have all been merged within G. pedunculosa (Skog, 2012). This species is endemic to Puerto Rico.
Conradia pedunculosa DC. var. lutea Bello in Anales Soc. Esp. Hist. Nat. 10: 288. 1881 (Rubiaceae) – Type: not designated.
= Gesneria pedunculosa (DC.) Fritsch in Engler & Prantl, Nat. Pflanzenfam. 4(3b): 184. 1894 ≡ Conradia pedunculosa DC., Prodr. 7: 525. 1839.See entry for Conradia pedunculosa DC. var. pallida Bello
for a taxonomic discussion of this name.
Atelandra laurina Bello in Anales Soc. Esp. Hist. Nat. 10: 289, fig. 5. 1881 (placed by Bello in Myrsinaceae, accepted name Primulaceae; Atelandra Lindl. belongs to Lami aceae) – Neotype (designated here): PUERTO RICO. Prope Utuado, y sylva primaeva ad los Angeles, 8 Feb 1887, P.E.E. Sintenis 6177 (NY barcode 00843756).
= Meliosma herbertii Rolfe var. martinicensis Krug & Urb. in Urban, Symb. Antill. 1: 516. 1900.Bello (1881) was not certain of the family of this taxon
and provisionally placed it in Atelandra Lindl. (accepted name Hemigenia R.Br., Lamiaceae; Guerin, 2008) under Myrsin-aceae even though Bello’s plant actually is a member of Sabi-aceae (Beusekom, 1971). It is uncertain why Bello selected this genus for placing this species since Atelandra is restricted to Australia. It has been claimed that “Atelandra Bello” is a legiti-mate name (Beusekom, 1971). We do not concur, as Bello did not use two asterisks to indicate that this was a new genus. Furthermore, within Atelandra, Bello described two species
339
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
(A. laurina and A. obtusifolia Bello—see below) without providing a full, independent description for a putative new genus. Therefore, we believe that a strict interpretation of the Code requires that these two species are placed in Atelandra. Still, based on Bello’s original descriptions (they include one plate for A. laurina) these species belong to Meliosma Blume (J. Zúñiga, pers. comm.). Urban (1910) examined one illustra-tion made by Krug, as well as plant material from Stahl’s col-lections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Stahl (1888) accepted A. laurina as a good species but Urban (1910) as well as all other taxonomists work-ing in the region (e.g., Britton & Wilson, 1924c; Liogier, 1994; Acevedo-Rodríguez & Strong, 2012) have assigned Bello’s species to M. herbertii, admittedly a later name (1893) as the transfer of Bello’s species to Meliosma is blocked by an ear-lier M. laurina Blume (1849).
Atelandra obtusifolia Bello in Anales Soc. Esp. Hist. Nat. 10: 289. 1881 ≡ Meliosma obtusifolia (Bello) Krug & Urb. in Ber. Deutsch. Bot. Ges. 13: 216. 1895 – Neotype (desig-nated here): PUERTO RICO. Prope Adjuntas in monte Jajuya [sic], 30 Apr 1886, P.E.E. Sintenis 4229 (LD barcode 1694725; isoneotypes: MO barcode MO-730512, P barcode P04348425).Urban (1910) examined one illustration made by Krug and
plant material from Stahl’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Previously, Krug and Urban (Urban, 1895b) accepted Bello’s specific epi-thet and made a new combination under Meliosma Blume. Stahl (1888) accepted Atelandra obtusifolia. All subsequent taxono-mists working in the region (e.g., Britton & Wilson, 1924c; Liogier, 1994; Acevedo-Rodríguez & Strong, 2012) have fol-lowed Urban’s taxonomic placement. Zúñiga (J. Zúñiga, pers. comm.) also agreed with this interpretation. The species is endemic to Puerto Rico.
Metastelma lineare Bello in Anales Soc. Esp. Hist. Nat. 10: 292. 1881 (Apocynaceae, placed by Bello in Asclepi-adeae) ≡ Cynanchum lineare (Bello) Alain in Bull. Tor-rey Bot. Club 90: 191. 1963 (Apocynaceae) nom. illeg., non N.E.Br. in Bull. Misc. Inform. Kew 1897: 273. 1897 ≡ Cyn anchum belloi P.T.Li in J. S. China Agric. Univ. 14(1): 57. 1993 – Neotype (designated here): PUERTO RICO. Prope Maricao ad montum Alegrillo, 23 Nov 1884, P.E.E. Sintenis 87 (GH barcode 00359289; isoneotypes: BM barcode BM001124485, LD barcode 1694741, MO barcode MO-877014, W Herb. Mus. Palat. Vindob. Aequ. 1900 No. 409).Urban (1910) examined one illustration and plant mate-
rial from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1) and accepted this spe-cies. Stahl (1888) considered the name a synonym of Metastelma filiforme (Griseb.) C.Wright although this species is restricted to Cuba (Krings & Endress, 2012). Subsequent taxonomists (e.g., Britton & Wilson, 1925a; Liogier, 1995; Acevedo-Rodríguez, 2005a; Krings & Endress, 2012; S. Liede, pers. comm.) have also accepted Bello’s name.
Bignonia odorata Bello in Anales Soc. Esp. Hist. Nat. 10: 293. 1881 (Bignoniaceae) – Neotype (designated here): PUERTO RICO. Coamo, circa los Baños, 4 Nov 1885, P.E.E. Sintenis 2116 (NY barcode 01113621).
= Amphilophium lactiflorum (Vahl) L.G.Lohmann in Hokche & al., Nuevo Cat. Fl. Vasc. Venezuela: 271. 2008 (Big-noniaceae) ≡ Bignonia lactiflora Vahl, Symb. Bot. 3: 80. 1794 ≡ Distictis lactiflora (Vahl) DC., Prodr. 9: 191. 1845 (Bignoniaceae) ≡ Macrodiscus lactiflorus (Vahl) Bureau ex K.Schum. in Engler & Prantl, Nat. Pflanzenfam. 4(3b): 216. 1894 (Bignoniaceae).Urban (1910) examined one illustration and plant mate-
rial from Krug’s collections that originally were labeled using Bello’s name (Electr. Suppl.: Table S1). He also received plant material from Stahl’s collections that was originally identified as Bignonia odorata (Electr. Suppl.: Table S1). However, Urban (1911) considered Bello’s species a synonym of Macrodiscus lactiflorus. Earlier, Stahl (1888) accepted B. odorata. Britton & Wilson (1925b) and Liogier (1995) assigned this species to synonymy under Distictis lactiflora. Acevedo-Rodríguez & Strong (2012) adopted Lohmann’s (Hokche & al., 2008) recent combination and considered Bello’s name a synonym of Amphilophium lactiflorum.
Bignonia caryophyllea Bello in Anales Soc. Esp. Hist. Nat. 10: 293. 1881 ≡ Tynanthus caryophylleus (Bello) Alain in Rho-dora 67: 352. 1965 (Bignoniaceae) – Neotype (designated here): PUERTO RICO. Forest near Humacao, 28 Feb 1926, N.L. Britton 8614 (NY barcode 00874601).
= Tynanthus polyanthus (Bureau) Sandwith in Kew Bull. 8: 465. 1954 ≡ Schizopsis polyantha Bureau in Adansonia 5: 378. 1865 (Bignoniaceae).Bello (1881) was uncertain about the proper genus for this
species. Urban (1911) did not examine any plant material or illustrations originally labeled with Bello’s name and listed this name as “B.? caryophyllea”. Stahl (1888) and Britton & Wilson (1925b) accepted Bignonia caryophyllea as a good species although with reservations on its generic placement. Acevedo-Rodríguez & Strong (2012) considered the species a synonym of Tynanthus polyanthus and we concur.
Spathodea portoricensis Bello in Anales Soc. Esp. Hist. Nat. 10: 293. 1881 (Bignoniaceae) – Neotype (designated here): PUERTO RICO. Maricao State Forest, N. side of road 120 near lookout tower, elev. ca. 850 m, 18 Apr 1984, G.R. Proctor 40465 (SJ barcode SJ 00045).
= Tabebuia haemantha (Bertero ex Spreng.) A.DC., Prodr. 9: 214. 1845 (Bignoniaceae) ≡ Bignonia haemantha Bertero ex Spreng., Syst. Veg. 2: 832. 1825 ≡ Tecoma haemantha (Bertero ex Spreng.) Griseb., Cat. Pl. Cub.: 194. 1866 (Bignoniaceae).Urban (1911) examined one illustration made by Krug and
plant material from Stahl’s and Krug’s collections that origi-nally were labeled with Bello’s name (Electr. Suppl.: Table S1). He considered this species to be a synonym of Tecoma haemantha. Other taxonomists working in the region (e.g., Britton & Wilson, 1925b; Liogier, 1995; Acevedo-Rodríguez
340
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
& Strong, 2012) have placed Bello’s name in synonymy under Tabebuia haemantha. In his monograph of Tabebuia Gomes ex DC., Gentry (1992) also placed S. portoricensis in synonymy under Tabebuia haemantha. Tabebuia haemantha is endemic to Puerto Rico.
Crescentia microcarpa Bello in Anales Soc. Esp. Hist. Nat. 10: 294. 1881 (Bignoniaceae) – Neotype (designated here): PUERTO RICO. Lajas [Cabo Rojo?], South of Boquerón, 13 Aug 1950, E.L. Little Jr. 13630 (UPR barcode UPR 00092).
= Crescentia linearifolia Miers in Trans. Linn. Soc. London 26: 172. 1868.Bello was uncertain about the placement of this species and
indicated that it could be a variety of Crescentia cujete L. Urban (1911) examined plant material from Stahl’s collections that origi-nally was labeled with Bello’s name (Electr. Suppl.: Table S1) and considered the name a synonym of C. linearifolia. Stahl (1888) indicated that Bello’s species resembled both C. cuneifolia Gardner and C. acuminata Kunth, now considered synonyms of C. cujete (Gentry, 1980). Other taxonomists working in the Antilles (e.g., Britton & Wilson, 1925b; Liogier, 1995; Acevedo-Rodríguez & Strong, 2012) have followed Urban’s taxonomic interpretation for this species. Here we follow Gentry (1980) who assigned C. microcarpa to synonymy under C. linearifolia.
Ipomoea caerulea Bello in Anales Soc. Esp. Hist. Nat. 10: 296. 1881 (Convolvulaceae), nom. illeg., non J.Koenig. ex Roxb. in Bot. Reg. 4: t. 276. 1818.
= Convolvulus cuspidatus Willd. ex Spreng., Syst. Veg. 1: 697. 1824.Urban (1910) examined one illustration made by Krug that
originally was labeled with Bello’s name (Electr. Suppl.: Table S1) and considered this species a synonym of Ipomoea meyeri. Stahl (1888) was not familiar with the species, and accepted Bello’s name (as “caerula”) although he did indicate that the species was morphologically similar to Jacquemontia tam-nifolia (L.) Griseb. (Convolvulaceae; as “tamifolia”). Britton & Wilson (1925a) and Acevedo-Rodríguez & Strong (2012) adopted Urban’s opinion. Austin (D. Austin, pers. comm.) and G. Staples (pers. comm.) agree that the protologue of I. cae-rulea fits better with I. meyeri than with J. tamnifolia.
Heliophytum portoricense Bello in Anales Soc. Esp. Hist. Nat. 10: 297. 1881 (Boraginaceae) – Neotype (designated here): PUERTO RICO. Salinas de Cabo-Rojo ad litora maris, 5 Feb 1885, P.E.E. Sintenis 585 (NY barcode 01352112).
= Heliotropium parviflorum L., Mant. Pl. 2: 201. 1771 ≡ Helio-phytum parviflorum (L.) DC., Prodr. 9: 553. 1845.Urban (1910) did not examine any plant material or illus-
tration originally assigned to this species, and yet considered Bello’s species to be a synonym of Heliotropium parviflorum.
Stahl (1888) was not familiar with the species either, but assigned it to Heliophytum parviflorum. Britton & Wilson (1925a) suggested that this species is a synonym of Schobera angiosperma. Most taxonomists (Liogier, 1995; Förther, 1998; Feuillet, 2012; F. Luebert, pers. comm.) assign Bello’s species to Heliotropium angiospermum.
Acnistus frutescens Bello in Anales Soc. Esp. Hist. Nat. 10: 299. 1881 (Solanaceae) – Neotype (designated here): PUERTO RICO. On roadside, between Barranquitas and Aibonito, 1 May 1979, A.H. Liogier & al. 28820 (UPR barcode UPR 04120).
= Acnistus arborescens (L.) Schltdl. in Linnaea 7: 67. 1832 ≡ Atropa arborescens L., Cent. Pl. II: 10. 1756 (Solanaceae).Urban (1911) examined one illustration from Krug’s col-
lections that originally was labeled with Bello’s name (Electr. Suppl.: Table S1) and accepted this species; however, he also indicated that it might be a variety of Acnistus arborescens. Stahl (1888) accepted A. frutescens although he was not famil-iar with this taxon. Other taxonomists working in the region (e.g., Britton & Wilson, 1925b; Liogier, 1995; Knapp, 2012) have followed Urban. However, both Britton & Wilson (1925b) and Liogier (1995) were uncertain about the taxonomic status of A. frutescens. In his taxonomic revision of Acnistus Schott, Hunziker (1982) agreed with Urban’s statement, and here we follow this taxonomic assessment.
Adhatoda tetramera Bello in Anales Soc. Esp. Hist. Nat. 10: 301. 1881 (Acanthaceae) – Neotype (designated here): PUERTO RICO. Quebradillas, Barrio Terranova, vicin-ity of El Puente Blanco, former railway bridge, 1.6 km due NE of intersection of highway 2 & road 485, elev. ca. 50 m, 22 Dec 1994, G.R. Proctor 49813 (SJ barcode SJ 00403).
= Justicia borinquensis Britton in Sci. Surv. Porto Rico & Vir-gin Islands 6: 217. 1925.Bello (1881) was not certain of the taxonomic placement
of this species and suggested that it might refer to Beloperone nemorosa (Sw.) Nees (Acanthaceae, accepted name Justicia nemorosa Sw.; Acevedo-Rodríguez & Strong, 2012). Urban (1911) examined one illustration made by Krug that originally was labeled with Bello’s name (Electr. Suppl.: Table S1). Accord-ingly, he placed this species in synonymy under J. sessilis. Stahl (1888) accepted Bello’s name but was not familiar with the spe-cies. Britton & Wilson (1925b) followed Urban’s taxonomic placement and more recently, Liogier (1997) and Acevedo- Rodríguez & Strong (2012) considered Adhatoda tetramera a synonym of Siphonoglossa sessilis. Daniel (T. Daniel, pers. comm.) indicated that the protologue description of A. tetratema fits well with S. sessilis. In addition, this is the only species of Siphonoglossa Oerst. occurring in Puerto Rico.
341
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
NAMES PUBLISHED BY DOMINGO BELLO Y ESPINOSA IN 1883
Rivina viridiflora Bello in Anales Soc. Esp. Hist. Nat. 12: 105. 1883 (Phytolaccaceae) ≡ Rivina laevis L. var. viridiflora (Bello) Voss, Vilm. Blumengärtn., ed. 3, 1: 877. 1895 – Neo-type (designated here): PUERTO RICO. Prope Rincon in fruticetis litoralibus, 30 Nov 1886, P.E.E. Sintenis 5527 (NY barcode 01029581)
= Rivina humilis L. Sp. Pl.: 121. 1753.= Rivina humilis L. var. canescens L., Sp. Pl.: 122. 1753.= Rivina humilis L. var. glabra L., Sp. Pl.: 122. 1753.= Rivina laevis L., Mant. Pl. 1: 41. 1767.= Rivina laevis L. var. pubescens Griseb., Fl. Brit. W. I.: 59.
1859.Urban (1905) did not study any plant material or illustration
originally labeled with Bello’s name, and yet regarded this spe-cies as a synonym of Rivina humilis. This placement has been followed by subsequent taxonomists working in the region (e.g., Britton & Wilson, 1924a; Acevedo-Rodríguez & Strong, 2012).
Rivina humilis L. var. canescens Bello in Anales Soc. Esp. Hist. Nat. 12: 105. 1883, nom. illeg., non L., Sp. Pl.: 122. 1753.
= Rivina humilis L., Sp. Pl.: 121. 1753.Bello (1883) published this variety with a Latin description
and without an authority; therefore, a strict interpretation of the Code made us understand that this was a new taxonomic entity published by Bello. No varieties are recognized within Rivina humilis (Acevedo-Rodríguez & Strong, 2012).
Alternanthera paniculata Bello in Anales Soc. Esp. Hist. Nat. 12: 106. 1883 (Amaranthaceae), nom. illeg., non Kunth in Humboldt & al., Nov. Gen. Sp. 2: 208. 1818.
= Iresine diffusa Humb. & Bonpl. ex Willd., Sp. Pl. 4: 765. 1806 (Amaranthaceae).
1806.= Iresine havanensis Kunth in Humboldt & al., Nov. Gen. Sp.
2: 199. 1818.= Iresine parvifolia Kunth in Humboldt & al., Nov. Gen. Sp.
2: 199. 1818.= Iresine celosia subsp. tomentosa de la Luz in Feddes Repert.
96: 553. 1985.Urban (1905) examined one illustration made by Krug
that referred to this taxon (Electr. Suppl.: Table S1), and con-cluded that Bello’s species was a synonym of Iresine panicu-lata (L.) Kuntze (as “panniculata”). Iresine paniculata has been considered a synonym of I. diffusa by some authors (e.g., Acevedo-Rodríguez & Strong, 2012). However, there are mor-phological differences between these two species (Sánchez del Pino & al., 1999; I. Sánchez del Pino, pers. comm.) and we follow this recommendation. There are two native spe-cies in Puerto Rico: I. diffusa and I. angustifolia Euphrasén. Alternanthera paniculata Bello is a synonym of the former; whereas A. linearis Bello (see below) is a synonym of the latter.
Acevedo-Rodríguez & Strong (2012) did not include Bello’s name in their checklist of seed plants of the West Indies.
Alternanthera linearis Bello in Anales Soc. Esp. Hist. Nat. 12: 107. 1883 – Neotype (designated here): PUERTO RICO. On roadside, Río Mar, Luquillo, 23 Jan 1979, A.H. Liogier & al. 28204 (UPR barcode UPR 01532).
= Iresine angustifolia Euphrasén, Beskr. St. Barthél.: 165. 1795.= Iresine elatior Rich. ex Willd., Sp. Pl. 4: 766. 1806.
Urban (1905) examined one illustration and plant material from Krug’s collections that originally were annotated with Bello’s name (Electr. Suppl.: Table S1). He regarded this species as a synonym of Iresine elatior. Britton & Wilson (1924a) also assigned Bello’s name to I. elatior. Other taxonomists working in the region and with this genus (e.g., Acevedo-Rodríguez & Strong, 2012; I. Sánchez del Pino, pers. comm.) placed Bello’s species in synonymy under I. angustifolia.
Pseudolmedia bucidifolia Bello in Anales Soc. Esp. Hist. Nat. 12: 109. 1883 (“bucidaefolia”) (Moraceae, placed by Bello in Urticaceae) – Neotype (designated here): PUERTO RICO. Seven miles S. of Caguas, moist soil, 24 Jan 1899, A.A. Heller 269 (NY barcode 00819160).
= Buchenavia tetraphylla (Aubl.) R.A.Howard in J. Arnold Arbor. 64: 266. 1983 (Combretaceae) ≡ Cordia tetraphylla Aubl., Hist. Pl. Guiane 1: 224. 1775 (Boraginaceae).
= Bucida capitata Vahl, Eclog. Amer. 1: 50. 1797 (Combret-aceae) ≡ Buchenavia capitata (Vahl) Eichler in Flora 49: 165. 1866.Bello (1883) was not certain about the generic position of
this species. Urban (1910) examined plant material and one illustration from Krug’s collections that originally was labelled with Bello’s name (Electr. Suppl.: Table S1), and regarded this species as a synonym of Buchenavia capitata, an opinion accepted by Britton & Wilson (1925a). Subsequent taxonomists working in the region (e.g., Liogier, 1994; Acevedo-Rodríguez & Strong, 2012) have also accepted Urban’s determination. In his monograph of neotropical Combretaceae, Stace & Alwan (2010) considered P. bucidifolia a synonym of B. tetraphylla.
Sponia stipellata Bello in Anales Soc. Esp. Hist. Nat. 12: 109. 1883 (Ulmaceae, placed by Bello in Urticaceae).This is an undetermined taxon.Urban (1903) included the following statement for this
species: “mihi omnino ignotum est. Vix hujus familiae?” (it is entirely unknown to me. Certainly the family?). Acevedo-Rodríguez & Strong (2012) considered this to be a “doubtful name”.
Xanthosoma sylvestre Bello in Anales Soc. Esp. Hist. Nat. 12: 114. 1883 (Araceae) – Neotype (designated here): PUERTO RICO. Prope Aguas—buenas, in graminosis montanis circa oppidium, 8 Sep 1885, P.E.E. Sintenis 2515 (US barcode 00686307).
= Caladium hortulanum Birdsey, Cult. Aroids: 42. 1951.Urban (1903) examined one illustration made by Krug that
originally was annotated with Bello’s name (Electr. Suppl.: Table S1). He considered this species to be a synonym of Cala-dium bicolor. Britton & Wilson (1923) assigned the name to synonymy under Cyrtospadix bicolor. Acevedo-Rodríguez & Nicolson (2005) and Acevedo-Rodríguez & Strong (2012) followed Urban’s interpretation for this name.
Pleurothallis coriacea Bello in Anales Soc. Esp. Hist. Nat. 12: 116. 1883 (Orchidaceae) – Neotype (designated here): PUERTO RICO. Prope Adjuntas ad arbores vetustas in fruticetis Coff. Arab. in monte Galsa, 9 Apr 1886, P.E.E. Sintenis 4251 (US barcode 00428328; isoneotypes: MO barcode MO-637503, P barcode P00485511).
= Pleurothallis pubescens Lindl. in Compan. Bot. Mag. 2: 355. 1837 ≡ Acianthera pubescens (Lindl.) Pridgeon & M.W.Chase in Lindleyana 16: 245. 2001 (Orchidaceae).Urban (1903) examined one illustration by Krug that origi-
nally was labeled with Bello’s name (Electr. Suppl.: Table S1). Both Urban (1903) and Britton & Wilson (1924a) accepted Bello’s name. However, Ackerman (1995, 2012) regarded Bello’s species as a synonym of Pleurothallis pubescenes.
Epidendrum kraenzlinii Bello in Anales Soc. Esp. Hist. Nat. 12: 116, pl. 1, fig. 1–3. 1883 (Orchidaceae) ≡ Psychilis kraenzlinii (Bello) Sauleda in Phytologia 65: 15. 1988 (Orchidaceae) ≡ Encyclia kraenzlinii (Bello) Ackerman in Lindleyana 2: 123. 1987 (Orchidaceae) – Lectotype (designated by Sauleda in Phytologia 65: 15. 1988): [icon] “Epidendrum Kraenzlinii” in Anales Soc. Esp. Hist. Nat. 12: t. 1, figs. 1–3. 1883.Bello (1883) published one plate for this species (Fig. 3).
Urban (1903) examined one additional illustration and plant material from Krug’s collections that originally were labeled with Bello’s name (Electr. Suppl.: Table S1). Urban (1903) con-sidered this species a synonym of Epidendrum papilionaceum Vahl. (accepted name Psychilis bifida (Aubl.) Sauleda; Sauleda, 1988). Britton & Wilson (1924a) followed Urban’s treatment. Ackerman (1995, 2012) regarded Bello’s species as a synonym of P. kraenzlinii. This species is endemic to Puerto Rico.
Epidendrum krugii Bello in Anales Soc. Esp. Hist. Nat. 12: 117. 1883 ≡ Psychilis krugii (Bello) Sauleda in Phytologia 65: 17. 1988 ≡ Encyclia krugii (Bello) Britton & P.Wilson in Sci. Surv. Porto Rico Virgin Islands 6: 532. 1930 – Lectotype (designated by Sauleda in Phytologia 65: 17. 1988): [icon] “E[pidendrum]. Krugii” in Anales Soc. Esp. Hist. Nat. 12: t. 1, fig. 4–7. 1883.Bello (1883) published one plate for this species (Fig. 3)
that was examined by Urban (1905) who did not have access to any plant material or additional illustrations originally labeled with Bello’s name; he regarded this species as a synonym of Epidendrum papilionaceum (accepted name Psychilis bifida; Sauleda, 1988). Britton & Wilson (1924a) followed Urban’s
treatment. Ackerman (1995, 2012) regarded Bello’s species as a synonym of P. krugii. This species is endemic to Puerto Rico.
Smilax coriacea Bello in Anales Soc. Esp. Hist. Nat. 12: 120. 1883 (Smilacaceae, placed by Bello in Liliaceae), nom. illeg., non Spreng., Syst. Veg. 2: 103. 1825
= Smilax coriacea Spreng., Syst. Veg. 2: 103. 1825.= Smilax ilicifolia Desv. ex Ham., Prodr. Pl. Ind. Occid.: 58.
1825.= Smilax coriacea Spreng. var. ilicifolia (Desv. ex Ham.)
O.E.Schulz in Urban, Symb. Antill. 4: 150. 1903.= Smilax havanensis Jacq. var. portoricensis A.DC. in Candolle
& Candolle, Monogr. Phan. 1: 124. 1878.= Smilax guianensis Vitman var. subarmata O.E.Schulz in
Urban, Symb. Antill. 4: 149. 1903 ≡ Smilax subarmata (O.E.Schulz) O.E.Schulz in Urban, Symb. Antill. 5: 28. 1904.Bello used one asterisk to refer to those names that he
regarded as new taxa; Smilax coriacea Bello is marked as such and therefore, this name falls within this category. In addition, Bello included a Latin description and does not refer to any other name authority. Smilax coriacea Bello is an illegitimate name as it is preceded by Smilax coriacea Spreng. However, taxonomically Smilax coriacea Bello corresponds with Smi-lax coriacea Spreng. Urban (1903) did not examine any plant material or illustrations originally assigned to Bello’s name, but he still concluded that Bello’s name was a synonym of Smilax coriacea. Acevedo-Rodríguez (2005b) followed Urban’s conclusion.
Smilax rotundifolia Bello in Anales Soc. Esp. Hist. Nat. 12: 120. 1883, nom. illeg., non L., Sp. Pl.: 1030. 1753.
= Smilax coriacea Spreng., Syst. Veg. 2: 103. 1825.Urban (1903) did not examine any plant material or illus-
trations originally assigned to Bello’s name, and was uncer-tain about the placement of this taxon, although he thought it was probably a synonym of Smilax guianensis var. subar-mata. Britton & Wilson (1923) were doubtful that Bello was referring to S. rotundifolia L. The latter does not occur in the West Indies. Acevedo-Rodríguez (2005b) and Acevedo-Rodríguez & Strong (2012) regarded Bello’s species as a syn-onym of S. coriacea.
Tillandsia ramosa Bello in Anales Soc. Esp. Hist. Nat. 12: 121. 1883 (Bromeliaceae) – Neotype (designated here): PUERTO RICO. Aguada, in arboribus ad Rio Grande, 18 Dec 1886, P.E.E. Sintenis 5777 (P barcode P00761280).
= Tillandsia utriculata L., Sp. Pl.: 286. 1753.= Tillandsia flexuosa Sw, var. pallida Lindl. in Bot. Reg. 9:
ad t. 749. 1823.= Tillandsia sintenisii Baker in J. Bot. 26: 12. 1888.
As indicated by Cedeño-Maldonado (2005), the name Til-landsia ramosa Sweet is a nomem nudum as it was published without a description. Urban (1903) studied an illustration made by Krug that was labeled with Bello’s name (Electr. Suppl.: Table S1), and regarded the name as a synonym of Tillandsia utricu-lata. Britton & Wilson (1923) and Acevedo-Rodríguez & Strong
343
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
(2012) followed Urban’s taxonomic interpretation for this spe-cies. Based on the description provided by Bello, J. Pinzón (pers. comm.) agreed with this taxonomic placement.
Caraguata macrostachya Bello in Anales Soc. Esp. Hist. Nat. 12: 122. 1883 (Bromeliaceae) ≡ Vriesea macrostachya (Bello) Mez in Candolle & Candolle, Monogr. Phan. 9: 601. 1896 (Bromeliaceae) ≡ Neovriesia macrostachya (Bello) Britton in Sci. Surv. Porto Rico Virgin Islands 5: 142 1923 (Bromeli aceae) – Neotype (designated by Grant in Nor-dic J. Bot. 17: 158. 1997): PUERTO RICO. Orocovis: Barrio Ala de la Piedra, Toro Negro Forest Reserve at the east end of Lago El Guineo at the beginning of the road to the dam, 26 Aug 1993, F.S. Axelrod 6935 (US barcode 00515148; isoneotypes: B barcodes B 10 0243372 & B 10 0243373, MARY barcodes MARY1000342 & MARY1000343, NY barcode 232839, SEL barcodes SEL001272 & SEL001273, UPRRP barcode UPRRP 09754).Urban (1903) considered this taxon a synonym of Vriesea
macrostrachya. Britton & Wilson (1923) also accepted Bello’s epithet but under Neovriesia Britton, and a new combination was proposed. Grant (1997) and taxonomists working in the region (e.g., Cedeño-Maldonado, 2005; Acevedo-Rodríguez & Strong, 2012) followed Urban’s treatment.
Tradescantia portoricensis Bello in Anales Soc. Esp. Hist. Nat. 12: 122. 1883 (Commelinaceae) – Neotype (desig-nated here): PUERTO RICO. Dorado forest, 28 Mar 1966, R.O. Woodbury s.n. (UPR barcode UPR 06057).
= Athyrocarpus pseudomonosperma Kuntze, Revis. Gen. Pl. 3(3): 319. 1898 (Commelinaceae) ≡ Phaeosphaerion pseu-domonosperma (Kuntze) Steyerm. in Fieldiana, Bot. 28(1): 152. 1951 (Commelinaceae).Urban (1903) studied one illustration made by Krug that
originally was assigned to this species. However, he considered Bello’s name to be a synonym of Athyrocarpus persicarii folius (Delile) Hemsl. Britton & Wilson (1923) followed Urban. Athyrocarpus persicariifolius is based on Commelina per-sicariifolia Delile, and, as indicated by Hunt (1994), has traditionally been considered a synonym of C. rufipes var. glabrata (e.g., Proctor, 2005; Acevedo-Rodríguez & Strong, 2012). However, Hunt (1994) also stated that it is not cer-tain if A. persicariifolius should be treated as a synonym of C. rufipes var. glabrata. Based on the species description provided by Bello (1883), D. Hunt (pers. comm.) indicated that Tradescantia portoricensis is a synonym of C. rufipes var. glabrata as reported here.
Rajania flexuosa Bello in Anales Soc. Esp. Hist. Nat. 12: 124. 1883 (Dioscoreaceae), nom. illeg., non Poir. in Lamarck, Encycl. 6: 59. 1804.
= Dioscorea alata L., Sp. Pl.: 1033. 1753 (Dioscoreaceae).Bello (1883) was not certain if this species should be
assigned to Rajania hastata L. Urban (1903) studied plant mate-rial and one illustration from Krug’s collections that originally were labeled using Bello’s name and considered this taxon a synonym of Dioscorea alata. Subsequent taxonomists have fol-lowed this placement (e.g., Britton & Wilson, 1924a; Acevedo-Rodríguez, 2005a, c; Acevedo-Rodríguez & Strong, 2012; Raz, pers. comm).
Paspalum affine Bello in Anales Soc. Esp. Hist. Nat. 12: 125. 1883 (Poaceae) nom. illeg., non Steudel, Syn. Pl. Glumac. 1: 24. 1853.
= Paspalum paniculatum L. var. minus Scribn. in Publ. Field Columbian Mus., Bot. Ser. 2(1): 24. 1800.
= Paspalum hemisphericum Poir. in Lamarck, Encycl. 5: 31. 1804.
= Paspalum strictum Pers., Syn. Pl. 1: 86. 1805.Paspalum affine Steud. does not occur in the West Indies
(Zuloaga & al., 2003; Peterson & al., 2012); therefore, Bello’s plant is not related to this species. Urban (1903) did not examine any plant material or illustration that referred to Bello’s species and he was uncertain as to its placement indicating that it was probably a synonym of P. hemisphericum. Chase (1929), Zulo-aga & al. (2003), and Peterson & al. (2012) considered Bello’s name a synonym of P. paniculatum. Zuloaga (F. Zuloaga, pers. comm.) agreed with this taxonomic placement.
DISCUSSION
In his two works, Bello (1881, 1883) published 88 new taxa (71 in 1881, and 17 in 1883), comprising 3 new unispecific gen-era (Homonoma, Psidiastrum, Stahlia [Fig. 5A]), 63 species, and 22 varieties. Only one of the genera (Stahlia) is currently accepted. Eleven of the species names and one variety are illegitimate (Electr. Suppl.: Table S1). The 52 legitimate species names published by Bello included 4 new combinations (Bom-bycella betulina, B. phoenicia, Heteropterys pubiflora, and Jussiaea plumeriana). Eugenia costata, Eugenia paniculata, Sida purpurea, Sponia stipellata, and Turpinia glandulosa are the only names that we could not assign to any species because of difficulties in interpreting their brief descriptions. Furthermore, two of these species, Eugenia costata and Eugenia paniculata are illegitimate. Three species are still in current use: Magnolia portoricensis (Fig. 5C), Metastelma lineare (Fig. 4B), and Opuntia repens (Fig. 5D). Five of Bello’s names (Atelandra obtusifolia [Fig. 4E], Caraguata macro-stachya [Fig. 4A], Cereus quadricostatus [Fig. 5B], Epiden-drum kraenzlinii [Fig. 4D], E. krugii [Fig. 4C]) are basionyms of names in current use. Six species (Bombycella phoenicia, Borreria alternans, Crescentia microcarpa, Diodia nitens, Geophila cordata, Gossypium janiphae folium) have Spanish descriptions, and the three new genera have Latin descriptions. Eight varietal descriptions are in Spanish (for the genera Cof-fea, Emilia, and Mangifera). None of the varieties are currently
344
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Fig. 5. Species that are widely accepted by taxonomists, either as they were originally published by Bello or as basionyms of other taxa. The unispecific genus Stahlia Bello (Fabaceae) was originally assigned to S. maritima by Bello in 1881, not being aware that the name Caesalpinia monosperma Tul. was previously assigned to this taxon by Tulasne in 1844. A, Stahlia monosperma (Tul.) Urb.; B, Leptocereus quadricostatus (Bello) Britton & Rose; C, Magnolia portoricensis Bello; D, Opuntia repens Bello. — Photo credit: A, M. Gardner; B & C, E. Santiago-Valentín; D, E. Cuevas and E. Medina).
345
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
accepted; in this work we regarded them as mere morphologi-cal variants without taxonomic ranking.
Ten of the species described by Bello (1881, 1883) referred to taxa endemic to Puerto Rico (Electr. Suppl.: Table S1). In addition, Bello’s works provided new descriptions for nine species restricted to the Caribbean Islands, including Puerto Rico (Electr. Suppl.: Table S1). The rest of the names are of non-endemic taxa, including six cultivated ones (Anacardium occidentale, Bignonia caryophyllea, Cajanus luteus, Coffea arabica, Gossypium janiphifolium, Mangifera indica). For the Caribbean island endemic Capparis breynia and for the Puerto Rico endemic Conradia pedunculosa Bello described two varieties each.
It is likely that five of the species published by Bello with descriptions but without authorities (Galactia filiformis (Jacq.) Benth., Fabaceae; Jacquemontia tamnifolia (L.) Griseb., Convol-vulaceae; Miconia pyramidalis (Desr.) DC., Melastomat aceae; Pimenta vulgaris Lind., Myrtaceae, accepted name P. dioica (L.) Merr.; Acevedo-Rodríguez & Strong, 2012; Rumex ber-landieri Meisn., Polygonaceae, accepted name R. chrysocarpus Moris; Mosyakin, 2005) were previously published by other taxonomists and that their authorships were mistakenly omit-ted in the Apuntes. Since these names were not marked with an asterisk by Bello (1881, 1883), we do not regard them as new taxonomic entities.
Landrum (1986) considered Pimenta vulgaris sensu Bello as P. racemosa (Mill.) J.W.Moore. It is worth mentioning that Galactia filiformis and Rumex chrysocarpus do not occur in Puerto Rico. Urban (1905) suggested that G. filiformis sensu Bello corresponds to G. striata var. tomentosa (accepted name G. striata (Jacq.) Urb.; Acevedo-Rodríguez & Strong, 2012). Acevedo-Rodríguez & Lewis (2012) also indicated that Bello’s description for G. filiformis referred to G. striata. The lat-ter is morphologically similar to G. dubia (Acevedo-Rodríguez, 2005a). Bello’s account for this name clearly stated that it had short racemes; therefore, it is likely that this species refers to the West Indian endemic G. dubia and not to G. striata (L. Rico, pers. comm.). Rumex crispus is the only species of the genus found in Puerto Rico (Acevedo-Rodríguez & Strong, 2012) and both Urban (1905) and Acevedo-Rodríguez & Strong (2012) suggested that R. berlandieri sensu Bello refers to this species.
Bello (1881, 1883) also reported five species with neither authorities nor taxonomic descriptions. These names do not seem to represent new taxonomic entities, as they were not coded with asterisks; therefore, we believe that omissions of their actual authorities were typographical errors. These five species are: Leria nutans (L.) DC. (Asteraceae, accepted name Chaptalia nutans (L.) Pol., Asteraceae; Robinson & Funk, 2012), Dorstenia contrajerva L. (Moraceae; “contrayerba” by Bello, 1883), Passiflora foetida L. (Passifloraceae), Poinsettia pulcherrima (Willd. ex Klotzsch) Graham (Euphorbiaceae, accepted name Euphorbia pulcherrima Willd. ex Klotzsch, Euphorbiaceae; Acevedo-Rodríguez & Strong, 2012), and Vol-kameria aculeata L. (Lamiaceae).
Bello published five infraspecific unranked names within the legumes Centrosema virginianum (L.) Benth. (α angustifolium, β ellipticum, γ ovatum) and Lablab vulgaris
Savi (accepted name Lablab purpureus (L.) Sweet; Lewis & Acevedo-Rodríguez, 2012) (α albiflorus, β purpureus) with-out descriptions and authorities. We believe that these five taxa refer to infraspecific names that were originally published by Candolle (1825) with no formal taxonomic rank and with Greek symbols as: C. virginiana α angustifolia, C. virginiana β ellip-tica, C. virginiana γ ovata, L. vulgaris γ albiflorus, and L. vul-garis β purpureus. Interestingly, Lewis & Acevedo-Rodríguez (2012) assigned variety categories (as Centrosema virginianum (L.) Benth. var. angustifolium DC., C. virginianum var. ellip-ticum DC., and C. virginianum var. ovatum DC.) to three of the infraspecific names that were unranked by Candolle (1825).
In the Apuntes, there are names that we consider as nomina nuda. For instance, Bello (1881) assigned the name Piriqueta longifolia to Augustin de Candolle; however, this botanist did not publish the name. The Apuntes do not provide any descrip-tion and there is no asterisk indicating this as a new species. Arbo (1995) considered this name a nomen nudum. Urban (1910) examined one illustration made by Krug originally labeled as P. longifolia and considered this species as P. cistoides (L.) Griseb. Urban (1898, 1911) was aware that Bello had limited bibliographic resources; his main botanical references were the works of Grisebach, Augustin de Candolle’s Prodomus, and Achille Richard’s Essai d’une flore de l’île de Cuba. There-fore, it is not surprising that many of Bello’s names are either homonyms or synonyms of other species (Urban, 1898, 1911).
Howard (1996) reported that Krug was dismayed when he found that Bello published his two floristic studies without hav-ing him as one of the authors, and without making any mention of his drawings or contributions. We have been unable to find additional references to support these claims. Urban (1898, 1902, 1911) provided biographical details for Krug and Bello, and he also discussed Bello’s works in length. However, none of these accounts made any mention of Krug’s disappointment with Bello because these works were not joint publications. Among the many species legitimately published by Bello (1883) there was the orchid Epidendrum krugii (accepted name Psychilis krugii (Bello) Sauleda, see above) that was named after Krug. The protologue of this species stated that it was dedicated to the “señor cónsul D. Leopoldo Krug, a quien profeso una antigua y sincera amistad, y se ocupa activamente de la Historia natural de Puerto-Rico” (Mr. Consul Leopoldo Krug, for whom I have an old and sincere friendship, and who is actively working on the natural history of Puerto Rico). In addition, Bello (1883) referred to Krug as an “excelente amigo” (excellent friend) when he thanked him for his help with the classification of the pteri-dophytes plants included in the second part of his Apuntes. A similar gesture of recognition was made by Bello to Agustín Stahl, to whom he dedicated the genus Stahlia. We know that after returning to the Canary Islands, Bello still maintained contact with some of the naturalists he had met in Puerto Rico. For instance, he facilitated the publication of a short paper on Canary Islands birds by J.C. Gundlach in one of the most influ-ential journals of the archipelago (Gundlach, 1879).
Although the contributions of the Apuntes toward advanc-ing knowledge on the flora of Puerto Rico could be regarded as limited quantitatively, this does not diminish its merit as
346
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Acevedo-Rodríguez, P. 2005a. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 51: 1–483.
Acevedo-Rodríguez, P. 2005b. Smilacaceae. Pp. 78–82 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Monocotyledons and gymno-sperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Acevedo-Rodríguez, P. 2005c. Dioscoreaceae. Pp. 82–92 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Monocotyledons and gymno-sperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Acevedo-Rodríguez, P. 2007. The extant botanical collections and nomenclatural types of Agustín Stahl, Puerto Rican botanist. Caribbean J. Sci. 43: 189–199.
Acevedo-Rodríguez, P. & Lewis, G.P. 2012. Leguminosae (Fabaceae). Pp. 392–471 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Cata-logue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Acevedo-Rodríguez, P. & Nicolson, D.H. 2005. Araceae. Pp. 21–52 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Monocotyledons and gymnosperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Acevedo-Rodríguez, P. & Strong, M.T. (eds.) 2005. Monocotyledons and gymnosperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Acevedo-Rodríguez, P. & Strong, M.T. (eds.) 2012. Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Ackerman, J.D. 1995. An orchid flora of Puerto Rico and the Virgin Islands. Mem. New York Bot. Gard. 73: 1–203.
Ackerman, J.D. 2012. Orchidaceae. Pp. 622–667 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Axelrod, F.S. 2011. A systematic vademecum to the vascular plants of Puerto Rico. Fort Worth: Botanical Research Institute of Texas.
Bello, D. 1879a. Un jardín canario. Revista de Canarias 1: 321–325.Bello, D. 1879b. Un jardín canario (continuación). Revista de Canarias
1: 340–345.Bello, D. 1879c. Un jardín canario (continuación). Revista de Canarias
1: 358–361.Bello, D. 1879d. Un jardín canario (continuación). Revista de Canarias
1: 369–372.
an early attempt to establish taxonomic order for plant diver-sity of that island. The nine still accepted taxa described by Bello belong to a wide assortment of unrelated genera and families including Apocynaceae, Bromeliaceae, Cactaceae, Fabaceae, Magnoliaceae, Orchidaceae, and Sabiaceae. This is a formidable taxonomic accomplishment, considering his lack of bibliographic references and the relative isolation from major botanical institutions. Bello’s relevance in the botanical history of this island has been recognized by the five species named after him: Cynanchum belloi P.T.Li (accepted name Metastelma lineare Bello, see above), Cordia bellonis Urb. (Boraginaceae, accepted name Varronia bellonis (Urb.) Britton, Boraginaceae; Feuillet, 2012), Dipholis bellonis Urb. (Sapotaceae, accepted name Sideroxylon portoricense Urb. subsp. portoricense, Sapo-taceae; Acevedo-Rodríguez & Strong, 2012), Eugenia bellonis Krug & Urb. (Myrtaceae, accepted name Mosiera longipes (O.Berg) Small, Myrtaceae; Axelrod, 2011), and Heteropterys bellonis Urb. (accepted name Heteropterys wydleriana Juss., see above).
ACKNOWLEDGMENTS
We dedicate this paper to Ezequiel Ballesteros and Clara Régulo for their support, friendship, and the enjoyable time we shared with them at La Laguna during our research in the Canary Islands. This project was funded by a CREST Grant (NSF-HRD 0734826) to the University of Puerto Rico (to E. Cuevas, E. Santiago-Valentín, and J. Rauscher). Part of this work was developed during a sabbati-cal leave granted to E. Santiago-Valentín by the University of Puerto Rico Botanical Garden and the Biology Department, UPR-Río Piedras Campus. Javier Francisco-Ortega thanks Fairchild Tropical Botanic Garden (FTBG) and the Louis J. Skinner Foundation for their sup-port of his summer research. Many specialists kindly helped us with the interpretation of the names published by Bello, and we wish to express our gratitude for their time and for sharing their knowl-edge and unpublished results: J. Ackerman (Orchid aceae), D. Atha (Rivina), D. Austin (Ipomoea), F. Barrie (Eugenia), O. Blanchard (Sida), M. Caraballo (Phoradendron), X. Cornejo (Cappar aceae), T. Croat (Xanthosoma), T. Daniel (Acanthaceae), A. Delgado Salinas (Phaseo-lus), M. Endress (Metastelma), R. Faden (Commelina), P. Fryxell (Malv-aceae), P. Griffith (Cactaceae), P. Hoch (Ludwigia), D. Hunt (Trades-cantia), W. Judd (Melastomataceae), S. Liede (Metastelma), F. Luebert (Heliotropium), F. Michelangeli (Melastomataceae), M. Nee (Caya-ponia), J. Pinzón (Tillandsia), L. Raz (Rajania), S. Renner (Caya-ponia), L. Rico (Fabaceae), I. Sánchez del Pino (Iresine), T. Schuster (Rumex), B. Sidoti (Tillandsia), G. Staples (Ipomoea), C. Taylor (Rubi-aceae), T.N. Taylor (Pilosocereus), D.C. Zappi (Pilosocereus), F. Zuloaga (Poaceae), and J. Zúñiga (Meliosma). We thank the following herbaria: B, BM, G, GH, K, LD, MO, NY, P, S, US, and W. A special thanks is extended to R. Vogt, Curator of the herbarium, and S. Bollendorf, Technical Herbarium Assistant, of the Botanischer Garten und Bota-nisches Museum Berlin-Dahlem, for the help and support during the research visit of E. Santiago-Valentín. Kanchi N. Gandhi, G. Moore, and D. Stevenson helped with the nomenclature. Jesús Duque, F. García- Talavera, M. Hernández González, and O. Rodríguez Delgado helped us locate relevant archival documents in Tenerife. Thanks to María del
Carmen Duque, Director of the Museo Municipal de Bellas Artes de Santa Cruz de Tenerife, and Jerónimo Cabrera, Chief Operations Man-ager of Cultural Affairs of the Council of Santa Cruz de Tenerife, for their support in the study of Bello’s manuscripts and illustrations. Silvia Aguiló and M. Colom provided help accessing archival documents in Puerto Rico. Gene Rosenberg provided translations of German texts published by Urban, and David Auerbach proof-read the manuscript. Melissa Abdo and E. Warschefsky provided technical assistance at Fairchild Tropical Botanic Garden. José Sustache and O. Monsegur provided important assistance during field studies. We also wish to express our gratitude to the Montgomery Botanical Center and FTBG for providing lodging and work facilities during the four research vis-its of E. Santiago-Valentín to Miami. This is contribution number 9 from the Herbarium of the University of Puerto Rico Botanical Garden and 290 from the Tropical Biology Program of Florida International University.
LITERATURE CITED
347
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
Bello, D. 1879e. Un jardín canario (continuación). Revista de Canarias 1: 390–393.
Bello, D. 1879f. Un jardín canario (continuación). Revista de Canarias 1: 407–409.
Bello, D. 1880a. Un jardín canario (continuación). Revista de Canarias 2: 1–4.
Bello, D. 1880b. Un jardín canario (continuación). Revista de Canarias 2: 22–24.
Bello, D. 1880c. Un jardín canario (continuación). Revista de Canarias 2: 33–37.
Bello, D. 1880d. Un jardín canario (conclusión). Revista de Canarias 2: 51–54.
Bello, D. 1881. Apuntes para la flora de Puerto-Rico. Primera parte. Anales Soc. Esp. Hist. Nat. 10: 231–304.
Bello, D. 1883. Apuntes para la flora de Puerto-Rico. Segunda parte. Anales Soc. Esp. Hist. Nat. 12: 103–130.
Beusekom, C.F. van 1971. Revision of Meliosma (Sabiaceae), section Lorenzanea excepted, living and fossil, geography and phylogeny. Blumea 19: 355–529.
Blanco, P., Puig-Samper, M.A., Zamudio, G., Valero, M. & Maldo-nado, J.L. 2000. Exploración botánica de las Islas de Barlovento: Cuba y Puerto Rico. Siglo XVIII. La obra de Martín de Sessé y José Estévez. Aranjuez: Ediciones Doce Calles & CSIC.
Borhidi, A. 2006. Rubiáceas de México. Budapest: Akadémiai Kiadó.Britton, N.L. & Rose, J.N. 1913. Studies in Cactaceae – 1. Contr. U.S.
Natl. Herb. 16: 239–242.Britton, N.L. & Wilson, P. 1923. Scientific survey of Porto Rico and
the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (part), vol. 5, part 1. New York: New York Academy of Sciences.
Britton, N.L. & Wilson, P. 1924a. Scientific survey of Porto Rico and the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (continued), vol. 5, part 2. New York: New York Academy of Sciences.
Britton, N.L. & Wilson, P. 1924b. Scientific survey of Porto Rico and the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (continued), vol. 5, part 3. New York: New York Academy of Sciences.
Britton, N.L. & Wilson, P. 1924c. Scientific survey of Porto Rico and the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (continued), vol. 5, part 4. New York: New York Academy of Sciences.
Britton, N.L. & Wilson, P. 1925a. Scientific survey of Porto Rico and the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (continued), vol. 6, part 1. New York: New York Academy of Sciences.
Britton, N.L. & Wilson, P. 1925b. Scientific survey of Porto Rico and the Virgin Islands: Botany of Porto Rico and the Virgin Islands; Descriptive flora—Spermatophyta (continued), vol. 6, part 2. New York: New York Academy of Sciences.
Brummitt, R.K. 2009. Report of the Nomenclature Committee for Vascular Plants: 60. Taxon 58: 280–292.
Candolle, A.P. de 1825. Prodromus systematis naturalis regni vegeta-bilis, vol. 2. Parisiis [Paris]: sumptibus sociorum Treuttel et Würtz. http://dx.doi.org/10.5962/bhl.title.286
Cedeño-Maldonado, J.A. 2005. Bromeliaceae. Pp. 199–231 in: Acevedo- Rodríguez, P. & Strong, M.T. (eds.), Monocotyledons and gymnosperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Chase, A. 1929. The North American species of Paspalum. Contr. U.S. Natl. Herb. 28: 1–310.
Cifre de Loubriel, E. 1995. La formación del pueblo puertorriqueño: La contribución de los isleño-canarios. San Juan, Puerto Rico: Centro de Estudios Avanzados de Puerto Rico y el Caribe.
Croat, T.B. 1976. Flora of Panama, part VI. Family 105. Staphyleaceae. Ann. Missouri Bot. Gard. 63: 393–397.
http://dx.doi.org/10.2307/2395280
Crook, R. & Mottram, R. 2002. Opuntia index. Part 8: R. Bradleya 20: 51–66.
Delgado-Salinas, A., Thulin, M., Pasquet, R., Weeden, N. & Lavin, M. 2011. Vigna (Leguminosae) sensu lato: The names and identi-ties of the American segregate genera. Amer. J. Bot. 98: 1694–1715. http://dx.doi.org/10.3732/ajb.1100069
Dorr, L.J. 2012. Malvaceae. Pp. 496–529 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Exell, A.W. 1944. Catalogue of the vascular plants of S. Tomé (with Principe and Annobon). London: Trustees of the British Museum.
Feuillet, C. 2012. Boraginaceae. Pp. 157–174 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Förther, H. 1998. Die infragenerische Gliederung der Gattung Heliotro-pium L. und ihre Stellung innerhalb der subfam. Heliotropioideae (Schrad.) Arn. (Boraginaceae). Sendtnera 5: 35–241.
Fryxell, P.A. 1979. The natural history of cotton tribe (Malvaceae, tribe Gossypieae). College Station: Texas A&M University Press.
Fryxell, P.A. & Berazaín, F.A. 2007. (1780) Proposal to reject the name Hibiscus brasiliensis (Malvaceae). Taxon 56: 611–612.
García Pulido, D. 2000. San Rafael y San Roque un camposanto con historia (1810–1916). Santa Cruz de Tenerife, Canary Islands: Ayun-tamiento de Santa Cruz.
Gentry, A.H. 1980. Bignoniaceae—Part I (Crescentieae and Tourret-tieae). Fl. Neotrop. Monogr. 25(1): 1–150.
González Escrig, J.L. 2000. Estudios botánico-forestales españoles de la Isla de Puerto Rico durante el siglo XIX. Llull: 23: 315–328.
Govaerts, R., Frodin, D.G. & Radcliffe-Smith, A. 2000. World check-list and bibliography of Euphorbiaceae (and Pandaceae). Kew: The Board of Trustees of the Royal Botanic Gardens, Kew.
Govaerts, R., Sobral, N., Ashton, P., Barrie, F., Holst, B.K., Landrum, L.L., Matsumoto, K., Fernanda Mazine, F., Nic Lughadha, E., Proença, C., Soares-Silva, L.H., Wilson, P.G. & Lucas, E. 2008. World checklist of Myrtaceae. Kew: Kew Pub-lishing, Royal Botanic Gardens.
Govaerts, R., Fernández Casas, F.J., Barker, C., Carter, S., Davies, S., Esser, H.-J., Gilbert, M., Hoffmann, P., Radcliffe-Smith, A., Steinmann, V., Welzen, P. van & Whitmoore, T. 2013a. World checklist of Euphorbiaceae. Facilitated by the Royal Botanic Gar-dens, Kew. http://apps.kew.org/wcsp/
Govaerts, R., Ruhsam, M., Andersson, L., Robbrecht, E., Bridson, D., Davis, A., Schanzer, I. & Sonké, B. 2013b. World checklist of Rubiaceae. Facilitated by the Royal Botanic Gardens, Kew.
http://www.kew.org/wcsp/rubiaceae/Grant, J.R. 1997. The identity of the West Indian endemic Vriesea
macrostachya (Bromeliaceae). Nordic J. Bot. 17: 157–159. http://dx.doi.org/10.1111/j.1756-1051.1997.tb00302.xGuerin, G.R. 2008. A taxonomic revision of Hemigenia section Mal-
Gundlach, J.C. 1879. Aves de las Islas Canarias. Revista de Canarias 1: 385–386.
Gundlach, J.C. 1880. Contribución a la herpetología cubana. Habana: Imprenta de G. Montiel y Compañía.
Hiepko, P. 1996. Collections in the Botanical Museum Berlin-Dahlem (B) of particular interest for the Flora of the Greater Antilles. Fl. Greater Antilles Newslett. 10.
http://www.nybg.org/bsci/fga/Newsletter/FGANno10.htm#CBMBHokche, O., Berry, P.E. & Huber, O. 2008. Nuevo catálogo de la flora
vascular de Venezuela. Caracas: Fundación Instituto Botánico de Venezuela Dr. Tobías Lasser.
Howard, R.A. 1996. Ignatz Urban and the “Symbolae Antillanae”. Fl. Greater Antilles Newslett. 10.
TAXON 64 (2) • April 2015: 323–349Santiago-Valentín & al. • Bello and the Flora de Puerto-Rico
Version of Record
Hunt, D. 2006. The new cactus lexicon. Milbourne Port: DH Books.Hunt, D.R. 1994. Commelina L. N.v: Hierba del pollo. Pp. 171–173 in:
Davidse, G., Sousa S., M. & Chater, A.O. (eds.), Flora Mesoameri-cana, vol. 6, Alismataceae a Cyperaceae. Mexico City: Universi-dad Nacional Autónoma de México.
Hunziker, A.T. 1982. Estudios sobre Solanaceae: 17. Revisión sinóptica de Acnistus. Kurtziana 15: 81–102.
Iltis, H.H. & Cornejo, X. 2010a. Studies in the Capparaceae XXVIII: The Quadrella cynophallophora complex. J. Bot. Res. Inst. Texas 4: 93–115.
Iltis, H.H. & Cornejo, X. 2010b. Studies in the Capparaceae XXIX: Synopsis of Quadrella, a Mesoamerican and West Indian genus. J. Bot. Res. Inst. Texas 4: 117–132.
IPNI 2013. The international plant name index. http://www.ipni.orgIzquierdo, E. 2005. Periodistas canarios. Siglos XVIII al XX, vol. 1.
La Laguna, Canary Islands: Consejería de Educación Cultura y Deportes, Gobierno de Canarias.
Johnston, I.M. 1949. The botany of San Jose Island (Gulf of Panama). Sargentia 8: 1–306.
Knapp, S. 2012. Solanaceae. Pp. 898–913 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Krings, A. & Endress, M. 2012. Apocynaceae. Pp. 36–54 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Kuijt, J. 2003. Monograph of Phoradendron. Syst. Bot. Monogr. 66: 1–643. http://dx.doi.org/10.2307/25011253
Kuijt, J. 2012. Santalaceae. Pp. 867–873 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Lavin, M. 1988. Systematics of Coursetia (Leguminosae-Papilion-oideae). Syst. Bot. Monographs 21: 1–167.
http://dx.doi.org/10.2307/25027701Lewis, G.P & Acevedo-Rodríguez, P. 2012. Leguminosae (Fab-
aceae) [Caesalpinoideae and Faboideae by Gwilym P. Lewis and P. Acevedo- Rodríguez; Mimosoideae by P. Acevedo-Rodríguez]. Pp. 392–471 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Linnaeus, C. 1763. Species plantarum, ed. 2, vol. 2. Holmiae [Stock-holm]: impensis direct. Laurentii Salvii.
Liogier, A.H. 1985. Descriptive flora of Puerto Rico and adjacent islands, vol. 1. Río Piedras, Puerto Rico: Editorial de la Universi-dad de Puerto Rico.
Liogier, A.H. 1988. Descriptive flora of Puerto Rico and adjacent islands, vol. 2. Río Piedras, Puerto Rico: Editorial de la Univer-sidad de Puerto Rico.
Liogier, A.H. 1994. Descriptive flora of Puerto Rico and adjacent islands, vol. 3. Río Piedras, Puerto Rico: Editorial de la Universi-dad de Puerto Rico.
Liogier, A.H. 1995. Descriptive flora of Puerto Rico and adjacent islands, vol. 4. Río Piedras, Puerto Rico: Editorial de la Univer-sidad de Puerto Rico.
Liogier, A.H. 1996. Botany and botanists in Puerto Rico. Pp. 41–53 in: Figueroa-Colón, J.C. (ed.), The scientific survey of Puerto Rico and the Virgin Islands: An eighty year reassessment of the islands natural history. Ann. New York Acad. Sci. 776: 1–273.
Liogier, A.H. 1997. Descriptive flora of Puerto Rico and adjacent islands, vol. 5. Río Piedras, Puerto Rico: Editorial de la Universi-dad de Puerto Rico.
Luckow, M. 1993. Monograph of Desmanthus (Leguminosae-Mimos-oideae). Syst. Bot. Monogr. 38: 1–166.
http://dx.doi.org/10.2307/25027822Magnolia Society International 2007. Classification of the Magnoli-
Michelangeli, F.A. & Bécquer-Granados, E.R. 2012. Melastomat-aceae. Pp. 531–562 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Mosyakin, S.L. 2005. Rumex. Pp. 489–533 in: Flora of North America Editorial Committee (ed.), Flora of North America north of Mexico, vol. 5. Oxford: Oxford University Press.
Padrón Acosta, S. 1968. Retablo canario del siglo XIX. Santa Cruz de Tenerife, Canary Islands: Aula de Cultura de Tenerife.
Peterson, P.M., Soreng, R.J. & Smith, S.F. 2012. Poaceae. Pp. 714–766 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Pinto, F.M. 1884. Don Domingo Bello y Espinosa. La Ilustración Canaria 15 (Febrero 15 de 1884): 121–123.
Proctor, G.R. 2005. Commelinaceae. Pp. 153–167 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Monocotyledons and gym-nosperms of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 52: 1–415.
Raven, P. 1963. The Old World species of Ludwigia (including Jussiaea), with a synopsis of the genus (Onagraceae). Reinwardtia 6: 327–427.
Robinson, H. & Funk, V.A. 2012. Asteraceae. Pp. 89–142 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Rodríguez Delgado, O. 1995. Bello y Espinosa, Domingo. P. 573 in: Macías Hernández, A.M. (ed.), Gran enciclopedia canaria, vol. 3. Santa Cruz de Tenerife, Canary Islands: Ediciones Canarias.
Royal Botanic Gardens, Kew 2013. Kew bibliographic databases. http://kbd.kew.org/kbd/searchpage.do
Rudd, V. 1955. The American species of Aeschynomene. Contr. U.S. Natl. Herb. 32: 1–172.
Salm-Reifferscheid-Dyck, J.F.M.A.H.I. 1834. Hortus dyckensis: ou, Catalogue des plantes cultivées dan les jardins de Dyck. Dussel-dorf: chez Arnz & Comp.
Sánchez del Pino, I., Flores Olvera, H. & Valdés, J. 1999. La familia Amaranthaceae en la flora halófita de gipsófila de México. Anales Inst. Biol. Univ. Nac. Autón. México. Bot. 70: 29–135.
Sandwith, N.Y. 1934. Contributions to the flora of tropical America: XIX. Bull. Misc. Inform. Kew 1934: 124–126.
Santiago-Valentín, E. & González López, R.M. 2002. Nuevos docu-mentos y comentarios acerca de las exploraciones de Juan Cristóbal Gundlach a Puerto Rico. Pp. 103–123 in: Dathe, W. & González López, R.M. (eds.), Johann Christoph Gundlach (1810–1896): Un naturalista en Cuba. Marburg an der Lahn: Basilikien Presse.
Santiago-Valentín, E., Sánchez-Pinto, L. & Francisco-Ortega, J. 2010. From the Canary Islands to the Greater Antilles: Domingo Bello y Espinosa and his contributions to the study of the plants of Puerto Rico. Tropical Gard. 65(3): 44–46.
Santiago-Valentín, E., Sánchez-Pinto, L. & Francisco-Ortega, J. 2014. Domingo Bello y Espinosa: Desde Canarias a las Antillas—Sus estudios de la flora Puerto Rico del siglo XIX. Makaronesia 15: 162–175.
Sauleda, R.P. 1988. A revision of the genus Psychilis Rafinesque (Orchid aceae). Phytologia 65: 1–33.
Skog, L. 2012. Gesneriaceae. Pp. 350–364 in: Acevedo-Rodríguez, P. & Strong, M.T. (eds.), Catalogue of seed plants of the West Indies. Smithsonian Contr. Bot. 98: 1–1192.
Stace, C. & Alwan, A.-R. 2010. Combretaceae: Terminalia and Buchenavia. Fl. Neotrop. Monogr. 107: 1–369.
Stahl, A. 1883. Estudios sobre la flora de Puerto Rico, folleto 1. Puerto Rico: El Asimilista.
Stahl, A. 1884. Estudios para la flora de Puerto Rico, folleto 2. Puerto Rico: González & Co.
349
Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015: 323–349
Version of Record
Stahl, A. 1885. Estudios para la flora de Puerto Rico, folleto 3. Puerto Rico: González & Co.
Stahl, A. 1886. Estudios para la flora de Puerto Rico, folleto 4. Puerto Rico: González & Co.
Stahl, A. 1887. Estudios para la flora de Puerto Rico, folleto 5. Puerto Rico: González & Co.
Stahl, A. 1888. Estudios para la flora de Puerto Rico, folleto 6. Puerto Rico: González & Co.
Steffen, M. 1948. Lexicología canaria (continuación). Nombres popula-res de las hipericáceas en la península ibérica. Revista de Historia 14: 414–457.
Steyermark, J.A. 1972. Rubiaceae. Pp. 227–832 in: Maguirre, B. (ed.), The botany of the Guayana Highland–part IX. Mem. New York Bot. Gard. 23: 1–832.
Taylor, C.M. 2001. Overview of the neotropical genus Notopleura (Rubiaceae: Psychotrieae), with the description of some new spe-cies. Ann. Missouri Bot. Gard. 88: 478–515.
Americanum Magna Hama Pars XXV: The nocturnally flowering Psychotria domingensis–Coussarea hondensis group plus three other Mesoamerican Psychotria species transfer to Palicourea. Novon 20: 481–492. http://dx.doi.org/10.3417/2009124
The Plant List 2013. The plant list. http://www.theplantlist.org/Tulasne, L.R. 1844. Légumineuses arborescentes de l’Amérique du
Sud. Arch. Mus. Hist. Nat. 4: 65–200.Urban, I. 1883. Monographiae der familie der Tureneraceen. Jahrb.
Königl. Bot. Gart. Berlin 2: 1–152.Urban, I. 1895a. Additamenta ad cognitionem florae Indiae Occidenta-
lis. Particula II. Myrtaceae. Bot. Jahrb. Syst. 19: 562–681.Urban, I. 1895b. Über die Sabiaceengattung Meliosma. Ber. Deutsch.
Bot. Ges. 13: 211–222.Urban, I. 1898. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 1, fasc. 1. Berlin: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1899. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 1, fasc. 2. Berlin: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144
Urban, I. 1900. Symbolae antillanae; seu Fundamenta florae Indiae Occidentalis, vol. 2, fasc. 2. Leipzig: Fratres Borntraeger.
http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1902. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 3, fasc. 1. Leipzig: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1903. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 4, fasc. 1. Leipzig: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1905. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 4, fasc. 2. Leipzig: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1910. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 4, fasc. 3. Leipzig: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Urban, I. 1911. Symbolae antillanae; seu Fundamenta florae Indiae
Occidentalis, vol. 4, fasc. 4. Leipzig: Fratres Borntraeger. http://dx.doi.org/10.5962/bhl.title.144Viera y Clavijo, J. de 1882a. Catálogo de plantas de las Islas Canarias
(inédito). Revista de Canarias 4: 33–38.Viera y Clavijo, J. de 1882b. Catálogo de plantas de las Islas Canarias
(inédito) (continuación). Revista de Canarias 4: 75–77.Viera y Clavijo, J. de 1882c. Catálogo de plantas de las Islas Canarias
(inédito) (conclusión). Revista de Canarias 4: 81–83.Wydler, H. 1830. Plantarum quarundam descriptiones. Linnaea 5:
423–430.Wiley, J.W., Frahnert, S., Aguilera-Román, R. & Eckhoff, P. 2014.
Juan Cristóbal Gundlach’s contributions to the knowledge of Puerto Rican birds and his influence on the development of natural history in Puerto Rico. Arch. Nat. Hist. 41: 251–269.
son, P.M., Soreng, P.M. & Judziewicz, E.J. 2003. Catalogue of New World grasses (Poaceae): III. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae. Contr. U.S. Natl. Herb. 46: 1–662.
Vol. 64 (2) • April 2015
International Journal of Taxonomy, Phylogeny and Evolution
Electronic Supplement to
Domingo Bello y Espinosa (11–1) and the new taxa published in his
Apuntes para la flora de Puerto-Rico Eugenio Santiago-Valentín, Lázaro Sánchez-Pinto & Javier Francisco-Ortega
Taxon : –
S1
Electr. Suppl. to: Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015
Table S1. Taxonomic index.Accepted name Name published by Belloa Illustrationsb Plant materialc Original distributionAcnistus arborescens (L.) Schltdl. A. frutescens Bello (1: 299)
[A.H. Liogier 28820, UPR]t. 157 In Furnias:
ex BelloNeotropical
Aeschynomene sensitiva Sw. var. sensitiva
A. fistulosa Bello (1: 259) [P.E.E. Sintenis 94, US]
t. 255, 256 Neotropical
Amphilophium lactiflorum (Vahl) L.G.Lohman
Bignonia odorata Bello (1: 293) [P.E.E. Sintenis 2116, NY]
t. 197 Bello! Stahl! Greater Antilles
Anacardium occidentale L. A. occidentale var. luteum Bello (1: 252)
t. 167 Bello! Stahl! Cultivated
A. occidentale A. occidentale var. rubrum Bello (1: 252)
t. 167 Bello! Stahl! Cultivated
Buchenavia tetraphylla (Aubl.) R.A.Howard
Pseudolmedia bucidaefolia Bello (2: 109) [A.A. Heller 269, NY]
t. 186 Bello! Neotropical
Cajanus cajan (L.) Huth C. luteus Bello (1: 260) [P.E.E. Sintenis 6015, US]
Dioscorea alata L. Rajania flexuosa Bello (2: 124)d t. 97 Bello! Tropical Asia
S2
Electr. Suppl. to: Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015
Table S1. Continued.Accepted name Name published by Belloa Illustrationsb Plant materialc Original distributionEmilia fosbergii Nicolson E. sonchifolia (L.) DC. var. rosea
Bello (1: 286)t. 276 Bello! Tropical Asia
Eugenia L. Psidiastrum Bello (1: 272) [type P. dubium Bello]
t. 227 Widespread
Eugenia sp. E. paniculata Bello (1: 271)d t. 228 Porto Rico: ex Bello
Eugenia sp. E. costata Bello (1: 272)d Prope Anones: ex Bello
E. axillaris (Sw.) Willd. Psidiastrum dubium Bello (1: 272) [L.E. Gregory 208, UPR]
t. 227 Neotropical
E. domingensis O.Berg E. calyculata Bello (1: 271) [E.L. Little Jr. 13596, UPR]
t. 225 Bello! Neotropical
Geophila repens (L.) I.M.Johnst. G. cordata Bello (1: 282)d t. 6 Neotropical
Gesneria pedunculosa (DC.) Fritsch Conradia pedunculosa DC. var. lutea Bello (1: 288)
t. 204 Bello! Puerto Rico
G. pedunculosa C. pedunculosa var. pallida Bello (1: 288)
I. diffusa Humb. & Bonpl. ex Willd. A. paniculata Bello (2: 106)d t. 203 Mexico, SE U.S.A., West Indies, Central America, and South America
Leptocereus quadricostatus (Bello) Britton & Rose (Fig. 5B)
Cereus quadricostatus Bello (1: 276) [P.E.E. Sintenis 4919, B and isoneotypes in BM, G, GH, K, W]
t. 6 Puerto Rico
Leptospron adenanthum (G.Mey.) A.Delgado
Phaseolus cochleatus Bello (1: 262)d t. 277 Bello! Mexico, Central America, West Indies, and South America
S3
Electr. Suppl. to: Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015
Table S1. Continued.Accepted name Name published by Belloa Illustrationsb Plant materialc Original distributionL. adenanthum P. cochleatus var. pallida Bello
(1: 262)Mexico, Central America, West Indies, and South America
L. adenanthum P. cochleatus var. violacea Bello (1: 262)
t. 277 Bello! Mexico, Central America, West Indies, and South America
Ludwigia erecta (L.) H.Hara Jussiaea plumeriana (DC.) Bello (1: 267)
Neotropical
Magnolia portoricensis Bello (Fig. 5C) M. portoricensis (1: 233) [P.E.E. Sintenis 4581, B and isoneotype in LD]
t. 254 Stahl! Puerto Rico
Malvaceae sp. Sida purpurea Bello (1: 239) t. 49 fig. 1 In Aguadilla: ex Bello
Mangifera indica L. M. indica var. armeniaca Bello (1: 253)
t. 166 Bello! Stahl! Cultivated
M. indica M. indica var. intermedia Bello (1: 253)
t. 166 Bello! Stahl! Cultivated
M. indica M. indica var. leiosperma Bello (1: 253)
t. 166 Bello! Stahl! Cultivated
M. indica M. indica var. macrocarpa Bello (1: 253)
t. 166 Bello! Stahl! Cultivated
M. indica M. indica var. viridis Bello (1: 253) t. 166 Bello! Stahl! Cultivated
Paspalum paniculatum L. Paspalum affine Bello (2: 125)d Neotropical
Phoradendron quadrangulare (Kunth) Griseb.
Viscum randiae Bello (1: 278) [P.E.E. Sintenis 885, US]
t. 239 Neotropical
S4
Electr. Suppl. to: Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015
Table S1. Continued.Accepted name Name published by Belloa Illustrationsb Plant materialc Original distributionPilosocereus royenii (L.) Byles & Rowley
Stahlia Bello Stahlia Bello (1: 255) [type S. maritima Bello]
t. 340 Greater Antilles
S. monosperma (Tul.) Urban (Fig. 5A) S. maritima Bello (1: 255) [P.E.E. Sintenis 3876, NY and neotypes in BM, G, GH, NY, P, W]
t. 340 Greater Antilles
Tabebuia haemantha (Bertero ex Spreng.) DC.
Spathodea portoricensis Bello (1: 293) [G.R. Proctor 40465, SJ]
t. 200 Bello! Stahl! Puerto Rico
Tetrapterys inaequalis Cav. T. paniculata Bello (1: 245) [A. Stahl 338, S]
Greater and Lesser Antilles
Tillandsia utriculata L. T. ramosa Bello (2: 121) [P.E.E. Sintenis 5777, P]
t. 116 Neotropical
Tynanthus polyanthus (Bureau ex Baill.) Sandwith
Bignonia caryophyllea Bello (1: 293) [N.L. Britton 8614, NY]
Cultivated
S5
Electr. Suppl. to: Santiago-Valentín & al. • Bello and the Flora de Puerto-RicoTAXON 64 (2) • April 2015
Table S1. Continued.Accepted name Name published by Belloa Illustrationsb Plant materialc Original distributionUndetermined taxon Sponia stipellata Bello (2: 109)
Undetermined taxon Turpinia glandulosa Bello (1: 250)
Vigna trichocarpa (C.Wright) A.Delgado
Phaseolus lanceolatus Bello (1: 262) [J.A. Stevenson 2097, NY]
t. 280 Bello! Neotropical
Vriesea macrostachya (Bello) Mez (Fig. 4A)
Caraguata macrostachya Bello (2: 122) [F.S. Axelrod 6935, US and isoneotypes in B, MARY, NY, SEL, UPRRP]
Greater Antilles
a Information inside parentheses is coded as follow: (Volume number of Bello’s work: page number). Types are indicated inside brackets. Notice that they refer to neotypes except when specified.
b Illustrations made by Krug that were examined by Urban.c Plant material belonging to Krug’s (including his illustrations) or Stahl’s collections that was examined by Urban. Locality of taxa reported by
Urban that was based on information from the Apuntes.d Nomen illegitimum.e Based on the short description provided by Bello it is not certain if this variety can be assigned to Cynophalla flexuosa or to Quadrella
