Research Article |
Corresponding author: Andreas Berger ( andi.berger@univie.ac.at ) Academic editor: Peter de Lange
© 2017 Andreas Berger.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Berger A (2017) Two new combinations, lectotypifications and a new name for Costa Rican Palicourea s.l. PhytoKeys 80: 53-63. https://doi.org/10.3897/phytokeys.80.13330
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Species of the complex and diverse genera Psychotria and Palicourea are common but little-known elements in many tropical forest ecosystems. DNA-phylogenetic studies and a re-evaluation of morphological characters have recently shown that species of Psychotria subg. Heteropsychotria are nested within Palicourea s.l., which was traditionally separated by exhibiting a bird-pollinated (vs. insect-pollinated) pollination syndrome. In order to render both genera monophyletic groups, species of subg. Heteropsychotria need to be transferred to Palicourea s.l. For Central American species, most of the necessary combinations have already been made. In the course of ongoing research on the phytochemical characterization of species and clades of Costa Rican Palicourea s.l., the nomenclature of Mesoamerican species was revised. As a result, two new combinations and a new name are proposed here: Palicourea horquetensis (Dwyer & Hayden) A. C. Berger & C. M. Taylor is based on Rudgea horquetensis Dwyer & Hayden, Palicourea tonduzii (K. Krause) A. C. Berger is based on Cephaelis tonduzii K. Kraus and Palicourea longiinvolucrata A. C. Berger replaces Psychotria hispidula Standl. In addition, two lectotypes are designated.
Palicourea , Psychotria , Rubiaceae , Mesoamerica, taxonomy
Species of the complex and diverse genera Psychotria L. (1759: 929) and Palicourea Aubl. (1775: 172–175) are prominent but little-known elements in tropical forest ecosystems. Both have long been considered closely related, and Palicourea was differentiated from Psychotria by characters associated with hummingbird rather than insect pollination. Species of Palicourea are typically found in the understory of rainforests and are especially frequent in high elevation habitats where Psychotria and other related genera are less speciose (
Both genera were traditionally classified in the tribe Psychotrieae. Recently, however, DNA-phylogenetic studies and a re-evaluation of morphological characters have shown that species of Psychotria subg. Heteropsychotria Steyerm. (1972: 484) are more closely related to Palicourea. Consequently, views shifted towards a narrower concept of Psychotria and Psychotrieae which peaked in the ongoing segregation of hundreds of species and the establishment of the sister tribe Palicoureeae (
Species of Psychotria subg. Heteropsychotria and Palicourea cannot be distinguished by vegetative or fruit characters. In addition, both groups show similar accumulation of alkaloids (e.g.,
By contrast, species of Palicourea are hummingbird-pollinated, frequently have long-pedunculate and open inflorescences, colored inflorescence axes, large and long pedicellate flowers and vividly colored corollas with well-developed tubes. Corollas have a gibbous, nectar-accumulating swelling at their base that is protected by an internal ring of hairs. In many plant groups, bird-pollinated flowers have repeatedly evolved in groups of bee-pollinated ancestors and are not phylogenetically informative at the generic level (e.g.,
In order to render both Palicourea and Psychotria monophyletic groups, most species of Psychotria subg. Heteropsychotria have to be transferred to Palicourea s.l. The combined group includes more than 800 species, is variable in flower characters, but is supported by vegetative and fruit characters as well as by DNA phylogenetic data (
In the course of ongoing research on the phytochemical characterization of species and clades of Costa Rican Palicourea s.l., the nomenclature of Mesoamerican species was revised and the need for a new name and two new combinations became apparent. These are proposed here. In addition, a complete synonymy, an enumeration of type specimens and two lectotypifications are provided for these three species.
The present work is based on an extensive study of herbarium specimens, digital images and relevant literature including regional (e.g., Manual de Plantas de Costa Rica,
For all names, protologues were checked to verify or revise author and page citations, information on collectors and localities. Subsequently, the type category applying to each name was assessed in accordance with the ICN (Melbourne code,
For the three nomenclatural novelties, both possible author abbreviations of my name are preoccupied: Berger stands for Ernst Friedrich Berger (1814–1853) and A. Berger stands for Alwin Berger (1871–1931). Therefore, I have adopted the abbreviation A. C. Berger that includes my second forename Christoph, which I have never used before in my publications.
Basionym. Rudgea horquetensis Dwyer & Hayden, Ann. Missouri Bot. Gard. 54(2): 145–146, 1967.—Type: PANAMA. Chiriquí: Distr. Boquete, Cerro Horqueta, ca. 1980 m, 26 Jul 1940, C. von Hagen & W. von Hagen 2156 (holotype: NY barcode 133202!).
=
Coussarea
nebulosa
Dwyer, Ann. Missouri Bot. Gard. 67(1): 131, 1980a. ≡ Psychotria nebulosa (Dwyer) C. M. Taylor, Novon 5(2): 205, 1995, nom. illeg. hom., non Psychotria nebulosa K. Krause, Bot. Jahrb. Syst. 57(1): 46–47, 1920. ≡ Palicourea nebulosa (Dwyer) C. M. Taylor, Novon 20(4): 488, 2010.—Type: PANAMA. Chiriquí: Monte Rey near Boquete, ca. 1170 m, 20 Jul 1971, T. B. Croat 15868 (lectotype, designated by
=
Rudgea
chiriquiensis
Dwyer, Ann. Missouri Bot. Gard. 67(2): 476, 1980b. ≡ Coussarea chiriquiensis (Dwyer) C. M. Taylor, Fieldiana, Bot., n.s. 33: 113, 1993.—Type: PANAMA. Chiriquí: Cerro Colorado, along road above San Félix, 29 km above bridge over Río San Félix, 7.9 km above turnoff to Escopeta, 1500 m, 14 Jul 1976, T. B. Croat 37071 (lectotype, designated by
Rudgea horquetensis Dwyer & Hayden was accepted as a species of Rudgea by most authors including
The protologue of Coussarea nebulosa states that the holotype is located at MO. At the time of publication two sheets of the type collection have been accessioned at MO, making the holotype designation ambiguous. In 1975, Dwyer annotated both sheets as Coussarea nebulosa, but did not specify what sheet he intended to be the holotype. Hence, both specimens represent syntypes (ICN, Art. 40.2 & Note 1; see also
A similar case of indirect lectotypification is found in Rudgea chiriquiensis. The species was described with reference to two collections at that time housed at MO, one of which was later distributed to PMA. Likewise, lectotypification (of the MO sheet) was achieved by
Palicourea horquetensis is only known from few sites in Costa Rica and Panama.
Basionym: Cephaelis tonduzii K. Krause, Bot. Jahrb. Syst. 54(3, Beibl. 119): 45–46, 1916. Non Psychotria tonduzii Standl., J. Wash. Acad. Sci. 15(13): 287, 1925b.—Type: COSTA RICA. Cartago: Tuis, 650 m, Nov 1897, A. Tonduz 11461 (lectotype, designated here: fragm. F barcode V0068631F! ex B; syntype, or possibly holotype: B † [photo: F neg. BN-773!]).
= Cephaelis discolor Pol., Linnaea 41(5–6): 572–573, 1877. ≡ Uragoga angosturensis Kuntze, Revis. Gen. Pl. 2: 954, 1891, nom. nov., non Uragoga discolor (Benth.) Kuntze, Revis. Gen. Pl. 2: 960, 1891. Non Psychotria discolor (Griseb.) Rolfe, Bull. Misc. Inform. Kew 1893: 258, 1893, nec Palicourea discolor K. Krause, Bot. Jahrb. Syst. 54(3, Beibl. 119): 40–41, 1916.—Type: COSTA RICA. Cartago: Angostura, Nov 1875, H. Polakowsky 384 (lectotype, designated here: fragm. F barcode V0068625F! ex B; syntype, or possibly holotype: B † [photo: F neg. BN-722!]).
= Evea guapilensis Standl., J. Wash. Acad. Sci. 15(5): 104–105, 1925a. ≡ Cephaelis guapilensis (Standl.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4(8): 295, 1929. ≡ Psychotria guapilensis (Standl.) Hammel, Selbyana 12: 139, 1991. ≡ Palicourea guapilensis (Standl.) Borhidi, Acta Bot. Hung. 59(1–2): 17, 2017a.—Type: COSTA RICA. Limón: Vicinity of Guápiles, 300–500 m, 12–13 Mar 1924, P. C. Standley 37025 (holotype: US barcode 00129829!).
= Evea nana Standl., J. Wash. Acad. Sci. 15(5): 105, 1925a. ≡ Cephaelis nana (Standl.) Standl., J. Wash. Acad. Sci. 17(7): 171, 1927. Type: PANAMA. Colón: Hills N of Frijoles, 19 Dec 1923, P. C. Standley 27550 (holotype: US barcode 1153871!).
= Cephaelis nicaraguensis Standl., Trop. Woods 16: 46, 1928.—Type: NICARAGUA. Atlántico Norte: Puerto Cabezas [Bragman’s Bluff], bank of Kukalaya River, 60 m, 08 Dec 1927, F. C. Englesing 58 (holotype: F barcode V0068627F!; isotype: fragm. G barcode G00300772! ex F).
The earliest published name for the species is Cephaelis discolor but combinations under Psychotria and Palicourea are preoccupied by Psychotria discolor (Griseb.) Rolfe and Palicourea discolor K. Krause. The next available name Cephaelis tonduzii K. Krause cannot be used in Psychotria because of the earlier Psychotria tonduzii Standl. Hence, the next name Evea guapilensis Standl. was adopted and the species became known as Psychotria guapilensis (Standl.) Hammel. Recently,
However, he overlooked that the earlier published name Cephaelis tonduzii K. Krause is still available under Palicourea and ought to have been adopted. The corresponding new combination is provided here. It is fortunate that the correct name for this showy species honors Swiss botanist Adolphe Tonduz (1862–1921), a long-term employee at the Museo Nacional de Costa Rica (
Cephaelis tonduzii was described with reference to an entire gathering by Tonduz. Krause did not cite a particular herbarium specimen and all possible duplicates are therefore syntypes (ICN, Art. 40.2 & Note 1; see also
The fragment originates from the type at B, as shown by information given on the label. It consists of a capsule with a leaf and part of an inflorescence, which is mounted together with a photograph (“Berlin negative”) of the original B specimen. The removed leaf is clearly recognizable on the photograph providing a definite link between both. Although not the best choice, the fragment at F appears to be the only extant original material and is here designated as lectotype.
A similar situation is found in Cephaelis discolor, which was based on a gathering of H. Polakowsky. Likewise, the only known original material is a fragment at F that originated from B. It consists of a leaf, some bracts and fruits in a capsule, which is mounted together with a photography of the B specimen, likewise destroyed. The fragment at F is here designated as lectotype.
Palicourea tonduzii is known from Nicaragua, Costa Rica, Panama and Ecuador (
Basionym/replaced synonym: Psychotria hispidula Standl. ex Steyerm., Acta Biol. Venez. 4(1): 97–98, 1964. ≡ Palicourea hispidula (Standl. ex Steyerm.) Borhidi, Acta Bot. Hung. 59(1–2):17, 2017a, nom. illeg. hom., non Palicourea hispidula Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 11(5): 227, 1936.—Type: COLOMBIA. Valle del Cauca: Río Calima, La Trojita, 5–50 m, 19 Feb–10 Mar 1944, J. Cuatrecasas 16359 (holotype: F barcode V0070450F!; isotypes: BC 623883!, U barcode 0006197!, US barcode 00138790!, VEN n.v.).
–
Psychotria
involucrata
sensu
– Psychotria hoffmannseggiana sensu Burger & Taylor (1993, p.p.), non (Roem. & Schult.) Müll. Arg.
Palicourea longiinvolucrata and related species, note diagnostic differences in bract arrangement and length. Palicourea longiinvolucrata, inflorescence (ABerger 1418) and infructescence (BBerger 1633); Palicourea hoffmannseggiana, infructescence (C unvouchered); Palicourea gracilenta, infructescence (DBerger 2055); Palicourea winkleri, infructescence (EBerger 1737). All photos by the author.
Palicourea longiinvolucrata is known from Belize to Bolivia, from Venezuela and from Brazil (
Special thanks are due to Charlotte M. Taylor (MO) who generously helped answering many questions in the course of the project. She pointed me to the earlier name Rudgea horquetensis and also curates the Rubiaceae in the invaluable TROPICOS database (http://www.tropicos.org). Further thanks goes to J. McNeill (E) for some help with nomenclature. Also, many thanks go to the curators and staff of the herbaria that helped me by checking their collections for type specimens and digitized them on request. In addition, I wish to thank the Andrew W. Mellon Foundation which funded the digitization of most type specimens currently available online. I wish to thank Rhys Gardner for reviewing the paper and providing some valuable suggestions on how to improve the manuscript. Fieldwork was supported by a grant by the Society for the Promotion of the La Gamba Field Station. Publication of this article was supported by the Open Access Publishing Fund of the University of Vienna.