VOL. 23, SUPPLEMENT December 2023 I N T E R N A T I O N A L J O U R N A L O N O R C H I D O L O G Y ISSN 1409-3871 THE LANKESTER CATALOGUE OF COSTA RICAN ORCHIDACEAE Franco PuPulin, Diego Bogarín and aDam P. Karremans INTERNATIONAL JOURNAL ON ORCHIDOLOGY LANKESTERIANA Editor-in-Chief (Director) Diego Bogarín Universidad de Costa Rica, Costa Rica diego.bogarin@ucr.ac.cr Associate Editors James D. ackerman University of Puerto Rico, U.S.A. germán carnevali Centro de Investigación Cientifica de Yucatán, Mexico PhilliP criBB Royal Botanic Gardens, Kew, U.K. carlos F. Fighetti The American Orchid Society, U.S.A. günter gerlach Botanischer Garten München-Nymphenburg, Germany heiko hentrich Deutsche Homöopathie-Union Karlsruhe, Germany Julián monge-náJera Universidad de Costa Rica, Costa Rica DaviD l. roBerts University of Kent, U.K. anDré schuiteman Royal Botanic Gardens, Kew, U.K. Jorge Warner Universidad de Costa Rica, Costa Rica Franco Bruno Universitá La Sapienza, Roma, Italia mark W. chase Royal Botanic Gardens, Kew. U.K. roBert l. Dressler† Universidad de Costa Rica, Costa Rica lauren garDiner Royal Botanic Gardens, Kew, U.K. eric hágsater Herbario AMO, Mexico Wesley e. higgins The American Orchid Society, U.S.A. alec m. PriDgeon Royal Botanic Gardens, Kew, U.K. gustavo a. romero Harvard University Herbaria, U.S.A. PhiliP seaton lucn/ssc Orchid Specialist Group, U.K. W. mark Whitten† Florida Museum of Natural History, U.S.A. Comité Científico / Scientific Committee Diego Bogarín Universidad de Costa Rica, Costa Rica aDam P. karremans Universidad de Costa Rica, Costa Rica gaBriela Jones román Universidad Estatal a Distancia, Costa Rica melania FernánDez Universidad de Costa Rica, Costa Rica víctor m. Jiménez Universidad de Costa Rica, Costa Rica Jorge Warner Universidad de Costa Rica, Costa Rica Consejo Editorial / Editorial Committee norris h. Williams† Florida Museum of Natural History, U.S.A melissa Díaz-morales Universidad de Costa Rica, Costa Rica melissa.diaz_m@ucr.ac.cr Franco PuPulin Universidad de Costa Rica, Costa Rica franco.pupulin@ucr.ac.cr Technical Editor noelia BelFort oconitrillo Universidad de Costa Rica, Costa Rica noelia.belfort@ucr.ac.cr lankesteriana international Journal on orchiDology Copyright © 2023 Lankester Botanical Garden, University of Costa Rica Effective publication date ISSN 2215-2067: December 29, 2023 Effective publication date ISSN 1409-3871 (printed): December 29, 2023 Layout: Jardín Botánico Lankester. Cover: Benzingia reichenbachiana (Schltr.) Dressler. Photograph by Franco Pupulin. Printer: MasterLitho. Printed copies: 200 Printed in Costa Rica / Impreso en Costa Rica R Lankesteriana / International Journal on Orchidology No. 1 (2001)-- . -- San José, Costa Rica: Editorial Universidad de Costa Rica, 2001-- v. ISSN-1409-3871 1. Botánica - Publicaciones periódicas, 2. Publicaciones periódicas costarricenses ISSN 1409-3871 VOL. 23, SUPPLEMENT DECEMBER 2023 I N T E R N A T I O N A L J O U R N A L O N O R C H I D O L O G Y The Lankester Catalogue of Costa Rican Orchidaceae The checklist The catalogue Phylogenetic arrangement and diversity Index of accepted names and synonyms Franco PuPulin, Diego Bogarín and aDam P. Karremans 1 18 40 176 180 Franco PuPulin1,2,3,6 Diego Bogarín1,4,5,6 & aDam P. Karremans1 aBstract. This study presents an updated comprehensive catalogue of the Costa Rican orchid flora. We have documented 1684 species, four subspecies, 14 natural hybrids, and seven forms across 201 genera and two nothogenera, × Bensteinia and × Cochlezella. All Orchidaceae subfamilies are represented except for Apostasioideae. The subtribe Pleurothallidinae accounts for about 35% of all the species, followed by Laeliinae (20%) and Oncidiinae (+12%). The most species-rich genera in Costa Rica are Epidendrum (245 spp.), Lepanthes (163 spp.), Stelis (109 spp.), and Pleurothallis (63 spp.). About 30% of all the species are endemic, primarily concentrated in Lepanthes (109 spp. = 67%), Epidendrum (76 spp. = 31%), Stelis (40 spp. = 37%), Telipogon (35 spp. = 88%), and Pleurothallis (31 spp. = 46%). All species are vouchered by specimens housed in major public herbaria, including CR, JBL, and USJ in Costa Rica, as well as AMES, K, SEL, US, and W, on the international scale, among others. The catalogue includes 5300 orchid names at the specific and subspecific ranks, nearly 70% of which are treated as synonyms. The new catalog presents an increase of 324 species compared to its predecessor, published twenty years ago, a net increment of almost 24%. The rise in the recorded richness of the Costa Rican orchid flora is mainly attributed to the discovery of species new to science and a lesser extent, taxa newly recorded for the country as the result of the systematic explorations carried out in Costa Rica during the present century. A phylogenetic arrangement of the genera recorded in Costa Rica, from the rank of subfamily to that of subtribe, is presented, including a synopsis of the diversity recorded for each genus. An enumerated checklist of the accepted species and a comprehensive index of accepted names and synonyms are also provided. The index is cross-referenced with both the checklist and the main catalogue. New combinations for Andreettaea fimbriata and Camaridium muscoides are proposed. Photographs of representative species from the majority of genera, based on Costa Rican vouchers, are included. We expect that this catalogue will inspire orchidological and botanical research in Costa Rica and other Neotropical regions, leading to additional discoveries in the upcoming years. resumen. Este estudio presenta un catálogo completo actualizado de la flora de orquídeas de Costa Rica. Hemos documentado 1684 especies, cuatro subespecies, 14 híbridos naturales y siete formas en 201 géneros y dos nothogéneros, × Bensteinia y × Cochlezella. Todas las subfamilias de Orchidaceae están representadas, excepto Apostasioideae. La subtribu Pleurothallidinae representa aproximadamente el 35% de todas las especies, seguida de Laeliinae (20%) y Oncidiinae (+12%). Los géneros más ricos en especies en Costa Rica son Epidendrum (245 spp.), Lepanthes (163 spp.), Stelis (109 spp.) y Pleurothallis (63 spp.). Aproximadamente el 30% de todas las especies son endémicas, principalmente concentradas en Lepanthes (109 spp. = 67%), Epidendrum (76 spp. = 31%), Stelis (40 spp. = 37%), Telipogon (35 spp. = 88%), y Pleurothallis (31 spp. = 46%). Todas las especies están respaldadas por ejemplares almacenados en herbarios públicos importantes, incluyendo CR, JBL y USJ en Costa Rica, así como AMES, K, SEL, US y W, a nivel internacional, entre otros. El catálogo incluye 5300 nombres de orquídeas en los rangos específicos y subespecíficos, casi el 70% de los cuales se tratan como sinónimos. El nuevo catálogo presenta un aumento de 324 especies en comparación con su predecesor, publicado hace veinte años, un aumento neto de casi 24%. El aumento en la riqueza registrada de la flora de orquídeas THE LANKESTER CATALOGUE OF COSTA RICAN ORCHIDACEAE 1 Centro de Investigación Jardín Botánico Lankester, Universidad de Costa Rica, P.O. Box 302-7050, Cartago, Costa Rica. 2 Harvard University Herbaria, Cambridge, Massachusetts 02138, U.S.A. 3 The Marie Selby Botanical Gardens, Sarasota, Florida 34236, U.S.A. 4 Naturalis Biodiversity Center, Leiden, 2333 CR, the Netherlands. 5 Herbario UCH, Universidad Autónoma de Chiriquí, 0427, David, Chiriquí, Panamá. 6 Authors for correspondence: franco.pupulin@ucr.ac.cr; diego.bogarin@ucr.ac.cr ORCID of the Authors: FP , DB , AK Received 17 October 2023; accepted for publication 27 December 2023. First published online: 30 December 2023. Licensed under a Creative Commons Attribution-NonCommercial-No Derivs 3.0 Costa Rica License. LANKESTERIANA 23(Supplement): 1–254. 2023. doi: http://dx.doi.org/10.15517/lank.v23iSupplement.58145 https://orcid.org/0000-0001-6780-8395 https://orcid.org/0000-0002-8408-8841 https://orcid.org/0000-0001-5987-7710 http://dx.doi.org/10.15517/lank.v23iSupplement.58145 LANKESTERIANA2 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. Introduction. An up-to-date version of the catalogue of Costa Rican orchids has been long overdue and we felt Lankester Botanical Garden’s 50th anniversary provided the perfect occasion for its publication. It is undoubtedly an opportunity offered by an important date in the history of our research center, just like publishing the first “working list” in 2002 marked the official beginning of Lankester Botanical Garden’s career as a research center of the University of Costa Rica (Pupulin 2002). At the time we believed we could update the catalogue, with its imperfections, every decade. For various reasons, however, it was not possible to do so, and more than two decades have elapsed since. Today we can finally announce the release of a new catalog. An update of the names included in the orchid flora of Costa Rica was necessary, not only to compile the taxonomic concepts that we consider most useful concerning the genera recorded in the country’s flora into but also to include a large number of species new to science or new to the country, which have accumulated over the last twenty years; often shifting from one genus to another on numerous occasions during these two decades. The last 20 or so years have witnessed a tremendous surge in orchid phylogenetic studies, which in some cases have confirmed past hypotheses while at the same time offering new interpretations of the relationships among different groups of these plants. Inevitably, these extensive investigations, spanning nearly all the groups of orchids, have generated new hypotheses regarding the boundaries of traditionally known genera and, in numerous cases, resulting in new genera, albeit not always universally accepted by the botanical community. The extent of these nomenclatural changes in Neotropical orchids, particularly those pertaining to the flora of Costa Rica, will become evident to the reader in this new edition of the catalogue. In fact, the sheer number of nomenclatural changes has compelled us to create a comprehensive taxonomic index. This index of both old and new names allows readers, especially non-specialists, to access the names of the species as they have traditionally been treated and identify which ones, among the various alternatives, are used in the catalogue to better align with the current phylogenetic hypotheses supported by diverse datasets. It is indeed appropriate to inquire at this point about the specific orchid taxonomy that the authors have chosen to adopt in creating this catalogue, considering the numerous possibilities that may be compatible with present and previous alternatives. Undoubtedly, modern systematics must reflect the natural history of the organisms in both the nomenclature system and the taxonomic hierarchies. Over the twenty-year gap between this version of the catalogue and its predecessor, studies based on DNA sequence comparisons have joined the traditional approach based on comparative morphology to infer the phylogenetic relationships among different orchid groups. These relationships, particularly within such a vast and diverse family as orchids, are sometimes obscured by numerous parallelisms - instances where independent forms have evolved analogous or almost identical morphological traits in response to similar evolutionary pressures. The new framework of orchid phylogenetics has been consolidated in the last two decades through Genera Orchidacearum (Pridgeon et al. 1999, 2001, 2003, 2005, 2009, 2014), a comprehensive work aimed at reflecting the evolutionary diversification of the family within a de Costa Rica se atribuye principalmente al descubrimiento de especies nuevas para la ciencia y en menor medida, a taxones recién registrados para el país como resultado de las exploraciones sistemáticas realizadas en el país durante el presente siglo. Se presenta un esquema filogenético de los géneros registrados en Costa Rica, desde el rango de subfamilia hasta el de subtribu, incluyendo una sinopsis de la diversidad registrada para cada género. También se proporciona una lista enumerada de las especies aceptadas y un índice completo de nombres aceptados y sinónimos. Éste índice enlaza la lista anotada de especies y el catálogo principal. Se proponen nuevas combinaciones para Andreettaea fimbriata y Camaridium muscoides. Se incluyen fotografías de especies representativas de la mayoría de los géneros basadas en ejemplares de Costa Rica. Esperamos que este catálogo inspire la investigación orquideológica y botánica en Costa Rica y otras regiones neotropicales, lo que llevará a descubrimientos adicionales en los próximos años. KeyworDs / PalaBras clave: biodiversidad, biodiversity, checklist, Costa Rican orchids, lista anotada, Neotropics, Neotrópico, orquídeas de Costa Rica LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 3PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae unified nomenclatural system. This monumental and unparalleled collective endeavor has finally provided botanists with an essential framework for family classification. It incorporates molecular analyses, comparative morphology of plants and flowers, and essential data derived from previous and recent studies on ultrastructure, chemistry, ecology, reproductive biology, and biogeography. This new edition of the catalogue of Costa Rican orchids is primarily structured based on the framework proposed by Genera Orchidacearum, with a few exceptions that can be broadly categorized into two essential types. The first type, which is relatively straightforward, arises from the emergence of more recent monographic studies. These studies have proposed different interpretations or uncovered critical relationships that were not resolved in previous studies, often due to the limited sample size. In other instances, corrections were deemed necessary due to new determinations of the material previously utilized for the analyses. These reevaluations often clarified issues that had limited resolution or provided evidence against previously improbable phylogenetic hypotheses. While less common, the second type of exception may warrant a more detailed explanation. Phylogenetic studies in Orchidaceae have clarified the relationships among groups primarily at the level of subfamilies, tribes, and subtribes (Pridgeon et al. 1999, 2001, 2003, 2005, 2009, 2014). However, determining the relationships at the genus and species levels remains challenging. This difficulty stems from two main factors, gene sampling, and species sampling. Firstly, phylogenetic reconstructions in the Orchidaceae have relied heavily on a limited number of molecular markers (plastid matK, rcbL, ycf1, and nuclear nrITS). Unfortunately, these markers sometimes fail to exhibit sufficient genetic divergence, particularly in the highly diverse and recently evolved neotropical orchid groups such as Cymbidieae and Pleurothallidinae (Pérez- Escobar et al. 2017). The advent of new genome sequencing techniques presents an opportunity to obtain hundreds of nuclear markers and complete plastid genomes, potentially enhancing the support values in molecular analyses and enabling better resolution of many orchid clades (Bogarín et al. 2018, Pérez- Escobar et al. 2021), but its application in species-rich orchid groups has been rather limited due to the cost of sequencing hundreds of genes for hundreds of taxa. The inadequate sampling of either genes or species remains a limitation in phylogenetic studies. Furthermore, the discovery of new species (and potential new clades) and clarification of taxonomic concepts improve the phylogenetic sampling of species. For example, we adopt the circumscription of Telipogon Kunth in a broad sense, which includes Dipterostele Schltr. and Stellilabium Schltr. as suggested in Genera Orchidacearum. This decision stems from the difficulties presented by phylogenetic analyses conducted on this group, characterized by low support values and limited resolution that hinder the confident differentiation among clades. Therefore, we advocate for a broad definition of Telipogon until future studies with broader gene and taxon sampling can effectively elucidate the clades within this group. A similar situation occurs with the adoption of a broad circumscription for the genus Stelis Sw. in the Pleurothallidinae (Karremans et al. 2013). Once the classification system aligns with all available data and ensures there are no contradictions with solid phylogenetic reconstructions, it is perhaps important to remember that the hypotheses of relationships derived from phylogenetic studies are not automatically reflected in a hierarchical classification system. The translation of the “objective” scheme of phylogenetic relationships, as shown in cladograms, into a formal hierarchical classification system remains primarily a human endeavor and thus is subject to a certain degree of subjectivity in decision-making. Even the less attentive reader may have noticed that the precise details in generic circumscriptions adopted in Genera Orchidacearum are not uniformly consistent across phylogenetic reconstructions of various groups. This variation in the definition of taxonomic units and their ranking untimely depends on the personal perspectives of individual taxonomists. While we have strived to stay faithful to the general framework of Genera Orchidacearum, this catalogue does present certain choices that reflect our “point of view” as botanists engaged in the daily interpretation of the rich orchid flora of Costa Rica. The index of names and their corresponding designations will undoubtedly assist readers in identifying these instances and independently selecting the nomenclatural system they consider most suitable for their specific requirements. LANKESTERIANA4 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. Materials and methods. This catalogue is based on previous checklists and taxonomic treatments of Orchidaceae from Costa Rica by Pupulin (2002), Dressler (2003), Ossenbach et al. (2007), and Ulloa Ulloa et al. (2023). We incorporated additions to the flora and updated the status of endemic species based on studies by Bogarín et al. (2008, 2013, 2015), Karremans et al. (2012), Fernández et al. (2014), and Belfort-Oconitrillo et al. (in prep). Additionally, we have incorporated information from recent literature and our continuous research on Costa Rican Orchidaceae (Pupulin 2022). Primary information was sourced from the following herbaria: AMES, BM, CR, F, JBL, K, MO, NY, SEL, US, USJ, and W. Additionally, we consulted various online databases, including Epidendra (2023), IPNI (2023), Tropicos (2023), and POWO (2023). This catalogue mainly adheres to the orchid phylogenetic classification and generic circumscriptions proposed in Genera Orchidacearum (Pridgeon et al. 1999, 2001, 2003, 2005, 2009, 2014). In some cases, we have integrated updates to these circumscriptions based on subsequent studies or our own botanical expertise and interpretation, especially within groups we have taxonomically reviewed. Data analysis and visualization were carried out using R (R Core Team 2023) with the following packages: dplyr (Wickham et al. 2023), ggplot2 (Wickham 2016), and viridis (Garnier et al. 2023). We also used Python (van Rossum & Drake 2009) with the libraries: pandas (McKinney 2011), matplotlib (Hunter 2007), and seaborn (Waskom et al. 2021). To explore the phylogenetic relationships among the genera of Costa Rican orchid flora, we retrieved the nrITS sequences for one representative of each genus from the NCBI GenBank database using the NCBI eUtils API. These sequences were then aligned using MAFFT v7.471 (Katoh & Standley 2013) with default settings and resulting alignments were manually inspected. Maximum Likelihood analyses were performed using IQ-TREE v2.2.2.6, with 1000 bootstrap replicates (Kalyaanamoorthy et al. 2017, Nguyen et al. 2015). The consensus phylogenetic tree and the species numbers per genus were visualized with the R packages APE, ggtree, and phytools (Paradis & Schliep 2019, Revell 2012, Yu et al. 2017). The final figures were edited in Adobe Illustrator 2021. Results. In this catalogue, we record 1684 species, four subspecies, 14 natural hybrids, and seven forms, across 201 genera and 2 nothogenera. All the subfamilies of Orchidaceae are represented, with the exception for Apostasioideae, whose genera are restricted to the tropical regions of Asia. The phylogenetic relationships among the major orchid groups in the country are shown in Fig. 1(1) and 1(2). The cladogram, which includes a representative species of all the orchid genera recorded in the country (except for × Bensteinia Christenson, × Cochlezella , J.M.H.Shaw, Psilochilus Barb.Rodr., Solenocentrum Schltr., and Uleiorchis Hoehne), provides valuable information on the natural history of the family in the country, reflecting its kinship with the floras of other regions, both from a chronological and geological point of view. The supra- specific taxonomy of the orchid flora of Costa Rica, based on its phylogenetically arrangement from DNA sequence data analyses, is summarized in a specific chapter of the catalogue. Epidendroideae is by far the most diverse orchid subfamily in Costa Rica, accommodating around 93% of all the species recorded in the country. Orchidoideae accounts for about 6% of the species, while Vanilloideae and Cypripedioideae represent only 1% and 0.1% of all the taxa included in the catalogue respectively (Fig. 2). The richest groups in the orchid flora of the country are subtribes Pleurothallidinae (ca. 600 spp.), Laeliinae (ca. 350 spp.), Oncidiinae (ca. 200 spp.), Maxillariinae (ca. 140 spp.), and Zygopetalinae (ca. 70 spp.). Aside from tribe Sobralieae (ca. 60 spp.), the remaining groups have less than 50 species in the country’s flora. Approximately 60% of all 21 subtribes have in Costa Rica less than 10 species (Fig. 3–4). The ten most species-rich orchid genera in Costa Rica are Epidendrum L. (245 spp.), Lepanthes Sw. (163 spp.), Stelis Sw. (109 spp.), Pleurothallis R.Br. (63 spp.), Camaridium Lindl. (51 spp.), Specklinia Lindl. (47 spp.), Sobralia Ruiz & Pav., and Telipogon Kunth (both with 40 spp.), Scaphyglottis Poepp. & Endl. (38 spp.), and Masdevallia Ruiz & Pav. (36 spp.) (Fig. 5). About 30% of the species recorded for the country are endemic to Costa Rica (Fig. 6), with the highest number of endemic species in Lepanthes (109 spp. = 67%), Epidendrum (76 spp. = 31%), Stelis (40 spp. = 37%), Telipogon (35 spp. = 88%), and Pleurothallis (31 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 5PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae spp. = 46%), Specklinia (14 spp. = 30%), Sobralia (14 spp. = 35%), and Masdevallia (13 spp. = 36%) (Fig. 7). The exotic flora includes the orchids Arundina graminifolia (D.Don) Hochr., Oeceoclades maculata (Lindl.) Lindl., Spathoglottis plicata Blume, and Phalaenopsis stuartiana Rchb.f. (Pupulin 2002, Bogarín & Pupulin 2021, Pupulin & Karremans 2022, Belfort-Oconitrillo et al. in prep). The catalogue includes 5300 orchid names at the specific and subspecific ranks, almost 70% of which are here treated as synonyms (Fig. 8–9). Also, taxonomic inflation is not uniform among genera and suprageneric groups. Discussion. The number of species recorded in this catalogue accounts for more than 5% of all known orchid species worldwide and approximately 11% of all Neotropical orchids, a remarkable percentage considering Costa Rica’s relatively small land size (Fig. 10–31). In fact, Costa Rica occupies only three- ten thousandths of the total land area on the planet. Regarding the orchid diversity index, which measures the relationship between diversity and country size, Costa Rica still holds the highest number of recorded orchid species and genera per area in the American tropics, if not worldwide. The number of orchid species documented in Costa Rica in relation to its area is more than double that of Ecuador, four times greater than Honduras, ten times greater than Mexico, and an astonishing seventy times greater than the orchid-rich Brazil. The total account of Costa Rican orchids recorded in this catalogue presents an increase of 324 species compared to its predecessor in 2002. This signifies a richness that is approximately 20% higher than that presented in the general catalogue of orchids from the Central American isthmus, which estimated that 1461 orchid species occur in Costa Rica (Ossenbach et al. 2007). The description and cataloging of orchid species in Costa Rica have experienced varied phases (Fig. 9). The orchidologists J. Lindley and H.G. Reichenbach were primary contributors to the initial species descriptions in Central America (Reichenbach 1866). Their work in Costa Rica intensified after 1846 with the arrival of prominent botanists and explorers, such as Augustus R. Endrés, C. Hoffmann, A.S. Oersted, J. von Warscewicz, and H. Wendland. Later, the number of published names saw significant growth in the early 1900s, peaking with the seminal contributions of Oakes Ames and Rudolf Schlechter by the mid- 1920s (Ames 1922, Schlechter 1921). This surge was synchronous with intensive botanical explorations by key figures like the brothers Brade, A.M. Brenes, C.H. Lankester, H. Pittier, A. Tonduz, and C. Wercklé. However, the subsequent decades, from the 1930s to the 1970s, observed a downturn in orchid research. A resurgence became evident in the early 1980s, marked by a wave of newly published orchid taxa. This resurgence mirrored the increased research endeavors of orchidologists such as J.T. Atwood, C.A. Luer, C.H. Dodson, and R.L. Dressler (Ossenbach 2009). The establishment of the Lankester Botanical Garden as part of the University of Costa Rica in 1973, further propelled this renewal. Throughout its 50-year legacy, JBL has been instrumental in fostering consistent progress in the study of Costa Rican orchids. The numerous nomenclatural changes resulting from improved taxonomic concepts might give the impression of numerous corrections compared to the edition published at the beginning of the millennium (Pupulin 2002). However, in essence – that is, as regards the identity of the species – the two lists align almost perfectly. There are certainly some exceptions and corrections are required for specimen identification, but the number disparity mainly stems from newly described species and those previously unknown in Costa Rica. These newly recorded species represent discoveries made in the past twenty years. In summary, it signifies a clear expansion of our understanding of the orchid diversity in Costa Rica. This increase in orchid species in the catalogue can be attributed mainly to exploration and research conducted by the staff of Lankester Botanical Garden over the past 20 years. It incorporates data from Bogarín (2011), who reported 34 new species and 24 new records since the publication of Orchids of the Central American isthmus (Ossenbach et al. 2007). Additionally, Karremans et al. (2012) compared the new state-of-art knowledge with the information compiled by Dressler (2003) in his orchid treatment for the Manual de plantas de Costa Rica. They found that roughly over approximately one decade, 130 species of orchids were described as new to science, LANKESTERIANA6 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. Apostasia Phragmipedium Cleistes Vanilla Epistephium Habenaria Goodyera Kreodanthus Microchilus Aspidogyne Platythelys Aa Gomphichis Prescottia Baskervilla Ponthieva Cranichis Pterichis Coccineorchis Cyclopogon Sarcoglottis Brachystele Pelexia Stenorrhynchos Mesadenella Pteroglossa Sacoila Aulosepalum Beloglottis Hapalorchis Eurystyles Lankesterella Greenwoodiella Schiedeella Spiranthes Funkiella Microthelys Triphora Monophyllorchis Elleanthus Sobralia Palmorchis Corymborkis Tropidia Calanthe Liparis Crossoglossa Malaxis Govenia Coelia Chysis Bletia Isochilus Nemaconia Arpophyllum Laelia Prosthechea Nidema Encyclia Caularthron Acrorchis Jacquiniella Barkeria Epidendrum Homalopetalum Scaphyglottis Dimerandra Brassavola Cattleya Guarianthe Myrmecophila Acianthera Brachionidium Myoxanthus Echinosepala Restrepia Dresslerella Pleurothallopsis Restrepiella Barbosella Anathallis Octomeria Zootrophion Karma Lankesteriana Lepanthopsis Trichosalpinx Stellamaris Draconanthes Lepanthes Trisetella Dracula Diodonopsis Masdevallia Pabstiella Pleurothallis Phloeophila Stelis Dryadella Andreettaea Scaphosepalum Platystele Specklinia 0 50 100 150 200 250 0 50 100 150 200 Number of species Number of species Figure 1(1). Genus-level phylogenetic tree (ML) based on nrITS sequences of representative genera of the Costa Rican orchid flora showing species numbers across genera. to Fig. 1(2) LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 7PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae Apostasia Phragmipedium Cleistes Vanilla Epistephium Habenaria Goodyera Kreodanthus Microchilus Aspidogyne Platythelys Aa Gomphichis Prescottia Baskervilla Ponthieva Cranichis Pterichis Coccineorchis Cyclopogon Sarcoglottis Brachystele Pelexia Stenorrhynchos Mesadenella Pteroglossa Sacoila Aulosepalum Beloglottis Hapalorchis Eurystyles Lankesterella Greenwoodiella Schiedeella Spiranthes Funkiella Microthelys Triphora Monophyllorchis Elleanthus Sobralia Palmorchis Corymborkis Tropidia Calanthe Liparis Crossoglossa Malaxis Govenia Coelia Chysis Bletia Isochilus Nemaconia Arpophyllum Laelia Prosthechea Nidema Encyclia Caularthron Acrorchis Jacquiniella Barkeria Epidendrum Homalopetalum Scaphyglottis Dimerandra Brassavola Cattleya Guarianthe Myrmecophila Acianthera Brachionidium Myoxanthus Echinosepala Restrepia Dresslerella Pleurothallopsis Restrepiella Barbosella Anathallis Octomeria Zootrophion Karma Lankesteriana Lepanthopsis Trichosalpinx Stellamaris Draconanthes Lepanthes Trisetella Dracula Diodonopsis Masdevallia Pabstiella Pleurothallis Phloeophila Stelis Dryadella Andreettaea Scaphosepalum Platystele Specklinia 0 50 100 150 200 250 0 50 100 150 200 Number of species Number of species LANKESTERIANA8 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. Bulbophyllum Campylocentrum Spathoglottis Arundina Polystachya Eulophia Oeceoclades Cyrtopodium Galeandra Catasetum Clowesia Dressleria Mormodes Cycnoches Lycaste Xylobium Inti Cryptocentrum Rhetinantha Ornithidium Nitidobulbon Heterotaxis Maxillaria Mormolyca Maxillariella Christensonella Camaridium Sauvetrea Mapinguari Trigonidium Peristeria Coeliopsis Gongora Paphinia Horichia Houlletia Trevoria Stanhopea Coryanthes Sievekingia Acineta Lacaena Kegeliella Polycycnis Eriopsis Cryptarrhena Galeottia Warrea Warreopsis Dichaea Huntleya Chondroscaphe Kefersteinia Benzingia Daiotyla Cochleanthes Stenotyla Warczewiczella Chaubardiella Pescatoria Trichocentrum Cohniella Lophiaris Cuitlauzina Rossioglossum Trichopilia Psychopsis Fernandezia Ornithocephalus Telipogon Lockhartia Oncidium Sigmatostalix Otoglossum Cyrtochiloides Miltoniopsis Cischweinfia Systeloglossum Mesospinidium Aspasia Brassia Rhynchostele Erycina Plectrophora Leochilus Rodriguezia Trizeuxis Comparettia Ionopsis Warmingia Macradenia Macroclinium Notylia 0 50 0 50 100 150 200 Number of species Number of species Figure 1(2). Genus-level phylogenetic tree (ML) based on nrITS sequences of representative genera of the Costa Rican orchid flora showing species numbers across genera. from Fig. 1(1) LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 9PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae Ep id en dr oi de ae O rc hi do id ea e Va ni llo id ea e C yp rip ed io id ea e 0 500 1000 1500 Nu m be r o f s pe cie s 1684 species 201 genera 2 nothogenera 4 subspecies 7 forms 14 natural hybrids Subfamily Number of species Tr ib e/ S ub tr ib e Figure 2. Distribution of species numbers across the orchid subfamilies within the Costa Rican orchid flora. Figure 3. Distribution of species numbers across the tribes/subtribes within the Costa Rican orchid flora. and a similar number of taxa were first recorded as part of the country’s flora. Considering the reevaluation of previously identified species, taxonomic changes (including additions, reductions, or nomenclatural changes) were observed in 61 out of the 174 genera of Costa Rican orchids featured in the Manual (Dressler 2003). This accounts for roughly one-third of all the genera. These findings highlight the dynamic nature of orchid taxonomy and the continuous updates and improvements in our understanding of the orchid flora in Costa Rica (Karremans et al. 2012). While the new catalogue introduces novelties across various major groups of known orchids in Costa Rica, the readers will likely notice that the most substantial increase is visible in those groups that researchers at Lankester Botanical Garden have studied most assiduously in recent decades. Similarly, it is apparent that many of the newly recorded floristic novelties in this edition of the catalogue pertain to small plants that had previously gone unnoticed during exploration activities. A great source of information regarding the country’s orchid diversity stems from ongoing floristic projects and the systematic exploration and inventory conducted by researchers at JBL in the national parks and protected areas of Costa Rica, with the support of the Ministry of Environment. Through meticulous LANKESTERIANA10 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 598 349 193 143 N um be r o f t ax a Subtribe Ma lax idi na e Figure 4. Distribution of taxa across the 10 most species-rich subtribes within the Costa Rican orchid flora. Genera N um be r o f t ax a Figure 5. Distribution of taxa across the 10 most species-rich genera within the Costa Rican orchid flora. and systematic surveys in recent years, we have not only improved previous generic identifications but also unveiled a hidden treasure of diversity that had never been appreciated (Pupulin 2022). It may not come as a surprise that Guayabo National Monument, encompassing a small area of only 3 km2 on the rich Caribbean side of the Cordillera Volcánica Central, harbors an impressive count of 120 orchids species, that is, an orchid flora comparable to that of Italy. Even more surprising is the fact that Manuel Antonio National Park situated on the Pacific coast of Costa Rica at sea level, covering only 4 km2, and traditionally considered an area of lower orchid diversity, is home to 39 orchid species (Pupulin 1988). In contrast, on the Caribbean coast, within the similarly sized Cahuita National Park we have recorded 51 orchid species (Pupulin & Bogarín 2018). Additionally, our inventory efforts in Barra Honda National Park, spanning 23 km2 in the middle of a sunburned savannah of the arid Península de Nicoya in Guanacaste, revealed a remarkable assemblage of 36 species across 23 genera. Notably, this included the LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 11PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae Endemic 30.4% 519 taxa Non-endemic 69.6% 1190 taxa Figure 6. Proportion of endemic taxa versus non-endemic taxa within the Costa Rican orchid flora. Figure 7. Proportion of endemic species for top 10 species-rich genera within the Costa Rican orchid flora. N um be r o f n am es Genera 63 222 24 491 245 552 Figure 8. Distribution of published and accepted names numbers within the Costa Rican orchid flora. discovery of a new genus, Tropidia Lindl., previously unknown to the flora of Costa Rica (Bogarín & Pupulin 2021). Strikingly, the pantropical Tropidia is not the only orchid genus newly recorded from Costa Rica since the last version of the catalogue. Also new to the country are Epistephium Kunth and Horichia Jenny. The first, a member of the Vanilloideae, is a South American genus with a single species reaching Central America: Epistephium ellipticum R.O.Williams & Summerh. The tiny E. ellipticum is a woodland plant that can be easily overlooked in the lowland forests of the Caribbean plains where it has been recently found (Karremans 2022, Karremans et al. 2012). While Horichia dressleri Jenny, the sole member of this monotypic genus, was previously known to occur LANKESTERIANA12 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. only in the in the Caribbean lowlands of western Panama but has since confirmed to occur in Costa Rica. This exceedingly beautiful orchid, belonging to the Stanhopeinae subtribe, is an example of an intrinsically rare orchid that would certainly not have gone unnoticed if it were more common (Pupulin & Karremans 2017). To these we may also add the Asian genus Spathoglottis Blume, of which one species, Spathoglottis plicata Blume, has now naturalized in Costa Rica (Pupulin & Karremans 2022). These exotic species are able to reproduce within the natural environments of Costa Rica. Nevertheless, there is presently no compelling evidence to categorize these species as invasive exotic plants or to suggest that they exert detrimental effects on natural ecosystems (Bogarín & Pupulin 2021). With regard to endemism, although it is expected that the number of endemic species will be greater in the most diverse groups such as Epidendrum, Lepanthes and Stelis, the high percentage within the genus Telipogon is notable, with 88% of its species endemic. The levels of endemism in these species-rich genera are comparable to those found in Panama, underscoring their floristic similarities (Bogarín et al. 2015). Further discoveries and improvements will surely continue to be made to this catalogue in the future and the reader may reasonably wonder about the level of completeness in terms of faithfully reflecting Costa Rican orchid diversity. In general terms, we can confidently answer that we have made great progress compared to the catalogue from twenty years ago. This progress is attributed to our extensive exploration efforts throughout the country and the meticulous work dedicated to accurately applying the names. Nevertheless, we are also aware that such a catalogue is always a work in progress. We confess to having several taxonomic novelties that await attention in our drawers at Lankester Botanical Garden, and have no doubt that more will be found as remote areas are explored and complex taxa are monographed. This catalogue provides an updated summary of the orchid flora of Costa Rica, reflecting our most current knowledge. Benefiting from an unparalleled documentary foundation, we believe that in terms of the completeness of direct sources, most of which we have digitized over the last two decades, it has no parallel among other Neotropical regions. We have accurately compared the protologues and available type materials for 5249 orchid names. Of these, over 3500 had to be included under the synonymy of other, accepted, taxa, encompassing both homotypic and heterotypic synonyms. With respect to the latter, we have taken an extremely conservative approach. We only included species in the catalogue that were described from Costa Rica or the immediate neighboring countries, and only those for which we had the opportunity to examine specimens collected at the type localities. Additionally, we incorporated taxa with a broad distribution spanning the dry, lowland regions prevalent across most tropical and subtropical American regions. Once we filter the excess of synonyms that resulted from an overly broad circumscription of the species, a fascinating and somewhat new phylogeographic perspective of the Costa Rican orchid flora emerges. While Costa Rica and the mountainous region of western Panama constitutes almost a biotic unit (together termed by us the “Central American Andes”), the relationships with the true Andean flora are less pronounced. Similarly, their affinity with the floras of southern Mexico and northern Central America is limited. The floristic affinities with the Antillean flora and the Guyana region are even more constrained with which the orchid flora of Costa Rica shares very few species with these areas. Finally, almost no species of Costa Rica are shared with the extremely orchid-rich regions Brazilian Atlantic Forest. As we noted before, taxonomic inflation has led to a large number of synonyms, accounting for nearly three-quarters of all names used in the flora of Costa Rica. Interestingly, these superfluous names are concentrated in those genera, such as Epidendrum, Maxillaria, and Pleurothallis, whose relative frequency attracted the taxonomic attention of botanists and naturalists during the initial exploration of the American tropics. In contrast, the more inconspicuous but still diverse groups, such as Lepanthes or Telipogon, comprised mostly of small plants have received systematic attention only in recent decades. As a result, the ratio between accepted names and synonyms is considerably lower (Fig. 9). We attribute this largely to the enhanced communication among scholars facilitated by the advent of information LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 13PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae technology. This advancement has enabled almost real-time access to taxonomic novelties published in specialized literature. Furthermore, recent improvements in documentation and illustration techniques, using more sophisticated tools, have made morphological comparisons more efficient and precise. As a result, the unnecessary duplication of names, which once troubled traditional taxonomy, has been significantly reduced. Unfortunately, the systematic work aimed at fine taxonomic revision has not been evenly distributed across the different groups of orchids in Costa Rica. However, we believe that the catalogue is a faithful representation of the country’s true diversity, particularly in the treatment of some groups that we have extensively studied over the past 20 years. We can confidently consider the catalogue complete or nearly complete in certain subtribes, including Angraecinae, Catasetinae, Maxillariinae, Oncidiinae, Stanhopeinae, and Zygopetalinae. Most of the groups within the subtribe Pleurothallidinae have been largely resolved, except for the genus Stelis and, in part, Lepanthes. Subtribe Laeliinae is almost entirely covered, although novelties will surely continue to be found in Epidendrum L. (Karremans et al. 2021). We have also made substantial progress with monographs on genera within the subfamilies Cypripedioideae and Vanilloideae, as well as with less diverse groups consisting of one or a few species, such as: Bletiinae, Bulpbophylinae, Coeliinae, Cyrtopodiinae, Eriopsidinae, among others. While we do not anticipate significant changes in tribe Sobralieae (with about 60 taxa), many other groups of the early diverging clades within Epidendroideae have not yet been thoroughly studied in depth. Monographic studies on mostly terrestrial Orchidoideae have only recently been initiated, and we may expect a considerable number of taxonomic novelties in groups such as Cranichidinae (ca. 30 known taxa), Goodyerinae, Spiranthinae (ca. 50), and Orchidinae (less than 100 species in total). Among the derived groups of Epidendroideae, the taxonomic challenge persists in the pleurothallids, which encompass approximately 600 species across 37 genera in Costa Rica. Within this group, there are some genera for which we have good reason to suspect that the diversity of names we use in this catalogue does not accurately reflect the actual species diversity in the flora of Costa Rica. One such a case is the genus Stelis, of which we have included 144 species here. However, determining the true extent of the diversity of this genus remains largely speculative, based on a broad morphological assessment of two thousand individuals grown in the collections at JBL. The photographic treatment of some broadly circumscribed morpho-species of Stelis offered in Pupulin (2022) exemplifies this situation. But there are also other genera for which the underestimation of diversity is not a mere suspicion but an absolute certainty. One notable example is Pleurothallis, particularly the species characterized by a small habit and small flowers, colloquially referred as “frogs”. We are currently completing the taxonomic revision of 1 10 100 18 45 18 50 18 55 18 60 18 65 18 70 18 75 18 80 18 85 18 90 18 95 19 00 19 05 19 10 19 15 19 20 19 25 19 30 19 35 19 40 19 45 19 50 19 55 19 60 19 65 19 70 19 75 19 80 19 85 19 90 19 95 20 00 20 05 20 10 20 15 20 20 Year of publication Nu m be r o f n am es p ub lis he d (L og ) Figure 9. Historical count of orchid names published by year for the flora of Costa Rica. LANKESTERIANA14 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. this group for the flora of Costa Rica. Even though the study could not be finished before the publication of this catalogue, in the case of frog-like Pleurothallis species, we are confident that the number of species recognized for Costa Rica must be at least quadrupled. However, it is important to note that the names of these taxa are not yet available to be presented to the reader. In terms of botanical exploration, we anticipate a rise in species within the highly diverse genera of Laeliinae, Oncidiinae, and Pleurothallidinae as we intensify our exploration of remote areas, including the Cordillera de Talamanca, Fila Matama, and upper volcanic areas of the Cordillera de Tilarán and Guanacaste. Having a more accurate and up-to-date catalogue of the Orchidaceae of Costa Rica is today perhaps more important than ever before. Not only is it critical to have basic information on the taxonomy and distribution of genera and species at the disposal of botanists, but this data is also increasingly leveraged by conservationists and policymakers to establish priorities and passing legislation. Costa Rica stands as a major orchid diversity hotspot, being home to a significant number of endemic species. It is our collective responsibility to safeguard this extraordinary biological richness. A primary factor contributing to this exuberant diversity is the range of ecosystems present across the country, emphasizing the crucial role that each biological reserve and protected natural area plays in safeguarding local species. From the well-known Parque Nacional Manuel Antonio on the Pacific coast to Parque Nacional Cahuita on the Caribbean coast, as well as the extensive and mostly unexplored highlands of Parque Internacional La Amistad in the heart of the country, all of these areas harbor unique orchid floras. Conclusion. Costa Rica boasts a unique orchid diversity, housing over 5% of the world’s known orchid species within its relatively small landmass. Recent phylogenomic studies on orchid biogeography suggest that southern Central America displays the highest speciation rates on a global scale, demonstrating that the orchid flora of Costa Rica underwent recent and rapid speciation (Pérez-Escobar et al. 2023). This positions Costa Rica as a key biological center of speciation in the Neotropics. Such studies are only possible with a well-established system of information and continuous research on orchid taxonomy, which forms the foundation of modern biological studies. With a legacy dating back to pioneering orchidologists in 1846 and sustained by institutions like the Lankester Botanical Gardens today, our knowledge of the country’s orchid flora has considerably grown. This is underscored by the recent catalogue update, which reflects a 24% rise in orchid species since the early 2000s. Costa Rica is now well on its way to register over 1700 orchid species, a staggering number that astonishes both locals and foreign visitors alike. Beyond being mere numbers, these species represent the diverse ecosystems and unique ecological niches of a highly biodiverse country with well-established conservation policies through its network of protected areas. Fortunately, the country has demonstrated its commitment to this cause through its public institutions for decades. This updated catalog will serve as an invaluable resource for students, botanists, conservationists, tourism professionals, and policymakers. We anticipate that this catalog will catalyze orchidological and botanical research not only in Costa Rica but also in other Neotropical regions, propelling further discoveries in the upcoming years. acKnowleDgments We sincerely express our gratitude to our friend and colleague, Jorge Warner, for his invaluable and unwavering support of the Flora Costaricensis projects under JBL’s direction. These research projects have been pivotal in updating the orchid checklist, providing a robust foundation for this catalogue. We are grateful to the Rectory and Vice-Rectory for Research of the University of Costa Rica for their constant support with equipment, facilities, and materials provided to the Microscopy and Photography laboratories at the Lankester Botanical Garden and for sponsoring the ongoing projects toward the completion of the illustrated orchid flora of Costa Rica (Flora Costaricensis), including the projects C2056, C2051, C2050, C1060, C1059, C1058, C1057, C1056, C1055, C0051, C0050, C2055, and C2056. Our sincere appreciation goes to the Ministerio de Ambiente y Energía (MINAE) and the Sistema Nacional de Áreas de Conservación (SINAC) of Costa Rica. Their support, including access to protected areas and the issuance of scientific permits, has been crucial to our research over the last decades. Our gratitude also extends to the dedicated individuals responsible for the care and maintenance of the living orchid collections at the Lankester Botanical Garden: Reinaldo Gómez, Giovanni Meza, Gustavo Rojas Aguilar, Nelson Solano, LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 15PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae literature citeD Ames, O. (1922). Orchidaceae quaedam Americanae. Schedulae Orchidianae, 1, 1–24. Bogarín, D. (2011). How many orchid species in Costa Rica? A review of the latest discoveries. In: A. M. Pridgeon & H. G. Navarrete Zambrano (Eds.), Proceedings of the Third Scientific Conference on Andean Orchids, Quito (pp. 185–205). Lankesteriana: International Journal on Orchidology, 11(3), 185–205. Bogarín, D., Karremans, A. & Pupulin, F. (2008). New species and records of Orchidaceae from Costa Rica. Lankesteriana: International Journal on Orchidology, 8(2), 53–74. https://doi.org/10.15517/lank.v0i0.7927 Bogarín, D., Pérez-Escobar, O. A., Groenenberg, D., Holland, S. D., Karremans, A. P., Lemmon, E. M., Lemmon, A. R., Pupulin, F., Smets, E. & Gravendeel, B. (2018). Anchored hybrid enrichment generated nuclear, plastid and mitochondrial markers resolve the Lepanthes horrida (Orchidaceae: Pleurothallidinae) species complex. Molecular Phylogenetics and Evolution, 129, 27–47. Bogarín, D. & Pupulin, F. (2021). The orchid flora of Barra Honda National Park, Nicoya, Guanacaste, Costa Rica. Harvard Papers in Botany, 26(1), 7–99. Bogarín, D., Pupulin, F., Arrocha, C. & Warner, J. (2013). Orchids without borders: studying the hotspot of Costa Rica and Panama. Lankesteriana: International Journal on Orchidology, 13(1–2). https://doi.org/10.15517/lank.v0i0.11529 Bogarín, D., Serracín, Z., Samudio, Z., Rincón, R. & Pupulin, F. (2015). An updated checklist of the Orchidaceae of Panamá. 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Rafael Trejos, and Gerson Villalobos. Additionally, we are indebted to those who actively participated in fieldwork or generously provided invaluable data on the species featured in this catalog. We express our sincerest thanks to Smith Abarca, Rafael Acuña, Leonardo Álvarez, Noelia Belfort-Oconitrillo, Mario A. Blanco, David Carazo, Alexis Castillo, Marco Cedeño, Isler Chinchilla, Jorge de la Cruz, Melissa Díaz, Melania Fernández, Reinaldo Gómez, Daniel Jiménez, Esteban Jiménez, Eberhard Kaes, Stephen Kirby, Miguel Méndez, Robert Müntz, Lizbeth Oses, Josué Pacheco, Raimundo Parra, Oscar A. Pérez-Escobar, Jeremy Quesada, José Manuel Quesada, Ana Cristina Rodríguez, Gustavo Rojas-Alvarado, Grettel Salguero, Rafael Trejos, and Gerson Villalobos. We are also grateful to all members of the orchid associations in Costa Rica, especially ACO, AJUCORI, ANARO, and ASOCOR for their valuable contributions. Special thanks to our colleague Isler Chinchilla for kindly reviewing and updating the Malaxidinae taxonomy and to three anonymous reviewers for their valuable observations that improved the manuscript. Finally, we extend our gratitude to Armando Estrada, Silvia Lobo, and Alexander Rodríguez from the Museo Nacional de Costa Rica (CR), along with Rafael Acuña and Mario A. Blanco (USJ), for their invaluable assistance during our visits to study herbarium materials. We also appreciate the cooperation of staff and curators at AMES, FLAS, INB, JBL, K, MO, NY, SEL, and W, who granted us access to their collections and shared valuable information. LANKESTERIANA16 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. Karremans, A. P., Bakker, F. T., Pupulin, F., Solano-Gomez, R. & Smulders, M. J. M. (2013). Phylogenetics of Stelis and closely related genera (Orchidaceae: Pleurothallidinae). Plant Systematics and Evolution, 29(1), 69–86. Katoh, K., & Standley, D. M. (2013). 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J., Li, L., Negrao, R., Oses, L., Phillips, C., Rincón, M., Salazar-Chavez, G., Simpson, L., Smidt, E., Solano-Gomez, R., Parra-Sánchez, E., Tremblay, R. L., van den Berg, C., Villanueva, B. S., Zuluaga, A., Chase, M. W., Fay, M. F., Condamine, F. L., Forest, F., Nargar, K., Renner, S. S., Baker, W. J. & Antonelli, A. (2023). The origin and speciation of orchids. BioRxiv, 2023.09.10.556973. https://doi.org/10.1101/2023.09.10.556973 Pérez-Escobar, O. A., Chomicki, G., Condamine, F. L., Karremans, A. P., Bogarín, D., Matzke, N. J., Silvestro, D. & Antonelli, A. (2017). Recent origin and rapid speciation of Neotropical orchids in the world’s richest plant biodiversity hotspot. New Phytologist, 215, 891–905. Pérez-Escobar, O. A., Dodsworth, S., Bogarín, D., Bellot, S., Balbuena, J. A., Schley, R. J., Kikuchi, I. A., Morris, S. K., Epitawalage, N., Cowan, R., Maurin, O., Zuntini, A., Arias, T., Serna-Sánchez, A., Gravendeel, B., Torres Jimenez, M. F., Nargar, K., Chomicki, G., Chase, M. W., Leitch, I. J., Forrest, F. & Baker, W. J. (2021). Hundreds of nuclear and plastid loci yield novel insights into orchid relationships. American Journal of Botany, 108(7), 1166–1180. https://doi. org/10.1002/ajb2.1702 POWO. (2023, December 26). Plants of the World Online. Facilitated by Kew Royal Botanic Gardens. Retrieved from https:// powo.science.kew.org/Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (1999). Genera Orchidacearum. Volume 1. Apostasioideae and Cypripedioideae. Oxford: Oxford University Press. Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (2001). Genera Orchidacearum. Volume 2. Orchidoideae (Part one). Oxford: Oxford University Press. Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (2003). Genera Orchidacearum. Volume 3. Orchidoideae (Part 2). Vanilloideae. Oxford: Oxford University Press. Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (2005). Genera Orchidacearum. Volume 4. Epidendroideae (Part one). Oxford: Oxford University Press. Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (2009). Genera Orchidacearum. Volume 5. Epidendroideae (Part two). Oxford: Oxford University Press. Pridgeon, A. M., Cribb, P. J., Chase, M. W. & Rasmussen, F. (2014). Genera Orchidacearum. Volume 6: Epidendroideae (Part three): Oxford, Oxford University Press. Pupulin, F. (1988). Orchid florula of Parque Nacional Manuel Antonio, Quepos, Costa Rica. Revista de Biología Tropical, 46(4), 961–1031. Pupulin, F. (2002). Catálogo revisado y anotado de las Orchidaceae de Costa Rica. Lankesteriana: International Journal on Orchidology, 4, 1–88. doi: https://doi.org/10.15517/lank.v2i2.23095 Pupulin, F. (2022). Inexhaustible diversity. In: F. Pupulin & collaborators (Eds.), Vanishing Beauty: Native Costa Rican Orchids. Vol. 3. Restrepia–Zootrophion and Appendices (pp. vii–xv). Oberreifenberg, Germany: Koeltz Botanical Books. LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 17PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae Pupulin, F. & Bogarín, D. (2018). Orchids of paradise. Exploring the lower Talamanca Sashores of Costa Rica, the “Coast of Plenty” of Columbus. Orchids (American Orchid Society Bulletin), 87, 846–857. Pupulin, F. & Karremans, A. P. (2017). Horichia dressleri. A new genus recorded for Costa Rica and a truly rare orchid? Lindleyana in Orchids (American Orchid Society Bulletin), 86(8), 610–617. Pupulin, F. & Karremans, A. P. (2022). Spathoglottis Blume. In: F. Pupulin & collaborators (Eds.), Vanishing Beauty: Native Costa Rican Orchids. Vol. 3. Restrepia–Zootrophion and Appendices (pp. 1114–1115). Oberreifenberg, Germany: Koeltz Botanical Books. R Core Team. (2023). R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. ISBN 3-900051-07-0. Retrieved from http://www.R-project.org/ Revell, L. J. (2012). phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3(2), 217–223. Reichenbach, H. G. (1866). Beitrage zu einer orchideenkunde Central-Amerika’s. Hamburg: T.G. Meissner. 97 pp. Schlechter, F. R. R. (1921). Orchidaceae novae et criticae. Decas LXIX. Additamenta ad Orchideologiam Central- Americanam. Repertorium Specierum Novarum Regni Vegetabilis Beihefte, 17(481–485), 138–144. Tropicos. (2023, December 26). Tropicos v3.4.2. Facilitated by the Missouri Botanical Garden. Retrieved from https:// tropicos.org Ulloa Ulloa, C., Hernández, H. M., Davidse, G., Barrie, F. R., Knapp, S. & Dressler, R. (2023). Flora Mesoamericana, Vol 7, Parte 2: Orchidaceae. Universidad Nacional Autónoma de México, Missouri Botanical Garden Press, and the Natural History Museum (London). Van Rossum, G. & Drake, F. L. (2009). Python 3 Reference Manual. Scotts Valley, CA: CreateSpace. Waskom, M. (2021). Seaborn: statistical data visualization. The Journal of Open Source Software, 6(60), 3021. https://doi. org/10.21105/joss.03021 Wickham, H. (2016). ggplot2: Elegant Graphics for Data Analysis. New York: Springer-Verlag. ISBN 978-3-319-24277-4, https://ggplot2.tidyverse.org. Wickham, H., François, R., Henry, L., Müller, K. & Vaughan, D. (2023). dplyr: A Grammar of Data Manipulation. https:// dplyr.tidyverse.org, https://github.com/tidyverse/dplyr. Yu, G., Smith, D. K., Zhu, H., Guan, Y. & Lam, T. T. (2017). ggtree: An R package for visualization and annotation of phylogenetic trees with their covariates and other associated data. Methods in Ecology and Evolution, 8(1), 28–36. doi: https://doi.org/10.1111/2041-210X.12628 LANKESTERIANA18 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 1. Aa paleacea (Kunth) Rchb.f. 2. Acianthera aberrans (Luer) Pupulin & Bogarín 3. Acianthera breedlovei Soto Arenas, Solano & Salazar 4. Acianthera brunnescens (Schltr.) Karremans * 5. Acianthera butcheri (L.O.Williams) Pridgeon & M.W.Chase 6. Acianthera cabiriae Pupulin, G.A.Rojas & J.D. Zúñiga * 7. Acianthera cachensis (Ames) Karremans * 8. Acianthera circumplexa (Lindl.) Pridgeon & M.W.Chase 9. Acianthera cogniauxiana (Schltr.) Pridgeon & M.W.Chase 10. Acianthera costaricensis (Schltr.) Pupulin & Karremans * 11. Acianthera crassilabia (Ames & C.Schweinf.) Luer * 12. Acianthera decipiens (Ames & C.Schweinf.) Pridgeon & M.W.Chase * 13. Acianthera ellipsophylla (L.O.Williams) Pridgeon & M.W.Chase 14. Acianthera erinacea (Rchb.f.) A.Doucette * 15. Acianthera fecunda Pupulin, G.A.Rojas & J.D. Zuñiga * 16. Acianthera geminicaulina (Ames) Pridgeon & M.W.Chase 17. Acianthera hamata Pupulin & G.A.Rojas * 18. Acianthera hondurensis (Ames) Pridgeon & M.W.Chase 19. Acianthera lanceana (G.Lodd. ex Lindl.) Pridgeon & M.W.Chase 20. Acianthera lepidota (L.O.Williams) Pridgeon & M.W.Chase 21. Acianthera oscitans (Ames) Pridgeon & M.W.Chase 22. Acianthera pantasmi (Rchb.f.) Pridgeon & M.W.Chase 23. Acianthera sicaria (Lindl.) Pridgeon & M.W.Chase 24. Acianthera sigmoidea (Ames & C.Schweinf.) A.Doucette * 25. Acianthera testifolia (Sw.) Solano 26. Acianthera verecunda (Schltr.) Pridgeon & M.W.Chase 27. Acineta densa Lindl. 28. Acineta sella-turcica Rchb.f. 29. Acrorchis roseola Dressler 30. Anathallis endresii (Luer) Pridgeon & M.W.Chase * 31. Anathallis funerea (Barb.Rodr.) Luer 32. Anathallis grayumii (Luer) Luer * 33. Anathallis lewisiae (Ames) Solano & Soto Arenas 34. Anathallis minutalis (Lindl.) Pridgeon & M.W.Chase 35. Anathallis sertularioides (Sw.) Pridgeon & M.W.Chase 36. Anathallis transtalamancana Karremans & Chinchilla * 37. Andreettaea cryptophyta (Barb.Rodr.) A.Doucette 38. Andreettaea divexa (Ames) A.Doucette 39. Andreettaea exesilabia (A.H.Heller & A.D.Hawkes) A.Doucette 40. Andreettaea fimbriata (Ames & C.Schweinf.) Pupulin, Bogarín & Karremans 41. Andreettaea herpestes (Luer) A.Doucette * 42. Andreettaea marginata (Rich.) A.Doucette 43. Andreettaea quinqueseta (Ames) A.Doucette 44. Andreettaea samacensis (Ames) A.Doucette 45. Andreettaea segregatifolia (Ames & C.Schweinf.) A.Doucette 46. Andreettaea strumosa (Ames) A.Doucette 47. Andreettaea tapantiensis (Mel.Fernández & Bogarín) A.Doucette * 48. Arpophyllum giganteum Hartw. ex Lindl. 49. Arundina graminifolia (D.Don) Hochr. 50. Aspasia epidendroides Lindl. 51. Aspasia principissa Rchb.f. 52. Aspidogyne clavigera (Rchb.f.) Meneguzzo 53. Aspidogyne costaricensis Ormerod & M.A.Blanco * 54. Aspidogyne grayumii Ormerod * 55. Aspidogyne roseoalba (Dressler) Ormerod 56. Aspidogyne stictophylla (Schltr.) Garay 57. Aspidogyne tuerckheimii (Schltr.) Garay 58. Aspidogyne utriculata (Dressler) Szlach. * 59. Aulosepalum pyramidale (Lindl.) M.A.Dix & M.W.Dix 60. Barbosella dolichorhiza Schltr. 61. Barbosella geminata Luer * 62. Barbosella orbicularis Luer THE CHECKLIST1 1 Taxa marked with an asterisk are here treated as endemic to Costa Rica. LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 19PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae 63. Barbosella prorepens (Rchb.f.) Schltr. 64. Barkeria lindleyana Bateman ex Lindl. * 65. Barkeria obovata (C.Presl) Christenson 66. Baskervilla colombiana Garay 67. Baskervilla leptantha Dressler * 68. Beloglottis costaricensis (Rchb.f.) Schltr. * 69. Beloglottis ecallosa (Ames & C.Schweinf.) Hamer & Garay 70. Beloglottis hameri Garay 71. Beloglottis subpandurata (Ames & C.Schweinf.) Garay 72. × Bensteinia ramonensis Pupulin * 73. Benzingia reichenbachiana (Schltr.) Dressler 74. Bletia campanulata Lex. in P. de La Llave & J.M. de Lexarza 75. Bletia purpurea (Lam.) A.DC. 76. Brachionidium cornu-nigricum Bogarín & Karremans * 77. Brachionidium cruziae L.O.Williams * 78. Brachionidium dressleri Luer * 79. Brachionidium folsomii Dressler 80. Brachionidium haberi Luer * 81. Brachionidium kirbyi Bogarín, Karremans & M.Muñoz * 82. Brachionidium minusculum Luer & Dressler * 83. Brachionidium polypodium Luer 84. Brachionidium pusillum Ames & C.Schweinf. 85. Brachionidium valerioi Ames & C.Schweinf. 86. Brachystele guayanensis (Lindl.) Schltr. 87. Brassavola acaulis Lindl. & Paxton 88. Brassavola grandiflora Lindl. 89. Brassavola nodosa (L.) Lindl. 90. Brassia caudata (L.) Lindl. 91. Brassia chlorops Endrés & Rchb.f. 92. Brassia gireoudiana Rchb.f. & Warsz. 93. Brassia longissima (Rchb.f.) Nash 94. Brassia suavissima Pupulin & Bogarín * 95. Brassia verrucosa Bateman ex Lindl. 96. Bulbophyllum aristatum (Rchb.f.) Hemsl. 97. Bulbophyllum pachyrachis (A.Rich.) Griseb. 98. Calanthe calanthoides (A.Rich. & Galeotti) Hamer & Garay 99. Camaridium adolphi Schltr. 100. Camaridium alfaroi (Ames & C.Schweinf.) M.A.Blanco * 101. Camaridium amabile (J.T.Atwood) M.A.Blanco * 102. Camaridium amparoanum Schltr. * 103. Camaridium ampliflorum (C.Schweinf.) M.A.Blanco 104. Camaridium anceps (Rchb.f.) M.A.Blanco 105. Camaridium aurantiacum (Schltr.) M.A.Blanco 106. Camaridium biolleyi (Schltr.) Schltr. 107. Camaridium bracteatum (Schltr.) Schltr. 108. Camaridium bradeorum Schltr. 109. Camaridium brenesii Schltr. 110. Camaridium brevilabium (Ames & Correll) M.A.Blanco 111. Camaridium burgeri (J.T.Atwood) M.A.Blanco * 112. Camaridium campanulatum (C.Schweinf.) M.A.Blanco 113. Camaridium cedralense (J.T.Atwood & Mora- Ret.) M.A.Blanco * 114. Camaridium costaricense Schltr. 115. Camaridium ctenostachys (Rchb.f.) Schltr. 116. Camaridium cucullatum (Lindl.) M.A.Blanco 117. Camaridium dendrobioides Schltr. 118. Camaridium dichotomum Schltr. 119. Camaridium exaltatum Kraenzl. 120. Camaridium falcatum (Ames & Correll) M.A.Blanco 121. Camaridium gomezianum (J.T.Atwood) M.A.Blanco 122. Camaridium haberi (J.T.Atwood) M.A.Blanco * 123. Camaridium horichii (Senghas) M.A.Blanco 124. Camaridium imbricatum Schltr. 125. Camaridium inauditum (Rchb.f.) M.A.Blanco 126. Camaridium lankesteri (Ames) M.A.Blanco 127. Camaridium latifolium Schltr. 128. Camaridium micranthum M.A.Blanco 129. Camaridium microphyton (Schltr.) M.A.Blanco 130. Camaridium minus Schltr. 131. Camaridium mombachoense (A.H.Heller ex J.T.Atwood) M.A.Blanco 132. Camaridium monteverdense (J.T.Atwood & Barboza) M.A.Blanco 133. Camaridium muscoides (J.T.Atwood) Bogarín, Pupulin & Karremans * 134. Camaridium neglectum (Schltr.) M.A.Blanco 135. Camaridium nutantiflorum Schltr. 136. Camaridium obscurum (Linden & Rchb.f.) M.A.Blanco 137. Camaridium paleatum (Rchb.f.) M.A.Blanco 138. Camaridium pygmaeum M.A.Blanco 139. Camaridium ramonense (Schltr.) M.A.Blanco 140. Camaridium sigmoideum (C.Schweinf.) M.A.Blanco 141. Camaridium standleyi M.A.Blanco LANKESTERIANA20 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 142. Camaridium stenophyllum (Schltr.) M.A.Blanco 143. Camaridium strumatum (Endrés & Rchb.f.) M.A.Blanco 144. Camaridium suaveolens (Barringer) M.A.Blanco * 145. Camaridium tigrinum (C.Schweinf.) M.A.Blanco 146. Camaridium tuberculare (J.T.Atwood) M.A.Blanco 147. Camaridium vaginale (Rchb.f.) M.A.Blanco 148. Camaridium valerioi (Ames & C.Schweinf.) M.A.Blanco 149. Camaridium vittariifolium (L.O.Williams) M.A.Blanco * 150. Campylocentrum amistadense Bogarín * 151. Campylocentrum brenesii Schltr. 152. Campylocentrum generalense Bogarín & Pupulin 153. Campylocentrum lankesteri Ames 154. Campylocentrum multiflorum Schltr. 155. Campylocentrum panamense Ames. 156. Campylocentrum poeppigii (Rchb.f.) Rolfe 157. Campylocentrum schiedei (Rchb.f.) Benth. ex Hemsl. 158. Campylocentrum stenanthum Schltr. 159. Campylocentrum tenellum Todzia 160. Catasetum maculatum Kunth 161. Cattleya dowiana Bateman & Rchb.f. subsp. dowiana f. dowiana Cattleya dowiana subsp. dowiana f. rosita (Pfau) Pupulin * Cattleya dowiana subsp. dowiana f. carmoniana Pupulin * 162. Caularthron bilamellatum (Rchb.f.) R.E.Schult. 163. Chaubardiella pacuarensis Jenny * 164. Chaubardiella subquadrata (Schltr.) Garay 165. Chondroscaphe atrilinguis Dressler 166. Chondroscaphe bicolor (Rolfe) Dressler 167. Chondroscaphe yamilethiae Pupulin * 168. Christensonella uncata (Lindl.) Szlach. 169. Chysis costaricensis Schltr. * 170. Chysis tricostata Schltr. 171. Cischweinfia dasyandra (Rchb.f.) Dressler & N.H.Williams 172. Cischweinfia donrafae Dressler & Dalström * 173. Cischweinfia gersonii Pupulin, Bogarín & Karremans * 174. Cleistes costaricensis Christenson 175. Cleistes rosea Lindl. 176. Clowesia warczewitzii (Lindl. & Paxton) Dodson 177. Coccineorchis bracteosa (Ames & C.Schweinf.) Garay 178. Coccineorchis cernua (Lindl.) Garay 179. Coccineorchis navarrensis (Ames) Garay 180. Coccineorchis standleyi (Ames) Garay 181. Coccineorchis warszewicziana Szlach. 182. Cochleanthes aromatica (Rchb.f.) R.E.Schult. & Garay 183. × Cochlezella costaricensis Pupulin * 184. Coelia macrostachya Lindl. 185. Coeliopsis hyacinthosma Rchb.f. 186. Cohniella ascendens (Lindl.) Christenson 187. Cohniella brachyphylla (Lindl.) Cetzal & Carnevali 188. Cohniella × francoi Cetzal & Carnevali * 189. Comparettia aurea (Schltr.) M.W.Chase & N.H.Williams 190. Comparettia falcata Poepp. & Endl. f. falcata Comparettia falcata f. alba Pupulin 191. Coryanthes hunteriana Schltr. 192. Coryanthes kaiseriana G.Gerlach 193. Coryanthes picturata Rchb.f. 194. Corymborkis flava (Sw.) Kuntze 195. Corymborkis forcipigera (Rchb.f. & Warsz.) L.O.Williams 196. Cranichis acuminatissima Ames & C.Schweinf. * 197. Cranichis ciliata Kunth 198. Cranichis curtii Schltr. 199. Cranichis diphylla Sw. 200. Cranichis irazuensis Schltr. 201. Cranichis lankesteri Ames 202. Cranichis muscosa Sw. 203. Cranichis reticulata Rchb.f. * 204. Cranichis saccata Ames * 205. Cranichis talamancana Dressler * 206. Cranichis wageneri Rchb.f. 207. Crossoglossa aurantilineata Pupulin * 208. Crossoglossa blephariglottis (Schltr.) Dressler ex Dodson 209. Crossoglossa dressleri Ormerod 210. Crossoglossa eustachys (Schltr.) Dressler ex Dodson 211. Crossoglossa fratrum (Schltr.) Dressler ex Dodson 212. Crossoglossa sotoana Pupulin & Karremans * 213. Crossoglossa tipuloides (Lindl.) Dodson 214. Cryptarrhena guatemalensis Schltr. 215. Cryptarrhena lunata R.Br. 216. Cryptocentrum calcaratum (Schltr.) Schltr. 217. Cryptocentrum flavum Schltr. 218. Cryptocentrum gracilipes Schltr. * 219. Cryptocentrum gracillimum Ames & C.Schweinf. 220. Cryptocentrum latifolium Schltr. LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 21PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae 221. Cryptocentrum lehmannii (Rchb.f.) Garay 222. Cryptocentrum standleyi Ames 223. Cuitlauzina convallarioides (Schltr.) Dressler & N.H.Williams 224. Cuitlauzina egertonii (Lindl.) Dressler & N.H.Williams 225. Cyclopogon comosus (Rchb.f.) Burns-Bal. & E.W.Greenw. 226. Cyclopogon elatus (Sw.) Schltr. 227. Cyclopogon fuscofloralis (Szlach.) Carnevali & G.A.Romero 228. Cyclopogon obliquus (J.J. Sm.) Szlach. 229. Cyclopogon olivaceus (Rolfe) Schltr. 230. Cyclopogon prasophylloides (Garay) Szlach. 231. Cyclopogon prasophyllus (Rchb.f.) Schltr. 232. Cycnoches amparoanum Schltr. 233. Cycnoches aureum Lindl. & Paxton 234. Cycnoches warszewiczii Rchb.f. 235. Cyrtochiloides ochmatochila (Rchb.f.) N.H.Williams & M.W.Chase 236. Cyrtochiloides panduriformis (Ames & C.Schweinf.) N.H.Williams & M.W.Chase 237. Cyrtopodium macrobulbon (Lex.) G.A.Romero & Carnevali 238. Daiotyla albicans (Rolfe) Dressler 239. Daiotyla crassa (Dressler) Dressler 240. Daiotyla rhodotyla Pupulin * 241. Dichaea acostae Schltr. 242. Dichaea acroblephara Schltr. 243. Dichaea amparoana Schltr. * 244. Dichaea auriculata Pupulin & Karremans * 245. Dichaea costaricensis Schltr. 246. Dichaea cryptarrhena Rchb.f. ex Kraenzl. 247. Dichaea dammeriana Kraenzl. 248. Dichaea dressleri Folsom 249. Dichaea eburnea (Dressler & Pupulin) Endrés ex Pupulin 250. Dichaea eligulata Folsom 251. Dichaea elliptica Dressler & Folsom 252. Dichaea elvirae Pupulin * 253. Dichaea filiarum Pupulin 254. Dichaea glauca (Sw.) Lindl. 255. Dichaea globosa Dressler & Pupulin 256. Dichaea gomez-lauritoi Pupulin * 257. Dichaea gracillima C.Schweinf. * 258. Dichaea hystricina Rchb.f. 259. Dichaea lankesteri Ames 260. Dichaea morrisii Fawc. & Rendle 261. Dichaea obovatipetala Folsom 262. Dichaea oxyglossa Schltr. 263. Dichaea panamensis Lindl. 264. Dichaea pendula (Aubl.) Cogn. 265. Dichaea poicillantha Schltr. 266. Dichaea sarapiquiensis Folsom * 267. Dichaea similis Schltr. 268. Dichaea squarrosa Lindl. 269. Dichaea trichocarpa (Sw.) Lindl. 270. Dichaea trulla Rchb.f. 271. Dichaea tuerckheimii Schltr. 272. Dichaea viridula Pupulin * 273. Dimerandra emarginata (G.Mey.) Hoehne 274. Diodonopsis erinacea (Rchb.f.) Pridgeon & M.W.Chase 275. Diodonopsis pygmaea (Kraenzl.) Pridgeon & M.W.Chase * 276. Draconanthes aberrans (Schltr.) Luer 277. Dracula astuta (Rchb.f.) Luer * 278. Dracula carlueri Hermans & P.J.Cribb * 279. Dracula erythrochaete (Rchb.f.) Luer 280. Dracula inexperata Pupulin * 281. Dracula maduroi Luer 282. Dracula pusilla (Rolfe) Luer 283. Dracula ripleyana Luer * 284. Dracula vespertilio (Rchb.f.) Luer 285. Dresslerella hispida (L.O.Williams) Luer 286. Dresslerella pilosissima (Schltr.) Luer 287. Dresslerella stellaris Luer & R.Escobar 288. Dressleria dilecta (Rchb.f.) Dodson 289. Dressleria eburnea (Rolfe) Dodson 290. Dressleria helleri Dodson 291. Dryadella fuchsii Luer 292. Dryadella gnoma (Luer) Luer 293. Dryadella greenwoodiana Soto Arenas, Salazar & Solano 294. Dryadella guatemalensis (Schltr.) Luer 295. Dryadella odontostele Luer 296. Dryadella sororcula Luer 297. Echinosepala alexandrae (Schltr.) Pupulin & Bogarín * 298. Echinosepala expolita Pupulin & Belfort * 299. Echinosepala glenioides Pupulin * 300. Echinosepala isthmica Pupulin * 301. Echinosepala lappiformis (A.H.Heller & L.O.Williams) Pridgeon & M.W.Chase 302. Echinosepala longipedunculata Pupulin & Karremans * LANKESTERIANA22 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 303. Echinosepala pan (Luer) Pridgeon & M.W.Chase 304. Echinosepala stonei (Luer) Pridgeon & M.W.Chase * 305. Echinosepala tomentosa (Luer) Pridgeon & M.W.Chase * 306. Echinosepala truncata Pupulin, Álv.-Alcázar & Bogarín * 307. Echinosepala vittata (Pupulin & M.A.Blanco) Luer * 308. Elleanthus aurantiacus (Lindl.) Rchb.f. 309. Elleanthus bradeorum Schltr. 310. Elleanthus caricoides Nash 311. Elleanthus carinatus Dressler & Bogarín * 312. Elleanthus cynarocephalus (Rchb.f.) Rchb.f. 313. Elleanthus glaucophyllus Schltr. 314. Elleanthus graminifolius (Barb.Rodr.) Løjtnant 315. Elleanthus hymenophorus (Rchb.f.) Rchb.f. 316. Elleanthus jimenezii (Schltr.) C.Schweinf. 317. Elleanthus lancifolius C.Presl 318. Elleanthus lentii Barringer 319. Elleanthus ligularis Dressler & Bogarín 320. Elleanthus longibracteatus (Lindl. ex Griseb.) Fawc. 321. Elleanthus muscicola Schltr. 322. Elleanthus poiformis Schltr. 323. Elleanthus stolonifer Barringer 324. Elleanthus tillandsioides Barringer. 325. Elleanthus tonduzii Schltr. * 326. Elleanthus tricallosus Ames & C.Schweinf. 327. Elleanthus wercklei Schltr. * 328. Encyclia alata (Bateman) Schltr. 329. Encyclia ceratistes (Lindl.) Schltr. 330. Encyclia chloroleuca (Hook.) Neumann. 331. Encyclia gravida (Lindl.) Schltr. 332. Encyclia macrochila (Hook.) Neumann 333. Encyclia mooreana (Rolfe) Schltr. 334. Encyclia ossenbachiana Pupulin * 335. Encyclia peraltensis (Ames) Dressler * 336. Encyclia stellata (Lindl.) Schltr. 337. Epidendrum aberrans Schltr. 338. Epidendrum acroamparoanum Hágsater & L.Sánchez * 339. Epidendrum acunae Dressler 340. Epidendrum adnatum Ames & C.Schweinf. 341. Epidendrum aeolicum Karremans * 342. Epidendrum alberti Schltr. 343. Epidendrum alfaroi Ames & C.Schweinf. 344. Epidendrum alfredii Schltr. * 345. Epidendrum alieniferum Karremans & Bogarín * 346. Epidendrum allenii L.O.Williams 347. Epidendrum allochronum Hágsater 348. Epidendrum anastasioi Hágsater * 349. Epidendrum anceps Jacq. 350. Epidendrum angustilobum Fawc. & Rendle 351. Epidendrum angustisegmentum (L.O.Williams) Hágsater 352. Epidendrum anoglossoides Ames & C.Schweinf. 353. Epidendrum anoglossum Schltr. 354. Epidendrum apatotylosum Hágsater * 355. Epidendrum arcuiflorum Ames & C.Schweinf. 356. Epidendrum astroselaginella Hágsater & E.Santiago 357. Epidendrum atrorugosum Hágsater 358. Epidendrum atwoodchlamys Hágsater * 359. Epidendrum atwoodii Hágsater & L.Sánchez * 360. Epidendrum atypicum Hágsater & E.Santiago * 361. Epidendrum barbae Rchb.f. 362. Epidendrum barbeyanum Kraenzl. 363. Epidendrum baumannianum Schltr. 364. Epidendrum belloi Hágsater 365. Epidendrum bicentenarium Karremans & Gil-Amaya * 366. Epidendrum bicuniculatum Hágsater & E.Santiago * 367. Epidendrum bilobatum Ames * 368. Epidendrum bisulcatum Ames 369. Epidendrum blepharistes Barker ex Lindl. 370. Epidendrum bogarinii Karremans & C.M.Sm. * 371. Epidendrum brachyclinium Hágsater & García- Cruz * 372. Epidendrum brachyrepens Hágsater 373. Epidendrum bracteostigma Hágsater & García-Cruz * 374. Epidendrum bracteosum Ames & C.Schweinf. * 375. Epidendrum brenesii Schltr. * 376. Epidendrum buenaventurae F.Lehm. & Kraenzl. 377. Epidendrum bugabense Hágsater 378. Epidendrum campbellstigma Hágsater & García- Cruz * 379. Epidendrum candelabrum Hágsater 380. Epidendrum capitalinum Karremans * 381. Epidendrum cardiochilum L.O.Williams 382. Epidendrum cardiophorum Schltr. 383. Epidendrum caroli Schltr. 384. Epidendrum centropetalum Rchb.f. 385. Epidendrum chalcochromum Hágsater * 386. Epidendrum chespiritorum Karremans 387. Epidendrum chirripoense Hágsater 388. Epidendrum chlorocorymbos Schltr. 389. Epidendrum ciliare L. LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 23PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae 390. Epidendrum circinatum Ames 391. Epidendrum cocoense Hágsater * 392. Epidendrum colibri Karremans * 393. Epidendrum concavilabium C.Schweinf. * 394. Epidendrum confertum Ames & C.Schweinf. 395. Epidendrum congestoides Ames & C.Schweinf. 396. Epidendrum congestum Rolfe 397. Epidendrum conservatorum Karremans * 398. Epidendrum cordiforme C.Schweinf. 399. Epidendrum coriifolium Lindl. 400. Epidendrum coronatum Ruiz & Pav. 401. Epidendrum crescentilobum Ames 402. Epidendrum criniferum Rchb.f. 403. Epidendrum cristatum Ruiz & Pav. 404. Epidendrum cryptanthum L.O.Williams 405. Epidendrum cymbiglossum Hágsater 406. Epidendrum cystosum Ames 407. Epidendrum davidsei Hágsater * 408. Epidendrum dentiferum Ames & C.Schweinf. 409. Epidendrum dolichochlamys Hágsater & E.Santiago * 410. Epidendrum eburneum Rchb.f. 411. Epidendrum elcimeyae Hágsater & García-Cruz * 412. Epidendrum endresii Rchb.f. 413. Epidendrum epidendroides (Garay) Mora-Ret. & García Castro * 414. Epidendrum erythrostigma Hágsater 415. Epidendrum estrellense Ames 416. Epidendrum exasperatum Rchb.f. 417. Epidendrum exiguum Ames & C.Schweinf. 418. Epidendrum firmum Rchb.f. 419. Epidendrum flexicaule Schltr. 420. Epidendrum flexuosum G.Mey. 421. Epidendrum fulfordianum Pupulin & Karremans * 422. Epidendrum gibbosum L.O.Williams 423. Epidendrum globuliferum Karremans * 424. Epidendrum glumarum Hamer & Garay 425. Epidendrum goniorhachis Schltr. * 426. Epidendrum grayumii Hágsater & E.Santiago * 427. Epidendrum guanacastense Ames & C.Schweinf. 428. Epidendrum gymnochlamys Hágsater & E.Santiago 429. Epidendrum haberi Hágsater & L.Sánchez * 430. Epidendrum hameri Hágsater & L.Sánchez 431. Epidendrum hammelii Hágsater & L.Sánchez 432. Epidendrum hartmanniorum Karremans & M.Díaz * 433. Epidendrum hawkesii A.H.Heller 434. Epidendrum hellerianum A.D.Hawkes 435. Epidendrum horichii Hágsater * 436. Epidendrum hunterianum Schltr. 437. Epidendrum incomptum Rchb.f. 438. Epidendrum ingramii Hágsater & García-Cruz * 439. Epidendrum inhibitiosum Karremans & Mel. Fernández * 440. Epidendrum insulanum Schltr. 441. Epidendrum intermixtum Ames & C.Schweinf. 442. Epidendrum iricolor Karremans * 443. Epidendrum isomerum Schltr 444. Epidendrum isthmi Schltr. 445. Epidendrum isthmoides Hágsater & E.Santiago * 446. Epidendrum jimenezii Hágsater 447. Epidendrum jorge-warneri Karremans & Hágsater 448. Epidendrum lacustre Lindl. 449. Epidendrum lagenocolumna Hágsater & L.Sánchez 450. Epidendrum lancilabium Schltr. 451. Epidendrum lankesteri Ames * 452. Epidendrum laterale Rolfe 453. Epidendrum laucheanum Bonhof ex Rolfe 454. Epidendrum lockhartioides Schltr. 455. Epidendrum longibracteatum Hágsater 456. Epidendrum macroclinium Hágsater 457. Epidendrum maduroi Hágsater & García-Cruz 458. Epidendrum magnibracteatum Ames 459. Epidendrum manta-birostris Karremans * 460. Epidendrum microanoglossum Hágsater & Karremans 461. Epidendrum microcardium Schltr. 462. Epidendrum microdendron Rchb.f. 463. Epidendrum microrigidiflorum Hágsater 464. Epidendrum mirabile Ames & C.Schweinf. * 465. Epidendrum miriamiae Karremans * 466. Epidendrum miserrimum Rchb.f. 467. Epidendrum modestiflorum Schltr. 468. Epidendrum monophlebium Hágsater * 469. Epidendrum × monteverdense (Pupulin & Hágsater) Hágsater * 470. Epidendrum montis-narae Pupulin & L.Sánchez 471. Epidendrum mora-retanae Hágsater 472. Epidendrum muricatisepalum Hágsater * 473. Epidendrum muscicola Schltr. 474. Epidendrum myodes Rchb.f. 475. Epidendrum nativitatis Karremans * 476. Epidendrum nervosiflorum Ames & C.Schweinf. * 477. Epidendrum × nocteburneum Hágsater & L.Sánchez 478. Epidendrum nocturnum Jacq. 479. Epidendrum notabile Schltr. LANKESTERIANA24 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 480. Epidendrum nutantirhachis Ames & C.Schweinf. 481. Epidendrum obliquifolium Ames, F.T.Hubb. & C.Schweinf. * 482. Epidendrum octomerioides Schltr. 483. Epidendrum odontochilum Hágsater 484. Epidendrum oerstedii Rchb.f. 485. Epidendrum orthodontum Hágsater & L.Sánchez, 486. Epidendrum oxyglossum Schltr. 487. Epidendrum pachyceras Hágsater & L.Sánchez 488. Epidendrum pachyrachis Ames 489. Epidendrum pachytepalum Hágsater & E.Santiago 490. Epidendrum pallens Rchb.f. 491. Epidendrum palmidium Hágsater * 492. Epidendrum pansamalae Schltr. 493. Epidendrum paradisicola Hágsater & García-Cruz * 494. Epidendrum paraguastigma Hágsater & García-Cruz 495. Epidendrum paranthicum Rchb.f. 496. Epidendrum parkinsonianum Hook. 497. Epidendrum parviexasperatum (Hágsater) Hágsater 498. Epidendrum paucifolium Schltr. 499. Epidendrum pendens L.O.Williams 500. Epidendrum penneystigma Hágsater & García-Cruz * 501. Epidendrum pentadactylum Rchb.f. * 502. Epidendrum pergameneum Rchb.f. 503. Epidendrum pfavii Rolfe 504. Epidendrum philowercklei Hágsater & E.Santiago 505. Epidendrum phragmites A.H.Heller & L.O.Williams 506. Epidendrum phyllocharis Rchb.f. 507. Epidendrum physodes Rchb.f. 508. Epidendrum piliferum Rchb.f. 509. Epidendrum pinniferum C.Schweinf. 510. Epidendrum platystigma Rchb.f. 511. Epidendrum platystomoides Hágsater & L.Sánchez * 512. Epidendrum polychlamys Schltr. 513. Epidendrum porpax Rchb.f. 514. Epidendrum powellii Schltr. 515. Epidendrum pseudepidendrum Rchb.f. 516. Epidendrum pseudobarbeyanum Pupulin & Karremans * 517. Epidendrum pseudoramosum Schltr. 518. Epidendrum pterocaulum Hágsater & E.Santiago * 519. Epidendrum pumilum Rolfe 520. Epidendrum purpurascens Focke 521. Epidendrum radicans Pav. ex Lindl. 522. Epidendrum rafael-lucasii Hágsater 523. Epidendrum ramonianum Schltr. * 524. Epidendrum ramosissimum Ames & C.Schweinf. * 525. Epidendrum ramosum Jacq. 526. Epidendrum repens Cogn. 527. Epidendrum resectum Rchb.f. 528. Epidendrum rigidiflorum Schltr. 529. Epidendrum rossanae Karremans * 530. Epidendrum rousseauae Schltr. 531. Epidendrum rudolfii Karremans 532. Epidendrum rugosum Ames 533. Epidendrum salpichlamys Hágsater & E.Santiago * 534. Epidendrum saltatrix Karremans * 535. Epidendrum sanchoi Ames 536. Epidendrum sandiorum Hágsater, Karremans & L.Sánchez * 537. Epidendrum santaclarense Ames 538. Epidendrum santaelenae Karremans * 539. Epidendrum scandens Karremans, Chinchilla & Rojas-Alv. * 540. Epidendrum scharfii Hágsater & Dodson 541. Epidendrum schumannianum Schltr. * 542. Epidendrum sculptum Rchb.f 543. Epidendrum selaginella Schltr. 544. Epidendrum serruliferum Schltr. 545. Epidendrum sidereum Karremans * 546. Epidendrum sigmoideum Hágsater * 547. Epidendrum sinac Karremans * 548. Epidendrum sotoanum Karremans & Hágsater 549. Epidendrum spiculiferum Karremans & Bogarín * 550. Epidendrum stamfordianum Bateman 551. Epidendrum stangeanum Rchb.f. 552. Epidendrum stellidifforme Hágsater & Dodson 553. Epidendrum stenoselaginella Hágsater & E.Santiago 554. Epidendrum stevensii Hágsater * 555. Epidendrum stolidium Hágsater 556. Epidendrum storkii Ames 557. Epidendrum strobiliferum Rchb.f. 558. Epidendrum subnutans Ames & C.Schweinf. 559. Epidendrum summerhayesii Hágsater 560. Epidendrum synchronum Hágsater * 561. Epidendrum talamancanum (J.T.Atwood) Mora- Ret. & García Castro 562. Epidendrum tetraceros Rchb.f. 563. Epidendrum thurstoniorum Hágsater * 564. Epidendrum tolimense Lindl. 565. Epidendrum trachythece Schltr LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 25PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae 566. Epidendrum trialatum Hágsater 567. Epidendrum triangulabium Ames & C.Schweinf. 568. Epidendrum trianthum Schltr. 569. Epidendrum tritropianthum Hágsater & E.Santiago 570. Epidendrum turialvae Rchb.f. 571. Epidendrum usurpator Karremans * 572. Epidendrum utyumii Karremans * 573. Epidendrum vallis-silentii M.Díaz & Karremans * 574. Epidendrum vexillium Hágsater 575. Epidendrum villegastigma Hágsater & García-Cruz * 576. Epidendrum villotae Hágsater & Dodson 577. Epidendrum vincentinum Lindl. 578. Epidendrum volutum Lindl. & Paxton 579. Epidendrum vulcanicola A.H.Heller 580. Epidendrum vulgoamparoanum Hágsater & L.Sánchez 581. Epidendrum wallisii Rchb.f. 582. Epidendrum wercklei Schltr. 583. Epidendrum zunigae Hágsater, Karremans & Bogarín * 584. Epistephium ellipticum R.O.Williams & Summerh. 585. Eriopsis wercklei Schltr. 586. Erycina crista-galli (Rchb.f.) N.H.Williams & M.W.Chase 587. Erycina pumilio (Rchb.f.) N.H.Williams & M.W.Chase 588. Erycina pusilla (L.) N.H.Williams & M.W.Chase 589. Eulophia alta (L.) Fawc. & Rendle 590. Eurystyles auriculata Schltr. 591. Eurystyles cornu-bovis Szlach. 592. Eurystyles standleyi Ames 593. Eurystyles uxoris Bogarín * 594. Fernandezia costaricensis (Ames & C.Schweinf.) M.W.Chase * 595. Fernandezia crystallina (Lindl.) M.W.Chase 596. Fernandezia hispidula (Rchb.f.) M.W.Chase 597. Fernandezia tica Mora-Ret. & García Castro 598. Funkiella valerioi (Ames & C.Schweinf.) Salazar & Soto Arenas 599. Funkiella versiformis Szlach. * 600. Galeandra arundinis Garay & G.A.Romero 601. Galeandra beyrichii Rchb.f. 602. Galeottia grandiflora A.Rich. 603. Gomphichis adnata (Ridl.) Schltr. 604. Gongora amparoana Schltr. * 605. Gongora armeniaca (Lindl.) Rchb.f. subsp. armeniaca Gongora armeniaca subsp. cornuta (Klotzsch) Stein 606. Gongora batemanii Henshall 607. Gongora boracayanensis Jenny * 608. Gongora claviodora Dressler 609. Gongora gibba Dressler 610. Gongora horichiana Fowlie 611. Gongora leucochila Lem. 612. Gongora unicolor Schltr. 613. Goodyera bradeorum Schltr. 614. Goodyera erosa (Ames & C.Schweinf.) Ames 615. Goodyera major Ames & Correll 616. Goodyera micrantha Schltr. * 617. Goodyera modesta Schltr. * 618. Goodyera ovatilabia Schltr. 619. Goodyera turialbae Schltr. * 620. Govenia ciliilabia Ames & C.Schweinf. 621. Govenia lagenophora Lindl. 622. Govenia liliacea (Lex.) Lindl. 623. Govenia quadriplicata Rchb.f. 624. Govenia viaria Dressler 625. Greenwoodiella wercklei (Schltr.) Salazar & R.Jiménez 626. Guarianthe aurantiaca (Bateman ex Lindl.) Dressler & W.E.Higgins 627. Guarianthe hennisiana (Rolfe) Van den Berg 628. Guarianthe skinneri (Bateman) Dressler & W.E.Higgins f. skinneri Guarianthe skinneri f. alba (Rchb.f.) Christenson 629. Habenaria alata Hook. 630. Habenaria aviculoides Ames & C.Schweinf. 631. Habenaria carlotae Dressler * 632. Habenaria costaricensis Schltr. 633. Habenaria distans Griseb. 634. Habenaria eustachya Rchb.f. 635. Habenaria heptadactyla Rchb.f. 636. Habenaria irazuensis Schltr. * 637. Habenaria lankesteri Ames 638. Habenaria macroceratitis Willd. 639. Habenaria mesodactyla Griseb. 640. Habenaria monorrhiza (Sw.) Rchb.f. 641. Habenaria pringlei B.L.Rob. 642. Habenaria repens Nutt. 643. Habenaria rodeiensis Barb.Rodr. 644. Habenaria strictissima Rchb.f. 645. Habenaria trifida Kunth 646. Habenaria wercklei Schltr. LANKESTERIANA26 LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 647. Hapalorchis pumila (C.Schweinf.) Garay 648. Heterotaxis maleolens (Schltr.) Ojeda & Carnevali 649. Heterotaxis sessilis (Sw.) F.Barros 650. Heterotaxis valenzuelana (A.Rich.) Ojeda & Carnevali 651. Homalopetalum pumilio (Rchb.f.) Schltr. 652. Horichia dressleri Jenny 653. Houlletia tigrina Linden ex Lindl. 654. Huntleya burtii (Endrés & Rchb.f.) Pfitzer 655. Inti bicallosa (Rchb.f.) M.A.Blanco 656. Inti chartacifolia (Ames & C.Schweinf.) M.A.Blanco 657. Ionopsis satyrioides (Sw.) Rchb.f. 658. Ionopsis utricularioides (Sw.) Lindl 659. Isochilus carnosiflorus Lindl. 660. Isochilus chiriquensis Schltr. 661. Isochilus latibracteatus A.Rich. & Galeotti 662. Isochilus linearis (Jacq.) R.Br. 663. Jacquiniella aporophylla (L.O.Williams) Dressler 664. Jacquiniella equitantifolia (Ames) Dressler 665. Jacquiniella globosa (Jacq.) Schltr. 666. Jacquiniella standleyi (Ames) Dressler 667. Jacquiniella teretifolia (Sw.) Britton & P.Wilson 668. Karma arbuscula (Lindl.) Karremans 669. Karma carinilabia (Luer) Karremans 670. Karma cedralensis (Ames) Karremans 671. Karma dura (Lindl.) Karremans 672. Karma fruticosa (Luer) Karremans 673. Karma membraniflora (C.Schweinf.) Karremans 674. Karma nana (Ames & C.Schweinf.) Karremans * 675. Karma parsonsii (Luer & Dod) Karremans * 676. Kefersteinia alata Pupulin 677. Kefersteinia alba Schltr. 678. Kefersteinia costaricensis Schltr. 679. Kefersteinia endresii Pupulin * 680. Kefersteinia excentrica Dressler & Mora-Ret. 681. Kefersteinia lactea (Rchb.f.) Schltr. 682. Kefersteinia microcharis Schltr. * 683. Kefersteinia orbicularis Pupulin 684. Kefersteinia parvilabris Schltr. 685. Kefersteinia retanae G.Gerlach * 686. Kefersteinia saccata Pupulin * 687. Kefersteinia wercklei Schltr. * 688. Kegeliella atropilosa L.O.Williams & A.H.Heller 689. Kegeliella kupperi Mansf. 690. Lacaena spectabilis (Klotzsch) Rchb.f. 691. Laelia lueddemanii (Prill.) L.O.Williams Laelia lueddemanii (Prill.) L.O.Williams subsp. costaricana (Rchb.f.) Pupulin 692. Laelia rubescens Lindl. 693. Lankesterella orthantha (Kraenzl.) Garay 694. Lankesteriana barbulata (Lindl.) Karremans 695. Lankesteriana casualis (Ames) Karremans 696. Lankesteriana caudatipetala (C.Schweinf.) Karremans 697. Lankesteriana cuspidata (Luer) Karremans 698. Lankesteriana duplooyi (Luer & Sayers) Karremans 699. Lankesteriana fractiflexa (Ames & C.Schweinf.) Karremans 700. Lankesteriana haberi (Luer) Karremans * 701. Leochilus candelariae (Kraenzl.) Pupulin * 702. Leochilus labiatus (Sw.) Kuntze 703. Leochilus leochilinus (Rchb.f.) M.W.Chase & N.H.Williams 704. Leochilus scriptus (Scheidw.) Rchb.f. 705. Leochilus tricuspidatus (Rchb.f.) Kraenzl. 706. Lepanthes acarina Luer 707. Lepanthes acoridilabia Ames & C.Schweinf. 708. Lepanthes adenophora Bogarín, M.Cedeño & Chinchilla * 709. Lepanthes amicitiae Bogarín & Pupulin 710. Lepanthes amplectens Luer & Hermans * 711. Lepanthes ankistra Luer & Dressler 712. Lepanthes antilocapra Luer & Dressler 713. Lepanthes arenasiana Bogarín & Mel.Fernández * 714. Lepanthes atrata Endrés ex Luer * 715. Lepanthes atwoodii Luer * 716. Lepanthes barbae Schltr. * 717. Lepanthes barbosae Luer * 718. Lepanthes blephariglossa Schltr. 719. Lepanthes blepharistes Rchb.f. 720. Lepanthes bogariniana Pupulin * 721. Lepanthes bradei Schltr. * 722. Lepanthes candida Endrés ex Luer 723. Lepanthes caroli-lueri Bogarín & Pupulin * 724. Lepanthes casasae Pupulin * 725. Lepanthes cascajalensis Ames 726. Lepanthes castilloae Bogarín & Kisel * 727. Lepanthes chameleon Ames * 728. Lepanthes chiriquensis Schltr. 729. Lepanthes ciliisepala Schltr. * 730. Lepanthes circularis Luer * 731. Lepanthes clareae Luer & Hermans * LANKESTERIANA 23(Supplement). 2023. © Universidad de Costa Rica, 2023. 27PuPulin, Bogarín & Karremans — Catalogue of Costa Rican Orchidaceae 732. Lepanthes coeloglossa Luer 733. Lepanthes comet-halleyi Luer * 734. Lepanthes confusa Ames & C.Schweinf. 735. Lepanthes costaricensis Schltr. 736. Lepanthes cribbii Pupulin * 737. Lepanthes crucitasensis Chinchilla, M.Cedeño & Bogarín * 738. Lepanthes cuspidata Luer * 739. Lepanthes daniel-jimenezii Bogarín & Pupulin * 740. Lepanthes davidsei Luer * 741. Lepanthes decipiens Ames & C.Schweinf. * 742. Lepanthes deformis Luer & Hirtz 743. Lepanthes demissa Luer 744. Lepanthes dichroma Luer * 745. Lepanthes dikoensis Bogarín & C.M.Sm. * 746. Lepanthes disticha (A.Rich. & Galeotti) Garay & R.E.Schult. 747. Lepanthes dolabriformis Luer * 748. Lepanthes dotae Endrés ex Luer * 749. Lepanthes droseroides Luer 750. Lepanthes durikäensis Pupulin & Bogarín * 751. Lepanthes edentula Luer 752. Lepanthes edwardsii Ames 753. Lepanthes elata Rchb.f. 754. Lepanthes elegans Luer 755. Lepanthes elusiva Bogarín, Kaes & M.Díaz * 756. Lepanthes elvirae Pupulin * 757. Lepanthes empticia Luer & Béhar * 758. Lepanthes erinacea Rchb.f. * 759. Lepanthes erubescens Bogarín, Pupulin & Karremans * 760. Lepanthes estrellensis Ames * 761. Lepanthes excedens Ames & Correll 762. Lepanthes eximia Ames. 763. Lepanthes expansilabia Bogarín & C.M.Sm. * 764. Lepanthes falcifera Luer * 765. Lepanthes falx-bellica Pupulin & Bogarín * 766. Lepanthes fascinata Luer * 767. Lepanthes ferrelliae Luer * 768. Lepanthes fimbriata Ames 769. Lepanthes forcipifera Luer * 770. Lepanthes fugiens Luer * 771. Lepanthes garciae Bogarín & D.Jiménez * 772. Lepanthes genetoapophantica Bogarín & Gravend. 773. Lepanthes gerardensis M.A.Blanco * 774. Lepanthes glicensteinii Luer * 775. Lepanthes gracillima Endrés ex Luer 776. Lepanthes grandiflora Ames & C.Schweinf. 777. Lepanthes gratiosa Pupulin & D.Jiménez * 778. Lepanthes guanacastensis Ames & C.Schweinf. 779. Lepanthes guardiana Endrés ex Luer * 780. Lepanthes hamulifera Luer * 781. Lepanthes helleri A.D.Hawkes 782. Lepanthes hermansii Luer * 783. Lepanthes horichii Luer * 784. Lepanthes horrida Rchb.f. 785. Lepanthes inaequiloba Ames & C.Schweinf. * 786. Lepanthes inescata Luer * 787. Lepanthes ingramii Luer * 788. Lepanthes inornata Schltr. * 789. Lepanthes jimenezii Schltr. 790. Lepanthes jugum Luer 791. Lepanthes kabebatae Bogarín, Karremans & Mel. Fernández * 792. Lepanthes kleinii Bogarín & Pupulin * 793. Lepanthes lancifolia Schltr. * 794. Lepanthes lappacea Luer * 795. Lepanthes latisepala Ames & C.Schweinf. * 796. Lepanthes leporina Pupulin & Bogarín * 797. Lepanthes limbellata Endrés ex Luer * 798. Lepanthes lindleyana Oerst. & Rchb.f. 799. Lepanthes luerorum B.T.Larsen 800. Lepanthes macalpinii Luer * 801. Lepanthes machogaffensis Pupulin & D.Jiménez * 802. Lepanthes mariposa Luer 803. Lepanthes marta-herrae Bogarín & Pupu