Webbia. Journal of Plant Taxonomy and Geography 77(1): 35-126, 2022 Firenze University Press www.fupress.com/webbia ISSN 0083-7792 (print) | ISSN 2169-4060 (online) | DOI: 10.36253/jopt-12092 W EB B IA Journal of Plant Taxonomy and Geography Citation: Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossen- bach, Diego Bogarín (2022) Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum. Webbia. Journal of Plant Taxonomy and Geography 77(1): 35-126. doi: 10.36253/jopt-12092 Received: September 14, 2021 Accepted: November 11, 2021 Published: April 20, 2022 Copyright: © 2022 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossen- bach, Diego Bogarín. This is an open access, peer-reviewed article published by Firenze University Press (http:// www.fupress.com/webbia) and distrib- uted under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, pro- vided the original author and source are credited. Data Availability Statement: All rel- evant data are within the paper and its Supporting Information files. Competing Interests: The Author(s) declare(s) no conflict of interest. Editor: Riccardo M. Baldini ORCID FP: https://orcid.org/0000-0001-6780- 8395 IFC: https://orcid.org/0000-0002-4250- 1642 GRA: https://orcid.org/0000-0002-8918- 0823 MF: https://orcid.org/0000-0001-6435- 4631 DB: https://orcid.org/0000-0002-8408- 8841 Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Franco Pupulin1,2,3,*, Isler F. Chinchilla1,4, Gustavo Rojas-Alvarado1, Melania Fernández1,5, Carlos Ossenbach1,6, Diego Bogarín1,7.8 1 Lankester Botanical Garden, University of Costa Rica. P.O. Box 302-7050 Cartago, Cos- ta Rica 2 Harvard University Herbaria, Cambridge, Massachusetts 02138, USA 3 The Marie Selby Botanical Gardens, Sarasota, Florida 34236, USA 4 Luis Fournier Origgi Herbarium, Universidad of Costa Rica, Apdo 11501-2060, San José, Costa Rica 5 Department of Plant & Soil Science, Texas Tech University, Lubbock, Texas 79409, U.S.A. 6 Orquideario 25 de mayo, Sabanilla de Montes de Oca, Costa Rica 7 Herbario UCH, Universidad Autónoma de Chiriquí, 0427, David, Chiriquí, Panamá 8 Naturalis Biodiversity Center, Evolutionary Ecology Group, 2333 CR Leiden, The Neth- erlands *Corresponding author. E-mail: franco.pupulin@ucr.ac.cr Abstract. The typification of 53 orchid species described by Rudolf Schlechter based on specimens gathered in Costa Rica by Paul (Pablo) Biolley, Juan José Cooper San- doval, Auguste R. Endrés, Carl Hoffmann, Emel Jiménez Segura, Otón Jiménez, Frie- drich Carl Lehmann, Ferdinand Nevermann, Richard Pfau, and Henry Pittier in the late 19th and early 20th centuries is discussed, and lectotypes are designated when necessary. Short biographical notes are provided for the main collectors whose Costa Rican orchid gatherings are presented here. Taxonomical and historical backgrounds are presented for the concerned taxa, and the rationale for their typifications is dis- cussed. Lectotypes are proposed for Epidendrum dolichostachyum, E. selaginella, Habe- naria jimenezii, Hexadesmia jimenezii, Masdevallia reflexa, Microstylis carpinterae, Not- ylia pittieri, Oncidium cabagrae, O. costaricense, Ornithidium biolleyi, Ornithocephalus xiphochilus, Physurus lehmannii, Platystele bulbinella, Pleurothallis pittieri, P. sororia, Sauroglossum nigricans, Scaphyglottis pauciflora, S. subulata, Sobralia pfavii, Solenocen- trum costaricense, Stelis coiloglossa, S. cooperi, S. cyclopetala, S. despectans, and S. ton- duziana. An epitype is designated for Gongora unicolor. Keywords: epitypification, flora of Costa Rica, history of botany, lectotypification, nomenclature, Orchidaceae. INTRODUCTION Rudolf Schlechter (1872–1925) (Figure 1) was arguably the most pro- ficient orchid taxonomist of the 20th century. With over 5,000 orchid taxa 36 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín described before his premature death, he proposed the largest number of new orchid genera and species among his contemporaries and gave birth to monographic revi- sions of genera and subtribes, as well as national and regional orchid floras. His interest in giving shape to orchid diversity spanned the entire world’s tropical flo- ras, from Africa to New Guinea, from Indonesia to South America, from Madagascar to China, from Cen- tral America to Japan, from the West Indies to Australia. In 1914 at the age of 42, and many years before ending his botanical activity, he produced an “encyclopedia” of the Orchidaceae, with notes on taxonomy and culture, under the title Orchideen, ihre Beschreibung, Kultur und Züchtung; Handbuch für Orchideenliebhaber, Züchter und Botaniker (“Orchids, their description, culture and breeding; manual for orchid lovers, breeders and bota- nists”, Schlechter 1914), a work that Senghas (2002) con- sidered the crowning moment of his career. From 1899, when he published his first orchid spe- cies from Guatemala and Mexico, based on plants col- lected by Georg Eduard Seler (1849–1922) and his wife Caecilie Seler-Sachs (1855–1933) and received for iden- tification at the Botanical Museum of Berlin-Dahlem (Schlechter 1899), he devoted a considerable part of his work to the study of the Orchidaceae from the Ameri- can isthmus (for a geographic definition of the region, see discussion in Ossenbach et al. 2007). In the next 25 years, he proposed new genera and species of orchids from Guatemala (Schlechter 1906a, 1906c, 1916, 1918a, 1920, 1921b, 1925), Mexico (Schlechter 1906c, 1914, 1915, 1916, 1918b, 1918c, 1925), Costa Rica (1906a, 1907a, 1907b, 1913a, 1920, 1921a, 1921b, 1923a, 1923b, 1923c, 1923d), Panama (Schlechter 1913a, 1921b, 1922), El Sal- vador (1913b), and Honduras (Schlechter 1918a). During the 1910’s and 1920’s, Schlechter was particularly fond of the orchid flora from Mesoamerica, a subject on which he maintained for a long time a fair academic competi- tion with his North American colleague, Prof. Oakes Ames (1874–1950) of Harvard University, who in that same period also devoted himself to a fervent study of the orchid flora of the American isthmus. It was Costa Rica, however, that truly represented that orchid “El Dorado” (Schlechter 1923c) that he need- ed to complete his ambitious project of describing a new species of orchid every day of his life (Reinikka 1995). Eventually, he came to describe from the small Central American republic almost four hundred taxa new to sci- ence, including 23 new genera, 382 new species, and five subspecific taxa. Without doubt, a combination of various factors contributed to this prodigious result. The position of Costa Rica in the central portion of the isthmus between two continents, in an area small enough to be affected by the climatic effects of both oceans, but large enough to host a complex system of mountain ranges of different origins that form a defined continental spine, is reflected in a particular number of different life zones and favors the maintenance of an extraordinarily diverse f lora. In terms of orchid diversity, Costa Rica has the high- est index in the American continent and possibly the highest globally (Karremans and Bogarín 2013), and the recent biogeographical assessment by Crain and Fernán- dez (2020) indicated the unique attributes underpinning diversity patterns and the occurrence of orchid hotspots. Furthermore, during the last decade of 19th centu- ry, Costa Rica saw the birth of a national science as the direct result of the educational reform inaugurated by President Bernardo Soto (1885–1889), who hired a group of European academics to staff the two new public high schools in the capital (Ossenbach 2009) (Figure 2). The Figure 1. Friedrich Wilhelm Rudolf Schlechter (1872–1925) in the Herbarium of the Botanical Museum in Berlin, 1909. Archives of Rudolf Jenny and courtesy of Dr. N. Kilian, Archives BGBM Berlin- Dahlem. 37Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum foundation of the National Museum in 1887 and the Instituto Físico-Geográfico in 1889 symbolized this sci- entific renaissance. With active botanical institutions and enthusiastic young botanists, early 20th century Costa Rica was in the perfect situation to begin the systematic exploration of its natural resources, and orchids were no exception. The work carried out by the staff of the Museo Nacional, with figures such as the Swiss Henri Francois Pittier (1857-1950), Paul Biolley (1861-1908), and Adolphe Tonduz (1862–1921), the Alsatian Karl Wercklé (1860– 1924) and the German brothers Alfred Brade (1867–1955) and Alexander Curt Brade (1881–1971), as well as those of national scholars such as Alberto M. Brenes (1870– 1948) and Otón Jiménez Luthmer (1895–1988) among others, had no equal in other Central American coun- tries (Standley 1937; Barringer 1986; Pupulin 2010a; Pupulin et al. 2016; Bogarín et al. in prep). Finally, it is worth mentioning the personal interest shown by the Cuban Amparo López-Calleja (1870-1951), wife of the notable Costa Rican ornithologist José Cás- tulo Zeledón (1846–1923), for the flora of her adoptive country, and in particular for orchids, which she culti- vated in her large garden in San José. Doña Amparo de Zeledón, as she was respectfully called, supported with her funds many of the field activities carried out by Ton- duz and Wercklé (who together collected almost 15,000 specimens of plants for the National Museum) (Ossen- bach 2009). Schlechter requested that she expressly arrange for Tonduz to press plants from her orchid gar- den and send out Wercklé on new collecting excursions, resulting in three shipments of orchid exsiccata sent to Schlechter between 1921 and 1923. Schlechter oppor- tunely acknowledged her commitment to creating in her honor the genus Amparoa Schltr. (= Rhynchostele Rchb.f.), baptizing several orchid species for her name, and dedicating to Doña Amparo a large chapter of his Beiträge zur Orchideenkunde von Zentralamerika, II. Additamenta ad Orchideologiam Costaricensem, under the title Orchidaceae Amparoanae (Schlechter 1923a). However, Schlechter’s love affair with the orchids of Costa Rica did not depend exclusively on the plants he received for identification from the National Museum, those provided through the interest of Doña Amparo, or the two later mailings by Guillermo Acosta (Schlech- ter 1923d). He maintained an active collaboration with the Boissier Herbarium, where the orchids that Adolphe Tonduz sent, alive from Costa Rica and subsequently cultivated in the Barbey-Boissier greenhouse Rivage (on the shores of Lake Geneva), were pressed (Pupulin et al. 2016). Furthermore, he visited the famous herbarium of Heinrich Gustav Reichenbach (1823–1889) in Vien- na shortly after it was made available again for study, where he studied the early Costa Rican collections by Karl Hoffmann Brehmer (1823–1859), August R. Endrés (1838–1874) Richard Pfau (1856–1897), and Friedrich G. Lehmann (1850–1903), among others, from which he eventually described several new orchid species. The interpretation of the outstanding work carried out by Schlechter on the orchid flora of Costa Rica has been greatly hampered by the fire of the herbarium at the Botanical Museum of Berlin during an Allied bomb- ing raid in 1943 (Ames 1944, Hiepko 1987) because most of the orchid types, together with Schlechter’s analytical sketches, were destroyed (Figure 3). Only those speci- mens that were moved to the Museum’s basements or were on loan to other institutions escaped the fire. Even though some of the orchid types from other regions of the world survived (Butzin 1978), all the type specimens of Orchidaceae from the Neotropics, including the spe- Figure 2. A, Building of the Colegio Superior de Señoritas for girls in 1909. By Vistas de Costa Rica. B, Building of the Liceo de Cos- ta Rica for boys in 1922. By Manuel Gómez Miralles, Documental Patrimonio Arquitectónico. 38 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín cies described by Schlechter, Fritz Kränzlin (1847–1934), and Rudolf Mansfeld (1901–1960) were lost (Butzin 1980). The only known exceptions are a syntype of Spi- ranthes goodyeroides Schltr. from Bolivia (Butzin 1978) and an isotype of Quekettia australis Kraenzl., collected in Brazil (Butzin 1980). It was undoubtedly a fortunate circumstance that the National Museum of Costa Rica kept duplicates of many of the collections made by its scientific staff, which were sent to Schlechter for determination. The German botanist used them as a basis for his descriptions of new Costa Rican orchid species. Many of these isotypes have served to typify the names of Schlechter’s orchids, whose original types have been lost (Barringer 1986; Lobo 2004; Pupulin 2010; Pupulin et al. 2016). But even more providential was the fact that during the two decades during which Schlechter devoted himself to describ- ing the orchids that came to him from his correspond- ents in Costa Rica, his colleague Oakes Ames (Figure 4), who had already developed a reputation of his own in orchidology working on the floras of Malaysia, Indone- sia, and the Philippine, directed his attention, with par- ticular emphasis, to the orchids of Mesoamerica. Ames (1908a) described his first orchids of the Central Ameri- can isthmus only two years after Schlechter, with his second Decas of new and critical orchids from Guatema- la (Schlechter 1906c), had begun his prolific series of sci- entific works on the Mesoamerican orchidaceous flora, which would have ended only at his death in 1925. Ames survived Schlechter and continued his work of eluci- dating Central American orchid flora, albeit with less emphasis, until the mid-1930s. The intense relationship between the two taxonomists was explored by Ossen- bach (2009), who highlighted their scientific rivalries and the deeply human aspects of solidarity and friend- ship that bound them. Not only did Ames contribute financially to the publication of part of Schlechter’s work on the orchid flora of the Andean countries after the German botanist had run out of funds for the remain- ing volumes, but he also supported Schlechter’s wife in paying the hospital bills during her husband’s illness (Ossenbach 2009). The amount of first-hand information that Schlechter had accumulated in his herbarium dur- ing the years of his relationships with the botanists of the Museo Nacional and the group sponsored by Doña Amparo de Zeledón was so fundamental to the under- standing of the rich flora of Costa Rica that Ames had several artists at once employed in Berlin to copy (under Schlechter’s supervision) the analytical sketches of new species made by the German taxonomist. In a few cas- es, the tracing was complemented with fragments of the holotype that Schlechter, and later his wife, made availa- ble to Ames for the Herbarium of the Botanical Museum at Harvard. After the loss of Schlechter’s types in the fire Figure 3. Part of the Berlin Herbarium destroyed during WWII, March 1943. Archives of Rudolf Jenny and courtesy of Dr. N. Kil- ian, Archives BGBM Berlin-Dahlem. Figure 4. Oakes Ames (1874–1950). Portrait by his wife, Blanche Ames. 39Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum of the Berlin Botanical Museum, these materials repre- sent the most precious source available today for schol- ars to give a face to the descriptions of the new orchid species published by Schlechter in over twenty years of activity, as they make up the only extant evidence of the original materials. The need for a solid framework that allows a consist- ent application of species’ names relative to the orchid flora of Costa Rica, as well as a critical examination of the taxonomic status of previously synonymized names, has become more and more compelling as the works devoted to the treatment of Orchidaceae for Flora Cos- taricensis are nearing completion. Cataloguing the infor- mation on type designations for Costa Rican orchid names and designating new types when appropriate has been particularly important and critical in the orchid species originally described by Schlechter due to the destruction of the main set of type specimens and the associated analytical drawings and notes. This patient work was inaugurated by Barringer, who in 1986 published a comprehensive paper on the typification of the Costa Rican orchids species described by Schlechter based on the extensive collections by Alberto M. Brenes (Barringer 1986). Pupulin (2010a) faced another large group of orchids described by R. Schlechter from plants collected in Costa Rica by Karl Wercklé, presenting a catalogue of 84 species and pro- viding lectotypification for 60 of them. Another impor- tant step was made in 2016, when Pupulin and collabo- rators typified the over 60 orchid names based on col- lections carried out by Adolphe Tonduz, proposing 36 lectotypes and two neotypes (Pupulin et al. 2016). Boga- rín et al. (in prep) devoted their attention to the orchids sent to Schlechter in 1921 by G. Acosta, upon which the taxonomist described 22 new species in 1923; they desig- nated 13 lectotypes and three neotypes. The present paper follows previous contributions of this nature. It is dedicated to the typification of orchid species based on Costa Rican material originally gath- ered by several different collectors and described by Schlechter between 1907 and 1923. The selection of the floral analyses made by Schlech- ter for lectotypification purposes, which has been adopted in previous papers (Barringer 1986; Mora and Atwood 1992, 1993; Atwood 1999; Pupulin 2010a; Pupu- lin et al. 2016), has been questioned by some authors, reviewers or online databases (i.e. Tropicos 2021) based on a supposed “posteriority” of the illustrations com- pared to the time in which the original materials were prepared (Hermans et al. 2020) or because these draw- ings are not considered original material and therefore it is an error to be corrected to neotype, Article 9.10 the Shenzhen code (Turland et al. 2018). This is a very strict interpretation of the Article 9.4 that defines origi- nal material as the material that “comprises the follow- ing elements: (a) those specimens and illustrations (both unpublished and published before the publication of the protologue) that the author associated with the taxon, and that were available to the author prior to, or at the time of, preparation of the description, diagnosis, or illustration with analysis (Articles 38.7 and 38.8) validat- ing the name. However, Article 9.4 per se does not del- egitimize the reproductions of original material since they are copies of the type material, and therefore, they are unequivocally original material. Indeed, the fact that Mansfeld published Schlechter’s drawings in 1931, five years after the death of the Ger- man taxonomist, is certainly undeniable. Likewise, it is unquestionable that the drawings traced by the artists hired by Prof. Ames, and made under the direct super- vision of Schlechter, were executed several years after Schlechter had prepared the original materials for the descriptions of his new species. Even more certain is that the negatives exposed in 1929 by J. Francis Macbride in the herbarium of the Berlin Botanical Museum and the pictures taken by Ames were actually printed on photo- graphic paper only years later and that Schlechter never even saw these “photographs”. However, no one should doubt that these materials are reproductions of the original material that Schlechter kept in his herbarium and subsequently deposited in the Berlin herbarium. The original drawings affixed to Schlechter’s herbarium sheets, immortalized by the negatives of Macbride and Ames, are indistinguishable from those published by Mansfeld (1931) and from the drawings traced for the herbarium of the Botanical Museum of Harvard Uni- versity (Figure 5). Questioning the conformity of these reproductions to the original materials, that is, question- ing their “veracity”, simply raises a long-standing and complex problem relating to the technical reproducibil- ity of illustrations, a technological and engineering issue that has continuously evolved and changed over time, to bring today to the apotheosis of electronic image and absolute reproducibility. We believe that no one would question the conformity to the “original” of an image taken today with a mobile phone camera, and the pos- sibility of using it in a publication indicating it as a “lec- totype”, even if an expert photographer would not miss the possible inconsistencies in the geometry and colors introduced into the image by the perspective, as well as the type of lens, sensor, and software used. The floral analyses published by Mansfeld are in no way “similar” or “inspired” to Schlechter’s originals, but their faith- ful reproduction according to the technical possibilities 40 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Figure 5. A, Drawing of Scaphyglottis subulata from the photo of the holotype taken in the Berlin Botanical Museum and printed in photo- graphic paper kept at AMES 39613. B, Drawings based on the type of S. subulata published by Mansfeld, 1931: Pl. 44, No. 176. C, drawings traced by the artists hired by Prof. Ames, and made under the direct supervision of Schlechter of Oncidium pittieri kept at AMES 24264. D, Drawings of the type of Oncidium pittieri published by Mansfeld, 1931: Pl. 74, No. 295. 41Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum allowed by the printing advances of those times (Figure 5B). The artists employed by Ames to work in Schlech- ter’s herbarium did not “imitate” his floral analyses but traced them under the watchful eye of the German bota- nist. These reproductions were indeed technically closer to the originals than were the engravings made for de Sancha’s press concerning the paintings originally made in Peru by the draughtsman of the botanical expedition of Ruiz and Pavón (1794, 1798), or the illustrations of Plumier’s Antillean collections published by Burman (in Plumier and Burman 1755), which were themselves cop- ied from what was already a hand-made copy (the Codex Boerhaavianus) of Plumier’s original drawings (Goethart 1910; Ossenbach 2016), and were used nonetheless to lec- totypify Linnaeus’s (1759) orchid species (e.g. McLeish et al. 1995). Plumier himself never saw the materials with which the species were lectotypified, for the simple rea- son that when Burman’s work was published in Amster- dam, he had been dead for nearly fifty years. And as for the quality of Burman’s copies compared to the original drawings made by Plumier, it might be useful to quote the opinion of John Lindley, who, in addition to being one of the greatest orchidologists of all time, was also an excellent illustrator: “Plumier’s Mss. appear, from the copy in Mr. Lambert’s Herbarium, to give a very clear account of this beautiful species [Epidendrum atropur- pureum]; yet Burman, with his usual skill, converted the figure into a caricature […]” (Lindley 1830–1840, p. 100). Some authors seem to favor using these same mate- rials – which are nothing but reproductions of the origi- nal analyses made by Schlechter – under a different type category, selecting them as neotypes rather than lectotypes or just using them as a reference for select- ing a neotype (Hermans et al. 2020). However subtle, the difference is certainly more than semantic. As it must be done among the materials that the original author knew and referred to in the description of a new taxon, the designation of a lectotype does not introduce any element of subjective judgment by subsequent authors on the identity of the taxon, except for the verification that the elements chosen for that purpose are in agree- ment with the protologue (because even the original authors may have made mistakes). On the contrary, the designation of a neotype virtually represents an entirely subjective interpretation of the original authors’ ideas and concepts because it uses materials that the author has never known or referred to it. There is undoubtedly a gradient of “certainty” in the various type categories that the Code visualizes to give stability to plant names. All the materials that can be selected to lectotypify a name (e.g. isotypes, syntypes, paratypes, original draw- ings and illustrations of the type, etc.) have in common the fact that the author of the name has identified them, and therefore adhere to “his” concept of species. Neo- typifications and epitypifications, on the other hand, must be viewed with greater caution since they make use of materials that were not identified by the original author and which correspond to the concept of the spe- cies according to “someone else”, however experienced. Also, the Tropicos database (https://www.tropicos. org) treated the lectotypifications based on the copy of Schlechter’s sketch of the holotype in Pupulin (2010a) and Pupulin et al. (2016) as neotypifications, erroneously stating that “corrected here to neotype because a depic- tion of the specimen is not considered original material” (Tropicos 2021). However, the Article 9.3 of the Shenz- hen code (Turland et al. 2018) states that “a lectotype is one specimen or illustration designated from the original material”. Also, Art. 9.12 states that “in lectotype desig- nation… if no isotype, syntype or isosyntype is extant… the lectotype must be chosen from among the uncited specimens and cited and uncited illustrations that com- prise the remaining original material”. Therefore, if an illustration meets the above requirements, it can be con- sidered original material and thus selected as a lectotype. For this reason, we choose, in the past, to use these reproductions to lectotypify the species described by Schlechter, and for the same reason, we keep this choice here. Also, other authors favored this view and selected the illustrations of the flower analysis made by Schlech- ter and posthumously published by Mansfeld (1931) as lectotypes (Guimarães et al. 2019). When none of the syntypes or isotypes has been preserved, these mate- rials must, in our opinion, be considered an integral part of the original materials as Schlechter knew them. They are technically reliable reproductions and certain- ly incomparably closer to his concepts than any “new” material we might select for typification purposes. It may be that a more orthodox application of the provi- sions of the Code – but also objectively less sensitive to the essential elements of botany – convinces readers that our lectotypifications must be “corrected” to neotypifica- tions. The Shenzhen code (Art. 9.10) (Turland et al. 2018) admits this possibility, recognizing the value of the con- ceptual discussion concerning typification and consider- ing our proposals as correctable errors without affecting the validity of the relative interpretations and choices. From our point of view, the choice of lectotypification is undoubtedly more consistent with the taxonomic story of the species discussed in this paper and preferable for nomenclatorial stability. Schlechter organized his magnum opus on the flora of Costa Rican orchids in various chapters, dedicated to those botanists, collectors, and patrons who provided the 42 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín most significant contributions, in terms of quantity and variety of materials, to his studies. “Orchidaceae Amp- aroanae”, “Orchidaceae Bradeanae Costaricenses”, “Orchi- daceae Brenesianae” commemorate some of these impor- tant figures (Schlechter 1923a, 1923b, 1923c). The names of other important figures of Costa Rican botany at the turn of the century, such as Tonduz and Wercklé, were recognized in the chapter dedicated to the “patroness” of their activities, Doña Amparo (Pupulin 2010a; Pupu- lin et al. 2016). The case of Guillermo Acosta, author of two important orchid shipments to Berlin in 1921, has been discussed by Bogarín et al. (in prep), highlighting his close and, in part, unrecognized relationship with Tonduz. The contributions of other collectors, smaller in quantity and perhaps less systematic in intentions, although not less botanically important, were gathered by Schlechter in a particular chapter dedicated to “Vari- ous collectors” (Schlechter 1923d). This chapter includes, as its main content in numerical terms, the collections that Guillermo Acosta sent to Schlechter in 1921, whose typification was dealt with by Bogarín and colleagues (in press). Alongside the Acosta orchids, however, there are numerous species of other botanists and naturalists active in Costa Rica at the beginning of the 20th century, whose collections reached Schlechter’s desk through the shipments to European specialists made by the curators of the Museo Nacional. For the sake of completeness, we have included in this paper of typification of the orchids collectorum variorum also those of some species that did not reach Schlechter’s hands directly, since they were not collected during the period of his botanical activity, but rather date back to the second half of the nineteenth cen- tury. Schlechter had the opportunity to study them in Vienna when the herbarium of Reichenbach was newly made accessible to the public after it had been closed for 25 years by the will of his testament. Between 1907 and 1923, he described a dozen new species based on the col- lections that these early travelers and explorers made in Costa Rica from 1857 to 1888 (Schlechter 1907a, 1907b, 1918a, 1920, 1921b, 1923d). But let us now have a closer look at the different fig- ures, in chronological order and importance, who make up the cast of the “various collectors” of Schlechter’s Costa Rican orchids. COLLECTORES VARII ORCHIDACEARUM COSTARICAE AB R. SCHLECHTER DESCRIPTAE Karl Hoffmann The Germans Karl Hoffmann (1833–1859) (Figure 6) and Alexander von Frantzius (1821–1877) came to Costa Rica in 1853, bearing letters of recommendation from Nees von Esenbeck, President of the German Academy, and Alexander von Humboldt for President Juan Rafael Mora. They arrived at Greytown (San Juan del Norte) as passengers of the brig Antoinette, together with a group of German immigrants, and continued to Costa Rica along the road of Sarapiquí (Hilje 2007). Frantzius was a professor at the Physiological Institute in Breslau, and Hoffmann was well-known for his practical and literary works during the cholera epidemics in Berlin during the years of 1848 and 1849. Soon they began to explore the country and collect specimens, mainly botanical. Hoffmann was later a physician in the Costa Rican army during the war against pro-slavery activist W. Walker. At the same time, Frantzius soon became a suc- cessful businessman and owner of a pharmacy. Hoff- mann and Frantzius spent their leisure time, the first dedicated to collecting plants and studying their natural distribution, the second to similar studies in mammals and birds. Hoffmann climbed two of Costa Rica’s most Figure 6. Karl Hoffmann (1823–1859). Courtesy of Luko Hilje. 43Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum important volcanoes: on May 5, 1855, Irazú near Cart- ago, and in August of the same year, Barva in the prov- ince of Heredia. Hoffmann intended to write a book with the title Flo- ra and Fauna of Costa Rica, but he had to abort this idea because of the war and his illness. After the war against Walker, Hoffmann retired to Puntarenas, where he died in 1859. His mortal remains were brought to San José in 1929, where they were buried with military honors. Hoffmann sent his collections to the herbarium of Berlin, to the renowned botanist Johann F. Klotzsch. They were later described by Reichenbach in 1866 in his Orchideae Hoffmannianae (Reichenbach 1866). One can find among them the types for three new species: Pelexia hoffmannii Rchb.f., Epidendrum (= Prosthechea) ionophlebium Rchb.f. and Ponera albida Rchb.f. Schlechter described an additional new orchid spe- cies collected by Hoffmann as Epidendrum hoffmannii (= Prosthechea ionophlebia (Rchb.f.) W.E. Higgins) after he was able to visit Reichenbach’s herbarium in Vienna after World War I. Hoffmann collected it in the small village of Curridabat, to the east of San José (misspelled by Schlech- ter as ‘Curidabal’) (Pupulin and Karremans 2007). Auguste R. Endrés Auguste R. Endrés (1838–1875) was perhaps the most proficient and dedicated botanist who ever studied the orchid flora of Costa Rica. His name appeared sporadi- cally in the  Gardener’s Chronicle  since 1871, associated with orchid novelties described by Reichenbach in Ham- burg. But unfortunately, we have no portrait of Endrés.  He was born in Herbitzheim, a village in the depart- ment of Bas-Rhin in Alsace, France, of a German fam- ily, and the roots of his culture were German. In 1855, Endrés moved to New York with his grandfather Auguste Reeb, where he was joined by the rest of the family two years later. Here he made the acquaintance of Isaac Buchanan, a well-known horticulturist, who  intro- duced  Endrés to several of the most famous names in orchidology of that time, such as William Hooker, George U. Skinner, James Bateman, Hugh Low, John Day, and Capt. John Dow, probably Endrés’ best friend dur- ing the years of his Costa Rican adventure. Skinner and Bateman decided to employ Endrés to collect orchids in Costa Rica, following a recommendation by Buchanan.  In 1866, with a commission to collect for Bateman and for Professor Reichenbach, Endrés arrived in Grey- town, Nicaragua. He  traveled along the  San Juan River to  neighboring Costa Rica  by canoe. His first known orchid, Dichaea trulla, was collected and illustrated that same year. During the next seven years spent  search- ing  for orchids, Endrés explored all  corners of  Costa Rica known (Ossenbach et al. 2010; Ossenbach 2013; Ossenbach and Pupulin 2013). Economic constraints forced him to collect orchids for horticultural purposes and even  work as the superintendent of the construc- tion of a new road, but he never stopped collecting for science. The Pleurothallid orchids were his main scien- tific interest, particularly the genus Lepanthes, of which Endrés discovered, described, and illustrated, over  two- thirds of all the species known from Costa Rica.  Endrés traveled to Europe in 1874 to discuss with Reichenbach – with whom he had a contrasting human and scientific relationship – the future of his research and the use of his materials. During their meeting in Hamburg, Endrés made the acquaintance with the great Czech collector Benedikt Roezl.  It was likely on his sug- gestion that he eventually sailed to Colombia, where he fell ill from pleurisy  while  traveling toward the high- lands of the Cordillera de Santa Marta and died in November of 1874.  What remains of his work  shows  that Endrés was planning a formal treatment of the orchids of Costa Rica, something  to resemble  a modern orchid f lora. To  this aim,  he committed himself  to explore,  collect and prepare specimens,  write  descriptions, and  made botanical illustrations of all the orchid species of Cos- ta Rica he  could  observe (Pupulin 2013).  However, with thousands of botanical drawings, accurate descrip- tions, and references to the collecting localities ready for the press, plus all the relative dried materials at hand, Reichenbach ended up publishing just a few of them, mainly in his own cryptic descriptive style.  After the death of Reichenbach in 1889, his her- barium, including all Endrés’ unpublished work, was bequeathed to the Natural History Museum of Vien- na.  There,  it remained closed for study for another 25 years  because of  the clauses of his will.  Finally, howev- er, Schlechter could study the immense labor left behind by Endrés in the recently opened orchid collection at the Hof Imperial Museum in Vienna during his visit around 1915 (Jenny, pers, comm. 2011), a few months before the beginning of the First World War. Here, among the plants collected 40 years before by Endrés, he described three as new to science (Schlechter 1921a). Schlechter named  Chondrorhyncha endresii  in honor of the great explorer and botanist. Friedrich Carl Lehmann As a collector for Hugh Low & Co. of London, Frie- drich Carl Lehmann (1850–1903) (Figure 7) went to South America in the late 1870s. Around 1878 he settled 44 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín in Popayán, Colombia, where he held the position of the consul of Germany until his death. He made significant discoveries of new Colombian orchids, especially in the genus Masdevallia, which was his favorite. In 1883 he was described as “the most important traveler and col- lector in the United States of Colombia and neighboring territories of our time” (Regel 1883). In 1878, Reichen- bach had published his Orchidaceae F. C. Lehmannianae ecuadorenses, where he described Lehmann’s collections in Ecuador from the year 1876. At the beginning of the 1880s, Lehmann traveled to Costa Rica, Panama, and Guatemala. Although his journey’s exact dates are not known, the first dated col- lection from our area is the type specimen of Catasetum blepharochilum (=Catasetum maculatum) (Lehmann 1061, Costa Rica), in December 1881. In a short time, he discovered numerous new Central American species, described later by Schlechter and Kränzlin. An important number of the orchids collected by Lehmann were described by him and F. Kränzlin in 1899 under the latter’s Orchidaceae Lehmannianae in Guatemala, Costarica, Columbia et Ecuador collectae, quas determinavit et descripsit (Kränzlin 1899). Lehmann liked to say: “I attribute my good health, and even my life mainly to two things: First, when in danger either from natives or, worse still, from lawless white men, I never produce a revolver or other weap- on… Secondly, I never drink water without first boiling it” (Taylor 1974, p. 176). His precautions did not help him. He shared the fate of many other explorers of these regions and died by drowning in 1903 while trying to cross the Timbiquí River to visit a gold mine in which he had interests. His widow sold his herbarium and his drawings to the herbarium at Kew. Lehmann was also an excellent illustrator. Many of the pencil drawings with which he accompanied his herbarium specimens are preserved at the herbarium at Kew (Cribb 2010). He also wrote the notes for the geo- graphical descriptions in the monograph of Masdeval- lia edited by the Marquis de Lothian and illustrated by Miss Woolward, where his extensive knowledge about the plants in their native habitats can be appreciated. A new genus was dedicated to him by Kränzlin: Neole- hmannia. Lehmann was quite generous in distributing his materials to several botanists and institutions, mostly in Europe. Notable among these were H. G. Reichenbach in Hamburg, R. A. Rolfe in Kew, H. N. Ridley at the British Museum, F. Kränzlin in Berlin (who eventually sold his materials to Hamburg) among others. Even though the largest part of Lehmann’s personal herbarium, together with his plant illustrations, were acquired by Kew from his widow in 1903 (Cribb 2010) and are now hosted at the Herbarium of the Royal Botanic Gardens, Kew, UK (K), we are aware of Costa Rican orchid specimens col- lected by Lehmann in several other herbaria, both in Europe and in the United States. Interestingly, although the types of some orchid species collected by Lehmann and later described by Schlechter were regarded as destroyed in the herbarium fire of the Berlin-Dahlem Botanical Museum (i.e. Garay 1978; Guimarães et al. 2019; Hágsater 2009; Luer 2017, 2018; Ormerod 2002, 2008), the presence of these speci- mens at B is highly doubtful. Lehmann used to court his contemporary botanists, sending them plants to provide names for his collections. Certainly, he could not have had a relationship with Schlechter, who was a generation younger, and whose interest in American orchids did not begin until the second decade of 1900, when Lehmann had been dead for over ten years. We know that dur- ing the last decade of the 19th century, Lehmann sent material to Fritz Kränzlin, then in Berlin, who in 1899 dedicated a long article to him in which he determined the collections received from Lehmann and described 107 new species, most of them under Lehmann’s joint Figure 7. Friedrich Carl Lehmann (1850–1903). Unknown artist. 45Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum authorship. This material was probably lent by Kränzlin to the Herbarium of the Berlin-Dahlem Botanical Muse- um. Still, after 1903 it had to be returned to Kew, which had acquired ownership of Lehmann’s collections (Cribb 2010). As to the materials of his private herbarium, prob- ably due to Kränzlin’s rivalry with Schlechter, these were eventually not bequeathed to the Berlin-Dahlem Museum, as it might have been expected, but were sold instead to the Herbarium Hamburgense, where they are still held today. In any case, since the article that Kränz- lin dedicated to Lehmann includes not only the descrip- tion of the new species, but also the identification of the remaining material received in Berlin, it is important to note that there is no mention of any of the species col- lected in Costa Rica and later described by Schlechter. For this reason, it seems reasonable to believe that it was not in Berlin where Schlechter studied Lehmann’s mate- rial but elsewhere. Although we have tried to answer the question con- cerning where Schlechter may have studied Lehmann’s Costa Rican plants, we have not reached an obvious con- clusion. We have been able to observe specimens of the relatively few orchids collected by Lehmann in Costa Rica between 1881 and 1882 in the herbaria of the Nat- ural History Museum in London (BM), Meise Botanic Garden, Belgium (BR), Geneva, Switzerland (G), the Herbarium Hamburgense, Germany (HBG), the United States National Herbarium at the Smithsonian Institu- tion, Washington, U.S.A. (US), and the Natural History Museum in Vienna, Austria (W) (herbaria acronyms according to Index Herbariorum). None of the speci- mens we studied, however, are annotated and determined in Schlechter’s unmistakable handwriting. We know with certainty that he used to annotate the samples that were sent to him for determination because the National Museum of Costa Rica has a rich series of duplicates of collections made by the scientific staff of the Museum, on which Schlechter affixed his own labels before return- ing the sheets. This leads us to believe that none of the surviving specimens, among those we have been able to trace, represent the holotype used by Schlechter for his descriptions or, even more so, to make his precise draw- ings of the plants and their floral analyses. Even though we cannot state this with absolute certainty, the possibility exists that the holotypes of these species described by Schlechter nevertheless exist in some herbarium, public or private, that we have not had the opportunity to examine. For this reason, in the paragraphs dedicated to the few Lehmannian orchids described by Schlechter, we preferred to indicate that the holotype has not been located. Even if, in the absence of specimens that can be unequivocally interpreted as holo- types, we have designated the relative lectotypes for rea- sons of nomenclatorial stability, we maintain the hope that such specimens may be found in the future making our lectotypifications superfluous. Richard Pfau A Swiss, Richard Pfau (1856–1897) founded a nurs- ery in San José, Costa Rica, in the final years of the XIX century, that sold a great variety of ornamental plants. He also collected native plants for export. Through his collections, we know that he was also in Panama and Colombia, and at least one of the new species described from plants sent to Europe by Pfau came from Mexico: Vanilla pfaviana Rchb.f. Pfau wrote the first work published in Costa Rica about the orchids of this country: New, Rare and Beauti- ful Orchids of Costa Rica (ca. 1895), of greater interest for horticulture than for botany. In this work, Pfau advises on how to grow and pack orchids for exportation and included a list of the species he had for sale in his nurs- ery (Figure 8). But Pfau’s voice was also one of the first to address the rising concern about the destruction of our nature when he describes one of our most beautiful orchids: “Cattleya skinneri, some ten years ago, was a common Orchid all over Central America; but in the last few years it has been exported by shiploads; and to-day – at least in Costa Rica – it has almost become rare” (Pfau ca. 1895). Pfau also wrote several articles about Central America and its orchids, such as “The climate of Cen- tral America, Orchid culture” (Pfau 1883), “Notes on the fertilization of Orchids in the Tropics” (Pfau 1894), and “Costa Rica and its Orchids” (Pfau 1896). As did Roezl and Wallis, Pfau sold his plants in Europe through the agency of Eduard Ortgies in Zurich. Schlechter described several orchids collected by Pfau in Costa Rica, such as Sobralia pfavii and Telipo- gon pfavii. Previously, Reichenbach had described other specimens collected by Pfau in Panamá (Pleurothallis pfavii and Trichocentrum pfavii), and Rolfe described from Costa Rica Epidendrum pfavii, of which a colored illustration by Pfau is preserved on the same sheet as the type specimen in Kew (Figure 9). Juan José Cooper Sandoval Henry Cooper, a British mining engineer, came to Costa Rica in 1825, under a contract with the gov- ernment to survey agricultural lands, claimed by the wealthy landowner Victoriano Fernández, in the north- 46 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín ern plains of San Carlos bordering a river that has been since then known as Río Cooper. A small hamlet in the same area is also known under the toponym of Cooper. However, it is often misspelled as Kopper, after a Ger- man family who settled in the region some thirty years later. Cooper then remained in the country, working in several mines in the hills of Aguacate. He eventually married a Costa Rican girl named Margarita Sandoval, and Juan José Cooper (1843-1911) (Figure 10A), their second son, was born in 1843 (Hilje 2014). Juan José Cooper was strongly drawn to the natu- ral sciences. Early in his life, he began to work as an assistant to Alexander von Frantzius at his pharmacy in San José. Several young men made their first expe- riences at von Frantzius’ pharmacy, such as Ernesto Rohrmoser, Gerhard Jäger, Manuel Carranza, and Juan José Cooper. They assisted von Frantzius and learned soon to prepare bird skins. Still, their enthusiasm soon diminished, to the point that von Frantzius complained in a letter to Wilhelm Peters, at the Zoology Museum in Berlin: “they behave like small children”! Things changed when young José Cástulo Zeledón (1846–1923) (Figure 10B), rightly called Costa Rica’s first natural- ist, became his pupil. Through von Frantzius’s recom- mendations, Zeledón was admitted to the Smithsonian Institute in Washington, where he spent several years before returning to Costa Rica as a qualified ornitholo- gist (Hilje 2018). Zeledón went in 1872 as a zoologist with William Gabb’s expedition to Talamanca, and Juan José Cooper went with him as his assistant. They returned to San José with an extensive collection of birds (May 2016). A few years later, when Robert Ridgway described and named the Pacific screech-owl as Megascops cooperi, he wrote, “I have named this species at the request of Mr. Zeledón, the collector of the type-specimens, after Mr. Juan Cooper, of Cartago, Costa Rica, a particular friend Figure 8. Cover of Richard Pfau’s book on the orchids of Costa Rica. Printed by the author, San José, ca. 1895. Figure 9. Epidendrum pfavii. Coloured illustration by Pfau on the same sheet of type specimen (K000463409). 47Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum of his, to whom he is much indebted for many interest- ing contributions to his collection.” In the early 1890s, Cooper was hired by the recently founded National Museum of Costa Rica as a botanist and taxidermist. Although he collected some 200 species of birds, in his older years, he dedicated himself more to botany, collecting mainly in the vicinity of Cartago, his city of residence. Schlechter named Stelis cooperi (col- lected in 1888) and Pleurothallis cooperi (collection date unknown) after him. Henri Pittier As part of an educational reform aimed at secular- izing public education, the government of President Ber- nardo Soto (1885–1889) hired a group of European aca- demics to staff the two new public high schools in the capital, San José. The arrival of these academics marks the beginning of a small scientific renaissance in Costa Rica. Two institutions symbolize this renaissance: the National Museum and the Instituto Físico-Geográfico (IFG), founded in 1887 and 1889, respectively. Among the hired teachers were Pablo Biolley (1861– 1908) and Henri Francois Pittier (1857-1950) (Figure 11), who respectively arrived in 1886 and 1887. Pittier lived in Costa Rica until 1905 and, during these years, conducted a systematic exploration of the Costa Rican flora that had no equal in his time in any country of tropical America. These efforts resulted in the publication of the Primitiae Florae Costaricensis, the first flora of Costa Rica, a work that unfortunately was not concluded. It was published in conjunction with a Belgian colleague, Téophile Alexis Durand (1855–1912), and appeared in three volumes and 12 fascicles, published from 1891 to 1905. According to Paul C. Standley (1937: 49), in his introduction to the Flora of Costa Rica, “Henri Pittier has undoubtedly gained a more intimate knowledge of the natural history and especially the botany of Central America and northwestern South America than has ever been possessed by any single person.” Although hired to teach at secondary schools, Pit- tier had more ambitious ideas. After he arrived in Costa Rica, he started to fight to form a meteorological obser- vatory and an institute. The Meteorological Institute was Figure 10. A, Juan José Cooper Sandoval (1843–1811). Courtesy of Luko Hilje. B, José Cástulo Zeledón (1846–1923). Unknown photogra- pher. 48 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín founded in April 1888 under the direction of Pittier. Pit- tier’s work at the Institute went parallel to the founda- tion of the National Museum, of which Anastasio Alfa- ro was named the first director. Pittier, who had been in Costa Rica for only two months, was named on the Board of Directors of the Museum, together with Pablo Biolley and José Cástulo Zeledón. The combined efforts of Pittier, Alfaro, Tonduz, Biolley, Wercklé, Brenes, and the Brade brothers resulted in the formation of the National Herbarium that counted initially with more than 5,000 species and “was unequaled below the Río Grande del Norte” (Standley 1937). Unfortunately, Alfaro and Pittier, the two promi- nent scientific entrepreneurs in the small country, never could agree on how to organize their operations. In 1889 the government consolidated the Museum and the Meteorological Institute into one center, the ‘Instituto Físico-Geográfico Nacional de Costa Rica’. This was a temporary triumph for Pittier, who was named director. However, integration only lasted a few months, and the Museum was again separated from the rest of the Institute. The inevitable outcome was that constant intrigues and lack of funds led to Pit- tier’s final falling out with the government. In 1905 he left the country to work in the United States and Pan- ama and a long and distinguished career in Venezuela until he died in 1950. The Instituto lost its creator and engine, and Costa Rica a dynamic and prolific scientist with his departure. Henri Pittier was always interested in orchids. While working on his Primitiae Flora Costaricensis, he sent a significant number of specimens to his friend Théophile Durand in Brussels, who passed them on to Schlechter in Berlin for identification. The orchids collected in Pan- ama during his work for the United States government went the same way. After initial differences (Schlechter, for some time refused to return the material sent by Pit- tier), Pittier worked together with Schlechter until the death of the German scientist in 1925. In 1906, Schlech- ter dedicated a new genus of orchid to Pittier: Pittierella (today a synonym of Cryptocentrum or Maxillaria s.l.) and several new orchid species, among them Cranichis pittieri, Epidendrum henrici, Lockhartia pittieri, Notylia pittieri, Oncidium pittieri, Scaphosepalum pittieri, and Vanilla pittieri. Paul (Pablo) Biolley Pablo Biolley (1862-1908) (Figure 12) was born in the Swiss town of Neuchâtel in 1862. He obtained his degree in natural sciences there and continued his stud- ies in the Netherlands and Germany. Biolley formed part of the first group of Swiss teachers hired by the govern- ment of Bernardo Soto and was appointed as professor of the recently founded ‘Liceo de Costa Rica’, where he began teaching in 1877. He established himself perma- nently in Costa Rica, obtaining Costa Rican nationality and marrying a Costa Rican. He died in 1908 at the age of forty-six. His sister Stella arrived in 1889 and was a teacher at the ‘Colegio Superior de Señoritas’ for many years. In Pablo Biolley’s honor, a village and a district in Costa Rica’s southern region were named ‘Biolley’. Also, an important height in the cordillera of Talamanca car- ries the name ‘Cerro Biolley’. Immediately after he arrived in Costa Rica, he became one of the scientists who gave generous impulse to the foundation and development of Costa Rica’s first scientific institutions and was named naturalist of the National Museum also occupying a chair on the Board of Directors (Díaz and Solano 2009). Figure 11. Henri Pittier (1857–1950) in 1903. Unknown photogra- pher. 49Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Biolley accompanied Pittier during many of his explorations, often in the company of Adolphe Tonduz, and was, for a short period (1904) director of the Insti- tuto Físico-Geográfico. He also went on botanical excur- sions with Charles H. Lankester to the Atlantic region, collecting in Turrialba and Peralta’s vicinity. Although Paul Biolley’s primary interest was in entomology, he made important contributions to the knowledge of the Costa Rican flora. To him, we owe, among others, the discovery of the types of Maxillaria biolleyi (Schltr.) L.O.Williams) and Telipogon biolleyi Schltr. An interesting species of the Costa Rican orchid flora, Epidendrum insulanum, was described by Schlech- ter from a collection by Pittier during an expedition in 1902 with Biolley to Cocos Island. Emel Jiménez Segura During the government of President Rafael Yglesias Castro, between 1894 and 1902, several young Costa Rican teachers were sent to complete their education at Santiago de Chile’s Pedagogical Institute. Among those who returned to Costa Rica at the turn of the century were several prominent educators such as José Fidel Tris- tán, Miguel Obregón Lizano, and Roberto Brenes Mesén. Miguel Obregón was named Consul of Costa Rica in Santiago in 1899, and received the commission of select- ing a Chilean professor to take over as director of the Liceo de Costa Rica, founded three years earlier in the aftermath of the educational reform of President Ber- nardo Soto Alfaro. Doctor Zacarías Salinas was selected and arrived in San José in 1900. He immediately went to work, beginning a profound reform of the school’s cur- ricular system. Salinas hired several of the teachers that had trained in Chile, such as José Fidel Tristán and Roberto Brenes Mesén, and was instrumental in selecting a new group of students that was sent to Santiago in 1901, among them Joaquín García Monge, Alberto Rudín (younger brother of Juan Rudín, brother-in-law of Henri Pittier) and Emel Jiménez Segura (1881–ca. 1960) (Figure 13), who returned from Chile in 1904 and formed part of the new staff of the Liceo. Emel Jiménez was put in charge of the Department of Natural Sciences. Botanist Otón Jimenéz Luthmer, who studied at the Liceo de Costa Rica and was a pupil of Emel Jiménez, described him as demanding and of strong character but praised his humanity and sense of justice. Jiménez taught his botany classes using live material whenever possible, and for this, he counted on the friendship he had established with Alfredo Brade, a German gardener who had a plant nursery in San José and supplied him with the necessary specimens (Jiménez 1959). Emel Jiménez continued at the Liceo de Costa Rica until well into the 1940s. In his last active years, between 1938 and 1946, he was joined at the Liceo by another renowned Costa Rican botanist, Rafael Lucas Rodrígues Figure 12. Paul Biolley (1862–1908). Unknown photographer. Figure 13. Emel Jiménez Segura (1881–ca. 1960) with wife Telma Royo. Courtesy of his grandson Gerardo Mora Jiménez. 50 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Caballero. Emel Jiménez and his generation of Costa Rican educators formed in Chile had a strong influence on Costa Rica’s educational system in the first half of the XX century. Schlechter described Epidendrum urostachyum from a collection by Emel Jiménez in 1913, in the hills of El Tablazo, to the south of San José. Ferdinand Nevermann Wilhelm Heinrich Ferdinand Nevermann (1881– 1938) (Figure 14) was born in Hamburg and arrived in October 1909, having graduated with honors as a mechanical engineer a few years earlier. After exploring the whole country and having started a family, in 1918 Nevermann acquired a farm which he called ‘Ham- burgo’, in El Cairo de Siquirres, in the Atlantic region of Costa Rica. While investigating the insects that attacked his banana plantations, Nevermann began an interest that led him to become one of Costa Rica’s most important entomologists but a world authority on this subject. After World War I, the call by the German gov- ernment to all its citizens living abroad to help refur- bish the collections of the German museums that had been destroyed led to Nevermann, sending insect col- lections to Germany with increasing frequency. In these years, he established close relations with Ber- lin’s Museum and Botanical Garden. Nevermann also sent plants. There is a beautiful white orchid, Coryan- thes nevermannii, which we owe to him” (Apuntes… 1938: 341). This reference is curious because there is no record of an orchid with the epithet ‘nevermannii’ in the international registers of botanical nomencla- ture. The answer to this riddle can be found in a let- ter by Rudolf Schlechter to Nevermann dated May 8, 1925: “The two orchids which were sent to me inter- ested me vividly. The double inflorescence with the big pendant flowers is a new species of Coryanthes, which I will soon describe as Coryanthes nevermannii Schltr. It is the first species of Coryanthes that until now I have known from Costa Rica. It is for me a special pleas- ure to dedicate this plant to you. Not smaller interest raised the slender-leaved Vanilla. This one also has not yet received a name. It will carry your name as Vanilla nevermannii Schltr.” (1925 May 8 letter from Schlech- ter to Nevermann). Schlechter died six months later, in November 1925, and the species dedicated to Never- mann were never published. When in 1943 Schlechter’s herbarium was destroyed, all evidence of Nevermann’s collections disappeared. Thus, we will never know for sure which species correspond to Coryanthes never- mannii and Vanilla nevermannii. In 1936 he took over the Chair of Entomology at the National School of Agriculture, but died shortly after- wards in an unfortunate accident. During the night of June 30, 1938, while studying the nocturnal behavior of a species of ant, he was shot by the son of a neighbor who mistook him for an intruder. Paul C. Standley visited Nevermann at his farm, where he collected several species of orchids, wrote in his Flora of Costa Rica: “To Mr. Ferdinand Nevermann there are special obligations for a most pleasant and profitable visit to his fincas in the lowlands along the Reventazón River. Enviable is the botanist who receives a welcome from so considerate a host, or visits the forest with so competent a guide” (Standley 1937: 59). Figure 14. Wilhelm Heinrich Ferdinand Nevermann (1881–1938). Courtesy of his granddaughter Helga Nevermann. 51Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Otón Jiménez Schlechter (1918a, p. 371) wrote: “a young collec- tor stands out lately in Costa Rica, O. Jiménez, who in a short period of activity has already found a series of new species and, through his efforts, promises to enrich significantly our knowledge about the f lora of that country, especially of the Orchidaceae.” Otón Jiménez (1895–1988) (Figure 15) had the good fortune to study at the Liceo de Costa Rica in its golden age, with teach- ers like Emel Jiménez, Dr. Michaud, and Paul Biolley. Of a precocious intelligence, he was only 17 years of age when he was appointed as director of the Herbarium of the National Museum, a position he held until 1914. He remembered his first encounter with Charles H. Lank- ester in 1911: “I still remember his smile while shaking hands with me, observing my youngster-look due to the short trousers, long socks and occasionally a sailor-type blouse, the usual attire of the students of those years […]” (Jiménez, 1967: 248). His friendship with Lankester, which lasted throughout their lives, converted him into a lover of orchids, accompanying the great Englishman on many of his collecting trips. Jiménez had the privilege to grow up during a period when the botanical explora- tion of Costa Rica was in full effervescence. “By 1914, Costa Rica had become the center of scientific research in tropical America” (Evans 1999, p. 20). Jiménez knew the great botanists of his time: Wercklé, Pittier, Tonduz, the Brade brothers, Donnell Smith, Pittier, Britton, Dr. Patiño (Colombia), Wilson Popenoe, Maxon, Standley, Williams, and Allen, and married a daughter of Anasta- sio Alfaro. Louis O. Williams, who went on a few excur- sions with Jiménez, described him as “one of the most vivacious and enjoyable gentlemen (and botanists) to be met anywhere” (Williams 1972: 206). In 1915, in a let- ter to J. Barnhart, Pittier described him as follows: “… a disciple of Tonduz and a pharmacy student, who has already done a large amount of collecting and may yet surpass his master.” Silvia Troyo, a granddaughter of Otón Jiménez, wrote in a personal letter of September 2003: “Because of the ups and downs of the Museum after the depar- ture of Pittier, and because of his studies in Pharmacy, ‘Oto’ could not continue with the botany, as he wished. However, during the remaining years of his life, he dedicated to botany as much time as he could (which unfortunately was not much). After this period, since the collections at the Museum were not well organized and sometimes were lost, he started to send his collec- tions abroad, I believe for the rest of his lifetime. I know that in the process, many got lost, especially those des- tined to Europe… his later work, besides collecting and taxonomy, was oriented to the investigation of the nour- ishing properties of certain plants, or to the study of certain drugs.” Together with Lankester, he had to suffer Oakes Ames’ impatience: “When may I expect the specimens that Jimenez has in hand? Now is the time to get this material under the lens.” “Otón’ has not sent me a scrap. I think it will be wise if you remind him of my needs and accompany him to the post office with the package.” (1923 August 24 and December 18 letters from Oakes Ames to Charles H. Lankester). But it was not Ames but Schlechter and several other scientists who really valued Jiménez’ work, naming in his honor a series of new spe- cies: Epidendrum jimenezii Hágsater, Epilyna jimenezii Schltr., Habenaria jimenezii Schltr., Lepanthes jimenezii Schltr., Pachystele jimenezii Schltr., Scaphyglottis jimen- ezii Schltr., and Stelis jimenezii Schltr. In addition to being an excellent botanist, Jiménez was a prolific writer, who left interesting articles about Figure 15. Otón Jiménez (1895–1988). Courtesy of Silvia Troyo. 52 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Von Frantzius, Humboldt, Wercklé, Tonduz, Brade, and Lankester; an important bibliographical source for the study of scientific life in Costa Rica during the XIX and the first half of the XX century. “It is much to be regret- ted that the demands of business affairs have precluded a greater amount of personal fieldwork on the part of one who has such a keen perception of facts and the ability to discover them in strange places.” (Standley 1937: 53). TYPIFICATION OF COSTA RICAN ORCHIDACEAE DESCRIBED BY RUDOLF SCHLECHTER Species collectoribus variis lectae 1. Catasetum blepharochilum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 158. 1920 Type: Cuenca? [(Costa Rica.) “An Rio Toro Amarillo in dichten feuchten Wäldern in der Ebene. Costa Rica. 20 Dezbr. 1881” / “Dans les forets touffues et humides de la plaine sur le Rio Toro Amarillo, 20 Décb. 1881”], F. C. Lehmann 1061. Holotype, not found. Isotype, designated as lectotype by Romero and Jenny (1993), G 00168805! (Figure 16). Schlechter’s f loral analysis published in Mansfeld (1929: Pl. 56, No. 216!). Figure 17. Catasetum blepharochilum is seldom recorded among the orchids of Costa Rica, even as a synonym of the wide- spread C. maculatum, despite the type specimen having been collected along the Toro Amarillo River on the Car- ibbean plains east to the Central Volcanic Cordillera in Costa Rica. It was cited neither in Pupulin’s catalogue of Costa Rican Orchidaceae (Pupulin 2002) nor in Dressler’s treatment of Catasetum for the Manual de plantas de Cos- ta Rica (Dressler 2003). The reason for this was an error made by Schlechter (1920), who dubiously assigned the type specimen to “Cauca?” (Colombia) and treated the species as an Andean member of the genus. It is regarded as a Colombian conspecific with C. maculatum in Ulloa Ulloa et al. (2017), as well as in the major taxonomic data- bases available on the net [e.g., the Global Biodiversity Information Facility (Döring 2017), Tropicos 2021, WCSP 2021]. The type locality is, however, correctly cited as Cos- ta Rica by IPNI (2020). The isotype at G, that Romero and Jenny (1993) designated as the lectotype, bears two origi- nal labels by Lehmann, in German and French, which clearly state that the type specimen was collected in the Caribbean plains of Costa Rica in December 1881, a date that coincides with the visit of Lehmann to the country from December 1881 to May 1882. The analytical sketch prepared by Schlechter (in Mansfeld 1929) shows the oblong opening (or “mouth”) of the deeply saccate, conical lip, provided with delicate hairs along the lateral margins, which is typical of the species and distinguishes it from the similar C. integerri- mum Hook. The illustration in Hoehne’s (1945, p. 79, No. II) Flo- ra Brasilica, which extends the occurrence of C. blepha- rochilum to Brazil, is simply a rearrangement of Schlech- ter’s original sketch posthumously published in the series of his analytical drawings of new orchid species from the Andean countries, edited by Mansfeld in 1929. 2. Chondrorhyncha endresii Schltr., Repert. Sp. Nov. Regni Veg. 17: 14. 1921 Type: Costa Rica. “Ohne nähere Standortsangabe”, A. R. Endrés 166. Holotype, W 0018830!; drawings of the type and descriptions (W 0018831!); drawings of the flower and details (W 0018833!); Endrés’ original description of his collection no. 166 (W 0018832!); copy of Schlechter’s sketch of the holotype, with a drawing of the plant habit and analysis of the flower (AMES 00106743!) (Figure 18). This name is a synonym of Chondrorhyncha bicolor Rolfe [≡ Chondroscaphe bicolor (Rolfe) Dressler] (Pupu- lin et al. 2009), a concept based on a Costa Rican collec- tion by Richard Pfau (Type: Costa Rica. Without specific locality, R. Pfau s.n., K). For other synonyms of the spe- cies see Pupulin (2010b). Dressler (2001, p. 47) consid- ered C. bicolor a “lost species,” not corresponding to any other Central American species of Chondroscaphe, but several of the anomalous features of this species noted in the protologue are attributable to Rolfe’s interpreta- tion of the poorly preserved type specimen, which is indistinguishable from Costa Rican populations treated as C. endresii (Pupulin 2010b). When Schlechter (1921a) described the forgotten collection kept in Reichenbach’s herbarium, together with Endrés’ drawings of the plant habit, the flower, and floral details, he did not suspect that it corresponded with the schematic description of C. bicolor provided by Rolfe. The shape of the lip, with its oblong, thick, apically bilobed callus, which Schlech- ter used to characterize C. endresii, is nonetheless identi- cal to that of C. bicolor, and the drawing of the rostel- lum of this species, made by Endrés, clearly illustrates the characteristically ligulate, abruptly introrse stig- matic arms that are typical of C. bicolor (Pupulin et al. 2009). Among the materials referable to the type at W is a manuscript name by Reichenbach, who intended to publish the species with the name “Chondrorhyncha umbonata”, and the name “umbonata”, in Reichenbach’s handwriting, is noted on an envelope mounted on the type sheet (Pupulin et al. 2011, 2013). 53Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Figure 16. Lectotype of Catasetum blepharochilum (G 00168805). Reproduced with the kind permission of the Director, Conservatoire et Jardin botaniques de la Ville de Genève. 54 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín 3. Chondrorhyncha reichenbachiana Schltr., Repert. Spec. Nov. Regni Veg. 17: 15. 1921 Type: Costa Rica. “Cataratas, blühend Marz-August”, A.R. Endrés 557. Holotype, W 0018829 / Rchb.Orch. 49753!; Endrés’ drawings of the type and descriptions, W 0018826 / Rchb.Orch. 28550!; floral analysis of the type, originally prepared by Schlechter, published by Mansfeld (1931: Pl. 63, No. 252!); tracing of Schlech- ter’s sketch of the holotype, with drawing of the plant habit and analysis of the flower, AMES 00106751! (Fig- ure 19). Among the materials that Endrés sent to Reichen- bach from Costa Rica was a specimen of an unknown “Zygopetalum” with a characteristic dark grey-green foli- age and a solitary flower born above the pendent leaves. Reichenbach knew it was a new species, and he anno- tated the correspondent sheet with the intended name of “Chondrorhyncha lamellata”, in allusion to the lamellate callus of the lip. He also used Endrés’ accurate sketches to prepare two botanical plates for his Xenia Orchidacea (published between 1858 and 1874, and then continued by Fritz Kränzlin until 1900), with the intended names “Chondrorhyncha lamellata” and “Zygopetalum lamel- latum”. However, they were never published, and like many other discoveries by Endrés, remained hidden in Reichenbach’s herbarium after his death in 1889 (Pupu- lin 2009). It was only around 1915, just a few months before the beginning of the First World War’s hostilities, when Schlechter visited the recently opened orchid col- lection at the Hof Imperial Museum in Vienna. Here, he found the plant collected 40 years before by Endrés, describing it in 1921 as Chondrorhyncha reichenbachiana in honour of his great predecessor. The large callus that occupies the whole centre of the lip from side to side, noted by Schlechter (1921) in the protologue and his drawing of the type, is diagnos- tic of the species, which has no close relatives in Cen- tral America. The name is the basionym of Benzingia reichenbachiana (Schltr.) Dressler. Figure 17. Schlechter’s floral analysis of Catasetum blepharochilum (in Masfeld 1929: Pl. 56, No. 216). Figure 18. Chondrorhyncha endresii, plant habit and analysis of the flower, traced from Schlechter’s sketch of the holotype (AMES 00106743). Courtesy of the Harvard University Herbaria, repro- duced with permission of the President and Fellows of Harvard College. 55Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum 4. Endresiella zahlbruckneriana Schltr., Repert. Spec. Nov. Regni Veg. 17: 14. 1921 Type: “Costa Rica. an der Strasse von San Ramon nach San Carlos, Legua. Blühend im September”, A.R. Endrés 512. Holotype, W 0019449 / Rchb.f. Orch. 43634!; sketches of the type specimen by Endrés, W 0020711 / Rchb.Orch. 37186!; copy of Schlechter’s sketch of the holotype (largely traced on Endrés’ drawings), with plant habit and analysis of the flower, prepared under his supervision, AMES 0099111! (Figure 20). Schlechter created the genus Endresiella in 1921 to accommodate a species with the habit similar to a small Stanhopea Frost ex Hook., and flowers similar to those of the genus Schlimmia Planch. & Linden ex Linden, but smaller. He dedicated the “very excellent new orchid genus to the well-known researcher of the orchid flora of Costa Rica, Endres”, noting that his vast collections, together with “marvelously executed drawings”, were still largely unpublished in the herbarium of Reichen- bach (Schlechter 1921). Schlechter found the imperfect specimen in Reichenbach’s Herbarium filed under the genus Sievekingia Rchb.f., fortunately, accompanied by excellent and detailed drawings (reproduced in Ossen- bach et al. 2013: 316, Figure 316). He dedicated the type species to the Austrian botanist Dr. Alexander Zahl- bruckner (1860–1938), curator of the herbarium at the Naturhistorisches Museum in Vienna, then director of the museum’s botany department. The densely packed inflorescence, the white flowers with yellow mesochile and the lateral sepals connate to the middle distinguish the species. This name is the bas- ionym of Trevoria zahlbruckneriana (Schltr.) Garay. 5. Epidendrum cardiophorum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 9(208–210): 214. 1911. Type: Costa Rica in dem Wäldern von Tsaki, Talamanca, ca. 200 m, blühend im April 1895, H. Pittier [s.n., (Herb. Figure 19. Tracings of Schlechter’s original sketch from the holo- type of Chondrorhyncha reichenbachiana (AMES 00106751). Cour- tesy of the Harvard University Herbaria, reproduced with permis- sion of the President and Fellows of Harvard College. Figure 20. Copy of Schlechter’s sketch of Endresiella zahlbruckne- riana holotype, prepared under his supervision (AMES 99111). Courtesy of the Harvard University Herbaria, reproduced with per- mission of the President and Fellows of Harvard College. 56 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Figure 21. Isolectotype of Epidendrum cardiophorum (BR 00000657435). Courtesy of the Meise Botanic Garden Herbarium. 57Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum instit. physicgeogr. nat. costaricensis, IFGN)] 9519. Holo- type, B, destroyed; lectotype, designated by Pupulin et al. (2016): CR 9519!; isolectotypes: BR 00000657435! (Figure 21); G 00168668!; US 815035 / 00316361!; copy of Schlechter’s drawing of the holotype, made under Schlechter’s supervision, AMES (HUH-00070175!). Pupulin et al. (2016) selected an isotype at CR, which is a complete and fertile specimen in excellent condition, as lectotype (Pupulin et al. 2016  : 289, Figure 17A). As Pupulin et al. (2016) noted, the quote of Pittier 9519 in the protologue and on the copy of Schlechter’s drawing of the type represents a misunderstanding of the numera- tion system used at the IFGN. The rhizomatous habit with scandent rhizome and stems produced far apart from each other, the ancipitous, short inflorescence, and the small flowers with the part of the perianth less than 1 cm long distinguish this species. Epidendrum cardiopho- rum ranges from Mexico to the northern Andes. 6. Epidendrum dolichostachyum Schltr., Repert. Spec. Nov. Regni Veg. 3: 79. 1906 Type: Costa Rica. [San José:] Bei La Palma [1550 m]; blühend im Sep 1896, H. Pittier (10311 Herb. Insti- tut. Costaric. [Herb. Nac. Costa Rica]). Holotype, B, destroyed [tracing of Schlechter’s drawing of the hol- otype, AMES 00070288! (Figure 22)]. Isotypes: BR 0000006574550!, designated here as the lectotype (Figure 23) (drawing of a flower, AMO, not seen; floral analysis from the type, prepared by Eric Hágsater, CR, two sheets with the same drawings); isolectotype: US, not seen; flo- ral analysis from the holotype, reproduced in Mansfeld (1931: Pl. 49, No. 194!). Atwood (1989) indicated that the “holotype” of Epi- dendrum dolichostachyum is conserved at CR, but we have not located it. According to Lobo (2003), it was probably never deposited at this herbarium. We found two sheets with copies of a floral analysis of the type of E. dolichostachyum prepared by Hágsater; these were probably the materials examined by Atwood (1989). According to the protologue, E. dolichostachyum is morphologically similar to E. laucheanum Rolfe, both having narrowly lanceolate, acuminate leaves, a termi- nal, pendent, many-f lowered, racemose inf lorescence with ancipitous peduncle, flowers of similar dimensions, with sepals and petals ocher-brown to purplish brown and lime-colored lip, the lip cordate to reniform with incurved sides and recurved at the apex, and the disc with a fleshy costa. Rolfe (1893) described the lip of E. laucheanum as having entire margins, while Schlechter (1906b) characterized the lip of E. dolichostachyum with subcrenulate margins. However, the flowers on the holo- type specimen of E. laucheanum (K-000463483) show a lip with minutely erose margins, as previously reported by Santiago and Hágsater (2010). We did not find evi- dence to separate these taxa, and therefore consider them as conspecific in agreement with Atwood (1989). A specimen collected by A. Tonduz [10388 Herb. Inst. Fis.-Geogr. Nac. (BR 0000006573195)] in December 1986, in Alto de Ochomogo (Cartago), Costa Rica, car- ries an envelope with the annotation: “H. Pittier 10311 (BR 0000006601348) Epidendrum dolichostachyum” (BR- 0000006573195), which coincides with the type collec- tion number of E. dolichostachyum. Unfortunately, we could not verify that the material contained in this enve- lope is part of the type of E. dolichostachyum. Figure 22. Epidendrum dolichostachyum, tracing of Schlechter’s drawing of the holotype (AMES 70288). Courtesy of the Harvard University Herbaria, reproduced with permission of the President and Fellows of Harvard College. 58 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Figure 23. Lectotype of Epidendrum dolichostachyum (BR 0000006574550). Courtesy of the Meise Botanic Garden Herbarium. 59Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum 7. Epidendrum hoffmannii Schltr., Repert. Spec. Nov. Regni Veg. 16: 444. 1920 Type: Costa Rica, [San José:] Curidabal [Curridabat], Mai 1857, C. Hoffmann 570. Holotype, W-Rchb.Orch. 51054! (Figure 24); tracing of Schlechter’s drawing of the holotype, AMES 70416! (Figure 25). The comprehensive work by Pupulin and Karremans (2007) revealed a series of details about the history of E. hoffmannii that illustrates the taxonomical conundrum in which this species is involved. During his expeditions across Costa Rica, the German physician Karl Hoffmann Brehmer collected two Epidendrum plants from the area of “Curidabal” [Curridabat], just a few kilometers out- side of the capital city of San José. Deliberately or not, one of the plants was assigned with the collection num- ber Hoffmann 570, while the other was left sine numero. The plants were sent to Heinrich Gustav Reichenbach, professor of botany and director of the Botanic Gar- dens at Hamburg University, who probably considered the individuals as belonging to the same species, as he described only one species under the name Epiden- drum ionophlebium, based on Hoffmann sine numero (Reichenbach 1866). The other plant, under field num- ber Hoffmann 570, was left to oblivion, until Schlechter had access to the materials sometime around 1915. His eye captured subtle differences between the two indi- viduals and described a new species, Epidendrum hoff- mannii, based on Hoffmann 570 (Schlechter 1920). Fur- thermore, the analysis carried out by Pupulin and Kar- remans (2007) also revealed that the specimen collected by Hoffmann under his field number 570 and saved at W was mistakenly annotated as E. ionophlebium, when it is actually the holotype of E. hoffmannii. Apparently, “this specimen is not annotated in Reichenbach’s hand- writing, and the identification as ‘Epidendrum ionophle- bium’ was affixed to it when the specimen was mounted in Vienna” (Pupulin and Karremans 2007: 456). While studying Reichenbach’s materials, Schlechter also prepared illustrations of the two Epidendrum. Both of these original drawings were destroyed during WWII, but copies prepared under Schlechter’s supervision are saved at AMES. This illustration of E. hoffmannii dis- tinctly shows a slender plant with ovoid pseudobulbs and narrowly elliptic leaves bearing a short inflorescence with two flowers, characters also seen in the dried speci- men. The dissection of the flower displays lanceolate sepals, elliptic petals with acuminate apices, and a lip with a broadly ovate lamina and acuminate apex. These characters, along with the ornate, velvety lip described in the protologue, largely agree with the concept of the widely distributed Prosthechea chacaoensis (Rchb.f.) W.E.Higgins and is considered a synonym of the latter. 8. Epidendrum insulanum Schltr., Beih. Bot. Centralbl., Abt. 2 36(2): 404. 1918 Type: Costa Rica, [Puntarenas:] Cordon littoral à Wafer Bay, Cocos Island (Pacific Ocean), Jan 1902, H. Pittier (16350 Herb. Nac. Costa Rica). Holotype, B, destroyed [Schlechter’s drawing of the holotype, reproduced in Mansfeld, 1931: Pl. 50, No. 199!; tracing at AMES (HUH 70447!) (Figure 26)]. Isotypes: GR 3580!, selected by Trusty and Blanco (2005) as the lectotype, AMES 73449!; GR 3579! (AMES 73450 / HUH-70446), isolec- totype, fruiting (Figure 27); GR 3581! (AMES 73448 / HUH 70445), isolectotype, sterile; CR 16350! (2 sheets), isolectotypes. Epidendrum insulanum is endemic to Cocos Island, an Oceanic Island situated more than 500 kilometers from the nearest continental point at Cabo Blanco, Península de Nicoya, Costa Rica. On the Island, the species is a common epiphyte in premontane rainfor- est, where the plants grow on exposed or shady con- ditions in both shrubs and trees, intermixed with E. cocoense (Bogarín et al. 2011). The copy of Schlechter drawing of the holotype well illustrates the single-flow- ered inflorescences, the erose apex of column lacking the two teeth present in other species of the Epiden- drum ramosum group, the lateral lobes of the lip that do not cover the apex of column in lateral view, and the triangular callus that runs through the entire lip to form an apical mucro, which are described in the protologue (Schlechter 1918a) and are diagnostic of the species. A modern botanical illustration of E. insu- lanum, based on a living specimen from the island, is provided in Bogarín et al. (2011). Schlechter compared it with E. repens Cogn., a species of broad distribution from Mexico to Venezuela and Colombia, and down to Bolivia along the Andes, which also belongs to the Epi- dendrum ramosum group. 9. Epidendrum paucifolium Schltr., Repert. Spec. Nov. Regni Veg. 3: 248–249. 1907 Type: Costa Rica. Bei Cuera de Tigre, blühened im Jan- uar 1897, H. Pittier 10515. Holotype, B, destroyed. Iso- types: BR 0000006573546!, selected by Santiago and Hágsater (2008) as lectotype (Figure 28); isolectotype: M-0226680!. 60 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Figure 24. Holotype of Epidendrum hoffmannii (W-Rchb. Orch. 51054). Courtesy of the Naturhistorisches Museum Wien. 61Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum According to the protologue and type material, Epi- dendrum paucifolium can be distinguished by the com- bination of non-pseudobulbous stems, 1–2 apical, erect, leaves, oblong to elliptic leaf blades; the apical inflores- cence, with ancipitous peduncle, longer than rachis, with 2 tubular, acuminate bracts approximately the same length as the internodes, ancipitous rachis, with concave, lanceo- late, perfoliate, acuminate floral bracts, longer than ovary; few-flowered, with 3–4 flowers opened simultaneously, clustered near the inflorescence apex, the slightly extend- ed oblong, acute sepals, recurved margins, obliquely sub- spatulate, obtuse petals, lip with the ovate, cordate, obtuse, short apiculate, blade; disc with 3 vertical keels extended to near the apex of the lip; the apex of the column with a pair of prominent digitate teeth on the back, and clinan- drium with denticulate margins (Schlechter 1907a). It also has fuchsia or magenta flowers with a column basally white (Santiago and Hágsater 2008). Epidendrum paucifo- lium ranges from Costa Rica to the western Panama. We were unfortunately unable to locate the type locality, “Cuera de Tigre” (or “Cuero de Tigre”) on a modern Costa Rican map. 10. Epidendrum selaginella Schltr., Repert. Spec. Nov. Regni Veg. 3: 48. 1906 Type: Costa Rica. [San José:] An feuchten Felsen auf dem Recreo, am Wege von Carillo, c. 1200 m, blühend im Juli 1888, J. Cooper 523. Holotype, B. destroyed [tracing of Schlechter’s drawing of the holotype, AMES 00070862! (Figure 29)]. Isotype: US 579506 / barcode 00093842!, designated here as the lectotype (Figure 30); floral analy- sis from the holotype reproduced in Mansfeld (1931: Pl. 57, No. 225!). Epidendrum selaginella belongs to the Epidanthus Group characterized by flat leaves, a tiny ligule opposite to the blade, entire lip, and the anther with four poles. The species is distinguished by having thin and apically laterally compressed stems, oblong to ovate, emargin- ate leaves, flowers congested in the apical third of the inflorescence, papillose ovary, abaxially papillose sepals, and unguiculate lip, with the subdeltate, subcordate and obtuse, blade without a keel. Epidendrum selaginella ranges from Costa Rica to central Panama. Figure 25. Tracing of Schlechter’s drawing from the holotype of Epidendrum hoffmannii (AMES 70416). Courtesy of the Harvard University Herbaria, reproduced with permission of the President and Fellows of Harvard College. Figure 26. Epidendrum insulanum, tracing of Schlechter’s drawing of the holotype at AMES (HUH 70447). Courtesy of the Harvard University Herbaria, reproduced with permission of the President and Fellows of Harvard College. 62 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín Figure 27. Isolectotype of Epidendrum insulanum [GR 3581 (AMES 73448)]. Courtesy of the Harvard University Herbaria, reproduced with permission of the President and Fellows of Harvard College. 63Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Figure 28. Lectotype of Epidendrum paucifolium (BR 0000006573546). Courtesy of the Meise Botanic Garden Herbarium. 64 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín The sketch of the holotype prepared under Schlech- ter’s supervision includes the plant habit and a floral analysis (Figure 29), the latter reproduced by Mans- feld (1931), clearly showing the diagnostic characters of E. selaginella. Three floral drawings that do not belong to the type collection but are instead associated with a collection by Maxon (no. 467) are placed on the lower- left corner of the sheet of the holotype drawing (AMES 00070862). Schlechter (1906a) stated that the vegeta- tive appearance of this species resembles Selaginella P. Beauv., a genus of lithophytic plants, hence its specific epithet. 11. Epidendrum tenuiflorum Schltr., Repert. Spec. Nov. Regni Veg. 3: 49. 1906 Type: Costa-Rica. [Cartago:] Bei Aguacaliente, ca. 1300 m; blühend am 2 Jan 1888, H. Pittier 38. Holotype, B, destroyed; tracing of Schlechter’s drawing of the holo- type, AMES 24105 / barcode 00070928!, selected by Mora and Atwood (1992) as lectotype (Figure 31); Schlechter’s floral analysis reproduced in Mansfeld (1931: Pl. 58, No. 229!). Mora and Atwood (1992, t. 1457) designated the tracings of Schlechter’s drawing of the holotype (AMES 24105) as the lectotype (cited originally as “type”). This is regarded as a formal lectotypification because the authors indicated by direct citation the term “type” (Art. 7.11) and specified the herbarium where the specimen is kept (Art. 40 note 1). Also, before 1 January 2001, it was not mandatory to include the typification statement phrase “designated here” (hic designatus) or an equiva- lent (Art. 7.11) and “lectotypus”, its abbreviation, or its equivalent in a modern language (Art. 9.23) (Turland et al. 2018). Thus, the lectotypification proposed by Santia- go and Hágsater (2006) is a superfluous type designation (Art. 10.5). Together with the floral analysis published by Mansfeld (1931), this drawing is the only copy of the original material associated with the protologue of this species. It also includes a sketch of the plant habit that was not included in Mansfeld’s compilation. The draw- ing shows a combination of diagnostic characters con- sistent with the protologue of E. tenuiflorum (Schlechter 1906a), such as the linear leaves and short inflorescence, the trilobed lip provided with short, lanceolate, acute lateral lobes, and a broadly obcuneate, deeply bilobed middle lobe with a tiny apicule, the elongate, shallowly trilobed clinandrium exceeding the column length, with minute lateral lobes and a widely ovate, apiculate mid- dle lobe. It is noteworthy that the lip’s lateral lobes were drawn recurved when they are incurved in living flow- ers, but this is probably because rehydrated material was used to prepare the sketches. Schlechter (1906a) suggested that E. tenuiflorum is morphologically similar to Epidendrum centropetalum Rchb.f. (Reichenbach 1852) but differs from the latter by the mostly trilobed clinandrium. However, the two taxa are indistinguishable when comparing the protologues. Therefore, Santiago and Hágsater (2006) consider them conspecific. 12. Epidendrum urostachyum Schltr., Beih. Bot. Cen- tralbl., Abt. 2. 36(3): 409-410. 1918 Type: Costa Rica. El Tablazo, près San José, 1900 m, Sept 1913, E. Jiménez s.n. (n. herb. Nac. Costa Rica 17651). Holotype, B, destroyed; traces of the original drawing of the holotype made under Schlechter’s supervision, an envelope putatively containing fragments of the holo- type, and flowers saved in glycerin, AMES 82254 / bar- code 00070965, designated here as the lectotype (Figure 32). Schlechter’s drawing of the holotype published in Mansfeld (1931: Pl. 58, No. 231!). Figure 29. Tracing of Schlechter’s drawing from the holotype of Epidendrum selaginella (AMES 00070862). Courtesy of the Harvard University Herbaria, reproduced with permission of the President and Fellows of Harvard College. 65Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Figure 30. Lectotype of Epidendrum selaginella (US 579506). Courtesy of the United States National Herbarium, Smithsonian Institution. 66 Franco Pupulin, Isler F. Chinchilla, Gustavo Rojas-Alvarado, Melania Fernández, Carlos Ossenbach, Diego Bogarín The herbarium specimen AMES 82254 is a mixed collection, comprising the tracings of the holotype draw- ing made under Schlechter’s supervision, an envelope supposedly containing “fragments of holotype”, and flowers preserved in a separate glycerin collection identi- fied with the same accession number. Since restrictions associated to the COVID-19 pandemic limited our possi- bilities to confirm and study the contents in the envelope and the glycerin material, we are basing our selection of the lectotype on the tracings of the holotype drawing. The tracings show a slender plant of narrow leaves and hanging inflorescence, which bears flowers with oblong sepals, oblique petals and orbiculate lips. The base of the lip is adnate to the column, while the lamina shows wavy margins and a callus extending to the middle. Schlechter distinguished E. urostachyum from the two morphologically similar species E. laucheanum and E. dolichostachyum by the smaller flowers and the shape of the lip (Schlechter 1918a). Later, several authors have considered the concept described by Schlechter as Epi- dendrum urostachyum under the synonymy of Epiden- drum laucheanum (Pupulin 2002, Hágsater 2010, Boga- rín et al. 2014), a variable species first found in Popayán, Colombia (Rolfe 1893). Epidendrum laucheanum is rec- ognized by a long, hanging inflorescence that arches towards the floor, with ocher-brown flowers and a green to orange or purple lip (Dressler 2003), which largely agrees with the original description and tracings of E. urostachyum. 13. Gongora unicolor Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 299. 1923 Syntypes: Costa Rica. Ohne nähere Standortsang- abe (kultiviert im Garten von Mr. C.W. Powell, Pana- ma), C. H. Lankester s.n. (B, destroyed). Costa Rica. [Limón:] Las Mercedes, Ebene von Limon, Nov 1921, F. Nevermann s.n. (B, destroyed); Schlechter ś floral anal- ysis of the holotype, reproduced in Mansfeld (1931: Pl. 62, No. 248!), designated here as lectotype (Figure 33). Figure 31. Lectotype of Epidendrum tenuiflorum (AMES 24105). Courtesy of the Harvard University Herbaria, reproduced with per- mission of the President and Fellows of Harvard College. Figure 32. Lectotype of Epidendrum urostachyum (AMES 82254). Courtesy of the Harvard University Herbaria, reproduced with per- mission of the President and Fellows of Harvard College. 67Typification of Costa Rican Orchidaceae described by Rudolf Schlechter. Species variorum collectorum Epitype, designated here (Figures 34–35): Costa Rica. Heredia: Sarapiquí, Horquetas, road to Rara Avis, devi- ation point to El Manú, ca. 1 km, 10º19’40” N 83º58’29” W, 120–170 m, tropical rain forest, secondary vegeta- tion and scattered trees along a small river, 27 Septem- ber 2003. Flowered in cultivation at Lankester Botani- cal Garden, 16 May 2014, F. Pupulin, H. León-Páez, C. Ossenbach & B. Arias 4954 (JBL-spirit D0992!); isoepi- type: JBL-spirit D0153!. When publishing Gongora unicolor, Schlechter cit- ed two specimens from Costa Rica. One of them was collected by Charles Lankester with no specific local- ity and cultivated by Powell in Panama. The second one was found by Ferdinand Nevermann, who collected the specimen in the plains of Limón in an area called “Las Mercedes” in the lowlands of the Caribbean watershed. At present, this locality refers to the town of Hambur- go de la Rita, Pococi, Limon, at approximately 50 m in elevation. In the original description, Schlechter failed to declare which of these materials was chosen as the type specimen, and the two specimens must be considered syntypes of G. unicolor. Unfortunately, both specimens were lost after the destruction of the Berlin herbarium, and no isosyntypes or other type materials are known. In his “Monograph of the genus Gongora”, Jenny (1993) cited the drawing reproduced in Mansfeld as an icono- type, but this term is not recognized by the International Code of Nomenclature. In the absence of other materials that can serve for lectotypification, the analytical sketch prepared by Schlechter and reproduced in Mansfeld (1931) is chosen as lectotype. Gongora unicolor is a com- plex species difficult to identify from herbarium materi- als as the main differences are based on flower color and scent. The immaculate flesh-colored to pale tan flowers with a particular strong scent of either fresh cornmeal (Dressler 1966, 2003) or “unpleasant odour” (Atwood 1987) are the main identifying characteristics of this species. Although not diagnostic, other characters as the distinctively concave base of the lip and the presence of a narrow groove running dorsally from near the base of the lip, are useful to distinguish this species in herbari- um material. The lectotype of G. unicolor only shows some floral characters and is taxonomically ambiguous, as it does not allow unequivocal interpretation of the features which are diagnostic of this taxon. Therefore, to favor the interpretation of the lectotype in accordance with Art. 9.9 of the International Code (Turland et al., 2018), we designated as epitype a specimen that was collected in the vicinity of the type locality of Gongora unicolor. 14. Habenaria jimenezii Schltr., Beih. Bot. Centralbl., Abt. 2. 36(2): 372. 1918 Type: Costa Rica: Río Virilla, Nov. 1912, O. Jiménez 631. Holotype: B, destroyed; copy of Schlechter’s sketch of the holotype, with a drawing of the plant habit and anal