A new nothogenus and nothospecies in the Zygopetalinae (Orchidaceae), with a note on euglossine pollination in the subtribe

33 A new nothogenus and nothospecies in the Zygopetalinae (Orchidaceae), with a note on euglossine pollination in the subtribe Franco Pupulin1,2,3 , Adam P. Karremans1, Elvira Salas Hidalgo4 1 Lankester Botanical Garden, University of Costa Rica, P.O.Box 302-7050 Cartago, Costa Rica 2 Harvard University Herbaria, Cambridge, MA, USA 3 The Marie Selby Botanical Gardens, Sarasota, FL, USA 4 Department of Biochemistry, School of Medicine, University of Costa Rica, San José, Costa Rica Corresponding author: Franco Pupulin ([email protected]) Copyright: © Franco Pupulin et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Natural hybridization in Neotropical orchids is summarized, and the frequency of nothogenera in the subtribe Zygopetalinae is highlighted. A new nothogenus discovered in Costa Rica, formed by the cross between species of Benzingia and Chondroscaphe, is described, and the new nothospecies ×Benzoscaphe stelleri is described and illustrated. Putative parents, as well as their derivative hybrid, are compared through composite digital plates. Cues are offered about the pollination mechanisms of the putative parents, based on the morphology of the flowers and pollinaria, which possibly led to the formation of the natural cross between different genera. Key words: Benzingia pollinaria, ×Benzoscaphe, Chondroscaphe pollinaria, natural hybridization, orchid hybrid pollinarium Introduction Hybrid taxa formed by a natural cross between species of the same genus have been recorded in several orchid groups in the Neotropics. Individuals belonging to a nothotaxon are not easy to detect, and their presence is usually documented on the basis of unique combinations in the morphology of plants and flowers, which present intermediate traits between already known sympatric species, whether in the same or in different genera. Nothospecies are known in subfamily Cypripedioideae (Cypripedium: Szlachetko et al. 2017; Phragmipedium: Pupulin and Díaz-Morales 2018; Díaz-Morales and Pupulin 2019), subfamily Vanilloideae (Vanilla: Pansarin 2025), and in many subtribes in the Epidendroideae. Among them, natural hybrids have been documented in Catasetinae (Catasetum: Cantuária et al. 2021; Romero-González and Carnevali 2024; Krahl et al. 2024), Dendrobiinae (Bulbophyllum: Borba and Semir 1998; Azevedo et al. 2006; Fiorini et al. 2023), Cyrtopodiinae (Cyrtopodium: Batista and Medeiros 2022), Laeliinae (Cattleya: Braem 1995; Castro Neto and Catharino 2004; van den Berg 2014; Barrera-Rojas and van den Berg 2025; Encyclia: Dressler and Pollard 1974; Sauleda and Adams 1990; Pérez-García and Hágsater 2012; Sauleda and Esperón 2016, 2019; León-Peralta et al. 2023; Epidendrum: Pupulin 2003; Pinheiro et al. 2010; Academic editor: Vincent Droissart Received: 22 May 2025 Accepted: 13 August 2025 Published: 7 November 2025 Citation: Pupulin F, Karremans AP, Salas Hidalgo E (2025) A new nothogenus and nothospecies in the Zygopetalinae (Orchidaceae), with a note on euglossine pollination in the subtribe. PhytoKeys 266: 33–52. https://doi.org/10.3897/ phytokeys.266.159892 PhytoKeys 266: 33–52 (2025) DOI: 10.3897/phytokeys.266.159892 34 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Bogarín et al. 2011; Laelia: Salazar et al. 2014; Solano et al. 2019; Cetzal-Ix et al. 2020; Prosthechea: Mó et al. 2014; Tetramicra: Soto Calvo et al. 2019), Oncidiinae (Cohniella: Cetzal-Ix et al. 2013; Ionopsis: Krahl et al. 2021; Lophiaris: Cetzal-Ix et al. 2018), Pleurothallidinae (Dracula: Zambrano and Solano-Gómez 2011; Masdevallia: Lipińska et al. 2019; Archila Morales et al. 2021; Pleurothallis: Pupulin et al. 2021), and Stanhopeinae (Gongora: Jiménez-Vázquez et al. 2023). Less frequent are hybrid taxa formed between species belonging to different genera. Among the Oncidiinae, natural hybrids between Comparettia and Rodriguezia were reported from Colombia (Sauleda 2015; Sauleda and Arrondeau 2020), while Cetzal-Ix et al. (2012) described the new nothogenus ×Cohnlophiaris, resulting from the cross between species of Cohniella and Lophiaris. However, according to alternative generic circumscriptions (i.e., Neubig et al. 2012), the latter nothogenus is considered a synonym of Trichocentrum. Chiron and Castro Neto (2002) described the hybrid genus ×Hoffmanncattleya, which includes natural hybrids between the genera Cattleya and Hoffmannseggella (Castro Neto and Menezes 2007). However, most authors consider the two genera synonymous. Compared to the paucity of recorded intergeneric hybrids among other orchid groups, the Zygopetalinae appears particularly interesting, with at least four nothospecies recorded in three natural nothogenera: ×Bensteinia Christenson (Christenson 2006; Pupulin 2007), ×Cochlezella J.M.H.Shaw (Pupulin 2015), and ×Pescatoscaphe Christenson (Christenson 2006). Here, we record another nothogenus in the subtribe, ×Benzoscaphe, resulting from the cross between Benzingia reichenbachiana (Schltr.) Dressler and a species of Chondroscaphe (Dressler) Senghas & G. Gerlach, which we hypothesize to be C. bicolor (Rolfe) Dressler. Interestingly, the hybrid between Benzingia Dodson and Chondroscaphe has never been produced artificially. The first genus has been hybridized with Kefersteinia Rchb.f. to produce ×Bensteinia, with two natural nothospecies occurring in Ecuador and Costa Rica (Christenson 2006; Pupulin 2007), and with Cochleanthes Raf. plus Kefersteinia to create the hybrid swarm Goldnerara (Royal Horticultural Society 2017). Chondroscaphe has been artificially crossed with Pescatoria Rchb.f. (= ×Pescatoscaphe, which today also includes the hybrids of the synonymous Bollea Lindl., previously treated as ×Bolleoscaphe), Cochleanthes (= ×Cochloscaphe), Warscewiczella Rchb.f. (= ×Warczewscaphe), and Zygopetalum Hook. (= ×Zygoscaphe) (Royal Horticultural Society 2017). When the astonishing size of the Orchidaceae is taken into account, likely exceeding 30,000 species (Kindlmann et al. 2023), the number of recorded naturally occurring hybrids is remarkably low. Nevertheless, multiple factors contribute to the paucity of nothotaxa reported to date. Orchids possess a complex system of barriers, both abiotic and biotic, to prevent pollen from being deposited on the female reproductive organs of the “wrong” species. Among the abiotic barriers, the most significant are undoubtedly the geographic isolation between species whose reproductive systems are mechanically compatible and the temporal barrier resulting from a chronological shift in the phenology of morphologically related species. The latter scenario is especially true in groups whose flowers are “generalist” in attracting pollinators, as is the case, for example, in the genus Sobralia and other groups characterized by ephemeral flowers and synchronous flowering. If these barriers are disrupted—as sometimes happens due to anthropogenic transformation of the natural landscape—other factors related to the biology, morphology, mechanics, and chemistry of flowers come into play to reduce 35 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) the possibility of erroneous gamete exchange and to ensure that the genetic coherence of the species is maintained. Species-specific attractive signals, precise shape and size of floral organs to limit interaction exclusively to the correct pollinator, highly specific positioning of the pollinarium, and its morphological and structural transformation after removal aimed at precisely orienting the pollen toward the stigmatic cavity are some of the functional strategies that orchids employ to prevent erroneous sexual crossing (Tremblay et al. 2005; Ackerman et al. 2023). When these are also thwarted by an error in the pollen vector, other chemical barriers are put in place to maintain species identity, such as incompatible signals that compromise the activation of pollen tubes or, ultimately, genetic incompatibility between sperm cells and the ovules that should be fertilized. Considering the large number and complex nature of the barriers that prevent interbreeding between different species, it is notable that they can be evaded in groups belonging to all the subfamilies of Orchidaceae recorded in the Neotropics. Documenting a naturally occurring interspecific or, even more so, intergeneric hybrid represents an event with infinitesimal statistical probability. In the field, this means encountering a hybrid individual that vegetatively closely resembles one or both of its parents, likely a unique individual that—due to its unique floral morphology—may not have left any offspring, collecting that individual entirely by chance, cultivating it to flower, and having prior knowledge of the putative species that may have generated it, allowing comparison and thus advancing a scientific hypothesis regarding its hybrid nature. Seen in this way, it is perhaps not too speculative to suppose that the relatively few nothospecies documented so far represent nothing more than the tip of an iceberg of hybrid individuals that coexist in nature with their ancestors. From an evolutionary perspective, the distinction between F1, subsequent filial generations, as well as backcrosses and combinations of these could be of potential interest, even though for the International Code of Nomenclature for algae, fungi, and plants (Turland et al. 2025), a nothotaxon includes all individuals derived from the crossing of representatives of the stated parent taxa (Art. H.11.1). Studying the role of natural hybridization in orchid evolution has certainly been hampered to date by the limited number of individuals with diagnosable morphological characteristics that allow their identification as such. More generally, the sporadic introduction of new genetic combinations into a population could lead to increased genetic variability, while repeated backcrossing could result in polyploidy and favor the incorporation of genes that accelerate adaptation. During the systematic documentation of Costa Rican species of Orchidaceae, mostly intended for the treatment of the family for Flora Costaricensis (Atwood and Mora-Retana 1999; Pupulin 2010), we have received considerable help from a large group of resident naturalists and plant enthusiasts, who share with the personnel of Lankester Botanical Garden their most unusual findings. This kind of cooperation, which in the recent history of our research center has made possible the discovery and scientific description of several rare and rarely seen orchid species for the flora of Costa Rica, is a true example of what citizen science can accomplish when a local scientific institution opens its doors to the contributions of trained naturalists. In the specific case of the new nothospecies described in this paper, one of the plants from a collection made on the southeastern slopes of the Central Volcanic Cordillera of Costa Rica was grown and flowered by Norman Steller, a keen orchid grower who collected the plant together with staff of Lankester Botanical Garden and brought it to our research department for documentation and study. 36 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Material and methods The plants used for this study were collected during routine field activities carried out by the scientific and horticultural staff of Lankester Botanical Garden (JBL, by its acronym in Spanish) under the scientific permit R-SINAC-SE-DT_PI-005-2024 issued by the Costa Rican Ministry of Environment and its National System of Protected Areas (SINAC, by its acronym in Spanish). The plants were cultivated in the research greenhouses of JBL and in private greenhouses until flowering, for documentation and study. Macro photographs were taken with Nikon D850 and D810 cameras fitted with a Micro-Nikkor AF 105 mm lens. Photomicrographs were taken with a Leica Z16 zoom microscope fitted with a Planapo 1× lens and recorded on a Nikon D810 camera via a Leica trinocular tube and dedicated photo tubes for a full-frame format sensor. Black-and-white line drawings were made on Apple iPad 9th generation tablets using the Procreate 5.3.15 raster graphics editor, laser-printed, and rendered with Rapidograph technical pens. As this nothospecies—as with most of the Zygopetalinae belonging to the “Chondrorhyncha complex”—produces only a single, subapical flower per inflorescence, the dissected parts of the perianth were stored in FAA to serve as type material. The voucher was then accessioned into the liquid collection of the Lankester Botanical Garden Herbarium (JBL). The original plant from which the type material was prepared is maintained in cultivation, in the hope that it can produce more flowers in the future. Even though these flowers will not have any significance in terms of typification (being prepared on a date different from that of the holotype, they do not belong to the original materials), they would nonetheless be a useful reference, as they originate from the same individual plant that served as the type. The plant and flower of the new nothospecies were compared with living material grown in the Center’s greenhouses and with conserved flowers in the JBL herbarium of both putative parental species. Data on the distribution range of the putative parents were extracted from the extensive georeferenced databases of orchid distribution maintained at the Center, and the resulting habitat-compatible regions were plotted on an orographic map of Costa Rica (see further under the section on Distribution). Phenological data from both field-collected and cultivated plants of the putative parents were gathered and examined to test the consistency of the hypothesis. FP conceptualized the paper; FP and APK wrote the original draft; and ES wrote the descriptions and reviewed and edited the manuscript. Taxonomy ×Benzoscaphe Pupulin, nothogen. nov. urn:lsid:ipni.org:names:77371638-1 Type species. ×Benzoscaphe stelleri Pupulin, Karremans & E.Salas Diagnosis. Nothogenus novum caracteribus intermediis inter Benzingiam Dodson et Chondroscaphen (Dressler) Senghas et G.Gerlach. Folia viridi-grisacea sicut Benzingiarum species sed erecta prod pendulis sicut Chondroscapharum species; sepalis lateralibus omnino reflexis subuncinatis, basi petalorum ad columna adnata, pede columnae elongato Benzingiae similiter; inflorescentia suberecta prod pendula, petalis subtrapezoideis, labello callis duobus ornato, anthera elongata anguste ovata, polliniis elongatis valde curvatis supra stipitem 37 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) rectangulo concavo-convexo viscidioque incrassato sicut Chondroscapharum species; floris magnitudo et forma labelli intermediis inter genera dignoscenda. Description. Epiphytic, caespitose, erect herbs with a tuft of leaves fan-like arranged. Roots slender, flexuous, whitish. Leaves narrowly elliptic, acuminate, with a short conduplicate petiole, grayish green, articulated with basal, ovate, conduplicate sheaths provided with hyaline margins. Inflorescence a single-flowered raceme, suberect, much shorter than the leaves. Floral bracts two, the outer bract larger, broadly ovate, loose, the inner bractlet lanceolate, smaller. Pedicellate ovary terete, gently curved toward the apex. Flower spreading, downward facing, the dorsal sepal porrect-recurved, the lateral sepals reflexed, subuncinate, the petals inserted along the basal side of the column, porrect with the apex gently recurved, the lip obscurely three-lobed, the lateral lobes erect, the midlobe geniculate-bent, with ruffled margins; disc with a median, irregularly dentate callus and a second thickening toward the apex of the midlobe. Column hemiterete, dilated around the stigma, provided with a long foot, the stigma transversal, slit-like, the clinandrium swallow, the anther ovate; pollinia 4 in two superposed pairs of different sizes, on a rectangular, erected stipe and a narrowly lanceolate, thick viscidium. The new nothogenus presents intermediate features between the genera Benzingia (Fig. 1) and Chondroscaphe (Fig. 2). The leaves are grayish green with the epidermis somewhat prismatic, like in Benzingia, but they are held erect as in species of Chondroscaphe. The size of the flower is intermediate between the smaller flower of Benzingia (natural spread of flower ca. 3 cm) and larger ones typical of Chondroscaphe (spread flowers >6 cm). The flower of the nothogenus is similar to that of Benzingia in the lateral sepals completely reflexed and uncinate at the apex, the base of the petals adnate to the margins of the column, and the elongate column foot. It resembles Chondroscaphe in the subtrapezoid petals, the lip provided with a basal callus and a distinct apical bulging, the narrowly ovate, elongate anther cap, and the shape of the pollinarium, with four elongate, distinctly curved pollinia lying over a rectangular, concave-convex stipe, attached to a thick viscidium. ×Benzoscaphe stelleri Pupulin, Karremans & E.Salas, nothosp. nov. urn:lsid:ipni.org:names:77371639-1 Type. Costa RiCa • Alajuela: Naranjo, San Juanillo, road to Zarcero, epiphytic in forest remnants at the right of the road, 10°08'40.4"N, 84°23'13.0"W, 1450 m, 14 March 2024, F. Pupulin 9146 & N. Steller (holotype, JBL-K0409). (Figs 3–5). Diagnosis. Nothospecies nova caracteribus intermediis inter Benzingiam reichenbachianam (Schltr.) Dressler et valde probabiliter Chondroscaphen bicolorem (Rolfe) Dressler dignoscenda. Folia viridi-grisacea sicut Benzingiae reichenbachianae sed erecta prod pendulis sicut Chondroscaphe bicolor; sepalis lateralibus omnino reflexis subuncinatis, basi petalorum ad columna adnata, pede columnae elongato Benzingiae similiter; inflorescentia suberecta prod pendula, petalis subtrapezoideis, labello callis duobus ornato, anthera elongata anguste ovata, polliniis elongatis valde curvatis supra stipitem rectangulo concavo-convexo viscidioque incrassato sicut Chondroscaphe bicolor; floris magnitudo et forma labelli intermediis inter duabus speciebus. 38 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Figure 1. Lankester Composite Digital Plate of Benzingia reichenbachiana. A. Habit; B. Flower; C. Dissected perianth; D. Ovary, column, and lip in three-quarters view; E. Lip in longitudinal section; F. column, several views; G. Pollinarium in dorsal, ventral, and lateral views: H. anther cap. Scale bars: 1 mm (G, H); 2 cm (D, E, F); 3 cm (C); 5 cm (A). Prepared by F. Pupulin from Pupulin 5159 (voucher at JBL-B0354). 39 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Figure 2. Lankester Composite Digital Plate of Chondroscaphe bicolor. A. Habit; B. Flower; C. Dissected perianth; D. Ovary, column, and lip in three-quarters view; E. Column in lateral and three-quarters views; F. Column in ventral view; G. Pollinarium in dorsal, ventral, and lateral views; H. Anther cap. Scale bars: 3 mm (G, H); 1 cm (D, E, F); 2 cm (B); 3 cm (C); 10 cm (A). Prepared by F. Pupulin from Bogarin 14717 (voucher at JBL-L0130). 40 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Figure 3. ×Benzoscaphe stelleri. A. Habit B. Flower C. Dissected perianth; D. Ovary, column, and lip in three-quarters view; E. Column in lateral, three-quarters, and ventral views (emasculated at the right); F. Pollinarium in dorsal, ventral, three-quarters, and lateral views; G. Anther cap. Scale bars: 3 mm (F, G); 5 mm (E); 1 cm (B, D); 3 cm (C); 10 cm (A). Drawn by S. Poltronieri from F. Pupulin 9146 (voucher at JBL-K0409). 41 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Figure 4. Lankester Composite Digital Plate of ×Benzoscaphe stelleri. A. Habit; B. Flower; C. Dissected perianth; D. Ovary, column, and lip in three-quarters view; E. Column in lateral, three-quarters, and ventral views (emasculated at the right); F. Pollinarium in dorsal, ventral, three-quarters, and lateral views; G. Anther cap. Scale bars: 3 mm (F, G); 5 mm (E); 1 cm (B, D); 3 cm (C); 10 cm (A). Prepared by F. Pupulin from F. Pupulin 9146 (voucher at JBL-K0409). 48 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Use of AI No use of AI was reported. Funding This work was supported by the Costa Rican Ministry of Environment and Energy (MINAE) and its National System of Conservation Areas (SINAC) and by the Vice-Presidency of Research of the University of Costa Rica. Author contributions Conceptualization: FP. Formal analysis: ESH. Investigation: APK, FP. Writing - original draft: APK, FP. Writing - review and editing: ESH. Author ORCIDs Franco Pupulin https://orcid.org/0000-0001-6780-8395 Adam P. Karremans https://orcid.org/0000-0001-5987-7710 Elvira Salas Hidalgo https://orcid.org/0000-0002-6342-421X Data availability All of the data that support the findings of this study are available in the main text. References Ackerman JD (1983) Euglossine bee pollination of the orchid, Cochleanthes lipscombiae: A food source mimic. American Journal of Botany 70: 830–834. https://doi. org/10.1002/j.1537-2197.1983.tb06420.x Ackerman JD, Phillips RD, Tremblay RL, Karremans A, Reiter N, Peter CI, Bogarín D, Pérez-Escobar OA, Liu H (2023) Beyond the various contrivances by which orchids are pollinated: Global patterns in orchid pollination biology. Botanical Journal of the Linnean Society 202(3): 295–324. https://doi.org/10.1093/botlinnean/boac082 Archila Morales FL, Lipińska MM, Szlachetko DL (2021) Masdevallia × urbanae (Orchidaceae)—A New, Natural Hybrid between M. floribunda and M. tuerckheimii from Guatemala. Diversity 13(3): 108. https://doi.org/10.3390/d13030108 Atwood JT, Mora Retana DE (1999) Family #39 Orchidaceae: Tribe Maxillarieae: subtribes Maxillariinae and Oncidiinae. In: Burger WC (Ed.) Flora Costaricensis. Fieldiana, Botany, n.s. 40: i–vi, 1–182. Azevedo CO, Borba EL, van den Berg C (2006) Evidence of natural hybridization and introgression in Bulbophyllum involutum Borba, Semir & F. Barros and B. weddellii (Lindl.) Rchb.f. (Orchidaceae) in the Chapada Diamantina, Brazil, by using allozyme markers. Revista Brasileira de Botânica. Brazilian Journal of Botany 29: 415–421. https://doi.org/10.1590/S0100-84042006000300008 Barrera-Rojas CH, van den Berg C (2025) A new natural hybrid of Cattleya (Orchidaceae: Laeliinae) from Bahia, Brazil. Phytotaxa 689(1): 139–145. https://doi.org/10.11646/ phytotaxa.689.1.9 Batista JAN, Medeiros ADS (2022) A new putative natural hybrid of Cyrtopodium (Orchidaceae) from the south coast of Brazil. Phytotaxa 536(3): 279–286. https://doi.org/10.11646/phytotaxa.536.3.8 Bogarín D, Warner J, Powell M, Savolainen V (2011) The orchid flora of Cocos Island National Park, Puntarenas, Costa Rica. Botanical Journal of the Linnean Society 166(1): 20–39. https://doi.org/10.1111/j.1095-8339.2011.01127.x 49 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Borba EL, Semir J (1998) Bulbophyllum ×cipoense (Orchidaceae), a new natural hybrid from the Brazilian “campos rupestres”: Description and biology. Lindleyana 13: 113–120. Braem GJ (1995) Cattleya ×tenuata P.Castro & Campacci. Orchis 99: 30–31. Cantuária PC, Krahl DRP, Krahl AH, Chiron C, Silva JBF (2021) Catasetum × sheyllae (Orchidaceae: Catasetinae), a new natural hybrid from Brazilian Amazon. Phytotaxa 527: 257–265. https://doi.org/10.11646/phytotaxa.527.4.3 Carnevali G, Cetzal-Ix W, Tapia-Muñoz JL, Romero-González G (2015) The world upside down: the first Kefersteinia (Orchidaceae: Zygopetalinae) with non-resupinate flowers. Phytotaxa 239(2): 165–173. https://doi.org/10.11646/phytotaxa.239.2.5 Castro Neto VP, Catharino ELM (2004) Un nouvel hybride de Cattleya de Bahia (Brésil). Richardiana 4: 42–46. Castro Neto VP, Menezes LC (2007) Un nouvel hybride naturel originaire de Bahia (Brésil), dans le genre Hoffmanncattleya (Orchidaceae). Richardiana 7: 70–74. Cetzal-Ix W, Balam-Narváez R, Carnevali G (2012) A new nothogenus and nothospecies in the Oncidiinae (Orchidaceae) from Quintana Roo, Mexico. Nordic Journal of Botany 30(1): 40–46. https://doi.org/10.1111/j.1756-1051.2011.01261.x Cetzal-Ix W, Carnevali G, Noguera-Savelli E, Jáuregui D (2013) Morphological and anatomical characterization of a new natural hybrid between Cohniella ascendens and C. brachyphylla (Oncidiinae: Orchidaceae). Phytotaxa 144(2): 45–55. https://doi. org/10.11646/phytotaxa.144.2.2 Cetzal-Ix W, Carnevali G, Noguera-Savelli E, Mó E, Cash-Arcia N, Basu SK (2018) Natural hybridization in Lophiaris (Orchidaceae: Oncidiinae). Systematic Botany 43(4): 930–949. https://doi.org/10.1600/036364418X697599 Cetzal-Ix W, Carnevali G, Jiménez-Machorro R, Pérez-García EA (2020) Laelia × meavei: a new natural hybrid between L. dawsonii fo. dawsonii and L. rubescens fo. peduncularis (Orchidaceae: Laeliinae) from Oaxaca, Mexico. Phytotaxa 446(2): 81–94. https://doi.org/10.11646/phytotaxa.446.2.1 Chiron GR, Castro Neto VP (2002) Révision des espèces brésiliènnes du genre Laelia Lindley. Richardiana 2: 4–28. Christenson EA (2006) Deux genres hybrids naturels chez les Zygopetalinae. Richardiana 6(3): 136–138. Díaz-Morales M, Pupulin F (2019) The strange case of Phragmipedium × talamancanum. An international story of conservation and taxonomy. Orchids (Bull. Amer. Orch. Soc.) 88(12): 906–915. Dressler RL (1972) Una Kefersteinia nueva del Ecuador. Orquideología 7(3): 135–137. Dressler RL, Pollard GE (1974) The Genus Encyclia in Mexico. Asociación Mexicana de Orquideología, A. C., Mexico. Fiorini CF, de Camargo Smidt E, Knowles LL, Leite Borba E (2023) Hybridization boosters diversification in a Neotropical Bulbophyllum (Orchidaceae) group. Molecular Phylogenetics and Evolution 186: 107858. https://doi.org/10.1016/j.ympev.2023.107858 Gerlach G, Schill R (1991) Composition of orchid scents attracting euglossine bees. Botanica Acta 104: 379–384. https://doi.org/10.1111/j.1438-8677.1991.tb00245.x Heide-Jørgensen HS (2021) Orchid pollination. Viscum, June 2021. https://viscum.dk/ orchid-pollination/ [consulted on April 2025] Jiménez-Vázquez A, Vega H, Alvarado A, Mó E (2023) First record of a natural hybrid of the genus Gongora (Stanhopeinae) and comments on the parent species. Molecular Phylogenetics and Evolution 186: 107858. https://doi.org/10.15517/lank. v23i1.54261 50 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Kindlmann P, Kull T, McCormick M (2023) The Distribution and Diversity of Orchids. Diversity 15(7): 810. https://doi.org/10.3390/d15070810 Krahl AH, Krahl DRP, Pansarin E (2021) Ionopsis × atalibae (Orchidaceae, Oncidiinae), a new natural hybrid from the Brazilian Amazon. Phytotaxa 478(2): 268–274. https://doi.org/10.11646/phytotaxa.478.2.8 Krahl DRP, de Oliveira MS, Schmal P, Krahl AH, Chiron G, da Silva JBF, Cantuária P (2024) Catasetum × vilhenense (Orchidaceae: Catasetinae), a new natural hybrid from the Amazon forest. Darwiniana 12(2): 308–317. https://doi.org/10.14522/darwiniana.2024.122.1251 León-Peralta CE, Salmerón-Barrera E, Pérez-García EA (2023) Encyclia × tixtlensis (Orchidaceae: Laeliinae), a possible natural hybrid between E. meliosma and E. atrorubens in Guerrero, México. Phytotaxa 613(3): 249–260. https://doi.org/10.11646/phytotaxa.613.3.4 Lipińska MM, Barabasz C (2024) Pollination of Guatemalan orchids – state of knowledge. In: Lipińska M, Lopez-Selva MM, Sierra JM (Eds) Biodiversity research in Central America. Neotropical Biology and Conservation 19(2): 199–221. https://doi. org/10.3897/neotropical.19.e119499 Lipińska MM, Archila Morales FL, Gilka W, Beuk PLT, Szlachetko DL (2019) First report on effective pollination of Masdevallia floribunda, M. tuerckheimii and their hybrid (Orchidaceae-Pleurothallidinae) by Zygothrica fruit flies (Diptera-Drosophilidae) in Guatemala. Phyton 59(1–2): 27–33. Mó E, Cetzal-Ix W, Carnevali G, Pérez-García E, Basu S (2014) A new natural hybrid between Prosthechea cochleata and P. radiata (Orchidaceae) from Alta Verapaz, Guatemala. Turkish Journal of Botany 38(5): 11. https://doi.org/10.3906/bot-1312-86 Neubig KM, Whitten WM, Williams NH, Blanco MA, Endara L, Burleigh JG, Silvera K, Cushman JC, Chase MW (2012) Generic recircumscriptions of Oncidiinae (Orchidaceae: Cymbidieae) based on maximum likelihood analysis of combined DNA datasets. Botanical Journal of the Linnean Society 168: 117–146. https://doi.org/10.1111/j.10958339.2011.01194.x Nunes CEP, Wolowski M, Pansarin ER, Gerlach G, Aximoff I, Vereecken NJ, Salvador MJ, Sazima M (2017) More than euglossines: The diverse pollinators and floral scents of Zygopetalinae orchids. Science and Nature 104: 92. https://doi.org/10.1007/s00114017-1511-3 Pansarin ER (2025) Vanilla × robusta (Orchidaceae: Vanilloideae), the first natural Vanilla hybrid for South America. Nordic Journal of Botany 2025(7): e04743. https://doi. org/10.1111/njb.04743 Pérez-García E, Hágsater E (2012) Encyclia × nizanburyi (Orchidaceae) un nuevo híbrido natural del Istmo de Tehuantepec, México. Lankesteriana 12(1): 1–8. https://doi. org/10.15517/lank.v12i1.18268 Pinheiro F, de Barros F, Palma-Silva C, Meyer D, Fay MF, Suzuki RM, Lexer C, Cozzolino S (2010) Hybridization and introgression across different ploidy levels in the Neotropical orchids Epidendrum fulgens and E. puniceoluteum (Orchidaceae). Molecular Ecology 19(18): 3981–3994. https://doi.org/10.1111/j.1365-294X.2010.04780.x Pupulin F (2003) A noteworthy new natural hybrid in the genus Oerstedella Rchb.f. (Orchidaceae: Laeliinae). Lankesteriana, 31–36. Pupulin F (2005) Chondrorhyncha Lindl. In: Pupulin F (Ed.) Vanishing Beauty—Native Costa Rican Orchids, Vol. 1: Acianthera-Kegeliella. Editorial de la Universidad de Costa Rica, San José, 100–109. Pupulin F (2007) ×Bensteinia ramonensis, a new natural hybrid in the Zygopetalinae. Selbyana 28(2): 112–116. 51 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Pupulin F (2009) Subtribe Zygopetalinae. In: Pridgeon AM et al. (Eds) Genera Orchidacearum. Vol. 5. Epidendroideae (Part two). Oxford University Press, Oxford, 456–546. Pupulin F (2010) Flora Costaricensis/Family #39 Orchidaceae: Tribe Cymbidieae: Subtribe Zygopetalinae. Fieldiana, Botany, n.s. 40: i–vi, 1–182. https://doi. org/10.3158/0015-0746-49.1.1 Pupulin F (2015) × Cochlezella costaricensis, a name for a rare natural hybrid in the Zygopetalinae (Orchidaceae). Harvard Papers in Botany 20(2): 199–208. https://doi. org/10.3100/hpib.v20iss2.2015.n8 Pupulin F, Díaz-Morales M (2018) On the meaning of Cypripedium × grande (Orchidaceae) and its taxonomic history, with a new name for the nothospecies occurring in Costa Rica and Panama. Phytotaxa 382(2): 167–181. https://doi.org/10.11646/ phytotaxa.382.2.2 Pupulin F, Aguilar J, Belfort-Oconitrillo N, Díaz-Morales M, Bogarín D (2021) Florae costaricensis subtribui Pleurothallidinis (Orchidaceae) prodromus II. Systematics of the Pleurothallis cardiothallis and P. phyllocardia groups, and other related groups of Pleurothallis with large vegetative habit. Harvard Papers in Botany 26(1): 203–295. https://doi.org/10.3100/hpib.v26iss1.2021.n14 Ramírez S, Dressler RL, Ospina M (2002) Abejas euglosinas (Hymenoptera: Apidae) de la región Neotropical: listado de especies con notas sobre su biología. Biota Colombiana 3(1): 7–118. Reynolds AA (2004) Diversity and rarity in an Andean cloud forest: deceit pollination and fruit set in the endemic orchid, Chondroscaphe embreei. Master of Science (MS) Thesis, University of Georgia. Romero-Gonzalez GA, Carnevali G (2024) Catasetum xsteyermarkii (Catasetinae: Orchidaceae). A new putative natural hybrid of Catasetum (Catasetinae: Orchidaceae) from the Venezuelan Guyana. Harvard Papers in Botany 29(1): 1–13. https://doi. org/10.3100/hpib.v29iss1.2024.n1 Royal Horticultural Society (2017) Sander’s list of genera and nothogenera with components. www.rhs.org.uk/plants/pdfs/plant-registration-forms/list-of-orchid-generawith-components.pdf [consulted November 2024] Salazar GA, Jimenez-Machorro R, Huerta HM, Hágsater E (2014) A new species and a new natural hybrid of Laelia (Orchidaceae) from Oaxaca, Mexico. Phytotaxa 178(3): 161–170. https://doi.org/10.11646/phytotaxa.178.3.1 Sauleda RP (2015) An intergeneric natural hybrid between two non-sympatric species. New World Orchidaceae – Nomenclatural Notes, Nomenclatural Note 16. [sine paginatio] Sauleda RP, Adams RM (1990) The Orchidaceae of the Bahama Archipelago: Additions, distributional extensions, and nomenclatural changes. Brittonia 42(4): 286–291. https://doi.org/10.2307/2806819 Sauleda RP, Arrondeau P (2020) A natural hybrid of Comparettia falcata Poepp. & Endl. and Rodriguezia lanceolata Ruiz & Pavon (Orchidaceae) is reported for Colombia. New World Orchidaceae – Nomenclatural Notes. Nomenclatural Note 67: 1–10. Sauleda RP, Esperón P (2016) A new natural hybrid from Eastern Cuba. New World Orchidaceae – Nomenclatural Notes, Nomenclatural Note 22: ePublished. Sauleda RP, Esperón P (2019) A new natural hybrid of Encyclia moebusii H. Ditr. (Orchidaceae) is described for Cuba. New World Orchidaceae – Nomenclatural Notes, Nomenclatural Note 57: ePublished. Sierra-Ariza MA, Harding P (2023) A new species of Kefersteinia (Orchidaceae: Zygopetalinae), with peculiar non-resupinate flowers. Harvard Papers in Botany 28(2): 727–733. https://doi.org/10.3100/hpib.v28iss2.2023.n10 52 PhytoKeys 266: 33–52 (2025), DOI: 10.3897/phytokeys.266.159892 Franco Pupulin et al.: A new nothogenus in the Zygopetalinae (Orchidaceae) Solano R, Huerta-Espinoza H, Cruz-Garcia G, Ortiz-Riveros F (2019) A new natural hybrid in the genus Laelia (Orchidaceae) from Oaxaca, Mexico. Phytotaxa 402(5): 232–240. https://doi.org/10.11646/phytotaxa.402.5.2 Soto Calvo MA, Esperón P, Sauleda RP (2019) A new natural hybrid of Tetramicra Lindl. (Orchidaceae: Laeliinae) from Isla de la Junventud (Isla de Pinos) is described. New World Orchidaceae – Nomenclatural Notes, Nomenclatural Note 59: ePublished. Szlachetko DL, Kolanowska M, Muller F, Vannini J, Rojek J, Gorniak M (2017) First Guatemalan record of natural hybridisation between Neotropical species of the Lady’s Slipper orchid (Orchidaceae, Cypripedioideae). PeerJ 5: e4162. https://doi.org/10.7717/ peerj.4162 Tremblay RL, Ackerman JD, Zimmerman JK, Calvo RN (2005) Variation in sexual reproduction in orchids and its evolutionary consequences: A spasmodic journey to diversification. Biological Journal of the Linnean Society. Linnean Society of London 84(1): 1–54. https://doi.org/10.1111/j.1095-8312.2004.00400.x Turland NJ, Wiersema JH, Barrie FR, Gandhi KN, Gravendyck J, Greuter W, Hawksworth DL, Herendeen PS, Klopper RR, Knapp S, Kusber W-H, Li D-Z, May TW, Monro AM, Prado J, Price MJ, Smith GF, Zamora Señoret JC (2025) International Code of Nomenclature for algae, fungi, and plants (Madrid Code). Regnum Vegetabile 162: XLVII, 303 pp. van den Berg C (2014) The importance of hybridisation for the evolution of Cattleya. Renziana 4: 74–79. Zambrano BJ, Solano-Gómez R (2011) Un nuevo híbrido natural de Dracula (Orchidaceae: Pleurothallidinae) del suroccidente de Ecuador. A new natural hybrid of Dracula (Orchidaceae: Pleurothallidinae) from southwestern Ecuador. Revista Mexicana de Biodiversidad 82(3): 758–761. https://doi.org/10.22201/ib.20078706e.2011.3.735