Thesis

Systematics and evoltution of the genus Pleurothallis R. Br. (Orchidaceae) in the Greater Antilles

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Abstract

Die antillanische Flora ist eine der artenreichsten der Erde. Trotz jahrhundertelanger floristischer Forschung zeigen jüngere Studien, daß der Archipel noch immer weiße Flecken beherbergt. Das trifft besonders auf die Familie der Orchideen zu, deren letzte Bearbeitung für Cuba mehr als ein halbes Jahrhundert zurückliegt. Die vorliegende Arbeit basiert auf der lang ausstehenden Revision der Orchideengattung Pleurothallis R. Br. für die Flora de Cuba. Mittels weiterer morphologischer, palynologischer, molekulargenetischer, phytogeographischer und ökologischer Untersuchungen auch eines Florenteils der anderen Großen Antillen wird die Genese der großantillanischen Pleurothallis-Flora rekonstruiert. Der Archipel umfaßt mehr als 70 Arten dieser Gattung, wobei die Zahlen auf den einzelnen Inseln sehr verschieden sind: Cuba besitzt 39, Jamaica 23, Hispaniola 40 und Puerto Rico 11 Spezies. Das Zentrum der Diversität liegt im montanen Dreieck Ost-Cuba - Jamaica - Hispaniola, einer Region, die 95 % der groß-antillanischen Arten beherbergt, wovon 75% endemisch auf einer der Inseln sind. Da die meisten Arten entweder endemisch oder pankaribisch verbreitet sind, bleiben die floristischen Bezüge zwischen den Inseln und zu den kontinentalen Nachbargebieten nur schwach ausgeprägt. Immerhin lassen sich einige Verbindungen unter den Inseln der Großen Antillen und besonders zu Mittelamerika erkennen. Diese Affinitäten steigen von Ost nach West. Molekulargenetische und (mikro-)morphologische Daten zeigen ein deutliches Muster der historischen Biogeographie. Danach lassen sich die antillanischen Arten hinsichtlich ihrer Genese in drei Gruppen einteilen. 25% der Arten sind pankaribisch verbreitet, wobei der Großteil der Inselpopulationen vom mittelamerikanischen Festland stammt. Ebenfalls aus dieser Region stammen weitere 25%, die jedoch auf den Inseln neue Arten gebildet haben. Die verbleibenden 50% der groß-antillanischen Sippen sind autochthon und das Ergebnis adaptiver Radiation auf den Inseln. Diese intensive Kladogenese beschränkt sich auf drei Verwandtschaftskreise innerhalb der Gattung Pleurothallis in den Untergattungen Antilla Luer und Specklinia Lindl. (2 Linien). Es stellte sich heraus, daß der überwiegende Anteil der Artbildungsprozesse allopatrischer Natur ist. Sympatrie konnte nur in einem einzigen Fall direkt belegt werden. Das Ergebnis der allopatrischen Speziation sind zwei Typen von Vikarianz, räumlich geographischer und geologischer. In Cuba sind überraschenderweise fast 80% der endemischen Arten an einen Gesteinstyp gebunden, überwiegend an Serpentin. West-Hispaniola, wo viele Schwesternarten cubanischer Sippen beheimatet sind, besteht fast ausschließlich aus Kalkstein. Geographische Vikarianz ist daher oft geologisch unterlegt, eine Bindung die für Epiphyten kaum vermutet wurde. Hinter der Geologie verbergen sich jedoch eher Bestäuberareale und weniger physiologische Anpassung als limitierender Faktor. Eine Verfrachtung in Vegetation auf anderem petrologischen Untergrund scheint damit der Hauptauslöser für Artbildungen gewesen zu sein. Ausgangspunkt waren höchstwahrscheinlich individuenarme Gründerpopulationen die den Bedingungen eines founder events ausgesetzt waren. Neben den reichen geologischen Verhältnissen im Dreieck Ost-Cuba - Jamaica - Hispaniola wird die intensive Artbildung durch weitere spezifisch lokale Bedingungen unterstützt. Karibische Wirbelstürme dürften entlang der Hauptrouten für eine häufige Verfrachtung von Samen oder Pflanzen von Mittelamerika auf die Großen Antillen sowie zwischen den Inseln selber verantwortlich sein. Ein zweiter günstiger Umstand für erfolgreiche Migration innerhalb des Dreiecks besteht in der räumlichen Nähe der Inselgebirge und deren optimalen klimatischen Bedingungen für die Besiedlung durch mikrophytische Epiphyten. Molekulargenetische Daten lieferten weiterhin wertvolle Informationen in Bezug auf die beiden aktuell diskutierten Systeme der Pleurothallidinae, einer streng morphologischen (Luer) und einer fast ausschließlich auf DNA-Sequenzen (Pridgeon & Chase) basierenden Klassifikation. DNA-Sequenzen der cubanischen Arten stützen das neue System von Pridgeon & Chase weitestgehend, zeigen aber Widersprüche bezüglich der Monophylie in einigen der neuen oder wieder errichteten Taxa. Angesicht dessen, daß die karibische Florenregion leider nicht nur durch ihre Biodiversität zu den zehn globalen hot spots zählt, sondern auch durch die großflächige Zerstörung von Primärvegetation, war es auch ein Anliegen der vorliegenden Arbeit, ein erstes detailliertes Bild von Genese und Verbreitung antillanischer Orchideen zu vermitteln. Diese Daten können direkt für die Gestaltung und das Management von karibischen Schutzgebieten eingesetzt werden, da Orchideen in der Naturschutzpolitik einen hohen Argumentationswert besitzen.

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... Tomzanonia represents one of a handful of Pleurothallidinae lineages endemic to the Caribbean that make the region important for studying the subtribe's evolution and biogeography. Research on the phylogeography of Pleurothallidinae in the Caribbean (Stenzel 2004) revealed that Caribbean Pleurothallis R.Br. in Aiton (1813: 211) is not monophyletic and colonized the Caribbean post-Pleistocene with several dispersals from Central America followed by subsequent anagenesis. In this paper we build on the data set generated by Stenzel (2004) by including the unusual species Pleurothallis quisqueyana, which is attributed to Pleurothallis subg. ...
... Research on the phylogeography of Pleurothallidinae in the Caribbean (Stenzel 2004) revealed that Caribbean Pleurothallis R.Br. in Aiton (1813: 211) is not monophyletic and colonized the Caribbean post-Pleistocene with several dispersals from Central America followed by subsequent anagenesis. In this paper we build on the data set generated by Stenzel (2004) by including the unusual species Pleurothallis quisqueyana, which is attributed to Pleurothallis subg. Antilla Luer (2000: 39) and shown by Stenzel's (2004) results to be closely related to a clade of Pleurothallis now recognized in the genus Acianthera. ...
... In this paper we build on the data set generated by Stenzel (2004) by including the unusual species Pleurothallis quisqueyana, which is attributed to Pleurothallis subg. Antilla Luer (2000: 39) and shown by Stenzel's (2004) results to be closely related to a clade of Pleurothallis now recognized in the genus Acianthera. ...
Article
A new species of Specklinia with a repent growth habit and minute purple flowers is described and illustrated. The new species is distinguished from Specklinia wrightii based on its morphological and molecular distinctness from that species. The phylogenetic placement of the new species is provided based on an nrITS tree. The species described here represents the first new orchid to be described from material originating from the Parc National Naturel Macaya in six years. New combinations are made in Acianthera for species of Kraenzlinella and Pleurothallis subgen. Antilla embedded within the genus. In particular Kraenzlinella rinkei is provided as a new synonym for Specklinia montezumae and Specklinia simpliciflora is transferred to Acianthera sect. Antilla based on morphology and geographic distribution.
... Not surprisingly, subsequent phylogenetic studies within the Pleurothallidinae have shown that the generic, subgeneric and sectional systematics of the subtribe were not fully resolved. Recircumscriptions and emendations were either made or at least suggested by several authors who used novel analytical methods and/or included a broader sampling of species (Stenzel 2004, Abele 2007, Karremans 2010, 2014, Chiron et al. 2012, Karremans et al. 2013a, 2013b, Wilson et al. 2013, Karremans & Rincón-González 2015, Chiron et al. 2016. Meanwhile, hundreds of species' names, either new species or combinations, and dozens of new genera have since then been proposed by Luer (2002aLuer ( , 2004Luer ( , 2005Luer ( , 2006Luer ( , 2007Luer ( , 2009 and others, mostly but not exclusively, on the basis of morphology. ...
... The proposed affinities are Acianthera (Ac), Dilomilis (Di), Lepanthes (Le), Masdevallia (Ma), Octomeria (Oc), Phloeophila (Ph), Pleurothallis (Pl), Restrepia (Re) and Specklinia (Sp); the oldest generic name is used here for the species within those clades (except Pleurothallis). This phylogenetic arrangement of the subtribe, despite taking only DNAbased studies into consideration , Stenzel 2004, Solano-Gómez 2005, Matuszkiewicz & Tukallo 2006, Abele 2007, Forster 2007, Meyer & Cameron 2009, Karremans 2010, 2014, Endara 2011, Bogarín et al. 2013, Chiron et al. 2012, Karremans et al. 2013a, 2013b, Wilson et al. 2013, in press, Pessoa et al. 2014, Karremans & Rincón-González 2015, McDaniel & Cameron 2015, Chiron et al. 2016 *At the time of preparation of this manuscript the total species number for genus Masdevallia had not been revised thoroughly. **The number of species attributed to Pleurothallis excludes those names that have not been placed elsewhere but most like do not belong in the genus. ...
... If Acianthera is to be maintained in its currently most accepted, broad sense (Pridgeon 2005), then it includes the generic concepts of Arthrosia, Brenesia, Cryptophoranthus, and Sarracenella, as proposed by and Chiron & van den Berg (2012). Nevertheless, it should also include Aberrantia, as suggested by Luer (2004) and proposed by Bogarín et al. (2008), Antilla, as shown by Stenzel (2004), Apoda-prorepentia, as shown by Stenzel (2004) and Karremans & Rincón-González (2015), Didactylus, as suggested by Luer (2004), Ogygia, as proposed by Solano-Gómez (2003;, Pleurobotryum, as shown by Chiron et al. (2012), Proctoria, as suggested by Luer (2006), and Unguella, as suggested by Luer (2004). To my knowledge, there are no DNA data available (published or unpublished) of species belonging to the monospecific genera Ogygia and Proctoria that can confirm their exact phylogenetic relationships. ...
Article
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Subtribe Pleurothallidinae with just over 5000 species is possibly the most species-rich of all orchids. It has been growing steadily for more than two centuries, but the last three decades have been especially active in terms of systematic and phylogenetic studies in the group. The growth in species numbers has been accompanied by the marked increase in generic and infrageneric concepts. Nevertheless, Pleurothallidinae are plagued with cases of convergent and divergent morphology, and phylogenetic relatedness is not always apparent. This opens the door to controversial changes in generic circumscriptions that are considered too inclusive by some and too exclusive by others. A grave consequence of these disagreements is the difficulty of assessing which and how many species actually belong to each genus. Here an attempt is made to place generic names among their close relatives as a first step to re-evaluating the whole subtribe.
... DNA data has consistently shown that P. restrepioides, type species of Elongatia, and its closest relatives belong in Pleurothallis rather than Stelis Wilson et al. 2013, Pérez-Escobar et al. 2017 (Fig. 32) Morphologically this taxon could be confused with a member of Stelis in the broad sense, and in fact many authors still place it in Stelis rather than Pleurothallis. Stenzel (2004) doubted the results of his own phylogenetic reconstruction in which two accessions of Pleurothallis ghiesbreghtiana A.Rich. & Galeotti (= P. quadrifida) were found sister to Pleurothallis rather than Stelis. ...
... & Galeotti (= P. quadrifida) were found sister to Pleurothallis rather than Stelis. However, except for the phylogenetic inference presented by Solano-Gómez (2005), all other DNA based studies consistently show that P. quadrifida, type species of Lalexia, is sister to the remaining species of Pleurothallis rather than Stelis (Stenzel 2004, Wilson et al. 2013, Pérez-Escobar et al. 2017). The exclusion from Stelis is supported by multi-gene genomic studies (Ponert et al. 2019). ...
Article
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Despite the availability of multiple sources of evidence and consistency in the support for a broadly circumscribed Stelis Sw. (Orchidaceae: Pleurothallidinae), some authors continue to be hesitant in its use. It is certain that the more typical species of Stelis, with their triangular, flattish flowers with very short fleshy petals and lip, form a monophyletic group that is easily recognized. However, it is likewise undisputed that they are not an isolated lineage in the subtribe and that several groups of species with a similar vegetative habit but that lack the typical Stelis flower are in fact very close relatives, sharing a relatively recent common ancestor. Those species groups need to be classified in a way that also reflects their own evolutionary history, and alternatives to a broadly circumscribed Stelis are possible yet neither straightforward or practical at this time. An infrageneric classification for the whole group is provided here in an attempt to clarify which species actually belong where in this highly complex affinity. Emphasis is made on the difficulty of diagnosing the less typical members of each proposed subgenus or section, and on the importance of floral convergence and divergence as a result of pollinator adaptation. As here defined, Stelis is the largest genus in the Pleurothallidinae, with 1243 species. Key words: convergence; evolutionary history; floral morphology; generic circumscription; Pleurothallidinae; pollinator adaptation
... The phylogenetic relationships of Pleurothallidinae were first studied by Pridgeon et al. (2001), who suggested that Octomeria is most closely related to Brachionidium Lindl., with moderate to weak support, forming a group that is sister to the rest of the subtribe. Stenzel (2004) and Forster (2007) also recovered Octomeria in a clade with Brachionidium, but also included the genus Atopoglossum Luer. More recently, Karremans (2016) provided an updated phylogenetic overview of the Pleurothallidinae, where Sansonia Chiron was placed along with Octomeria, Atopoglossum and Brachionidium, forming a basal clade that the author referred to as "Affinity Octomeria". ...
Article
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Octomeria is a well‐represented basal genus in the subtribe Pleurothallidinae, the main myophile group of the Orchidaceae. The systematics of the genus is based in leaf shape and degree of connation of sepals. In this study, we analyse comparatively the flower micromorphology of 15 species through light microscopy, histochemical tests and scanning electron microscopy, including most of the infrageneric groups proposed for the genus. The epidermal structures of sepals and petals are very similar, with oblong or isodiametric cells, stomata and trichome‐like structures adaxially. The lip presents a uniseriate epidermis with ovate to irregular cells and mesophyll with homogeneous parenchyma and idioblasts with raphides. Calluses and grooves in the lip were shown to be secretory regions. Histochemical tests in fresh flowers were positive for starch in the sepals, petals and lip and positive for osmophores in the lip. Three types of waxes were observed; one of them is described for the first time. Our study shows that despite similarities in the perianth, floral micromorphology indicated differences in the lip and column surfaces. Thus, this approach can be an important source of phylogenetic information for the genus.
... The pioneering molecular phylogenetic studies of on Pleurothallidinae demonstrated that many of the genera recognized at that time were either polyphyletic or paraphyletic and that additional sampling and sequencing would need to be done in the different groups in order to characterize monophyletic genera. In the last decade and a half, a number of molecular phylogenetic studies have been published on multiple genera in Pleurothallidinae, including: Acianthera Scheidweiler (1842: 292) (Stenzel 2004;Chiron et al. 2012;Karremans & Rincón-González 2015;Karremans et al. 2016b); Anathallis Barbosa Rodrigues (1877: 23) (Chiron et al. 2012;Karremans 2014Karremans , 2015Pessoa et al. 2014); Masdevallia Ruiz & Pavón (1794: 122) (Matuszkiewicz & Tukallo, 2006;Abele 2007); Pabstiella Brieger & Senghas (1976: 195) (Chiron et al. 2012); Phloeophila (Chiron et al. 2016); Specklinia Lindley (1830: 8) (Bogarín et al. 2013;Karremans et al. 2013b;Karremans et al. 2015a;Karremans et al. 2015b;Karremans et al. 2016a); and Stelis Swartz (1799ba: 239) (Solano-Gomez 2005; Karremans 2010; Karremans et al. 2013a). Other molecular phylogenetic studies are in progress and preliminary reports have been published, including Dracula Luer (1978: 190) (Meyer & Cameron, 2009); Masdevallia (Doucette et al. 2014.); ...
Article
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Most of the species studied in this paper have previously been placed in either Pleurothallis or Lepanthes. However, at one time or another, members of the group have also been placed in the genera Andinia, Brachycladium, Lueranthos, Masdevalliantha, Neooreophilus, Oreophilus, Penducella, Salpistele and Xenosia. Phylogenetic analyses of nuclear ITS and plastid matK sequences indicate that these species form a strongly supported clade that is only distantly related to Lepanthes and is distinct from Pleurothallis and Salpistele. Since this clade includes the type species of Andinia, A. dielsii, and it has taxonomic precedence over all other generic names belonging to this group, Andinia is re-circumscribed and expanded to include 72 species segregated into five subgenera: Aenigma, Andinia, Brachycladium, Masdevalliantha and Minuscula. The required taxonomic transfers are made herein. We hypothesize that convergent evolution towards a similar pollinator syndrome involving deceit pollination via pseudocopulation by Diptera resulted in a similar floral morphology between species of subgenus Brachycladium and species of Lepanthes; hence the prior placement of the species of subgenus Brachycladium in Lepanthes. Species of the re-circumscribed Andinia are confined exclusively to the Andes, ranging from about 1,200 to 3,800 m, from Colombia south to Bolivia, making the generic name very apt. Elevational distributions of the individual clades are discussed in relation to the possible evolutionary diversification of the most species-rich clade, subgenus Brachycladium.
... After the alignments had been edited, additional sequences were obtained from Hagen Stenzel (Stenzel 2004), and from NCbI genbank, the latter using nblASt. Echinosepala aspasicensis (reichenbach 1855: 73) Pridgeon & Chase (2002: 101) was used as outgroup in all cases, as this taxon has been suggested to be the most earliest-branching lineage of all included species . ...
Article
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The phylogenetic relationships within Specklinia (Pleurothallidinae; Orchidaceae) and related genera are re-evaluated using Bayesian analyses of nrITS and chloroplast matK sequence data of a wide sampling of species. Specklinia is found paraphyletic in the DNA based trees, with species alternatively assigned to Muscarella proven distinct, monophyletic and easily recognizable. Specklinia as such includes about 100 morphologically highly diverse species. Their phenotypic differences have prompted the creation of up to eleven generic names within this relatively small group. Here we show not only that these morphologically divergent species are closely related, but also that they can still be recognized by certain conserved morphological traits. The genera Acostaea, Areldia, Empusella, Cucumeria, Gerardoa, Pseudoctomeria, Sarcinula, Sylphia, Tribulago and Tridelta are found embedded within Specklinia, and therefore reduced under the synonymy of the latter. Specklinia is confirmed as sister to a clade that includes Platystele, Scaphosepalum and Teagueia. Five well-supported subgenera are proposed for Specklinia and are characterized both geographically and morphologically. The species belonging to each subgenus are listed. Incaea is synonymized with Dryadella, while Rubellia is reduced under Platystele. New combinations for several species are proposed. The criteria for the generic delimitation of Specklinia and other genera in the Pleurothallidinae are discussed.
... The three independently extracted and sequenced accessions of Phloeophila nummularia form a highly supported cluster (PP=1). The two obtained from plants of Brazilian origin being almost identical to each other (Karremans 5959 and Tesch sn), whereas a third accession obtained by Stenzel (2004) from Cuban material is sister to these and has some variation in sequence. ...
Article
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Two of us found some years ago that Phloeophila nummularia, type species of the genus Phloeophila, was placed in amolecular phylogentic study in Pabstiella. In this article, we show with the inclusion of new sequences of Phloeophila that this result was not correct. Phloeophila forms a unique clade in Pleurothallidinae and is not closely related to Pabstiella. In addition, the exclusion of three species, one transferred to Dryadella and two others to Acianthera, is proposed.
... Acianthera melanochthoda (Luer & Hirtz in Luer 1996: 169) Pridgeon & Chase (2001: 244) was the only species assigned to Apoda-prorepentia sequenced by . A second species, Acianthera testifolia (Swartz 1788: 122) Solano in Solano et al. (2011, was included by Stenzel (2004) in his phylogenetic analysis of the Pleurothallidinae of the Antilles. In both cases it was clear that the species of Apoda-prorepentia were embedded within Acianthera and closely related to other Central American and Andean species of the genus. ...
Article
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A new species of Acianthera, A. serratifolia, is described and illustrated based on material from Colombia. The species belongs to a small group of species that has alternatively been placed in the genus Apoda-prorepentia. Nevertheless, Apoda-prorepentia is embedded within Acianthera and therefore reduced to synonymy. The required new combinations are proposed.
... Since the publication of the reclassification of subtribe Pleurothallidinae by, based on, several subsequent studies have placed hundreds of additional pleurothallids in a dNA–based phylogenetic context (Stenzel 2004, Abele 2007, Chiron et al. 2012, Karremans 2014). Together those studies suggest that although refinement is necessary in many groups, the generic framework presented by Pridgeon (2005) is maintained in general terms. ...
Article
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