GENERIC REALIGNMENTS IN MAXILLARIINAE (ORCHIDACEAE)

2007

The orchid genus Maxillaria is one of the largest and most common of neotropical orchid genera, but its current generic boundaries and relationships have long been regarded as artificial. Phylogenetic relationships within subtribe Maxillariinae sensu Dressler (1993) with emphasis on Maxillaria s.l. were inferred using parsimony analyses of individual and combined DNA sequence data. We analyzed a combined matrix of nrITS DNA, the plastid matK gene and flanking trnK intron, and the plastid atpB-rbcL intergenic spacer for 619 individuals representing ca. 354 species. The plastid rpoC1 gene (ca. 2600 bp) was sequenced for 84 selected species and combined in a more limited analysis with the other data sets to provide greater resolution. In a wellresolved, supported consensus, most clades were present in more than one individual analysis. All the currently recognized minor genera of ''core'' Maxillariinae (Anthosiphon, Chrysocycnis, Cryptocentrum, Cyrtidiorchis, Mormolyca, Pityphyllum, and Trigonidium) are embedded within a polyphyletic Maxillaria s.l. Our results support the recognition of a more restricted Maxillaria, of some previously published segregate genera (Brasiliorchis, Camaridium, Christensonella, Heterotaxis, Ornithidium, Sauvetrea), and of several novel clades at the generic level. These revised monophyletic generic concepts should minimize further nomenclatural changes, encourage monographic studies, and facilitate more focused analyses of character evolution within Maxillariinae.

American Journal of Botany, 2000

The monophyly of and phylogenetic relationships within the orchid tribe Maxillarieae Pfitzer were evaluated using parsimony analyses of combined nuclear ribosomal and plastid DNA sequence data of ITS 1 and 2, matK, and the trnL intron and the trnL-F intergene spacer. Each of the separate analyses produced highly congruent but weakly supported patterns (by the bootstrap), so these were combined in a single analysis. Analysis of 90 ingroup taxa (representing ϳ35% of currently recognized genera) and four outgroup taxa produced resolved and highly supported cladograms. Based on the cladograms, we recognize six subtribes: Eriopsidinae, Oncidiinae (including Pachyphyllinae, Ornithocephalinae, and Telipogoninae), Stanhopeinae, Coeliopsidinae, Maxillariinae (including Lycastinae and Bifrenariinae), and Zygopetalinae (including Cryptarrheninae, Dichaeinae, Huntleyinae, and Warreinae). Stanhopeinae were sampled most intensively; their generic relationships were highly resolved in the analysis and largely agree with currently accepted generic concepts based on morphology. Coeliopsidinae (Coeliopsis, Lycomormium, Peristeria) are sister to Stanhopeinae. Correlations are drawn among phylogeny, pollination mechanisms, and life history traits.

2000

The monophyly of and phylogenetic relationships within the orchid tribe Maxillarieae Pfitzer were evaluated using parsimony analyses of combined nuclear ribosomal and plastid DNA sequence data of ITS 1 and 2, matK, and the trnL intron and the trnL-F intergene spacer. Each of the separate analyses produced highly congruent but weakly supported patterns (by the bootstrap), so these were combined in a single analysis. Analysis of 90 ingroup taxa (representing ϳ35% of currently recognized genera) and four outgroup taxa produced resolved and highly supported cladograms. Based on the cladograms, we recognize six subtribes: Eriopsidinae, Oncidiinae (including Pachyphyllinae, Ornithocephalinae, and Telipogoninae), Stanhopeinae, Coeliopsidinae, Maxillariinae (including Lycastinae and Bifrenariinae), and Zygopetalinae (including Cryptarrheninae, Dichaeinae, Huntleyinae, and Warreinae). Stanhopeinae were sampled most intensively; their generic relationships were highly resolved in the analysis and largely agree with currently accepted generic concepts based on morphology. Coeliopsidinae (Coeliopsis, Lycomormium, Peristeria) are sister to Stanhopeinae. Correlations are drawn among phylogeny, pollination mechanisms, and life history traits.

Lankesteriana, 2008

A recent phylogenetic analysis of four DNA regions for ca. 354 species of core Maxillariinae strongly indicate that the genus Maxillaria, as traditionally circumscribed, is grossly polyphyletic. We present a new phylogenetic classification for core Maxillariinae that recognizes 17 genera. Necessary realignments include: 1) resurrection of the genera Camaridium, Heterotaxis, and Ornithidium; 2) recognition of the recent segregates Brasiliorchis (=Maxillaria sect. Repentes), Christensonella (=Maxillaria sect. Urceolatae), Nitidobulbon (in press), and a recircumscribed Sauvetrea (=Maxillaria sect. Trigonae); 3) adoption of the new genera Inti (=Maxillaria sect. Polyphyllae), Mapinguari, Maxillariella (=Maxillaria sections Ebulbes and Erectae), and Rhetinantha; 4) transfers from Maxillaria sect. Reflexae to Ornithidium, and Maxillaria sect. Rufescens to Mormolyca; and 5) synonymizing of the genera Adamanthus, Pseudomaxillaria, Psittacoglossum, and Sepalosaccus (under Camaridium), Anthosiphon (under Cryptocentrum), Chrysocycnis (under Mormolyca), Dicrypta, Marsupiaria, and Pentulops (under Heterotaxis), and Laricorchis, Neo-urbania, and Siagonanthus (under Ornithidium). Some new synonyms at the specific level are also presented.

American Journal of Botany, 2007

The orchid genus Maxillaria is one of the largest and most common of neotropical orchid genera, but its current generic boundaries and relationships have long been regarded as artificial. Phylogenetic relationships within subtribe Maxillariinae sensu Dressler (1993) with emphasis on Maxillaria s.l. were inferred using parsimony analyses of individual and combined DNA sequence data. We analyzed a combined matrix of nrITS DNA, the plastid matK gene and flanking trnK intron, and the plastid atpB-rbcL intergenic spacer for 619 individuals representing ca. 354 species. The plastid rpoC1 gene (ca. 2600 bp) was sequenced for 84 selected species and combined in a more limited analysis with the other data sets to provide greater resolution. In a wellresolved, supported consensus, most clades were present in more than one individual analysis. All the currently recognized minor genera of ''core'' Maxillariinae (Anthosiphon, Chrysocycnis, Cryptocentrum, Cyrtidiorchis, Mormolyca, Pityphyllum, and Trigonidium) are embedded within a polyphyletic Maxillaria s.l. Our results support the recognition of a more restricted Maxillaria, of some previously published segregate genera (Brasiliorchis, Camaridium, Christensonella, Heterotaxis, Ornithidium, Sauvetrea), and of several novel clades at the generic level. These revised monophyletic generic concepts should minimize further nomenclatural changes, encourage monographic studies, and facilitate more focused analyses of character evolution within Maxillariinae.

Kew Bulletin, 2011

The type specimen of Maxillaria grandiflora (Kunth) Lindl. was collected by Alexander von Humboldt and Aimé Bonpland during their scientific expedition to Tropical America. Its type locality, which has been the subject of confusion, is identified as a place on the eastern part of La Cruz municipality in the province of Nariño, Colombia. The type of M. grandiflora is shown to be a mixed collection and a lectotype is designated; the protologue illustration represents an amalgamation of both species. The type of Heterotaxis valenzuelana (A. Rich.) Ojeda & Carnevali was collected by José María Valenzuela but has been erroneously cited as Wright 3314 in the recent literature.

Botanical Journal of the Linnean Society, 2015

Since the last classification of Orchidaceae in 2003, there has been major progress in the determination of relationships, and we present here a revised classification including a list of all 736 currently recognized genera. A number of generic changes have occurred in Orchideae (Orchidoideae), but the majority of changes have occurred in Epidendroideae. In the latter, almost all of the problematic placements recognized in the previous classification 11 years ago have now been resolved. In Epidendroideae, we have recognized three new tribes (relative to the last classification): Thaieae (monogeneric) for Thaia, which was previously considered to be the only taxon incertae sedis; Xerorchideae (monogeneric) for Xerorchis; and Wullschlaegelieae for achlorophyllous Wullschlaegelia, which had tentatively been placed in Calypsoeae. Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid grade, with Neottieae sister to the remainder of Epidendroideae. Arethuseae are sister to the rest of the higher Epidendroideae, which is unsurprising given their mostly soft pollinia. Tribal relationships within Epidendroideae have been much clarified by analyses of multiple plastid DNA regions and the low-copy nuclear gene Xdh. Four major clades within the remainder of Epidendroideae are recognized: Vandeae/Podochileae/Collabieae, Cymbidieae, Malaxideae and Epidendreae, the last now including Calypsoinae (previously recognized as a tribe on its own) and Agrostophyllinae s.s. Agrostophyllinae and Collabiinae were unplaced subtribes in the 2003 classification. The former are now split between two subtribes, Agrostophyllinae s.s. and Adrorhizinae, the first now included in Epidendreae and the second in Vandeae. Collabiinae, also probably related to Vandeae, are now elevated to a tribe along with Podochileae. Malaxis and relatives are placed in Malaxidinae and included with Dendrobiinae in Malaxideae. The increased resolution and content of larger clades, recognized here as tribes, do not support the 'phylads' in Epidendroideae proposed 22 years ago by Dressler.

Harvard Papers in Botany, 2008

Few orchid species have experienced such a complicated taxonomic history as Ornithidium pendulum (Poepp. & Endl.) Cogn. (Fig. 1). This species has been described under six different names from three (or four) countries. It has been known by a misapplied name (Maxillaria ramosa Ruiz & Pav.) for over 40 years, and it has been confused with a hitherto undescribed species from Venezuela and the Guianas. McIllmurray and Oakeley (2004) unraveled much of the confusion, but their paper has remained relatively unknown among orchid taxonomists. The present paper aims to further clarify the identity of O. pendulum.

Since the last classification of Orchidaceae in 2003, there has been major progress in the determination of relationships, and we present here a revised classification including a list of all 736 currently recognized genera. A number of generic changes have occurred in Orchideae (Orchidoideae), but the majority of changes have occurred in Epidendroideae. In the latter, almost all of the problematic placements recognized in the previous classification 11 years ago have now been resolved. In Epidendroideae, we have recognized three new tribes (relative to the last classification): Thaieae (monogeneric) for Thaia, which was previously considered to be the only taxon incertae sedis; Xerorchideae (monogeneric) for Xerorchis; and Wullschlaegelieae for achlorophyllous Wullschlaegelia, which had tentatively been placed in Calypsoeae. Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid grade, with Neottieae sister to the remainder of Epidendroideae. Arethuseae are sister to the rest of the higher Epidendroideae, which is unsurprising given their mostly soft pollinia. Tribal relationships within Epidendroideae have been much clarified by analyses of multiple plastid DNA regions and the low-copy nuclear gene Xdh. Four major clades within the remainder of Epidendroideae are recognized: Vandeae/Podochileae/Collabieae, Cymbidieae, Malaxideae and Epidendreae, the last now including Calypsoinae (previously recognized as a tribe on its own) and Agrostophyllinae s.s. Agrostophyllinae and Collabiinae were unplaced subtribes in the 2003 classification. The former are now split between two subtribes, Agrostophyllinae s.s. and Adrorhizinae, the first now included in Epidendreae and the second in Vandeae. Collabiinae, also probably related to Vandeae, are now elevated to a tribe along with Podochileae. Malaxis and relatives are placed in Malaxidinae and included with Dendrobiinae in Malaxideae. The increased resolution and content of larger clades, recognized here as tribes, do not support the 'phylads' in Epidendroideae proposed 22 years ago by Dressler.

2005

Schoot ofArts and Seience, Uhivensity of7bdy,o, Kbmaba, 7bkyo, J53-8902, JZu)an, Molecular phylogenetic analyscs were performed using data sets derived from DNA sequences ofthe plastid genome (matK and trnK introns) and the nuelear genome (rDNA ITS) in an examination ofrelationships of all sections ofPhataenqpsis and closely related gcnera. The fo11owing insights were provided: (1) The genera Lesliea and IVbthodoritis are nested within Phalaenopsis, (2) Phalaenopsis subgenus Aphyilae and section EsmeJ'aldd, often treated as thc independent genera Kirrgidium and Doritis respectively, are also nested within Phalaenqpsis. (3) Two subgenera of Phalaenqpsis, namely, Phalaenopsis and 1larishianae, are not monophyletic. (4) Phalaenopsis sections Deliciosae, SZautqglottis, Amboinenses and Zehrinae are not monophyletic. (5) lnconsistencies bctween the plastid and nuclear lineages indicate a hybrid origin ofPhalaenopsis minus and Phalaenopsis phitmpinensis. (6) In light of these findings, and to accommodate phylogenetic integrity and stability in nomenclature, we adopt a broadly defincd Doritis characterized by the possession of fbur pollinia, an explicit character state.