Nataly O'Leary

A molecular phylogeny and classification of Verbenaceae.

UNLABELLED PREMISE OF THE STUDY Verbenaceae consist of trees, shrubs, lianas, and herbs distributed primarily in Latin America, where they occur in a wide array of ecosystems. A second center of diversity exists in Africa. Competing morphology-based classifications that rely on different traits conflict in significant ways. A broad phylogenetic study was undertaken to assess those classifications and to examine the historical geography of the family. • METHODS Analysis of seven chloroplast DNA regions for 109 species, representing all genera except one monotypic genus, provide inference into evolutionary relationships in Verbenaceae. • KEY RESULTS The phylogeny shows that none of the traditional classifications reflect phylogenetic relationships very well. Eight clades are recognized as tribes (Casselieae, Citharexyleae, Duranteae, Lantaneae, Neospartoneae trib. nov., Petreeae, Priveae, and Verbeneae). Two genera, Dipyrena and Rhaphithamnus, remain unplaced in these larger clades. Petreeae, which consist of Neotropical lianas, are sister to the rest of the family. Lantaneae and Verbeneae together form a derived clade that comprises approximately two-thirds of the species in Verbenaceae. • CONCLUSIONS We present a new tribal classification, including one new tribe, Neospartoneae trib. nov., to accommodate three small genera of Argentine species (Diostea, Neosparton, and Lampaya). Phylogenetic inference suggests a South American origin for Verbenaceae, with approximately six colonization events having given rise to the Old World species.

Reassignment of Species of Paraphyletic Junellia s. l. to the New Genus Mulguraea (Verbenaceae) and New Circumscription of Genus Junellia: Molecular and Morphological Congruence

Abstract Verbenaceae tribe Verbeneae includes three major genera, Verbena, Glandularia, and Junellia, which form a recently diversified group. Junellia is a South American genus, whereas Verbena and Glandularia are distributed in temperate regions of both South and North America. Seven noncoding chloroplast regions were sequenced including intergenic spacers and/or introns in trnD-trnT, trnS-trnG, trnS-trnfM, trnT-trnL, trnG, trnL, and trnL-trnF; the nuclear ITS and ETS regions were also sequenced. Together with previous studies, these results suggest that Junellia, as traditionally conceived, is a paraphyletic group of two separate clades. Junellia should be restricted to the clade containing the type, J. micrantha, which also includes Glandularia subgenus Paraglandularia and the genus Urbania. Consequently Urbania and Glandularia subgenus Paraglandularia are reduced to synonymy under Junellia, and eight new combinations in Junellia are proposed: Junellia ballsii, J. crithmifolia, J. fasciculata, J. hookeriana, J. lucanensis, J. occulta, J. pappigera and J. origenes. The remaining Junellia species form a monophyletic group here designated as the new genus Mulguraea, with 13 new combinations: Mulguraea arequipense, M. asparagoides, M. aspera var. aspera, M. aspera var. longidentata, M. cedroides, M. cinerascens, M. echegarayi, M. hystrix, M. ligustrina var. ligustrina, M. ligustrina var. lorentzii, M. scoparia, M. tetragonocalyx and M. tridens . Verbena and Glandularia s. s. are both monophyletic based on ETS/ITS data, but neither is monophyletic based on cpDNA data. Relationships within each genus are still not wholly resolved, nevertheless there is evidence that South and North American Verbena might both be monophyletic. Verbena and Glandularia are sister groups, and together they are sister to the reconstituted Junellia. Mulguraea is sister to the group comprising all the three genera, Verbena, Glandularia, and Junellia.

Evolution of morphological traits in Verbenaceae

PREMISE OF THE STUDY A new infrafamilial circumscription of the Verbenaceae with eight tribes: Casselieae, Citharexyleae, Duranteae, Lantaneae, Neospartoneae, Petreeae, Priveae, and Verbeneae, has been recently proposed, on the basis of molecular phylogenetic studies. Two genera, Dipyrena and Rhaphithamnus, remain unplaced. The aim of this work is to reconstruct the evolutionary history of morphological characters traditionally employed in the classification of the Verbenaceae, with special attention to tribes Verbeneae and Lantaneae. METHODS Twenty-one characters, related to habit and vegetative morphology, inflorescence and floral morphology, ovary and fruit morphology, as well as chromosome number, were optimized over a molecular phylogeny of Verbenaceae. KEY RESULTS All tribes are supported by at least one morphological trait except tribes Duranteae and Citharexyleae. Suffrutescent habit, sessile flowers, and four cluses are synapomorphies for tribe Verbeneae. Gynoecium with short style and entire stigma are synapomorphic traits for tribe Lantaneae. Sessile flowers and unicarpellate ovaries are morphological synapomorphies for the new tribe Neospartoneae. Suffrutescent habit is a synapomorphic trait for tribe Priveae. Homothetic pleiobotrya and absence of the adaxial staminode are synapomorphic traits for tribe Casselieae. Undivided fleshy fruits are probably a synapomorphic trait for tribe Petreeae. Putative plesiomorphies for the ancestor of the Verbenaceae are discussed as well as synapomorphic traits within other Verbenaceae clades. CONCLUSIONS Many of the characters traditionally employed in classification have proven to be very homoplastic, or have been shown not to support relationships within the family. Moreover, traditional assumptions concerning character polarity have in some cases been shown to be incorrect.

Chromosome studies on eight species of Eryngium L. (Saniculoideae, Apiaceae) from Argentina

Abstract A cytological analysis of eight Argentinean species of Eryngium L. (Saniculoideae, Apiaceae) is carried out. Karyotypes of eight species and original chromosome counts for three of them (indicated with an asterisk) are notified: E. coronatum*Hook. et Arn. (2n=2x-16; 14m + 2sm), E. echinatum Urb. (2n=2x=16; 12m + 2m-sm + 2sm), E. nudicaule Lam. (2n=2x=14; 8m + 2m-sm + 4sm), E. eburneum Decne. (2n=2x=16; 8m + 2m-sm + 6sm), E. horridum Malme (2n=2x=16; 12m + 4sm), E. megapotamicum*Malme (2n=4x=32; 4m + 6m-sm + 22sm), E. mesopotamicum* Pedersen (2n=6x=48; 30m + 16sm +2st), and E. pandanifolium Cham. et Schlechtd. (2n=6x=48; 26m + 4m-sm + 14sm + 4st). The first three species belong to the Section Foetida, while the remaining five, belong to Panniculata. Both Sections are easily differentiated by morphology; our chromosomal study shows that these sections can also be recognized karyologically. All the species, except E. nudicaule (x=7), present x=8 which is the most common basic chromosome number in the genus and in the subfamily Saniculoideae. The karyotype analysis made on the eight species mainly shows metacentric and submetacentric chromosomes differing in proportion between species; only E. pandanifolium and E. mesopotamicum show subtelocentric pairs. These two species only differ in the color of their inflorescences; besides, differences in their karyotypes were negligible. These facts agree with the suggestion that they would be varieties of the same species. Two different phenomena seem to have occurred during karyotype evolution in the genus Eryngium: aneuploidy within Section Foetida, and polyploidy within Section Panniculata.

TAXONOMIC REVISION OF ALOYSIA (VERBENACEAE, LANTANEAE) IN SOUTH AMERICA

Abstract Aloysia Paláu is the third largest genus of tribe Lantaneae, after Lippia L. and Lantana L., in the Verbenaceae. Recent molecular phylogenetic studies have circumscribed genus Aloysia as 31 species, with the transfer of most species of Acantholippia Griseb. and the inclusion of the monotypic Xeroaloysia Tronc., as well as the exclusion of several North American Aloysia species that nest within a Lippia–Lantana clade. Newly circumscribed Aloysia are found mostly in South America, where the genus is represented by 28 species and six varieties. Only four Aloysia species are found in North America, A. coalcomana Siedo, A. macrostachya (Torr.) Moldenke, A. wrightii A. Heller, and A. gratissima (Gillies & Hook.) Tronc. var. gratissima, this last being the only taxon found in both North and South America. A taxonomic revision of the genus Aloysia for South America is provided with detailed morphological descriptions, as well as keys for taxonomic identification, illustrations or indication of iconography, and distribution and herbarium specimen lists. The genus Xeroaloysia Tronc. is here considered a synonym of Aloysia, and nine new taxonomic synonyms are here established. Lectotypification is designated for Verbena L. sect. Aloysioides Walp., and neotypification is designated for V. salviifolia Hook. & Arn.

Resolving the Genera Aloysia and Acantholippia within Tribe Lantaneae (Verbenaceae), using Chloroplast and Nuclear Sequences

Abstract Species belonging to the genera Aloysia and Acantholippia are difficult to place within Lantaneae due to gene tree incongruence and limited sampling in previous studies. We use an expanded sample of both genera, and DNA sequence data from six loci, to reveal that Aloysia and Acantholippia species occur in five consistently inferred, well-supported clades. The precise relationships of these clades to one another are still enigmatic, due to gene tree incongruence. However, coalescent-based species tree inference supports the inclusion of most of Acantholippia in an expanded Aloysia sensu lato, with a 4-lobed calyx as its defining feature. Five new combinations are proposed to reflect this relationship: Aloysia deserticola, Aloysia riojana, Aloysia salsoloides, Aloysia tarapacana, and Aloysia trífida. Geographic range shifts from subtropical South America to North America have occurred at least twice in Aloysia. Shifts between determinate and indeterminate inflorescence arrangement have occurred at least twice independently. The elongate, lax inflorescence, which is characteristic of most of Aloysia, is hypothesized to be derived from a condensed inflorescence.

Phylogenetic Position of the Monotypic Genus Verbenoxylum (Verbenaceae) and New Combination Under Recordia

Abstract In spite of the recent studies on the phylogeny of Verbenaceae, the position of the monotypic Verbenoxylum, endemic to the Atlantic rainforest in southeastern Brazil, remains unsolved. Molecular data were here analyzed to infer the phylogenetic placement of this genus; furthermore morphological data was studied in order to examine traits that support relationships among taxa. Sequences of the plastid regions of ndhF gene and trnL—trnF intergenic spacer were analyzed to conduct phylogenetic studies with maximum parsimony, maximum likelihood, and Bayesian inference. Morphological traits that had been traditionally used to distinguish tribes within Verbenaceae, as well as those employed to characterize Verbenoxylum, were examined. Verbenoxylum is nested within the tribe Duranteae, sister to Recordia, a monotypic genus endemic to Bolivia, a placement never reported before. The morphological traits analyzed prove not to be useful to distinguish tribes but are important at lower taxonomic levels. Based on the sister relationship and morphological similarities between the genera Verbenoxylum and Recordia, we propose the inclusion of Verbenoxylum reitzii into Recordia, forming the new combination Recordia reitzii .

A taxonomic revision of Casselia (Verbenaceae), a genus endemic to the South American Cerrado and Mata Atlántica biogeographic provinces1

Abstract A taxonomic revision of the genus Casselia is provided. Casselia is a small genus of Verbenaceae, placed in the tribe Casselieae, along with Tamonea and Parodianthus. It is represented by six species, distributed in South America, along central Brazil, eastern Bolivia and northern Paraguay. Five new synonyms and two lectotypifications are proposed. Detailed morphological descriptions are given for each taxon, as well as a key for their identification, illustrations, actualised synonymy, distribution, and a discussion about the relationship between closely related taxa.

Taxonomic revision of Ocimum (Lamiaceae) in Argentina1

Abstract In Argentina Ocimum is represented by four species: Ocimum campechianum, Ocimum nudicaule, Ocimum ovatum, and Ocimum selloi. Complete descriptions for each of these taxa and illustrations, as well as a key for their identification, together with distribution maps in Argentina are given. Four new synonyms are here proposed, and lectotypes are here designated for the following species: Lumnitzera carnosa, Ocimum balansae, Ocimum hassleri var. acutatum, Ocimum micranthum, Ocimum neurophyllum, O. nudicaule, O. ovatum, O. selloi, O. selloi var. angustifolium, O. selloi f. serratum, and O. selloi f. subintegrifolium.

Typification of the name Duranta mutisii (Verbenaceae)

Fil: Moroni, Pablo Daniel. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Instituto de Botanica Darwinion. Academia Nacional de Ciencias Exactas, Fisicas y Naturales. Instituto de Botanica Darwinion; Argentina