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Geographic distributions and inflorescence types mapped on to the *BEAST tree. A. Condensed inflorescence; terminal inflorescence absent. B. Lax inflorescence; terminal inflorescence absent. C. Lax inflorescence; terminal inflorescence present. D. Condensed inflorescence; terminal inflorescence absent. E. Condensed inflorescence; terminal inflorescence present.
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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-...
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... (Fig. 3), but this relation- ship is not found in the analyses of individual loci (Figs. S2- S4); they are not sister to one another in the species tree, but support for their positions is low (Fig. 4). At least two inde- pendent range shifts into North America are evident in Aloysia (excluding the species nesting within the Lantana-Lippia clade; Fig. 5), but it is unclear from these results whether North American distributions are due to northward migration via the Andes, or to long-distance dispersal. Individuals identified morphologically as A. scorodonioides and A. virgata do not form monophyletic groups, confirming the suspicion that the boundaries of these species are not yet ...
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... clade; Fig. 5), but it is unclear from these results whether North American distributions are due to northward migration via the Andes, or to long-distance dispersal. Individuals identified morphologically as A. scorodonioides and A. virgata do not form monophyletic groups, confirming the suspicion that the boundaries of these species are not yet Fig. 5), and occur in semi-arid to arid habitats in subtropical South America, near the borders between Argentina, Chile, and Bolivia. This clade is also predicted to include Acantholippia tarapacana Botta and Acantholippia riojana Hieron. ex Moldenke, which share morphological features (such as inflorescence arrangement and spiny branches) ...
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... TRIFIDA-This species is reconstructed as dis- crete from any other clade. It is superficially similar to mem- bers of the A. salsoloides clade, but lacks spines, and its condensed inflorescences are axillary only (homothetic pleiobotrya sensu O' Leary et al. 2012; Fig. 5). Acantholippia trifida is endemic to north-central Chile, ranging just across the border into ...
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... in a sister relationship with the Lantana-Lippia clade. It possesses xerophytic adaptations in common with other species of Acantholippia, such as reduced leaves, but several characters unite it morphologically with the Lantana-Lippia clade: its inflorescences are condensed and axillary only (homothetic pleiobotrya sensu O' Leary et al. 2012; Fig. 5), and its calyx is bilabiate (Botta 1980). Acantholippia seriphioides is widespread and abundant in dry habitats in southern ...
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... BARBATA AND RELATIVES-This subclade comprises five species (only three sampled here) with condensed inflo- rescences featuring conspicuous floral bracts, and bifid caly- ces, indicative of their common ancestry with the rest of the Lantana-Lippia clade (Fig. 5). It is unclear why these species have been considered members of Aloysia; the first so named was transferred from Lippia without accompanying justifica- tion by Moldenke (1940), who then described the remainder under Aloysia. All five are endemic to Mexico; two (A. nahuire Gentry & Moldenke and its segregate, A. coalcomana Siedo) are ...
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... pleiobotrya sensu O' Leary et al. (2012) are found in the Lantana-Lippia clade, the A. gratissima clade, A. catamarcensis + A. polystachya, and Acantholippia trifida (Fig. 5). This pattern was interpreted as resulting from two parallel losses of the terminal inflorescence, based on chloroplast topol- ogy and limited sampling, where one shift from heterothetic to homothetic pleiobotrya was interpreted as a synapomorphy for the A. gratissima clade + Lantana-Lippia clade (O' Leary et al. 2012). Our results, ...
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... inflorescence, based on chloroplast topol- ogy and limited sampling, where one shift from heterothetic to homothetic pleiobotrya was interpreted as a synapomorphy for the A. gratissima clade + Lantana-Lippia clade (O' Leary et al. 2012). Our results, based on increased data and sam- pling, suggest a more complicated pattern of trait evolution (Fig. 5), involving at least three shifts between heterothetic and homothetic ...
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... on the results presented here, the condensed inflo- rescence found in Acantholippia species, Aloysia polystachya + Aloysia catamarcensis, and the Lantana-Lippia clade (Fig. 5) is most parsimoniously interpreted as representative of the ancestral condition in core Lantaneae (excluding Coelocarpum). Both of our combined data analyses suggest that the lax inflo- rescence characteristic of Aloysia as traditionally circumscribed is derived twice independently: in the A. gratissima clade, and in the A. citrodora ...
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Citations
... Lantana species depicted a close phylogenetic relationship by grouping in the same clade on the basis of ccsA, matK, ndhF, rbcL, rpoC2, rps3 and trnL-F markers . Lu-Irving et al., (2014) also studied the close phylogenetic relationship of Lantana species based on the chloroplast regions trnT-trnL, rpl32-trnL and trnQ-rps16 and nuclear regions ETS, PPR 123 and PPR 81. The close relationship of Lantana was also studied by Thode et al., (2013) on the basis of chloroplast markers ndhF and trnL-trnF. ...
... Since the work of Marx et al. (2010), many publications based on morphology and/or molecular data have proposed important changes within the family, including several generic realignments. In tribe Duranteae, Verbenoxylum has been included within Recordia (Thode et al. 2013); in tribe Citharexyleae, Baillonia has been included within Citharexylum (Christenhusz et al. 2018); in tribe Lantaneae Acantholippia and Xeroaloysia were included in Aloysia (Lu-Irving et al. 2014); while Diphyllocalyx and Isidroa were described as new genera (Greuter & Rodríguez 2016), and in tribe Petreae, Xolocotzia has been included within Petrea (Christenhusz et al. 2018). The inclusion of Diostea in Dipyrena by Ravenna (2008) is contradicted by phylogenetic evidence and has not been accepted. ...
... These studies suggest the need for redefining the generic boundaries within Lantaneae. As mentioned above, Aloysia and related genera have already been recircumscribed (Lu-Irving et al. 2014;, with Aloysia now including taxa previously recognized as Acantholippia and Xeroaloysia. The remainder of Lantaneae forms a large clade comprising seven genera in which five from among them are embedded within the two large genera, Lantana and Lippia, neither of which is monophyletic (Lu-Irving et al. in press). ...
Background and aims-The last comprehensive study that estimated the number of Verbenaceae genera and species was published in 2004, and included 34 genera and around 1200 species. Since then, several publications based on morphology and/or molecular data have proposed important changes within the family. Due to the lack of updated literature to cite when referring to the number of Verbenaceae taxa, a review of these estimates is necessary. Key results and conclusion-We present a detailed list of genera currently accepted in Verbenaceae with the number of species contained in each and compare our numbers with the previous estimate. In addition, we indicate the geographic distribution and the most recent important taxonomic or phylogenetic works for each genus. Our compilation shows that Verbenaceae have 32 genera and 800 species currently accepted. This work provides up-to-date numbers and brings a holistic view of the family.
... Only A. gratissima is found in both hemispheres . Considered monophyletic, the genus is characterized by the morphological synapomorphy of an evenly 4-lobed calyx (Lu-Irving et al. 2014). Ten species are recorded in Brazil, all of them occurring in the Atlantic Forest, of which six are endemic (Moroni & O'Leary 2020a). ...
Verbenaceae is represented in Brazil by 15 genera and ca. 290 species, with most of its richness in the Cerrado and Atlantic Forest domains. The state of Espírito Santo has an area of 46,184.1 km 2 , wholly embedded in the Atlantic Forest. Nonetheless, due to the intense environmental degradation in the state, only 10.5% of its original vegetation remains. The present study is part of the "Flora do estado do Espírito Santo" project and aims to provide a taxonomic treatment to Verbenaceae in the state, as well as to analyze its distribution patterns, together with preliminary conservation assessments for each species. It is based on morphological analysis of herbaria collections, field expeditions, and literature compilation. Distribution maps were made for each species, while the preliminary conservation assessments followed the criteria proposed by the IUCN. Verbenaceae is represented in the state of Espírito Santo by 31 species arranged in 11 genera: Aloysia (2 spp.), Bouchea (1 sp.). Casselia (1 sp.), Citharexylum (1 sp.), Glandularia (1 sp.), Lantana (10 spp.), Lippia (4 spp.), Petrea (1 sp.), Priva (2 spp.), Stachytarpheta (6 spp.) and Verbena (2 spp.). Eight species occurring in Espírito Santo are threatened with extinction. Three new records are verified, including species of the genus Lantana. The dense rainforest, which covers most of the state's territory, presents the most significant number of species (20 spp.), followed by pioneer vegetations (12 spp.), seasonal semideciduous forests (11 spp.), inselbergs (8 spp.), and ecological refuges (2 spp.). The main richness centers for Verbenaceae in the state of Espírito Santo are the municipalities of Linhares, Santa Teresa, Vitória, Vila Velha, and Guarapari in this order. In contrast, the family is noticeably little represented in the northern region of the state, as well as near its borders, where the forest remnants are highly fragmented and scattered. The current results contribute to the taxonomic and biogeographic knowledge of Verbenaceae, to the creation of conservation strategies for threatened species in the State of Espírito Santo, and reinforce the need of fieldwork in several areas of the state.
... Lantaneae is a monophyletic tribe of Verbenaceae that currently comprises about 280 species, arranged in nine genera (Marx et al. 2010, Lu-Irving et al. 2014, Greuter & Rodríguez 2016. Lantana Linnaeus (1753: 626) and Lippia Linnaeus (1753: 633) are the largest genera of this tribe and comprise together approximately 80% of its species (Marx et al. 2010, Lu-Irving & Olmstead 2013. ...
Lantana and Lippia are two large and taxonomically complicated genera of Verbenaceae. During studies of these genera for the “Flora do Brasil 2020” project some taxonomic problems were identified. Here, we propose the reestablishment of the species Lippia subracemosa, nine new synonyms and three lectotypes for Lippia jangadensis, Lippia thymoides, and Lippia vernonioides, the last one also with the designation of an epitype.
... In this study, we explore the origin of four lineages of Verbenaceae ( Aloysia , Citharexylum , Glandularia , and Verbena ) found in North American arid habitats. Because Verbenaceae arose and diversifi ed fi rst in South America, North American lineages are all inferred to have South American ancestry ( Yuan and Olmstead, 2008a ;Marx et al., 2010 ;Lu-Irving and Olmstead, 2013 ;Olmstead, 2013 ;Lu-Irving et al., 2014 ). However, several biogeographic patterns are possible for the North American lineages, including (1) long-distance dispersal from South American arid zones, (2) a migration route connecting South American and North American arid zones, or (3) de novo evolution of plants in North American arid zones from wet tropical forest ancestors. ...
... However, several biogeographic patterns are possible for the North American lineages, including (1) long-distance dispersal from South American arid zones, (2) a migration route connecting South American and North American arid zones, or (3) de novo evolution of plants in North American arid zones from wet tropical forest ancestors. We review previous evidence from Lantaneae and Verbeneae ( Yuan and Olmstead, 2008a , b ;O'Leary et al., 2009 ;Lu-Irving et al., 2014 ) and present new evidence from Verbena and Citharexylum to improve our understanding of the origin of North American arid-zone Verbenaceae. ...
... Because this study integrates previously published results ( Aloysia : Lu-Irving et al., 2014 ), a previously studied group for which additional sampling was obtained (Verbeneae: Yuan and Olmstead, 2008a ;O'Leary et al., 2009 ), and a group ( Citharexylum ) not previously subject to phylogenetic analysis, sampling and DNA regions vary among the diff erent case studies. For a table of vouchers, herbaria housing voucher specimens, and GenBank accession numbers accompanying sequence data used in this study, see Appendix S1 in the Supplemental Data with this article. ...
Premise of the study:
Verbenaceae originated and initially diversified in South America in wet forest habitats. They have diversified extensively in arid habitats in both South and North America. This study aims to understand the origin of the North American arid-land members of Verbenaceae.
Methods:
A phylogenetic approach is used to examine four genera (Aloysia, Citharexylum, Glandularia, Verbena) in three distinct clades with representatives in North American deserts and disjunct South and North American distributions. Phylogenetic analyses were conducted using maximum likelihood and Bayesian approaches. Analyses included both plastid and nuclear DNA regions and include the first study of Citharexylum and an expanded sampling of tribe Verbeneae (Glandularia and Verbena). Ancestral areas were reconstructed for each group.
Key results:
North American desert species of Aloysia and Glandularia were likely derived from ancestors in arid temperate South America, perhaps by long-distance dispersal. The pattern for Verbena was less clear, with evidence from plastid DNA implicating an Andean dispersal route to the North American clade, whereas nuclear data suggest that the Andean and North American species resulted from independent dispersals from southern South America. A previously unrecognized clade of Andean Verbeneae was discovered, raising the possibility of an Andean origin of Verbena or Verbena and Glandularia. North American desert species of Citharexylum represent multiple, independent origins from mesic habitat ancestors in Mesoamerica.
Conclusions:
North American arid-zone Verbenaceae are derived from South and Central American ancestors via multiple avenues, including long-distance, amphitropical dispersal, Andean migration corridors, and in situ evolution of desert-adapted species.
... Similar convergent evolution of habit features has recently been revealed by a phylogenetic study of the Acantholippia-Aloysia complex and has led to the transfer to Aloysia of a number of species formerly included in Acantholippia Griseb. (Lu-Irving & al. 2014). Willdenowia 46 -2016 Nashia inaguensis, which provides the type of the generic name, has fruits of the same type as Lantana. ...
The seven species currently placed in the genus Nashia are revised, based on the near totality of the (scant) material found in the world's herbaria. A morphological analysis of, in particular, the calyx and fruit reveals the heterogeneity of that genus. As a result, Nashia is reduced to its single original species, N. inaguensis, described from Great Inagua, Bahamas (where it is here considered to have been introduced by man), and recently found on Puerto Rico and the Virgin Islands (St Croix). Nashia spinifera, of Hispaniola, is placed in Isidroa, a monotypic new genus. The five Cuban endemics, plus two newly described here, make up the new genus Diphyllocalyx, formerly Lippia sect. Diphyllocalyx of Grisebach. The formal treatment of all nine taxa comprises synonymy, type citation, detailed descriptions, specimen citation, distribution maps, and a key for identification. Possible evolutionary scenarios as well as conservation status are discussed. Citation: Greuter W. & Rankin Rodríguez R. 2016: Revision of the Caribbean endemics currently placed in Nashia (Verbenaceae) [Novitiae florae cubensis 50], — Willdenowia 46: 5–22. doi: http://dx.doi.org/10.3372/wi.46.46101 Version of record first published online on 4 January 2016 ahead of inclusion in April 2016 issue.
Recent phylogenetic results strongly suggest that neither Lantana nor Lippia are monophyletic. This has a direct impact on taxonomy and nomenclature in the tribe Lantaneae and it underscores the need to revise its classification. However, any generic recircumscription following the International Code of Nomenclature for algae, fungi, and plants will require substantial nomenclatural changes. As a solution, a phylogenetic classification following the PhyloCode is applied to the whole Lantaneae, and 28 rank‐free names are proposed and defined for clades. Clade / Lantaneae is defined to exclude Coelocarpum . Traditional taxon names previously recognized at the ranks of tribe, genus, and section are maintained when associated with clades. Discussion about fruit and corolla morphology, the basis of most previous classifications, within the group is presented along with a key to the clades of / Lantaneae . A useful classification of the plant diversity in tribe Lantaneae consistent with the phylogenetic relationships within the group is provided. This rank‐free system promotes name stability as new data become available and avoids name changes to well‐known species.
Premise:
As a family of Neotropical origin and primarily Neotropical distribution, the Verbenaceae are a good but understudied system with which to understand Neotropical evolution. Tribe Citharexyleae comprises three genera: Baillonia, Citharexylum-one of the largest genera in Verbenaceae-and Rehdera. A molecular phylogenetic approach was taken to resolve intergeneric relationships in Citharexyleae and infrageneric relationships in Citharexylum. The phylogeny is used to elucidate character evolution in a widespread, morphologically diverse Neotropical genus.
Methods:
Seven plastid regions, two nuclear ribosomal spacers, and six low-copy nuclear loci were analyzed for 64 species of Citharexyleae. Phylogenetic analyses were conducted using maximum likelihood, Bayesian inference, and multispecies coalescent approaches. Habit, presence or absence of thorns, inflorescence architecture, flower color, fruit color, and geography were examined to identify diagnostic character states for clades within Citharexylum.
Results:
Rehdera is resolved as sister to Citharexylum, and Baillonia nested within Citharexylum. Two species, C. oleinum and C. tetramerum, are not closely related to tribe Citharexyleae, but may be related to members of tribe Duranteae instead. Seven clades within Citharexylum are inferred, each characterized by a combination of geography, fruit color and/or maturation, and inflorescence architecture. There is evidence of correlated evolution between habit, axillary inflorescences, and flower number per inflorescence. Shrubs with reduced inflorescences have evolved repeatedly.
Conclusions:
A subgeneric classification for Citharexylum is proposed. Although suites of associated traits are found, character morphology has been labile throughout Citharexylum's evolutionary history. Morphological diversity may be related to adaptation to differing mesic and xeric habitats.
Premise:
Lantana and Lippia (Verbenaceae) are two large Linnean genera whose classification has been based on associated fruit traits: fleshy vs. dry fruits and one vs. two seed-bearing units. We reconstruct evolutionary relationships and the evolution of the two fruit traits to test the validity of these traits for classification.
Methods:
Previous studies of plastid DNA sequences provided limited resolution for this group. Consequently, seven nuclear loci, including ITS, ETS, and five PPR loci, were sequenced for 88 accessions of the Lantana/Lippia clade and three outgroups.
Results:
Neither Lantana nor Lippia is monophyletic. Burroughsia, Nashia, Phyla, and several Aloysia species are included within the clade comprising Lantana and Lippia. We provide a hypothesis for fruit evolution and biogeographic history in the group and their relevance for classification.
Conclusions:
Fleshy fruits evolved multiple times in the Lantana/Lippia clade and thus are not suitable taxonomic characters. Several sections of Lantana and Lippia and the small genera are monophyletic, but Lippia section Zappania is broadly paraphyletic, making circumscription of genera difficult. Lippia sect. Rhodolippia is a polyphyletic group characterized by convergence in showy bracts. Species of Lantana sect. Sarcolippia, previously transferred to Lippia, are not monophyletic. The clade originated and diversified in South America, with at least four expansions into both Central America and the Caribbean and two to Africa. The types species of Lantana and Lippia occur in small sister clades, rendering any taxonomy that retains either genus similar to its current circumscription impossible.
Aloysia Paláu is an important herbal genus from the Verbenaceae family and possesses numerous remedial properties in the folk medicine of Asian, European, and, in particular, South American countries. Only a few reports have discussed some phytochemical characteristics associated with Aloysia species. Right the lack of an exhaustive report prompted us to organize this review article. Accordingly, besides the ethnobotanical knowledge of Aloysia species, their essential oil profiles, phytochemistry of the polar isolated fractions, and the relevant biological activities are discussed in detail.
