Phylogenetics, biogeography, and staminal evolution in the tribe Mentheae (Lamiaceae)

? Premise of the study: The mint family (Lamiaceae) is the sixth largest family of flowering plants, with the tribe Mentheae containing about a third of the species. We present a detailed perspective on the evolution of the tribe Mentheae based on a phylogenetic analysis of cpDNA and nrDNA that is the most comprehensive to date, a biogeographic set of analyses using a fossil-calibrated chronogram, and an examination of staminal evolution. ? Methods: Data from four cpDNA and two nrDNA markers representing all extant genera within the tribe Mentheae were analyzed using the programs BEAST, Lagrange, S-DIVA, and BayesTraits. BEAST was used to simultaneously estimate phylogeny and divergence times, Lagrange and S-DIVA were used for biogeographical reconstruction, and BayesTraits was used to infer staminal evolution within the tribe. ? Key results: Currently accepted subtribal delimitations are shown to be invalid and are updated. The Mentheae and all five of its subtribes have a Mediterranean origin and have dispersed to the New World multiple times. The vast majority of New World species of subtribe Menthinae are the product of a single dispersal event in the mid-late Miocene. At least four transitions from four stamens to two stamens have occurred within Mentheae, once in the subtribe Salviinae, once in the subtribe Lycopinae, and at least twice in the subtribe Menthinae. ? Conclusions: Worldwide cooling trends probably played a large role in the diversification and present day distribution of the tribe Mentheae. Additional work is needed to ascertain relationships within some Mentheae genera, especially in the subtribe Menthinae.     

Palynological evolutionary trends within the tribe Mentheae with special emphasis on subtribe Menthinae (Nepetoideae: Lamiaceae)

The pollen morphology of subtribe Menthinae sensu Harley et al. [In: The families and genera of vascular plants VII. Flowering plants·dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae). Springer, Berlin, pp 167–275, 2004] and two genera of uncertain subtribal affinities (Heterolamium and Melissa) are documented in order to complete our palynological overview of the tribe Mentheae. Menthinae pollen is small to medium in size (13–43 μm), oblate to prolate in shape and mostly hexacolpate (sometimes pentacolpate). Perforate, microreticulate or bireticulate exine ornamentation types were observed. The exine ornamentation of Menthinae is systematically highly informative particularly at generic level. The exine stratification in all taxa studied is characterized by unbranched columellae. Orbicules are consistently absent in Menthinae. Our palynological data are interpreted in a phylogenetic context at tribal level in order to assess the systematic value of pollen characters and to evaluate the existing molecular phylogenies for this group. Pollen morphology suggests Heterolamium as a close relative of subtribe Nepetinae and supports the molecular affinity of Melissa to subtribe Salviinae.     

Staminal evolution in the genus Salvia (Lamiaceae): molecular phylogenetic evidence for multiple origins of the staminal lever

BACKGROUND AND AIMS: The genus Salvia has traditionally included any member of the tribe Mentheae (Lamiaceae) with only two stamens and with each stamen expressing an elongate connective. The recent demonstration of the non-monophyly of the genus presents interesting implications for staminal evolution in the tribe Mentheae. In the context of a molecular phylogeny, the staminal morphology of the various lineages of Salvia and related genera is characterized and an evolutionary interpretation of staminal variation within the tribe Mentheae is presented. METHODS: Two molecular analyses are presented in order to investigate phylogenetic relationships in the tribe Mentheae and the genus Salvia. The first presents a tribal survey of the Mentheae and the second concentrates on Salvia and related genera. Schematic sketches are presented for the staminal morphology of each major lineage of Salvia and related genera. KEY RESULTS: These analyses suggest an independent origin of the staminal elongate connective on at least three different occasions within the tribe Mentheae, each time with a distinct morphology. Each independent origin of the lever mechanism shows a similar progression of staminal change from slight elongation of the connective tissue separating two fertile thecae to abortion of the posterior thecae and fusion of adjacent posterior thecae. A monophyletic lineage within the Mentheae is characterized consisting of the genera Lepechinia, Melissa, Salvia, Dorystaechas, Meriandra, Zhumeria, Perovskia and Rosmarinus. CONCLUSIONS: Based on these results the following are characterized: (1) the independent origin of the staminal lever mechanism on at least three different occasions in Salvia, (2) that Salvia is clearly polyphyletic, with five other genera intercalated within it, and (3) staminal evolution has proceeded in different ways in each of the three lineages of Salvia but has resulted in remarkably similar staminal morphologies.     

A systematic histological study of palm fruits. V. Subtribe Archontophoeniciae (Arecaceae)

Pericarp histology in the Archontophoenicinae provides little to characterize the subtribe as a whole, revealing instead two separate trends with parallels in other subtribes of the Areceae. The data support a close relationship among the three genera occurring in New Caledonia:Chambeyronia, Actinokentia, andKentiopsis, in which there is a complex endocarp consisting of short, oblique fibrous bundles embedded in a thick mantle of brachysclereids, and a loose endocarp of heavily fibrous, flattened vascular bundles adjacent to a relatively thin locular epidermis. The data also support a close relationship between the two genera of the New Zealand/Tasman Sea region:Hedyscepe andRhopalostylis, in which the pericarp is more or less fibrous throughout, with purely fibrous bundles in the outer pericarp and heavily fibrous vascular bundles in the inner pericarp. These results confirm relationships revealed by other morphological data.Archontophoenix appears to be most like the New Caledonian genera in its pericarp structure, with a similar mantle of short fibrous bundles embedded in a a mantle of brachysclereids in the outer pericarp, although it differs significantly in other aspects of morphology and anatomy.     

Phylogenetic position of the genus Ferula (Apiaceae) and its placement in tribe Scandiceae as inferred from nrDNA ITS sequence variation

Recent molecular systematic investigations suggested that Ferula, an umbellifer genus of about 170 species, is polyphyletic, with its members placed in the apioid superclade and within tribe Scandiceae. We analyzed ITS sequence variation from 134 accessions of Apiaceae, including 83 accessions (74 species) of Ferula to ascertain the phylogenetic position of the genus within the family. Phylogenetic analyses of these data using maximum parsimony, Bayesian, and neighbor-joining methods support the monophyly of Ferula upon the addition of Dorema and Leutea (as Ferula sensu lato) and its placement in tribe Scandiceae. Ferula sensu is closely allied with other major lineages of Scandiceae, corresponding to subtribes Scandicinae, Daucinae, and Torilidinae. Therefore, we recognize the Ferula clade as subtribe Ferulinae. Another addition to tribe Scandiceae is a clade composed of genera Glaucosciadium and Mozaffariania. The three accessions of Ferula misplaced in the apioid superclade represent a species of Silaum.     

Phylogeny of Celastraceae tribe Euonymeae inferred from morphological characters and nuclear and plastid genes

The phylogeny of Celastraceae tribe Euonymeae (∼230 species in eight genera in both the Old and New Worlds) was inferred using morphological characters together with plastid (matK, trnL-F) and nuclear (ITS and 26S rDNA) genes. Tribe Euonymeae has been defined as those genera of Celastraceae with generally opposite leaves, isomerous carpels, loculicidally dehiscent capsules, and arillate seeds (except Microtropis). Euonymus is the most diverse (129 species) and widely cultivated genus in the tribe. We infer that tribe Euonymeae consists of at least six separate lineages within Celastraceae and that a revised natural classification of the family is needed. Microtropis and Quetzalia are inferred to be distinct sister groups that together are sister to Zinowiewia. The endangered Monimopetalum chinense is an isolated and early derived lineage of Celastraceae that represents an important component of phylogenetic diversity within the family. Hedraianthera is sister to Brassiantha, and we describe a second species (Brassiantha hedraiantheroides A.J. Ford) that represents the first reported occurrence of this genus in Australia. Euonymus globularis, from eastern Australia, is sister to Menepetalum, which is endemic to New Caledonia, and we erect a new genus (Dinghoua R.H. Archer) for it. The Madagascan species of Euonymus are sister to Pleurostylia and recognized as a distinct genus (Astrocassine ined.). Glyptopetalum, Torralbasia, and Xylonymus are all closely related to Euonymus sensu stricto and are questionably distinct from it. Current intrageneric classifications of Euonymus are not completely natural and require revision.     

Genetic divergence in the tribe Electronini (Myctophidae)

To reveal evolutionary relationships and the rate of divergence of lanternfishes in the tribe Electronini (Myctophidae), analysis of sequence of cox 1 mtDNA fragment in three genera (12 species) of the tribe and in a closely related genus Myctophum (9 species) was performed. The results support the tribe monophyly; however, the divergence within it appeared to be not profound: genetic distances between genera and subgenera are smaller than between species within Myctophum and other genera of myctophids. In subgenera of Protomyctophum, species formation has obviously not completed since interspecies distances in them (0.5–4.7%) are comparable to estimates of intraspecific variation in other lanternfishes. Minimal genetic differences were found between species P. (Hierops) arcticum and P. (H.) subparallelum, whose ranges at the present time are isolated in the northern and southern hemispheres. After calibration of the molecular clock, the time of isolation of these two species can be dated to the Middle Pleistocene. Weak tribe diversification indicates its evolutionary youth, which can be related to the absence of efficient hydrographic barriers for isolation. The data obtained do not permit to consider tribe members as the most primitive in the family Myctophidae.     

PEP car☐ykinase containing species in the Brachiaria group of the subfamily panicoideae

In the Gramineae, a survey of species among the Brachiaria group of the subfamily Panicoideae, tribe Paniceae revealed that they are PEP car?ykinase containing species. This group includes the genera Brachiaria, Eriochloa and Urochloa. With the exception of the genus Panicum, these are the only genera within the Panicoideae found to contain PEP car?ykinase species. It is suggested that the PEP car?ykinase species of the genus Panicum, P. fasciculatum, P. maximum, P. molle and P. texanum, might be best placed in the Brachiaria group.     

Fruit and Seed Anatomy of Chenopodium and Related Genera (Chenopodioideae,Chenopodiaceae/Amaranthaceae): Implications for Evolution and Taxonomy

A comparative carpological study of 96 species of all clades formerly considered as the tribe Chenopodieae has been conducted for the first time. The results show important differences in the anatomical structure of the pericarp and seed coat between representatives of terminal clades including Chenopodium s.str.+Chenopodiastrum and the recently recognized genera Blitum, Oxybasis and Dysphania. Within Chenopodium the most significant changes in fruit and seed structure are found in members of C. sect. Skottsbergia. The genera Rhagodia and Einadia differ insignificantly from Chenopodium. The evolution of heterospermy in Chenopodium is discussed. Almost all representatives of the tribe Dysphanieae are clearly separated from other Chenopodioideae on the basis of a diverse set of characteristics, including the small dimensions of the fruits (especially in Australian taxa), their subglobose shape (excl. Teloxys and Suckleya), and peculiarities of the pericarp indumentum. The set of fruit and seed characters evolved within the subfamily Chenopodioideae is described. A recent phylogenetic hypothesis is employed to examine the evolution of three (out of a total of 21) characters, namely seed color, testa-cell protoplast characteristics and embryo orientation.     

Phylogenetic relationships of Combretoideae (Combretaceae) inferred from plastid,nuclear gene and spacer sequences

Phylogenetic relationships of the subfamily Combretoideae (Combretaceae) were studied based on DNA sequences of nuclear ribosomal internal transcribed spacer (ITS) regions, the plastid rbcL gene and the intergenic spacer between the psaA and ycf3 genes (PY-IGS), including 16 species of eight genera within two traditional tribes of Combretoideae, and two species of the subfamily Strephonematoideae of Combretaceae as outgroups. Phylogenetic trees based on the three data sets (ITS, rbcL, and PY-IGS) were generated by using maximum parsimony (MP) and maximum likelihood (ML) analyses. Partition-homogeneity tests indicated that the three data sets and the combined data set are homogeneous. In the combined phylogenetic trees, all ingroup taxa are divided into two main clades, which correspond to the two tribes Laguncularieae and Combreteae. In the Laguncularieae clade, two mangrove genera, Lumnitzera and Laguncularia, are shown to be sister taxa. In the tribe Combreteae, two major clades can be classified: one includes three genera Quisqualis, Combretum and Calycopteris, within which the monophyly of the tribe Combreteae sensu Engler and Diels including Quisqualis and Combretum is strongly supported, and this monophyly is then sister to the monotypic genus Calycopteris; another major clade includes three genera Anogeissus, Terminalia and Conocarpus. There is no support for the monophyly of Terminalia as it forms a polytomy with Anogeissus. This clade is sister to Conocarpus. Electronic Publication     

Monophyly and relationships of the tribe Exaceae (Gentianaceae) inferred from nuclear ribosomal and chloroplast DNA sequences

Both chloroplast trnL (UAA) intron and nuclear ribosomal ITS sequences highly confirmed the monophyly of the tribes of the Gentianaceae defined by the recent classification, and revealed the tribe Exaceae as a basal clade just next to the basal-most lineage, the tribe Saccifolieae. Within the tribe Exaceae, Sebaea (except Sebaea madagascariensis) appeared as the most basal clade as the sister group to the rest of the tribe. The Madagascan endemic genera Gentianothamnus and Tachiadenus were very closely related to each other, together standing as sister to a clade comprising Sebaea madagascariensis, Ornichia, and Exacum. The saprophytic genus Cotylanthera nested deeply inside Exacum. Sebaea madagascariensis was shown closer to the Madagascan endemic genus Ornichia than to any other sampled Sebaea species. Exacum appeared as the most derived taxon within this tribe. The topology of the phylogenetic trees conform with the Gondwana vicariance hypothesis regarding the biogeography of Exaceae. However, no evidence for matching the older relationships within the family to the tectonic history could be corroborated with various divergence time analyses. Divergence dating estimated a post-Gondwana diverging of the Gentianaceae about 50 million years ago (MYA), and the tribe Exaceae as about 40 MYA. The Mozambique Channel land-bridge could have played an important role in the biogeographic history of the tribe Exaceae.     

Pericarp strength of sunflower and its value for plant defense against the sunflower moth,Homoeosoma electellum

Sunflower pericarps provide a barrier against seed feeding by larvae of the sunflower moth, Homoeosoma electellum. Pericarp hardening is thought to be accelerated by a phytomelanin layer beneath the hypodermis, but among germplasm with phytomelanin, broad variation in sunflower pericarp strength exists. To facilitate the use of pericarp strength in sunflower breeding, resistance to mechanical puncture was assessed for diverse sunflower germplasm, and feeding tests were used to evaluate whether differences in pericarp strength prevent H. electellum larvae from penetrating achenes. Test on field-grown sunflowers indicates that public restorer lines have lower pericarp strengths compared with maintainer lines and commercial hybrids at 14 days after the start of anthesis. Interspecific crosses or plant introduction (PI) accessions believed to be resistant to H. electellum, including PI 170415, comprised a group with exceptionally high pericarp strength relative to other germplasm. In subsequent tests on greenhouse-grown sunflowers, overall results were similar, but using field-grown plants provided greater statistical power. In choice tests with achenes that differed in pericarp strength, 7- to 9-day-old sunflower moth larvae fed more often on seed protected by a weaker pericarp, at a more than 5-to-1 ratio, while 10-day-old larvae fed indiscriminately. Pericarp strength data contradict previous published results for individual entries and heterotic groups, but support the generalization that improved physical resistance to the sunflower moth is possible. To use pericarp strength in PI 170415 or similar sources, the inheritance of high pericarp strength and potential trade-offs between pericarp strength and other agronomic traits need to be understood.     

Flavonoid patterns in Leptodactylon and Linanthus

Twenty-seven flavonoids were found among three species of Leptodactylon and sixteen species of Linanthus, of which only three were identical between the two genera. This argues strongly for the maintenance of two genera; however, the underlying similarities in coumarins, flavonol glycosides, chrysoeriol and glycoflavones suggest that the two genera are closely related. The flavonoid data also suggest that Leptodactylon and Linanthus of the tribe Gilieae may actually be closer to Phlox and Microsteris of the tribe Polemonieae, than to other Gilieae.     

Salvia (Lamiaceae) is not monophyletic: implications for the systematics, radiation, and ecological specializations of Salvia and tribe Mentheae

Salvia, with over 900 species from both the Old and New World, is the largest genus in the Lamiaceae. Unlike most members of the subfamily Nepetoideae to which it belongs, only two stamens are expressed in Salvia. Although the structure of these stamens is remarkably variable across the genus, generally each stamen has an elongate connective and divergent anther thecae, which form a lever mechanism important in pollination. In a preliminary investigation of infrageneric relationships within Salvia, the monophyly of the genus and its relationship to other members of the tribe Mentheae were investigated using the chloroplast DNA regions rbcL and trnL-F. Significant conclusions drawn from the data include: Salvia is not monophyletic, Rosmarinus and Perovskia together are sister to an Old World clade of Salvia, the section Audibertia is sister to subgenus Calosphace or the monotypic Asian genus Dorystaechas, and the New World members of section Heterosphace are sister to section Salviastrum. Owing to the non-monophyly of Salvia, relationships at the next clearly monophyletic level, tribe Mentheae, were investigated.     

Analysis of the taxonomic structure of the crane fly family Limoniidae (Diptera) based on the larval characters

A morphological study of the larval characters in the genera Elephantomyia, Helius, Microlimonia, Lipsothrix, and Teucholabis was conducted. Significant differences between Elephantomyia and Helius were discovered; Elephantomyia should be placed in the subfamily Hexatominae, while Helius cannot be assigned to any known subfamily. The genus Microlimonia has nothing in common with other representatives of the tribe Limoniini, and the genera Lipsothrix and Teucholabis, with those of Gonomyini; these genera cannot be included in any of the existing tribes. The presently accepted subdivision of the Limoniidae into 4 subfamilies does not reflect its actual diversity.     

天门冬科黄精族细胞学研究进展

在全面收集和整理黄精族染色体数据的基础上,对国内外有关黄精族各类群间的染色体数目和倍性的变化规律进行了总结,并从染色体的多倍化和非整倍化与系统发育关系和地理分布方面探讨了黄精族内各属的起源和演化关系问题。黄精族包括黄精属、舞鹤草属、异黄精属和竹根七属,共约100余种,其中舞鹤草属(x=18)、异黄精属(x=16)和竹根七属(x=20)的染色体基数稳定,而黄精属染色体基数波动较大,主要为x=8~16,既有多倍化也有非整倍化现象。染色体数据表明黄精族4个属的染色体进化模式各不相同,揭示了黄精族内染色体从高基数向低基数演化的规律;各属内染色体的演化主要是体现在二倍体水平上的核型变异,多倍化在本族中不占主导地位;仅黄精属内伴有非常强烈的非整倍化现象;细胞学证据与分子系统发育的结果比较吻合,为黄精族内属间以及属下的系统发育与进化提供了重要的参考资料。     

Pericarp structure of the Caryopteris group (Lamiaceae subfam. Ajugoideae)

Pericarp and seed structure was investigated in 13 out of the 16 species of the old Caryopteris s.l. and in a few species of the related Rubiteucris , Amethystea , Trichostema and Ajuga . The pericarp structure was found to be very variable, and the variation largely matches the most recent classification where Caryopteris s.l. is divided into six genera: Caryopteris s.s., Discretitheca , Pseudocaryopteris , Rubiteucris , Schnabelia and Tripora . All the six genera but Rubiteucris and Schnabelia can be distinguished from each other by differences in pericarp structure, and Caryopteris s.s. was found to have a particularly divergent structure. Similarities in pericarp structure also suggest that Schnabelia , Rubiteucris and Tripora may be closely related.     

A systematic histological study of palm fruits. VIII. Subtribe Dypsidinae (Arecaceae)

Analysis of the pericarp structure in the four genera of the palm subtribe Dypsidinae reveals tissues similar to those in other taxa within the pseudomonomerous Indo-Pacific arecoid palms, but generally in unspecialized configurations consistent with their presumed basal position within this group. Unique tissues within some members of genus Dypsis include thin-walled, tannin-filled fibers around the vascular bundles. Large-fruited members of the presumably related genera Lemurophoenix, Masoala, and Marojejya show more distinctive arrangements of protective tissues and are quite different from one another. Only Marojejya appears to be closely related to Dypsis. Lemurophoenix and Masoala, by possession of both unsheathed vascular bundles and bundles with heavy fibrous sheaths, show possible affinities with genera well-removed from Dypsis.     

Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al. (1988): phylogenetic systematics and revised classification

Taxonomic schemes for the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al., (1988) have been unstable due to the large number of genera and the paucity of known reliable characters. Reliable characters are essential when using phylogenetic inference in developing a natural classification. Morphological and developmental studies using light, scanning and transmission electron microscopy have revealed the new characters of host response, en face patterns, phasmid structure and female cuticular layers. These techniques also gave us insight into the homoplasy and polarity of many characters, revealed previously undetected character states and clarified misinterpreted character states. A matrix with the 19 most reliable characters is proposed for 20 operational taxonomic units (OTUs) and we employ this matrix for comparing computer generated phylogenetic analyses of the PHYLIP and PAUP packages. PAUP was deemed the more reliable parsimony algorithm for phylogenetic analysis of the Heteroderinae (Fink, 1986; Platnick, 1987). Monophyly of Atalodera + Sherodera + Thecavermiculatus (tribe Ataloderini), and Cactodera + Heterodera + Afenestrata, as well as Punctodera + Globodera + Dolichodera is supported by both programs. Most importantly, analyses strongly support monophyly of all cyst-forming genera (tribe Heteroderini) contrary to previous hypotheses of repeated evolution of the cyst (Wouts, 1985). In addition, monophyly of the Heteroderini with the Ataloderini is demonstrated. PAUP indicates monophyly of Sarisodera + Rhizonema + Bellodera + Hylonema and Ekphymatodera (tribe Sarisoderini new rank). Monophyly of the Sarisoderini was at first only weakly supported, but, subsequently, the reduced width of the submedial lips of second stage juveniles and males was recognized as a synapomorphy which strengthened subsequent PAUP trees and monophyly of the tribe. The present study rejects as paraphyletic or polyphyletic several previously proposed combinations, including Thecavermiculatus sequoiae (versus Rhizonema sequoiae), Sarisodera africana (versus Afenestrata africana), Dolichodera andinus (versus Thecavermiculatus andinus). The question whether T. andinus is a distinct genus, was not resolved due to insufficient data. PAUP supports our previous observations that Cactodera betulae is intermediate in a transformation series between other Cactodera and Heterodera: it also indicates these species as bring monophyletic with Heterodera + Afenestrata, but not with other Cactodera. Although these phylogenetic analyses strongly support some relationships, they indicate unresolved alternative hypotheses for others. Meloidodera (tribe Meloidoderini) and Cryphodera (tribe Cryphoderini) must be investigated for consideration of a possible synapomorphy not included in the present data matrix. Future studies are proposed to more clearly define the monophyly of the Heteroderini, as well as the Sarisoderini. Tests are also proposed to clarify questions of the monophyly of Verutus (tribe Verutini new rank) with the Heteroderinae versus other Tylenchida.