Phylogenetic relationships within the fern genus Hymenophyllum s.l. (Hymenophyllaceae, Filicopsida): contribution of morphology and cytology

The phylogenetic relationships of Hymenophyllum and its segregate genera Cardiomanes, Hymenoglossum, Rosenstockia, Serpyllopsis and Microtrichomanes are addressed, using 31 morphological characters of the sporophyte and one cytological character. As expected, this study reveals considerable morphological heterogeneity within the genus sensu lato, but several apomorphic changes allow support for some clades. Four unresolved taxa, Cardiomanes, Hymenoglossum, Diplophyllum and Mecodium pro parte are probably the most basal elements in Hymenophyllum. The analysis also suggests the polyphyly of Mecodium, and two unexpected associations: Sphaerocionium together with Microtrichomanes; and a broad clade composed of subg. Hymenophyllum, Hemicyatheon and Craspedophyllum, genera Rosenstockia and Serpyllopsis, and subsect. Leptocionium and Amphipterum. These associations appear justified by morphological, cytological or geographical data, and most of them are in agreement with preliminary molecular results.     

A global molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy

To establish a basis for future taxonomic revisions and to infer the evolutionary traits of Trichomanes s.l. , one of two large filmy fern genera, molecular phylogenetic analyses were conducted using chloroplast rbcL sequences. Sampling covered most of the global distribution range of this genus throughout the tropical and temperate zones, as well as all taxonomically significant species by the addition of 51 samples principally from Asia. The evolution of eight selected characters, which were adopted as taxonomic keys and/or putatively reflect morphological regression, was optimized on the retrieved most parsimonious rbcL trees, together with new observations on the stem anatomy of 45 species. The eight robust clades retrieved within Trichomanes in the rbcL phylogeny conflict with the existing classifications. Our results also suggest that the cup-shaped sorus, the primary diagnostic character of the genus, appears in parallel in the Hymenophyllum lineage, as demonstrated by Pleuromanes (typified by Trichomanes pallidum ) included in the lineage. The suggestion that the regressive character states are homoplastic apomorphies in the evolution of the frond size, stem thickness, root system, and stem anatomy (stele and cortex) probably illustrates a strong tendency towards adaptive evolutionary transition in Trichomanes . © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 1–27.     

Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae)

Background and Aims

Pteris (Pteridaceae), comprising over 250 species, had been thought to be a monophyletic genus until the three monotypic genera Neurocallis, Ochropteris and Platyzoma were included. However, the relationships between the type species of the genus Pteris, P. longifolia, and other species are still unknown. Furthermore, several infrageneric morphological classifications have been proposed, but are debated. To date, no worldwide phylogenetic hypothesis has been proposed for the genus, and no comprehensive biogeographical history of Pteris, crucial to understanding its cosmopolitan distribution, has been presented.

Methods

A molecular phylogeny of Pteris is presented for 135 species, based on cpDNA rbcL and matK and using maximum parsimony, maximum likelihood and Bayesian inference approaches. The inferred phylogeny was used to assess the biogeographical history of Pteris and to reconstruct the evolution of one ecological and four morphological characters commonly used for infrageneric classifications.

Key Results

The monophyly of Pteris remains uncertain, especially regarding the relationship of Pteris with Actiniopteris + Onychium and Platyzoma. Pteris comprises 11 clades supported by combinations of ecological and morphological character states, but none of the characters used in previous classifications were found to be exclusive synapomorphies. The results indicate that Pteris diversified around 47 million years ago, and when species colonized new geographical areas they generated new lineages, which are associated with morphological character transitions.

Conclusions

This first phylogeny of Pteris on a global scale and including more than half of the diversity of the genus should contribute to a new, more reliable infrageneric classification of Pteris, based not only on a few morphological characters but also on ecological traits and geographical distribution. The inferred biogeographical history highlights long-distance dispersal as a major process shaping the worldwide distribution of the species. Colonization of different niches was followed by subsequent morphological diversification. Dispersal events followed by allopatric and parapatric speciation contribute to the species diversity of Pteris.     

Molecular phylogeny and evolution of Monilinia (Sclerotiniaceae) based on coding and noncoding rDNA sequences

The nuclear internal transcribed spacers, the 5.8S subunit, ~560 bp of the small subunit, and ~320 bp of the large subunit of the nuclear ribosomal DNA repeat from 17 species of Monilinia and eight species of closely related genera were sequenced. Phylogenies were constructed using maximum parsimony. The results support the hypothesis that Monilinia is not monophyletic. A fundamental distinction was found between the section Junctoriae and the section Disjunctoriae. Four evolutionary lineages were identified within the Disjunctoriae: one species on Crataegus, one group of species on dry stone fruits of rosaceous hosts, one group of species on capsular fruits of ericaceous hosts, and one group of species on sweet berry fruits of ericaceous hosts. Comparisons between branching topologies of hosts and Monilinia species suggest that although cospeciation among hosts and parasites has been the rule, several host jumps have taken place. Sclerotinia pirolae was determined to be a true member of the Disjunctoriae. The closest taxon groups to the Junctoriae were found to be Botrytis and Sclerotinia, with Ciborinia being the closest taxon group to the Disjunctoriae. There is evidence of an increased rate of ssrRNA evolution in the lineage of species that attack ericaceous berries.     

Molecular systematics and evolutionary history of the genus Carabus (Col. Carabidae)

Despite the number of evolutionary, ecological and conservation studies that are conducted on Carabus, the global evolutionary history of the genus remains poorly understood. Here, we analysed 7.5 kilobases of DNA sequence data (six mitochondrial and four nuclear genes) from a worldwide sample of 45% of the known subgenera (99 species and 14 subspecies). We compared the nuclear and mitochondrial phylogenies obtained from Maximum likelihood and Bayesian analyses through topological tests of congruence and dating analyses. Our results mostly corroborate the monophyly of the morphological subgroups of Carabus. However, current morphological and molecular data appear unable to accurately infer the deep branchings within the genus. We show that Carabus originated ca. 16.7-25.1Ma, approximately 25Myr later than previously estimated. Major groups of Carabus are subdivided into clades that diverged from each other in a relatively short period of time around 10Ma (6.6-14.8). This time frame suggests that the present-day distribution of Carabus subgroups may be explained by isolation resulting from Eurasian forest fragmentation brought on by Miocene climate changes and by mountain orogenesis. Finally, we highlight several conflicts between mitochondrial and nuclear topologies that may be explained by mitochondrial introgression.     

Molecular systematics of the genus Artibeus (Chiroptera: Phyllostomidae)

A molecular phylogeny of the genus Artibeus using 19 of the 20 recognized species, many with samples from a broad geographic range, is presented. The analysis shows a clear distinction between the two subgenera (or genera), the 'large'Artibeus and the 'small'Dermanura, in both mitochondrial and nuclear genes. The placement and status of A. concolor remains inconclusive and is presented as the third subgenus Koopmania. The phylogenies and divergence time estimates show a marked influence of the Andes in the formation of the subgenera and the main lineages inside each subgenus. Nuclear genes showed a highly incomplete lineage sorting among species inside subgenera Artibeus and Dermanura. Indeed, shared alleles were also found between Artibeus and Koopmania, which are presumed to have split apart during the Miocene, showing that great care should be taken in using these markers. Cytochrome-b gene divergences and monophyly analyses suggest that A. lituratus and A. intermedius are indeed conspecifics. These analyses also suggested the existence of at least four 'new' species revealing a significant cryptic diversity inside the genus.     

Molecular systematics of the genus Allodontichthys (Cyprinodontiformes: Goodeidae)

Phylogenetic analyses of cytochrome c oxidasesubunit I (627bp) and mitochondrial controlregion (338bp aligned) sequences for all knownspecies of Allodontichthys wereperformed. Allodontichthys was recoveredas monophyletic, and A. hubbsi wassupported as the most primitive member of thegenus in all analyses. Combined-data analysessuggest that A. polylepis is sister to aclade comprising A. tamazulae and A. zonistius; however, a strict consensus of COIand control region cladograms results in acollapse of this node. The two species thatare broadly sympatric, A. hubbsi and A. tamazulae, are not sister taxa.Allodontichthys was found the sistergroup of a clade including Ilyodon andXenotaenia. These three genera are theonly goodeids to occupy high-gradient riversystems. Ilyodon and Allodontichthys share similar distributions;however, Allodontichthys exhibits greaterthan an order of magnitude more geneticvariation in COI than Ilyodon incomparisons of individuals from the samelocalities in the Ameca and Armeria drainagesystems. This is interpreted to mean Allodontichthys arrived in some westerndrainages significantly before Ilyodon.The Rio Coahuayana appears to be the center oforigin of the genus Allodontichthys. This river basin contains the most primitivemember of the genus, and appears to haveconnected with the Mesa Central, the center of endemism of the Goodeidae. A calibrated rateof molecular change in COI reveals Allodontichthys began diverging approximately6.2 million years ago (mya). Remainingspeciation in the group appears to haveoccurred about 2.7 to 3.3 mya. The hypothesisthat a hard polytomy comprising A. zonistius, A. polylepis, and A. tamazulae could be explained by speciation ofA. polylepis and A. tamazulae fromdifferent A. zonistius ancestors ispresented. Stream capture is evident but notfrequent in the history of Allodontichthys, and speciation does notclearly correlate with documented geologicactivity of the Mexican Transvolcanic Belt.     

Molecular systematics and biogeography of the amphibious genus Littorella (Plantaginaceae)

Littorella (Plantaginaceae) is a disjunct, amphibious genus represented by three closely related species. Littorella uniflora occurs in Europe including Iceland and the Azores, L. americana is found in temperate North America, and L. australis grows in temperate South America. Littorella has been recognized in numerous floristic treatments, but its status as a genus has recently been questioned. Rahn (Botanical Journal of the Linnean Society 120: 145-198, 1996) proposed a new phylogeny for Plantaginaceae based on morphological, embryological, and chemical data in which he reduced Littorella to a subgenus of Plantago. This article compares the phylogeny proposed by Rahn to one based on DNA sequence data from the internal transcribed spacer (ITS) region. In our analysis, Littorella forms a strongly supported monophyletic clade sister to Plantago and its recognition at the generic rank appears warranted. Littorella australis is sister to L. americana, and this clade is sister to the European L. uniflora. This more distant relationship between L. uniflora and L. americana provides support for maintaining both taxa at the specific rank and suggests a European origin for Littorella. Our studies also indicate that the monotypic genus Bougueria is deeply nested within Plantago and that its inclusion within Plantago as proposed by Rahn appears justified.     

Molecular systematics and global phylogeography of angel sharks (genus Squatina)

Angel sharks of the genus Squatina represent a group comprising 22 extant benthic species inhabiting continental shelves and upper slopes. In the present study, a comprehensive phylogenetic reconstruction of 17 Squatina species based on two mitochondrial markers (COI and 16S rRNA) is provided. The phylogenetic reconstructions are used to test biogeographic patterns. In addition, a molecular clock analysis is conducted to estimate divergence times of the emerged clades. All analyses show Squatina to be monophyletic. Four geographic clades are recognized, of which the Europe–North Africa–Asia clade is probably a result of the Tethys Sea closure. A second sister group relationship emerged in the analyses, including S. californica (eastern North Pacific) and S. dumeril (western North Atlantic), probably related to the rise of the Panamanian isthmus. The molecular clock analysis show that both lineage divergences coincide with the estimated time of these two geological events.     

Periodicities in coding and noncoding regions of the genes

Gene population statistical studies of protein coding genes and introns have identified two types of periodicities on the purine/pyrimidine alphabet: (i) the modulo 3 periodicity or coding periodicity (periodicity P3) in protein coding genes of eukaryotes, prokaryotes, viruses, chloroplasts, mitochondria, plasmids and in introns of viruses and mitochondria, and (ii) the modulo 2 periodicity (periodicity P2) in the eukaryotic introns. The periodicity study is herein extended to the 5' and 3' regions of eukaryotes, prokaryotes and viruses and shows: (i) the periodicity P3 in the 5' and 3' regions of eukaryotes. Therefore, these observations suggest a unitary and dynamic concept for the genes as for a given genome, the 5' and 3' regions have the genetic information for protein coding genes and for introns: (1) In the eukaryotic genome, the 5' (P2 and P3) and 3' (P2 and P3) regions have the information for protein coding genes (P3) and for introns (P2). The intensity of P3 is high in 5' regions and weak in 3' regions, while the intensity of P2 is weak in 5' regions and high in 3' regions. (2) In the prokaryotic genome, the 5' (P3) and 3' (P3) regions have the information for protein coding genes (P3). (3) In the viral genome, the 5' (P3) and 3' (P3) regions have the information for protein coding genes (P3) and for introns (P3). The absence of P2 in viral introns (in opposition to eukaryotic introns) may be related to the absence of P2 in 5' and 3' regions of viruses.     

Phylogenetic relationships in the orchid genus Serapias L. based on noncoding regions of the chloroplast genome

A molecular phylogenetic analysis was performed on 14 species of the Mediterranean unrewarding orchid genus Serapias using sequences of four noncoding regions of chloroplast DNA. This study has led to a new interpretation of the evolutionary relationships in this genus. The well-defined phylogenetic tree supports a division of taxa into two main clades, each including two minor groups. The molecular relationships found in this study differ from those defined by traditional systematic morphological assessments. By comparing the variation in sequence to variations in floral traits, we propose that the split in the two main lineages reflects an early differentiation of flower size, perhaps due to the shift from allo- to self-pollination. Conversely, the relationships within each minor group do not reflect floral size variation; therefore, we presume that this diversification resulted from genetic drift, local selection forces, and multiple, independent transitions towards self-pollination and polyploidy.     

Molecular systematics and Holarctic phylogeography of cestodes of the genus Anoplocephaloides Baer, 1923 s. s. (Cyclophyllidea,Anoplocephalidae) in lemmings (Lemmus,Synaptomys)

The present molecular systematic and phylogeographic analysis is based on sequences of cytochrome c oxidase subunit 1 (cox1) (mtDNA) and 28S ribosomal DNA and includes 59 isolates of cestodes of the genus Anoplocephaloides Baer, 1923 s. s. (Cyclophyllidea, Anoplocephalidae) from arvicoline rodents (lemmings and voles) in the Holarctic region. The emphasis is on Anoplocephaloides lemmi (Rausch 1952) parasitizing Lemmus trimucronatus and Lemmus sibiricus in the northern parts of North America and Arctic coast of Siberia, and Anoplocephaloides kontrimavichusi (Rausch 1976) parasitizing Synaptomys borealis in Alaska and British Columbia. The cox1 data, 28S data and their concatenated data all suggest that A. lemmi and A. kontrimavichusi are both non‐monophyletic, each consisting of two separate, well‐defined clades, that is independent species. As an example, the sister group of the clade 1 of A. lemmi, evidently representing the ‘type clade’ of this species, is the clade 1 of A. kontrimavichusi. For A. kontrimavichusi, it is not known which one is the type clade. There is also fairly strong evidence for the non‐monophyly of Anoplocephaloides dentata (Galli‐Valerio, 1905)‐like species, although an earlier phylogeny suggested that this multispecies assemblage may be monophyletic. The results suggest a deep phylogenetic codivergence of Lemmus spp. and A. lemmi, primarily separating the two largely allopatric host and parasite species at the Kolyma River in east Siberia. There are also two allopatric sublineages within each main clade/species of A. lemmi and Lemmus, but the present distributions of the sublineages within the eastern L. trimucronatus and clade 1 of A. lemmi are not concordant. This discrepancy may be most parsimoniously explained by an extensive westward distributional shift of the easternmost parasite subclade. The results further suggest that the clade 1 of A. kontrimavichusi has diverged through a host shift from the precursor of L. trimucronatus to S. borealis.     

Molecular phylogenetics,character evolution and systematics of the genus Micranthes (Saxifragaceae)

With c. 85 species, the genus Micranthes is among the larger genera of the Saxifragaceae. It is only distantly related to the morphologically similar genus Saxifraga, in which it has frequently been included as Saxifraga section Micranthes. To study the molecular evolution of Micranthes, we analysed nuclear ribosomal (internal transcribed spacer, ITS) and plastid (trnL–trnF) DNA sequences in a comprehensive set of taxa comprising c. 75% of the species. The molecular phylogenetic tree from the combined dataset revealed eight well‐supported clades of Micranthes. These clades agree in part with previously acknowledged subsections or series of Saxifraga section Micranthes. As these eight groups can also be delineated morphologically, we suggest that they should be recognized as sections of Micranthes. New relationships were also detected for some species and species groups, e.g. section Davuricae sister to sections Intermediae and Merkianae, and M. micranthidifolia as a member of section Micranthes. Species proposed to be excluded from the genus Micranthes for morphological reasons were resolved in the molecular tree in Saxifraga. Many morphological characters surveyed were homoplasious to varying extents. Micromorphological characters support comparatively well the clades in the phylogenetic tree. An updated nomenclature and a taxonomic conspectus of sections and species of Micranthes are provided. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 47–66.     

Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae)

We present the first phylogenetic study on the widespread Middle American microhylid frog genus Hypopachus. Partial sequences of mitochondrial (12S and 16S ribosomal RNA) and nuclear (rhodopsin) genes (1275 bp total) were analyzed from 43 samples of Hypopachus, three currently recognized species of Gastrophryne, and seven arthroleptid, brevicipitid and microhylid outgroup taxa. Maximum parsimony (PAUP), maximum likelihood (RAxML) and Bayesian inference (MrBayes) optimality criteria were used for phylogenetic analyses, and BEAST was used to estimate divergence dates of major clades. Population-level analyses were conducted with the programs NETWORK and Arlequin. Results confirm the placement of Hypopachus and Gastrophryne as sister taxa, but the latter genus was strongly supported as paraphyletic. The African phrynomerine genus Phrynomantis was recovered as the sister taxon to a monophyletic Chiasmocleis, rendering our well-supported clade of gastrophrynines paraphyletic. Hypopachus barberi was supported as a disjunctly distributed highland species, and we recovered a basal split in lowland populations of Hypopachus variolosus from the Pacific versant of Mexico and elsewhere in the Mesoamerican lowlands. Dating analyses from BEAST estimate speciation within the genus Hypopachus occurred in the late Miocene/early Pliocene for most clades. Previous studies have not found bioacoustic or morphological differences among these lowland clades, and our molecular data support the continued recognition of two species in the genus Hypopachus.     

Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences.

Based on extensive phenetic analyses, bats of the genus Myotis have been classically subdivided into four major subgenera each of which comprise many species with similar morphological and ecological adaptations. Each subgenus thus corresponds to a distinct "ecomorph" encompassing bat species exploiting their environment in a similar fashion. As three of these subgenera are cosmopolitan, regional species assemblages of Myotis usually include sympatric representatives of each ecomorph. If species within these ecomorphs are monophyletic, such assemblages would suggest extensive secondary dispersal across geographic areas. Conversely, these ecomorphological adaptations may have evolved independently through deterministic processes, such as adaptive radiation. In this case, phylogenetic reconstructions are not expected to sort species of the same ecomorph into monophyletic clades. To test these predictions, we reconstructed the phylogenetic history of 13 American, 11 Palaearctic, and 6 other Myotis species, using sequence data obtained from nearly 2 kb of mitochondrial genes (cytochrome b and nd1). Separate or combined analyses of these sequences clearly demonstrate the existence of several pairs of morphologically very similar species (i.e., sibling species) which are phylogenetically not closely related. None of the three tested subgenera constitute monophyletic units. For instance, Nearctic and Neotropical species currently classified into the three subgenera were clustered in a single, well-supported monophyletic clade. These species thus evolved independently of their ecological equivalents from the Palaearctic region. Independent adaptive radiations among species of the genus Myotis therefore produced strikingly similar evolutionary solutions in different parts of the world. Furthermore, all phylogenetic reconstructions based on mtDNA strongly supported the existence of an unsuspected monophyletic clade which included all assayed New World species plus M. brandtii (from the Palaearctic Region). This "American" clade thus radiated into a morphologically diverse species assemblage which evolved after the first Myotis species colonized the Americas. Molecular reconstructions support paleontological evidence that species of the genus Myotis had a burst of diversification during the late Miocene-early Pliocene epoch.