Testing eco-evolutionary predictions using fossil data: Phyletic evolution following ecological opportunity*

Fossil sequences provide observations of phenotypes within a lineage over time and represent essential data for increasing our understanding of phyletic evolution beyond microevolutionary timescales. I investigate if fossil time series of the diatom Stephanodiscus niagarae/yellowstonensis follow evolutionary dynamics compatible with hypotheses for how the adaptive landscape changes when a population enters a new environment. The lineage—which has a remarkably detailed stratigraphic record—invaded Yellowstone Lake immediately after recession of ice from the basin 14,000 years ago. Several phyletic models portraying different types of evolutionary dynamics—both compatible and not compatible with changes in the adaptive landscape following ecological opportunity—were fitted to the fossil times-series of S. niagarae/yellowstonensis. Different models best describe the three analyzed traits. Two of the models (a new model of decelerated evolution and an Ornstein–Uhlenbeck model) capture trait dynamics compatible with an event of ecological opportunity, whereas the third model (random walk) does not. Entering a new environment may accordingly affect trait dynamics for thousands of years, but the effects can vary across phenotypes. However, tests of model adequacy reveal shortcomings in all three models explaining the trait dynamics, suggesting model development is needed to more fully understand the phyletic evolution in S. niagarae/yellowstonensis.     

FIVE NEW GOMPHONEMA SPECIES (BACILLARIOPHYCEAE) FROM RIVERS IN SOUTH AFRICA AND SWAZILAND1

Diatom material from South Africa and Swaziland was examined with light and scanning electron microscopy. Five new taxa are proposed: Gomphonema crocodilei, G. quasicrocodilei, G. venusta, G. latistigmata, and G. cholnokyi. Gomphonema venusta has been previously misidentified as Gomphonema clevei Fricke and G. cholnokyi as G. subclavatum (Grunow) Grunow. All of the species possess biseriate striae, areolae with external vela, a true stigma, and marginal laminae underlying the valve margin and mantle. Compared to the other members of gomphonemoid lineage, the five new species show greatest similarity to Gomphoneis mesta Passy-Tolar & Lowe, G. magna Kociolek & Stoermer, and G. rhombica (Fricke) Merino, García, Hernández-Mariné, & Fernández and to the Herculeana lineage of Gomphoneis sensu Kociolek and Stoermer. The most recent phylogenetic analysis of the genus Gomphoneis showed that the two lineages of the genus. Herculeana and Elegans, are not monophyletic. The Herculeana lineage is more closely allied to Gomphonema than to the Elegans lineage. Since the Elegans lineage contains the generitype of Gomphoneis (G. elegans (Grunow) Cheve), and the species described here are more closely related to the Herculeana lineage, we have assigned them to Gomphonema.     

Multilocus phylogeny and taxonomy of East Asian voles Alexandromys (Rodentia,Arvicolinae)

Phylogenetic relationships, taxonomy and nomenclature issues within East Asian voles Alexandromys were addressed using comprehensive species samples, including all 12 valid species of the genus, and multilocus analysis. We examined the mitochondrial cytochrome b (cytb) gene and three nuclear genes in 36 specimens. Additionally, we examined a data set of 106 specimens using only the cytb gene. Our results did not confirm the aggregation of A. kikuchii, A. montebelli and A. oeconomus into a separate clade, namely the subgenus Pallasiinus. Analysis of incomplete lineage sorting using JML software highlighted both the cases of mitochondrial introgression and incomplete lineage sorting within the genus. Thus, the sister position of A. sachalinensis and A. maximowiczii in mitochondrial trees could be explained by mitochondrial introgression, while the sister position of A. limnophilus and A. fortis in mitochondrial trees could be successfully explained by incomplete lineage sorting. Very short genetic distances, together with an absence of monophyly, of the three species, A. evoronensis, A. mujanensis and A. maximowiczii, is supported by multiple morphological data, which indicates that these three taxa should be one species—A. maximowiczii. Analysis of genetic distances and tree topology revealed that three species of short‐tailed voles—A. middendorffii, A. mongolicus and A. gromovi—are more closely related to each other than to other established species of Alexandromys. The lacustrine vole, A. limnophilus, is closely related to the group of short‐tailed voles. Analysis of the type specimens of limnophilus and flaviventris confirmed that these taxa form one species together with A. l. malygini. Our results suggest that the mountains of western Mongolia are inhabited by a new taxon of short‐tailed voles of the same rank as middendorffii, mongolicus and gromovi—A. m. alpinus ssp. n.     

GRACILARIOPSIS MCLACHLANII SP. NOV. AND GRACILARIOPSIS PERSICA SP. NOV. OF THE GRACILARIACEAE (GRACILARIALES,RHODOPHYCEAE) FROM THE INDIAN OCEAN1

Two new species of Gracilariopsis from the Indian Ocean are proposed—Gracilariopsis (Gp.) mclachlanii Buriyo, Bellorin et M. C. Oliveira sp. nov. from Tanzania and Gracilariopsis persica Bellorin, Sohrabipour et E. C. Oliveira sp. nov. from Iran—based on morphology and DNA sequence data (rbcL gene and SSU rDNA). Both species fit the typical features of Gracilariopsis: axes cylindrical throughout, freely and loosely ramified up to four orders, with an abrupt transition in cell size from medulla to cortex, cystocarps lacking tubular nutritive cells and superficial spermatangia. Nucleotide sequence comparisons of rbcL and SSU rDNA placed both species into the Gracilariopsis clade as distinct species from all the accepted species for this genus, forming a deeply divergent lineage together with some species from the Pacific. The new species are very difficult to distinguish on morphological grounds from other species of Gracilariopsis, stressing the importance of homologous molecular marker comparisons for the species recognition in this character‐poor genus.     

Msx1creERT2 knock‐In allele: A useful tool to target embryonic and adult cardiac valves

Summary : Heart valve development begins with the endothelial‐to‐mesenchymal transition (EMT) of endocardial cells. Although lineage studies have demonstrated contributions from cardiac neural crest and epicardium to semilunar and atrioventricular (AV) valve formation, respectively, most valve mesenchyme derives from the endocardial EMT. Specific Cre mouse lines for fate‐mapping analyses of valve endocardial cells are limited. Msx1 displayed expression in AV canal endocardium and cushion mesenchyme between E9.5 and E11.5, when EMT is underway. Additionally, previous studies have demonstrated that deletion of Msx1 and its paralog Msx2 results in hypoplastic AV cushions and impaired endocardial signaling. A knock‐in tamoxifen‐inducible Cre line was recently generated (Msx1^CreERT2) and characterized during embryonic development and after birth, and was shown to recapitulate the endogenous Msx1 expression pattern. Here, we further analyze this knock‐in allele and track the Msx1‐expressing cells and their descendants during cardiac development with a particular focus on their contribution to the valves and their precursors. Thus, Msx1^CreERT2 mice represent a useful model for lineage tracing and conditional gene manipulation of endocardial and mesenchymal cushion cells essential to understand mechanisms of valve development and remodeling. genesis 53:337–345, 2015. © 2015 Wiley Periodicals, Inc.     

Taxonomy,phylogeography and climate relations of the Western Palaearctic spurge hawkmoth (Lepidoptera,Sphingidae, Macroglossinae)

Hundsdoerfer, AK., Mende, MB., Kitching, IJ. & Cordellier, M. (2011). Taxonomy, phylogeography and climate relations of the Western Palaearctic spurge hawkmoth (Lepidoptera, Sphingidae, Macroglossinae). —Zoologica Scripta, 40, 403–417. The primary aim of this study was to analyse mitochondrial genetic variability within the Hyles euphorbiae lineage in the Western Palaearctic, where observations of morphology have indicated much greater differentiation than is implied by the current taxonomy. The scope of the study is broadened further by including an analysis of the possible influences of climatic factors on lineage distributions. Mitochondrial sequences from three genes totalling an unambiguous alignment of 2284 bp were obtained for 356 individuals of the H. euphorbiae complex (HEC) s.s. Genetic admixture, network and climatic factor analyses were used to analyse the data. Unexpected mitochondrial variability was uncovered within the distribution range of H. euphorbiae in southern Europe, with two additional distinct clusters being found, both with rather patchy occurrences. The results of the genetic analysis presented here confirm a distinct H. cretica on Crete and the Dodecanese Islands and a distinct H. robertsi in Iran that hybridises with H. euphorbiae. Comparison of the molecular data with larval morphology shows only limited correlation of genetic clusters with pattern elements. The genetic network of the HEC s.s. shows strong incongruence with traditional taxonomy. Ecological niche modelling based on climate data revealed that the three best‐sampled clusters (H. tithymali, the well‐known H. euphorbiae lineage from Europe, and the previously unknown mitochondrial H. euphorbiae lineage that occurs mainly in Italy) could all occur in southern Italy and on the Mediterranean Islands. Biotic interactions appear to explain the observed ranges better than abiotic factors alone.     

IDENTIFICATION OF CRYPTIC SPECIES IN THE LESSONIA NIGRESCENS COMPLEX (PHAEOPHYCEAE,LAMINARIALES)1

The kelp Lessonia nigrescens Bory is the most ecologically and economically important seaweed in rocky intertidal and shallow subtidal habitats along the temperate Pacific South American coasts. Recent molecular studies suggest the existence of two lineages, one (northern lineage) from 17° S to 30° S and a second (central lineage) from 29° S to 41° S. To identify and name these lineages we performed morphological, nomenclatural and field studies. Four external and three internal anatomical traits permitted a morphological separation of the two lineages. The internal structure of both lineages was different from the isolectotype of Lessonia nigrescens. It is therefore concluded that the name Lessonia nigrescens should not be used for the Chilean material. Chordaria spicata Suhr appears as the oldest available name for the central lineage, while Lessonia berteroana Montagne is the oldest name for the northern lineage. In both cases, the type material consisted of small‐sized, apical branches of larger plants. The new combination Lessonia spicata (Suhr) Santelices is proposed for the central lineage and we reinstate Lessonia berteroana for the northern lineage. Laminaria scissa Suhr is reduced to synonym of L. spicata. Representative specimens of Lessonia nigrescens were not found during new visits to its type locality in Cape Horn and along Chile. Future studies should verify the status of this species.     

Phylogeography and cryptic variation within the Lacerta viridis complex (Lacertidae, Reptilia)

It is well known that the current genetic pattern of many European species has been highly influenced by climatic changes during the Pleistocene. While there are many well known vertebrate examples, knowledge about squamate reptiles is sparse. To obtain more data, a range‐wide sampling of Lacerta viridis was conducted and phylogenetic relations within the L. viridis complex were analysed using an mtDNA fragment encompassing part of cytochrome b, the adjacent tRNA genes and the noncoding control region. Most genetic divergence was found in the south of the distribution range. The Carpathian Basin and the regions north of the Carpathians and Alps are inhabited by the same mitochondrial lineage, corresponding to Lacerta viridis viridis. Three distinct lineages occurred in the south‐eastern Balkans — corresponding to L. v. viridis, L. v. meridionalis, L. v. guentherpetersi— as well as a fourth lineage for which no subspecies name is available. This distribution pattern suggests a rapid range expansion of L. v. viridis after the Holocene warming, leading to a colonization of the northern part of the species range. An unexpected finding was that a highly distinct genetic lineage occurs along the western Balkan coast. Phylogenetic analyses (Bayesian, maximum likelihood, maximum parsimony) suggested that this west Balkan lineage could represent the sister taxon of Lacerta bilineata. Due to the morphological similarity of taxa within the L. viridis complex this cryptic taxon was previously assigned to L. v. viridis. The distribution pattern of several parapatric, in part highly, distinct genetic lineages suggested the existence of several refuges in close proximity on the southern Balkans. Within L. bilineata sensu stricto a generally similar pattern emerged, with a high genetic diversity on the Apennine peninsula, arguing for two distinct refuges there, and a low genetic diversity in the northern part of the range. Close to the south‐eastern Alps, three distinct lineages (L. b. bilineata, L. v. viridis, west Balkan taxon) occurred within close proximity. We suggest that the west Balkan lineage represents an early offshoot of L. bilineata that was isolated during a previous Pleistocene glacial from the more western L. bilineata populations, which survived in refuges on the Apennine peninsula.     

Matrilineal Spirits,Descent and Territorial Power in Northern Thailand

This paper examines matrilineal spirits (phii puu nyaa) in northern Thailand based on field research recently undertaken in a village in Chiang Mai province. The paper suggests that the puu nyaa spirits remain locally significant—despite previous statements about their demise—and that matrilineal linkages are ideologically and practically important in the constitution of the groups. Nevertheless, there are alternative points of reference—to fathers, spouses and localities—that can attenuate attachments to matrilineal kin and introduce alternative sources of spiritual power. It becomes clear that phii puu nyaa spirit beliefs and practices are malleable and provide a basis for various orientations to spiritual power. The intermingling of different types of spiritual power is well illustrated by the presence of protective spirits locally referred to as aahak. In local perception, there is a very close relationship between phii puu nyaa and aahak, but the two entities appear to reflect quite different orientations: in simple terms the aahak represent a masculine, territorial and outward looking form of power which is contrasted with the female‐focused and lineage‐derived potency of the puu nyaa spirits. The paper argues that the outward orientation of the aahak provides some valuable insights into local perceptions of power, demonstrating how the supposedly peripheral and parochial draws regional power into more intimate domains.     

Phylogenetic relationships of the Chinese Labeoninae (Teleostei,Cypriniformes) derived from two nuclear and three mitochondrial genes

Zheng, L‐P., Yang, J‐X., Chen, X‐Y. & Wang, W‐Y. (2010) Phylogenetic relationships of the Chinese Labeoninae (Teleostei, Cypriniformes) derived from two nuclear and three mitochondrial genes. —Zoologica Scripta, 39, 559–571. The majority of genera within the Labeoninae occur in the South of China and the phylogeny of the subfamily Labeoninae has been a controversial topic over the years. The early and more recent phylogenetic results based on morphology are not in agreement, and some of the molecular analyses contradict those based on morphology. However, none of the previous studies has included an extensive sampling of Labeoninae genera. In this study, partial sequences of two nuclear (exon 3 of recombination activating protein 1 and rhodopsin) and three mitochondrial genes (cytochrome b, cytochrome oxidase subunit I and 16S ribosomal RNA) from 39 ingroup taxa and 11 outgroup taxa were used analysed to provide a hypothesis of phylogenetic relationships within the Chinese Labeoninae. The results supported the monophyly of Labeoninae and refuted the subdivision within Labeoninae based on the oromandibular morphology. It also further confirmed that the presence of a disc on lower lip arose through convergent evolution. Labeo was the basal clade and Osteochilus was closely related to Cirrhinus, forming the second basal clade. The monophyly of Garra and Bangana was refuted in this study, and they were both subdivided into two lineages. One lineage of Garra had closer relationships with Crossocheilus and Akrokolioplax, and the other lineage had closer relationships with Placocheilus. One lineage of Bangana represented the species with broadly interrupted postlabial groove; these species represented the true Bangana. The other lineage of Bangana included the species with continuous postlabial groove, and these species should be assigned to a new genus. Discogobio, as currently conceived, was paraphyletic, but it rendered monophyletic with the inclusion of Discocheilus. These results indicated that Discocheilus should be synonymised with Discogobio. Hongshuia and Sinocrossocheilus, respectively, formed independent lineages related to a lineage consisting of Pseudocrossocheilus plus seven other nominal genera. The relationships received weak support, but the repeatability of the clades could form the basis of recognising only the three genera mentioned above.     

Dr Jekyll and Mrs Hyde: Risky hybrid sex by amphibian‐parasitizing chytrids in the Brazilian Atlantic Forests

In their article in this issue of Molecular Ecology, Jenkinson et al. ( 2016 ) and colleagues address a worrying question—how could arguably the most dangerous pathogen known to science, Batrachochytrium dendrobatidis (Bd), become even more virulent? The answer: start having sex. Jenkinson et al. present a case for how the introduction into Brazil of the globally invasive lineage of Bd, BdGPL, has disrupted the relationship between native amphibians and an endemic Bd lineage, BdBrazil. BdBrazil is hypothesized to be native to the Atlantic Forest and so have a long co‐evolutionary history with biodiverse Atlantic Forest amphibian community. The authors suggest that this has resulted in a zone of hybrid Bd genotypes which are potentially more likely to cause fatal chytridiomycosis than either parent lineage. The endemic–nonendemic Bd hybrid genotypes described in this study, and the evidence for pathogen translocation via the global amphibian trade presented, highlights the danger of anthropogenic pathogen dispersal. This research emphasizes that biosecurity regulations may have to refocus on lineages within species if we are to mitigate against the danger of new, possibly hypervirulent genotypes of pathogens emerging as phylogeographic barriers are breached.     

Two new taxa of Achnanthidium and Encyonema (Bacillariophyceae) from the Yom River,Thailand, with special reference to the areolae occlusions implying ontogenetic relationship

We discovered two new taxa of Achnanthidium and Encyonema occurring with high abundance in the upstream of the Yom River, which is one of four major rivers in Northern Thailand. Achnanthidium pseudoconspicuum var. yomensis var. nov. closely resembles Achnanthidium pseudoconspicuum var. pseudoconspicuum in scanning electron microscopy (SEM), but has clearly different valve morphometry under the light microscopy (LM). Achnanthidium pseudoconspicuum var. yomensis is also similar to A. japonicum in valve morphology under LM, but differs to this species based on SEM observation. Encyonema yuwadeeanum sp. nov. shows similarities to Encyonema leei and Encyonema subkukenanum, but differs in valve shape and/or polar fissure shape. We discuss the possible valve ontogenetic relationship between Achnanthidium and Cymbellales based on the detailed areolae structures of these two new taxa.     

To be biased or to be Neotropical: systematic reassessment of a megadiverse lineage of rove-beetles (Philonthina,Staphylinini, Staphylininae)

Classifications in the world's tropics often involve an early and sustained adoption of Holarctic-based patterns. Such is the case of the megadiverse subtribe Philonthina and its Neotropical (NT) members, for which generic limits are ill-defined due to an alleged high level of homoplasy. Although a recent total-evidence study confirmed the monophyly of a NT lineage, most of its species are assigned to the speciose genera Belonuchus Nordmann and Paederomimus Sharp, neither of them monophyletic. Here, we aim to reveal internal relationships within the NT lineage by the reassessment of characters from traditional morphology-based systematics. Specific objectives are to test the monophyly of Belonuchus with regards to its only junior synonym, Musicoderus Sharp, as well as the placement of the six South American species of Hesperus Fauvel, a genus of Holarctic origin. We performed a phylogenetic analysis of the subtribe with focus on its NT lineage based on 132 morphological characters (50 of them novel) including 79 taxa from genera and/or species groups relevant to our study. Most novel characters assessed herein supported clades across Philonthina and its NT lineage. We found that the NT lineage diversified into at least seven clades, each of which provides a framework for future taxonomic studies. Among them, three clades containing the type species of Belonuchus, Paederomimus and Musicoderus (respectively) appear to be well supported and not closely related. The currently known South American species of Hesperus, however, are recovered within the NT lineage. We propose to resurrect Musicoderus from synonymy with Belonuchus and to transfer Hesperus novoteutonicus to Paederomimus as a new combination: Paederomimus novoteutonicus (Wendeler), comb.n .     

Post-larval and larval shells ofJuranomia Fürsich & Werner 1989, andAnomia Linnaeus 1758 (Anomüdae, Bivalvia)

Post-larval and larval shells ofJuranomia calcibyssata from the Bathonian to Callovian of Poland and RecentAnomia membranacea from the Mediterranean are described and compared to other fossil and Recent members of the family Anomüdae. The stratigraphic range of the monospecific genusJuranomia, which, up to now, was only known from the Kimmeridgian, can be extended to include the Lower Bathonian. The state of preservation of the fossil species allows recognition of an internal aragonitic, branching and complex cross-lamellar shell layer in the post-larval left valve, which previous studies could only assume to be present.Anomia membranacea is a member of theA. ephippium lineage as proved, among other characters, by the presence of an outer calcitic prismatic layer in its right valve. It possesses an anterior pedal retractor in the left valve, which, in the original discussion of the phylogenetic affinities ofJuranomia, was thought to be lacking in species ofAnomia. Consequently, the generaJuranomia andAnomia only differ in two important shell characters: closeness or distance between the three central muscles and thickness of the inner aragonitic shell layer of the left valve. Larval shells ofJuranomia are similar to those of Recent anomiids in shape, size, the presence of a byssal notch in the right valve, and an external sinus and internal shelly process in the left valve. The last three features are parts of a single character which is considered as an autapomorphy of the stem species of the Anomiiudae. The small P I size ofJuranomia calcibyssata suggests a purely planktic-planktotrophic development and thus, high potential of dispersal, just as its modern counterparts. Irrespective of the general similarity in shell size, the mean dimensions of the P II are likely species-specific.     

Description of Conticribra tricircularis,a new genus and species of Thalassiosirales,with a discussion on its relationship to other continuous cribra species of Thalassiosira Cleve (Bacillariophyta) and its freshwater origin

A new diatom genus Conticribra is erected to accommodate C. tricircularis, described from a freshwater Pliocene deposit in Trout Creek, Oregon (USA). The genus accommodates species possessing: (i) loculate areolae with (semi-) continuous cribra; (ii) non-plicated valve face; (iii) rimoportula located on the valve mantle, replacing a fultoportula. Conticribra tricircularis has no valve face fultoportulae and can easily be distinguished by its marginal fultoportulae with four satellite pores arranged in three rings. Three species are transferred to the new genus from Thalassiosira sensu lato. Using evidence from the fossil record and recent molecular data, a hypothesis concerning the freshwater origin of Conticribra is discussed.     

Phylogeny and systematics of Melanesia’s most diverse gecko lineage (Cyrtodactylus,Gekkonidae, Squamata)

Oliver, P.M., Richards, S.J. & Sistrom, M. (2012). Phylogeny and systematics of Melanesia’s most diverse gecko lineage (Cyrtodactylus, Gekkonidae, Squamata). —Zoologica Scripta, 41, 437–454. The systematics and biogeographical history of the diverse fauna of New Guinea and surrounding islands (Melanesia) remain poorly known. We present a phylogeny for 16 of the 21 recognised Melanesian bent‐toed geckos in the genus Cyrtodactylus based on mitochondrial sequence data. These analyses reveal two divergent lineages of Cyrtodactylus within Melanesia. One includes a single recognised species with clear affinities to sampled taxa from Asia. The other comprises a relatively diverse radiation (likely 30+ species), not closely related to sampled extralimital taxa and centred on the Melanesian region (including Australia). Many taxa within this second lineage are endemic to islands surrounding New Guinea, and dispersal and speciation on peripheral islands appears to have played an important role in the accumulation of species diversity within this clade. In contrast, little diversity is centred upon montane areas, although we do identify at least one lineage closely associated with hill and lower montane forest that probably dates to at least the late Miocene. Our phylogenetic analyses also reveal numerous divergent lineages that require taxonomic attention, including at least two widespread taxa that are likely to be composite, additional specimens of Cyrtodactylus capreoloides (until recently known only from the holotype) and several divergent and completely novel lineages, two of which we introduce herein: Cyrtodactylus arcanus sp. n. and Cyrtodactylus medioclivus sp. n.     

Bayesian Poisson tree processes and multispecies coalescent models shed new light on the diversification of Nawab butterflies in the Solomon Islands (Nymphalidae,Charaxinae, Polyura)

Butterflies of the genus Polyura form a widespread tropical group distributed from Pakistan to Fiji. The rare endemic Polyura epigenes Godman & Salvin, 1888 from the Solomon Islands archipelago represents a case of marked island polymorphism. We sequenced museum specimens of this species across its geographic range to study the phylogeography and genetic differentiation of populations in the archipelago. We used the Bayesian Poisson tree processes and multispecies coalescent models, to study species boundaries. We also estimated divergence times to investigate the biogeographic history of populations. Our molecular species delimitation and nuclear DNA network analyses unambiguously indicate that Malaita populations form an independent metapopulation lineage, as defined in the generalized lineage concept. This lineage, previously ranked as a subspecies, is raised to species rank under the name Polyura bicolor Turlin & Sato, 1995  stat. nov. Divergence time estimates suggest that this lineage split from its sister taxon in the late Pleistocene. At this time, the bathymetric isolation of Malaita from the rest of the archipelago probably prevented gene flow during periods of lower sea level, thereby fostering allopatric speciation. The combination of molecular species delimitation methods, morphological comparisons, and divergence time estimation is useful to study lineage diversification across intricate geographic regions.