The endocast from Dana Aoule North (DAN5/P1): A 1.5 million year-old human braincase from Gona,Afar, Ethiopia

The nearly complete cranium DAN5/P1 was found at Gona (Afar, Ethiopia), dated to 1.5–1.6 Ma, and assigned to the species Homo erectus. Its size is, nonetheless, particularly small for the known range of variation of this taxon, and the cranial capacity has been estimated as 598 cc. In this study, we analyzed a reconstruction of its endocranial cast, to investigate its paleoneurological features. The main anatomical traits of the endocast were described, and its morphology was compared with other fossil and modern human samples. The endocast shows most of the traits associated with less encephalized human taxa, like narrow frontal lobes and a simple meningeal vascular network with posterior parietal branches. The parietal region is relatively tall and rounded, although not especially large. Based on our set of measures, the general endocranial proportions are within the range of fossils included in the species Homo habilis or in the genus Australopithecus. Similarities with the genus Homo include a more posterior position of the frontal lobe relative to the cranial bones, and the general endocranial length and width when size is taken into account. This new specimen extends the known brain size variability of Homo ergaster/erectus, while suggesting that differences in gross brain proportions among early human species, or even between early humans and australopiths, were absent or subtle.     

A polyphasic approach to the delimitation of diatom species: a case study for the genus Pinnularia (Bacillariophyta)

Diatoms are one of the most abundant and arguably the most species‐rich group of protists. Diatom species delimitation has often been based exclusively on the recognition of morphological discontinuities without investigation of other lines of evidence. Even though DNA sequences and reproductive experiments have revealed several examples of (pseudo)cryptic diversity, our understanding of diatom species boundaries and diversity remains limited. The cosmopolitan pennate raphid diatom genus Pinnularia represents one of the most taxon‐rich diatom genera. In this study, we focused on the delimitation of species in one of the major clades of the genus, the Pinnularia subgibba group, based on 105 strains from a worldwide origin. We compared genetic distances between the sequences of seven molecular markers and selected the most variable pair, the mitochondrial cox1 and nuclear encoded LSU rDNA, to formulate a primary hypothesis on the species limits using three single‐locus automated species delimitation methods. We compared the DNA‐based primary hypotheses with morphology and with other available lines of evidence. The results indicate that our data set comprised 15 species of the P. subgibba group. The vast majority of these taxa have an uncertain taxonomic identity, suggesting that several may be unknown to science and/or members of (pseudo)cryptic species complexes within the P. subgibba group.     

Geographic and stratigraphic distributions of the Caribbean species of Cladocora (Scleractinia, Faviidae)

A complete account of the faviid genus Cladocora within the Caribbean is presented. In the Caribbean this genus represents an extant group that had its earliest occurrence during the Campanian-Maastrichtian of Jamaica. Recent forms have been reported throughout the Caribbean. The following forms were found (with stratigraphic ranges in the Caribbean): C. arbuscula (Pliocene-Recent), C. debilis (Pleistocene-Recent), C. gracilis (Middle-Upper Maastrichtian), C. jamaicaensis (Campanian-Maastrichtian and Eocene), C. johnsoni (Pliocene), and C. recrescens (Middle-Upper Oligocene). The occurrence of the genus Cladocora in the Caribbean is largely continuous from the Campanian to Recent, during the majority of the Caribbean species show affinities to European assemblages. For the time intervals Paleocene, Lower Oligocene, and Miocene the taxon has not been reported from the Caribbean.     

Systematics and phylogeny of Vasseuromys (Mammalia,Rodentia, Gliridae) with a description of a new species from the late Miocene of eastern Europe

Vasseuromys is a species‐rich genus of small‐ to medium‐sized glirids spanning the latest Oligocene to late Miocene of Europe and western Asia. Despite extensive discoveries over the past 50 years, little phylogenetic work has been done on Vasseuromys. This study presents the first phylogenetic analysis of the genus that includes all the described species and a new taxon Vasseuromys tectus sp. nov. from the late Miocene of eastern Europe, providing the first insights into the evolutionary relationships within the clade. Results suggest that the genus is clearly paraphyletic. Two strongly supported genus‐level clades are recognized within ‘Vasseuromys’: a restricted Vasseuromys clade (containing the three species, V. pannonicus, V. rugosus and V. tectus) and the Glirulus clade that includes ‘Vasseuromys’ duplex. The remaining ‘Vasseuromys’ species are found to constitute a set of paraphyletic taxa, with the polyphyletic ‘Ramys’ nested within it. The genus Gliruloides is synonymized with Glirulus. Vasseuromys tectus sp. nov. is the most derived member of the genus in having a greater number of cheek teeth ridges including constantly present anterotrope, centrotrope, second prototrope on M1–2, third metatrope on M2, two to three posterotropids on p4 and strong ectolophids on lower molars. The results of the study confirm a European origin for Vasseuromys while suggesting that the late Miocene species of the genus dispersed from the east in the early Turolian.     

Phylogenetic affinities and species limits within the genus Megadontomys (Rodentia: Cricetidae) based on mitochondrial sequence data

The evolutionary history of the genus Megadontomys, a group of mice allopatrically distributed along the cool‐humid forest in the highlands of México, is controversial. In this study, we examined phylogenetic relationships within the genus using sequences data from the complete mitochondrial cytochrome b gene. This information also allowed us to corroborate species limits, geographic boundaries of taxonomic entities and assess genetic variation within each taxon. The results of the phylogenetic analyses based on maximum likelihood, Bayesian inference and maximum parsimony were largely congruent in that M. nelsoni and M. thomasi were more closely related relative to M. cryophilus. These results are concordant with previous studies based on morphology and allozyme variation. However, testing of the alternative hypothesis of a closer evolutionary affinity between M. nelsoni–M. cryophilus did not produce a significantly less likely tree. The lack of unambiguous support towards one of these previously proposed contending hypotheses is congruent with the alternative scenario of an almost simultaneous diversification of the three species. Application of the phylogenetic species concept and the genetic species concept supports the recognition of three distinct taxonomic entities at the specific level. M. nelsoni inhabits the Sierra Madre Oriental (Hidalgo, Veracruz, and Puebla) including the Sierra Mazateca (Oaxaca); M. cryophilus is restricted to the Sierra de Juárez (Oaxaca); and M. thomasi occurs in portions of the Sierra Madre del Sur (Guerrero) and the Sierra Mixteca (Oaxaca). Our data show that M. thomasi is formed by two genetically distinct lineages that potentially may represent distinct Evolutionary Significant Units.     

A new species of Zelkova (Ulmaceae,Ulmoideae) with leaves and fruits from the Oligocene of South China and its biogeographical implications

A new fossil species of Zelkova is described from the Oligocene Ningming Formation of Guangxi, South China. Zelkova ningmingensis sp. nov. is characterized by leaves with craspedodromous venation pattern and drupaceous fruit type. The new species possesses elliptical to ovate leaves, bearing 7–12 pairs of secondary and simple toothed margin, as well as epidermal cells with straight or rounded anticlinal walls. The species is compared with extant and other fossil species hitherto reported of the genus. It is most similar to the living Zelkova schneideriana in the leaf gross morphology and epidermal characters, which may be suggested to be the ancestral type of Zelkova schneideriana. The discovery of Zelkova ningmingensis sp. nov. in Guangxi indicates that Zelkova has already existed in southern China as early as the Oligocene. In combination with Zelkova material from the Oligocene of Europe, it can be inferred that Eurasian Zelkova had begun to diversify by at least the Oligocene. Because China is the biodiversity centre of modern Zelkova, the fossil herein provides new insights into Zelkova biogeography.     

Lagomorphs from the Miocene of Sandelzhausen (southern Germany)

Three genera of lagomorphs, Prolagus, Lagopsis, and “Amphilagus,” were identified during a revision of the lagomorph material from Sandelzhausen (MN5, Early/Middle Miocene boundary, southern Germany). Evidence of two morphological and dimensional classes were observed at some tooth positions in Prolagus (some p3 show an unmistakable P. oeningensis morphology, others closely resemble P. crusafonti), but not at other tooth positions (e.g., M1–2). Insufficient data from Sandelzhausen precludes identification of two different species of Prolagus from this locality, and to define the characteristics of the possible P. crusafonti-like species. Thus, all Prolagus specimens have been classified as P. aff. oeningensis. The genus Lagopsis is represented by L. cf. penai, whose presence is compatible with a MN5 age. The relative abundance of Lagopsis to Prolagus may indicate relatively cool and wet palaeoclimatic conditions. The largest primitive lagomorph species from continental Europe is present at Sandelzhausen. Morphological and dimensional comparisons with other European primitive lagomorphs exclude any affinity with the genera Eurolagus and Titanomys and with the species included in “Amphilagus ulmensis”. Some common features with “Amphilagus antiquus” were observed, although they are not sufficient for the attribution to this taxon. Until there is a general revision of European primitive lagomorphs, the Sandelzhausen giant lagomorph is classified as “Amphilagus” sp. Its origins, whether from evolution within Europe or migration from Asia, remain unknown.      

A new dugong species (Sirenia,Dugongidae) from the Eocene of Catalonia (NE Iberian Peninsula)

A new stem dugongid species, Prototherium ausetanum sp. nov. (Dugongidae, Halitheriinae), is described based on a cranium from the middle Eocene of Mas Vilageliu (Tona, NE Iberian Peninsula). The new species displays a combination of features that enables its distinction from other halitheriines, including Prototherium veronense (type species of the genus), Prototherium? intermedium (which likely belongs to a different genus), and Prototherium? montserratense (here considered a nomen dubium), as well as Eotheroides spp. In overall morphology (e.g., dolichocephaly) the new species more closely resembles species previously included in Prototherium. However, a cladistic analysis based on craniodental features recovers the new species as the sister taxon of Eotheroides aegyptiacum (type species of this genus), further constituting a polytomy with P. veronense, Eotheroides lambondrano and the remaining Halitheriinae. Our analysis further indicates that P.? intermedium is more derived than other species of Prototherium, but it does not conclusively resolve the phylogenetic relationships between the included species of Prototherium and Eotheroides. A deeper taxonomic revision of these two genera would be required in order to better resolve the phylogeny of early dugongids.     

Two new species of Phyllonema (Rivulariaceae,Cyanobacteria) with an emendation of the genus

Two untapered, heterocytous species were observed and collected from the intertidal and supratidal zones of the Mexican coastline of the Pacific Ocean near Oaxaca and from the Gulf of Mexico. These populations were highly similar in morphology to the freshwater taxon Petalonema incrustans in the Scytonemataceae. However, 16S rRNA sequence data and phylogenetic analysis indicated that they were sister taxa to the epiphyllic, Brazilian species Phyllonema aveceniicola in the Rivulariaceae, described from culture material. While genetic identity between the two new species was high, they differed significantly in morphology, 16S rRNA gene sequence identity, and sequence and structure of the 16S–23S ITS region. Their morphology differed markedly from the generitype of the previously monotypic Phyllonema, which has tapered, heteropolar, single‐false branched trichomes with very thin or absent sheath. The two new species, Phyllonema ansata and Phyllonema tangolundensis, described from both culture and environmental material, have untapered, isopolar, geminately false branched trichomes with thick, lamellated sheaths, differences so significant that the species would not be placed in Phyllonema without molecular corroboration. The morphological differences are so significant that a formal emendation of the genus is required. These taxa provide a challenge to algal taxonomy because the morphological differences are such that one would logically conclude that they represent different genera, but the phylogenetic evidence for including them all in the same genus is conclusive. This conclusion is counter to the current trend in algal taxonomy in which taxa with minor morphological differences have been repeatedly placed in separate genera based primarily upon DNA sequence evidence.     

Is Cyclocardia (Conrad) a wastebasket taxon? Exploring the phylogeny of the most diverse genus of the Carditidae (Archiheterodonta,Bivalvia)

The carditid genus Cyclocardia is currently the most diverse genus of the family, including nearly 180 nominal species encompassing wide stratigraphical (Cretaceous–Recent) and geographical (Antarctica, South and North America, Europe, Africa, Alaska, Russia, Japan and New Zealand) ranges. Due to the lack of autapomorphies in the diagnosis of the genus and its large account of species, we re-evaluate the systematic and phylogenetic status of Cyclocardia. We applied three approaches: bibliographic revision, phylogenetic analysis and an exploration of morphological disparity. We used a shell–character matrix comprising 65 taxa (2 outgroups, 29 non-Cyclocardia carditids and 28 species of Cyclocardia) for phylogenetic and disparity analyses. Maximum Observable Rescaled Distances was used to construct a distance matrix to compare Cyclocardia species and other carditid groups. According to our results, Cyclocardia represents a non-monophyletic taxon and is thus a ‘wastebasket taxon’, chiefly because its diagnosis was based mainly on plesiomorphic characters. The European species C. kickxi and C. chameformis are placed within Scalaricardita, and the previously proposed genus Crassicardia is monophyletic (including C. crassidens, C. crebricostata, C. isaotakii and C. rjabininae). Three new genera are proposed for new groups identified by the phylogenetic analysis: South American Oesterheldia gen. nov. (including O. cannada and O. dalek), western North American Coanicardita gen. nov. (including C. ventricosa and C. occidentalis), and North Pacific Hippocampocardia gen. nov. (including H. barbarensis, H. hamiltonensis and H. yakatagensis). The newly defined monophyletic Cyclocardia is restricted to the Atlantic Ocean species C. borealis, C. novangliae and C. compressa.     

Stable climate corridors promote gene flow in the Cape sand snake species complex (Psammophiidae)

Climate shifts during the Quaternary Period have driven changes in regional range dynamics for many species, influencing population structure of species and in some cases promoting speciation. Within southern Africa, the psammophine snakes Psammophis trinasalis and P. namibensis were historically considered subspecies of P. leightoni but were elevated to species rank based on ecological differences. Preliminary phylogenetic analyses suggested intraspecific, not interspecific genetic variation between these taxa, but this finding was based on very limited data and could not be confirmed. To assess the level of genetic differentiation within the P. leightoni species complex, we explored the evolutionary history of these snakes by combining phylogenetic analyses, species distribution modelling and an examination of morphology. We generated a comprehensive, multi-gene phylogeny for Psammophis that included wider geographic sampling of the three species in the complex. Using this phylogeny, Bayesian and distance-based species delimitation analyses showed intraspecific, not interspecific divergences between taxa in the complex, suggesting that they collectively represent a single taxon. Furthermore, non-metric multidimensional scaling analysis of scalation characters showed no differences between the species. Moreover, palaeo-modelling at three time periods since the last interglacial period suggest that there have been varying levels of connectivity between these taxa, which has likely facilitated gene flow between them. Given the evidence, we propose that the P. leightoni complex represents a single species and therefore formally synonymise the three species.     

Cutting the Gordian Knot: Phylogenetic and ecological diversification of the Mesalina brevirostris species complex (Squamata,Lacertidae)

Mesalina are small lacertid lizards occurring in the Saharo‐Sindian deserts from North Africa to the east of the Iranian plateau. Earlier phylogenetic studies indicated that there are several species complexes within the genus and that thorough taxonomic revisions are needed. In this study, we aim at resolving the phylogeny and taxonomy of the M. brevirostris species complex distributed from the Middle East to the Arabian/Persian Gulf region and Pakistan. We sequenced three mitochondrial and three nuclear gene fragments, and in combination with species delimitation and species‐tree estimation, we infer a time‐calibrated phylogeny of the complex. The results of the genetic analyses support the presence of four clearly delimited species in the complex that diverged approximately between the middle Pliocene and the Pliocene/Pleistocene boundary. Species distribution models of the four species show that the areas of suitable habitat are geographically well delineated and nearly allopatric, and that most of the species have rather divergent environmental niches. Morphological characters also confirm the differences between the species, although sometimes minute. As a result of all these lines of evidence, we revise the taxonomy of the Mesalina brevirostris species complex. We designate a lectotype for Mesalina brevirostris Blanford, 1874; resurrect the available name Eremias bernoullii Schenkel, 1901 from the synonymy of M. brevirostris; elevate M. brevirostris microlepis (Angel, 1936) to species status; and describe Mesalina saudiarabica, a new species from Saudi Arabia.     

Morphofunctional analysis of Svobodaina species (Brachiopoda,Heterorthidae) from south‐western Europe

Geometric morphometric methods applied to the ventral muscle field outline of the various species of Svobodaina from south‐western Europe identify criteria for discriminating among this key cluster of brachiopod species. These data indicate the close relationship between the patterns of the ventral muscle field in the Svobodaina species and the environmental conditions where each inhabited; the fields are better developed in species related to high‐energy environments. For example, S. armoricana, with the smallest diductor scars, would have inhabited the quiet marine environments of the lower offshore or within protected lagoonal settings. S. feisti would have inhabited the upper offshore, a more energetic environment than S. armoricana. Finally, S. havliceki, with the largest diductor scars, would have thrived in the most energetic environments among all the south‐western European Svobodaina species, living just above the fair‐weather wave base in the lower shoreface. The palaeoecological results suggest a distribution of Svobodaina species during the Late Ordovician along an onshore–offshore transect across the shallow marine platforms of the Mediterranean margin of Gondwana. On the other hand, the occurrence in some localities of several species with overlapping ranges or within the same assemblage indicates that the biostratigraphical efficacy of the genus is restricted. Thus, the previously defined taxon‐range biozones characterized by Svobodaina species of the north Gondwanan margin are in need of reassessment. The morphology of Svobodaina may be a considerable aid to environmental analyses rather than to precise biostratigraphical correlations.     

A taxonomic revision of Encyclia (Orchidaceae: Laeliinae) in Costa Rica

A taxonomic revision of Encyclia (Orchidaceae) in Costa Rica is presented. The taxonomic history of the genus and its phylogenetic position are discussed. Characters of vegetative and floral morphology are described and their taxonomic significance is discussed. The genus is treated as comprising nine species in the country and a key to species is provided. Each taxon is described on the basis of Costa Rican material, illustrated in a composite plate, and its distribution within the country is assessed. Distribution maps for all the taxa are given. Overall distribution, derivation of name, synonymy, notes on species ecology and diagnostic features are presented for each taxon. The names Encyclia tonduziana and Epidendrum mooreanum are typified. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 395–448.     

The signal crayfish is not a single species: cryptic diversity and invasions in the Pacific Northwest range of Pacifastacus leniusculus

1. We used historical sources, morphology‐based taxonomy and mtDNA sequence data to address questions about the signal crayfish Pacifastacus leniusculus. These included evaluating unrecognised cryptic diversity and investigating the extent to which P. leniusculus may have been introduced within its presumed native range in the Pacific Northwest region of North America. Our study builds and expands on Pacific Northwest phylogeographic knowledge, particularly related to patterns of glacial refugia for freshwater species. 2. Extensive collections (824 specimens) from British Columbia (Canada), Idaho, Nevada, Oregon and Washington (United States) were used to characterise P. leniusculus at the mitochondrial 16S rRNA gene. Genetic groups within the species were elucidated by phylogenetics and amova ; evolutionary relationships within the most common and diverse group were investigated using a statistical parsimony haplotype network, a nested amova , and tests of isolation by distance. Morphological measurements were used to relate findings of molecular analyses to three historically recognised P. leniusculus subspecies and characterise cryptic diversity by morphology. 3. We found substantial cryptic diversity, with three groups highly distinct from P. leniusculus in discrete geographic regions: the Chehalis River glacial refugium, Central Oregon and the Okanagan Plateau. Disjunct distributions of P. leniusculus relative to these cryptic groups and known patterns of Pleistocene glaciation and landscape evolution cast doubt on whether P. leniusculus is native to some areas such as coastal drainages of northern Washington and southern British Columbia. Morphological traits previously used to characterise P. leniusculus subspecies still persist but may be incapable of distinguishing P. leniusculus from newly discovered cryptic groups. 4. Cryptic diversity found within P. leniusculus highlights the pressing need for a thorough investigation of the genus Pacifastacus using data based on more extensive gene and taxon sampling. It also warrants conservation attention, as introductions of P. leniusculus within the Pacific Northwest may carry risks of hybridisation and introgression for cryptic groups. Owing to high genetic diversity and limited dispersal capacity relative to more vagile organisms like freshwater fish, crayfish of the genus Pacifastacus offer powerful potential insights into the geological history and phylogeography of the Pacific Northwest region. Finally, by shedding light on the long‐neglected native range of P. leniusculus, our results should also better inform our understanding of potential source populations for, and the ecology of, this important invasive species in regions including Europe, Japan and elsewhere in North America.