Evolution and biogeography of primates: a new model based on molecular phylogenetics,vicariance and plate tectonics

Heads, M. Evolution and biogeography of primates: a new model based on molecular phylogenetics, vicariance and plate tectonics. —Zoologica Scripta, 39, 107–127. The ages of the oldest fossils suggest an origin for primates in the Paleocene (～56 Ma). Fossil‐calibrated molecular clock dates give Cretaceous dates (～80–116 Ma). Both these estimates are minimum dates although they are often ‘transmogrified’ and treated as maximum or absolute dates. Oldest fossils can underestimate ages by tens of millions of years and instead of calibrating the time‐course of evolution with a scanty fossil record, the geographical boundaries of the main molecular clades of primates are calibrated here with radiometrically dated tectonic events. This indicates that primates originated when a globally widespread ancestor (early Archonta) differentiated into a northern group (Plesiadapiformes, extinct), a southern group (Primates), and two south‐east Asian groups (Dermoptera and Scandentia). The division occurred with the breakup of Pangea in the Early Jurassic and the opening of the central Atlantic (～185 Ma). Within primates, the strepsirrhines and haplorhines diverged with volcanism and buckling on the Lebombo Monocline, a volcanic rifted margin in south‐east Africa (Early Jurassic, ～180 Ma). Within strepsirrhines, lorises and galagos (Africa and Asia) and lemurs (Madagascar) diverged with the formation of the Mozambique Channel (Middle Jurassic, ～160 Ma). Within haplorhines, Old World monkeys and New World monkeys diverged with the opening of the Atlantic (Early Cretaceous, ～130 Ma). The main aspects of primate distribution are interpreted as the result of plate tectonics, phylogeny and vicariance, with some subsequent range expansion leading to secondary overlap. Long‐distance, trans‐oceanic dispersal events are not necessary. The primate ancestral complex was already widespread globally when sea‐floor spreading, strike‐slip rifting and orogeny fractured and deformed distributions through the Jurassic and Cretaceous, leading to the origin of the modern clades. The model suggests that the topology of the phylogenetic tree reflects a sequence of differentiation in a widespread ancestor rather than a series of dispersal events.     

Plywood‐like shell microstructures in hyoliths from the middle Cambrian (Drumian) Gowers Formation,Georgina Basin,Australia

Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators.     

Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy

Popov, L.E., Bassett, M.G. & Holmer, L.E. 2012: Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy. Lethaia, Vol. 45, pp. 566–573. The early ontogeny of Palaeozoic Craniiformea (Brachiopoda) remains controversial, with conflicting reports of evidence indicating lecithotrophic versus planktotrophic larval stages. Further compelling evidence for lecithotrophy in Palaeozoic craniiforms is described here. Newly obtained, well‐preserved Silurian specimens of craniiforms, including Craniops (Craniopsida), and Lepidocrania? and Orthisocrania (Craniida) from Gotland and the St. Petersburg region, form the basis for this study. The new material demonstrates that the characters of shell structure and shell formation provide evidence of early differentiation of an adult dorsal mantle, and the presence of a distinctive primary layer with a characteristic lath‐like pattern indicates that these craniiforms underwent a lecithotrophic larval stage, more or less identical to that of living. □Novocrania. Brachiopoda, Craniiformea, ontogeny, phylogeny, Early Palaeozoic.     

Buoyancy of some Palaeozoic ammonoids and their hydrostatic properties based on empirical 3D‐models

The interpretation of the function of the ammonoid phragmocone as a buoyancy device is now widely accepted among ammonoid researchers. During the 20^th century, several theoretical models were proposed for the role of the chambered shell (phragmocone); accordingly, the phragmocone had hydrostatic properties, which enabled it to attain neutral buoyancy, presuming it was partially filled with gas. With new three‐dimensional reconstructions of ammonoid shells, we are now able to test these hypothetical models using empirical volume data of actual ammonoid shells. We investigated three Palaeozoic ammonoids (Devonian and Carboniferous), namely Fidelites clariondi, Diallagites lenticulifer and Goniatites multiliratus, to reconstruct their hydrostatic properties, their syn vivo shell orientation and their buoyancy. According to our models, measurements and calculations, these specimens had aperture orientations of 19°, 64° and 125° during their lives. Although none of our results coincide with the aperture orientation of the living Nautilus, they do verify the predictions for shell orientations based on published theoretical models. Our calculations also show that the shorter the body chamber, the poorer was the hydrodynamic stability of the animal. This finding corroborates the results of theoretical models from the 1990s. With these results, which are based on actual specimens, we favour the rejection of hypotheses suggesting a purely benthonic mode of life of ammonoids. Additionally, it is now possible to assess hydrodynamic properties of the shells through ontogeny and phylogeny, leading to insights to validate theoretical modes of life and habitat through the animal's life.     

Origin of Mediterranean insular endemics in the Boraginales: integrative evidence from molecular dating and ancestral area reconstruction

Aim The presence of numerous reliable fossils and the occurrence of many endemic island species make the Boraginales particularly suitable for integrative biogeographical studies. In this paper we aim to elucidate the time frame and events associated with the origin of selected borages endemic to the Mediterranean climate zone. More specifically, we describe and examine the alternative palaeo‐ and neoendemic hypotheses for their origin. Location Corsica and Sardinia (continental fragment islands) and the Canary Islands (an oceanic island archipelago). Methods Eighty‐nine accessions, representing 30 genera from five families ascribed to the Boraginales, were examined for six chloroplast DNA regions. We used an integrative approach including phylogenetic analyses (Mr Bayes ), Bayesian molecular dating (T3 package) with four fossil constraints on nodes, and biogeographical reconstructions (diva ) to elucidate the temporal and spatial origins of the Corso‐Sardinian and Canary Island endemics. Results Species of Echium endemic to the Canary Islands diverged from their continental sister clade during the Miocene (15.3 ± 5.4 Ma), probably after the rise of the oldest islands (c. 20 Ma). Corso‐Sardinian endemics of Borago diverged from their primarily North African sister clade during the late Miocene‐Pliocene (c. 6.9 ± 3.6 Ma), well after the initial fragmentation of the islands (c. 30 Ma). Similarly, Corso‐Sardinian endemics of Anchusa diverged from the South African Anchusa capensis during the Pliocene–Pleistocene (c. 2.7 ± 2.1 Ma). Main conclusions The present study reveals an Anatolian origin for Anchusa, Borago and Echium and underlines the importance of the Eastern Mediterranean region as a possible reservoir for plant evolution in the Mediterranean Basin. For Anchusa and Borago, the divergence from their respective sister clades on the two types of islands post‐dated the formation of the islands, thus supporting the neo‐endemic hypothesis, whereas the dating results for the origin of Echium endemics were less conclusive.     

Telemetered cephalopod energetics: swimming, soaring, and blimping

Cephalopods are uniquely suited to field energetic studies.Their hollow mantles that pump water for respiration and jettingalso can accommodate differential transducer-transmitters. Thesetransmitters indicate pressure-flow power output, which canbe calibrated against oxygen consumption by swim-tunnel respirometry.Radio-acoustic positioning telemetry (RAPT) records pressure-flowpower and animal movements with meter accuracy in nature. Despiteinherent inefficiencies, jetting is the primary mode of locomotionfor both primitive nautilus and powerful, migratory oceanicsquids. In between, large-finned squid and cuttlefish mix jettingwith fin undulation in complex gaits that increase locomotorefficiency. Our studies show that the complex nervous systemscephalopods evolved to control mixed gaits are also sensitiveto flow and density fields in nature and that they use theseto further reduce locomotion costs. Buoyed up by evacuated shells,nautilus and cuttlefish live in boundary layers and navigatecheaply through them like balloonists. Large-finned, negativelybuoyant squid soar like eagles in rising currents, but losecontrol in currents above one body length per second. Many muscularsquids have life histories linked to current systems. Neutrallybuoyant ammoniacal cephalopods in the mesopelagic are a limitingcase in need of study. The small density differential betweenseawater and isotonic ammonium chloride trebles their volume,making them blimp-like with very low power densities. Some specieslive entirely in this restricted habitat, but most become ammoniacallate in ontogeny, as they approach semelparous reproduction.Ammonium retained for buoyancy as carbon is terminally mobilizedfrom muscle protein for gametes and energy, compensates forlost muscle power.     

Latitudinal diversity gradients for brachiopod genera during late Palaeozoic time: links between climate, biogeography and evolutionary rates

Aim The latitudinal diversity gradient, in which taxonomic richness is greatest at low latitudes and declines towards the poles, is a pervasive feature of the biota through geological time. This study utilizes fossil data to examine how the latitudinal diversity gradient and associated spatial patterns covaried through the major climate shifts at the onset and end of the late Palaeozoic ice age. Location Data were acquired from fossil localities from around the world. Methods Latitudinal patterns of diversity, mean geographical range size and macroevolutionary rates were constructed from a literature‐derived data base of occurrences of fossil brachiopod genera in space and time. The literature search resulted in a total of 18,596 occurrences for 991 genera from 2320 localities. Results Climate changes associated with the onset of the late Palaeozoic ice age (c. 327 Ma) altered the biogeographical structure of the brachiopod fauna by the preferential elimination of narrowly distributed, largely tropical genera when glaciation began. Because the oceans were left populated primarily with widespread genera, the slope of the diversity gradient became gentle at this time, and the gradient of average latitudinal range size weakened. In addition, because narrowly distributed genera had intrinsically high rates of origination and extinction, the gradients of both of these macroevolutionary rates were also reduced. These patterns were reversed when the ice age climate abated in early Permian time (c. 290 Ma): narrowly distributed genera rediversified at low latitudes, restoring steep gradients of diversity, average latitudinal range size and macroevolutionary rates. Main conclusions During late Palaeozoic time, these latitudinal gradients for brachiopods may have been linked by the increased magnitude of seasonality during the late Palaeozoic ice age. Pronounced seasonality would have prevented the existence of genera with narrow latitudinal ranges. These results for the late Palaeozoic ice age suggest a climatic basis for the present‐day latitudinal diversity gradient.     

Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny

The fossil record suggests a rapid radiation of placental mammals following the Cretaceous-Paleogene (K-Pg) mass extinction 65 million years ago (Ma); nevertheless, molecular time estimates, while highly variable, are generally much older. Early molecular studies suffer from inadequate dating methods, reliance on the molecular clock, and simplistic and over-confident interpretations of the fossil record. More recent studies have used Bayesian dating methods that circumvent those issues, but the use of limited data has led to large estimation uncertainties, precluding a decisive conclusion on the timing of mammalian diversifications. Here we use a powerful Bayesian method to analyse 36 nuclear genomes and 274 mitochondrial genomes (20.6 million base pairs), combined with robust but flexible fossil calibrations. Our posterior time estimates suggest that marsupials diverged from eutherians 168-178 Ma, and crown Marsupialia diverged 64-84 Ma. Placentalia diverged 88-90 Ma, and present-day placental orders (except Primates and Xenarthra) originated in a ～20 Myr window (45-65 Ma) after the K-Pg extinction. Therefore we reject a pre K-Pg model of placental ordinal diversification. We suggest other infamous instances of mismatch between molecular and palaeontological divergence time estimates will be resolved with this same approach.     

Major issues in the origins of ray‐finned fish (Actinopterygii) biodiversity

Ray‐finned fishes (Actinopterygii) dominate modern aquatic ecosystems and are represented by over 32000 extant species. The vast majority of living actinopterygians are teleosts; their success is often attributed to a genome duplication event or morphological novelties. The remainder are ‘living fossils’ belonging to a few depauperate lineages with long‐retained ecomorphologies: Polypteriformes (bichirs), Holostei (bowfin and gar) and Chondrostei (paddlefish and sturgeon). Despite over a century of systematic work, the circumstances surrounding the origins of these clades, as well as their basic interrelationships and diagnoses, have been largely mired in uncertainty. Here, I review the systematics and characteristics of these major ray‐finned fish clades, and the early fossil record of Actinopterygii, in order to gauge the sources of doubt. Recent relaxed molecular clock studies have pushed the origins of actinopterygian crown clades to the mid‐late Palaeozoic [Silurian–Carboniferous; 420 to 298 million years ago (Ma)], despite a diagnostic body fossil record extending only to the later Mesozoic (251 to 66 Ma). This disjunct, recently termed the ‘Teleost Gap’ (although it affects all crown lineages), is based partly on calibrations from potential Palaeozoic stem‐taxa and thus has been attributed to poor fossil sampling. Actinopterygian fossils of appropriate ages are usually abundant and well preserved, yet long‐term neglect of this record in both taxonomic and systematic studies has exacerbated the gaps and obscured potential synapomorphies. At the moment, it is possible that later Palaeozoic‐age teleost, holostean, chondrostean and/or polypteriform crown taxa sit unrecognized in museum drawers. However, it is equally likely that the ‘Teleost Gap’ is an artifact of incorrect attributions to extant lineages, overwriting both a post‐Palaeozoic crown actinopterygian radiation and the ecomorphological diversity of stem‐taxa.     

Taxonomic review of the New World tamarins (Primates: Callitrichidae)

Twelve generic names have been ascribed to the New World tamarins but all are currently placed in just one: Saguinus Hoffmannsegg, 1807. Based on geographical distributions, morphology, and pelage patterns and coloration, they have been divided into six species groups: (1) nigricollis, (2) mystax, (3) midas, (4) inustus, (5) bicolor and (6) oedipus. Molecular phylogenetic studies have validated five of these groups; each are distinct clades. Saguinus inustus is embedded in the mystax group. Genetic studies show that tamarins are sister to all other callitrichids, diverging 15?13 Ma. The small‐bodied nigricollis group diverged from the remaining, larger tamarins 11?8 Ma, and the mystax group diverged 7?6 Ma; these radiations are older than those of the marmosets (Callithrix, Cebuella, Mico), which began to diversify 6?5 Ma. The oedipus group diverged from the midas and bicolor groups 5?4 Ma. We review recent taxonomic changes and summarize the history of the generic names. Taking into account the Late Miocene divergence time (11?8 Ma) between the large‐ and small‐bodied tamarin lineages, the small size of the nigricollis group species when compared with other tamarins, and the sympatry of the nigricollis group species with the larger mystax group species, we argue that the nigricollis group be recognized as a distinct genus: Leontocebus Wagner, 1839.     

The thermohaline expressway: the Southern Ocean as a centre of origin for deep‐sea octopuses

Understanding how environmental forcing has generated and maintained large‐scale patterns of biodiversity is a key goal of evolutionary research and critical to predicting the impacts of global climate change. We suggest that the initiation of the global thermohaline circulation provided a mechanism for the radiation of Southern Ocean fauna into the deep sea. We test this hypothesis using a relaxed phylogenetic approach to coestimate phylogeny and divergence times for a lineage of octopuses with Antarctic and deep‐sea representatives. We show that the deep‐sea lineage had their evolutionary origins in Antarctica, and estimate that this lineage diverged around 33 million years ago (Ma) and subsequently radiated at 15 Ma. Both of these dates are critical in development of the thermohaline circulation and we suggest that this has acted as an evolutionary driver enabling the Southern Ocean to become a centre of origin for deep‐sea fauna. This is the first unequivocal molecular evidence that deep‐sea fauna from other ocean basins originated from Southern Ocean taxa and this is the first evidence to be dated. © The Willi Hennig Society 2008.     

Beak from the body chamber of an Early Carboniferous shelled longiconic coleoid cephalopod from Arkansas,USA

Here we report the discovery of an Early Carboniferous (Late Visean) 3D cephalopod beak displaying significant similarity to the lower beak of Recent coleoids. It was uncovered in a fragmentarily preserved, longiconic shell from the Moorefield Formation in Arkansas, USA. This shell comprises a fractured 29‐mm‐long body chamber having a maximum diameter of ~14 mm and showing an indistinct pro‐ostracum‐like structure. The beak‐bearing shell could easily have been mistaken for a bactritid or orthocerid if it were not for a coleoid‐type, weakly mineralized, evidently organic‐rich shell wall which shows a lamello‐columnar ultrastructure of a bulk of shell wall thickness and plate ultrastructure of thin outer layer. The specimen is assigned to an as‐yet unnamed shelled coleoid of a so far unknown high‐level taxonomic group. A partially exposed, 4.0‐mm‐long portion of the beak is the lower beak in oblique view from its left side. It exhibits fractured anthracite‐like black, apparently originally chitin material, helmet‐like general shape, broad hood with narrow shallow median groove and small notch posteriorly, pronounced pointed, non‐biomineralized upside belt rostrum, high shoulder and about a 90–100 degrees jaw angle. A broad hood and massive rostrum emphasize its similarity to the lower mandible of Recent Vampyroteuthis and signify that its unique, among living coleoids, structure has been existed for at least since Late Visean time (~333 my).     

Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny

Popov, L.E., Bassett, M.G., Holmer, L.E., Skovsted, C.B. & Zuykov, M.A. 2010: Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny. Lethaia, Vol. 43, pp. 323–333. Well preserved specimens of the Early Palaeozoic craniiform brachiopods Orthisocrania and Craniops retain clear evidence of a lecithotrophic larval stage, indicating the loss of planktotrophy early in their phylogeny. The size of the earliest mineralized dorsal shell was <100 μm across, and the well preserved shell structure in these fossil craniiforms allows their earliest ontogeny to be compared directly with that of living Novocrania, in which the first mineralized dorsal shell (metamorphic shell) is secreted only after settlement of the lecithotrophic larvae. Immediately outside this earliest shell (early post‐metamorphic or brephic shell) and in the rest of the dorsal valve the primary layer in both fossil and living craniiforms has characteristic radially arranged laths, which are invariably lacking in the earliest dorsal shell. The ventral valve of the fossil specimens commonly preserves traces of an early attachment scar (cicatrix), which is equal in size to the dorsal metamorphic shell, and the brephic post‐metamorphic ventral valve also has a primary shell with radially arranged laths. However, a primary shell with radial laths is completely lacking in the ventral valve of living Novocrania, indicating that heterochrony may have been involved in the origin of the encrusting mode of life in living craniids; the entire ventral valve of Recent craniids (with the possible exception of Neoancistrocrania) may correspond to the earliest attachment scar of some fossil taxa such as Orthisocrania. It is also probable that the unique absence of an inner mantle lobe as well as the absence of lobate cells in Novocrania could be the result of heterochronic changes. The dorsal valve of both fossil and living craniiforms has a marked outer growth ring, around 500 μm across, marking the transition to the adult, and a significant change in regime of shell secretion. The earliest craniiform attachment is considered to be homologous to the unique attachment structures described recently in polytoechioids (e.g. Antigonambonites) and other members of the strophomenate clade. However, unlike the craniiforms, polytoechioids and strophomenates all have planktotrophic larvae, and planktotrophy is most probably a plesiomorphic character for all Brachiopoda. □Brachiopoda, Craniiformea, Early Palaeozoic, ontogeny, phylogeny.     

A NEW RECONSTRUCTION OF ONYCHOSELACHE TRAQUAIRI, COMMENTS ON EARLY CHONDRICHTHYAN PECTORAL GIRDLES AND HYBODONTIFORM PHYLOGENY

Abstract:  A new, third, specimen of Onychoselache traquairi from the Viséan (Holkerian) of Scotland allows a significant revision of the anatomy of this stem-group elasmobranch. This first report of material from the Mumbie Quarry exposure of the Glencartholm fish beds presents a new reconstruction of Onychoselache showing broad-based cephalic and nuchal spines, and exceptionally large pectoral fins. Details of the jaws, braincase and postcranial skeleton demonstrate that Onychoselache is a well-characterized member of the Hybodontiformes. Comparisons of the pectoral skeleton with other early chondrichthyan examples, including new material of Tristychius arcuatus and Plesioselachus macracanthus , highlight a range of early chondrichthyan conditions that are incorporated into a revised hybodontiform phylogeny. Close resemblance between Onychoselache and Mesozoic and late Palaeozoic hybodonts implies that these clades diverged within the Carboniferous and Permian. Major differences between Onychoselache and the coeval Tristychius (a modified reconstruction of which is included) indicate that the Neoselachii-Hybodontiformes split is probably Late Devonian, consistent with records of isolated teeth. The pectoral fins of Onychoselache , while unique among Palaeozoic forms, resemble those of Recent bamboo and epaulette sharks (Orectolobiformes). The functional corollary of this convergence is that Onychoselache represents an instance of a non-tetrapod early vertebrate with a near-walking gait.     

Microstructural disparity between basal micrabaciids and other Scleractinia: new evidence from Neogene Stephanophyllia

Recent molecular analyses based on mitochondrial and nuclear markers place the Micrabaciidae in the basal clade of scleractinian corals. The molecular distinctiveness of micrabaciids is supported by a set of unique morphological characters, among which the microstructure of thickening deposits is the most characteristic one. In all extant and well‐preserved Mesozoic micrabaciids (extinct Micrabacia, and still living Letepsammia, Rhombopsammia, Stephanophyllia, Leptopenus), thickening deposits consist of irregular meshwork of small chip‐like bundles of fibres. Here, we document Neogene (Miocene and Pliocene) forms identified as Stephanophyllia whose thickening deposits consist of long and thin parallel fibres that, instead of bundles (like in majority of Scleractinia), form layers of thatch‐like structures that thicken the septa. This microstructural pattern distinguishes Neogene Stephanophyllia from all examined so far micrabaciids and suggests that mechanisms of biologically controlled mineralization within this clade were more diverse. Nonetheless, the group as a whole is still clearly separated microstructurally from other scleractinians. Despite their basal position in scleractinian phylogeny, the fossil record of Micrabaciidae starts only in the Lower Cretaceous. No Palaeozoic, Triassic or Jurassic forms that could be considered ancestral to micrabaciids and would share some microstructural or morphological (e.g. septal insertion pattern) characters have yet been found. Possible explanations of such morphological disparity of micrabaciids from other scleractinians are either sudden emergence by skeletonization of long evolved, soft‐bodied group of basal hexacorallians or migration of their skeletonized, deep‐water ancestors to shallow‐waters.     

Electrophoretic identification of poritid species (Anthozoa: Scleractinia)

Electrophoretic surveys of 13 enzyme-coding loci distinguished unambiguously five morphologically defined species of Porites and two species of Goniopora. Each species was identifiable solely by unique, qualitative banding patterns at 1–6 loci. Genetic distances give preliminary estimates that these Porites species diverged from common ancestors 8–22 Ma during the Miocene, and that the two Goniopora species diverged about 3.5 Ma in the Pliocene, assuming Porites evolved from Goniopora 55 million years ago (Ma). Correspondence to: R. L. Garthwaite     

Phylogenetic characterization of three morphs of mussels (Bivalvia, Mytilidae) inhabiting isolated marine environments in Palau Islands

Marine lakes in the Palau Islands are known to harbor unique marine fauna that have remained isolated since the formation of the lakes after the Last Glacial Maximum. We analyzed mussels from marine lakes located on different islands and conducted morphological, phylogenetic and population genetic characterization to clarify their evolutionary history. The mussels were morphologically classified into three differentiated morphs: NS, ON, and MC. Their common characteristics were consistent with the Brachidontes-Hormomya complex of the Mytilidae family. Phylogenetic analysis based on the nuclear 18S ribosomal RNA gene supported the taxonomic position of the mussels among the Mytilidae. In the mitochondrial cytochrome c oxidase subunit I (COI) gene lineage, NS-and MC-morphs were highly diverged from each other; their estimated time of divergence dates back to the mid-Pleistocene. ON-morph was more closely related to MC-morph, although the shell morphologies of ON- and MC-morphs were easily distinguishable. Population genetic analysis revealed the coexistence of highly diverged haplotypes within a population of ON-morph, indicating introgression of mtDNA among the morphs. Our data suggest that morphological differentiation of marine lake mussels can occur in a relatively short period under different environmental conditions. Thus, the marine lakes provide a unique site for the study of diversification in mussels.     

The ancient Britons: groundwater fauna survived extreme climate change over tens of millions of years across NW Europe

Global climate changes during the Cenozoic (65.5–0 Ma) caused major biological range shifts and extinctions. In northern Europe, for example, a pattern of few endemics and the dominance of wide‐ranging species is thought to have been determined by the Pleistocene (2.59–0.01 Ma) glaciations. This study, in contrast, reveals an ancient subsurface fauna endemic to Britain and Ireland. Using a Bayesian phylogenetic approach, we found that two species of stygobitic invertebrates (genus Niphargus) have not only survived the entire Pleistocene in refugia but have persisted for at least 19.5 million years. Other Niphargus species form distinct cryptic taxa that diverged from their nearest continental relative between 5.6 and 1.0 Ma. The study also reveals an unusual biogeographical pattern in the Niphargus genus. It originated in north‐west Europe approximately 87 Ma and underwent a gradual range expansion. Phylogenetic diversity and species age are highest in north‐west Europe, suggesting resilience to extreme climate change and strongly contrasting the patterns seen in surface fauna. However, species diversity is highest in south‐east Europe, indicating that once the genus spread to these areas (approximately 25 Ma), geomorphological and climatic conditions enabled much higher diversification. Our study highlights that groundwater ecosystems provide an important contribution to biodiversity and offers insight into the interactions between biological and climatic processes.     

Torsion in Patella caerulea (Mollusca, Patellogastropoda): ontogenetic process, timing, and mechanisms

Abstract. Torsion is a process in gastropod ontogenesis where the visceral body portion rotates 180° relative to the head/foot region. We investigated this process in the limpet Patella caerulea by using light microscopy of living larvae, as well as scanning electron microscopy (SEM) of larvae fixed during the torsion process. The completion of the 180° twist takes considerably less time in larvae of Patella caerulea than previously described for other basal gastropod species. At a rearing temperature of 20–22°C, individuals complete ontogenetic torsion in ?2 h. Furthermore, the whole process is monophasic, i.e., carried out at a constant speed, without any evidence of distinct ‘fast” or ‘slow” phases. Both larval shell muscles—the main and the accessory larval retractor—are already fully contractile before the onset of torsion. During the torsion process both retractors perform cramp‐like contractions at ～30 s intervals, which are followed by hydraulic movements of the foot. However, retraction into the embryonic shell occurs only after torsion is completed. The formation of the larval operculum is entirely in‐dependent from ontogenetic torsion and starts before the onset of rotation, as does the mineralization of the embryonic shell. The reported variability regarding the timing (mono‐ versus biphasic; duration) of torsion in basal gastropod species precludes any attempt to interpret these data phylogenetically. The present findings indicate that the torsion process in Patella caerulea, and probably generally in basal gastropods, is primarily caused by contraction of the larval shell muscles in combination with hydraulic activities. In contrast, the adult shell musculature, which is independently formed after torsion is completed, does not contribute to ontogenetic torsion in any way. Thus, fossil data relying on muscle scars of adult shell muscles alone appear inappropriate to prove torted or untorted conditions in early Paleozoic univalved molluses. Therefore, we argue that paleontological studies dealing with gastropod phylogeny require data other than those based on fossilized attachment sites of adult shell muscles.     

Divergence time estimates for the early history of animal phyla and the origin of plants, animals and fungi

In the past, molecular clocks have been used to estimate divergence times among animal phyla, but those time estimates have varied widely (1200-670 million years ago, Ma). In order to obtain time estimates that are more robust, we have analysed a larger number of genes for divergences among three well-represented animal phyla, and among plants, animals and fungi. The time estimate for the chordate-arthropod divergence, using 50 genes, is 993 +/- 46 Ma. Nematodes were found to have diverged from the lineage leading to arthropods and chordates at 1177 +/- 79 Ma. Phylogenetic analyses also show that a basal position of nematodes has strong support (p > 99%) and is not the result of rate biases. The three-way split (relationships unresolved) of plants, animals and fungi was estimated at 1576 +/- 88 Ma. By inference, the basal animal phyla (Porifera, Cnidaria, Ctenophora) diverged between about 1200-1500 Ma. This suggests that at least six animal phyla originated deep in the Precambrian, more than 400 million years earlier than their first appearance in the fossil record.