A Paedomorphic parasite associated with a neotenic amphibian host: phylogenetic evidence suggests a revised systematic position for Sphyranuridae within anuran and turtle Polystomatoineans

The phylogenetic relationships of the families Polystomatidae and Sphyranuridae (subclass Polystomatoinea) within tetrapod monogenean parasites were investigated using partial 18S rDNA sequences. About 600 nucleotides of 11 species were sequenced, including 7 species of the most common subfamilies of Polystomatidae found in anurans and turtles, 1 species of the family Sphyranuridae parasitizing exclusively urodelans, and 3 species of the subclass Oligonchoinea infesting teleostean fishes. The phylogenetic analyses were performed using three reconstruction methods: neighbor-joining, maximum-parsimony, and maximum-likelihood. Polystomatoineans but not polystomatids were shown to be monophyletic. Within the polystomatoineans there are two clades: one includes the amphibian monogeneans (anuran polystomatids and urodelan sphyranurids) and the other includes the turtle polystomatids. Polystomatoineans may have coevolved with amphibian hosts, and an ancestral "polystome" dispersed at least 200 million years ago, either from the basal stem of lissamphibians or from an anuran ancestral stock, to freshwater turtles. Furthermore, the urodelan genus Sphyranura, initially assigned to the family Sphyranuridae on the basis of morphological and ontogenetic evidence, is clearly nested within polystomatids, suggesting that its systematic status must be revised. This supports recent findings which argue that species of the family Sphyranuridae may be paedomorphic parasites exclusively infesting neotenic mudpuppies.     

Phylogeny and a revised classification of the Monogenoidea Bychowsky, 1937 (Platyhelminthes)

A hypothesis (CI=57.3%) on the evolutionary relationships of families comprising the class Monogenoidea is proposed based on 141 character states in 47 homologous series and employing phylogenetic systematics. Based on the analysis, three subclasses, the Polyonchoinea, Polystomatoinea and Oligonchoinea, are recognised. The analysis supports independent origins of the Montchadskyellidae within the Polyonchoinea and of the Neodactylodiscidae and Amphibdellatidae within the order Dactylogyridea (Polyonchoinea); the suborder Montchadskyellinea is raised to ordinal status and new suborders Neodactylodiscinea and Amphibdellatinea are proposed to reflect these origins. The Gyrodactylidea (Polyonchoinea) is supported by three synapomorphies and comprises the Gyrodactylidae, Anoplodiscidae, Tetraonchoididae and Bothitrematidae. The analysis supports recognition of the Polystomatoinea comprising Polystomatidae and Sphyranuridae. Evolutionary relationships within the Oligonchoinea indicate independent origins of three ordinal taxa, the Chimaericolidea (monotypic), Diclybothriidea (including Diclybothriidae and Hexabothriidae) and Mazocraeidea (with five suborders). The suborder Mazocraeinea comprises the Plectanocotylidae, Mazocraeidae and Mazoplectidae, and is characterised by two synapomorphies. The suborder Gastrocotylinea, characterised by presence of accessory sclerites in the haptoral sucker, is divided into two infraorders, the monotypic Anthocotylina infraorder novum and Gastrocotylina. Two superfamilies of the Gastrocotylina are recognised, the Protomicrocotyloidea and Gastrocotyloidea; the Pseudodiclidophoridae is considered incertae sedis within the Gastrocotylina. The suborder Discocotylinea comprises the Discocotylidae, Octomacridae and Diplozoidae and is supported by four synapomorphies. The monotypic Hexostomatinea suborder novum is proposed to reflect an independent origin of the Hexostomatidae within the Mazocraeidea. The terminal suborder Microcotylinea comprises four superfamilies, the Microcotyloidea, Allopyragraphoroidea, Diclidophoroidea and Pyragraphoroidea. The analysis supports incorporation of the Pterinotrematidae in the Pyragraphoroidea and rejection of the monotypic order Pterinotrematidea. The following taxa are also rejected for reasons of paraphyly and/or polyphyly: Articulonchoinea, Bothriocotylea, Eucotylea, Monoaxonematidea, Tetraonchidea, Gotocotyloidea, Anchorophoridae and Macrovalvitrematidae. The Sundanonchidae, Iagotrematidae and Microbothriidae were not included in the analysis because of lack of pertinent information regarding character states.     

Suspension and optical properties of the crystalline lens in the eyes of basal vertebrates

We have investigated the apparatus suspending the crystalline lens in the eyes of basal vertebrates. Data are presented for Holocephali (Chondrichthyes) and the actinopterygians Polypteriformes, Polyodontidae (Acipenseriformes), Lepisosteiformes, Amiiformes, and one teleost species, the banded archerfish (Toxotes jaculatrix). We also studied the optical properties of the lens in Polypteriformes, Lepisosteiformes, and the archerfish. Together with previously published results, our findings show that there are three basic types of lens suspension in vertebrates. These are i) a rotationally symmetric suspension (Petromyzontida, lampreys; Ceratodontiformes, lungfishes; Tetrapoda), ii) a suspension with a dorso‐ventral axis of symmetry and a ventral papilla (all Chondrichthyes and Acipenseriformes), and iii) an asymmetric suspension with a ventral muscle and a varying number of ligaments (all Actinopterygii except for Acipenseriformes). Large eyes with presumably high spatial resolution have evolved in all groups. Multifocal lenses creating well‐focused color images are also present in all groups studied. Stable and exact positioning of the lens, in many cases in combination with accommodative changes in lens position or shape, is achieved by all three types of lens suspension. It is somewhat surprising that lens suspensions are strikingly similar in Chondrichthyes and Acipenseriformes (Actinopterygii), while the suspension apparatus in Polypteriformes, usually being regarded as an actinopterygian group more basal than Acipenseriformes, are considerably more teleostean‐like. This study completes a series of investigations on lens suspensions in nontetrapod vertebrates, covering all major groups except for the rare and highly derived coelacanths. J. Morphol. 275:613–622, 2014. © 2013 Wiley Periodicals, Inc.     

Sturgeons,sharks, and rays have multifocal crystalline lenses and similar lens suspension apparatuses

Crystalline lenses with multiple focal lengths in monochromatic light (multifocal lenses) are present in many vertebrate groups. These lenses compensate for chromatic aberration and create well‐focused color images. Stabilization of the lens within the eye and the ability to adjust focus are further requirements for vision in high detail. We investigated the occurrence of multifocal lenses by photorefractometry and lens suspension structures by light and electron microscopy in sturgeons (Acipenseriformes, Chondrostei) as well as sharks and rays (Elasmobranchii, Chondrichthyes). Multifocal lenses were found in two more major vertebrate groups, the Chondrostei represented by Acipenseriformes and Chondrichthyes represented by Elasmobranchii. The lens suspension structures of sturgeons, sharks, and rays are more complex than described previously. The lens is suspended by many delicate suspensory fibers in association with a ventral papilla in all groups studied. The arrangements of the suspensory fibers are most similar between sturgeons and sharks. In rays, the lens is suspended by a smaller ventral papilla and the suspensory fibers are arranged more concentrically to the lens. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Phylogenetics of the Monogenea--evidence from a medley of molecules

Nuclear ribosomal DNA sequences of Monogenea from both complete small and partial large (D1-D2) subunits were determined and added to previously published sequences in order to best estimate the molecular phylogeny of the group. A total of 35 ssrDNA, 100 D1 lsrDNA and 51 D2 lsrDNA monogenean sequences were used, representing a total of 27 families. From these sequences different data sets were assembled and analysed to make the best use of all available molecular phylogenetic information from the taxa. Maximum parsimony and minimum evolution trees for each data partition were rooted against published sequences from the Cestoda, forcing the Monogenea to appear monophyletic. There was broad agreement between tree topologies estimated by both methods and between genes. Well-supported nodes were restricted to deeply diverging major groupings and more derived taxa with the lsrDNA data but were at most nodes with ssrDNA. The Polyonchoinea showed the greatest resolution with a general pattern of ((Monocotylidae(Capsalidae(Udonellidae+Gyrodactylidea)))((Anoplodiscidae+Sundanonchidae)(Pseudomurraytrematidae+Dactylogyridae))). The Heteronchoinea readily split into the Polystomatoinea+Oligonchoinea, and Chimaericolidae and Hexabothriidae were successively the most basal of oligonchoinean taxa. Relationships within the Mazocraeidea, comprising 27 families of which 15 were sampled here, were largely unresolved and appear to reflect a rapid radiation of this group that is reflected in very short internal branches for ssrDNA and D1 lsrDNA, and highly divergent D2 lsrDNA. A reduced morphological matrix, employing only those families represented by molecules, contrasted sharply with respect to polyonchoinean interrelationships. Deep branches of the Heteronchoinea were similar for both classes of data but also showed that the interrelationships of the mazocraeidean families are labile and susceptible to sampling.     

A view of early vertebrate evolution inferred from the phylogeny of polystome parasites (Monogenea: Polystomatidae)

The Polystomatidae is the only family within the Monogenea to parasitize sarcopterygians such as the Australian lungfish Neoceratodus poisteri and freshwater tetrapods (lissamphibians and chelonians). We present a phylogeny based on partial 18S rDNA sequences of 26 species of Polystomatidae and three taxon from the infrasubclass Oligonchoinea (= Polyopisthocotylea) obtained from the gills of teleost fishes. The basal position of the polystome from lungfish within the Polystomatidae suggests that the family arose during the evolutionary transition between actinopterygians and sarcopterygians, ca. 425 million years (Myr) ago. The monophyly of the polystomatid lineages from chelonian and lissamphibian hosts, in addition to estimates of the divergence times, indicate that polystomatids from turtles radiated ca. 191 Myr ago, following a switch from an aquatic amniote presumed to be extinct to turtles, which diversified in the Upper Triassic. Within polystomatids from lissamphibians, we observe a polytomy of four lineages, namely caudatan, neobatrachian, pelobatid and pipid polystomatid lineages, which occurred ca. 246 Myr ago according to molecular divergence-time estimates. This suggests that the first polystomatids of amphibians originated during the evolution and diversification of lissamphibian orders and suborders ca. 250 Myr ago. Finally, we report a vicariance event between two major groups of neobatrachian polystomes, which is probably linked to the separation of South America from Africa ca. 100 Myr ago.     

On the ontogeny of the haptor and the evolution of the Monogenea

Data on the ontogeny of the posterior haptor of monogeneans were obtained from more than 150 publications and summarised. These data were plotted into diagrams showing evolutionary capacity levels based on the theory of a progressive evolution of marginal hooks, anchors and other attachment components of the posterior haptor in the Monogenea (Malmberg, 1986). 5 + 5 unhinged marginal hooks are assumed to be the most primitive monogenean haptoral condition. Thus the diagrams were founded on a 5 + 5 unhinged marginal hook evolutionary capacity level, and the evolutionary capacity levels of anchors and other haptoral attachement components were arranged according to haptoral ontogenetical sequences. In the final plotting diagram data on hosts, type of spermatozoa, oncomiracidial ciliation, sensilla pattern and protonephridial systems were also included. In this way a number of correlations were revealed. Thus, for example, the number of 5 + 5 marginal hooks correlates with the most primitive monogenean type of spermatozoon and with few sensillae, many ciliated cells and a simple protonephridial system in the oncomiracidium. On the basis of the reviewed data it is concluded that the ancient monogeneans with 5 + 5 unhinged marginal hooks were divided into two main lines, one retaining unhinged marginal hooks and the other evolving hinged marginal hooks. Both main lines have recent representatives at different marginal hook evolutionary capacity levels, i.e. monogeneans retaining a haptor with only marginal hooks. For the main line with hinged marginal hooks the name Articulon-choinea n. subclass is proposed. Members with 8 + 8 hinged marginal hooks only are here called Proanchorea n. superord. Monogeneans with unhinged marginal hooks only are here called Ananchorea n. superord. and three new families are erected for its recent members: Anonchohapteridae n. fam., Acolpentronidae n. fam. and Anacanthoridae n. fam. (with 7 + 7, 8 + 8 and 9 + 9 unhinged marginal hooks, respectively). Except for the families of Articulonchoinea (e.g. Acanthocotylidae, Gyrodactylidae, Tetraonchoididae) Bychowsky's (1957) division of the Monogenea into the Oligonchoinea and Polyonchoinea fits the proposed scheme, i.e. monogeneans with unhinged marginal hooks form one old group, the Oligonchoinea, which have 5 + 5 unhinged marginal hooks, and the other group form the Polyonchoinea, which (with the exception of the Hexabothriidae) has a greater number (7 + 7, 8 + 8 or 9 + 9) of unhinged marginal hooks. It is proposed that both these names, Oligonchoinea (sensu mihi) and Polyonchoinea (sensu mihi), will be retained on one side and Articulonchoinea placed on the other side, which reflects the early monogenean evolution. Except for the members of Ananchorea [Polyonchoinea], all members of the Oligonchoinea and Polyonchoinea have anchors, which imply that they are further evolved, i.e. have passed the 5 + 5 marginal hook evolutionary capacity level (Malmberg, 1986). There are two main types of anchors in the Monogenea: haptoral anchors, with anlages appearing in the haptor, and peduncular anchors, with anlages in the peduncle. There are two types of haptoral anchors: peripheral haptoral anchors, ontogenetically the oldest, and central haptoral anchors. Peduncular anchors, in turn, are ontogenetically younger than peripheral haptoral anchors. There may be two pairs of peduncular anchors: medial peduncular anchors, ontogentically the oldest, and lateral peduncular anchors. Only peduncular (not haptoral) anchors have anchor bars. Monogeneans with haptoral anchors are here called Mediohaptanchorea n. superord. and Laterohaptanchorea n. superord. or haptanchoreans. All oligonchoineans and the oldest polyonchoineans are haptanchoreans. Certain members of Calceostomatidae [Polyonchoinea] are the only monogeneans with both (peripheral) haptoral and peduncular anchors (one pair). These monogeneans are here called Mixanchorea n. superord. Polyonchoineans with peduncular anchors and unhinged marginal hooks are here called the Pedunculanchorea n. superord. The most primitive pedunculanchoreans have only one pair of peduncular anchors with an anchor bar, while the most advanced have both medial and lateral peduncular anchors; each pair having an anchor bar. Certain families of the Articulonchoinea, the Anchorea n. superord., also have peduncular anchors (parallel evolution): only one family, the Sundanonchidae n. fam., has both medial and lateral peduncular anchors, each anchor pair with an anchor bar. Evolutionary lines from different monogenean evolutionary capacity levels are discussed and a new system of classification for the Monogenea is proposed.In agreeing to publish this article, I recognise that its contents are controversial and contrary to generally accepted views on monogenean systematics and evolution. I have anticipated a reaction to the article by inviting senior workers in the field to comment upon it: their views will be reported in a future issue of this journal. EditorIn agreeing to publish this article, I recognise that its contents are controversial and contrary to generally accepted views on monogenean systematics and evolution. I have anticipated a reaction to the article by inviting senior workers in the field to comment upon it: their views will be reported in a future issue of this journal. Editor     

Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

Molecular phylogeny of early vertebrates: monophyly of the agnathans as revealed by sequences of 35 genes

Extant vertebrates are divided into three major groups: hagfishes (Hyperotreti, myxinoids), lampreys (Hyperoartia, petromyzontids), and jawed vertebrates (Gnathostomata). The phylogenetic relationships among the groups and within the jawed vertebrates are controversial, for both morphological and molecular studies have rendered themselves to conflicting interpretations. Here, we use the sequences of 35 nuclear protein-encoding genes to provide definitive evidence for the monophyly of the Agnatha (jawless vertebrates, a group encompassing the hagfishes and lampreys). Our analyses also give a strong support for the separation of Chondrichthyes (cartilaginous fishes) before the divergence of Osteichthyes (bony fishes) from the other gnathostomes.     

Evolution of the facial musculature in basal ray-finned fishes

Background

The facial musculature is a remarkable anatomical complex involved in vital activities of fishes, such as food capture and gill ventilation. The evolution of the facial muscles is largely unknown in most major fish lineages, such as the Actinopterygii. This megadiverse group includes all ray-finned fishes and comprises approximately half of the living vertebrate species. The Polypteriformes, Acipenseriformes, Lepisosteiformes, Amiiformes, Elopiformes, and Hiodontiformes occupy basal positions in the actinopterygian phylogeny and a comparative study of their facial musculature is crucial for understanding the cranial evolution of bony fishes (Osteichthyes) as a whole.

Results

The facial musculature of basal actinopterygians is revised, redescribed, and analyzed under an evolutionary perspective. We identified twenty main muscle components ontogenetically and evolutionarily derived from three primordial muscles. Homologies of these components are clarified and serve as basis for the proposition of a standardized and unifying myological terminology for all ray-finned fishes. The evolutionary changes in the facial musculature are optimized on the osteichthyan tree and several new synapomorphies are identified for its largest clades, including the Actinopterygii, Neopterygii, and Teleostei. Myological data alone ambiguously support the monophyly of the Holostei. A newly identified specialization constitutes the first unequivocal morphological synapomorphy for the Elopiformes. The myological survey additionally allowed a reinterpretation of the homologies of ossifications in the upper jaw of acipenseriforms.

Conclusions

The facial musculature proved to be extremely informative for the higher-level phylogeny of bony fishes. These muscles have undergone remarkable changes during the early radiation of ray-finned fishes, with significant implications for the knowledge of the musculoskeletal evolution of both derived actinopterygians and lobe-finned fishes (Sarcopterygii).

     

Evolution of jaw depression mechanics in aquatic vertebrates: insights from Ghondrichthyes

The widely accepted phylogenctic position of Chondrichthyes as the sister group to all other living gnathostomes makes biomechanical analyses of this group of special significance for estimates of skull function in early jawed vertebrates. We review key findings of recent experimental research on the feeding mechanisms of living elasmobranchs with respect to our understanding of jaw depression mechanisms in gnathostome vertebrates. We introduce the possibility that the ancestral jaw depression mechanism in gnathostomes was mediated by the coracomandibularis muscle and that for hyoid depression by the coracohyoideus muscle, as in modern Chondrichthyes and possibly placoderms. This mechanism of jaw depression appears to have been replaced by the sternohyoideus (homologous to the coracohyoideus) coupling in Osteichthycs following the split of this lineage from Chondrichthyes. Concurrent with the replacement of the branchiomandibularis (homologous to the coracomandibularis) coupling by the sternohyoideus coupling as the dominant mechanism of jaw depression in Osteichthyes was the fusion and shift in attachment of the intcrhyoideus and intermandibularis muscles (producing the protractor hyoideus muscle, mistakenly refereed to as the geniohyoideus), which resulted in a more diversified role of the sternohyoideus coupling in Osteichthyes. The coracohyoideus coupling appears to have been already present in vertebrates where it functioned in hyoid depression, as in modern Chondrichthyes, before it acquired the additional role of jaw depression in Osteichthyes.     

Evolutionary origin of the Otx2 enhancer for its expression in visceral endoderm

In the mouse, the Otx2 gene has been shown to play essential roles in the visceral endoderm during anterior-posterior axis formation and head induction. While these are primary processes in vertebrate embryogenesis, the visceral endoderm is a tissue unique to mammals. Two enhancers (VE and CM) have been previously found to direct Otx2 expression during early embryogenesis. This study demonstrates that in anterior visceral endoderm the CM enhancer does not have an activity by itself, but enhances the activity of the VE enhancer. These two enhancers also cooperate for the activities in anterior mesendoderm and cephalic mesenchyme. Comparative studies suggest that VE enhancer function was most likely established before the divergence of sarcopterygians into Actinistia, Dipnoi and tetrapods, while the nucleotide sequence corresponding to the VE enhancer was already present in the last common ancestor of bony fishes. The CM enhancer sequence and function would have been also established in ancestral sarcopterygians. The VE/CM enhancers and their gene cascades in the ancestral sarcopterygian head organizer would then have been co-opted by amphibian deep endoderm cells and mammalian visceral endoderm cells for the head development.     

The American Paddlefish Genome Provides Novel Insights into Chromosomal Evolution and Bone Mineralization in Early Vertebrates

Sturgeons and paddlefishes (Acipenseriformes) occupy the basal position of ray-finned fishes, although they have cartilaginous skeletons as in Chondrichthyes. This evolutionary status and their morphological specializations make them a research focus, but their complex genomes (polyploidy and the presence of microchromosomes) bring obstacles and challenges to molecular studies. Here, we generated the first high-quality genome assembly of the American paddlefish (Polyodon spathula) at a chromosome level. Comparative genomic analyses revealed a recent species-specific whole-genome duplication event, and extensive chromosomal changes, including head-to-head fusions of pairs of intact, large ancestral chromosomes within the paddlefish. We also provide an overview of the paddlefish SCPP (secretory calcium-binding phosphoprotein) repertoire that is responsible for tissue mineralization, demonstrating that the earliest flourishing of SCPP members occurred at least before the split between Acipenseriformes and teleosts. In summary, this genome assembly provides a genetic resource for understanding chromosomal evolution in polyploid nonteleost fishes and bone mineralization in early vertebrates.     

Ventral gill arch muscles and the interrelationships of gnathostomes, with a new classification of the Vertebrata

The ventral gill arch muscles of chondrichthyans, Latimeria , dipnoans, larval amphibians and actinopterygians are described and compared. Muscle patterns are characterized as primitive or derived and the derived patterns are used to test various hypotheses of the interrelationships of these griathostome groups. Gnathostomes differ from agnathans in having branchial muscles associated with the ventral gill arches. The monophyly of chondrichthyans is corroborated by the presence of coracobranchiales of hypobranchial origin. The monophyly of Recent teleostomes (Osteichthyes) is indicated by the presence of discrete transversi ventrales and interarcuales ventrales, and by the loss of the fifth superficial constrictor. A monophyletic Sarcopterygii which includes Latimeria is refuted by three paired branchial muscles found in dipnoans, Recent choanates and actinopterygians, but missing in Latimeria: pharyngoclaviculares, obliqui ventrales 1 and transversi ventrales 4. A new name, Euosteichthyes, is proposed for the group including dipnoans, choanates and actinopterygians. Sarcopterygii is restricted to include only dipnoans and choanates (among Recent organisms). Actinistia, including Latimeria , is proposed as the sister-group of all other Recent osteictithyans. Brachiopterygians (polypterids) are placed within Actinopterygii. A phylogenetic hypothesis supporting this classification is presented.     

Evolution of the Vertebrate Resistin Gene Family

Resistin (encoded by Retn) was previously identified in rodents as a hormone associated with diabetes; however human resistin is instead linked to inflammation. Resistin is a member of a small gene family that includes the resistin-like peptides (encoded by Retnl genes) in mammals. Genomic searches of available genome sequences of diverse vertebrates and phylogenetic analyses were conducted to determine the size and origin of the resistin-like gene family. Genes encoding peptides similar to resistin were found in Mammalia, Sauria, Amphibia, and Actinistia (coelacanth, a lobe-finned fish), but not in Aves or fish from Actinopterygii, Chondrichthyes, or Agnatha. Retnl originated by duplication and transposition from Retn on the early mammalian lineage after divergence of the platypus, but before the placental and marsupial mammal divergence. The resistin-like gene family illustrates an instance where the locus of origin of duplicated genes can be identified, with Retn continuing to reside at this location. Mammalian species typically have a single copy Retn gene, but are much more variable in their numbers of Retnl genes, ranging from 0 to 9. Since Retn is located at the locus of origin, thus likely retained the ancestral expression pattern, largely maintained its copy number, and did not display accelerated evolution, we suggest that it is more likely to have maintained an ancestral function, while Retnl, which transposed to a new location, displays accelerated evolution, and shows greater variability in gene number, including gene loss, likely evolved new, but potentially lineage-specific, functions.     

Cephalic muscles of Cyclostomes (hagfishes and lampreys) and Chondrichthyes (sharks,rays and holocephalans): comparative anatomy and early evolution of the vertebrate head muscles

Living vertebrate diversity comprises hagfishes and lampreys (Cyclostomata), elasmobranchs and holocephalans (Chondrichthyes), and bony fish which include tetrapods (Osteichthyes). Based on dissections and an extensive comparative analysis, we provide an updated overview of the anatomy, homologies and evolution of cyclostome and chondrichthyan cephalic muscles, with osteichthyans as primary comparative taxa. The analysis also infers plesiomorphic conditions for vertebrates and gnathostomes. We follow a uniform myological terminology for the Gnathostomata to demonstrate that the last common ancestor of extant vertebrates probably had a single intermandibularis and other mandibular muscles (labial muscles), some constrictores hyoidei and branchiales, and epibranchial and hypobranchial muscle sheets. The division of the cucullaris into levatores arcuum branchialium and protractor pectoralis is an osteichthyan synapomorphy and reflects an evolutionary trend towards a greater separation between the head and pectoral girdle that culminated in the formation of the tetrapod neck. Hence, this paper addresses a long‐standing, central issue regarding vertebrate comparative anatomy. It thus provides a valuable basis for future evolutionary, developmental and functional studies of vertebrates and/or of specific vertebrate subgroups/model organisms. © 2014 The Linnean Society of London     

Evolutionary divergence in freshwater insects with contrasting dispersal capacity across a sea of desert

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Age at maturity of some fish species distributed in Turkish marine waters (Actinopterygii and Elasmobranchii)

We collected data on the age at maturity (t_m) and maximum reported age (t_max) for 153 stocks of marine fishes in Turkey, belonging to 59 species, 24 families and 2 classes (Actinopterygii and Elasmobranchii). Among Actinopterygii t_m had an average of 1.8 years (1 to 4 years) while among Elasmobranchii it had an average of 11.9 years (2 to 11.9 years). Overall, t_max ranged between two years (for Sarda sarda) and 34 years (for Squalus acanthias). Mean t_max was found to be 6.24 years for Actinopterygii and 10.11 years for Elasmobranchii. t_m showed a positive linear correlation with t_max for both Actinopterygii and Elasmobranchii. Mean t_m?t_max did not differ significantly with sex within the Actinopterygii and Elasmobranchii. The ratio t_m?t_max was found to be significantly lower for Actinopterygii than for Elasmobranchii.     

The relative influence of natural selection and geography on gene flow in guppies

Two general processes may influence gene flow among populations. One involves divergent selection, wherein the maladaptation of immigrants and hybrids impedes gene flow between ecological environments (i.e. ecological speciation). The other involves geographic features that limit dispersal. We determined the relative influence of these two processes in natural populations of Trinidadian guppies (Poecilia reticulata). If selection is important, gene flow should be reduced between different selective environments. If geography is important, gene flow should be impeded by geographic distance and physical barriers. We examined how genetic divergence, long-term gene flow, and contemporary dispersal within a watershed were influenced by waterfalls, geographic distance, predation, and habitat features. We found that waterfalls and geographic distance increased genetic divergence and reduced dispersal and long-term gene flow. Differences in predation or habitat features did not influence genetic divergence or gene flow. In contrast, differences in predation did appear to reduce contemporary dispersal. We suggest that the standard predictions of ecological speciation may be heavily nuanced by the mating behaviour and life history strategies of guppies.