Developmental Analysis of the Wing Disc in the Mutant Engrailed of DROSOPHILA MELANOGASTER

The engrailed (en) mutation leads to the transformation of the posterior structures of the dorsal mesothoracic disc into those characteristic of the anterior region of the same disc. Similar posterior-anterior duplications have been detected in dorsal as well as ventral structures of all the thoracic segments. —Genetic combinations of en with other pattern mutants have shown their synergistic effect on the posterior wing pattern.—A clonal analysis of the en wing disc shows that en affects its development in a characteristic way. The genetic change, by induced mitotic recombination, of en⁺ into en cells is followed by the corresponding transformation, except when it takes place some cell divisions prior to differentiation.—The en posterior wing disc cells show positive affinities with normal anterior wing disc cells in aggregates.—The mode of action of the en⁺ locus controlling wing disc development is discussed.     

Ancestor–descendant relationships in evolution: origin of the extant pygmy right whale,Caperea marginata

Ancestor–descendant relationships (ADRs), involving descent with modification, are the fundamental concept in evolution, but are usually difficult to recognize. We examined the cladistic relationship between the only reported fossil pygmy right whale, †Miocaperea pulchra, and its sole living relative, the enigmatic pygmy right whale Caperea marginata, the latter represented by both adult and juvenile specimens. †Miocaperea is phylogenetically bracketed between juvenile and adult Caperea marginata in morphologically based analyses, thus suggesting a possible ADR—the first so far identified within baleen whales (Cetacea: Mysticeti). The †Miocaperea–Caperea lineage may show long-term morphological stasis and, in turn, punctuated equilibrium.     

Morphological divergence in giant fossil dormice

Insular gigantism—evolutionary increases in body size from small-bodied mainland ancestors—is a conceptually significant, but poorly studied, evolutionary phenomenon. Gigantism is widespread on Mediterranean islands, particularly among fossil and extant dormice. These include an extant giant population of Eliomys quercinus on Formentera, the giant Balearic genus †Hypnomys and the exceptionally large †Leithia melitensis of Pleistocene Sicily. We quantified patterns of cranial and mandibular shape and their relationships to head size (allometry) among mainland and insular dormouse populations, asking to what extent the morphology of island giants is explained by allometry. We find that gigantism in dormice is not simply an extrapolation of the allometric trajectory of their mainland relatives. Instead, a large portion of their distinctive cranial and mandibular morphology resulted from the population- or species-specific evolutionary shape changes. Our findings suggest that body size increases in insular giant dormice were accompanied by the evolutionary divergence of feeding adaptations. This complements other evidence of ecological divergence in these taxa, which span predominantly faunivorous to herbivorous diets. Our findings suggest that insular gigantism involves context-dependent phenotypic modifications, underscoring the highly distinctive nature of island faunas.     

A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early ray-finned fishes

Gliding adaptations in thoracopterid flying fishes represent a remarkable case of convergent evolution of overwater gliding strategy with modern exocoetid flying fishes, but the evolutionary origin of this strategy was poorly known in the thoracopterids because of lack of transitional forms. Until recently, all thoracopterids, from the Late Triassic of Austria and Italy and the Middle Triassic of South China, were highly specialized ‘four-winged’ gliders in having wing-like paired fins and an asymmetrical caudal fin with the lower caudal lobe notably larger than the upper lobe. Here, we show that the new genus Wushaichthys and the previously alleged ‘peltopleurid’ Peripeltopleurus, from the Middle Triassic (Ladinian, 235–242 Ma) of South China and near the Ladinian/Anisian boundary of southern Switzerland and northern Italy, respectively, represent the most primitive and oldest known thoracopterids. Wushaichthys, the most basal thoracopterid, shows certain derived features of this group in the skull. Peripeltopleurus shows a condition intermediate between Wushaichthys and Thoracopterus in having a slightly asymmetrical caudal fin but still lacking wing-like paired fins. Phylogenetic studies suggest that the evolution of overwater gliding of thoracopterids was gradual in nature; a four-stage adaption following the ‘cranial specialization–asymmetrical caudal fin–enlarged paired fins–scale reduction’ sequence has been recognized in thoracopterid evolution. Moreover, Wushaichthys and Peripeltopleurus bear hooklets on the anal fin of supposed males, resembling those of modern viviparious teleosts. Early thoracopterids probably had evolved a live-bearing reproductive strategy.     

Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution

Background

Integration of diverse data (molecules, fossils) provides the most robust test of the phylogeny of cetaceans. Positioning key fossils is critical for reconstructing the character change from life on land to life in the water.

Methodology/Principal Findings

We reexamine relationships of critical extinct taxa that impact our understanding of the origin of Cetacea. We do this in the context of the largest total evidence analysis of morphological and molecular information for Artiodactyla (661 phenotypic characters and 46,587 molecular characters, coded for 33 extant and 48 extinct taxa). We score morphological data for Carnivoramorpha, †Creodonta, Lipotyphla, and the †raoellid artiodactylan †Indohyus and concentrate on determining which fossils are positioned along stem lineages to major artiodactylan crown clades. Shortest trees place Cetacea within Artiodactyla and close to †Indohyus, with †Mesonychia outside of Artiodactyla. The relationships of †Mesonychia and †Indohyus are highly unstable, however - in trees only two steps longer than minimum length, †Mesonychia falls inside Artiodactyla and displaces †Indohyus from a position close to Cetacea. Trees based only on data that fossilize continue to show the classic arrangement of relationships within Artiodactyla with Cetacea grouping outside the clade, a signal incongruent with the molecular data that dominate the total evidence result.

Conclusions/Significance

Integration of new fossil material of †Indohyus impacts placement of another extinct clade †Mesonychia, pushing it much farther down the tree. The phylogenetic position of †Indohyus suggests that the cetacean stem lineage included herbivorous and carnivorous aquatic species. We also conclude that extinct members of Cetancodonta (whales + hippopotamids) shared a derived ability to hear underwater sounds, even though several cetancodontans lack a pachyostotic auditory bulla. We revise the taxonomy of living and extinct artiodactylans and propose explicit node and stem-based definitions for the ingroup.     

Evolution of median fin modules in the axial skeleton of fishes

Detailed examples of how hierarchical assemblages of modules change over time are few. We found broadly conserved phylogenetic patterns in the directions of development within the median fins of fishes. From these, we identify four modules involved in their positioning and patterning. The evolutionary sequence of their hierarchical assembly and secondary dissociation is described. The changes in these modules during the evolution of fishes appear to be produced through dissociation, duplication and divergence, and co-option. Although the relationship between identified median fin modules and underlying mechanisms is unclear, Hox addresses may be correlated. Comparing homologous gene expression and function in various fishes may test these predictions.The earliest actinopterygians likely had dorsal and anal fins that were symmetrically positioned via a positioning module. The common patterning (differentiation) of skeletal elements within the dorsal and anal fins may have been set into motion by linkage to this positioning module. Frequent evolutionary changes in dorsal and anal fin position indicate a high level of dissociability of the positioning module from the patterning module. In contrast, the patterning of the dorsal and anal fins remains linked: In nearly all fishes, the endo- and exoskeletal elements of the two fins co-differentiate. In all fishes, the exoskeletal fin rays differentiate in the same directions as the endoskeletal supports, indicating complete developmental integration. In acanthopterygians, a new first dorsal fin module evolved via duplication and divergence. The median fins provide an example of how basic modularity is maintained over 400 million years of evolution.     

Genetic data from the extinct giant rat from Tenerife (Canary Islands) points to a recent divergence from mainland relatives

Evolution of vertebrate endemics in oceanic islands follows a predictable pattern, known as the island rule, according to which gigantism arises in originally small-sized species and dwarfism in large ones. Species of extinct insular giant rodents are known from all over the world. In the Canary Islands, two examples of giant rats, †Canariomys bravoi and †Canariomys tamarani, endemic to Tenerife and Gran Canaria, respectively, disappeared soon after human settlement. The highly derived morphological features of these insular endemic rodents hamper the reconstruction of their evolutionary histories. We have retrieved partial nuclear and mitochondrial data from †C. bravoi and used this information to explore its evolutionary affinities. The resulting dated phylogeny confidently places †C. bravoi within the African grass rat clade (Arvicanthis niloticus). The estimated divergence time, 650 000 years ago (95% higher posterior densities: 373 000–944 000), points toward an island colonization during the Günz–Mindel interglacial stage. †Canariomys bravoi ancestors would have reached the island via passive rafting and then underwent a yearly increase of mean body mass calculated between 0.0015 g and 0.0023 g; this corresponds to fast evolutionary rates (in darwins (d), ranging from 7.09 d to 2.78 d) that are well above those observed for non-insular mammals.     

?Kenyaichthyidae fam. nov. and ?Kenyaichthys gen. nov. – First Record of a Fossil Aplocheiloid Killifish (Teleostei,Cyprinodontiformes)

The extant Cyprinodontiformes (killifishes) with their two suborders Cyprinodontoidei and Aplocheiloidei represent a diverse and well-studied group of fishes. However, their fossil record is comparatively sparse and has so far yielded members of the Cyprinodontoidei only. Here we report on cyprinodontiform fossils from the upper Miocene Lukeino Formation in the Tugen Hills of the Central Rift Valley of Kenya, which represent the first fossil record of an aplocheiloid killifish. A total of 169 specimens - mostly extraordinarily well preserved - and a sample of ten extant cyprinodontiform species were studied on the basis of morphometrics, meristics and osteology. A phylogenetic analysis using PAUP was also conducted for the fossils. Both the osteological data and the phylogenetic analysis provide strong evidence for the assignment of the fossils to the Aplocheiloidei, and justify the definition of the new family †Kenyaichthyidae, the new genus †Kenyaichthys and the new species †K. kipkechi sp. nov. The phylogenetic analysis unexpectedly places †Kenyaichthys gen. nov. in a sister relationship to the Rivulidae (a purely Neotropical group), a probable explanation might be lack of available synapomorphies for the Rivulidae, Nothobranchiidae and Aplocheilidae. The specimens of †K. kipkechi sp. nov. show several polymorphic characters and large overlap in meristic traits, which justifies their interpretation as a species flock in statu nascendi. Patterns of variation in neural and haemal spine dimensions in the caudal vertebrae of †Kenyaichthys gen. nov. and the extant species studied indicate that some previously suggested synapomorphies of the Cyprinodontoidei and Aplocheiloidei need to be revised.     

A new Pseudophoxinus (Teleostei,Cyprinidae) species from Southwestern Anatolia,with remarks on the distribution of the genus in western Anatolia

Pseudophoxinus burduricus sp. n. is described from drainages of Salda and Burdur lakes, southwestern Turkey. It is distinguished from other Anatolian Pseudophoxinus by a combination of characters: lateral line incomplete, with 21–39 (commonly 26–37) perforated scales and 47–57+1-2 scales in lateral series; 10½–12½ scale rows between lateral line and dorsal fin origin, 3–4(5) scale rows between lateral line and the pelvic fin origin; dorsal fin commonly with 7½ branched rays; anal fin commonly with 6½ branched rays; 7–8(9) gill rakers on the first branchial arch; a faint and diffuse epidermal black stripe from eye to caudal fin base in alive and preserved individuals; mouth slightly subterminal, tip of mouth cleft on about level of lower margin of eye; snout rounded, its length greater than eye diameter. Comparison is given with all Pseudophoxinus species from western Anatolia.     

Parallel evolutionary trajectories underlie the origin of giant suspension-feeding whales and bony fishes

Giant suspension feeders such as mysticete whales, basking and whale sharks, and the extinct (indicated by ‘†’) †pachycormiform teleosts are conspicuous members of modern and fossil marine vertebrate faunas. Whether convergent anatomical features common to these clades arose along similar evolutionary pathways has remained unclear because of a lack of information surrounding the origins of all groups of large-bodied suspension feeders apart from baleen whales. New investigation reveals that the enigmatic ray-finned fish †Ohmdenia, from the Lower Jurassic (Toarcian, 183.0–175.6 Ma) Posidonia Shale Lagerstätte, represents the immediate sister group of edentulous †pachycormiforms, the longest lived radiation of large vertebrate suspension feeders. †Ohmdenia bisects the long morphological branch leading to suspension-feeding †pachycormiforms, providing information on the sequence of anatomical transformations preceding this major ecological shift that can be compared to changes associated with the origin of modern mysticetes. Similarities include initial modifications to jaw geometry associated with the reduction of dentition, followed by the loss of teeth. The evolution of largest body sizes within both radiations occurs only after the apparent onset of microphagy. Comparing the fit of contrasting evolutionary models to functionally relevant morphological measurements for whales and †pachycormiform fishes reveals strong support for a common adaptive peak shared by suspension-feeding members of both clades.     

Morphological responses of a stream fish to water impoundment

Water impoundment imposes fundamental changes on natural landscapes by transforming rivers into reservoirs. The dramatic shift in physical conditions accompanying the loss of flow creates novel ecological and evolutionary challenges for native species. In this study, we compared the body shape of Cyprinella venusta collected from eight pairs of river and reservoir sites across the Mobile River Basin (USA). Geometric morphometric analysis of the body shape showed that river populations differ from reservoir populations. Individuals inhabiting reservoirs are deep-bodied and have a smaller head, a more anterior dorsal fin, a shorter dorsal fin base and a more ventral position of the eye than C. venusta in streams. The direction of shape divergence within reservoir–river pairs was consistent among pairs of sites, and the shape of C. venusta in reservoirs is strongly correlated with reservoir size. These findings show that physical characteristics of reservoirs drive changes in the morphological attributes of native fish populations, indicating that water impoundment may be an important, yet largely unrecognized, evolutionary driver acting on aquatic biodiversity.     

Allometric growth of the trunk leads to the rostral shift of the pelvic fin in teleost fishes

The pelvic fin position among teleost fishes has shifted rostrally during evolution, resulting in diversification of both behavior and habitat. We explored the developmental basis for the rostral shift in pelvic fin position in teleost fishes using zebrafish (abdominal pelvic fins) and Nile tilapia (thoracic pelvic fins). Cell fate mapping experiments revealed that changes in the distribution of lateral plate mesodermal cells accompany the trunk-tail protrusion. Presumptive pelvic fin cells are originally located at the body wall adjacent to the anterior limit of hoxc10a expression in the spinal cord, and their position shifts rostrally as the trunk grows. We then showed that the differences in pelvic fin position between zebrafish and Nile tilapia were not due to changes in expression or function of gdf11. We also found that hox-independent motoneurons located above the pelvic fins innervate into the pelvic musculature. Our results suggest that there is a common mechanism among teleosts and tetrapods that controls paired appendage positioning via gdf11, but in teleost fishes the position of prospective pelvic fin cells on the yolk surface shifts as the trunk grows. In addition, teleost motoneurons, which lack lateral motor columns, innervate the pelvic fins in a manner independent of the rostral-caudal patterns of hox expression in the spinal cord.     

Mechanosensation in an adipose fin

Adipose fins are found on approximately 20% of ray-finned fish species. The apparently rudimentary anatomy of adipose fins inspired a longstanding hypothesis that these fins are vestigial and lack function. However, adipose fins have evolved repeatedly within Teleostei, suggesting adaptive function. Recently, adipose fins were proposed to function as mechanosensors, detecting fluid flow anterior to the caudal fin. Here we test the hypothesis that adipose fins are mechanosensitive in the catfish Corydoras aeneus. Neural activity, recorded from nerves that innervate the fin, was shown to encode information on both movement and position of the fin membrane, including the magnitude of fin membrane displacement. Thus, the adipose fin of C. aeneus is mechanosensitive and has the capacity to function as a ‘precaudal flow sensor’. These data force re-evaluation of adipose fin clipping, a common strategy for tagging fishes, and inform hypotheses of how function evolves in novel vertebrate appendages.     

Characterizing the Evolutionary Path(s) to Early Homo

Numerous studies suggest that the transition from Australopithecus to Homo was characterized by evolutionary innovation, resulting in the emergence and coexistence of a diversity of forms. However, the evolutionary processes necessary to drive such a transition have not been examined. Here, we apply statistical tests developed from quantitative evolutionary theory to assess whether morphological differences among late australopith and early Homo species in Africa have been shaped by natural selection. Where selection is demonstrated, we identify aspects of morphology that were most likely under selective pressure, and determine the nature (type, rate) of that selection. Results demonstrate that selection must be invoked to explain an Au. africanus—Au. sediba—Homo transition, while transitions from late australopiths to various early Homo species that exclude Au. sediba can be achieved through drift alone. Rate tests indicate that selection is largely directional, acting to rapidly differentiate these taxa. Reconstructions of patterns of directional selection needed to drive the Au. africanus—Au. sediba—Homo transition suggest that selection would have affected all regions of the skull. These results may indicate that an evolutionary path to Homo without Au. sediba is the simpler path and/or provide evidence that this pathway involved more reliance on cultural adaptations to cope with environmental change.     

The Jaw Adductor Muscle Complex in Teleostean Fishes: Evolution,Homologies and Revised Nomenclature (Osteichthyes: Actinopterygii)

The infraclass Teleostei is a highly diversified group of bony fishes that encompasses 96% of all species of living fishes and almost half of extant vertebrates. Evolution of various morphological complexes in teleosts, particularly those involving soft anatomy, remains poorly understood. Notable among these problematic complexes is the adductor mandibulae, the muscle that provides the primary force for jaw adduction and mouth closure and whose architecture varies from a simple arrangement of two segments to an intricate complex of up to ten discrete subdivisions. The present study analyzed multiple morphological attributes of the adductor mandibulae in representatives of 53 of the 55 extant teleostean orders, as well as significant information from the literature in order to elucidate the homologies of the main subdivisions of this muscle. The traditional alphanumeric terminology applied to the four main divisions of the adductor mandibulae – A₁, A₂, A₃, and Aω – patently fails to reflect homologous components of that muscle across the expanse of the Teleostei. Some features traditionally used as landmarks for identification of some divisions of the adductor mandibulae proved highly variable across the Teleostei; notably the insertion on the maxilla and the position of muscle components relative to the path of the ramus mandibularis trigeminus nerve. The evolutionary model of gain and loss of sections of the adductor mandibulae most commonly adopted under the alphanumeric system additionally proved ontogenetically incongruent and less parsimonious than a model of subdivision and coalescence of facial muscle sections. Results of the analysis demonstrate the impossibility of adapting the alphanumeric terminology so as to reflect homologous entities across the spectrum of teleosts. A new nomenclatural scheme is proposed in order to achieve congruence between homology and nomenclature of the adductor mandibulae components across the entire Teleostei.     

New genus and species of the family stichaeidae from Hokkaido,Japan

Specimens of a new genus and species of the stichaeid fish,Leptostichaeus pumilus, were collected from the Okhotsk Sea off Hokkaido in Japan. The present new genus and species clearly differs from all the other genera and species of the stichaeid fishes in the following characters: 3 or 4 pectoral fin rays; 10 or fewer caudal principal rays; 79–82 dorsal spines; no pelvic fin; last interneural spine supporting a single dorsal spine; infraorbital, occipital and lateral line canals absent; moderate size of dorsal spine shorter than eye diameter; membranes of dorsal and anal fins widely connected with caudal fin; a large black spot divided by a yellow band present just above gill cover.     

Sequence Analysis of the Gene Encoding Amylosucrase from Neisseria polysaccharea and Characterization of the Recombinant Enzyme

The Neisseria polysaccharea gene encoding amylosucrase was subcloned and expressed in Escherichia coli. Sequencing revealed that the deduced amino acid sequence differs significantly from that previously published. Comparison of the sequence with that of enzymes of the α-amylase family predicted a (β/α)₈-barrel domain. Six of the eight highly conserved regions in amylolytic enzymes are present in amylosucrase. Among them, four constitute the active site in α-amylases. These sites were also conserved in the sequence of glucosyltransferases and dextransucrases. Nevertheless, the evolutionary tree does not show strong homology between them. The amylosucrase was purified by affinity chromatography between fusion protein glutathione S-transferase–amylosucrase and glutathione-Sepharose 4B. The pure enzyme linearly elongated some branched chains of glycogen, to an average degree of polymerization of 75.     

Female bias for enlarged male body and dorsal fins in Xiphophorus variatus

Female preference for male fin elaborations in Poeciliid fishes may be driven by a sensory bias for increased lateral projection area (LPA) that has existed since the lineages diverged from a common ancestor. Previous research supports this hypothesis demonstrating female Poecilia latipinna, Poecilia mexicana, and Poecilia reticulata prefer males of larger body and dorsal fin size, but exhibit no such preferences when controlling for total LPA. In the current study, we further tested this hypothesis by presenting female platys, Xiphophorus variatus, with pairs of dummy males differing in: (1) body size (holding dorsal fin size constant); (2) dorsal fin size (holding body size constant); and (3) dorsal fin: body size ratio (holding total LPA constant). Females spent more time near dummies of greater body and dorsal fin size; however, in the third experiment, neither fin size, body size, nor any particular dorsal fin + body size combination was preferred. These results provide additional support for the LPA and sensory bias hypotheses, demonstrating that female X. variatus not only prefer males with “swords”, but sailfin-like dorsal fins as well when body size is held constant. Shared preference for increased LPA is consistent with common ancestry of the sensory/neural systems in females of all four species.     

A new Pseudophoxinus (Teleostei,Cyprinidae) species from Asi River Drainage (Turkey)

Pseudophoxinus turani sp. n. is described from the İncesu Spring (Hassa-Hatay) drainage of Asi River, Turkey. It is distinguished from other Eastern Mediterranean Region Pseudophoxinus species by a combination of characters: lateral line incomplete, with 12–25 (commonly 16–21) perforated scales and 38–46+2-3 scales in lateral series (commonly 41–44+2-3); 10–11 scale rows between the lateral line and dorsal-fin origin; 3–4 scale rows between the lateral line and the pelvic–fin origin; dorsal fin with 7½ branched rays; anal fin commonly with 7½ branched rays; 8-11gill rakers on the first branchial arch; dorsal profile markedly convex with marked hump at the nape, ventral profile less convex than dorsal profile; a small, irregular, black blotch on the base of the caudal fin; mouth terminal, with slightly distinct chin, its corner not reaching vertical through anterior margin of eye; snout somewhat long, with rounded tip; and its length greater than eye diameter.     

Stress-induced mutagenesis and complex adaptation

Because mutations are mostly deleterious, mutation rates should be reduced by natural selection. However, mutations also provide the raw material for adaptation. Therefore, evolutionary theory suggests that the mutation rate must balance between adaptability—the ability to adapt—and adaptedness—the ability to remain adapted. We model an asexual population crossing a fitness valley and analyse the rate of complex adaptation with and without stress-induced mutagenesis (SIM)—the increase of mutation rates in response to stress or maladaptation. We show that SIM increases the rate of complex adaptation without reducing the population mean fitness, thus breaking the evolutionary trade-off between adaptability and adaptedness. Our theoretical results support the hypothesis that SIM promotes adaptation and provide quantitative predictions of the rate of complex adaptation with different mutational strategies.