An integrative approach to examining a homology question: shell structures in soft‐shell turtles

Understanding the patterns of shell reduction in turtles is relevant when examining both fossils and living forms. The soft‐shelled turtles (Trionychidae) are characterized by the general reduction of the peripheral bony elements of the carapace, and some species possess structures of contested homology. By examining Remane's ‘principal criteria’, we addressed the primary homology of the prenuchal and the posterior peripheral ossicles (= PPOs) of the Asian flapshell turtles, Lissemys spp., thus evaluating their topological equivalence, their structural quality, and the presence of intermediate forms in ontogeny and phylogeny. We conducted an analysis of gross morphology, bone histology, and ontogeny of these elements in a large sample of living and fossil trionychids and their sister‐group, the carettochelyids. We conclude that the prenuchal comprises a neomorphic structure that does not fulfil any of the homology criteria examined. The assessment of the homology of PPOs is less straightforward because of the presence of partly conflicting evidence. Nevertheless, PPOs and standard peripherals share an antero‐posterior polarity of the ossification pattern, which we interpret as a significant shared underlying developmental pattern. Depending on the phylogenetic position of Lissemys in trionychid phylogeny, the hypothesis of PPOs homology with standard peripherals is a straightforward one or, alternatively, one involving homologous developmental processes at other levels of the hierarchy, resulting in similar microstructural characteristics of these bony shell features. In this respect, we consider the antero‐posterior polarity of the ossification pattern of both PPOs and standard peripherals as providing potential evidence for the homology of the genetic control regulating the expression of both these structures, and therefore we interpret these structures as homologues on the basis of a deeply homologous underlying developmental process. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 462–476.     

Sixteen years of winter stress: an assessment of cold hardiness,growth performance and survival of hybrid poplar clones at a boreal planting site

In recent years, thousands of hectares of hybrid poplar plantations have been established in Canada for the purpose of carbon sequestration and wood production. However, boreal planting environments pose special challenges that may compromise the long‐term survival and productivity of such plantations. In this study, we evaluated the effect of winter stress, that is, frequent freeze‐thaw and extreme cold events, on growth and survival of 47 hybrid poplar clones in a long‐term field experiment. We further assessed physiological and structural traits that are potentially important for cold tolerance for a selected set of seven clones. We found that trees with narrow xylem vessels showed reduced freezing‐induced embolism and showed superior productivity after 16 growing seasons. With respect to cold hardiness of living tissues, we only observed small differences among clones in early autumn, which were nonetheless significantly correlated to growth. Maximum winter cold hardiness and the timing of leaf senescence and budbreak were not related to growth or survival. In conclusion, our data suggest that reduction of freezing‐induced embolism due to small vessel diameters is an essential adaptive trait to ensure long‐term productivity of hybrid poplar plantations in boreal planting environments.     

Altered growth and improved resistance of Arabidopsis against Pseudomonas syringae by overexpression of the basic amino acid transporter AtCAT1

Amino acid transporters in plants are crucial for distributing amino acids between plant organs and cellular compartments. The H⁺‐coupled plasma membrane transporter CAT1 (cationic amino acid transporter 1) facilitates the high‐affinity uptake of basic amino acids. The uptake of lysine (Lys) via the roots was not altered in loss‐of‐function mutants, in accordance with the minor expression of CAT1 in roots, but plants ectopically overexpressing CAT1 incorporated Lys at higher rates. Exogenous Lys inhibited the primary root of Arabidopsis, whereas lateral roots were stimulated. These effects were augmented by the presence or absence of CAT1. Furthermore, the total biomass of soil‐grown plants ectopically overexpressing CAT1 was reduced and the time to flowering was accelerated. These effects were accompanied by only minor changes in the overall amino acid profile. Interestingly, CAT1 belongs to a specific small cluster of nitrogen‐containing metabolite transporter genes that are rapidly up‐regulated upon infection with Pseudomonas syringae and that may participate in the systemic response of plants to pathogen attack. The overexpression of CAT1 indeed enhanced the resistance to the hemibiotrophic bacterial pathogen P. syringae via a constitutively activated salicylic acid (SA) pathway, which is consistent with the developmental defects and the resistance phenotype.     

Implications of life-history transitions on the population genetic structure of the toxigenic marine dinoflagellate Alexandrium tamarense

Genotypic or phenotypic markers for characterization of natural populations of marine microalgae have typically addressed questions regarding differentiation among populations, usually with reference to a single or few clonal isolates. Based upon a large number of contemporaneous isolates from the same geographical population of the toxigenic species Alexandrium tamarense from the North Sea, we uncovered significant genetic substructure and low but significant multilocus linkage disequilibrium (LD) within the planktonic population. Between the alternative molecular genotyping approaches, only amplified fragment length polymorphism (AFLP) revealed cryptic genetic population substructure by Bayesian clustering, whereas microsatellite markers failed to yield concordant patterns. Both markers, however, gave evidence for genetic differentiation of population subgroups as defined by AFLP. A considerable portion of multilocus LD could be attributed to population subdivision. The remaining LD within population subgroups is interpreted as an indicator of frequency shifts of clonal lineages during vegetative growth of planktonic populations. Phenotypic characters such as cellular content and composition of neurotoxins associated with paralytic shellfish poisoning (PSP) and allelochemical properties may contribute to intra- or inter-annual differentiation of planktonic populations, if clonal lineages that express these characters are selectively favoured. Nevertheless, significant phenotypic differentiation for these characters among the genetically differentiated subgroups was only detected for PSP toxin content in two of the four population subgroups. By integrating the analysis of phenotypic and genotypic characteristics, we developed a conceptual population genetic model to explain the importance of life-cycle dynamics and transitions in the evolutionary ecology of these dinoflagellates.     

Male reproductive success and its behavioural correlates in a polygynous mammal,the Galápagos sea lion (Zalophus wollebaeki)

Sexual selection theory predicts competitive males and choosy females. Nevertheless, since molecular marker‐based studies, paternity outside the expected mating patterns has increasingly been described. Even in highly polygynous systems, where paternity is expected to be strongly skewed towards large, dominant males, alternative mating tactics have been suggested. We examined reproductive success in the polygynous Galápagos sea lion (Zalophus wollebaeki). Semiaquatic territoriality allows females to move freely and may lower the degree of polygyny otherwise suggested by both territorial behaviour and strong sexual dimorphism. We assigned paternities with 22 microsatellites and analysed how male reproductive success was related to size, dominance status, intra‐sexual agonistic behaviour, proximity to females, and attendance in the colony. Male behaviour was consistent across two seasons for all parameters under consideration. Attendance was by far the most important determinant of paternal success. Skew in reproductive success towards large, dominant males was weak and dominance status played no role. This appears to be caused by an extremely long reproductive season lasting five or more months, making it difficult for any male to monopolize receptive females. Females seem to choose displaying males that were present in the colony for a long time rather than dominance per se. Sexual dimorphism in Galápagos sea lions may thus be more influenced by selection for fasting than fighting ability. Our data provide further evidence for alternative mating tactics, as several males gained relatively high reproductive success despite short attendance and hardly any involvement in agonistic interactions.     

Small shelly fossils and trace fossils near the Precambrian-Cambrian boundary in the Yukon Territory, Canada

In the past an ‘explosion’ in diversity and abundance of small shelly fossils and of trace fossils has served to mark the base of the Cambrian. However, no evidence has been presented to prove that the ‘explosions’ of the two groups were synchronous. We describe small shelly fossils and trace fossils from the same phosphatic limestone beds that indicate that the two events were separate in time. The small shelly fossils are Anabarites trisulcatus, Hyolithellus cf. H. isiticus, Microcornus? sp., Protohertzina anabarica, P. unguliformis, P. sp. A, Pseudorthotheca sp. A, Rushtonia? sp. A, four types of tuberculate plates and one type of reticulate plate. These fossils represent a restricted, ‘pre-explosion’ fauna and are assigned to the Anabarites-Circotheca-Protohertzina Assemblage Zone, an uppermost Precambrian zone in the Meishucun Stage, Yunnan Province, China. A point at the top of this zone has received strong international endorsement for future designation as the base of the Cambrian. Associated with the small shelly fossils are the trace fossils Cruziana sp. A, Cruziana? sp. B, Rusophycus sp. A, Palaeophycus rubdark and arthropod scratch marks. If found in isolation, this trace fossil assemblage would be considered as post-Precambrian because it includes large, highly organized arthropod traces that are traditionally accepted as occurring above the trace fossil ‘explosion’. We therefore conclude that the trace fossil ‘explosion’ predates the small shelly fossil ‘explosion’. If the proposed location of the base of the Cambrian in Yunnan is accepted, the small shelly fossil ‘explosion’ concept and its relationship to the boundary would not be greatly modified. The trace fossil ‘explosion’, however, would no longer indicate the base of the Cambrian and the ranges of some trace fossils would be extended into the Precambrian.     

Evolution and history of hummingbirds (Aves: Trochilidae) from the Juan Fernandez Islands, Chile

The Juan Fernandez Firecrown Sephanoides fernandensis is an endangered endemic hummingbird that inhabits the Juan Fernandez Islands, 667 km off the coast of Chile. Its population has decreased from several thousand in the early part of this century to approximately 250–400 individuals at present. The reasons for its decline include habitat degradation by anthropogenic forest clearance and the introduction of grazing mammals and rodents. Another hummingbird, the Green-backed Firecrown Sephanoides sephaniodes, inhabits the Juan Fernandez Islands but is also found on the Chilean mainland. It currently numbers several thousand on the Juan Fernandez Islands but was considered rare during the late 19th and early 20th centuries. The sister relationship between the two species has not been critically tested, and so their evolutionary histories on the Juan Fernandez Islands remain uncertain. With the use of mtDNA cytochrome b and ND2 phylogenetic reconstructions, our study supports the two species as sister taxa. Moreover, the molecular data suggest that the genus Sephanoides is closely related to the higher altitude Andean hummingbirds typical of the paramo and puna habitats. The molecular divergence between the two species of Sephanoides indicates they may have become isolated from each other less than 1 million years ago, suggesting that S. fernandensis evolved in situ on the Juan Fernandez Islands. We find no evidence of genetic subdivision between populations of S. sephaniodes from the Juan Fernandez Islands and the mainland. In addition, high genetic variation of the Juan Fernandez Islands population does not indicate a long period of isolation of a limited number of S. sephaniodes but instead suggests a recent colonization event, perhaps from several mainland populations. As a result of molecular, morphological and apparent ecological similarities, we suggest that competition by S. sephaniodes may be an additional factor stifling the recovery of S. fernandensis. Possible conservation strategies include habitat restoration and the removal of introduced mammals; immediate implementation of such conservation management plans are necessary to save this species from extinction.