No abstract

Microscopic observation of the skin of Plestiodon lizards, which have body stripes and blue tail coloration, identified epidermal melanophores and three types of dermal chromatophores: xanthophores, iridophores, and melanophores. There was a vertical combination of these pigment cells, with xanthophores in the uppermost layer, iridophores in the intermediate layer, and melanophores in the basal layer, which varied according to the skin coloration. Skin with yellowish-white or brown coloration had an identical vertical order of xanthophores, iridophores, and melanophores, but yellowish-white skin had a thicker layer of iridophores and a thinner layer of melanophores than did brown skin. The thickness of the iridophore layer was proportional to the number of reflecting platelets within each iridophore. Skin showing green coloration also had three layers of dermal chromatophores, but the vertical order of xanthophores and iridophores was frequently reversed. Skin showing blue color had iridophores above the melanophores. In addition, the thickness of reflecting platelets in the blue tail was less than in yellowish-white or brown areas of the body. Skin with black coloration had only melanophores.

The holotype (MHM-K2) of the Eocene cheloniine Tasbacka danica is arguably one of the best preserved juvenile fossil sea turtles on record. Notwithstanding compactional flattening, the specimen is virtually intact, comprising a fully articulated skeleton exposed in dorsal view. MHM-K2 also preserves, with great fidelity, soft tissue traces visible as a sharply delineated carbon film around the bones and marginal scutes along the edge of the carapace. Here we show that the extraordinary preservation of the type of T. danica goes beyond gross morphology to include ultrastructural details and labile molecular components of the once-living animal. Haemoglobin-derived compounds, eumelanic pigments and proteinaceous materials retaining the immunological characteristics of sauropsid-specific β-keratin and tropomyosin were detected in tissues containing remnant melanosomes and decayed keratin plates. The preserved organics represent condensed remains of the cornified epidermis and, likely also, deeper anatomical features, and provide direct chemical evidence that adaptive melanism – a biological means used by extant sea turtle hatchlings to elevate metabolic and growth rates – had evolved 54 million years ago.

To provide histological foundation for studying the genetic mechanisms of color-pattern polymorphisms, we examined light reflectance profiles and cellular architectures of pigment cells that produced striped, nonstriped, and melanistic color patterns in the snake Elaphe quadrivirgata. Both, striped and nonstriped morphs, possessed the same set of epidermal melanophores and three types of dermal pigment cells (yellow xanthophores, iridescent iridophores, and black melanophores), but spatial variations in the densities of epidermal and dermal melanophores produced individual variations in stripe vividness. The densities of epidermal and dermal melanophores were two or three times higher in the dark-brown-stripe region than in the yellow background in the striped morph. However, the densities of epidermal and dermal melanophores between the striped and background regions were similar in the nonstriped morph. The melanistic morph had only epidermal and dermal melanophores and neither xanthophores nor iridophores were detected. Ghost stripes in the shed skin of some melanistic morphs suggested that stripe pattern formation and melanism were controlled independently. We proposed complete- and incomplete-dominance heredity models for the stripe-melanistic variation and striped, pale-striped, and nonstriped polymorphisms, respectively, according to the differences in pigment-cell composition and its spatial architecture.

Juveniles of numerous lizard species have a vividly blue-coloured tail that likely serves to deflect predator attacks toward the autotomizable tail rather than the lizard's body. The shades of blue colour in the tails of juvenile Plestiodon latiscutatus lizards vary across populations, most notably among those island populations with different predator assemblages. Here, we determine if this intraspecific variation is associated with the differences in colour vision capabilities of lizard predator species. If associated, it would be evidence for local adaptation of tail colour phenotypenatural selection is maximizing the conspicuousness of the tail to the dominant predator species to increase the chance of successfully deflecting attacks. We also use transmission electron microscopy (TEM) to determine the proximate cellular mechanisms that produce the shades of blue in different populations. We revealed that lizard tails with vivid blue reflectance evolved in communities with either weasel or snake predators, two groups of animals with the ability to detect blue wavelengths. However, lizard tail UV reflectance was much higher in populations with only snake predators; that snakes can detect UV, yet weasels cannot, suggests that high UV reflectance is an adaptation to increase tail conspicuousness specifically to snake predators. Finally, a cryptic brown tail evolved independently on the islands where birds are the primary lizard predator. We suggest that because birds have keen visual acuity; a brown, camouflaged phenotype is more advantageous. We also determined through TEM that the thickness of light reflecting platelets in iridophores, and densities of iridophores and xanthophores, predicted the wavelengths and intensity of light reflected by the lizard tail. For example, blue coloration was produced by selective reflection of short wavelengths of light by the thin light reflecting platelets of the iridophore. Greater iridophore density increased light reflectance, while greater xanthophore density decreased light reflectance.

Characterizing the spatial variation in the CO2 flux at both large and small scales is essential for precise estimation of an ecosystem's CO2 sink strength. However, little is known about small-scale CO2 flux variations in an ecosystem. We explored these variations in a Kobresia meadow ecosystem on the Qinghai-Tibetan plateau in relation to spatial variability in species composition and biomass. We established 14 points and measured net ecosystem production (NEP), gross primary production (GPP), and ecosystem respiration (Re) in relation to vegetation biomass, species richness, and environmental variables at each point, using an automated chamber system during the 2005 growing season. Mean light-saturated NEP and GPP were 30.3 and 40.5 micromol CO2 m(-2) s(-1) [coefficient of variation (CV), 42.7 and 29.4], respectively. Mean Re at 20 degrees C soil temperature, Re(20), was -10.9 micromol CO2 m(-2) s(-1) (CV, 27.3). Re(20) was positively correlated with vegetation biomass. GPP(max) was positively correlated with species richness, but 2 of the 14 points were outliers. Vegetation biomass was the main determinant of spatial variation of Re, whereas species richness mainly affected that of GPP, probably reflecting the complexity of canopy structure and light partitioning in this small grassland patch.

Aim We infer the biogeography and colonization history of a dispersal-limited terrestrial vertebrate, the Japanese four-lined ratsnake (Elaphe quadrivirgata), to reveal the number of times mainland populations have invaded the Izu Archipelago of Japan, the mainland sources of these colonists, and the timescale of colonization. We compare these results with those of past studies in an attempt to uncover general biogeographical patterns. Moreover, we briefly examine the significance of colonization history when evaluating the evolution of body size and melanism of the Izu Island E. quadrivirgata populations.Location The Izu Islands (Oshima, Toshima, Niijima, Shikine, Kozu, Tadanae and Mikura), a volcanic archipelago off the Pacific coast of central Japan.Methods We obtained DNA sequences of the mitochondrial cytochrome b gene (1117 base pairs) from 373 individual snakes sampled from seven of the Izu Islands and 25 mainland localities. We employed partitioned Bayesian phylogenetic analyses assuming a relaxed molecular clock to estimate phylogenetic relationships among extant haplotypes and to give an explicit temporal scale to the timing of clade divergence, colonization history and tempo of body-size evolution. Moreover, we employed model-based biogeographical analysis to calculate the minimum number of times E. quadrivirgata colonized the Izu Islands.Results We found evidence that three separate regions of the Izu Archipelago have been colonized independently from mainland ancestors within the past 0.58-0.20 Ma. The Izu Peninsula plus Oshima and Mikura were both colonized independently from lineages inhabiting eastern mainland Japan. The Toshima, Niijima, Shikine, Kozu and Tadanae populations all derive from a single colonization from western mainland Japan. Oshima has been subject to three or possibly four colonizations.Main conclusions These results support the hypothesis that the extreme body-size disparity among island populations of this ratsnake evolved in situ. Moreover, the fact that the dwarf, melanistic population inhabiting Oshima descends from multiple mainland colonization events is evidence of an extremely strong natural selection pressure resulting in the rapid evolution of this unique morphology. These results contrast with theoretical predictions that natural selection pressures should play a decreased role on islands close to the mainland and/or subject to frequent or recent immigration.

Predation may create strong natural selection pressure on the phenotype and life history characteristics of prey species. The Izu scincid lizards (Plestiodon latiscutatus) that inhabit the four Japanese Izu Islands with only bird predators are drab brown, mature later, lay small clutches of large eggs, and hatch large neonates. In contrast, skinks on seven islands with both snake and bird predators are conspicuously colored, mature early, lay large clutches of small eggs, and hatch small neonates. We test the hypothesis that these suites of traits have evolved independently on each island via natural selection pressures from one of two predator regimes – birds-only and birds + snakes. Using two mtDNA genes and a nuclear locus, we infer a time-calibrated phylogeny of P. latiscutatus that reveals a basal split between Mikura and all islands south, and Miyake, all islands north, and the Izu Peninsula. Populations inhabiting Miyake, Niijima, Shikine, and Toshima are not monophyletic, suggesting either multiple colonizations or an artifact of incomplete lineage sorting (ILS). We therefore developed novel phylogenetic comparative analyses that assume either a multiple colonization or more restrictive single colonization ILS scenario and found 1) statistically significant support for the of different suites of phenotypic and life history characteristics with the presence of bird-only or bird + snake predator assemblages, and 2) strong phylogenetic support for at least two independent derivations of either the “bird-only” or “snakes + birds” phenotypes regardless of colonization scenario. Finally, our time-calibrated phylogeographic analysis supports the conclusion that the ancestor to modern Izu Island P. latiscutatus dispersed from the mainland to the Izu proto-islands between 3–7.6 million years ago (Ma). These lineages remained present in the area during successive formation of the islands, with one lineage re-colonizing the mainland 0.24-0.7 Ma.