Polyphenism predicts actuarial senescence and lifespan in tiger salamanders.

Actuarial senescence (called 'senescence' hereafter) often shows broad variation at the intraspecific level. Phenotypic plasticity likely plays a central role in among-individual heterogeneity in senescence rate (i.e. the rate of increase in mortality with age), although our knowledge on this subject is still very fragmentary. Polyphenism-the unique sub-type of phenotypic plasticity where several discrete phenotypes are produced by the same genotype-may provide excellent study systems to investigate if and how plasticity affects the rate of senescence in nature. In this study, we investigated whether facultative paedomorphosis influences the rate of senescence in a salamander, Ambystoma mavortium nebulosum. Facultative paedomorphosis, a unique form of polyphenism found in dozens of urodele species worldwide, leads to the production of two discrete, environmentally induced phenotypes: metamorphic and paedomorphic individuals. We leveraged an extensive set of capture-recapture data (8948 individuals, 24 years of monitoring) that were analysed using multistate capture-recapture models and Bayesian age-dependent survival models. Multistate models revealed that paedomorphosis was the most common developmental pathway used by salamanders in our study system. Bayesian age-dependent survival models then showed that paedomorphs have accelerated senescence in both sexes and shorter adult lifespan (in females only) compared to metamorphs. In paedomorphs, senescence rate and adult lifespan also varied among ponds and individuals. Females with good body condition and high lifetime reproductive success had slower senescence and longer lifespan. Late-breeding females also lived longer but showed a senescence rate similar to that of early-breeding females. Moreover, males with good condition had longer lifespan than males with poor body condition, although they had similar senescence rates. In addition, late-breeding males lived longer but, unexpectedly, had higher senescence than early-breeding males. Overall, our work provides one of the few empirical cases suggesting that environmentally cued polyphenism could affect the senescence of a vertebrate in nature, thus providing insights on the ecological and evolutionary consequences of developmental plasticity on ageing.

The interaction between heterochrony and mechanical forces as main driver of floral evolution.

Heterochrony acts as a fundamental process affecting the early development of organisms in creating a subtle shift in the timing of initiation or the duration of a developmental process. In flowers this process is linked with mechanical forces that cause changes in the interaction of neighbouring floral organs by altering the timing and rate of initiation of organs. Heterochrony leads to a delay or acceleration of the development of neighbouring primordia, inducing a change in the morphospace of the flowers. As changes in the timing of development may affect organs differently at different stages of development, these shifts eventually lead to major morphological changes such as altered organ positions, fusions, or organ reductions with profound consequences for floral evolution and the diversification of flowers. By concentrating on early developmental stages in flowers it is possible to understand how heterochrony is responsible for shifts in organ position and the establishment of a novel floral Bauplan. However, it remains difficult to separate heterochrony as a process from pattern, as both are intimately linked. Therefore it is essential to connect different patterns in flowers through the process of developmental change.Examples illustrating the importance of heterochronic shifts affecting different organs of the flower are presented and discussed. These cover the transition from inflorescence to flower through the interaction of bracts and bracteoles, the pressure exercised by the perianth on the androecium and gynoecium, the inversed influence of stamens on petals, and the centrifugal influence of carpels on the androecium. Different processes are explored, including the occurrence of obdiplostemony, the onset of common primordia, variable carpel positions, and organ reduction and loss.

Ontogeny of the autopodial skeleton of the gecko Tarentola (Squamata: Phyllodactylidae).

Squamates exhibit evident diversity in their limb morphology. Gekkotans are a particularly diverse group in this respect. The appearance of toepads in gekkotans usually cooccurs with the reduction or loss of claws. The gecko Tarentola (Phyllodactylidae) shows a unique combination of features among geckos, with toepads, hyperphalangy, and dimorphism of claw expression (claws are retained on digits III and IV, but lost (manus) or strongly reduced (pes) on the remaining digits). Despite being a candidate model for studying embryonic skeletal development of the autopodium, no studies have investigated the autopodial development of the gecko Tarentola in detail. Here, we aim to follow up the development of the autopodial skeleton in T. annularis and T. mauritanica using acid-free double staining. The results indicate that the terminal phalanges of claw-bearing digits III and IV ossify earlier than in the remaining digits. This confirms the differential ossification as a result of claw regression in Tarentola. The strongly reduced second phalanges of digits IV in both the fore- and hindlimbs are the last ossifying phalanges. Such late ossification may precede the evolutionary loss of this phalanx. If this is correct, the autopodia of Tarentola would be an interesting example of both the hyperphalangy in digit I and the process of phalanx loss in digit IV. Delay in ossification of the miniaturised phalanx probably represents an example of paedomorphosis.

The neoteny goldilocks zone: The evolution of neoteny in Ambystoma.

Neoteny is a developmental strategy wherein an organism reaches sexual maturity without associated adult characteristics. In salamanders, neoteny takes the form of individuals retaining aquatic larval characteristics such as external gills upon maturation. Mole salamanders (Ambystoma) occupy a wide range of habitats and areas across the North American continent, and display examples of non-neotenic, facultatively neotenic and obligate neotenic species, providing high variation for investigating the factors influencing the evolution of neoteny. Here, we use phylogenetic comparative methods to test existing hypotheses that neoteny is associated with elevational and latitudinal distribution, cave-associated isolation, and hybridisation-related polyploidy. We also test if neoteny influences the diversity of habitats a species can occupy, since the restriction to an aquatic life should constrain the availability of different niches. We find that neoteny tends to occur in a narrow latitudinal band between 20-30° North, with particularly narrow latitudinal ranges for obligate compared to facultative neotenic species (16-52° North). We also find that facultatively neotenic species occur at elevations more than twice as high as other species on average, and that species with a higher frequency of neoteny typically have lower habitat diversity. Our results suggest that evolutionary transitions between non-neotenic and facultative neoteny states occur at relatively high and approximately equal rates. Moreover, we estimate that obligate neoteny cannot evolve directly from non-neotenic species (and vice versa), such that facultative neoteny acts as an evolutionary 'stepping stone' to and from obligate neoteny. However, our transition rate estimates suggest that obligate neoteny is lost >4-times faster than it evolves, partly explaining the rarity of obligate species. These results support the hypothesis that low latitudes favour the evolution of neoteny, presumably linked to more stable (aquatic) environments due to reduced seasonality, but once evolved it may constrain the diversity of habitats.

Redescription of Ochmacanthus batrachostoma (Miranda-Ribeiro, 1912) (Siluriformes: Trichomycteridae): a possible case of incipient paedomorphism

The lepidophagous stegophiline catfish Ochmacanthus batrachostoma (Miranda-Ribeiro, 1912) is endemic to the Paraná-Paraguai basin and is the only member of its genus in that drainage. It remains a poorly-known taxon mainly due to the historical scarcity of study specimens. The species is herein redescribed on the basis of type and non-type specimens from Brazil, Argentina and Paraguay. Ochmacanthus batrachostoma can be distinguished from congeners by the presence of long maxillary and rictal barbels, reduction of the posterior end of the caudal peduncle, a tadpole-like caudal fin and peduncle, among other morphometric traits and features of internal anatomy. Comparisons show that a number of unique traits of adult O. batrachostoma resemble conditions seen in juvenile specimens of other members of Stegophilinae. Such traits are also more pronounced in juveniles of O. batrachostoma itself. The species also has the smallest maximum body size in Ochmacanthus and among the smallest of any stegophiline. In combination, such observations suggest that the species is paedomorphic, although to a degree less extreme than seen in some other trichomycterids. This is the first possible case of paedomorphosis identified for stegophilines.

O bagre lepidofágico Stegophilinae Ochmacanthus batrachostoma (Miranda-Ribeiro, 1912) é endêmico da bacia Paraná-Paraguai e é único membro do gênero presente na drenagem. Permanece um táxon pouco conhecido principalmente devido à escassez histórica de espécimes para estudo. A espécie é aqui redescrita com base em espécimes tipo e não tipo provenientes do Brasil, Argentina e Paraguai. Ochmacanthus batrachostoma pode ser diferenciado dos congêneres pela presença de longos barbilhões maxilares e rictais, extremidade posterior do pedúnculo caudal reduzida, nadadeira e pedúnculo caudal com forma semelhante a cauda de um girino, além de outras características morfométricas e da anatomia interna. Comparações mostram que certas características exclusivas do adulto de O. batrachostoma se assemelham às condições observadas nos espécimes juvenis de outros Stegophilinae. Essas características são mais pronunciadas nos próprios juvenis de O. batrachostoma. A espécie também apresenta o menor comprimento corporal máximo entre os congêneres e é um dos menores Stegophilinae. Em combinação, essas observações sugerem que a espécie é pedomórfica, mas em grau menos extremo do que observado em outros trichomicterídeos. Esse é o primeiro caso possível de pedomorfose identificado em Stegophilinae.