Weather anomalies more important than climate means in driving insect phenology.

Studies of long-term trends in phenology often rely on climatic averages or accumulated heat, overlooking climate variability. Here we test the hypothesis that unusual weather conditions are critical in driving adult insect phenology. First, we generate phenological estimates for Lepidoptera (moths and butterflies) across the Eastern USA, and over a 70 year period, using natural history collections data. Next, we assemble a set of predictors, including the number of unusually warm and cold days prior to, and during, the adult flight period. We then use phylogenetically informed linear mixed effects models to evaluate effects of unusual weather events, climate context, species traits, and their interactions on flight onset, offset and duration. We find increasing numbers of both warm and cold days were strong effects, dramatically increasing flight duration. This strong effect on duration is likely driven by differential onset and termination dynamics. For flight onset, impact of unusual climate conditions is dependent on climatic context, but for flight cessation, more unusually cold days always lead to later termination particularly for multivoltine species. These results show that understanding phenological responses under global change must account for unusual weather events, especially given they are predicted to increase in frequency and severity.

Comparative Phylogenetics of Papilio Butterfly Wing Shape and Size Demonstrates Independent Hindwing and Forewing Evolution.

The complex forces that shape butterfly wings have long been a subject of experimental and comparative research. Butterflies use their wings for flight, camouflage, mate recognition, warning, and mimicry. However, general patterns and correlations among wing shape and size evolution are still poorly understood. We collected geometric morphometric measurements from over 1400 digitized museum specimens of Papilio swallowtails and combined them with phylogenetic data to test two hypotheses: 1) forewing shape and size evolve independently of hindwing shape and size and 2) wing size evolves more quickly than wing shape. We also determined the major axes of wing shape variation and discovered that most shape variability occurs in hindwing tails and adjacent areas. We conclude that forewing shape and size are functionally and biomechanically constrained, whereas hindwings are more labile, perhaps in response to disruptive selective pressure for Batesian mimicry or against predation. The development of a significant, re-usable, digitized data resource will enable further investigation on tradeoffs between flight performance and ecological selective pressures, along with the degree to which intraspecific, local-scale selection may explain macroevolutionary patterns. [Batesian mimicry; Lepidoptera; geometric morphometrics; museum specimens.].

Reconnecting cell and animal lineages: what do cell lineages tell us about the evolution and development of Spiralia?

Cell lineage studies in the clade Eutrochozoa, and especially the Spiralia, remains a rich and relatively untapped source for understanding broad evolutionary developmental problems; including (1) the utility of cell timing formation for phylogenetic hypotheses; (2) the evolution of cell timing changes and its relation to heterochronic patterns; (3) stereotypy or lack thereof in rates of change of cell growth during evolution and its relation to both evolutionary history and current usage; and (4) how mosaic cleavage timing variation may be expected to differ from other groups. A compilation of available cell timing information was made from previous studies where each division was explicitly followed and the total number of cells followed was greater than 24. From that compilation, we performed a series of heuristic and quantitative analyses, including a phylogenetic analysis using cell timing data as characters and analyses of timing variation across all taxa. Our results show that: (1) cell lineage data reconstructs a phylogenetic hypothesis that has similarities, especially among the Mollusca. to the patterns found in morphological and molecular analyses; (2) the mesentoblast (4d) is a unique cell compared to other cell in that it speeds up and slows down relative to other cells in taxa with both unequal and equal cell sizes; (3) some cells that form in the same quartet at the same point in the cell lineage hierarchy have much lower variations than analogous other cells, arguing for architectural constraint or stabilizing selection acting on those cells; and (4) although variation in cell timing generally increases during development, timing of formation of progeny cells in the first quartet has lower variation than the parent cells, arguing that some regulation-like behavior might be present.

Historical and biomechanical analysis of integration and dissociation in molluscan feeding, with special emphasis on the true limpets (Patellogastropoda: Gastropoda).

Modifications of the molluscan feeding apparatus have long been recognized as a crucial feature in molluscan diversification, related to the important process of gathering energy from the environment. An ecologically and evolutionarily significant dichotomy in molluscan feeding kinematics is whether radular teeth flex laterally (flexoglossate) or do not (stereoglossate). In this study, we use a combination of phylogenetic inference and biomechanical modeling to understand the transformational and causal basis for flexure or lack thereof. We also determine whether structural subsystems making up the feeding system are structurally, functionally, and evolutionarily integrated or dissociated. Regarding evolutionary dissociation, statistical analysis of state changes revealed by the phylogenetic analysis shows that radular and cartilage subsystems evolved independently. Regarding kinematics, the phylogenetic analysis shows that flexure arose at the base of the Mollusca and lack of flexure is a derived condition in one gastropod clade, the Patellogastropoda. Significantly, radular morphology shows no change at the node where kinematics become stereoglossate. However, acquisition of stereoglossy in the Patellogastropoda is correlated with the structural dissociation of the subradular membrane and underlying cartilages. Correlation is not causality, so we present a biomechanical model explaining the structural conditions necessary for the plesiomorphic kinematic state (flexoglossy). Our model suggests that plesiomorphically the radular teeth must flex laterally as they pass over the bending plane as a result of the mechanical restrictions in the flexible but inelastic subradular membrane and close association between subradular membrane and cartilages. Relating this model to the specific character states of the clades, we conclude that lack of flexure in patellogastropods is caused by the dissociation of the subradular membrane and cartilage supports.

INTEGRATING DEVELOPMENTAL EVOLUTIONARY PATTERNS AND MECHANISMS: A CASE STUDY USING THE GASTROPOD RADULA.

Determining the connection between ontogeny and phylogeny continues to be a major theme in biology. However, few studies have combined dissection of pattern and process that lead to transformation of complex morphological structures. Here we examine the patterns and processes of shape change in a model system-the gastropod radula. This system is a simple one having only two processes: initial secretion and postsecretional movement of teeth. However, it produces a tremendous amount of shape variability and fusion patterns. To determine both pattern and mechanism of shape change in an evolutionary context, we use three complementary approaches and datasets. First, we use a phylogenetic hypothesis to determine the polarity of developmental events. Second, we perform a morphometric analysis of shape change using relative warp analysis that allows us to locate and compare the direction and magnitude of ontogenetic and phylogenetic shape divergence. These comparisons are the basis for testing hyptheses of heterochrony and heterotopy, and we show how our results do not conform to expectations of pure heterochrony. The rejection of heterochrony as a hypothesis is based on empirically demonstrating (1) initial shape differs in each taxon; (2) a single dimension of shape variability does not simultaneously describe ontogenetic and evolutionary shape changes; and (3) a significantly different shape and size covariance between taxa. This rejection is probably based on spatial changes in initial conditions and not spatial changes caused by the process itself. Finally, we construct a mechanistic model that explains how shape change happens based on the sequence of events during ontogeny. By using the parameters in the model as characters in the phylogenetic dataset, we show that different parts of the system have arisen at different times and become co-opted into the process. By integrating our analyses together we show that spatial process parameters can be responsible for our nonspatial patterns and that different ontogenetic processes can create similar end morphologies.