Maximum body size in a radiating clade as a function of time

The behavior of the maximum body size (body length) in an evolving clade is exemplified by the evolutionary histories of Bivalvia, Cetacea, and Camerata (Crinoidea). Changes of the maximum size with time track closely diversification history: when a clade diversifies exponentially, the maximum size also increases exponentially, and when the number of species changes irregularly (at varying rates), the maximum size also changes in that manner. However, within any given clade, the maximum body size changes at lower rates than diversity does. The observed shifts in maximum body size approximate the rate of diversification per unit of time to the power of about 0.5.     

Limited role of character displacement in the coexistence of congeneric Anelosimus spiders in a Madagascan montane forest

Evolutionary and ecological theory predicts that closely related and similar species should coexist infrequently because speciation is more likely to occur allopatrically than sympatrically, and because co‐occurring species with similar traits may compete for limited resources, leading to competitive exclusion or character displacement. Here we study the unusual coexistence of 10 similar congeneric species of Anelosimus spiders within a small forest fragment in Madagascar. We asked if these species radiated in sympatry or allopatry, and if there was evidence for local‐scale character displacement in body size and other species‐level traits. We sampled ～ 350 colonies (6346 individuals) along a 2800 m transect. We identified colonies using morphology and DNA barcoding, and tested the monophyly of local and regional species assemblages with time‐calibrated phylogenies. We used null model analysis and phylogenetic signal inference to test for patterns of segregation in body size, microhabitat, phenology, and seasonality of coexisting species. We found that all species belong to a Madagascan clade that radiated during the Pliocene, but that contemporary local assemblages are non‐monophyletic. This is consistent with allopatric speciation during periods of global cooling and expansion of grasslands, and subsequent species assembly as forest fragments re‐expanded and coalesced. We found no evidence for character displacement, except for overdispersion and even spacing in phenology: species were segregated by instars in a manner consistent with resource partitioning or maintenance of reproductive isolation. Overdispersion or even spacing in phenology may contribute to coexistence either through resource partitioning or mate recognition. However, there was no support for a scenario of resource partitioning and divergence of body size or other correlated morphological characters. These traits are better explained by evolutionary forces operating during speciation, rather than ecological forces operating during local community assembly.     

Partitioning variance in a physiological trait: desiccation resistance in keratin beetles (Coleoptera, Trogidae)

1. Although variation in physiological traits forms the substance of evolutionary physiology, the way that this variation is partitioned among different hierarchical levels (e.g. population, species, genus) is not well known.
2. In this study variance partitioning is examined in body size, water content, lipid content, rate of water loss, maximum tolerable water loss and survival time at four levels (individual, population, species, genus) in southern African keratin beetles.
3. It is found that most variance in body size, and the physiological traits that are strongly influenced by body size (water and lipid content, maximum tolerable water loss, rate of water loss), is partitioned at the generic level (50–70%), then at the species level (20–50%) and finally at the population (1–9%) and individual (2–18%) levels.
4. On the other hand, variance in survival time, and variance in rate of maximum water loss once the effects of body size have been taken into account, are partitioned mostly at the species level (40–70%), whereas maximum tolerable water loss and lipid and water content show greatest variance at the individual level (63–75%). This is largely a consequence of differences in the extent of scaling of these traits.
5. The results suggest that where possible the effects of phylogeny should be controlled for when examining desiccation resistance in insects using comparative methods, but that confidence in conclusions from previous studies, which have used mass-specific data, is not unwarranted.     

How is dispersal integrated in life histories: a quantitative analysis using butterflies

As dispersal plays a key role in gene flow among populations, its evolutionary dynamics under environmental changes is particularly important. The inter-dependency of dispersal with other life history traits may constrain dispersal evolution, and lead to the indirect selection of other traits as a by-product of this inter-dependency. Identifying the dispersal's relationships to other life-history traits will help to better understand the evolutionary dynamics of dispersal, and the consequences for species persistence and ecosystem functioning under global changes. Dispersal may be linked to other life-history traits as their respective evolutionary dynamics may be inter-dependent, or, because they are mechanistically related to each other. We identify traits that are predicted to co-vary with dispersal, and investigated the correlations that may constrain dispersal using published information on butterflies. Our quantitative analysis revealed that (1) dispersal directly correlated with demographic traits, mostly fecundity, whereas phylogenetic relationships among species had a negligible influence on this pattern, (2) gene flow and individual movements are correlated with ecological specialisation and body size, respectively and (3) routine movements only affected short-distance dispersal. Together, these results provide important insights into evolutionary dynamics under global environmental changes, and are directly applicable to biodiversity conservation.     

An interspecific comparison between morphology and swimming performance in cyprinids

Flow regimes are believed to be of major evolutionary significance in fish. The flow regimes inhabited by cyprinids vary extensively from still flow regimes to riptide flow regimes. To test (i) whether flow‐driven swimming performance and relevant morphological differentiation are present among fish species and (ii) whether evolutionary shifts between high‐flow and low‐flow habitats in cyprinids are associated with evolutionary trade‐offs in locomotor performance, we obtained data on both steady and unsteady swimming performance and external body shape for 19 species of cyprinids that typically occur in different flow regimes (still, intermediate and riptide). We also measured the routine energy expenditure (RMR) and maximum metabolic rate (MMR) and calculated the optimal swimming speed. Our results showed that fish species from riptide groups tend to have a higher critical swimming speed (U_crit), maximum linear velocity (V_max) and fineness ratio (FR) than fish from the other two groups. However, there was no correlation between the reconstructed changes in the steady and unsteady swimming performance of the 19 species. According to the phylogenetically independent contrast (PIC) method, the U_crit was actively correlated with the MMR. These results indicated that selection will favour both higher steady and unsteady swimming performance and a more streamlined body shape in environments with high water velocities. The results suggested that steady swimming performance was more sensitive to the flow regime and that for this reason, changes in body shape resulted more from selective pressure on steady swimming performance than on unsteady swimming performance. No evolutionary trade‐off was observed between steady and unsteady swimming performance, although U_crit and MMR were found to have coevolved. However, a further analysis within each typically occurring habitat group suggested that the trade‐off that may exist between steady and unsteady swimming performance may be concealed by the effect of habitat.     

Evolutionary changes of nonlinear reaction norms according to thermal adaptation: a comparison of two Drosophila species

While the adaptive significance of discontinuous reaction norms is generally accepted, the evolutionary interpretation of continuous response curves remains speculative, and the occurrence of internal constraints is often suggested as an explanation of experimental observations. In Drosophila melanogaster, various morphometrical traits exhibit convex reaction norms to growth temperature, with a maximum value within the developmental thermal range. We compared a cold-adapted species (D. subobscura) with a mid thermal range at 16 °C, to the warm-adapted D. melanogaster (mid thermal range at 22 °C) for three different morphometrical traits: wing and thorax length in both sexes and ovariole number in females. Maximum value temperatures were ordered in the same way for the three traits in both species: ovariole number > thorax length > wing length. Significant differences were also observed between the two species for the curvature parameter of the quadratic adjustment. The major observation was a significant lateral shift in the reaction norms: maximum values were observed at much lower temperatures in the cold-adapted species than in the warm-adapted one. The parallelism between mid thermal range variation and the position of the maximum value strongly suggests an adaptive displacement of the response curves. Natural selection may thus act not only on trait mean values but also on phenotypic plasticity and on the shape of reaction norms.     

Reliability of leg muscle electromyography in vertical jumping

In this study we aimed to determine the reliability of the surface electromyography (EMG) of leg muscles during vertical jumping between two test sessions, held 2 weeks apart. Fifteen females performed three maximal vertical jumps with countermovement. The displacement of the body centre of mass (BCM), duration of propulsion phase (time), range of motion (ROM) and angular velocity of the knee and surface EMG of four leg muscles (rectus femoris, vastus medialis. biceps femoris and gastrocnemius) were recorded during the jumps. All variables were analysed throughout the propulsion and mid-propulsion phases. Intraclass correlation coefficients (ICC) for the rectus femoris, vastus medialis, biceps femoris and gastrocnemius were calculated to be 0.88, 0.70, 0.24 and 0.01, respectively. BCM, ROM and time values all indicated ICC values greater than 0.90, and the mean knee angular velocity was slightly lower, at 0.75. ICCs between displacement of the BCM and integrated EMG (IEMG) of the muscles studied were less than 0.50. The angular velocity of the knee did not correlate well with muscle activity. Factors that may have affected reliability were variations in the position of electrode replacement, skin resistance, cross-talk between muscles and jump mechanics. The results of this study suggest that while kinematic variables are reproducible over successive vertical jumps, the degree of repeatability of an IEMG signal is dependent upon the muscle studied.     

Plasticity of ontogenetic trajectories in cyprinids: a source of evolutionary novelties

Research in evolutionary developmental (evo‐devo) biology is making an increasingly important contribution to our understanding of the molecular mechanisms underlying the establishment of complex morphological traits. Deciphering the ontogenetic trajectories leading to the differentiation of sister species (and the existence of hybrids) is a new challenge in our understanding of speciation processes. In the present study, we characterized the ontogenetic trajectory of lower lip morphology in two cyprinid species and their hybrids. Chondrostoma toxostoma has an arched lower lip and a generalist diet. Chondrostoma nasus has a straight lower lip and a specialist diet. An analysis of 99 C. toxostoma, 99 C. nasus and 25 first‐generation (F₁) hybrid individuals demonstrated that the difference between arched and straight lip morphology was found to depend strongly on the height/width ratio of the lower lip. A comparison of the ontogenetic trajectories of these morphometric traits showed that the height of the lower lip was isometric to body length in both species, whereas developmental changes involving an acceleration and a hypermorphosis of the widening of the lower lip led to a straight lip morphology in C. nasus. F₁ hybrids were characterized by an extreme phenotype resulting from a rate of lower lip widening slower than that in the two parent species. Therefore, we rejected a codominance hypothesis and concluded that the first stage of hybridization provides the foundations of evolutionary novelty. These results have important evolutionary implications because lower lip shape is linked to dietary behaviour in many fish species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 342–355.     

Similarity of growth among Great tits (Parus major) and Blue tits (P. caeruleus)

Differences in morphology among species are proximately caused by changes in the ontogeny of individuals. It is therefore of importance to analyse possible differences in growth parameters among closely related species in order to understand what parameters are most and least likely, respectively, to change in evolution. In this paper I analyse growth in two closely related sympatric species, namely Great tit, Parus major, and Blue tit, P. caeruleus. The former is considerably larger than the latter in all external traits. The growth rates of the two species were found to be very similar for all traits, thus excluding differences in growth rate as a potential cause of evolutionary size changes. Offset of growth occurred at relatively similar times in the two species, excluding this factor as a major cause of the final size differences. However, size differences at hatching were pronounced and remained so throughout ontogeny, pointing to initial size (egg size or hatching size) as the target of factors promoting change. Bivariate allometric relations of traits vs. body size (mass) were similar between the two species at all ontogenetic stages. There was a high correlation among traits especially at intermediate age stages (5 and 8 days), but these correlations became weaker at older age and approached the low pattern of integration found in adults. All this suggests the operation of a general growth factor affecting all parts of the phenotype simultaneously, which has its major influence at the time of maximal growth. If closely related species in general have highly similar growth patterns, strong evolutionary allometry as found in many avian taxa is to be expected.     

Insulin-like growth factor-1 is associated with life-history variation across Mammalia

Despite the diversity of mammalian life histories, persistent patterns of covariation have been identified, such as the ‘fast–slow’ axis of life-history covariation. Smaller species generally exhibit ‘faster’ life histories, developing and reproducing rapidly, but dying young. Hormonal mechanisms with pleiotropic effects may mediate such broad patterns of life-history variation. Insulin-like growth factor 1 (IGF-1) is one such mechanism because heightened IGF-1 activity is related to traits associated with faster life histories, such as increased growth and reproduction, but decreased lifespan. Using comparative methods, we show that among 41 mammalian species, increased plasma IGF-1 concentrations are associated with fast life histories and altricial reproductive patterns. Interspecific path analyses show that the effects of IGF-1 on these broad patterns of life-history variation are through its direct effects on some individual life-history traits (adult body size, growth rate, basal metabolic rate) and through its indirect effects on the remaining life-history traits. Our results suggest that the role of IGF-1 as a mechanism mediating life-history variation is conserved over the evolutionary time period defining mammalian diversification, that hormone–trait linkages can evolve as a unit, and that suites of life-history traits could be adjusted in response to selection through changes in plasma IGF-1.     

Evolution of conspicuous colouration, body size and gregariousness: a comparative analysis of lepidopteran larvae

Protective colouration in animals includes camouflage (i.e., crypsis), that decreases the risk of detection, and conspicuous colouration, which is often used in combination with chemical defences to deter predators from attacking. Experiments have shown that the efficacy of conspicuous colouration increases with increasing size of pattern elements and larger body size. Prey species that have acquired avoidance inducing colouration therefore may be exposed to selection for larger body size, and such colouration may more easily evolve in large than in small prey species. Here we test for a difference in body size between species with different colouration modes and perform a comparative analysis based on phylogenetically independent contrasts to examine if evolutionary shifts in colour pattern have been associated with evolutionary changes in body size, using data for 578 species of moths. Larval body size did not differ between species with signalling and non-signalling larvae, and results from the comparative analysis suggest that these two traits have not evolved in parallel. The lack of association between evolutionary changes in colouration and body size may reflect a confounding influence of lifestyle, because evolutionary shifts from solitary to group-living larvae were associated with decreased larval body length and adult wing span. Because evolutionary changes in larval body size were associated with evolutionary changes in adult wing span the predicted association between colouration and size may have been confounded also by conflicting selection on body size in larvae and adults.     

What is comparable in comparative cognition?

To understand how complex, or 'advanced' various forms of cognition are, and to compare them between species for evolutionary studies, we need to understand the diversity of neural-computational mechanisms that may be involved, and to identify the genetic changes that are necessary to mediate changes in cognitive functions. The same overt cognitive capacity might be mediated by entirely different neural circuitries in different species, with a many-to-one mapping between behavioural routines, computations and their neural implementations. Comparative behavioural research needs to be complemented with a bottom-up approach in which neurobiological and molecular-genetic analyses allow pinpointing of underlying neural and genetic bases that constrain cognitive variation. Often, only very minor differences in circuitry might be needed to generate major shifts in cognitive functions and the possibility that cognitive traits arise by convergence or parallel evolution needs to be taken seriously. Hereditary variation in cognitive traits between individuals of a species might be extensive, and selection experiments on cognitive traits might be a useful avenue to explore how rapidly changes in cognitive abilities occur in the face of pertinent selection pressures.     

The evolutionary rate of leaf osmotic strength drives diversification of Primulina species in karst regions

Leaf water storage capacity and osmotic strength are important traits enabling species to adapt to environments that are often moisture limited. However, whether these drought tolerance traits are correlated with the species diversification rate (DR) of plant lineages is yet to be determined. In this study, we selected a species-rich genus (Primulina) of plants widely distributed in karst regions in which species frequently experience variable periods of drought. We measured water storage capacity-related traits (including leaf thickness and water content per mass) and saturated osmotic strength in the leaves of 58 Primulina species growing in a common garden. Subsequently, using phylogenetic methods, we examined the relationships between the rate of species diversification and the drought tolerance traits and between the species DR and evolutionary rates of these traits. We found that neither water storage capacity nor saturated osmotic strength showed significant relationships with the rate of species diversification. However, the evolutionary rate of saturated osmotic strength showed a significant correlation with the species DR, although no comparable significant relationship was detected regarding the evolutionary rate of water storage capacity. Our study indicates that the diversification among Primulina species has typically been accompanied by an extensive divergence of leaf osmotic strength but not a divergence in leaf water storage capacity. These findings will enhance our current understanding of how drought tolerance influences the diversification of plant species in karst regions.     

Early diversification of sperm size in the evolutionary history of the old world leaf warblers (Phylloscopidae)

Sperm morphological traits are highly variable among species and are commonly thought to evolve by post‐copulatory sexual selection. However, little is known about the evolutionary dynamics of sperm morphology, and whether rates of evolutionary change are variable over time and among taxonomic groups. Here, we examine sperm morphology from 21 species of Old World leaf warblers (Phylloscopidae), a group of generally dull, sexually monochromatic birds, which are known to have high levels of extra‐pair paternity. We found that sperm length differs markedly across species, spanning about 40% of the range observed across a larger selection of passerine birds. Furthermore, we found strong support for an ‘early‐burst’ model of trait evolution, implying that the majority of divergence in sperm length has occurred early in the evolutionary history of this clade with subsequent evolutionary stasis. This large early divergence matches the early divergence reported in ecological traits (i.e. body size and feeding behaviour). Our findings demonstrate that rates of evolution in sperm morphology can change over time in passerine taxa, and that evolutionary stasis in sperm traits can occur even in species exhibiting characteristics consistent with moderate‐to‐high levels of sperm competition. It remains a major challenge to identify the selection mechanisms and possible constraints responsible for these variable rates of sperm evolution.     

The role of interspecific interference competition in character displacement and the evolution of competitor recognition

The extent to which interspecific interference competition has contributed to character evolution is one of the most neglected problems in evolutionary biology. When formerly allopatric species come into secondary contact, aggressive interactions between the species can cause selection on traits that affect interspecific encounter rates (e.g. habitat preferences, activity schedules), competitor recognition (e.g. colouration, song), and fighting ability (e.g. weaponry, body size). We define agonistic character displacement (ACD) as the process of phenotypic evolution in a population caused by interference competition with one or more sympatric species and which results in shifts in traits that affect the rate, intensity or outcome of interspecific aggression. After clarifying the relationships between ACD and other evolutionary processes that may occur when species come into secondary contact, we develop an individual‐based, quantitative genetic model to examine how traits involved in competitor recognition would be expected to evolve under different secondary contact scenarios. Our simulation results show that both divergence and convergence are possible outcomes, depending on the intensity of interspecific exploitative competition, the costs associated with mutual versus unilateral recognition, and the extent of phenotypic differences prior to secondary contact. We then devise a set of eight criteria for evaluating putative examples of ACD and review the empirical literature to assess the strength of existing evidence and to identify promising avenues for future research. Our literature search revealed 33 putative examples of ACD across insects, fishes, bats, birds, lizards, and amphibians (15 divergence examples; 18 convergence examples). Only one example satisfies all eight criteria for demonstrating ACD, but most case studies satisfy four or more criteria. The current state of the evidence for ACD is similar to the state of the evidence for ecological character displacement just 10 years ago. We conclude by offering suggestions for further theoretical and empirical research on ACD.     

Comparing different approaches for determining joint torque parameters from isovelocity dynamometer measurements

Strength, or maximum joint torque, is a fundamental factor governing human movement, and is regularly assessed for clinical and rehabilitative purposes as well as for research into human performance. This study aimed to identify the most appropriate protocol for fitting a maximum voluntary torque function to experimental joint torque data. Three participants performed maximum isometric and concentric-eccentric knee extension trials on an isovelocity dynamometer and a separate experimental protocol was used to estimate maximum knee extension angular velocity. A nine parameter maximum voluntary torque function, which included angle, angular velocity and neural inhibition effects, was fitted to the experimental torque data and three aspects of this fitting protocol were investigated. Using an independent experimental estimate of maximum knee extension angular velocity gave lower variability in the high concentric velocity region of the maximum torque function compared to using dynamometer measurements alone. A weighted root mean square difference (RMSD) score function, that forced the majority (73-92%) of experimental data beneath the maximum torque function, was found to best account for the one-sided noise in experimental torques resulting from sub-maximal effort by the participants. The suggested protocol (an appropriately weighted RMSD score function and an independent estimate of maximum knee extension angular velocity) gave a weighted RMSD of between 11 and 13 Nm (4-5% of maximum isometric torque). It is recommended that this protocol be used in generating maximum voluntary joint torque functions in all torque-based modelling of dynamic human movement.     

The life history legacy of evolutionary body size change in carnivores

We estimate the body sizes of direct ancestors of extant carnivores, and examine selected aspects of life history as a function not only of species' current size, but also of recent changes in size. Carnivore species that have undergone marked recent evolutionary size change show life history characteristics typically associated with species closer to the ancestral body size. Thus, phyletic giants tend to mature earlier and have larger litters of smaller offspring at shorter intervals than do species of the same body size that are not phyletic giants. Phyletic dwarfs, by contrast, have slower life histories than nondwarf species of the same body size. We discuss two possible mechanisms for the legacy of recent size change: lag (in which life history variables cannot evolve as quickly as body size, leading to species having the 'wrong' life history for their body size) and body size optimization (in which life history and hence body size evolve in response to changes in energy availability); at present, we cannot distinguish between these alternatives. Our finding that recent body size changes help explain residual variation around life history allometries shows that a more dynamic view of character change enables comparative studies to make more precise predictions about species traits in the context of their evolutionary background.     

Allometry of reproduction in two species of gekkonid lizards (Gehyra): effects of body size miniaturization on clutch and egg sizes

In squamate reptiles there is an allometric pattern for small-bodied females to have smaller clutches and proportionally larger eggs than large-bodied females, and this pattern occurs both among and within species. The allometric patterns in two species of the gecko Gehyra were studied to see how evolutionary reductions in adult body size affect fecundity and offspring size among species, and how these changes affect allometric relationships within species. Gehyra dubia has two eggs per clutch (the typical clutch size for gekkonid lizards), whereas the smallerbodied G. variegata has a single egg per clutch. Within both species, egg size increased with female body size. The data are consistent with at least two mechanistic hypotheses: (1) that the width of the pelvis constrains egg size; and (2) in species with invariant clutch sizes, larger females can only allocate additional energy towards egg size and not number. More direct tests of these hypotheses are warranted. Miniaturization of body sizes in Gehyra is correlated with a clutch size reduction of 50% (from two to one), and a large (1.7-fold) compensatory increase in relative egg mass. However, the small-bodied G. variegata (one egg per clutch) had a lower relative clutch mass than did G. dubia. These findings have implications for understanding the influence of evolutionary reductions in body size on reproductive traits, and for allometric trends in squamate reptiles in general.     

Evolution of larval segment position across 12 Drosophila species*

Many developmental traits that are critical to the survival of the organism are also robust. These robust traits are resistant to phenotypic change in the face of variation. This presents a challenge to evolution. In this article, we asked whether and how a well-established robust trait, Drosophila segment patterning, changed over the evolutionary history of the genus. We compared segment position scaled to body length at the first-instar larval stage among 12 Drosophila species. We found that relative segment position has changed many times across the phylogeny. Changes were frequent, but primarily small in magnitude. Phylogenetic analysis demonstrated that rates of change in segment position are variable along the Drosophila phylogenetic tree, and that these changes can occur in short evolutionary timescales. Correlation between position shifts of segments decreased as the distance between two segments increased, suggesting local control of segment position. The posterior-most abdominal segment showed the highest magnitude of change on average, had the highest rate of evolution between species, and appeared to be evolving more independently as compared to the rest of the segments. This segment was exceptionally elongated in the cactophilic species in our dataset, raising questions as to whether this change may be adaptive.