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Models based on the Ornstein–Uhlenbeck process have become standard for the comparative study of adaptation. Cooper et al. (2016) have cast doubt on this practice by claiming statistical problems with fitting Ornstein–Uhlenbeck models to comparative data. Specifically, they claim that statistical tests of Brownian motion may have too high Type I error rates and that such error rates are exacerbated by measurement error. In this note, we argue that these results have little relevance to the estimation of adaptation with Ornstein–Uhlenbeck models for three reasons. First, we point out that Cooper et al. (2016) did not consider the detection of distinct optima (e.g. for different environments), and therefore did not evaluate the standard test for adaptation. Second, we show that consideration of parameter estimates, and not just statistical significance, will usually lead to correct inferences about evolutionary dynamics. Third, we show that bias due to measurement error can be corrected for by standard methods. We conclude that Cooper et al. (2016) have not identified any statistical problems specific to Ornstein–Uhlenbeck models, and that their cautions against their use in comparative analyses are unfounded and misleading. [adaptation, Ornstein–Uhlenbeck model, phylogenetic comparative method.]

The males of some species of the rove beetle genus Bledius Leach (1819) have a pronounced horn protruding from the anterior margin of the pronotum. The horn function is not known, however in many other horn-bearing beetles, these structures are implicated in male-male combat over females. In this study, scaling relationships are investigated for three species; B. spectabilis Kraatz, 1857, B. frisius Lohse, 1978 and B. tricornis (Herbst, 1784). Based on data collected from museum specimens, we fitted and compared different allometric models on pronotal width versus horn length in each species. A simple linear allometric model best describes the scaling relationship of horns in all three species. The horn allometry in the three Bledius species do not accordingly follow a ‘switch point’ model or a curved allometric relationship, models that are commonly reported to explain the scaling relationship of horns in many other beetle species. We therefore found no evidence of male dimorphism (minor/major phenotypes) in the species examined. The potential function of the horn in Bledius is discussed in light of similar studies.

The increasing availability of 3D-imaging technology provides new opportunities for measuring morphology. Photogrammetry enables easy 3D-data acquisition compared to conventional methods and here we assess its accuracy for measuring the size of deer antlers, a complex morphological structure. Using a proprietary photogrammetry software, we generated 3D images of antlers for 92 individuals from 29 species of cervids that vary widely in antler size and shape and used these to measure antler volume. By repeating the process, we found that the relative error averaged 8.5% of object size. Errors in converting arbitrary voxel units into real volumetric units accounted for 70% of the measurement variance and can therefore be reduced by replicating the conversion. We applied the method to clay models of known volume and found no indication of bias. The estimation was robust against variation in imaging device, distance and operator, but approximately 40 images per specimen were necessary to achieve good precision. We used the method to show that conventional measures of main-beam length are relatively poor estimators of antler volume. Using loose antlers of known weight, we also showed that the volume may be a relatively poor predictor of antler weight due to variation in bone density across species. We conclude that photogrammetry can be an efficient and accurate tool for measuring antlers, and likely many other complex morphological traits. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

11 pages, 3 figures, 1 table, supplementary material https://doi.org/10.1126/sciadv.abm7452.-- Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper, the Supplementary Materials, Dryad, and Zenodo, Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic, This project is funded by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement no. 724324 to L.H.L.). Extra funding for specific sampling expeditions were provided by the National Science Centre of Poland (grant PANIC/2016/23/B/ST10/01936 to P.K.), the ASSEMBLE Plus (Horizon 2020), the Leverhulme Trust (Research Project Award RPG-2016-429 to A.W.), the Russian Science Foundation (grant 18-14-00086 to A.N.O.), Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil [PQ-CNPq 308768/2018-3], Fundação de Amparo à Pesquisa do Estado de São Paulo, Brazil [FAPESP 19/17721-9] and the DISTANTCOM project (CTM2013-42667/ANT to C. Avila). B.F. was supported by Beatriu de Pinós (2019-BP-00183), funded by the AGAUR (Government of Catalonia) and by the Horizon 2020 programme under the Marie Skłodowska-Curie grant (no. 801370). The CAML-CEAMARC cruise of RV Aurora Australis (IPY project no. 53) were supported by the Australian Antarctic Division, the Japanese Science Foundation, the French Polar Institute IPEV, and the Muséum national d’Histoire naturelle, led by voyage leader, M. Riddle, With the institutional support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

The concept of evolvability—the capacity of a population to produce and maintain evolutionarily relevant variation—has become increasingly prominent in evolutionary biology. Paleontology has a long history of investigating questions of evolvability, but paleontological thinking has tended to neglect recent discussions, because many tools used in the current evolvability literature are challenging to apply to the fossil record. The fundamental difficulty is how to disentangle whether the causes of evolutionary patterns arise from variational properties of traits or lineages rather than being due to selection and ecological success. Despite these obstacles, the fossil record offers unique and growing sources of data that capture evolutionary patterns of sustained duration and significance otherwise inaccessible to evolutionary biologists. Additionally, there exist a variety of strategic possibilities for combining prominent neontological approaches to evolvability with those from paleontology. We illustrate three of these possibilities with quantitative genetics, evolutionary developmental biology, and phylogenetic models of macroevolution. In conclusion, we provide a methodological schema that focuses on the conceptualization, measurement, and testing of hypotheses to motivate and provide guidance for future empirical and theoretical studies of evolvability in the fossil record.

Morphometrics has become an indispensable component of the statistical analysis of size and shape variation in biological structures. Morphometric data has traditionally been gathered through low-throughput manual landmark annotation, which represents a significant bottleneck for morphometric-based phenomics. Here we propose a machine-learning-based high-throughput pipeline to collect high-dimensional morphometric data in images of semi rigid biological structures.The proposed framework has four main strengths. First, it allows for dense phenotyping with minimal impact on specimens. Second, it presents landmarking accuracy comparable to manual annotators, when applied to standardized datasets. Third, it performs data collection at speeds several orders of magnitude higher than manual annotators. And finally, it is of general applicability (i.e., not tied to a specific study system).State-of-the-art validation procedures show that the method achieves low error levels when applied to three morphometric datasets of increasing complexity, with error varying from 0.5% to 2% of the structure’s length in the automated placement of landmarks. As a benchmark for the speed of the entire automated landmarking pipeline, our framework places 23 landmarks on 13,686 objects (zooids) detected in 1684 pictures of fossil bryozoans in 3.12 minutes using a personal computer.The proposed machine-learning-based phenotyping pipeline can greatly increase the scale, reproducibility and speed of data collection within biological research. To aid the use of the framework, we have developed a file conversion algorithm that can be used to leverage current morphometric datasets for automation, allowing the entire procedure, from model training all the way to prediction, to be performed in a matter of hours.

Is speciation generally a "special time" in morphological evolution, or are lineage-splitting events just "more of the same" where the end product happens to be two separate lineages? Data on evolutionary dynamics during anagenetic and cladogenetic events among closely related lineages within a clade are rare, but the fossil record of the bryozoan genus Metrarabdotos is considered a textbook example of a clade where speciation causes rapid evolutionary change against a backdrop of morphological stasis within lineages. Here, we point to some methodological and measurement theoretical issues in the original work on Metrarabdotos. We then reanalyze a subset of the original data that can be meaningfully investigated using quantitative statistical approaches similar to those used in the original studies. We consistently fail to find variation in the evolutionary process during within-lineage evolution compared with cladogenetic events: the rates of evolution, the strength of selection, and the directions traveled in multivariate morphospace are not different when comparing evolution within lineages and at speciation events in Metrarabdotos, and genetic drift cannot be excluded as a sufficient explanation for the morphological differentiation within lineages and during speciation. Although widely considered the best example of a punctuated mode of evolution, morphological divergence and speciation are not linked in Metrarabdotos.

1. Comparing relative fit of different models of evolutionary dynamics to time series of phyletic change is a common tool when interpreting the fossil record. However, a measure of relative fit is no guarantee the preferred model describes the data well. Selecting a good model is essential for robust inferences, but we are currently lacking tools to investigate if a model of phyletic evolution represents an adequate description of trait dynamics in fossil data. 2. This study develops a general statistical framework implemented in R for assessing the adequacy of the three most commonly used models of evolution in the fossil record; stasis, directional change and random walk. The statistical framework is applied to 300 fossil time series in order to assess how often the three models represent adequate descriptions of evolutionary dynamics in the fossil record. 3. The model that showed the best relative fit to a particular fossil time series (using AICc) passed all adequacy tests in 219 out of 300 cases (73%, directional trend = 76%, stasis = 64%, random walk = 81%). It is therefore not uncommon that the best model according to AICc does not adequately describe the trait dynamics in a fossil time series. 4. Statistical tests of model adequacy ease evaluation of whether a particular model is a good descriptor of phyletic evolution and can assist in making meaningful inferences of model parameters (e.g., as rates of evolution) and interpretations of evolution in the fossil record.

Bryozoans are a moderately diverse, mostly marine phylum with a fossil record extending to the Early Ordovician. Compared to other phyla, little is known about their phylogenetic relationships at both lower and higher taxonomic levels. Hence, an effort is being made to elucidate their phylogenetic relationships. Here, we present newly sequenced nuclear and mitochondrial genes for 21 cheilostome bryozoans. Combining these data with existing orthologous molecular data, we focus on reconstructing the phylogenetic relationships of Fenestrulina and Microporella, two species-rich genera. They are currently placed in Microporellidae, defined by having a semicircular primary orifice and a proximal ascopore. Our six-gene phylogenetic analysis reveals that the genera Fenestrulina and Microporella are each monophyletic, with the sister clade to Microporella comprising non-microporellids. Our result hence supports the reinstatement of the family Fenestrulinidae Jullien, 1888 for Fenestrulina and genera with comparable frontal shield and ooecial morphologies. Our well-supported phylogeny, based on independent molecular data, lends credit to existing phylogenetic hypotheses based on morphological observations but does not conform to the current classification of these bryozoans. This illustrates the general need for a rethink of bryozoan higher level systematics, ideally based on both morphological and molecular data.

Real-time observation of adaptive evolution in the wild is rare and limited to cases of marked, often anthropogenic, environmental change. Here we present the case of a small population of reed warblers (Acrocephalus scirpaceus) over a period of 19 years (1996–2014) after colonizing a restored wetland habitat in Malta. Our data show a population decrease in body mass, following a trajectory consistent with a population ascending an adaptive peak, a so-called Ornstein–Uhlenbeck process. We corroborate these findings with genetic and ecological data, revealing that individual survival is correlated with body mass, and more than half of the variation in mean population fitness is explained by variation in body mass. Despite a small effective population size, an adaptive response has taken place within a decade. A founder event from a large, genetically variable source population to the southern range margin of the reed warbler distribution likely facilitated this process., Rapid local adaptation could potentially facilitate the recolonization of restored habitats. Here, the authors show that reed warblers have undergone substantial adaptive change in body mass in only 19 years after colonizing a restored wetland in Malta.

Fossil sequences provide observations of phenotypes within a lineage over time and represent essential data for increasing our understanding of phyletic evolution beyond microevolutionary timescales. I investigate if fossil time series of the diatom Stephanodiscus niagarae/yellowstonensis follow evolutionary dynamics compatible with hypotheses for how the adaptive landscape changes when a population enters a new environment. The lineage—which has a remarkably detailed stratigraphic record—invaded Yellowstone Lake immediately after recession of ice from the basin 14,000 years ago. Several phyletic models portraying different types of evolutionary dynamics—both compatible and not compatible with changes in the adaptive landscape following ecological opportunity—were fitted to the fossil times‐series of S. niagarae/yellowstonensis. Different models best describe the three analyzed traits. Two of the models (a new model of decelerated evolution and an Ornstein–Uhlenbeck model) capture trait dynamics compatible with an event of ecological opportunity, whereas the third model (random walk) does not. Entering a new environment may accordingly affect trait dynamics for thousands of years, but the effects can vary across phenotypes. However, tests of model adequacy reveal shortcomings in all three models explaining the trait dynamics, suggesting model development is needed to more fully understand the phyletic evolution in S. niagarae/yellowstonensis.

The dominating view of evolution based on the fossil record is that established species remain more or less unaltered during their existence. Substantial evolution is on the other hand routinely reported for contemporary populations, and most quantitative traits show high potential for evolution. These contrasting observations on long and short time scales are often referred to as the paradox of stasis, which rests on the fundamental assumption that periods of morphological stasis in the fossil record represent minimal evolutionary change. Investigating 450 fossil time series, I demonstrate that the non-accumulating morphological fluctuations during stasis travel similar distances in morphospace space compared to lineages showing directional change. Hence, lineages showing stasis are commonly undergoing considerable amounts of evolution, but this evolution does not accumulate to produce large net evolutionary changes over time. Rates of evolutionary change across modes in the fossil record may be more homogenous than previously assumed and advocated, supporting the claim that substantial evolution is not exclusively or causally linked to the process of speciation. Instead of exemplifying minimal evolution, stasis likely represents information on the dynamics of the adaptive landscape on macroevolutionary time scales, including the persistence of adaptive zones and ecological niches over millions of years. This article is protected by copyright. All rights reserved

The tight brain-body allometry across mammals and primates has motivated and informed many hypotheses about brain evolution in humans and other taxa. While a 2/3 or a 3/4 scaling is often at the core of such research, such exponents are derived from estimates based on particular statistical and evolutionary assumptions without careful consideration of how either may influence findings. Here we quantify primate brain-body allometry using phylogenetic comparative methods based on models of both adaptive and constrained evolution, and estimate and account for observational error in both response and predictor variables. Our results supported an evolutionary model in which brain size is directly constrained to evolve in unison with body size, rather than adapting to changes in the latter. The effects of controlling for phylogeny and observation error were substantial, and our analysis yielded a novel 3/5 scaling exponent for primate brain-body evolutionary allometry. Using this exponent with the latest brain- and body-size estimates to calculate new encephalization quotients for apes, humans, and fossil hominins, we found early hominins were substantially more encephalized than previously thought.

In the threespine stickleback Gasterosteus aculeatus model system, phenotypes are often classified into three morphs according to lateral plate number. Morph identity has been shown to be largely genetically determined, but substantial within‐morph variation in plate number exists. In this study, we test whether plate number has a plastic component in response to salinity in the low‐plated morph using a split‐clutch experiment where families were split in two, one half raised in water at 0 and the other at 30 ppt salt. We find a small salinity‐induced plastic effect on plate number in an unexpected direction, opposite to what we predicted: Fish raised in freshwater on average have slightly more plates than fish raised in saltwater. Our results confirm that heritability of plate number is high. Additionally, we find that variance in plate number at the family level can be predicted from other family level traits, which might indicate that epistatic interactions play a role in creating the observed pattern of lateral plate number variation.

Intralocus sexual conflicts arise whenever the fitness optima for a trait expressed in both males and females differ between the sexes and shared genetic architecture constrains the sexes from evolving independently towards their respective optima. Such sexual conflicts are commonplace in nature, yet their long-term evolutionary consequences remain unexplored. Using a Bayesian phylogenetic comparative framework, we studied the macroevolutionary dynamics of intersexual trait integration in stalk-eyed flies (Diopsidae) spanning a time frame of more than 25 Myr. We report that increased intensity of sexual selection on male eyestalks is associated with reduced intersexual eyestalk integration, as well as sex-specific rates of eyestalk evolution. Despite this, lineages where males have been under strong sexual selection for millions of years still exhibit high levels of intersexual trait integration. This low level of decoupling between the sexes may indicate that exaggerated female eyestalks are in fact adaptive—or alternatively, that there are strong constraints on reducing trait integration between the sexes. Future work should seek to clarify the relative roles of constraints and selection in contributing to the varying levels of intersexual trait integration in stalk-eyed flies, and in this way clarify whether sexual conflicts can act as constraints on adaptive evolution even on macroevolutionary time scales.

The allometric relationship between brain and body size among vertebrates is often considered a manifestation of evolutionary constraints. However, birds and mammals have undergone remarkable encephalization, in which brain size has increased without corresponding changes in body size. Here, we explore the hypothesis that a reduction of phenotypic integration between brain and body size has facilitated encephalization in birds and mammals. Using a large dataset comprising 20,213 specimens across 4,587 species of jawed vertebrates, we show that the among-species (evolutionary) brain-body allometries are remarkably constant, both across vertebrate classes and across taxonomic levels. Birds and mammals, however, are exceptional in that their within-species (static) allometries are shallower and more variable than in other vertebrates. These patterns are consistent with the idea that birds and mammals have reduced allometric constraints that are otherwise ubiquitous across jawed vertebrates. Further exploration of ontogenetic allometries in selected taxa of birds, fishes and mammals reveals that birds and mammals have extended the period of fetal brain growth compared to fishes. Based on these findings, we propose that avian and mammalian encephalization has been contingent on increased variability in brain growth patterns.

Long-term phenotypic stasis is frequently observed in the fossil record, but not readily predicted from microevolutionary theory. To test competing explanations for stasis on macroevolutionary timescales we need reliably estimated parameters from appropriate evolutionary models that adequately describe the evolutionary trait dynamics. Here, we develop tests to assess the adequacy of the most commonly used stasis model in evolutionary biology and apply them to time series of phenotypic traits from fossil lineages. Of the 572 fossil time series we analyzed from the literature, 263 time series showed a better fit to the stasis model relative to alternative models, but only 172 of those fitted the stasis model in both relative and absolute terms. The estimated trait variances from these 172 time series do not correlate with rough proxies of effective population size. Our preliminary investigation of the fixed-optimum hypothesis hence fails to give empirical support to the idea that genetic drift around a constant trait optimum is an explanation for stasis in the fossil record. We argue that optima following stationary processes on the adaptive landscape is a viable hypothesis for stasis that needs further investigation. We end by discussing how investigations of model adequacy can be a valuable approach for increasing our understanding of the dynamics of the adaptive landscape on macroevolutionary timescales.

The cheilostome bryozoan family Steginoporellidae in New Zealand comprises seven living species of Steginoporella. Three of these are new to science—Steginoporella discors n. sp., Steginoporella lineata n. sp. and Steginoporella modesta n. sp.—and one (Steginoporella magnifica) additionally occurs as a Plio-Pleistocene fossil. A new Early Pleistocene fossil species, Steginoporella tiara n. sp., is also recognised. The living species exhibit the full range of colonial morphologies known for the genus, and two of the new deep-shelf taxa described herein have the smallest known colonies, both linear, not exceeding 5 mm in width and 22 mm in length. One species has a recorded depth range down to 615 m, apparently the deepest known for the genus. Zooidal proportions vary, with a length:width ratio in the seven living species ranging from 1.31 to 1.81, exceeded only by that in the new fossil taxon, which has very elongate zooids. Notwithstanding the conspicuous differences in colonial and zooidal morphology, four of the living species appear to be closely related, sharing distinctive reticulation of opercular sclerites, a similar morphology of the median process and no B-zooid morphs. Only one New Zealand taxon has B-zooids. Biogeographically, all the species except S. magnifica (also known from Tonga) are nominally endemic, but it is possible that some of the deeper-water taxa may eventually be found outside the boundary of the New Zealand Exclusive Economic Zone. The operculum in Steginoporella species is initially a single thin layer continuous with the membranous frontal wall, becoming two-layered when fully functioning in feeding zooids and mandibulate B-zooids.

Competition is an important biotic interaction that influences survival and reproduction. While competition on ecological timescales has received great attention, little is known about competition on evolutionary timescales. Do competitive abilities change over hundreds of thousands to millions of years? Can we predict competitive outcomes using phenotypic traits? How much do traits that confer competitive advantage and competitive outcomes change? Here we show, using communities of encrusting marine bryozoans spanning more than 2 million years, that size is a significant determinant of overgrowth outcomes: colonies with larger zooids tend to overgrow colonies with smaller zooids. We also detected temporally coordinated changes in average zooid sizes, suggesting that different species responded to a common external driver. Although species-specific average zooid sizes change over evolutionary timescales, species-specific competitive abilities seem relatively stable, suggesting that traits other than zooid size also control overgrowth outcomes and/or that evolutionary constraints are involved.

Understanding the genetic basis of adaptation is one of the main enigmas of evolutionary biology. Among vertebrates, hemoglobin has been well documented as a key trait for adaptation to different environments. Here, we investigate the role of hemoglobins in adaptation to ocean depth in the diverse teleost order Gadiformes, with species distributed at a wide range of depths varying in temperature, hydrostatic pressure and oxygen levels. Using genomic data we characterized the full hemoglobin (Hb) gene repertoire for subset of species within this lineage. We discovered a correlation between expanded numbers of Hb genes and ocean depth, with the highest numbers in species occupying shallower, epipelagic regions. Moreover, we demonstrate that the Hb genes have functionally diverged through diversifying selection. Our results suggest that the more variable environment in shallower water has led to selection for a larger Hb gene repertoire and that Hbs have a key role in adaptive processes in marine environments.

Host-intrinsic factors as well as environmental changes are known to be strong evolutionary drivers defining the genetic foundation of immunity. Using a novel set of teleost genomes and a time-calibrated phylogeny, we here investigate the family of Toll-like receptor ( TLR ) genes and address the underlying evolutionary processes shaping the diversity of the first-line defence. Our findings reveal remarkable flexibility within the evolutionary design of teleost innate immunity characterized by prominent TLR gene losses and expansions. In the order of Gadiformes, expansions correlate with the loss of major histocompatibility complex class II ( MHCII ) and diversifying selection analyses support that this has fostered new immunological innovations in TLR s within this lineage. In teleosts overall, TLRs expansions correlate with species latitudinal distributions and maximum depth. By contrast, lineage-specific gene losses overlap with well-described changes in palaeoclimate (global ocean anoxia) and past Atlantic Ocean geography. In conclusion, we suggest that the evolvability of the teleost immune system has most likely played a prominent role in the survival and successful radiation of this lineage.

Morphological allometry refers to patterns of covariance between body parts resulting from variation in body size. Whether measured during growth (ontogenetic allometry), among individuals at similar developmental stage (static allometry), or among populations or species (evolutionary allometry), allometric relationships are often tight and relatively invariant. Consequently, it has been suggested that allometries have low evolvability and could constrain phenotypic evolution by forcing evolving species along fixed trajectories. Alternatively, allometric relationships may result from natural selection for functional optimization. Despite nearly a century of active research, distinguishing between these alternatives remains difficult, partly due to wide differences in the meaning assigned to the term allometry. In particular, a broad use of the term, encompassing any monotonic relationship between body parts, has become common. This usage breaks the connection to the proportional growth regulation that motivated Huxley's original narrow-sense use of allometry to refer to power-law relationships between traits. Focusing on the narrow-sense definition of allometry, we review here evidence for and against the allometry-as-a-constraint hypothesis. Although the low evolvability and the evolutionary invariance of the static allometric slope observed in some studies suggest a possible constraining effect of this parameter on phenotypic evolution, the lack of knowledge about selection on allometry prevents firm conclusions.

Morphological traits often covary within and among species according to simple power laws referred to as allometry. Such allometric relationships may result from common growth regulation, and this has given rise to the hypothesis that allometric exponents may have low evolvability and constrain trait evolution. We formalize hypotheses for how allometry may constrain morphological trait evolution across taxa, and test these using more than 300 empirical estimates of static (within-species) allometric relations of animal morphological traits. Although we find evidence for evolutionary changes in allometric parameters on million-year, cross-species time scales, there is limited evidence for microevolutionary changes in allometric slopes. Accordingly, we find that static allometries often predict evolutionary allometries on the subspecies level, but less so across species. Although there is a large body of work on allometry in a broad sense that includes all kinds of morphological trait-size relationships, we found relatively little information about the evolution of allometry in the narrow sense of a power relationship. Despite the many claims of microevolutionary changes of static allometries in the literature, hardly any of these apply to narrow-sense allometry, and we argue that the hypothesis of strongly constrained static allometric slopes remains viable.

The evolution of threespine sticklebacks in freshwater lakes constitutes a well-studied example of a phenotypic radiation that has produced numerous instances of parallel evolution, but the exact selective agents that drive these changes are not yet fully understood. We present a comparative study across 74 freshwater populations of threespine stickleback in Norway to test whether evolutionary changes in stickleback morphology are consistent with adaptations to physical parameters such as lake depth, lake area, lake perimeter and shoreline complexity, variables thought to reflect different habitats and feeding niches. Only weak indications of adaptation were found. Instead, populations seem to have diversified in phenotypic directions consistent with allometric scaling relationships. This indicates that evolutionary constraints may have played a role in structuring phenotypic variation across freshwater populations of stickleback. We also tested whether the number of lateral plates evolved in response to lake calcium levels, but found no evidence for this hypothesis.

The Late Cretaceous appearance of grasses, followed by the Cenozoic advancement of grasslands as dominant biomes, has contributed to the evolution of a range of specialized herbivores adapted to new diets, as well as to increasingly open and arid habitats. Many mammals including ruminants, the most diversified ungulate suborder, evolved high-crowned (hypsodont) teeth as an adaptation to tooth-wearing diets and habitats. The impact of different causes of tooth wear is still a matter of debate, and the temporal pattern of hypsodonty evolution in relation to the evolution of grasslands remains unclear. We present an improved time-calibrated molecular phylogeny of Cetartiodactyla, with phylogenetic reconstruction of ancestral ruminant diets and habitats, based on characteristics of extant taxa. Using this timeline, as well as the fossil record of grasslands, we conduct phylogenetic comparative analyses showing that hypsodonty in ruminants evolved as an adaptation to both diet and habitat. Our results demonstrate a slow, perhaps constrained, evolution of hypsodonty toward estimated optimal states, excluding the possibility of immediate adaptation. This augments recent findings that slow adaptation is not uncommon on million-year time scales.

Background: Bergmann’s rule proposes that animals in cold habitats will be larger than those in warm habitats. This prediction has been tested thoroughly at the intraspecific level, but few studies have investigated the hypothesis with interspecific data using phylogenetic comparative approaches. Many clades of mammals have representatives in numerous distinct biomes, making this order highly suitable for a large-scale interspecific assessment of Bergmann’s rule. Here, we evaluate Bergmann’s rule within 22 mammalian families—with a dataset that include ~35 % of all described species—using a phylogenetic comparative approach. The method is based on an Ornstein-Uhlenbeck model of evolution that allows for joint estimation of adaptation and constraints (phylogenetic inertia) in the evolution of a trait. We use this comparative method to investigate whether body mass evolves towards phenotypic optima that are functions of median latitude, maximum latitude or temperature. We also assess the closely related Allen’s rule in five families, by testing if relative forelimb length evolves as a function of temperature or latitude. Results: Among 22 mammalian families, there was weak support for Bergmann’s rule in one family: A decrease in temperature predicted increased body mass in Canidae (canids). We also found latitude and temperature to significantly predict body mass in Geomyidae (pocket gophers); however, the association went in the opposite direction of Bergmann’s predictions. Allen’s rule was supported in one of the five examined families (Pteropodidae; megabats), but only when forelimb length evolves towards an optimum that is a function of maximum latitude, not median latitude or temperature. Conclusions: Based on this exhaustive assessment of Bergmann’s rule, we conclude that factors other than latitude and temperature are the major drivers of body mass evolution at the family level in mammals. © The Author(s). 2016. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.

Julian Huxley showed that within-species (static) allometric (power-law) relations can arise from proportional growth regulation with the exponent in the power law equaling the factor of proportionality. Allometric exponents may therefore be hard to change and act as constraints on the independent evolution of traits. In apparent contradiction to this, many empirical studies have concluded that static allometries are evolvable. Many of these studies have been based, however, on a broad definition of allometry that includes any monotonic shape change with size, and do not falsify the hypothesis of constrained narrow-sense allometry. Here, we present the first phylogenetic comparative study of narrow-sense allometric exponents based on a reanalysis of data on eye span and body size in stalk-eyed flies (Diopsidae). Consistent with a role in sexual selection, we found strong evidence that male slopes were tracking "optima" based on sexual dimorphism and relative male trait size. This tracking was slow, however, with estimated times of 2-3 million years for adaptation to exceed ancestral influence on the trait. Our results are therefore consistent with adaptive evolution on million-year time scales, but cannot rule out that static allometry may act as a constraint on eye-span adaptation at shorter time scales.

Summary • Changes in chromosome number as a result of fission and fusion in holocentrics have direct and immediate effects on the recombination rate. We investigate the support for the classic hypothesis that environmental stability selects for increased recombination rates. • We employed a phylogenetic and cytogenetic data set from one of the most diverse angiosperm genera in the world, which has the largest nonpolyploid chromosome radiation (Carex, Cyperaceae; 2n = 12–124; 2100 spp.). We evaluated alternative Ornstein–Uhlenbeck models of chromosome number adaptation to the environment in an information-theoretic framework. • We found moderate support for a positive influence of lateral inflorescence unit size on chromosome number, which may be selected in a stable environment in which resources for reproductive investment are larger. We found weak support for a positive influence on chromosome number of water-saturated soils and among-month temperature constancy, which would be expected to be negatively select for pioneering species. Chromosome number showed a strong phylogenetic signal. • We argue that our finding of small but significant effects of life history and ecology is compatible with our original hypothesis regarding selection of optima in recombination rates: low recombination rate is optimal when inmediate fitness is required. By contrast, high recombination rate is optimal when stable environments allow for evolutionary innovation.

Homoploid hybrid speciation in animals is poorly understood, mainly because of the scarcity of well-documented cases. Here, we present the results of a multilocus sequence analysis on the house sparrow (Passer domesticus), Spanish sparrow (P. hispaniolensis) and their proposed hybrid descendant, the Italian sparrow (P. italiae). The Italian sparrow is shown to be genetically intermediate between the house sparrow and Spanish sparrow, exhibiting genealogical discordance and a mosaic pattern of alleles derived from either of the putative parental species. The average variation on the Z chromosome was significantly reduced compared with autosomal variation in the putative parental species, the house sparrow and Spanish sparrow. Additionally, divergence between the two species was elevated on the Z chromosome relative to the autosomes. This pattern of variation and divergence is consistent with reduced introgression of Z-linked genes and/or a faster-Z effect (increased rate of adaptive divergence on the Z). F(ST) -outlier tests were consistent with the faster-Z hypothesis: two of five Z-linked loci (CHD1Z and PLAA) were identified as candidates for being subject to positive, divergent selection in the putative parental species. Interestingly, the two latter genes showed a mosaic pattern in the (hybrid) Italian sparrow; that is, the Italian sparrow was found to be fixed for Spanish sparrow alleles at CHD1Z and to mainly have house sparrow alleles at PLAA. Preliminary evidence presented in this study thus suggests that sex chromosomes may play a significant role in this case of homoploid hybrid speciation.

The parietal, or third, eye is a photosensory organ situated in the middle of the skull of many lizards. Despite many hypotheses, its exact ecological functions are still unclear. Studies have compared the presence and absence of a functioning parietal eye, although there are no quantitative studies of parietal-eye traits in relation to ecology, physiology or behaviour. In the present study, we report the first comparative study of relative parietal-eye size in relation to climatic and thermophysiological variables. We studied thirty species of Liolaemus, a genus of South-American lizards inhabiting a range of climatic conditions, but found little evidence for adaptation to thermal environment, in that parietal-eye size did not vary meaningfully with latitude, altitude or any measures of environmental temperature. Neither did it relate to thermophysiology; there was a weak relation to thermal tolerance, although this was partially confounded with body size, which explained 23% of the among-species variance after controlling for within-species variation. The negative results obtained could not be explained by phylogenetic constraints because we found no evidence of phylogenetic inertia. We also observed high intraspecific variation indicating that parietal-eye size may not be under strong selection for accuracy. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 870–883.

A new genus, Altihoratosphaga, is erected for species formerly assigned to Horatosphaga Schaum, 1853, and a new species is described. Four species are included in Altihoratosphaga: Altihoratosphaga nomima (Karsch, 1896), Altihoratosphaga montivaga (Sjostedt, 1909), Altihoratosphaga nou (Hemp, 2007) and Altihoratosphaga hanangensis sp. nov. All four species are restricted to Tanzanian localities, and, except for A. nomima, for which no ecological data are available, are confined to montane forest habitats. Data on ecology, acoustics, chromosomes, and molecular relationships are provided, as well as a key to Altihoratosphaga species. The present-day distribution of Altihoratosphaga species suggests former migration events at times when wetter and colder climatic fluctuations favoured connections between montane forest communities, which today are isolated, enabling flightless taxa such as Altihoratosphaga and Monticolaria to spread. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158, 66–82.

Two species currently belong to the genus MonticolariaSjostedt, 1909, both of them restricted to submontane and montane forest, forest edge and clearings. In this study a new Monticolaria species is described which inhabits montane forest edge and forest clearings on Mt. Hanang and the Nou Forest reserve (Manyara escarpment) in north-western Tanzania. The first description of the female of Monticolaria kilimandjarica is also provided and a key for the genus is presented. Segments of the mitochondrial gene cytochrome oxidase subunit I (COI) were sequenced to infer phylogeographic patterns. These analyses confirm the genus as a monophyletic group. The calling song and data on the morphology of the stridulatory file of M. kilimandjarica are presented. The evolutionary history of Monticolaria is discussed in light of biogeographical, ecological and morphological data. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 494–506.

Proxy climate data for the Pacific Ocean show that the most intense El Nino-Southern Oscillation (ENSO) activity during the last millennium coincides with the most recent period of defor- estation on Easter Island. We therefore raise the question as to whether an abrupt change in ENSO variability and its associated impact on the marine resources surrounding Easter Island might have been—for a population already under strain from a number of different pressures—the final stressor causing social collapse. Studies showing how this hypothesized interaction between climate, marine resources and human history on Easter Island could be investigated are discussed.

A multitude of hypotheses claim that abiotic factors are the main drivers of macroevolutionary change. By contrast, Van Valen's Red Queen hypothesis is often put forward as the sole representative of the view that biotic forcing is the main evolutionary driver. This imbalance of hypotheses does not reflect our current knowledge: theoretical work demonstrates the plausibility of biotically driven long-term evolution, whereas empirical work suggests a central role for biotic forcing in macroevolution. We call for a more pluralistic view of how biotic forces may drive long-term evolution that is compatible with both phenotypic stasis in the fossil record and with non-constant extinction rates. Promising avenues of research include contrasting predictions from relevant theories within ecology and macroevolution, as well as embracing both abiotic and biotic proxies while modelling long-term evolutionary data. By fitting models describing hypotheses of biotically driven macroevolution to data, we could dissect their predictions and transcend beyond pattern description, possibly narrowing the divide between our current understanding of micro- and macroevolution.

Biological diversity is, to a large extent, a matter of variation in size. Proportional (isometric) scaling, where large and small individuals are magnified versions of each other, is often assumed to be the most common way morphological traits scale relative to overall size within species. However, the many traits showing nonproportional (allometric) scaling have motivated some of the most discussed hypotheses on scaling relationships in biology, like the positive allometry hypothesis for secondary sexual traits and the negative allometry hypothesis for genitals. I evaluate more than 3,200 allometric parameters from the literature and find that negative allometry, not isometry, is the expected scaling relationship of morphological traits within species. Slopes of secondary sexual traits are more often steeper compared with other traits, but slopes larger than unity are also common for traits not under sexual selection. The steepness of the allometric slope is accordingly a weak predictor of past and present patterns of selection. Scaling of genitals varies across taxonomic groups, but negative allometry of genitals in insects and spiders is a consistent pattern. Finally, I find indications that terrestrial organisms may have a different scaling of morphological traits overall compared with aquatic species. © 2015 University of Chicago Press

Morphological allometry refers to patterns of covariance between body parts resulting from variation in body size. Whether measured during growth (ontogenetic allometry), among individuals at similar developmental stage (static allometry), or among populations or species (evolutionary allometry), allometric relationships are often tight and relatively invariant. Consequently, it has been suggested that allometries have low evolvability and could constrain phenotypic evolution by forcing evolving species along fixed trajectories. Alternatively, allometric relationships may result from natural selection for functional optimization. Despite nearly a century of active research, distinguishing between these alternatives remains difficult, partly due to wide differences in the meaning assigned to the term allometry. In particular, a broad use of the term, encompassing any monotonic relationship between body parts, has become common. This usage breaks the connection to the proportional growth regulation that motivated Huxley's original narrow-sense use of allometry to refer to power-law relationships between traits. Focusing on the narrow-sense definition of allometry, we review here evidence for and against the allometry-as-a-constraint hypothesis. Although the low evolvability and the evolutionary invariance of the static allometric slope observed in some studies suggest a possible constraining effect of this parameter on phenotypic evolution, the lack of knowledge about selection on allometry prevents firm conclusions.

Morphological traits often covary within and among species according to simple power laws referred to as allometry. Such allometric relationships may result from common growth regulation, and this has given rise to the hypothesis that allometric exponents may have low evolvability and constrain trait evolution. We formalize hypotheses for how allometry may constrain morphological trait evolution across taxa, and test these using more than 300 empirical estimates of static (within-species) allometric relations of animal morphological traits. Although we find evidence for evolutionary changes in allometric parameters on million-year, cross-species time scales, there is limited evidence for microevolutionary changes in allometric slopes. Accordingly, we find that static allometries often predict evolutionary allometries on the subspecies level, but less so across species. Although there is a large body of work on allometry in a broad sense that includes all kinds of morphological trait-size relationships, we found relatively little information about the evolution of allometry in the narrow sense of a power relationship. Despite the many claims of microevolutionary changes of static allometries in the literature, hardly any of these apply to narrow-sense allometry, and we argue that the hypothesis of strongly constrained static allometric slopes remains viable.

Ecological interactions affect the survival and reproduction of individuals. However, ecological interactions are notoriously difficult to measure in extinct populations, hindering our understanding of how the outcomes of interactions such as competition vary in time and influence long-term evolutionary changes. Here, the outcomes of spatial competition in a temporally continuous community over evolutionary timescales are presented for the first time. Our research domain is encrusting cheilostome bryozoans from the Wanganui Basin of New Zealand over a ca 2 Myr time period (Pleistocene to Recent). We find that a subset of species can be identified as consistent winners, and others as consistent losers, in the sense that they win or lose interspecific competitive encounters statistically more often than the null hypothesis of 50%. Most species do not improve or worsen in their competitive abilities through the 2 Myr period, but a minority of species are winners in some intervals and losers in others. We found that conspecifics tend to cluster spatially and interact more often than expected under a null hypothesis: most of these are stand-off interactions where the two colonies involved stopped growing at edges of encounter. Counterintuitively, competitive ability has no bearing on ecological dominance.

Homoploid hybrid speciation in animals is poorly understood, mainly because of the scarcity of well-documented cases. Here, we present the results of a multilocus sequence analysis on the house sparrow (Passer domesticus), Spanish sparrow (P. hispaniolensis) and their proposed hybrid descendant, the Italian sparrow (P. italiae). The Italian sparrow is shown to be genetically intermediate between the house sparrow and Spanish sparrow, exhibiting genealogical discordance and a mosaic pattern of alleles derived from either of the putative parental species. The average variation on the Z chromosome was significantly reduced compared with autosomal variation in the putative parental species, the house sparrow and Spanish sparrow. Additionally, divergence between the two species was elevated on the Z chromosome relative to the autosomes. This pattern of variation and divergence is consistent with reduced introgression of Z-linked genes and/or a faster-Z effect (increased rate of adaptive divergence on the Z). F(ST) -outlier tests were consistent with the faster-Z hypothesis: two of five Z-linked loci (CHD1Z and PLAA) were identified as candidates for being subject to positive, divergent selection in the putative parental species. Interestingly, the two latter genes showed a mosaic pattern in the (hybrid) Italian sparrow; that is, the Italian sparrow was found to be fixed for Spanish sparrow alleles at CHD1Z and to mainly have house sparrow alleles at PLAA. Preliminary evidence presented in this study thus suggests that sex chromosomes may play a significant role in this case of homoploid hybrid speciation.

Many East African mountains are characterized by an exceptionally high biodiversity. Here we assess the hypothesis that climatic fluctuations during the Plio-Pleistocene led to ecological fragmentation with subsequent genetic isolation and speciation in forest habitats in East Africa. Hypotheses on speciation in savannah lineages are also investigated. To do this, mitochondrial DNA sequences from a group of bush crickets consisting of both forest and savannah inhabiting taxa were analysed in relation to Plio-Pleistocene range fragmentations indicated by palaeoclimatic studies. Coalescent modelling and mismatch distributions were used to distinguish between alternative biogeographical scenarios. The results indicate two radiations: the earliest one overlaps in time with the global spread of C4 grasslands and only grassland inhabiting lineages originated in this radiation. Climatically induced retraction of forest to higher altitudes about 0.8 million years ago, promoting vicariant speciation in species inhabiting the montane zone, can explain the second radiation. Although much of the biodiversity in East Africa is presently threatened by climate change, past climatic fluctuations appear to have contributed to the species richness observed in the East African hot spots. Perceiving forests as centres of speciation reinforces the importance of conserving the remaining forest patches in the region.