Niche differentiation, reproductive interference, and range expansion.

Understanding species distributions and predicting future range shifts requires considering all relevant abiotic factors and biotic interactions. Resource competition has received the most attention, but reproductive interference is another widespread biotic interaction that could influence species ranges. Rubyspot damselflies (Hetaerina spp.) exhibit a biogeographic pattern consistent with the hypothesis that reproductive interference has limited range expansion. Here, we use ecological niche models to evaluate whether this pattern could have instead been caused by niche differentiation. We found evidence for climatic niche differentiation, but the species that encounters the least reproductive interference has one of the narrowest and most peripheral niches. These findings strengthen the case that reproductive interference has limited range expansion and also provide a counterexample to the idea that release from negative species interactions triggers niche expansion. We propose that release from reproductive interference enables species to expand in range while specializing on the habitats most suitable for breeding.

Predator-induced prey dispersal can cause hump-shaped density-area relationships in prey populations.

Predation can both reduce prey abundance directly (through density-dependent effects) and indirectly through prey trait-mediated effects. Over the years, many studies have focused on describing the density-area relationship (DAR). However, the mechanisms responsible for the DAR are not well understood. Loss and fragmentation of habitats, owing to human activities, creates landscape-level spatial heterogeneity wherein patches of varying size, isolation and quality are separated by a human-modified "matrix" of varying degrees of hostility and has been a primary driver of species extinctions and declining biodiversity. How matrix hostility in combination with trait-mediated effects influence DAR, minimum patch size, and species coexistence remains an open question. In this paper, we employ a theoretical spatially explicit predator-prey population model built upon the reaction-diffusion framework to explore effects of predator-induced emigration (trait-mediated emigration) and matrix hostility on DAR, minimum patch size, and species coexistence. Our results show that when trait-mediated response strength is sufficiently strong, ranges of patch size emerge where a nonlinear hump-shaped prey DAR is predicted and other ranges where coexistence is not possible. In a conservation perspective, DAR is crucial not only in deciding whether we should have one large habitat patch or several-small (SLOSS), but for understanding the minimum patch size that can support a viable population. Our study lends more credence to the possibility that predators can alter prey DAR through predator-induced prey dispersal.

Ecological release and patch geometry can cause nonlinear density-area relationships.

A primary driver of species extinctions and declining biodiversity is loss and fragmentation of habitats owing to human activities. Many studies spanning a wide diversity of taxa have described the relationship between population density and habitat patch area, i.e., the density-area relationship (DAR), as positive, neutral, negative or some combination of the three. However, the mechanisms responsible for these relationships remain elusive. We employ a theoretical spatially explicit population model built upon the reaction-diffusion framework with absorbing boundary conditions to model a habitat specialist dwelling in islands of habitat surrounded by a hostile matrix. We consider patches with a convex or non-convex geometry. Our results show that a single species following logistic-type population growth exhibits a strictly positive and continuous DAR. However, when multiple asymptotically stable steady states are preset in the system, a discontinuous DAR arises. In the case of two species governed by diffusive Lotka-Volterra growth and competitive interactions, we observe that overall DAR structure can be either (1) positive, (2) positive for small areas and neutral for large, or (3) hump-shaped, i.e., positive for area below a threshold and negative for area above. Patch complexity such as non-convex geometry can cause discontinuities in DAR slope for a single species and create qualitatively different patterns in a competitive system as compared to a convex patch. We also compared our theoretical results with two empirical studies (Anolis lizards on islands and crossbills and pine squirrels in forest fragments) where the pragmatic view of DAR fails to give a mechanistic understanding of what was observed. Close qualitative agreement between theoretical and observed DAR indicates that our model gives a reasonable explanation of the mechanisms underpinning DAR found in those studies. From a conservation perspective, the DAR is crucial to the identification of valuable habitat fragments that favor high abundance and the design of a reserve for a target species. When it comes to protecting a single species, these results suggest that there is unlikely to be a simple solution and that conservation decisions should always be made on a case-by-case basis.

Strength of the 'island rule' in birds is positively associated with absence of avian predators.

The similar characteristics shared by island environments have been shown to lead to common patterns of adaptations in island species, commonly referred to as the 'insularity syndrome'. A well-known example is the 'island rule', where large species become smaller on islands and small species become larger, leading to well-known cases of dwarfism and gigantism. This pattern was recently verified on a global scale, but the mechanisms underlying it have been poorly investigated. Here, we focused on the role of released pressure from predation and competition experienced by island birds. Using 120 pairs of endemic island species and their mainland sister relatives, we first verified that the island rule was detected in our dataset, and then evaluated the effects of the numbers of raptors and interspecific competitors on the evolution of the insular species' body mass. We found a strong effect of predation on body mass evolution, with a stronger island rule for species occurring on islands with no raptors, while the pattern disappears in their presence. However, we did not find an effect of competition on this pattern. Our study shows the importance of considering ecological interactions for understanding patterns of body size evolution, and the exceptions to those patterns.

Latitudinal gradient in species diversity provides high niche opportunities for a range-expanding phytophagous insect.

When species undergo poleward range expansions in response to anthropogenic change, they likely encounter less diverse communities in new locations. If low diversity communities provide weak biotic interactions, such as reduced competition or predation, range-expanding species may experience high niche opportunities. Here, we investigated if oak gall wasp communities follow a latitudinal diversity gradient (LDG) and if lower diversity communities provide weaker interactions at the poles for a range-expanding community member, Neuroterus saltatorius. We performed systematic surveys of gall wasps on a dominant oak, Quercus garryana, throughout most of its range, from northern California to Vancouver Island, British Columbia. On 540 trees at 18 sites, we identified 23 oak gall wasp morphotypes in three guilds (leaf detachable, leaf integral, and stem galls). We performed regressions between oak gall wasp diversity, latitude, and other abiotic (e.g. temperature) and habitat (e.g. oak patch size) factors to reveal if gall wasp communities followed an LDG. To uncover patterns in local interactions, we first performed partial correlations of gall wasp morphotype occurrences on trees within regions). We then performed regressions between abundances of co-occurring gall wasps on trees to reveal if interactions are putatively competitive or antagonistic. Q. garryana-gall wasp communities followed an LDG, with lower diversity at higher latitudes, particularly with a loss of detachable leaf gall morphotypes. Detachable leaf gall wasps, including the range-expanding species, co-occurred most on trees, with weak co-occurrences on trees in the northern expanded region. Abundances of N. saltatorius and detachable and integral leaf galls co-occurring on trees were negatively related, suggesting antagonistic interactions. Overall, we found that LDGs create communities with weaker associations at the poles that might facilitate ecological release in a range-expanding community member. Given the ubiquity of LDGs in nature, poleward range-expanding species are likely moving into low diversity communities. Yet, understanding if latitudinal diversity pattern provides weak biotic interactions for range-expanding species is not well explored. Our large-scale study documenting diversity in a related community of phytophagous insects that co-occur on a host plant reveals that LDGs create high niche opportunities for a range-expanding community member. Biogeographical patterns in diversity and species interactions are likely important mechanisms contributing to altered biotic interactions under range-expansions.

Cuando las especies se expanden hacia los polos en respuesta al cambio antropogénico, es probable que las encuentren comunidades menos diversas en sus nuevos lugares. Si las comunidades de baja diversidad proporcionan interacciones bióticas débiles, como competencia reducida o depredación, las especies que se expanden pueden tener 'oportunidades de alto nicho'. Aquí, investigamos si las comunidades de avispas de las agallas muestran un gradiente de diversidad latitudinal (LDG) y si las comunidades de menor diversidad proporcionan interacciones antagónicas más débiles en los polos para las comunidades que se expanden. Específicamente, investigamos estas relaciones en la comunidad de Neuroterus saltatorius. Realizamos estudios sistemáticos de las avispas de las agallas en un roble dominante, Quercus garryana en la mayor parte de su área de distribución, desde el norte de California hasta la isla de Vancouver, en la Columbia Británica. Sobre 540 árboles en 18 sitios y 6 regiones, nos identificamos 23 mofotipos de avispas de las agallas en tres gremios (hoja desmontable, hoja integral y agallas del tallo). Ejecutamos regresiones entre la diversidad de avispas de las agallas, la latitud, y otros factores abióticos (p. ej., temperatura) y de hábitat (p. ej., tamaño del parche de roble) para revelar si las comunidades de avispas de las agallas siguieron un LDG. Para descubrir tendencias en las interacciones locales, primero ejecutamos correlaciones parciales de ocurrencias de morfotipos de avispas de las agallas en árboles dentro de las regiones. Luego ejecutamos regresiones entre las coocurrencias de las abundancias deavispas de las agallas en los árboles para revelar si las interacciones son supuestamente competitivas o antagónicas. Las comunidades de avispas Q. garryana-gall, siguieron un LDG, con menor diversidad en latitudes más altas, particularmente con una pérdida de morfotipos de agallas de hoja desmontable. Las avispas de las agallas de las hojas desprendibles, incluidas a las especies que se expanden su hábitat, se encontraron en sobre todos los árboles, con coocurrencias débiles en los árboles de la región expandida del norte. Las coocurrencias de las las abundancias de N. saltatorius,las agallas foliares desmontables, e agallas integrales en los árboles se relacionaron negativamente, lo que sugiere interacciones antagónicas. En general, encontramos que los LDG crean comunidades con asociaciones más débiles en los polos que podrían facilitar la liberación ecológica en un miembro de la comunidad que se expanden. Debido de la ubicuidad de los LDG en la naturaleza, es probable que las especies que se expanden hacia los polos se estén trasladando a comunidades de baja diversidad. Sin embargo, no se explora bien la comprensión de si las tendenciasde diversidad latitudinal proporcionan interacciones bióticas débiles para las especies que se expanden. Nuestro estudio degran escala documenta la diversidad en una comunidad relacionada de insectos fitófagos que coexisten en una planta hospedante. Como resultado, revela que los LDG pueden crear oportunidades de grandes nichos para un miembro de la comunidad que se expande. Los patrones biogeográficos en la diversidad y las interacciones entre especies son probablemente mecanismos importantes que contribuyen a las interacciones bióticas alteradas bajo la expansión del hábitat.

Does ecological release from distantly related species affect phenotypic divergence in brook charr?

Ecological opportunity occurs when a resource becomes available through a decrease of interspecific competition and another species colonizes the vacant niche through phenotypic plasticity and intraspecific competition. Brook charr exhibit a resource polymorphism in some Canadian Shield lakes, where a littoral ecotype feeds mainly on zoobenthos and a pelagic ecotype feeds mostly on zooplankton. The objectives of this study were to test that (i) resource polymorphism is common in these brook charr populations, (ii) the presence creek chub and white sucker, two introduced species competing with brook charr for littoral resources, will decrease the phenotypic divergence between the two brook charr ecotypes, and (iii) the ecological release from introduced species will increase population and/or individual niche widths in brook charr. The study was based on 27 lakes and five indicators of resource use (stomach content, liver δ13C, muscle astaxanthin concentration, pyloric caecum length, and gill raker length). Our results indicate that within-lake differences in resource use by both ecotypes are common and stable through time. When facing interspecific competition, both littoral and pelagic brook charr incorporated more pelagic prey into their diet but maintained the amplitude of their differences in resource use, which contradicts our second prediction. Finally, we did not find any significant effect of introduced species on population and individual niche widths of brook charr. We suggest that the difference in feeding mode among distantly related competitors could prevent the complete exclusion of a species from a given niche and explain the lack of response to the ecological release.

Phylogeographic and phenotypic outcomes of brown anole colonization across the Caribbean provide insight into the beginning stages of an adaptive radiation.

Some of the most important insights into the ecological and evolutionary processes of diversification and speciation have come from studies of island adaptive radiations, yet relatively little research has examined how these radiations initiate. We suggest that Anolis sagrei is a candidate for understanding the origins of the Caribbean Anolis adaptive radiation and how a colonizing anole species begins to undergo allopatric diversification, phenotypic divergence and, potentially, speciation. We undertook a genomic and morphological analysis of representative populations across the entire native range of A. sagrei, finding that the species originated in the early Pliocene, with the deepest divergence occurring between western and eastern Cuba. Lineages from these two regions subsequently colonized the northern Caribbean. We find that at the broadest scale, populations colonizing areas with fewer closely related competitors tend to evolve larger body size and more lamellae on their toepads. This trend follows expectations for post-colonization divergence from progenitors and convergence in allopatry, whereby populations freed from competition with close relatives evolve towards common morphological and ecological optima. Taken together, our results show a complex history of ancient and recent Cuban diaspora with populations on competitor-poor islands evolving away from their ancestral Cuban populations regardless of their phylogenetic relationships, thus providing insight into the original diversification of colonist anoles at the beginning of the radiation. Our research also supplies an evolutionary framework for the many studies of this increasingly important species in ecological and evolutionary research.

Patch-scale culls of an overabundant bird defeated by immediate recolonization.

Overabundant native animals cause a variety of human-wildlife conflicts that can require management to reduce their social, environmental, or economic impacts. Culling is an intuitively attractive management response to overabundance, but poor monitoring of results and costs means that evidence for successful outcomes is often lacking. Furthermore, many culls worldwide have been ineffective or counterproductive due to ecological release mechanisms or compensatory responses by the overabundant species. We completed a controlled, replicated, costed, and rigorously monitored experimental cull of the endemic Australian honeyeater, the Noisy Miner (Manorina melanocephala). Aggressive exclusion of birds from remnant woodland patches by overabundant Noisy Miners is listed as a Key Threatening Process under Australian conservation legislation due to its impacts on threatened birds. The problem is particularly prevalent in the highly modified agricultural landscapes of eastern Australia. The species impacts avian assemblages at low densities (0.6-0.8 birds/ha) and at a subcontinental scale (>1 million km2 ). Some ecologists recommend culling as the only management response capable of timely reversal of declines of threatened small woodland birds. We monitored Noisy Miner abundance before and for 12 months after a culling program and found that immediate recolonization from the surrounding landscape negated the impact of the cull. We hypothesize that this is due to a vacuum effect; whereby, birds resident in more marginal habitat around treatment patches move into the vacant territory post-cull. Modeled mean abundance of Noisy Miners declined by 22% in treatment sites compared to an increase of 4% in control sites in the post-cull period. Abundance in all sites, however, remained three to five times higher than published ecological impact thresholds. Return on investment analysis indicated no relationship between culling effort and reduction in Noisy Miner abundance. We conclude that culling at a patch scale is not an efficient method of reducing Noisy Miner abundance to levels unlikely to impact threatened woodland birds in the highly modified study landscape, despite estimated costs 18 times lower than another potential management response of revegetation. Our study highlights the importance of building empirical evidence before intuitively attractive but not necessarily ecologically effective management responses are applied more widely.

Solitary ecology as a phenomenon extending beyond insular systems: exaptive evolution in Anolis lizards.

The mechanisms driving phenotypic evolution have been of interest to biologists since Darwin. Ecological release-wherein adaptive evolution occurs following relaxation of constraining selective pressures-and environmental filtering-wherein exaptive traits allow colonization of a new area-have been studied in several insular cases. Anolis lizards, which may exist in solitude or sympatry with multiple congeners, are an excellent system for evaluating whether ecological release and environmental filtering are associated with phenotypic shifts across phylogenetic and geographical scales. Insular solitary Anolis exhibit phenotypic differentiation in body size and sexual size dimorphism-SSD-through exaptive and adaptive evolution, respectively. But, the generality of these effects has not yet been addressed. Here, we analyse the evolution of body size and SSD relative to sympatry in mainland Anolis. We found that mainland species co-occurring with few congeners exhibit uniform body size and greater SSD relative to other random mainland assemblages, consistent with the insular solitary pattern. The locations of evolutionary shifts for both traits do not coincide with evolutionary transitions to decreased levels of sympatry. These results are consistent with exaptive environmental filtering but not adaptive ecological release. Future studies should be conducted at local scales to evaluate the role of these factors in the evolution of solitary existence in mainland and island species.

Land colonisation by fish is associated with predictable changes in life history.

The colonisation of new environments is a central evolutionary process, yet why species make such transitions often remains unknown because of the difficulty in empirically investigating potential mechanisms. The most likely explanation for transitions to new environments is that doing so conveys survival benefits, either in the form of an ecological release or new ecological opportunity. Life history theory makes explicit predictions about how traits linked to survival and reproduction should change with shifts in age-specific mortality. We used these predictions to examine whether a current colonisation of land by fishes might convey survival benefits. We found that blenny species with more terrestrial lifestyles exhibited faster reproductive development and slower growth rates than species with more marine lifestyles; a life history trade off that is consistent with the hypothesis that mortality has become reduced in younger life stages on land. A plausible explanation for such a shift is that an ecological release or opportunity on land has conveyed survival benefits relative to the ancestral marine environment. More generally, our study illustrates how life history theory can be leveraged in novel ways to formulate testable predictions on why organisms might make transitions into novel environments.

The ubiquity of alpine plant radiations: from the Andes to the Hengduan Mountains.

Alpine plant radiations are compared across the world's major mountain ranges and shown to be overwhelmingly young and fast, largely confined to the Pliocene and Pleistocene, and some of them apparently in the early explosive phase of radiation. Accelerated diversification triggered by island-like ecological opportunities following the final phases of mountain uplift, and in many cases enabled by the key adaptation of perennial habit, provides a general model for alpine plant radiations. Accelerated growth form evolution facilitated by perenniality provides compelling evidence of ecological release and suggests striking parallels between island-like alpine, and especially tropicalpine radiations, and island radiations more generally. These parallels suggest that the world's mountains offer an excellent comparative system for explaining evolutionary radiation.

Ecological release exposes genetically based niche variation.

The evolutionary trajectories of ecological niches have profound impacts on community, population and speciation dynamics, yet the underlying causes of niche lability vs. stasis are poorly understood. Here, we conducted a field experiment to quantify the effects of competition and, conversely, competitive release on the microevolutionary processes driving microhabitat niche evolution in an annual plant population restricted to California vernal pool wetlands. Removing competitors generated a strong increase in mean fitness, the exposure of genetically based niche variation and directional selection for niche evolution in the experimental population. In contrast, genetic variation in the microhabitat niche and directional selection for niche evolution were not detected in individuals growing with competitors. These results indicate that ecological opportunity (here, the removal of competitors) can trigger the immediate expression of latent, heritable niche variation that is necessary for rapid evolutionary responses; conversely, competitors may restrict niche evolution, contributing to niche conservatism in saturated communities.

Crossing the line: increasing body size in a trans-Wallacean lizard radiation (Cyrtodactylus, Gekkota).

The region between the Asian and Australian continental plates (Wallacea) demarcates the transition between two differentiated regional biotas. Despite this striking pattern, some terrestrial lineages have successfully traversed the marine barriers of Wallacea and subsequently diversified in newly colonized regions. The hypothesis that these dispersals between biogeographic realms are correlated with detectable shifts in evolutionary trajectory has however rarely been tested. Here, we analyse the evolution of body size in a widespread and exceptionally diverse group of gekkotan lizards (Cyrtodactylus), and show that a clade that has dispersed eastwards and radiated in the Australopapuan region appears to have significantly expanded its body size 'envelope' and repeatedly evolved gigantism. This pattern suggests that the biotic composition of the proto-Papuan Archipelago provided a permissive environment in which new colonists were released from evolutionary constraints operating to the west of Wallacea.

Mortality dynamics of a polyphagous invasive herbivore reveal clues in its agroecosystem success.

BACKGROUND: The population dynamics of polyphagous pests such as Bemisia argentifolii (B. tabaci MEAM1) are governed by complex, interacting factors involving its cultivated and wild host plants, seasonality, movement and demography. To understand mechanisms contributing to population development and pest success within the agroecosystem, contiguous multi-host field sites were established in three environmentally distinct areas in Arizona. Life tables quantified and partition models described mortality sources and rates for immature insect stages on each host plant. RESULTS: Predation and dislodgement were the largest sources of marginal mortality, supplied the highest irreplaceable mortality and predation was the key factor. Rates of mortality were best predicted, in order, by source, temperature, host plant and season. Marginal mortality was highest for fourth-stage nymphs followed by eggs. Mortality rates were predicted in descending order by stage, temperature and season. Survivorship patterns varied among host plants, and generational mortality averaged 70% on spring cantaloupes but nearly 95% on all other hosts. Population density varied seasonally, persisting at low levels on winter hosts and expanding beginning in the spring; perennial hosts and weeds bridge populations year-round. CONCLUSION: Survival on winter hosts such as broccoli, albeit low, enables population continuity, whereas unusually high survivorship on spring crops like cantaloupe is an ecological release propelling population growth and driving regional dynamics in the summer and fall. This detailed understanding of mortality dynamics provides clues to the success of this invasive pest in our agroecosystems and facilitates opportunities for improved pest management at a broader landscape scale. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.

Effects of species interactions on the potential for evolution at species' range limits.

Species' ranges are limited by both ecological and evolutionary constraints. While there is a growing appreciation that ecological constraints include interactions among species, like competition, we know relatively little about how interactions contribute to evolutionary constraints at species' niche and range limits. Building on concepts from community ecology and evolutionary biology, we review how biotic interactions can influence adaptation at range limits by impeding the demographic conditions that facilitate evolution (which we term a 'demographic pathway to adaptation'), and/or by imposing evolutionary trade-offs with the abiotic environment (a 'trade-offs pathway'). While theory for the former is well-developed, theory for the trade-offs pathway is not, and empirical evidence is scarce for both. Therefore, we develop a model to illustrate how fitness trade-offs along biotic and abiotic gradients could affect the potential for range expansion and niche evolution following ecological release. The model shows that which genotypes are favoured at species' range edges can depend strongly on the biotic context and the nature of fitness trade-offs. Experiments that characterize trade-offs and properly account for biotic context are needed to predict which species will expand their niche or range in response to environmental change. This article is part of the theme issue 'Species' ranges in the face of changing environments (Part II)'.

The Evolution of 'Ecological Release' into the 21st Century.

Ecological release, originally conceived as niche expansion following a reduction in interspecific competition, may prompt invasion success, morphological evolution, speciation, and other ecological and evolutionary outcomes. However, the concept has not been recently reviewed. Here, we trace the study of 'ecological release' from its inception through the present day and find that current definitions are broad and highly varied. Viewing this development as a potential impediment to clear communication and hypothesis testing, we suggest a consensus definition for ecological release: niche expansions and shifts when a constraining interspecific interaction is reduced or removed. In rationalizing this definition, we highlight the various ways ecological release can unfold and address its potential evolutionary consequences.

Niche Differentiation at Multiple Spatial Scales on Large and Small Mediterranean Islands for the Endemic Silene velutina Pourr. ex Loisel. (Caryophyllaceae).

The aim of this work is to investigate niche variations in endemic Silene velutina (Caryophyllaceae, Angiosperms) on Mediterranean islands that differ in size. Six populations on both large and small islands were sampled across the geographic range of the species. For each population, 10 plots (1 × 2 m, with a 25 cm grill) were randomly placed to quantify environmental (abiotic and biotic factors and disturbance) and population (demographic structure and reproductive success) parameters. Niche parameters related to substrate, plant cover, community diversity and composition and disturbance showed significant variation in relation to island size. At the regional scale, we detected a broader niche on large islands associated with spatial heterogeneity and island size. In contrast, at the local scale, populations on small islands showed a broader niche, potentially due to a release from competition (low diversity and plant cover and absence of phanerophytes). Populations on large islands had a demographic structure biased towards vegetative individuals (seedlings and juveniles) with few reproductive individuals, while those on small islands had a majority of adults. Together, the results on niche breadth and demographic structure concord with the idea of a strategy based on adult persistence on small islands.

Shell colour diversification induced by ecological release: A shift in natural selection after a migration event.

Ecological release is often attributed to the rapid adaptive diversification of phenotypic traits. However, it is not well understood how natural selection changes its strength and direction through the process of ecological release. Herein, we demonstrated how shell colour of the Japanese land snail Euhadra peliomphala simodae has diversified via a shift in natural selection due to ecological release after migration from the mainland to an island. This snail's shell colour diversified on the island due to disruptive selection after migration from the mainland. We used trail camera traps to identify the cause of natural selection on both the mainland and the island. We then conducted a mark-recapture experiment while collecting microhabitat use data. In total, we captured and marked around 1,700 snails on the mainland, some of which were preyed upon by an unknown predator. The trail camera traps showed that the predator is the large Japanese field mouse Apodemus speciosus, and the predatory frequency was higher on the mainland than on the island. However, this predation did not correlate with shell colour. Microhabitat use on the island was more extensive than on the mainland, with snails on the island using both ground and arboreal microhabitats. A Bayesian estimation showed that the stabilizing selection on shell colour came from factors other than predation. Our results suggest that the course of natural selection was modified due to ecological release after migration from the mainland, explaining one cause of the phenotypic diversification.

Large-bodied sabre-toothed anchovies reveal unanticipated ecological diversity in early Palaeogene teleosts.

Many modern groups of marine fishes first appear in the fossil record during the early Palaeogene (66-40 Ma), including iconic predatory lineages of spiny-rayed fishes that appear to have originated in response to ecological roles left empty after the Cretaceous/Palaeogene extinction. The hypothesis of extinction-mediated ecological release likewise predicts that other fish groups have adopted novel predatory ecologies. Here, we report remarkable trophic innovation in early Palaeogene clupeiforms (herrings and allies), a group whose modern representatives are generally small-bodied planktivores. Two forms, the early Eocene (Ypresian) †Clupeopsis from Belgium and a new genus from the middle Eocene (Lutetian) of Pakistan, bear conspicuous features indicative of predatory ecology, including large size, long gapes and caniniform dentition. Most remarkable is the presence of a single, massive vomerine fang offset from the midline in both. Numerous features of the neurocranium, suspensorium and branchial skeleton place these taxa on the engraulid (anchovy) stem as the earliest known representatives of the clade. The identification of large-bodied, piscivorous anchovies contributes to an emerging picture of a phylogenetically diverse guild of predatory ray-finned fishes in early Palaeogene marine settings, which include completely extinct lineages alongside members of modern marine groups and taxa that are today restricted to freshwater or deep-sea environments.