Hypotheses in urban ecology: building a common knowledge base.

Urban ecology is a rapidly growing research field that has to keep pace with the pressing need to tackle the sustainability crisis. As an inherently multi-disciplinary field with close ties to practitioners and administrators, research synthesis and knowledge transfer between those different stakeholders is crucial. Knowledge maps can enhance knowledge transfer and provide orientation to researchers as well as practitioners. A promising option for developing such knowledge maps is to create hypothesis networks, which structure existing hypotheses and aggregate them according to topics and research aims. Combining expert knowledge with information from the literature, we here identify 62 research hypotheses used in urban ecology and link them in such a network. Our network clusters hypotheses into four distinct themes: (i) Urban species traits & evolution, (ii) Urban biotic communities, (iii) Urban habitats and (iv) Urban ecosystems. We discuss the potentials and limitations of this approach. All information is openly provided as part of an extendable Wikidata project, and we invite researchers, practitioners and others interested in urban ecology to contribute additional hypotheses, as well as comment and add to the existing ones. The hypothesis network and Wikidata project form a first step towards a knowledge base for urban ecology, which can be expanded and curated to benefit both practitioners and researchers.

Global Protected Areas as refuges for amphibians and reptiles under climate change.

Protected Areas (PAs) are the cornerstone of biodiversity conservation. Here, we collated distributional data for >14,000 (~70% of) species of amphibians and reptiles (herpetofauna) to perform a global assessment of the conservation effectiveness of PAs using species distribution models. Our analyses reveal that >91% of herpetofauna species are currently distributed in PAs, and that this proportion will remain unaltered under future climate change. Indeed, loss of species' distributional ranges will be lower inside PAs than outside them. Therefore, the proportion of effectively protected species is predicted to increase. However, over 7.8% of species currently occur outside PAs, and large spatial conservation gaps remain, mainly across tropical and subtropical moist broadleaf forests, and across non-high-income countries. We also predict that more than 300 amphibian and 500 reptile species may go extinct under climate change over the course of the ongoing century. Our study highlights the importance of PAs in providing herpetofauna with refuge from climate change, and suggests ways to optimize PAs to better conserve biodiversity worldwide.

The Ecologist's Career Compass: A game to explore career paths.

One of the most challenging endeavors for students is choosing a career path that best fits their interests, wills and skills, and setting their professional goals accordingly. Such decisions are often made from within the culture of academia, in which mentors and peers are mainly familiar with the academic job market and lack the knowledge necessary to consult about other types of careers. We aimed to address this gap for ecology and related fields by creating an engaging and effective tool to help students and professionals to familiarize themselves with the diversity of potential career paths available to ecologists. The tool is an applied card game - the Ecologist's Career Compass - which is provided here freely. The game is played as a trump card game and includes 33 cards, each representing a combination of one of four job-market sectors and one of nine types of positions. Each card indicates the level of seven skill categories required to likely be hired and succeed in the focal position at the focal sector, as well as more specific examples for typical jobs in the focal combination. The information in the game largely relies on input from a global survey we conducted among 315 ecologists from 35 countries. While the challenges faced by early-career ecologists in developing their professional path are substantial and diverse, this game can assist in gaining a broad comparative overview of the whole ecology job market and the skills required to likely excel in different paths. We hope this applied game will act as a conversation starter about the diversity of aspirations and opportunities in ecology classrooms and labs.

Urban affinity and its associated traits: A global analysis of bats.

Urbanization is a major contributor to the loss of biodiversity. Its rapid progress is mostly at the expense of natural ecosystems and the species inhabiting them. While some species can adjust quickly and thrive in cities, many others cannot. To support biodiversity conservation and guide management decisions in urban areas, it is important to find robust methods to estimate the urban affinity of species (i.e. their tendency to live in urban areas) and understand how it is associated with their traits. Since previous studies mainly relied on discrete classifications of species' urban affinity, often involving inconsistent assessments or variable parameters, their results were difficult to compare. To address this issue, we developed and evaluated a set of continuous indices that quantify species' urban affinity based on publicly available occurrence data. We investigated the extent to which a species' position along the urban affinity gradient depends on the chosen index and how this choice affects inferences about the relationship between urban affinity and a set of morphological, sensory and functional traits. While these indices are applicable to a wide range of taxonomic groups, we examined their performance using a global set of 356 bat species. As bats vary in sensitivity to anthropogenic disturbances, they provide an interesting case study. We found that different types of indices resulted in different rankings of species on the urban affinity spectrum, but this had little effect on the association of traits with urban affinity. Our results suggest that bat species predisposed to urban life are characterized by low echolocation call frequencies, relatively long call durations, small body size and flexibility in the selection of the roost type. We conclude that simple indices are appropriate and practical, and propose to apply them to more taxa to improve our understanding of how urbanization favours or filters species with particular traits.

Specialist versus Generalist at the Intraspecific Level: Functional Morphology and Substrate Preference of Mediodactylus kotschyi Geckos.

Populations of the same species occupying different microhabitats can either exhibit generalized traits across them or display intraspecific variability, adapting to each microhabitat in order to maximize performance. Intraspecific variability contributes to the generation of diversity, following selection and adaptation, and understanding such variability is important for comprehending how individuals choose their microhabitats. Compared with interspecific variability, however, intraspecific variability in functional morphology and its relationship with microhabitat preference and use have been relatively little studied. Here we examined whether populations of the gecko Mediodactylus kotschyi that differ in the substrates they occupy display habitat-specific behaviors and differing morphologies associated with functional adaptation to their microhabitats. We collected 207 geckos from under or on rocks or on trees from seven populations in Greece. On large islands individuals occupy both substrates; whereas small islets are devoid of trees and the geckos are restricted to rocks, while on the mainland they are only found on trees. We determined gecko substrate preferences in the laboratory, together with their clinging abilities to the different substrates. We measured their limbs, digits, and claws and assessed how these measurements relate to clinging ability. Geckos from all populations preferred the tree made available to them, but this preference was not statistically significant. Geckos from both large and small islands clung better to the tree than to the rock in the laboratory, while those from the mainland clung similarly to both substrates. Geckos collected from trees had longer manual digits and hind limbs. Geckos collected from large and small islands had taller (longer on the dorso-ventral axis; henceforth "deeper") claws. Longer digits and deeper but shorter claws were associated with a better ability to cling to rocks. Our findings suggest that while M. kotschyi is potentially preferentially arboreal, due to the great variation and plasticity it possesses, it can successfully also exploit the habitats available on the smallest, treeless islets in the Aegean Sea. Our study suggests that the dichotomous use of generalist versus specialist in describing species' habitat use is oversimplified, and we suggest the use of a generalist-specialist gradient instead.

Different solutions lead to similar life history traits across the great divides of the amniote tree of life.

Amniote vertebrates share a suite of extra-embryonic membranes that distinguish them from anamniotes. Other than that, however, their reproductive characteristics could not be more different. They differ in basic ectothermic vs endothermic physiology, in that two clades evolved powered flight, and one clade evolved a protective shell. In terms of reproductive strategies, some produce eggs and others give birth to live young, at various degrees of development. Crucially, endotherms provide lengthy parental care, including thermal and food provisioning-whereas ectotherms seldom do. These differences could be expected to manifest themselves in major differences between clades in quantitative reproductive traits. We review the reproductive characteristics, and the distributions of brood sizes, breeding frequencies, offspring sizes and their derivatives (yearly fecundity and biomass production rates) of the four major amniote clades (mammals, birds, turtles and squamates), and several major subclades (birds: Palaeognathae, Galloanserae, Neoaves; mammals: Metatheria and Eutheria). While there are differences between these clades in some of these traits, they generally show similar ranges, distribution shapes and central tendencies across birds, placental mammals and squamates. Marsupials and turtles, however, differ in having smaller offspring, a strategy which subsequently influences other traits.

Multilocus phylogeny and coalescent species delimitation in Kotschy's gecko, Mediodactylus kotschyi: Hidden diversity and cryptic species.

Kotschy's Gecko, Mediodactylus kotschyi, is a small gecko native to southeastern Europe and the Levant. It displays great morphological variation with a large number of morphologically recognized subspecies. However, it has been suggested that it constitutes a species complex of several yet unrecognized species. In this study, we used multilocus sequence data (three mitochondrial and three nuclear gene fragments) to estimate the phylogenetic relationships of 174 specimens from 129 sampling localities, covering a substantial part of the distribution range of the species. Our results revealed high genetic diversity of M. kotschyi populations and contributed to our knowledge about the phylogenetic relationships and the estimation of the divergence times between them. Diversification within M. kotschyi began approximately 15 million years ago (Mya) in the Middle Miocene, whereas the diversification within most of the major clades have been occurred in the last 5 Mya. Species delimitation analysis suggests there exists five species within the complex, and we propose to tentatively recognize the following taxa as full species: M. kotschyi (mainland Balkans, most of Aegean islands, and Italy), M. orientalis (Levant, Cyprus, southern Anatolia, and south-eastern Aegean islands), M. danilewskii (Black Sea region and south-western Anatolia), M. bartoni (Crete), and M. oertzeni (southern Dodecanese Islands). This newly recognized diversity underlines the complex biogeographical history of the Eastern Mediterranean region.

Publisher Correction: The global distribution of tetrapods reveals a need for targeted reptile conservation.

In this Article originally published, owing to a technical error, the author 'Laurent Chirio' was mistakenly designated as a corresponding author in the HTML version, the PDF was correct. This error has now been corrected in the HTML version. Further, in Supplementary Table 3, the authors misspelt the surname of 'Danny Meirte'; this file has now been replaced.

The global distribution of tetrapods reveals a need for targeted reptile conservation.

The distributions of amphibians, birds and mammals have underpinned global and local conservation priorities, and have been fundamental to our understanding of the determinants of global biodiversity. In contrast, the global distributions of reptiles, representing a third of terrestrial vertebrate diversity, have been unavailable. This prevented the incorporation of reptiles into conservation planning and biased our understanding of the underlying processes governing global vertebrate biodiversity. Here, we present and analyse the global distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness patterns of the other three tetrapod classes are good spatial surrogates for species richness of all reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. Hotspots of total and endemic lizard richness overlap very little with those of other taxa. Moreover, existing protected areas, sites of biodiversity significance and global conservation schemes represent birds and mammals better than reptiles. We show that additional conservation actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile knowledge to a global complementarity conservation priority scheme identifies many locations that consequently become important. Notably, investing resources in some of the world's arid, grassland and savannah habitats might be necessary to represent all terrestrial vertebrates efficiently.

Intraspecific competition, not predation, drives lizard tail loss on islands.

Tail autotomy is mainly considered an antipredator mechanism. Theory suggests that predation pressure relaxes on islands, subsequently reducing autotomy rates. Intraspecific aggression, which may also cause tail loss, probably intensifies on islands due to the higher abundance. We studied whether tail autotomy is mostly affected by predation pressure or by intraspecific competition. We further studied whether predator abundance or predator richness is more important in this context. To test our predictions, we examined multiple populations of two gecko species: Kotschy's gecko (Mediodactylus kotschyi; mainland and 41 islands) and the Mediterranean house gecko (Hemidactylus turcicus; mainland and 17 islands), and estimated their abundance together with five indices of predation. In both species, autotomy rates are higher on islands and decline with most predation indices, in contrast with common wisdom, and increase with gecko abundance. In M. kotschyi, tail-loss rates are higher on predator and viper-free islands, but increase with viper abundance. We suggest that autotomy is not simply, or maybe even mainly, an antipredatory mechanism. Rather, such defence mechanisms are a response to complex direct and indirect biotic interactions and perhaps, in the case of tail autotomy in insular populations, chiefly to intraspecific aggression.

Sex determination, longevity, and the birth and death of reptilian species.

Vertebrate sex-determining mechanisms (SDMs) are triggered by the genotype (GSD), by temperature (TSD), or occasionally, by both. The causes and consequences of SDM diversity remain enigmatic. Theory predicts SDM effects on species diversification, and life-span effects on SDM evolutionary turnover. Yet, evidence is conflicting in clades with labile SDMs, such as reptiles. Here, we investigate whether SDM is associated with diversification in turtles and lizards, and whether alterative factors, such as lifespan's effect on transition rates, could explain the relative prevalence of SDMs in turtles and lizards (including and excluding snakes). We assembled a comprehensive dataset of SDM states for squamates and turtles and leveraged large phylogenies for these two groups. We found no evidence that SDMs affect turtle, squamate, or lizard diversification. However, SDM transition rates differ between groups. In lizards TSD-to-GSD surpass GSD-to-TSD transitions, explaining the predominance of GSD lizards in nature. SDM transitions are fewer in turtles and the rates are similar to each other (TSD-to-GSD equals GSD-to-TSD), which, coupled with TSD ancestry, could explain TSD's predominance in turtles. These contrasting patterns can be explained by differences in life history. Namely, our data support the notion that in general, shorter lizard lifespan renders TSD detrimental favoring GSD evolution in squamates, whereas turtle longevity permits TSD retention. Thus, based on the macro-evolutionary evidence we uncovered, we hypothesize that turtles and lizards followed different evolutionary trajectories with respect to SDM, likely mediated by differences in lifespan. Combined, our findings revealed a complex evolutionary interplay between SDMs and life histories that warrants further research that should make use of expanded datasets on unexamined taxa to enable more conclusive analyses.

Erratum: Sex determination, longevity, and the birth and death of reptilian species.

[This corrects the article DOI: 10.1002/ece3.2277.].

Home is where the shell is: predicting turtle home range sizes.

Home range is the area traversed by an animal in its normal activities. The size of home ranges is thought to be tightly linked to body size, through size effect on metabolic requirements. Due to the structure of Eltonian food pyramids, home range sizes of carnivores are expected to exceed those of herbivorous species. The habitat may also affect home range size, with reduced costs of locomotion or lower food abundance in, for example, aquatic habitats selecting for larger home ranges. Furthermore, home range of males in polygamous species may be large due to sexual selection for increased reproductive output. Comparative studies on home range sizes have rarely been conducted on ectotherms. Because ectotherm metabolic rates are much lower than those of endotherms, energetic considerations of metabolic requirements may be less important in determining the home range sizes of the former, and other factors such as differing habitats and sexual selection may have an increased effect. We collected literature data on turtle home range sizes. We used phylogenetic generalized least squares analyses to determine whether body mass, sex, diet, habitat and social structure affect home range size. Turtle home range size increases with body mass. However, body mass explains relatively little of the variation in home range size. Aquatic turtles have larger home ranges than semiaquatic species. Omnivorous turtles have larger home ranges than herbivores and carnivores, but diet is not a strong predictor. Sex and social structure are unrelated to home range size. We conclude that energetic constraints are not the primary factor that determines home range size in turtles, and energetic costs of locomotion in different habitats probably play a major role.

Cope's Rule and the Universal Scaling Law of Ornament Complexity.

Luxuriant, bushy antlers, bizarre crests, and huge, twisting horns and tusks are conventionally understood as products of sexual selection. This view stems from both direct observation and from the empirical finding that the size of these structures grows faster than body size (i.e., ornament size shows positive allometry). We contend that the familiar evolutionary increase in the complexity of ornaments over time in many animal clades is decoupled from ornament size evolution. Increased body size comes with extended growth. Since growth scales to the quarter power of body size, we predicted that ornament complexity should scale according to the quarter power law as well, irrespective of the role of sexual selection in the evolution and function of the ornament. To test this hypothesis, we selected three clades (ammonites, deer, and ceratopsian dinosaurs) whose species bore ornaments that differ in terms of the importance of sexual selection to their evolution. We found that the exponent of the regression of ornament complexity to body size is the same for the three groups and is statistically indistinguishable from 0.25. We suggest that the evolution of ornament complexity is a by-product of Cope's rule. We argue that although sexual selection may control size in most ornaments, it does not influence their shape.

Global effects of land use on local terrestrial biodiversity.

Human activities, especially conversion and degradation of habitats, are causing global biodiversity declines. How local ecological assemblages are responding is less clear--a concern given their importance for many ecosystem functions and services. We analysed a terrestrial assemblage database of unprecedented geographic and taxonomic coverage to quantify local biodiversity responses to land use and related changes. Here we show that in the worst-affected habitats, these pressures reduce within-sample species richness by an average of 76.5%, total abundance by 39.5% and rarefaction-based richness by 40.3%. We estimate that, globally, these pressures have already slightly reduced average within-sample richness (by 13.6%), total abundance (10.7%) and rarefaction-based richness (8.1%), with changes showing marked spatial variation. Rapid further losses are predicted under a business-as-usual land-use scenario; within-sample richness is projected to fall by a further 3.4% globally by 2100, with losses concentrated in biodiverse but economically poor countries. Strong mitigation can deliver much more positive biodiversity changes (up to a 1.9% average increase) that are less strongly related to countries' socioeconomic status.

The number of competitor species is unlinked to sexual dimorphism.

Sexual size dimorphism (SSD) can allow males and females of the same species to specialize on different sized food items and therefore minimize intraspecific competition. Interspecific competition, however, is thought to limit sexual dimorphism, as larger competitors in the community will prevent the larger sex from evolving larger size, and smaller species may prevent the smaller sex from becoming even smaller. We tested this prediction using data on the sexual size dimorphism of lizards, and mammalian carnivores, on islands world-wide. Because insular communities are depauperate, and guilds are species-poor, it is often assumed that enhanced sexual size dimorphism is common on islands. The intensity of interspecific competition, hindering enhanced dimorphism, is thought to increase with competitor richness. We tested whether intraspecific sexual size dimorphism of mammalian carnivores and lizards decreases with increasing island species richness. We further computed the average sexual dimorphism of species on islands and tested whether species-rich islands are inhabited by relatively monomorphic species. Within families and guilds across carnivores and lizards, and with both intraspecific and interspecific approaches, we consistently failed to find support for the notion that species-poor islands harbour more sexually dimorphic individuals or species. We conclude that either interspecific competition does not affect the sexual size dimorphism of insular lizards and carnivores (i.e. character displacement and species sorting are rare in these taxa), or that the number of species in an assemblage or guild is a poor proxy for the intensity of interspecific competition in insular assemblages.