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Showing papers in "Ecography in 2020"


Journal ArticleDOI
TL;DR: This work proposes a standard protocol for reporting SDMs, and introduces a structured format for documenting and communicating the models, ensuring transparency and reproducibility, facilitating peer review and expert evaluation of model quality, as well as meta-analyses.
Abstract: Species distribution models (SDMs) constitute the most common class of models across ecology, evolution and conservation. The advent of ready-to-use software pack - ages and increasing availability of digital geoinformation have considerably assisted the application of SDMs in the past decade, greatly enabling their broader use for informing conservation and management, and for quantifying impacts from global change. However, models must be fit for purpose, with all important aspects of their development and applications properly considered. Despite the widespread use of SDMs, standardisation and documentation of modelling protocols remain limited, which makes it hard to assess whether development steps are appropriate for end use. To address these issues, we propose a standard protocol for reporting SDMs, with an emphasis on describing how a study’s objective is achieved through a series of model - ing decisions. We call this the ODMAP (Overview, Data, Model, Assessment and Prediction) protocol, as its components reflect the main steps involved in building SDMs and other empirically-based biodiversity models. The ODMAP protocol serves two main purposes. First, it provides a checklist for authors, detailing key steps for model building and analyses, and thus represents a quick guide and generic workflow for modern SDMs. Second, it introduces a structured format for documenting and communicating the models, ensuring transparency and reproducibility, facilitating peer review and expert evaluation of model quality, as well as meta-analyses. We detail all elements of ODMAP, and explain how it can be used for different model objectives and applications, and how it complements efforts to store associated metadata and define modelling standards. We illustrate its utility by revisiting nine previously published case studies, and provide an interactive web-based application to facilitate its use. We plan to advance ODMAP by encouraging its further refinement and adoption by the scientific community.

309 citations


Journal ArticleDOI
TL;DR: This study shows no particular benefit to using ensembles over individual tuned models, and suggests that further robust testing of performance is required for situations where models are used to predict to distant places or environments.
Abstract: Predictive performance is important to many applications of species distribution models (SDMs). The SDM ‘ensemble’ approach, which combines predictions across different modelling methods, is believed to improve predictive performance, and is used in many recent SDM studies. Here, we aim to compare the predictive performance of ensemble species distribution models to that of individual models, using a large presence–absence dataset of eucalypt tree species. To test model performance, we divided our dataset into calibration and evaluation folds using two spatial blocking strategies (checkerboard‐pattern and latitudinal slicing). We calibrated and cross‐validated all models within the calibration folds, using both repeated random division of data (a common approach) and spatial blocking. Ensembles were built using the software package ‘biomod2’, with standard (‘untuned’) settings. Boosted regression tree (BRT) models were also fitted to the same data, tuned according to published procedures. We then used evaluation folds to compare ensembles against both their component untuned individual models, and against the BRTs. We used area under the receiver‐operating characteristic curve (AUC) and log‐likelihood for assessing model performance. In all our tests, ensemble models performed well, but not consistently better than their component untuned individual models or tuned BRTs across all tests. Moreover, choosing untuned individual models with best cross‐validation performance also yielded good external performance, with blocked cross‐validation proving better suited for this choice, in this study, than repeated random cross‐validation. The latitudinal slice test was only possible for four species; this showed some individual models, and particularly the tuned one, performing better than ensembles. This study shows no particular benefit to using ensembles over individual tuned models. It also suggests that further robust testing of performance is required for situations where models are used to predict to distant places or environments.

142 citations



Journal ArticleDOI
TL;DR: In this paper, a conservation spatial plan is proposed to minimize extinction risk in the tropics using data on 289 219 species and modeling two future greenhouse gas concentration pathways (RCP2.6 and 8.5) while varying the extent of terrestrial protected land and conserved areas.
Abstract: Limiting climate change to less than 2°C is the focus of international policy under the climate convention (UNFCCC), and is essential to preventing extinctions, a focus of the Convention on Biological Diversity (CBD). The post‐2020 biodiversity framework drafted by the CBD proposes conserving 30% of both land and oceans by 2030. However, the combined impact on extinction risk of species from limiting climate change and increasing the extent of protected and conserved areas has not been assessed. Here we create conservation spatial plans to minimize extinction risk in the tropics using data on 289 219 species and modeling two future greenhouse gas concentration pathways (RCP2.6 and 8.5) while varying the extent of terrestrial protected land and conserved areas from <17% to 50%. We find that limiting climate change to 2°C and conserving 30% of terrestrial area could more than halve aggregate extinction risk compared with uncontrolled climate change and no increase in conserved area.

90 citations



Journal ArticleDOI
TL;DR: In this article, the authors analyzed natural disturbances in boreal and temperate forest ecosystems for the period 2001-2014, aiming to quantify their within-and between-biome variation and compare the climate sensitivity of disturbances across biomes.
Abstract: Disturbance regimes are changing in forests across the world in response to global climate change. Despite the profound impacts of disturbances on ecosystem services and biodiversity, assessments of disturbances at the global scale remain scarce. Here, we analyzed natural disturbances in boreal and temperate forest ecosystems for the period 2001-2014, aiming to 1) quantify their within- and between-biome variation and 2) compare the climate sensitivity of disturbances across biomes. We studied 103 unmanaged forest landscapes with a total land area of 28.2 x 10(6) ha, distributed across five continents. A consistent and comprehensive quantification of disturbances was derived by combining satellite-based disturbance maps with local expert knowledge of disturbance agents. We used Gaussian finite mixture models to identify clusters of landscapes with similar disturbance activity as indicated by the percent forest area disturbed as well as the size, edge density and perimeter-area-ratio of disturbed patches. The climate sensitivity of disturbances was analyzed using Bayesian generalized linear mixed effect models and a globally consistent climate dataset. Within-biome variation in natural disturbances was high in both boreal and temperate biomes, and disturbance patterns did not vary systematically with latitude or biome. The emergent clusters of disturbance activity in the boreal zone were similar to those in the temperate zone, but boreal landscapes were more likely to experience high disturbance activity than their temperate counterparts. Across both biomes high disturbance activity was particularly associated with wildfire, and was consistently linked to years with warmer and drier than average conditions. Natural disturbances are a key driver of variability in boreal and temperate forest ecosystems, with high similarity in the disturbance patterns between both biomes. The universally high climate sensitivity of disturbances across boreal and temperate ecosystems indicates that future climate change could substantially increase disturbance activity.

79 citations


Journal ArticleDOI
TL;DR: NicheMapR as discussed by the authors is an R package that includes a suite of programs for the mechanistic modelling of heat, water, energy and mass exchange between any kind of ectothermic organism and its environment.
Abstract: Mechanistic niche models characterise the fundamental niche of an organism by determining thermodynamic constraints on its heat, water and nutritional budget, and the consequences of this for growth, development and reproduction. They can thus quantify constraints on survival, activity and, ultimately, the vital rates that determine population growth, given a sequence of environmental conditions and the key morphological, physiological and behavioural functional traits. Here we introduce and document the ectotherm model of NicheMapR, an R package that includes a suite of programs for the mechanistic modelling of heat, water, energy and mass exchange between any kind of ectothermic organism and its environment. The NicheMapR ectotherm model is based on a Fortran program originally developed by Porter, Mitchell and Beckman for predicting core body temperature and evaporative water loss as a function of microclimatic conditions and behavioural thermoregulation. The model includes routines for computing steady state body temperature and evaporative water loss given two extreme microclimates (minimum and maximum shade) as computed by the NicheMapR microclimate model. Behavioural options include posture and colour change, shade-seeking, panting, climbing and retreating underground. Here we configure the program to be called from R as part of the NicheMapR package and describe the model in detail including new functionality for modelling whole life-cycle energy and water budgets using Dynamic Energy Budget theory. We include scripts for core operation of the ectotherm model as well as stand-alone R scripts for running the DEB model. Example applications are provided in the paper and in the associated vignettes. The integrated microclimate and ectotherm models should provide a strong thermodynamic basis for determining the effects of environmental change on the behaviour, distribution and abundance of ectothermic organisms.

78 citations


Journal ArticleDOI
TL;DR: In this paper, the authors used occurrence information from eBird to measure the survey completeness of the world's birds in this database at three temporal resolutions and four spatial resolutions across the annual cycle during the period 2002 to 2018.
Abstract: Measuring the completeness of survey inventories created by citizen‐science initiatives can identify the strengths and shortfalls in our knowledge of where species occur geographically. Here, we use occurrence information from eBird to measure the survey completeness of the world's birds in this database at three temporal resolutions and four spatial resolutions across the annual cycle during the period 2002 to 2018. Approximately 84% of the earth's terrestrial surface contained bird occurrence information with the greatest concentrations occurring in North America, Europe, India, Australia and New Zealand. The largest regions with low levels of survey completeness were located in central South America, northern and central Africa, and northern Asia. Across spatial and temporal resolutions, survey completeness in regions with occurrence information was 55–74% on average, with the highest values occurring at coarser temporal and coarser spatial resolutions and during spring migration within temperate and boreal regions. Across spatial and temporal resolutions, survey completeness exceeded 90% within ca 4–14% of the earth's terrestrial surface. Survey completeness increased globally from 2002 to 2018 across all months of the year at a rate of ca 3% yr–¹. The slowest gains occurred in Africa and in montane regions, and the most rapid gains occurred in India and in tropical forests after 2012. Thus, occurrence information from a global citizen‐science program for a charismatic and well‐studied taxon was geographically broad but contained heterogeneous patterns of survey completeness that were strongly influenced by temporal and especially spatial resolution. Our results identify regions where the application of additional effort would address current knowledge shortfalls, and regions where the maintenance of existing effort would benefit long‐term monitoring efforts. Our findings highlight the potential of citizen science initiatives to further our knowledge of where species occur across space and time, information whose applications under global change will likely increase.

67 citations


Journal ArticleDOI
TL;DR: In this article, the authors analyzed scientific survey data using a three-tiered analytical approach to provide an overview of changes in distribution for 19 northeast Atlantic fish species encompassing 73 commercial stocks over 30 years.
Abstract: Changes in fish distribution are being observed across the globe. In Europe's Common Fisheries Policy, the share of the catch of each fish stock is split among management areas using a fixed allocation key known as ‘Relative Stability’: in each management area, member states get the same proportion of the total catch each year. That proportion is largely based on catches made by those member states in the 1970s. Changes in distribution can, therefore, result in a mismatch between quota shares and regional abundances within management areas, with potential repercussions for the status of fish stocks and the fisheries that depend on them. Assessing distribution changes is crucial to ensure adequate management and sustainable exploitation of our fish resources. We analysed scientific survey data using a three‐tiered analytical approach to provide, for the first time, an overview of changes in distribution for 19 northeast Atlantic fish species encompassing 73 commercial stocks over 30 yr. All species have experienced changes in distribution, five of which did so across management areas. A cross‐species analysis suggested that shifts in areas of suitable thermal habitat, and density‐dependent use of these areas, are at least partly responsible for the observed changes. These findings challenge the current use of relative stability to allocate quotas.

62 citations


Journal ArticleDOI
TL;DR: All species were screened for relevance of physiological data on each species’ response to hypoxia or high elevations and the physiology of the lungs of high-altitude ducks and geese was analyzed.
Abstract: s were screened for relevance of physiological data on each species’ response to hypoxia or high elevations. Data base Biosis Citation Index, Zoological Record, Medline Terms included in Boolean search TS = ((\"bar-headed goose\" OR \"anser indicus\") And (\"highaltitude\" OR \"hypox*\" OR \"high elevation\")) TS = ((\"canada goose\" OR \"branta canadensis\") And (\"highaltitude\" OR \"hypox*\" OR \"high elevation\")) TS = ((\"evening grosbeak\" OR \"Coccothraustes vespertinus\") And (\"highaltitude\" OR \"hypox*\" OR \"high elevation\")) 0 Selection Criteria 1. Must be a published article. No meeting abstracts or presentations. 2. Must include the species in question as one of the primary species being reported on. Reporting on another species and simple comparisons (e.g. hemoglobin structure of a species not of interest compared to species of interest) do not meet the requirements. 3. Be associated with high elevation/altitude or hypoxia research. Bar-headed Goose (Anser indicus) Systematic Search Results: Bakkeren, C. et al. 2020. A morphometric analysis of the lungs of high-altitude ducks and geese. J Anat in press. Bishop, C. M. et al. 2015. The roller coaster flight strategy of bar-headed geese conserves energy during Himalayan migrations. Sci New York N Y 347: 250–4. Black, C. P. and Tenney, S. M. 1980a. Oxygen transport during progressive hypoxia in highaltitude and sea-level waterfowl. Resp Physiol 39: 217–239. Black, C. P. and Tenney, S. M. 1980b. Pulmonary hemodynamic responses to acute and chronic hypoxia in two waterfowl species. Comp Biochem Physiology Part Physiology 67: 291–293. Black, C. P. et al. 1978. Oxygen transport during progressive hypoxia in bar-headed geese (Anser indicus) acclimatized to sea level and 5600 meters. Respiratory function in birds, adult and embryonic. -Springer, Berlin, Heidelberg: 79–83. Butler, P. J. 2010. High fliers: The physiology of bar-headed geese. Comp Biochem Physiology Part Mol Integr Physiology 156: 325–329. Butler, P. J. 2016. The physiological basis of bird flight. Philosophical Transactions Royal Soc Lond Ser B Biological Sci 371: 20150384. Faraci, F. M. and Fedde, M. R. 1986. Regional circulatory responses to hypocapnia and hypercapnia in bar-headed geese. Am J Physiology-regulatory Integr Comp Physiology 250: R499–R504. Faraci, F. M. et al. 1984a. Attenuated pulmonary pressor response to hypoxia in bar-headed geese. Am J Physiology-regulatory Integr Comp Physiology 247: R402–R403. Faraci, F. M. et al. 1984b. Oxygen delivery to the heart and brain during hypoxia: Pekin duck vs. bar-headed goose. Am J Physiology-regulatory Integr Comp Physiology 247: R69–R75. Faraci, F. M. et al. 1985. Blood flow distribution during hypocapnic hypoxia in pekin ducks and bar-headed geese. Resp Physiol 61: 21–30. Fedde, M. R. 1987. Wonders of the bar-headed goose. Vigorous exercise in a low-oxygen environment. Explorer 29: 31–34. Fedde, M. 1990. High-Altitude Bird Flight: Exercise in a Hostile Environment. Physiology 5: 191–193. Fedde, M. R. et al. 1989. Cardiopulmonary function in exercising bar-headed geese during normoxia and hypoxia. Resp Physiol 77: 239–252. Harter, T. S. et al. 2015. Validation of the i-STAT and HemoCue systems for the analysis of blood parameters in the bar-headed goose, Anser indicus. Conserv Physiol 3: cov021. Hawkes, L. A. et al. 2011. The trans-Himalayan flights of bar-headed geese (Anser indicus). Proc National Acad Sci 108: 9516–9519. Hawkes, L. A. et al. 2012. The paradox of extreme high-altitude migration in bar-headed geese Anser indicus. Proc Biological Sci Royal Soc 280: 20122114. Hawkes, L. A. et al. 2014. Maximum Running Speed of Captive Bar-Headed Geese Is Unaffected by Severe Hypoxia. Plos One 9: e94015. Hawkes, L. A. et al. 2017. Do Bar-Headed Geese Train for High Altitude Flights? Integr Comp Biol 57: 240–251. Hiebl, I. and Braunitzer, G. 1988. Adaption of the hemoglobins of Bar-headed goose (Anser indicus), Andean goose (Chloephaga melanoptera) and Ruppells griffon (Gyps rueppellii) to life under hypoxic conditions. Journal Fur Ornithologie 2: 217–226. Hiebl, I. et al. 1986. High-altitude respiration of birds. The primary structures of the alpha Dchains of the Bar-headed Goose (Anser indicus), the Greylag Goose(Anser anser) and the Canada Goose (Branta canadensis). Biol Chem H-s 367: 591–9. Jendroszek, A. et al. 2018. Allosteric mechanisms underlying the adaptive increase in hemoglobin-oxygen affinity of the bar-headed goose. J Exp Biology 221: jeb185470. Knight, K. 2016. High-altitude bar-headed geese outperform Vancouver cousins. J Exp Biology 219: 1933.2-1934. Kumar, S. et al. 2010. A brief note on bar-headed geese fitted with satellite transmitters. Telemetry in Wildlife Science, ENVIS Bulletin: 131–134. Laguë, S. L. 2017. High-altitude champions: birds that live and migrate at altitude. J Appl Physiol 123: 942–950. Lague, S. L. et al. 2016. Altitude matters: differences in cardiovascular and respiratory responses to hypoxia in bar-headed geese reared at high and low altitudes. J Exp Biology 219: 1974– 84. Lague, S. L. et al. 2017. Divergent respiratory and cardiovascular responses to hypoxia in barheaded geese and Andean birds. J Exp Biology 220: 4186–4194. Lee, S. Y. et al. 2008. Have wing morphology or flight kinematics evolved for extreme high altitude migration in the bar-headed goose? Comp Biochem Physiology Toxicol Pharmacol Cbp 148: 324–31. Liang, Y. et al. 2001. The crystal structure of bar-headed goose hemoglobin in deoxy form: the allosteric mechanism of a hemoglobin species with high oxygen affinity. J Mol Biol 313: 123–37. Liu, X.-Z. et al. 2001. Avian haemoglobins and structural basis of high affinity for oxygen: structure of bar-headed goose aquomet haemoglobin. Acta Crystallogr Sect D Biological Crystallogr 57: 775–783. Llanos, A. J. et al. 2011. Counterpoint: high altitude is not for the birds! J Appl Physiology Bethesda Md 1985 111: 1515–8. McCracken, K. G. et al. 2010. Phylogenetic and structural analysis of the HbA (alphaA/betaA) and HbD (alphaD/betaA) hemoglobin genes in two high-altitude waterfowl from the Himalayas and the Andes: Bar-headed goose (Anser indicus) and Andean goose (Chloephaga melanoptera). Mol Phylogenet Evol 56: 649–58. Meir, J. U. and Milsom, W. K. 2013. High thermal sensitivity of blood enhances oxygen delivery in the high-flying bar-headed goose. J Exp Biol 216: 2172–2175. Meir, J. et al. 2019. Reduced metabolism supports hypoxic flight in the high-flying bar-headed goose (Anser indicus). eLife in press. Milsom, W. K. 2011. Cardiorespiratory support of avian flight. J Exp Biol 214: 4071–4072. Milsom, W. K. 2017. Different solutions to restoring oxygen delivery at altitude. Acta Physiol 222: e12926. Mu, C.-Y. et al. 2016. The complete mitochondrial genome of Anser indicus (Aves, Anseriformes, Anatidae ). Mitochondrial Dna Part 27: 4588–4589. Natarajan, C. et al. 2018. Molecular basis of hemoglobin adaptation in the high-flying barheaded goose. Plos Genet 14: e1007331. Nice, P. V. et al. 1980. A comparative study of ventilatory responses to hypoxia with reference to hemoglobin o2-affinity in llama, cat, rat, duck and goose. Comp Biochem Physiology Part Physiology 66: 347–350. Parr, N. et al. 2019. Tackling the Tibetan Plateau in a down suit: insights into thermoregulation by bar-headed geese during migration. J Exp Biology 222: jeb203695. Petschow, D. et al. 1977. Causes of high blood O2 affinity of animals living at high altitude. J Appl Physiol 42: 139–143. Prins, H. and Namgail, T. 2017. Bird migration across the Himalayas: wetland functioning amidst mountains and glaciers. Saunders, D. K. and Fedde, M. R. 1991. Physical conditioning: Effect on the myoglobin concentration in skeletal and cardiac muscle of bar-headed geese. Comp Biochem Physiology Part Physiology 100: 349–352. Scott, G. R. and Milsom, W. K. 2007. Control of breathing and adaptation to high altitude in the bar-headed goose. Am J Physiology-regulatory Integr Comp Physiology 293: R379–R391. Scott, G. R. et al. 2008. Body temperature depression and peripheral heat loss accompany the metabolic and ventilatory responses to hypoxia in low and high altitude birds. J Exp Biol 211: 1326–1335. Scott, G. R. et al. 2009a. Control of respiration in flight muscle from the high-altitude bar-headed goose and low-altitude birds. Am J Physiology-regulatory Integr Comp Physiology 297: R1066–R1074. Scott, G. R. et al. 2009b. Evolution of muscle phenotype for extreme high altitude flight in the bar-headed goose. Proc Royal Soc B Biological Sci 276: rspb20090947 3653. Scott, G. R. et al. 2011a. Molecular evolution of cytochrome C oxidase underlies high-altitude adaptation in the bar-headed goose. Mol Biol Evol 28: 351 363. Scott, G. R. et al. 2011b. Point: high altitude is for the birds! J Appl Physiology Bethesda Md 1985 111: 1514–5. Scott, G. R. et al. 2015. How Bar-Headed Geese Fly Over the Himalayas. Physiology 30: 107– 115. Snyder, G. K. et al. 1982. Development and Metabolism during Hypoxia in Embryos of High Altitude Anser indicus versus Sea Level Branta canadensis Geese. Physiol Zool 55: 113– 123. Snyder, G. K. et al. 1984. Effects of hypoxia on tissue capillarity in geese. Resp Physiol 58: 151–160. Spivey, R. J. and Bishop, C. M. 2014. An implantable instrument for studying the long-term flight biology of migratory birds. Rev Sci Instruments 85: 014301. Storz, J. F. et al. 2010. Phenotypic plasticity and genetic adaptation to high-altitude hypoxia in vertebrates. J Exp Biology 213: 4125 4136. Sunnucks, P. et al. 2017. Integrative Approaches for Studying Mitochondrial and Nuclear Genome Co-evolution in Oxidative Phosphorylation. Frontiers Genetics 8: 25. Wang, H.-C. et al. 2000. Crystallization and preliminary crystallographic studies of bar-headed goose fluoromethaemoglobin with inositol hexaphosphate. Acta Crystallogr Sect D Biological Crystallogr 56: 1183–1184. Wang, W. et al. 2020. First de novo whole genome sequencing and assembly of the bar-headed goose. Pee

59 citations




Journal ArticleDOI
TL;DR: Keitt et al. as discussed by the authors published an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Abstract: 1 –––––––––––––––––––––––––––––––––––––––– © 2020 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Subject Editor: Timothy Keitt Editor-in-Chief: Robert Holt Accepted 26 December 2019 43: 1–12, 2020 doi: 10.1111/ecog.04962 43 1–12

Journal ArticleDOI
TL;DR: This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Abstract: 60 –––––––––––––––––––––––––––––––––––––––– © 2019 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Subject Editor: Christine Meynard Editor-in-Chief: Miguel Araújo Accepted 11 September 2019 43: 60–74, 2020 doi: 10.1111/ecog.04630 doi: 10.1111/ecog.04630 43 60–74



Journal ArticleDOI
TL;DR: In this article, the Marie Curie Clarin COFUND program of the Principality of Asturias (ACB17•26) and the regional grant IDI/2018/000151 for research groups.
Abstract: BJ‐A thanks the Marie Curie Clarin‐COFUND program of the Principality of Asturias‐EU (ACB17‐26) and the regional grant IDI/2018/000151 for research groups.

Journal ArticleDOI
TL;DR: Mönkkönen et al. as mentioned in this paper published an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Abstract: 1 –––––––––––––––––––––––––––––––––––––––– © 2020 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Subject Editor: Mikko Mönkkönen Editor-in-Chief: Miguel Araújo Accepted 17 March 2020 43: 1–13, 2020 doi: 10.1111/ecog.05053 43 1–13

Journal ArticleDOI
TL;DR: In this paper, the authors used a series of simulations to explore how well models can discriminate between variables with different influence and calibrate the magnitude of influence relative to an omniscient model.
Abstract: Models of species’ distributions and niches are frequently used to infer the importance of range‐ and niche‐defining variables. However, the degree to which these models can reliably identify important variables and quantify their influence remains unknown. Here we use a series of simulations to explore how well models can 1) discriminate between variables with different influence and 2) calibrate the magnitude of influence relative to an ‘omniscient’ model. To quantify variable importance, we trained generalized additive models (GAMs), Maxent and boosted regression trees (BRTs) on simulated data and tested their sensitivity to permutations in each predictor. Importance was inferred by calculating the correlation between permuted and unpermuted predictions, and by comparing predictive accuracy of permuted and unpermuted predictions using AUC and the continuous Boyce index. In scenarios with one influential and one uninfluential variable, models failed to discriminate reliably between variables when training occurrences were 0.5, spatial extent was small, environmental data had coarse resolution and spatial autocorrelation was low, or when pairwise correlation between environmental variables was |r| > 0.7. When two variables influenced the distribution equally, importance was underestimated when species had narrow or intermediate niche breadth. Interactions between variables in how they shaped the niche did not affect inferences about their importance. When variables acted unequally, the effect of the stronger variable was overestimated. GAMs and Maxent discriminated between variables more reliably than BRTs, but no algorithm was consistently well‐calibrated vis‐a‐vis the omniscient model. Algorithm‐specific measures of importance like Maxent's change‐in‐gain metric were less robust than the permutation test. Overall, high predictive accuracy did not connote robust inferential capacity. As a result, requirements for reliably measuring variable importance are likely more stringent than for creating models with high predictive accuracy.

Journal ArticleDOI
TL;DR: These results represent the first model‐based monarch distributional estimates for the Mexican migration route and should provide foundations for future conservation work and demonstrates the potential benefits of using SDM‐derived richness estimates and phenological information for biotic factors affecting species distributions.
Abstract: Although long‐standing theory suggests that biotic variables are only relevant at local scales for explaining the patterns of species' distributions, recent studies have demonstrated improvements to species distribution models (SDMs) by incorporating predictor variables informed by biotic interactions. However, some key methodological questions remain, such as which kinds of interactions are permitted to include in these models, how to incorporate the effects of multiple interacting species, and how to account for interactions that may have a temporal dependence. We addressed these questions in an effort to model the distribution of the monarch butterfly Danaus plexippus during its fall migration (September–November) through Mexico, a region with new monitoring data and uncertain range limits even for this well‐studied insect. We estimated species richness of selected nectar plants (Asclepias spp.) and roosting trees (various highland species) for use as biotic variables in our models. To account for flowering phenology, we additionally estimated nectar plant richness of flowering species per month. We evaluated three types of models: climatic variables only (abiotic), plant richness estimates only (biotic) and combined (abiotic and biotic). We selected models with AICc and additionally determined if they performed better than random on spatially withheld data. We found that the combined models accounting for phenology performed best for all three months, and better than random for discriminatory ability but not omission rate. These combined models also produced the most ecologically realistic spatial patterns, but the modeled response for nectar plant richness matched ecological predictions for November only. These results represent the first model‐based monarch distributional estimates for the Mexican migration route and should provide foundations for future conservation work. More generally, the study demonstrates the potential benefits of using SDM‐derived richness estimates and phenological information for biotic factors affecting species distributions.


Journal ArticleDOI
TL;DR: In this article, the authors analyzed the relationship between the nestedness of the scavenger assemblages and climatic variables (i.e., temperature, precipitation, temperature variability and precipitation variability), ecosystem productivity and biomass (e.g., NDVI) using 53 study sites in 22 countries across five continents.
Abstract: The organization of ecological assemblages has important implications for ecosystem functioning, but little is known about how scavenger communities organize at the global scale. Here, we test four hypotheses on the factors affecting the network structure of terrestrial vertebrate scavenger assemblages and its implications on ecosystem functioning. We expect scavenger assemblages to be more nested (i.e. structured): 1) in species-rich and productive regions, as nestedness has been linked to high competition for carrion resources, and 2) regions with low human impact, because the most efficient carrion consumers that promote nestedness are large vertebrate scavengers, which are especially sensitive to human persecution. 3) We also expect climatic conditions to affect assemblage structure, because some scavenger assemblages have been shown to be more nested in colder months. Finally, 4) we expect more organized assemblages to be more efficient in the consumption of the resource. We first analyzed the relationship between the nestedness of the scavenger assemblages and climatic variables (i.e. temperature, precipitation, temperature variability and precipitation variability), ecosystem productivity and biomass (i.e. NDVI) and degree of human impact (i.e. human footprint) using 53 study sites in 22 countries across five continents. Then, we related structure (i.e. nestedness) with its function (i.e. carrion consumption rate). We found a more nested structure for scavenger assemblages in regions with higher NDVI values and lower human footprint. Moreover, more organized assemblages were more efficient in the consumption of carrion. However, our results did not support the prediction that the structure of the scavenger assemblages is directly related to climate. Our findings suggest that the nested structure of vertebrate scavenger assemblages affects its functionality and is driven by anthropogenic disturbance and ecosystem productivity worldwide. Disarray of scavenger assemblage structure by anthropogenic disturbance may lead to decreases in functionality of the terrestrial ecosystems via loss of key species and trophic facilitation processes.

Journal ArticleDOI
TL;DR: Comparing a diversity of climatic data for the studied populations showed that while CDL is under a tight photoperiodic regulation linked with latitude, its length depends also on climatic factors determining the growing season length.
Abstract: Reproductive diapause is a primary mechanism used by arthropods to synchronize their life cycle with seasonal changes in temperate regions. Our study species, Drosophila montana, represents the northern insect species where flies enter reproductive diapause under short day conditions and where the precise timing of diapause is crucial for both survival and offspring production. We have studied clinal variation in the critical day length for female diapause induction (CDL) and their overall susceptibility to enter diapause (diapause incidence), as well as the temperature sensitivity of these traits. The study was performed using multiple strains from four latitudinal clines of the species – short clines in Finland and Alaska and long clines in the Rocky Mountains and the western coast of North America – and from one population in Kamchatka, Russia. CDL showed strong latitudinal clines on both continents, decreasing by one hour per five degrees decline in latitude, on average. CDL also decreased in all populations along with an increase in fly rearing temperature postponing the diapause to later calendar time, the effects of temperature being stronger in southern than in northern population. Female diapause incidence was close to 100% under short day/low temperature conditions in all populations, but decreased below 50% even under short days in 19°C in the southern North American western coast populations and in 22°C in most populations. Comparing a diversity of climatic data for the studied populations showed that while CDL is under a tight photoperiodic regulation linked with latitude, its length depends also on climatic factors determining the growing season length. Overall, the study deepens our understanding of how spatial and environmental parameters affect the seasonal timing of an important biological event, reproductive diapause and helps to estimate the evolutionary potential of insect populations to survive in changing climatic conditions.

Journal ArticleDOI
TL;DR: In this paper, the authors used an extensive database of fuel hazard assessments to determine the extent to which environmental variables (climatic conditions and soil type) and disturbance (fire) can predict fuel hazard in native vegetation across south-eastern Australia.
Abstract: © 2019 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos Understanding spatial variation in wildland fuel is central to predicting wildfire behaviour as well as current and future fire regimes. Vegetation (plant material) – both live (biomass) and dead (necromass) – constitutes most aspects of wildland fuel (hereafter ‘fuel’). It therefore is likely that factors influencing vegetation structure and composition – climate, soils, disturbance – also will influence fuel structure and associated hazard. Nonetheless, these relationships are poorly understood in temperate environments. In this study, we used an extensive database of fuel hazard assessments to determine the extent to which environmental variables (climatic conditions and soil type) and disturbance (fire) can predict fuel hazard in native vegetation across south-eastern Australia. Fuel hazard ratings are based on the horizontal and vertical continuity of fine fuels (dead plant material < 6 mm thick, and live plant material < 3 mm thick) that burn in the flaming front of a fire. These scores are used widely by fire managers in Australia. We used environmental and disturbance variables to develop models to predict spatial patterns of hazard for each fuel stratum (surface, near-surface, elevated and bark) and the height of two fuel strata (near-surface, elevated). Soil, climate and time since fire were strong predictors of fuel hazard for at least one stratum, and soil predictors were the strongest predictors of fuel hazard across all strata. We used models to predict fuel hazard by stratum at a fixed time since fire in two regions with contrasting environments in south-eastern Australia to better understand the spatial arrangement of fuel hazard. Fuel hazard varied within and between regions, emphasising the complexity and heterogeneity of fuel patterns that affect fuel hazard from local to landscape extents. The models improve the basis for analysing fuel hazard patterns and therefore increase the capacity to predict fire regimes under future climates.

Journal ArticleDOI
TL;DR: Agarwal et al. as discussed by the authors published an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Abstract: 1090 –––––––––––––––––––––––––––––––––––––––– © 2020 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Subject Editor and Editor-in-Chief: Miguel Araújo Accepted 18 February 2020 43: 1090–1106, 2020 doi: 10.1111/ecog.04423 43 1090–1106


Journal ArticleDOI
TL;DR: In this paper, the authors examined the importance of climate and soil variables to explaining above-ground biomass distribution across the Australian continent using publicly available biomass data from 3130 mature forest sites, in 6 broad ecoregions, encompassing tropical, subtropical and temperate biomes.
Abstract: Above-ground biomass in forests is critical to the global carbon cycle as it stores and sequesters carbon from the atmosphere. Climate change will disrupt the carbon cycle hence understanding how climate and other abiotic variables determine forest biomass at broad spatial scales is important for validating and constraining Earth System models and predicting the impacts of climate change on forest carbon stores. We examined the importance of climate and soil variables to explaining above-ground biomass distribution across the Australian continent using publicly available biomass data from 3130 mature forest sites, in 6 broad ecoregions, encompassing tropical, subtropical and temperate biomes. We used the Random Forest algorithm to test the explanatory power of 14 abiotic variables (8 climate, 6 soil) and to identify the best-performing models based on climate-only, soil-only and climate plus soil. The best performing models explained ~50% of the variation (climate-only: R2 = 0.47 ± 0.04, and climate plus soils: R2 = 0.49 ± 0.04). Mean temperature of the driest quarter was the most important climate variable, and bulk density was the most important soil variable. Climate variables were consistently more important than soil variables in combined models, and model predictive performance was not substantively improved by the inclusion of soil variables. This result was also achieved when the analysis was repeated at the ecoregion scale. Predicted forest above-ground biomass ranged from 18 to 1066 Mg ha−1, often under-predicting measured above-ground biomass, which ranged from 7 to 1500 Mg ha−1. This suggested that other non-climate, non-edaphic variables impose a substantial influence on forest above-ground biomass, particularly in the high biomass range. We conclude that climate is a strong predictor of above-ground biomass at broad spatial scales and across large environmental gradients, yet to predict forest above-ground biomass distribution under future climates, other non-climatic factors must also be identified.

Journal ArticleDOI
TL;DR: In this paper, the authors assessed the spatiotemporal patterns of masting across the latitudinal distribution of a widely distributed dryland conifer species, pinon pine Pinus edulis.
Abstract: Masting, or the synchronous and irregular production of seed crops, is controlled by environmental conditions and resource budgets. Increasing temperatures and shifting precipitation regimes may alter the frequency and magnitude of masting, especially in species that experience chronic resource stress. Yet the effects of a changing climate on seed production are unlikely to be uniform across populations, particularly those that span broad abiotic gradients. In this study, we assessed the spatiotemporal patterns of masting across the latitudinal distribution of a widely distributed dryland conifer species, pinon pine Pinus edulis. We quantified seed cone production from 2004 to 2017 using cone abscission scars in 187 trees from 28 sites along an 1100 km latitudinal gradient to investigate the spatiotemporal drivers of seed cone production and synchrony across populations. Populations from chronically hot and dry areas (greater climatic water deficits and less monsoonal precipitation) tended to have greater interannual variability in seed cone production and smaller crop sizes. Mast years generally followed years with low vapor pressure deficits and high precipitation during key periods of the reproductive process, but the strength of these relationships varied across the region. Populations that received greater monsoonal precipitation were less sensitive to late summer vapor pressure deficits during seed cone initiation yet more sensitive to spring vapor pressure deficits during pollination. Spatially correlated patterns of vapor pressure deficit better predicted synchrony in seed cone production than geographic distance, and these patterns were conserved at distances up to 500 km. These results demonstrate that aridity drives spatiotemporal variability in seed cone production. As a result, projected increases in aridity are likely to decrease the frequency and magnitude of masting in these dry forests and woodlands. Declines in seed production may compound climatic limitations to recruitment and impede tree regeneration, with cascading effects for numerous wildlife species.


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TL;DR: In this paper, the authors used estuaries as a model system to test how landscape transformation alters functional diversity in coastal seascapes, and measured how variation in level of urbanisation, catchment modification and habitat loss influenced fish diversity across thirty-nine estuarial in eastern Australia.
Abstract: The ecological impacts of landscape modification and urbanisation have transformed the composition of plant and animal assemblages, and altered the condition of ecosystems globally. Landscape transformation influences the spatial distribution of species and ecological functions by selecting for generalist species with wide ecological niches, which can adapt to opportunities in highly-modified environments. These effects of landscape modification can shape functional diversity on land, but it is not clear whether they have similar functional consequences in the sea. We used estuaries as a model system to test how landscape transformation alters functional diversity in coastal seascapes, and measured how variation in level of urbanisation, catchment modification and habitat loss influenced fish diversity across thirty-nine estuaries in eastern Australia. Fish were surveyed with baited remote underwater video stations and functional diversity was indexed with three metrics that describe variation in the functional traits and niche space of assemblages. The extent of landscape transformation in the catchment of each estuary was associated with variation in the functional diversity of estuarine fish assemblages. These effects were, however, not what we expected as functional diversity was highest in modified estuaries that supported a large area of both urban and grazing land in their catchments, were bordered by a small area of natural terrestrial vegetation and that contained a moderate area of mangroves. Zoobenthivores and omnivores dominated assemblages in highly-modified estuaries, and piscivorous fishes were common in natural waterways. Our results demonstrate, that the modification and urbanisation of ecosystems on land can alter functional diversity in the sea. Intense landscape transformation appears to select for abundant generalists with wide trophic niches, and against species with specialised diets, and we suggest that these changes might have fundamental consequences for ecosystem functioning in estuaries, and other highly modified seascapes.