The multi-species conservation approaches starting from Vane-Wright et al. (1991) search for a bi... more The multi-species conservation approaches starting from Vane-Wright et al. (1991) search for a biologically valid basis to evaluate biodiversity. Such evaluations can be used to optimize reserve boundaries so reserves contain a high level of biodiversity. However, these optimization procedures do not minimize the future loss of biodiversity. We provide a method that can be used to minimize the loss of biodiversity. The method integrates an evolutionary species evaluation with an ecological multi-species risk analysis to estimate the expected loss of phylogenetic diversity. A minimization of this loss will optimize the preservation of phylogenetic diversity.
Narwhals are one of three highly specialized whale species that are endemic to the Arctic ([ 1 ][... more Narwhals are one of three highly specialized whale species that are endemic to the Arctic ([ 1 ][1]). The global narwhal population may number more than 100,000 individuals, but the species persists as a complex, highly divided meta-population, with limited or no exchange between neighboring subpopulations ([ 1 ][1]). Several narwhal populations in Greenland are suffering from low and declining numbers, and unsustainable hunting is putting the species at risk of local extirpation ([ 1 ][1]–[ 5 ][2]). Narwhal conservation requires human activities to be managed at the scale of subpopulations, each of which has its own environmental conditions and exploitation history. In Melville Bay, the number of narwhals killed by hunters has likely been unsustainable for a decade or more. From 2007 to 2019, the size of the area used by narwhals in Melville Bay has shrunk by 84% from 16,000 to 2600 km2 ([ 6 ][3]). The North Atlantic Marine Mammal Commission (NAMMCO) and the Canada-Greenland Joint Commission on Conservation and Management of Narwhal and Beluga recommended a limit of 280 removals between 2015 and 2019 ([ 2 ][4]), but the estimated accumulated removal during this period was at least 423 narwhals ([ 7 ][5]). In Southeast Greenland, the Scientific Committee of NAMMCO recommended in 2017—and reiterated in 2018—that annual catches should be reduced to fewer than 20 narwhals and that no narwhal should be taken south of 68°N ([ 3 ][6], [ 4 ][7]). When improved population modeling outputs became available in 2019, the Scientific Committee changed its recommendation to a moratorium on narwhal hunting throughout Southeast Greenland ([ 5 ][2]). Even so, the catches from 2017 to 2019 totaled 268 animals ([ 5 ][2]). The effects of the ongoing overharvesting can be seen in the population composition: a decreased proportion of females, an overrepresentation of old males, and an absence of calves and juveniles ([ 5 ][2]). The quota for 2020 is set at 58 narwhals ([ 8 ][8], [ 9 ][9]); this level of harvest could put the long-term existence of the narwhal stocks in Southeast Greenland in jeopardy. The narwhal is regarded as the most sensitive of all Arctic endemic marine mammals to climate change because of its adaptations to a narrow sea-temperature niche, dependence on sea ice, specialized feeding habits, relatively restricted range, and general sensitivity to ocean noise and other forms of anthropogenic disturbance ([ 1 ][1]). It is vital for authorities in Greenland to accept scientific advice regarding regional narwhal population declines and take the need for responsible management seriously. Given the extreme site fidelity of narwhals ([ 1 ][1]), individuals from other populations are unlikely to recolonize localities where the species has been extirpated. The loss of a local narwhal population from a specific fjord system is likely to be permanent. 1. [↵][10]1. R. C. Hobbs et al ., Mar. Fish. Rev. 81, 1 (2019). [OpenUrl][11] 2. [↵][12]NAMMCO, “Report of the 22nd Scientific Committee meeting” (2015). 3. [↵][13]NAMMCO, “Report of the 24th Scientific Committee meeting” (2017). 4. [↵][14]NAMMCO, 2018. “Report of the 25th Scientific Committee meeting” (2018). 5. [↵][15]NAMMCO, “Report of the 26th Scientific Committee meeting” (2019). 6. [↵][16]1. R. G. Hansen et al ., “Trends in abundance and distribution of narwhals ( Monodon monoceros ) on the summering grounds in Inglefield Bredning and Melville Bay, Greenland from 2007–2019” (NAMMCO–JCNB Joint Working Group on Narwhals and Belugas, 2020). 7. [↵][17]Government of Greenland, “Recommendations, quotas, and catches for the most important species” (2020); [in Danish]. 8. [↵][18]Government of Greenland, “2020 quotas for belugas and narwhals” (2020); [in Danish]. 9. [↵][19]Government of Greenland, Press release 15/6/2020 from the Department of Fisheries, Hunting and Agriculture, Journal number 2020-1593, File 13998768 (2020) [in Danish]. [1]: #ref-1 [2]: #ref-5 [3]: #ref-6 [4]: #ref-2 [5]: #ref-7 [6]: #ref-3 [7]: #ref-4 [8]: #ref-8 [9]: #ref-9 [10]: #xref-ref-1-1 "View reference 1 in text" [11]: {openurl}?query=rft.jtitle%253DMar.%2BFish.%2BRev.%26rft.volume%253D81%26rft.spage%253D1%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [12]: #xref-ref-2-1 "View reference 2 in text" [13]: #xref-ref-3-1 "View reference 3 in text" [14]: #xref-ref-4-1 "View reference 4 in text" [15]: #xref-ref-5-1 "View reference 5 in text" [16]: #xref-ref-6-1 "View reference 6 in text" [17]: #xref-ref-7-1 "View reference 7 in text" [18]: #xref-ref-8-1 "View reference 8 in text" [19]: #xref-ref-9-1 "View reference 9 in text"
With 100,000 narwhals inhabiting populations in the Canadian Arctic Archipelago, the Northwest Pa... more With 100,000 narwhals inhabiting populations in the Canadian Arctic Archipelago, the Northwest Passage runs through the largest aggregation of narwhals in the world. I use a metapopulation analysis that covers eight of the narwhal populations in the area to estimate that about 25,000 narwhals have emigrated from Eclipse Sound to Admiralty Inlet since 2007, causing almost total population collapse in Eclipse Sound following shipping of iron ore through Milne Inlet and Eclipse Sound. It is estimated that up to about 30,000 narwhals may vanish in the long run, unless shipping is increasingly regulated, or narwhals adapt to the disturbance and reinhabits the abandoned areas. Reflecting only a local disturbance, these population effects warn against intense shipping in other fjords with narwhals, and against a potentially much larger impact from increased shipping through the Northwest Passage in the future.
I use 56,214 life history data to estimate equilibrium life history models for birds and mammals ... more I use 56,214 life history data to estimate equilibrium life history models for birds and mammals with body mass estimates. Missing parameters were estimated by allometric correlations at the lowest taxonomic level (genus, family, order, class) with data. The estimation is optimised to predict the existing data, with precision estimated separately for the different taxonomic levels of the estimator. This provides complete life history models for 9,488 species of birds, and 4,865 species of mammals. Each model includes estimates of metabolism, net assimilated energy, individual growth, mortality, fecundity, age of reproductive maturity, generation time, life span, home range, population density, biomass, population consumption, and a relative measure of intra-specific interactive competition, providing 387,531 parameter estimates in total.
I use the North American Breeding Bird Survey (Sauer et al. 2017) to construct 462 population tra... more I use the North American Breeding Bird Survey (Sauer et al. 2017) to construct 462 population trajectories with about 50 yearly abundance estimates each. Applying AIC model-selection, I find that selection-regulated population dynamics is 25,000 (95%:0.42-1.7e17) times more probable than density-regulated growth. Selection is essential in 94% of the best models explaining 82% of the population dynamics variance across the North American continent. Similar results are obtained for 111, 215, and 420 populations of British birds (BTO 2020), Danish birds (DOF 2020), and birds and mammals in the Global Population Dynamic Database (GPDD 2010). The traditional paradigm---that the population dynamic growth rate is a function of the environment, with maximal per-capita growth at low population densities, and sub-optimal reproduction from famine at carrying capacities with strong competition for limited resources---is not supported. Selection regulation generates a new paradigm where the worl...
Since neo‐Darwinism arose from the work of Darwin and Mendel evolution by natural selection has b... more Since neo‐Darwinism arose from the work of Darwin and Mendel evolution by natural selection has been seen as contingent and historical being defined by an a posteriori selection process with no a priori laws that explain why evolution on Earth has taken the direction of the major evolutionary trends and transitions instead of any other direction. Recently, however, major life‐history trends and transitions have been explained as inevitable because of a deterministic selection that unfolds from the energetic state of the organism and the density‐dependent competitive interactions that arise from self‐replication in limited environments. I describe differences and similarities between the historical and deterministic selection processes, illustrate concepts using life‐history models on large body masses and limited reproductive rates, review life‐history evolution with a wider focus on major evolutionary transitions, and propose that biotic evolution is driven by a universal natural s...
Biodiversity conservation is confronted with two major problems: how to define and measure biodiv... more Biodiversity conservation is confronted with two major problems: how to define and measure biodiversity, and how to optimize the in situ conservation of biodiversity. Here we outline a conceptual framework for biodiversity conservation that is directed towards these problems. The framework combines a phylogenetic evaluation with a multi-species risk analysis and defines the objective of conservation biology as the minimization of the future loss of biodiversity.
In their Science paper, Desforges et al. (2018) address PCB pollution in killer whales, predictin... more In their Science paper, Desforges et al. (2018) address PCB pollution in killer whales, predicting a decline in calf survival and an associated collapse of killer whale populations worldwide. I refute the collapse, showing that it follows from a flawed model parametrisation. The result is not questioning contamination problems in killer whales, but only the bold prediction of global collapse.
This paper combines the catch allocation model for narwhals in East Canada and West Greenland wit... more This paper combines the catch allocation model for narwhals in East Canada and West Greenland with Bayesian population modelling of the eight summer aggregations of narwhals in the region. The catch allocation model allocates the catches in different hunting areas and seasons to the different summer aggregations, and the population models analyse the impact of these catches on the population dynamics of the eight narwhal aggregations. The population models run from 1970, and the catch allocation model needs population trajectories from 1970 to the present in order to estimate the catches taken from the different summer aggregations during this period. In an initial run it uses linear transitions between the available abundance estimates; but more elaborate population trajectories are estimated by the fit of the population models to the abundance data. The two models are therefore run in an iterative manner until the catch histories that are estimated by the allocation model, and the...
I show that the natural selection of metabolism and mass is selecting for the major life history ... more I show that the natural selection of metabolism and mass is selecting for the major life history and allometric transitions that define lifeforms from viruses, over prokaryotes and larger unicells, to multicellular animals with sexual reproduction. The proposed selection is driven by a mass specific metabolism that is selected as the pace of the resource handling that generates net energy for self-replication. This implies that an initial selection of mass is given by a dependence of mass specific metabolism on mass in replicators that are close to a lower size limit. A maximum dependence that is sublinear is shown to select for virus-like replicators with no intrinsic metabolism, no cell, and practically no mass. A maximum superlinear dependence is instead selecting for prokaryote-like self-replicating cells with asexual reproduction and incomplete metabolic pathways. These self-replicating cells have selection for increased net energy, and this generates a gradual unfolding of a p...
I simulate the evolution of metabolism and mass to explain the curvature in the metabolic allomet... more I simulate the evolution of metabolism and mass to explain the curvature in the metabolic allometry for placental and marsupial mammals. I assume that the release of inter-specific competition by the extinction of dinosaurs 65 million years ago made it possible for each clade to diversity into a multitude of species across a wide range of niches. The natural selection of metabolism and mass was then fitted to explain the maximum observed body masses over time, as well as the current inter-specific allometry for metabolism. The estimated selection of mass specific metabolism was found to bend the metabolic allometry over time, with the strongest curvature in the placental clade. The rate of exponential increase in mass specific metabolism for placentals was estimated to 9.3x10^-9 (95% CI: 7.3x10^-9 - 1.1x10^-8) on the per generation time-scale. This is an order of magnitude larger than the estimate for marsupials, in agreement with an average metabolism that is 30% larger in placenta...
The multi-species conservation approaches starting from Vane-Wright et al. (1991) search for a bi... more The multi-species conservation approaches starting from Vane-Wright et al. (1991) search for a biologically valid basis to evaluate biodiversity. Such evaluations can be used to optimize reserve boundaries so reserves contain a high level of biodiversity. However, these optimization procedures do not minimize the future loss of biodiversity. We provide a method that can be used to minimize the loss of biodiversity. The method integrates an evolutionary species evaluation with an ecological multi-species risk analysis to estimate the expected loss of phylogenetic diversity. A minimization of this loss will optimize the preservation of phylogenetic diversity.
Narwhals are one of three highly specialized whale species that are endemic to the Arctic ([ 1 ][... more Narwhals are one of three highly specialized whale species that are endemic to the Arctic ([ 1 ][1]). The global narwhal population may number more than 100,000 individuals, but the species persists as a complex, highly divided meta-population, with limited or no exchange between neighboring subpopulations ([ 1 ][1]). Several narwhal populations in Greenland are suffering from low and declining numbers, and unsustainable hunting is putting the species at risk of local extirpation ([ 1 ][1]–[ 5 ][2]). Narwhal conservation requires human activities to be managed at the scale of subpopulations, each of which has its own environmental conditions and exploitation history. In Melville Bay, the number of narwhals killed by hunters has likely been unsustainable for a decade or more. From 2007 to 2019, the size of the area used by narwhals in Melville Bay has shrunk by 84% from 16,000 to 2600 km2 ([ 6 ][3]). The North Atlantic Marine Mammal Commission (NAMMCO) and the Canada-Greenland Joint Commission on Conservation and Management of Narwhal and Beluga recommended a limit of 280 removals between 2015 and 2019 ([ 2 ][4]), but the estimated accumulated removal during this period was at least 423 narwhals ([ 7 ][5]). In Southeast Greenland, the Scientific Committee of NAMMCO recommended in 2017—and reiterated in 2018—that annual catches should be reduced to fewer than 20 narwhals and that no narwhal should be taken south of 68°N ([ 3 ][6], [ 4 ][7]). When improved population modeling outputs became available in 2019, the Scientific Committee changed its recommendation to a moratorium on narwhal hunting throughout Southeast Greenland ([ 5 ][2]). Even so, the catches from 2017 to 2019 totaled 268 animals ([ 5 ][2]). The effects of the ongoing overharvesting can be seen in the population composition: a decreased proportion of females, an overrepresentation of old males, and an absence of calves and juveniles ([ 5 ][2]). The quota for 2020 is set at 58 narwhals ([ 8 ][8], [ 9 ][9]); this level of harvest could put the long-term existence of the narwhal stocks in Southeast Greenland in jeopardy. The narwhal is regarded as the most sensitive of all Arctic endemic marine mammals to climate change because of its adaptations to a narrow sea-temperature niche, dependence on sea ice, specialized feeding habits, relatively restricted range, and general sensitivity to ocean noise and other forms of anthropogenic disturbance ([ 1 ][1]). It is vital for authorities in Greenland to accept scientific advice regarding regional narwhal population declines and take the need for responsible management seriously. Given the extreme site fidelity of narwhals ([ 1 ][1]), individuals from other populations are unlikely to recolonize localities where the species has been extirpated. The loss of a local narwhal population from a specific fjord system is likely to be permanent. 1. [↵][10]1. R. C. Hobbs et al ., Mar. Fish. Rev. 81, 1 (2019). [OpenUrl][11] 2. [↵][12]NAMMCO, “Report of the 22nd Scientific Committee meeting” (2015). 3. [↵][13]NAMMCO, “Report of the 24th Scientific Committee meeting” (2017). 4. [↵][14]NAMMCO, 2018. “Report of the 25th Scientific Committee meeting” (2018). 5. [↵][15]NAMMCO, “Report of the 26th Scientific Committee meeting” (2019). 6. [↵][16]1. R. G. Hansen et al ., “Trends in abundance and distribution of narwhals ( Monodon monoceros ) on the summering grounds in Inglefield Bredning and Melville Bay, Greenland from 2007–2019” (NAMMCO–JCNB Joint Working Group on Narwhals and Belugas, 2020). 7. [↵][17]Government of Greenland, “Recommendations, quotas, and catches for the most important species” (2020); [in Danish]. 8. [↵][18]Government of Greenland, “2020 quotas for belugas and narwhals” (2020); [in Danish]. 9. [↵][19]Government of Greenland, Press release 15/6/2020 from the Department of Fisheries, Hunting and Agriculture, Journal number 2020-1593, File 13998768 (2020) [in Danish]. [1]: #ref-1 [2]: #ref-5 [3]: #ref-6 [4]: #ref-2 [5]: #ref-7 [6]: #ref-3 [7]: #ref-4 [8]: #ref-8 [9]: #ref-9 [10]: #xref-ref-1-1 "View reference 1 in text" [11]: {openurl}?query=rft.jtitle%253DMar.%2BFish.%2BRev.%26rft.volume%253D81%26rft.spage%253D1%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [12]: #xref-ref-2-1 "View reference 2 in text" [13]: #xref-ref-3-1 "View reference 3 in text" [14]: #xref-ref-4-1 "View reference 4 in text" [15]: #xref-ref-5-1 "View reference 5 in text" [16]: #xref-ref-6-1 "View reference 6 in text" [17]: #xref-ref-7-1 "View reference 7 in text" [18]: #xref-ref-8-1 "View reference 8 in text" [19]: #xref-ref-9-1 "View reference 9 in text"
With 100,000 narwhals inhabiting populations in the Canadian Arctic Archipelago, the Northwest Pa... more With 100,000 narwhals inhabiting populations in the Canadian Arctic Archipelago, the Northwest Passage runs through the largest aggregation of narwhals in the world. I use a metapopulation analysis that covers eight of the narwhal populations in the area to estimate that about 25,000 narwhals have emigrated from Eclipse Sound to Admiralty Inlet since 2007, causing almost total population collapse in Eclipse Sound following shipping of iron ore through Milne Inlet and Eclipse Sound. It is estimated that up to about 30,000 narwhals may vanish in the long run, unless shipping is increasingly regulated, or narwhals adapt to the disturbance and reinhabits the abandoned areas. Reflecting only a local disturbance, these population effects warn against intense shipping in other fjords with narwhals, and against a potentially much larger impact from increased shipping through the Northwest Passage in the future.
I use 56,214 life history data to estimate equilibrium life history models for birds and mammals ... more I use 56,214 life history data to estimate equilibrium life history models for birds and mammals with body mass estimates. Missing parameters were estimated by allometric correlations at the lowest taxonomic level (genus, family, order, class) with data. The estimation is optimised to predict the existing data, with precision estimated separately for the different taxonomic levels of the estimator. This provides complete life history models for 9,488 species of birds, and 4,865 species of mammals. Each model includes estimates of metabolism, net assimilated energy, individual growth, mortality, fecundity, age of reproductive maturity, generation time, life span, home range, population density, biomass, population consumption, and a relative measure of intra-specific interactive competition, providing 387,531 parameter estimates in total.
I use the North American Breeding Bird Survey (Sauer et al. 2017) to construct 462 population tra... more I use the North American Breeding Bird Survey (Sauer et al. 2017) to construct 462 population trajectories with about 50 yearly abundance estimates each. Applying AIC model-selection, I find that selection-regulated population dynamics is 25,000 (95%:0.42-1.7e17) times more probable than density-regulated growth. Selection is essential in 94% of the best models explaining 82% of the population dynamics variance across the North American continent. Similar results are obtained for 111, 215, and 420 populations of British birds (BTO 2020), Danish birds (DOF 2020), and birds and mammals in the Global Population Dynamic Database (GPDD 2010). The traditional paradigm---that the population dynamic growth rate is a function of the environment, with maximal per-capita growth at low population densities, and sub-optimal reproduction from famine at carrying capacities with strong competition for limited resources---is not supported. Selection regulation generates a new paradigm where the worl...
Since neo‐Darwinism arose from the work of Darwin and Mendel evolution by natural selection has b... more Since neo‐Darwinism arose from the work of Darwin and Mendel evolution by natural selection has been seen as contingent and historical being defined by an a posteriori selection process with no a priori laws that explain why evolution on Earth has taken the direction of the major evolutionary trends and transitions instead of any other direction. Recently, however, major life‐history trends and transitions have been explained as inevitable because of a deterministic selection that unfolds from the energetic state of the organism and the density‐dependent competitive interactions that arise from self‐replication in limited environments. I describe differences and similarities between the historical and deterministic selection processes, illustrate concepts using life‐history models on large body masses and limited reproductive rates, review life‐history evolution with a wider focus on major evolutionary transitions, and propose that biotic evolution is driven by a universal natural s...
Biodiversity conservation is confronted with two major problems: how to define and measure biodiv... more Biodiversity conservation is confronted with two major problems: how to define and measure biodiversity, and how to optimize the in situ conservation of biodiversity. Here we outline a conceptual framework for biodiversity conservation that is directed towards these problems. The framework combines a phylogenetic evaluation with a multi-species risk analysis and defines the objective of conservation biology as the minimization of the future loss of biodiversity.
In their Science paper, Desforges et al. (2018) address PCB pollution in killer whales, predictin... more In their Science paper, Desforges et al. (2018) address PCB pollution in killer whales, predicting a decline in calf survival and an associated collapse of killer whale populations worldwide. I refute the collapse, showing that it follows from a flawed model parametrisation. The result is not questioning contamination problems in killer whales, but only the bold prediction of global collapse.
This paper combines the catch allocation model for narwhals in East Canada and West Greenland wit... more This paper combines the catch allocation model for narwhals in East Canada and West Greenland with Bayesian population modelling of the eight summer aggregations of narwhals in the region. The catch allocation model allocates the catches in different hunting areas and seasons to the different summer aggregations, and the population models analyse the impact of these catches on the population dynamics of the eight narwhal aggregations. The population models run from 1970, and the catch allocation model needs population trajectories from 1970 to the present in order to estimate the catches taken from the different summer aggregations during this period. In an initial run it uses linear transitions between the available abundance estimates; but more elaborate population trajectories are estimated by the fit of the population models to the abundance data. The two models are therefore run in an iterative manner until the catch histories that are estimated by the allocation model, and the...
I show that the natural selection of metabolism and mass is selecting for the major life history ... more I show that the natural selection of metabolism and mass is selecting for the major life history and allometric transitions that define lifeforms from viruses, over prokaryotes and larger unicells, to multicellular animals with sexual reproduction. The proposed selection is driven by a mass specific metabolism that is selected as the pace of the resource handling that generates net energy for self-replication. This implies that an initial selection of mass is given by a dependence of mass specific metabolism on mass in replicators that are close to a lower size limit. A maximum dependence that is sublinear is shown to select for virus-like replicators with no intrinsic metabolism, no cell, and practically no mass. A maximum superlinear dependence is instead selecting for prokaryote-like self-replicating cells with asexual reproduction and incomplete metabolic pathways. These self-replicating cells have selection for increased net energy, and this generates a gradual unfolding of a p...
I simulate the evolution of metabolism and mass to explain the curvature in the metabolic allomet... more I simulate the evolution of metabolism and mass to explain the curvature in the metabolic allometry for placental and marsupial mammals. I assume that the release of inter-specific competition by the extinction of dinosaurs 65 million years ago made it possible for each clade to diversity into a multitude of species across a wide range of niches. The natural selection of metabolism and mass was then fitted to explain the maximum observed body masses over time, as well as the current inter-specific allometry for metabolism. The estimated selection of mass specific metabolism was found to bend the metabolic allometry over time, with the strongest curvature in the placental clade. The rate of exponential increase in mass specific metabolism for placentals was estimated to 9.3x10^-9 (95% CI: 7.3x10^-9 - 1.1x10^-8) on the per generation time-scale. This is an order of magnitude larger than the estimate for marsupials, in agreement with an average metabolism that is 30% larger in placenta...
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