Harvest and decimation affect genetic drift and the effective population size in wild reindeer.

Harvesting and culling are methods used to monitor and manage wildlife diseases. An important consequence of these practices is a change in the genetic dynamics of affected populations that may threaten their long-term viability. The effective population size (N e) is a fundamental parameter for describing such changes as it determines the amount of genetic drift in a population. Here, we estimate N e of a harvested wild reindeer population in Norway. Then we use simulations to investigate the genetic consequences of management efforts for handling a recent spread of chronic wasting disease, including increased adult male harvest and population decimation. The N e/N ratio in this population was found to be 0.124 at the end of the study period, compared to 0.239 in the preceding 14 years period. The difference was caused by increased harvest rates with a high proportion of adult males (older than 2.5 years) being shot (15.2% in 2005-2018 and 44.8% in 2021). Increased harvest rates decreased N e in the simulations, but less sex biased harvest strategies had a lower negative impact. For harvest strategies that yield stable population dynamics, shifting the harvest from calves to adult males and females increased N e. Population decimation always resulted in decreased genetic variation in the population, with higher loss of heterozygosity and rare alleles with more severe decimation or longer periods of low population size. A very high proportion of males in the harvest had the most severe consequences for the loss of genetic variation. This study clearly shows how the effects of harvest strategies and changes in population size interact to determine the genetic drift of a managed population. The long-term genetic viability of wildlife populations subject to a disease will also depend on population impacts of the disease and how these interact with management actions.

Population dynamics and seasonal migration patterns of Spodoptera exigua in northern China based on 11 years of monitoring data.

Background: The beet armyworm, Spodoptera exigua (Hübner), is an important agricultural pest worldwide that has caused serious economic losses in the main crop-producing areas of China. To effectively monitor and control this pest, it is crucial to investigate its population dynamics and seasonal migration patterns in northern China. Methods: In this study, we monitored the population dynamics of S. exigua using sex pheromone traps in Shenyang, Liaoning Province from 2012 to 2022, combining these data with amigration trajectory simulation approach and synoptic weather analysis. Results: There were significant interannual and seasonal variations in the capture number of S. exigua, and the total number of S. exigua exceeded 2,000 individuals in 2018 and 2020. The highest and lowest numbers of S. exigua were trapped in September and May, accounting for 34.65% ± 6.81% and 0.11% ± 0.04% of the annual totals, respectively. The average occurrence period was 140.9 ± 9.34 days during 2012-2022. In addition, the biomass of S. exigua also increased significantly during these years. The simulated seasonal migration trajectories also revealed varying source regions in different months, primarily originated from Northeast China and East China. These unique insights into the migration patterns of S. exigua will contribute to a deeper understanding of its occurrence in northern China and provide a theoretical basis for regional monitoring, early warning, and the development of effective management strategies for long-range migratory pests.

Comparing the long-term persistence of different Wolbachia strains after the release of bacteria-carrying mosquitoes.

This paper proposes a bidimensional modeling framework for Wolbachia invasion, assuming imperfect maternal transmission, incomplete cytoplasmic incompatibility, and direct infection loss due to thermal stress. Our model adapts to various Wolbachia strains and retains all properties of higher-dimensional models. The conditions for the durable coexistence of Wolbachia-carrying and wild mosquitoes are expressed using the model's parameters in a compact closed form. When the Wolbachia bacterium is locally established, the size of the remanent wild population can be assessed by a direct formula derived from the model. The model was tested for four Wolbachia strains undergoing laboratory and field trials to control mosquito-borne diseases: wMel, wMelPop, wAlbB, and wAu. As all these bacterial strains affect the individual fitness of mosquito hosts differently and exhibit different levels of resistance to temperature variations, the model helped to conclude that: (1) the wMel strain spreads faster in wild mosquito populations; (2) the wMelPop exhibits lower resilience but also guarantees the smallest size of the remanent wild population; (3) the wAlbB strain performs better at higher ambient temperatures than others; (4) the wAu strain is not sustainable and cannot persist in the wild mosquito population despite its resistance to high temperatures.

Bringing traits back into the equation: A roadmap to understand species redistribution.

Ecological and evolutionary theories have proposed that species traits should be important in mediating species responses to contemporary climate change; yet, empirical evidence has so far provided mixed evidence for the role of behavioral, life history, or ecological characteristics in facilitating or hindering species range shifts. As such, the utility of trait-based approaches to predict species redistribution under climate change has been called into question. We develop the perspective, supported by evidence, that trait variation, if used carefully can have high potential utility, but that past analyses have in many cases failed to identify an explanatory value for traits by not fully embracing the complexity of species range shifts. First, we discuss the relevant theory linking species traits to range shift processes at the leading (expansion) and trailing (contraction) edges of species distributions and highlight the need to clarify the mechanistic basis of trait-based approaches. Second, we provide a brief overview of range shift-trait studies and identify new opportunities for trait integration that consider range-specific processes and intraspecific variability. Third, we explore the circumstances under which environmental and biotic context dependencies are likely to affect our ability to identify the contribution of species traits to range shift processes. Finally, we propose that revealing the role of traits in shaping species redistribution may likely require accounting for methodological variation arising from the range shift estimation process as well as addressing existing functional, geographical, and phylogenetic biases. We provide a series of considerations for more effectively integrating traits as well as extrinsic and methodological factors into species redistribution research. Together, these analytical approaches promise stronger mechanistic and predictive understanding that can help society mitigate and adapt to the effects of climate change on biodiversity.

Effectiveness of population-based recovery actions for threatened southern mountain caribou.

Habitat loss is affecting many species, including the southern mountain caribou (Rangifer tarandus caribou) population in western North America. Over the last half century, this threatened caribou population's range and abundance have dramatically contracted. An integrated population model was used to analyze 51 years (1973-2023) of demographic data from 40 southern mountain caribou subpopulations to assess the effectiveness of population-based recovery actions at increasing population growth. Reducing potential limiting factors on threatened caribou populations offered a rare opportunity to identify the causes of decline and assess methods of recovery. Southern mountain caribou abundance declined by 51% between 1991 and 2023, and 37% of subpopulations were functionally extirpated. Wolf reduction was the only recovery action that consistently increased population growth when applied in isolation, and combinations of wolf reductions with maternal penning or supplemental feeding provided rapid growth but were applied to only four subpopulations. As of 2023, recovery actions have increased the abundance of southern mountain caribou by 52%, compared to a simulation with no interventions. When predation pressure was reduced, rapid population growth was observed, even under contemporary climate change and high levels of habitat loss. Unless predation is reduced, caribou subpopulations will continue to be extirpated well before habitat conservation and restoration can become effective.

Population Structures and Dynamics of Rhododendron Communities with Different Stages of Succession in Northwest Guizhou, China.

To explore the population structures and dynamics of Rhododendron shrub communities at different stages of succession in northwest Guizhou, China, this study examined the populations of Rhododendron annae and Rhododendron irroratum shrub with two different stages. A space-for-time substitution was employed to establish the diameter class/height structures, static life tables, and survival/mortality rate/disappearance rate curves of both Rhododendron populations with different orders of succession. Their structural and quantitative dynamics were analyzed, and their development trends were predicted. The results showed that, quantitatively, the populations of R. annae and R. irroratum in the two Rhododendron communities with different orders of succession were dominated by age classes one, two, and three as well as height classes i, ii, and iii. The number of Rhododendron plants at the three age classes and the three height classes accounted for 97.61-100% of the total. The quantitative dynamic indices of R. annae and R. irroratum were both greater than 0, with and without considering external interference. In terms of age class and height structures, both Rhododendron populations were expanding populations, presenting "inverted-J-shaped" and irregular pyramid patterns. There was a sufficient number of young individuals, but few or no old individuals. Both survival curves of the populations of R. annae and R. irroratum in the two Rhododendron communities with different orders of succession belonged to the Deevy-II type. In the late stage of succession, the mortality curves and disappearance curves of both Rhododendron populations in these communities presented a trend of increasing first and then decreasing with increasing age class. This result indicates that at each age class, R. annae and R. irroratum showed a trend of gradual increase after two, four, and six years. In brief, the populations of R. annae and R. irroratum have rich reserves of seedlings and saplings, but high mortality and disappearance rates. In this context, it is necessary to reduce human interference and implement targeted conservation measures to promote the natural renewal of Rhododendron populations.

Neural correlates of perceptual similarity masking in primate V1.

Visual detection is a fundamental natural task. Detection becomes more challenging as the similarity between the target and the background in which it is embedded increases, a phenomenon termed 'similarity masking'. To test the hypothesis that V1 contributes to similarity masking, we used voltage sensitive dye imaging (VSDI) to measure V1 population responses while macaque monkeys performed a detection task under varying levels of target-background similarity. Paradoxically, we find that during an initial transient phase, V1 responses to the target are enhanced, rather than suppressed, by target-background similarity. This effect reverses in the second phase of the response, so that in this phase V1 signals are positively correlated with the behavioral effect of similarity. Finally, we show that a simple model with delayed divisive normalization can qualitatively account for our findings. Overall, our results support the hypothesis that a nonlinear gain control mechanism in V1 contributes to perceptual similarity masking.

The influence of gene flow on population viability in an isolated urban caracal population.

Wildlife populations are becoming increasingly fragmented by anthropogenic development. Small and isolated populations often face an elevated risk of extinction, in part due to inbreeding depression. Here, we examine the genomic consequences of urbanization in a caracal (Caracal caracal) population that has become isolated in the Cape Peninsula region of the City of Cape Town, South Africa, and is thought to number ~50 individuals. We document low levels of migration into the population over the past ~75 years, with an estimated rate of 1.3 effective migrants per generation. As a consequence of this isolation and small population size, levels of inbreeding are elevated in the contemporary Cape Peninsula population (mean FROH = 0.20). Inbreeding primarily manifests as long runs of homozygosity >10 Mb, consistent with the effects of isolation due to the rapid recent growth of Cape Town. To explore how reduced migration and elevated inbreeding may impact future population dynamics, we parameterized an eco-evolutionary simulation model. We find that if migration rates do not change in the future, the population is expected to decline, though with a low projected risk of extinction. However, if migration rates decline or anthropogenic mortality rates increase, the potential risk of extinction is greatly elevated. To avert a population decline, we suggest that translocating migrants into the Cape Peninsula to initiate a genetic rescue may be warranted in the near future. Our analysis highlights the utility of genomic datasets coupled with computational simulation models for investigating the influence of gene flow on population viability.

Intraspecific demographic and trait responses to environmental change drivers are linked in two species of ciliate.

BACKGROUND: Over the past decade, theory and observations have suggested intraspecific variation, trait-based differences within species, as a buffer against biodiversity loss from multiple environmental changes. This buffering effect can only occur when different populations of the same species respond differently to environmental change. More specifically, variation of demographic responses fosters buffering of demography, while variation of trait responses fosters buffering of functioning. Understanding how both responses are related is important for predicting biodiversity loss and its consequences. In this study, we aimed to empirically assess whether population-level trait responses to multiple environmental change drivers are related to the demographic response to these drivers. To this end, we measured demographic and trait responses in microcosm experiments with two species of ciliated protists. For three clonal strains of each species, we measured responses to two environmental change drivers (climate change and pollution) and their combination. We also examined if relationships between demographic and trait responses existed across treatments and strains. RESULTS: We found different demographic responses across strains of the same species but hardly any interactive effects between the two environmental change drivers. Also, trait responses (summarized in a survival strategy index) varied among strains within a species, again with no driver interactions. Demographic and trait responses were related across all strains of both species tested in this study: Increasing intrinsic growth and self-limitation were associated with a shift in survival strategy from sit-and-wait towards flee. CONCLUSIONS: Our results support the existence of a link between a population's demographic and trait responses to environmental change drivers in two species of ciliate. Future work could dive deeper into the specifics of phenotypical trait values, and changes therein, related to specific life strategies in different species of ciliate and other zooplankton grazers.

Study protocol: improving response to malaria in the Amazon through identification of inter-community networks and human mobility in border regions of Ecuador, Peru and Brazil.

INTRODUCTION: Understanding human mobility's role in malaria transmission is critical to successful control and elimination. However, common approaches to measuring mobility are ill-equipped for remote regions such as the Amazon. This study develops a network survey to quantify the effect of community connectivity and mobility on malaria transmission. METHODS: We measure community connectivity across the study area using a respondent driven sampling design among key informants who are at least 18 years of age. 45 initial communities will be selected: 10 in Brazil, 10 in Ecuador and 25 in Peru. Participants will be recruited in each initial node and administered a survey to obtain data on each community's mobility patterns. Survey responses will be ranked and the 2-3 most connected communities will then be selected and surveyed. This process will be repeated for a third round of data collection. Community network matrices will be linked with each country's malaria surveillance system to test the effects of mobility on disease risk. ETHICS AND DISSEMINATION: This study protocol has been approved by the institutional review boards of Duke University (USA), Universidad San Francisco de Quito (Ecuador), Universidad Peruana Cayetano Heredia (Peru) and Universidade Federal Minas Gerais (Brazil). Results will be disseminated in communities by the end of the study.

Range-Wide Phylogeography and Ecological Niche Modeling Provide Insights into the Evolutionary History of the Mongolian Racerunner (Eremias argus) in Northeast Asia.

The Mongolian racerunner, Eremias argus, is a small lizard endemic to Northeast Asia that can serve as an excellent model for investigating how geography and past climate change have jointly influenced the evolution of biodiversity in this region. To elucidate the processes underlying its diversification and demography, we reconstructed the range-wide phylogeographic pattern and evolutionary trajectory, using phylogenetic, population genetic, landscape genetic, Bayesian phylogeographic reconstruction and ecological niche modeling approaches. Phylogenetic analyses of the mtDNA cyt b gene revealed eight lineages that were unbounded by geographic region. The genetic structure of E. argus was mainly determined by geographic distance. Divergence dating indicated that E. argus and E. brenchleyi diverged during the Mid-Pliocene Warm Period. E. argus was estimated to have coalesced at~0.4351 Ma (Marine Isotope Stage 19). Bayesian phylogeographic diffusion analysis revealed out-of-Inner Mongolia and rapid colonization events from the end of the Last Interglacial to the Last Glacial Maximum, which is consistent with the expanded suitable range of the Last Glacial Maximum. Pre-Last Glacial Maximum growth of population is presented for most lineages of E. argus. The Glacial Maximum contraction model and the previous multiple glacial refugia hypotheses are rejected. This may be due to an increase in the amount of climatically favorable habitats in Northeast Asia. Furthermore, E. argus barbouri most likely represents an invalid taxon. The present study is the first to report a range-wide phylogeography of reptiles over such a large region in Northeast Asia. Our results make a significant contribution towards understanding the biogeography of the entire Northeast Asia.

The angiogenic growth of cities.

Describing the space-time evolution of urban population is a fundamental challenge in the science of cities, yet a complete theoretical treatment of the underlying dynamics is still missing. Here, we first reconstruct the evolution of London (UK) over 180 years and show that urban growth consists of an initial phase of diffusion-limited growth, followed by the development of the railway transport network and a consequential shift from central to suburban living. Such dynamics-which are analogous to angiogenesis in biological systems-can be described by a minimalist reaction-diffusion model coupled with economic constraints and an adaptive transport network. We then test the generality of our approach by reproducing the evolution of Sydney, Australia, from 1851 to 2011. We show that the rail system coevolves with urban population, displaying hierarchical characteristics that remain constant over time unless large-scale interventions are put in place to alter the modes of transport. These results demonstrate that transport schemes are first-order controls of long-term urbanization patterns and efforts aimed at creating more sustainable and healthier cities require careful consideration of population-transport feedbacks.

Redefining 'state-of-the-art' for integrated population models with immigration.

Research Highlight: Christian, M., Oosthuizen, W. C., Bester, M. N., & de Bruyn, P. N. (2024). Robustly estimating the demographic contribution of immigration: Simulation, sensitivity analysis and seals. Journal of Animal Ecology. https://doi.org/10.1111/1365-2656.14053. Immigration can have profound consequences for local population dynamics and demography, but collecting data to accurately quantifying it is challenging. The recent rise of integrated population models (IPMs) offers an alternative by making it possible to estimate immigration without the need for explicit data, and to quantify its contribution to population dynamics through transient Life Table Response Experiments (tLTREs). Simulation studies have, however, highlighted that this approach can be prone to bias and overestimation. In their new study, Christian et al. address one of the root causes of this issue by improving the estimation of time variation in vital rates and immigration using Gaussian processes in lieu of traditionally used temporal random effects. They demonstrate that IPM-tLTRE frameworks with Gaussian processes produce more accurate and less biased estimates of immigration and its contribution to population dynamics and illustrate the applicability of this approach using a long-term data set on elephant seals (Mirounga leonida). Results are validated with a simulation study and suggest that immigration of breeding females has been central for population recovery of elephant seals despite the species' high female site fidelity. Christian et al. thus present new insights into population regulation of long-lived marine mammals and highlight the potential for using Gaussian process priors in IPMs. They also illustrate a suite of 'best practices' for state-of-the-art IPM-tLTRE analyses and provide an inspirational example for the kind of ecological modelling workflow that can be invaluable not just as a starting point for fellow ecologists picking up or improving their own IPM-tLTRE analyses, but also for teaching and in contexts where model estimates are used for informing management and conservation decision-making.

An increase in management actions has compensated for past climate change effects on desert locust gregarization in western Africa.

In response to high population density, the desert locust, Schistocerca gregaria, becomes gregarious and forms swarms that can cause significant damage to crops and pastures, threatening food security of human populations from western Africa to India. This switch from solitary to gregarious populations is highly dependent on favorable weather conditions. Climate change, which has been hypothesized to shift conditions towards increasing risks of gregarization, is therefore likely to have significant impacts on the spatial distribution and likelihood of outbreak events. However, the desert locust is intensely managed at large scales, which possibly counteracts any increased risk of outbreaks due to a more favorable climate. Consequently, understanding the changes in risks in the future involves teasing out the effects of climate change and management actions. Here we studied the dynamics of gregarization at the very early stages of potential outbreaks, in parallel with trends in climate and management, between 1985 and 2018 in western Africa. We used three different spatial scales, with the goal to have a better understanding of the potential effects of climate change per se while controlling for management. Our first approach was to look at a regional scale, where we observed an overall decrease in gregarization events. However, this scale includes very heterogeneous environments and management efforts. To consider this heterogeneity, we divided the area into a grid of 0.5° cells. For each cell, a climate analysis was performed for rainfall and temperature, with trends obtained by a harmonic decomposition model on monthly data. Analyses of gregarization showed only a few significant trends, both positive and negative, mainly found in western Mauritania where management effort has increased. To improve the statistical power, these cells were then grouped into larger homogeneous climatic clusters, i.e. groups of cells with similar climatic conditions and similar climatic trends over the study period. At this scale, gregarization events depend on the intersection between climate conditions and management efforts. The clusters where gregarization increased were also the ones with the highest increase of management. These results highlight the important effect of preventive management, which may counteract the positive effects of climate change on locust proliferation.

Decadal demographic shifts and size-dependent disturbance responses of corals in a subtropical warming hotspot.

Long-term demographic studies at biogeographic transition zones can elucidate how body size mediates disturbance responses. Focusing on subtropical reefs in eastern Australia, we examine trends in the size-structure of corals with contrasting life-histories and zoogeographies surrounding the 2016 coral bleaching event (2010-2019) to determine their resilience and recovery capacity. We document demographic shifts, with disproportionate declines in the number of small corals and long-term persistence of larger corals. The incidence of bleaching (Pocillopora, Turbinaria) and partial mortality (Acropora, Pocillopora) increased with coral size, and bleached corals had greater risk of partial mortality. While endemic Pocillopora experienced marked declines, decadal stability of Turbinaria despite bleaching, coupled with abundance increase and bleaching resistance in Acropora indicate remarkable resilience of these taxa in the subtropics. Declines in the number of small corals and variable associations with environmental drivers indicate bottlenecks to recovery mediated by inhibitory effects of thermal extremes for Pocillopora (heat stress) and Acropora (heat and cold stress), and stimulatory effects of chlorophyll-a for Turbinaria. Although our study reveals signs of resilience, it foreshadows the vulnerability of subtropical corals to changing disturbance regimes that include marine heatwaves. Disparity in population dynamics suggest that subtropical reefs are ecologically distinct from tropical coral reefs.

Cost-benefit analysis of ecosystem modeling to support fisheries management.

Mathematical and statistical models underlie many of the world's most important fisheries management decisions. Since the 19th century, difficulty calibrating and fitting such models has been used to justify the selection of simple, stationary, single-species models to aid tactical fisheries management decisions. Whereas these justifications are reasonable, it is imperative that we quantify the value of different levels of model complexity for supporting fisheries management, especially given a changing climate, where old methodologies may no longer perform as well as in the past. Here we argue that cost-benefit analysis is an ideal lens to assess the value of model complexity in fisheries management. While some studies have reported the benefits of model complexity in fisheries, modeling costs are rarely considered. In the absence of cost data in the literature, we report, as a starting point, relative costs of single-species stock assessment and marine ecosystem models from two Australian organizations. We found that costs varied by two orders of magnitude, and that ecosystem model costs increased with model complexity. Using these costs, we walk through a hypothetical example of cost-benefit analysis. The demonstration is intended to catalyze the reporting of modeling costs and benefits.

An approximation of populations on a habitat with large carrying capacity.

We consider stochastic dynamics of a population which starts from a small colony on a habitat with large but limited carrying capacity. A common heuristics suggests that such population grows initially as a Galton-Watson branching process and then its size follows an almost deterministic path until reaching its maximum, sustainable by the habitat. In this paper we put forward an alternative and, in fact, more accurate approximation which suggests that the population size behaves as a special nonlinear transformation of the Galton-Watson process from the very beginning.

Temporal development in the impacts of plant invasions: search for the underlying mechanisms.

Many invasive plants have negative impacts on native populations and communities, but there remains much uncertainty about how these impacts develop over time. In this review, I describe the mechanisms that promote the initial dominance of invaders, the characteristic associated with large negative impacts, and present the processes that contribute to changes in invader abundance and impacts over time. Together with ecological processes such as ecosystem engineering or enemy accumulation, I show that temporal variation in impacts can be linked to evolution in both native and invasive species. I also show that multiple processes operating in the same invasion system can jointly shape long-term impacts. Finally, I present the framework of modern of coexistence theory as a tool for predicting the effects of invaders on native populations, and how these effects change with processes ongoing within invaded communities.

Syncopation and synchrony: Phenological dynamics of Pyropia nereocystis (Bangiophyceae) in central California.

Pyropia nereocystis is an annual northeastern Pacific-bladed bangialean species whose macroscopic stage epiphytized the annual canopy forming bull kelp Nereocystis luetkeana. I examined three in situ facets of these epiphyte-host dynamics in the central California region: (1) spatial and temporal variation in the presence of P. nereocystis epiphytes as a function of host density, (2) the relationship between individual host morphology and epiphytic P. nereocystis biomass, and (3) the ecophysiological growth ramifications for subtidal transplants of both life stages of P. nereocystis. Swath canopy surveys and whole host collections were conducted at five sites between November 2017 and February 2019. Additionally, transplants of P. nereocystis gametophytes and sporophytes were conducted across multiple subtidal depths. I observed temporal changes in the proportions of hosts epiphytized by P. nereocystis, with differences in seasonal persistence of P. nereocystis among sites and between years. Biomass of P. nereocystis was positively correlated with individual host stipe length, stipe surface area, and the primary principal component (PC) of stipe morphometrics denoted by principal component analysis (PCA). Gametogenesis in P. nereocystis epiphytes was spatially heterogeneous and limited for the 2018-2019 cohort due to comprehensive removal of hosts by the February 2019 sampling period. Transplants of P. nereocystis gametophytes yielded similar growth responses among depths, and sporophyte (conchocelis) transplant areal growth was positively correlated with transplant depth. These findings detail spatiotemporal complexity and multi-scale (individual, site, and whole region) phenological nuances for central Californian P. nereocystis epiphytes.

The GARD Prebiotic Reproduction Model Described in Order and Complexity.

Early steps in the origin of life were necessarily connected to the unlikely formation of self-reproducing structures from chaotic chemistry. Simulations of chemical kinetics based on the graded autocatalysis replication domain (GARD) model demonstrate the ability of a micellar system to become self-reproducing units away from equilibrium. Even though they may be very rare in the initial state of the system, the property of their endogenous mutually catalytic networks being dynamic attractors greatly enhanced reproduction propensity, revealing their potential for selection and Darwinian evolution processes. In parallel, order and complexity have been shown to be crucial parameters in successful evolution. Here, we probe these parameters in the dynamics of GARD-governed entities in an attempt to identify characteristic mechanisms of their development in non-covalent molecular assemblies. Using a virtual random walk perspective, a value for consecutive order is defined based on statistical thermodynamics. The complexity, on the other hand, is determined by the size of a minimal algorithm fully describing the statistical properties of the random walk. By referring to a previously published diagonal line in an order/complexity diagram that represents the progression of evolution, it is shown that the GARD model has the potential to advance in this direction. These results can serve as a solid foundation for identifying general criteria for future analyses of evolving systems.