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The Silent Hemorrhage: A Global Assessment of Anthropogenic Genetic Erosion and the Erasure of Evolutionary Potential


Abstract


The biodiversity crisis has traditionally been cataloged through the binary lens of species extinction—the complete cessation of a lineage. However, a far more insidious and widespread phenomenon precedes species loss: the erosion of genetic diversity within surviving populations. This "cryptic extinction" removes the evolutionary fuel required for adaptation to a rapidly changing biosphere, leaving species demographically present but genetically impoverished—the "living dead." This report provides an exhaustive synthesis of the landmark 2025 global meta-analysis by Shaw et al., published in Nature, which synthesized over 80,000 records to quantify this loss. We integrate these empirical findings with cutting-edge theoretical frameworks, specifically the Mutation-Area Relationship (MAR) and the Genetic Diversity-Area Relationship (GDAR), and advanced simulation methodologies like WFmoments. The analysis reveals that while genetic diversity is declining globally—particularly among mammals and birds—the rate of loss follows complex, non-equilibrium dynamics characterized by significant time lags, or "drift debt." Crucially, the data indicates that this erosion is not thermodynamic inevitability; active, genetically informed conservation interventions have demonstrably reversed declines in specific case studies. We further explore the operational challenges of the Kunming-Montreal Global Biodiversity Framework (GBF) indicators, the pervasive geographic biases in current genomic monitoring, and the technological renaissance offered by portable sequencing in bridging the "genomic gap" between the Global North and South.


1. Introduction to the Field: The Invisible Crisis



1.1 The Architecture of Evolutionary Potential


Biodiversity is hierarchical, existing at the level of ecosystems, species, and genes. While ecosystem degradation and species extinction garner visceral public attention and immediate policy responses, the loss of genetic diversity—the fundamental variation within DNA sequences among individuals of a population—remains largely invisible to the naked eye. Yet, it is this variation that constitutes the architecture of evolutionary potential.

Genetic diversity is not merely a record of evolutionary history; it is the functional toolkit for future survival. It manifests in the variability of the Major Histocompatibility Complex (MHC), which dictates immune response to novel pathogens; in the polygenic scores determining drought tolerance in plants; and in the behavioral plasticity of mammals facing urbanization. When this diversity is lost, a species loses its "options." A population may remain stable in census numbers ($N_c$) for generations, yet if it has been stripped of its rare alleles and heterozygosity, it becomes brittle, unable to pivot in response to the stochastic shocks of the Anthropocene.1

The scientific community has long theorized that the "Sixth Mass Extinction" is accompanied by a "Genetic Mass Extinction." Theoretical predictions based on the relationship between habitat loss and genetic variability suggested that at least 10% of genetic diversity might have already vanished from many plant and animal species.3 However, verifying this empirically has been a monumental challenge due to the lack of historical baselines and the difficulty of synthesizing disparate genetic datasets spanning decades.


1.2 The Paradigm Shift: From Demographic to Genomic Monitoring


Conservation biology is currently undergoing a profound paradigm shift. For decades, the discipline focused on maximizing the number of individuals (demography). The assumption was that if population numbers were kept high, genetic health would follow. We now know this to be dangerously simplistic. A population can be demographically recovered from a bottleneck of a few individuals—such as the Northern Elephant Seal—to tens of thousands, yet remain genetically monochromatic, vulnerable to a single viral strain that could bypass the identical immune defenses of every individual.

This realization has elevated the concept of Effective Population Size ($N_e$) over Census Population Size ($N_c$). $N_e$ describes the size of an idealized population (one that meets the assumptions of the Wright-Fisher model: random mating, constant size, no selection) that would experience the same rate of genetic drift as the actual population being studied. In almost all wild populations, $N_e$ is significantly smaller than $N_c$, often by an order of magnitude or more. This discrepancy means that a herd of 5,000 antelope might drift genetically as if it were a population of only 500, losing variation at an accelerated rate.4


1.3 The 2025 Watershed: Shaw et al.


The publication of the global meta-analysis by Shaw et al. in Nature (2025) marks a pivotal moment in this field. By synthesizing temporal genetic data from over three decades of research (1985–2019), covering 628 species across the tree of life, this study moves the conversation from theoretical alarm to empirical quantification. It provides the first robust global signal that within-population genetic diversity is actively eroding over timescales impacted by human activity.1

This report dissects the findings of this meta-analysis, not in isolation, but as the keystone of a broader theoretical edifice that includes the mathematics of spatial population genetics, the power laws of diversity-area relationships, and the practicalities of global policy implementation.


2. Technical Methodologies: Quantifying the Unseen



2.1 The Meta-Analytical Engine: Filtering the Noise


The robustness of the Shaw et al. findings rests on a rigorous systematic review protocol. The challenge in meta-analyzing genetic diversity is the extreme heterogeneity of the data. Genetic studies over the last 40 years have utilized a succession of marker technologies, from allozymes (proteins) to RFLP (Restriction Fragment Length Polymorphisms), microsatellites (Short Tandem Repeats), and finally, genome-wide Single Nucleotide Polymorphisms (SNPs).


2.1.1 The PRISMA Protocol


The researchers began with a massive corpus of 80,271 records identified through systematic literature searches. The filtration process, guided by the PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-Analyses) framework, was designed to isolate "temporal" studies—those rare gems that sampled the same population at different points in time, or compared historical samples (museum specimens) with contemporary ones.2

This rigorous culling resulted in a final dataset of 628 species. The taxonomic breadth is notable, though uneven:

  • Animals: 84.7% of the dataset (59.2% vertebrates, 25.5% invertebrates).

  • Plants: 12.7%.

  • Fungi: 1.9%.

  • Chromists: 0.6%.6

To handle the diversity of metrics (e.g., Expected Heterozygosity $H_e$ vs. Allelic Richness $A_r$), the authors utilized standardized effect size calculations, such as Hedge's $g$ or log-response ratios, allowing them to aggregate trends across different biological systems and genotyping technologies. The analysis was "agnostic" to the direction of change during extraction—meaning they did not cherry-pick studies showing decline, but included all studies reporting temporal metrics.2


2.2 Simulation Methodologies: The WFmoments Engine


Empirical data is often noisy and sparse. To understand the mechanisms driving the observed trends, modern conservation genomics relies heavily on simulation. A key innovation discussed in the context of this research is the WFmoments framework, developed by Spence and utilized by Exposito-Alonso's group.7


2.2.1 Beyond Individual-Based Models (IBMs)


Traditional simulations, such as those run in SLiM (Selection on Linked Mutations), are Individual-Based Models. They track every single gene copy in every individual across a landscape. While biologically realistic, they are computationally expensive, making it difficult to explore vast parameter spaces (e.g., testing 10,000 combinations of migration rates and habitat loss patterns).

WFmoments (Wright-Fisher Moments) takes a different mathematical approach. Instead of simulating individuals, it simulates the statistical moments of the allele frequency distribution.


2.2.2 The Mathematics of Moments


The dynamics of genetic diversity (specifically nucleotide diversity, $\pi$) are determined by the probability that two alleles sampled from the population are different. In a metapopulation (a group of connected subpopulations), this depends on the variance and covariance of allele frequencies between subpopulations.

WFmoments solves a linear system of Ordinary Differential Equations (ODEs) that describe how the first and second moments of allele frequencies evolve over time under forces of drift, migration, and mutation.8

The linear system can be conceptualized as follows:

The change in the second moment (which relates to homozygosity) in deme $i$ is a function of:

  1. Drift: Increases homozygosity within the deme (rate $\propto 1/2N_e$).

  2. Migration: Introduces alleles from neighbor $j$, reducing the difference between $i$ and $j$.

  3. Mutation: Introduces new variation (usually ignored in short-term ecological models or treated as negligible compared to drift).

By solving these ODEs, WFmoments can predict the trajectory of $\pi$ in complex, non-equilibrium scenarios—such as "edge contraction" where habitat is destroyed from the outside in—without the computational overhead of tracking individuals. This allows researchers to generate "pre-computed scenarios" or look-up tables to inverse-model observed genetic changes.9


2.3 Diversity Arrays Technology (DArT) and GDAR


In contexts where whole-genome sequencing is unavailable, techniques like Diversity Arrays Technology (DArT) provide a bridge. DArT is a microarray hybridization-based technique (and later DArTseq, a sequencing-based variant) that reduces genome complexity using restriction enzymes. It creates a representation of the genome, allowing for the interrogation of thousands of markers without a reference genome.11

This technology has been instrumental in generating the raw data for calculating Genetic Diversity-Area Relationships (GDAR) in crops and wild relatives (e.g., Vigna subterrenea and Brassica species), providing the empirical basis for the power laws discussed in Section 3.12


3. Theoretical Underpinnings: The Physics of Genetic Loss



3.1 The Theory of Island Biogeography Applied to Genomes


The fundamental theoretical advance in recent years is the unification of ecological theory (Species-Area Relationships, SAR) with population genetics. Just as the number of species scales with habitat area ($S = cA^z$), the amount of genetic variation within a species scales with the area it occupies.

However, genetic variation is not a single metric. It comes in two primary flavors that behave very differently under habitat loss:

  1. Allelic Richness ($A_r$): The total count of unique alleles in the population.

  2. Heterozygosity ($H_e$ / $\pi$): The probability that two randomly chosen alleles are different (or the average pairwise sequence divergence).


3.2 The Tale of Two Exponents: MAR vs. GDAR


Recent theoretical work has defined two distinct power laws governing these metrics, distinguished by their scaling exponent $z$.8


3.2.1 Mutation-Area Relationship (MAR)


  • Formula: $M \propto A^{z_{MAR}}$

  • Exponent: $z_{MAR} \approx 0.2 - 0.4$ (often centered around 0.3).

  • Biological Mechanism: Allelic richness is driven by the presence of rare alleles. Rare alleles are often spatially private (endemic to a specific subpopulation) or exist at very low frequencies. When habitat is destroyed, these rare variants are the first to vanish. Because they are numerous but spatially restricted, their loss scales rapidly with area loss. A $z$ value of 0.3 implies that losing 90% of habitat results in a loss of roughly 50% of the total alleles (since $0.1^{0.3} \approx 0.5$).14


3.2.2 Genetic Diversity-Area Relationship (GDAR)


  • Formula: $\pi \propto A^{z_{GDAR}}$

  • Exponent: $z_{GDAR} \approx 0.01 - 0.05$.

  • Biological Mechanism: Heterozygosity is dominated by common, intermediate-frequency alleles. These alleles are typically widespread across the species' range. When a specific patch of habitat is destroyed, these common alleles usually persist in the remaining patches. Therefore, heterozygosity is extremely resilient to the initial phases of habitat loss. It takes a massive reduction in area to significantly dent global heterozygosity. A $z$ value of 0.05 implies that even with 90% habitat loss, a species might retain nearly 90% of its heterozygosity ($0.1^{0.05} \approx 0.89$).8


3.3 The "Drift Debt" and Time Lags


The discrepancy between the high sensitivity of Allelic Richness ($A_r$) and the low sensitivity of Heterozygosity ($\pi$) creates a dangerous illusion of safety. A species may lose half its habitat and appear "genetically healthy" because its heterozygosity has barely moved. However, it has likely hemorrhaged a vast number of rare alleles (high MAR loss) that might have been critical for adaptation to future conditions (e.g., a rare heat-tolerance allele restricted to a warm-edge population that was destroyed).16

Furthermore, genetic loss is not instantaneous. It follows Non-Equilibrium Dynamics.

When a population is fragmented, it enters a state of "Drift Debt." The loss of heterozygosity is governed by the rate of decay:

$H_t = H_0 (1 - \frac{1}{2N_e})^t$

If $N_e$ is large (e.g., 10,000), the term $(1 - 1/20,000)$ is very close to 1, meaning the decay is glacial. The population carries a debt of genetic loss that will be "paid" over hundreds of generations as it drifts toward a new, lower equilibrium. The Shaw et al. meta-analysis, by finding a 6% loss already, suggests we are seeing only the tip of the iceberg—the early payments on a massive evolutionary debt.8


4. Recent Advances: The Empirical Reality (2024-2025)



4.1 Global Trends from the Shaw et al. Meta-Analysis


The Shaw et al. (2025) study provides the empirical anchor for these theoretical concerns. Analyzing the 628 species, the authors found a global signal of decline.

  • The 6% Figure: Across populations of 91 well-studied animal species, there was an average loss of 6% of genetic diversity over the past century. While this number might seem small, in the context of the GDAR power law ($z \approx 0.05$), a 6% loss in $\pi$ implies a catastrophic underlying loss of habitat and effective population size.3

  • Heterogeneity of Loss: The loss is not uniform. "Threats impacted two-thirds of the populations analyzed," but the response varied significantly by taxonomy.1


4.2 Taxonomic Disparities: The Mammal/Bird Crisis


The meta-analysis revealed a stark contrast between vertebrate classes:

  • Mammals and Birds (Aves and Mammalia): These groups showed the most severe and consistent declines in genetic diversity.

  • Fish and Insects: Groups like Actinopterygii (ray-finned fishes) and Insecta showed significantly less detectable loss, or even stability.6

Interpretation: This does not necessarily mean insects are safe. Rather, it reflects the mechanics of $N_e$. Insects and marine fish typically have massive effective population sizes (often $10^5$ to $10^7$). As established in Section 3.3, the rate of genetic drift is inversely proportional to population size ($1/2N_e$). A massive population loses diversity so slowly that a 30-year study window may fail to detect the signal, even if the population has crashed by 90% in census size. They are in the "lag" phase. Mammals and birds, with historically smaller populations, pay their drift debt faster, making the loss visible to current monitoring methods.


4.3 The Role of Conservation Interventions


Perhaps the most significant finding of the Shaw et al. study is the statistical validation of conservation interventions. The data showed that populations subject to active management—specifically translocations (moving individuals to simulate migration), connectivity restoration, and supplementation—did not follow the global trend of decline. In many cases, these populations maintained or even increased their genetic diversity.2

This finding transitions the field from "documenting the decline" to "prescribing the cure." It validates the theoretical prediction that artificial migration can offset the effects of fragmentation and drift.


5. Case Studies in Genetic Rescue: Mechanisms of Recovery


To understand how conservation halts genetic erosion, we must examine specific biological narratives highlighted in the recent literature. These cases illustrate the transition from the theoretical "vortex of extinction" to the practical "evolutionary rescue."


5.1 The Scandinavian Arctic Fox (Vulpes lagopus)


The Collapse: The Arctic Fox in Scandinavia (Sweden/Norway) was driven to the brink by overhunting for the fur trade in the early 20th century. Even after protection, the population failed to recover. It was trapped in a classic "Extinction Vortex": the small population size ($N_c < 100$) led to severe inbreeding. Inbreeding exposes deleterious recessive alleles (genetic load), reducing fitness (breeding success and juvenile survival), which further shrinks the population, increasing drift and inbreeding in a downward spiral.22

The Mechanism of Decline: By the early 2000s, the population was fragmented into four isolated subpopulations. Genetic analysis revealed a 25% loss of diversity over the last century and significant genetic structure ($F_{ST}$) between patches, indicating a complete cessation of gene flow.22

The Rescue: Conservationists implemented a two-pronged strategy:

  1. Ecological Support: Supplemental feeding and culling of the competitor Red Fox (Vulpes vulpes).

  2. Genetic Intervention: Release of captive-bred foxes and facilitating migration.The arrival of just three immigrant males in 2010/2011 (released from a captive breeding program) had a profound "Genetic Rescue" effect. These males introduced new alleles, masking the deleterious recessives fixed in the native population.The Outcome: The inbreeding coefficient ($F_{IS}$) declined by 43% within 5 years. The population size more than doubled. First-generation ($F_1$) offspring of the immigrants had 1.9x higher juvenile survival than inbred natives. This case is a textbook example of how restoring gene flow can jump-start demographic recovery.24


5.2 The Golden Bandicoot (Isoodon auratus)


The Context: Once widespread across mainland Australia, the Golden Bandicoot contracted to a few island refugia (e.g., Barrow Island) due to predation by invasive cats and foxes. Island populations are notoriously genetically depauperate due to the founder effect and isolation.

The Intervention: Translocations were undertaken to re-establish populations on the mainland (in fenced, predator-free exclosures) and other islands.

The Outcome: The Shaw et al. study highlights this as a success story. By carefully selecting source populations and managing the number of founders, conservationists successfully maintained high levels of genetic diversity in the new populations, preventing the severe bottlenecks usually associated with founder events.27


5.3 The Greater Prairie Chicken (Tympanuchus cupido)


The Vortex: This grouse species in North America suffered catastrophic habitat loss (conversion of prairie to agriculture). The Illinois population crashed to fewer than 50 birds in the 1990s. The result was a manifestation of "Inbreeding Depression" so severe it was visible: fertility rates dropped, and eggs failed to hatch. The population had lost allelic richness and was fixating deleterious alleles.27

The Rescue: Birds were translocated from large, genetically healthy populations in Kansas and Nebraska to Illinois.

The Outcome: The "genetic rescue" was immediate. Hatching success rebounded as genetic variation was restored. The population avoided immediate extinction, illustrating that demographic recovery is often impossible without genetic recovery.27


5.4 The Dusky Gopher Frog (Lithobates sevosus)


The Crisis: Restricted to a few ponds in Mississippi, this frog is critically endangered. Small population sizes led to low genetic diversity.

The Intervention: "Headstarting"—collecting egg masses, rearing tadpoles in protected tanks to metamorphosis, and releasing them—was combined with habitat management (prescribed burns to maintain open pine forests).

The Outcome: While headstarting is primarily a demographic tool (bypassing high larval mortality), when combined with managing multiple ponds, it stabilized the effective population size ($N_e$) and halted the erosion of diversity, allowing the population to begin a slow recovery.27


6. The Geography of Ignorance: Bias and the Global South



6.1 The Wallacean Shortfall in Genomics


Despite the "global" label of the Shaw et al. meta-analysis, the data reveals a profound geographic bias, often termed the "Wallacean Shortfall" (a lack of knowledge on the geographic distribution of species). The systematic review covered 141 countries, yet the density of data is heavily skewed toward the Global North (Europe and North America) and Australia/New Zealand.2

The Statistics of Bias:

  • Europe/North America: Dominant in long-term temporal genetic studies.

  • The Tropics (Global South): Significantly underrepresented.This bias is structural. Long-term genetic monitoring requires infrastructure—museum archives, continuous funding, and stable institutions—that has historically been concentrated in wealthy nations.


6.2 The Tropical "Blind Spot"


This bias is not merely a statistical nuisance; it is a biological crisis. Biodiversity is not distributed evenly; it follows a Latitudinal Diversity Gradient, peaking in the tropics.

  • High Sensitivity: Tropical species often have narrower thermal niches (Janzen's Hypothesis) and more specialized ecological interactions than temperate species.

  • Genetic Structure: Tropical mountains often generate high levels of "beta diversity" (differentiation between populations) due to climatic stratification.Implication: By basing our global estimates of genetic loss (e.g., the ~6% figure) largely on temperate species, we may be severely underestimating the global rate of loss. Temperate species, having recolonized the north after the last Ice Age, are often generalists with large ranges. Tropical specialists facing deforestation likely experience sharper, faster genetic erosion (higher $z_{MAR}$ exponents) than their temperate counterparts.29


7. Policy, Indicators, and the Future: Bridging the Gap



7.1 The Kunming-Montreal Global Biodiversity Framework (GBF)


The adoption of the Kunming-Montreal GBF marked a historic turning point. For the first time, genetic diversity was explicitly codified in international law under Goal A and Target 4.

  • Target 4: Calls for urgent management to "maintain and restore the genetic diversity within and between populations of all species" to maintain their adaptive potential.30


7.2 The Headline Indicator: $N_e > 500$


To operationalize this target, the GBF adopted a "Headline Indicator": The proportion of populations within species with an effective population size ($N_e$) > 500.4

This threshold is derived from the "50/500 rule" originally proposed by Franklin and Soulé in 1980:

  • $N_e = 50$: The minimum to avoid immediate inbreeding depression in the short term.

  • $N_e = 500$: The minimum to balance the loss of variation due to drift with the gain of variation due to mutation, thereby preserving long-term evolutionary potential.32


7.3 The Operational Challenge: The $N_e/N_c$ Ratio


The central challenge for nations implementing the GBF is that calculating $N_e$ usually requires DNA data, which is unavailable for most species. Consequently, the framework allows for the use of a "Census Proxy."

  • The Proxy: Estimate $N_e$ by multiplying the Census Size ($N_c$) by a ratio.

  • The Ratio: The default recommended ratio is 0.1 (i.e., $N_e \approx 0.1 \times N_c$). Therefore, a population is considered "safe" ($N_e > 500$) if it has more than 5,000 mature individuals.4

The Controversy: This 0.1 ratio is a blunt instrument.

  • Vertebrates: The ratio often ranges from 0.1 to 0.3.

  • High Fecundity Species (e.g., Fish/Trees): The ratio can be much lower ($10^{-3}$ to $10^{-5}$) due to "sweepstakes reproduction," where a few lucky individuals produce almost all the offspring.

  • Implication: Using the 0.1 proxy for a marine fish could lead to a catastrophic false sense of security, designating a population as genetically safe when it is actually drifting rapidly.5


7.4 Technological Bridges: Democratizing Genomics


To move beyond proxies and address the Global South data gap, technology is evolving.

  • Nanopore Sequencing: Recent studies (e.g., Gygax et al., 2025) have demonstrated the viability of "lab-in-a-backpack" approaches. Using portable Oxford Nanopore sequencers (MinION), researchers in remote locations (like Luambe National Park, Zambia) can perform end-to-end genetic monitoring—from DNA extraction to sequencing and analysis—without needing to export samples or access a mainframe. This circumvents the logistical and legal (Nagoya Protocol) hurdles of traditional genomics.34

  • Environmental DNA (eDNA): While traditionally used for species detection, eDNA is maturing into a population genetics tool. By sequencing nuclear markers from water or soil samples, researchers can estimate allele frequencies (and thus $\pi$ and $N_e$) without capturing the animals. This non-invasive approach is crucial for rare, elusive, or endangered species.37


Conclusion


The 2025 global meta-analysis by Shaw et al. serves as both a tombstone and a roadmap. It confirms that the "invisible extinction" of genetic diversity is a global reality, affecting two-thirds of monitored populations and ticking away as a "drift debt" that will inevitably lead to future fitness declines if left unchecked. The 6% average loss of diversity observed is merely the initial tremor of a larger earthquake, obscured by the time-lag inherent in genetic drift and the resilience of heterozygosity ($z_{GDAR}$) compared to the fragility of allelic richness ($z_{MAR}$).

However, the fatalism often associated with the biodiversity crisis is refuted by the data on conservation interventions. Genetic erosion is not an unstoppable physical force; it is a biological consequence of isolation and population reduction that can be arrested through connectivity, translocation, and habitat restoration. The cases of the Arctic Fox and the Greater Prairie Chicken prove that "genetic rescue" is not just theoretical—it is a practical, executable strategy.

The mandate for the post-2025 conservation era is clear: we must move beyond counting heads ($N_c$) and start managing the evolutionary potential ($N_e$) contained within them. By integrating the theoretical rigor of MAR/GDAR power laws with the operational targets of the GBF, and by deploying portable sequencing technologies to illuminate the blind spots of the Global South, we can halt the silent hemorrhage of the world's DNA. The cost of inaction is the permanent erasure of the library of life, leaving us with a world that is not only poorer in species, but frailer in its capacity to endure the future.


Table 1: Summary of Key Theoretical Frameworks in Conservation Genomics


Framework

Metric Modeled

Scaling Exponent (z)

Sensitivity to Habitat Loss

Biological Mechanism

MAR (Mutation-Area Relationship)

Allelic Richness ($A_r$), Segregating Sites ($M$)

High ($z \approx 0.2 - 0.4$)

High

Rare alleles are spatially localized or low-frequency; lost rapidly with area reduction.

GDAR (Genetic Diversity-Area Relationship)

Heterozygosity ($H_e$), Nucleotide Diversity ($\pi$)

Low ($z \approx 0.01 - 0.05$)

Low (Short-term)

Dominated by common alleles which persist in fragments; declines slowly until high habitat loss ($>90\%$) occurs.

WFmoments (Wright-Fisher Moments)

Variance/Covariance of Allele Frequencies

N/A (Dynamic Simulation)

Variable

Simulates the "lag" (drift debt) by solving ODEs for the first and second moments of frequency distribution over time.


Table 2: Comparative Conservation Outcomes from Genetic Interventions (Shaw et al. 2025)


Species

Threat

Intervention

Genetic Outcome

Demographic Outcome

Arctic Fox (Vulpes lagopus)

Inbreeding, Isolation

Translocation, Supplementation

$F_{IS}$ decreased 43%; diversity restored

Population doubled; survival increased

Golden Bandicoot (Isoodon auratus)

Invasive Predators

Translocation to predator-free sites

Maintenance of source diversity

Establishment of secure populations

Greater Prairie Chicken (T. cupido)

Habitat Loss, Inbreeding

Translocation of diverse individuals

Reduced inbreeding depression

Fertility rebounded (hatching success)

Dusky Gopher Frog (Lithobates sevosus)

Small $N_e$, Larval Mortality

Headstarting, Habitat Management

Stabilization of $N_e$

Slow population recovery

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