ZGSA Spring Research Symposium Abstracts 2018

Scientific Talks

Sam Straus (PhD Candidate, ​Avilés​ Lab)

Kleptoparasite loads on social spider colonies along an elevation gradient

Group living animals are likely to attract more parasites than solitary ones. Parasite loads, however, should also depend on
environmental conditions and on host characteristics and behaviors. Previous work has found that social spider colonies harbor
communities of kleptoparasitic spiders. We examined parasite loads and host hygiene behaviors in colonies of social and subsocial
spiders in the genus Anelosimus along an elevation gradient in eastern Ecuador. We found that kleptoparasite loads declined with
elevation. Host hygiene behaviors also declined with elevation, but within elevations species with more frequent hygiene behaviors
appeared more successful at keeping parasites at bay. Contrary to our predictions, parasite density declined with host nest and
colony size. The decline in parasite loads with elevation likely reflects a lower rate of energy exchange between colonies and their
environment, where colder temperatures mean fewer and smaller prey for colonies to process. The decline in parasite density with
host colony size likely reflects a declining surface area to volume ratio, and thus number of prey caught per capita, as the spiders’
three-dimensional webs grew in size. Social immunity, whereby a social host uses social behaviour to fight against parasites, is well
established in ants and humans, but in few species between. This study allows us to separately examine the effects of environment
and host characteristics on parasite loads of a social, but not eusocial organism.
 Keila Stark (Honours Student, O’Connor Lab)
Metacommunity archetypes driving epifaunal community structure in eelgrass meadows along the BC Coast

The abundance, diversity, and distribution of organisms across space are driven by some combination of dispersal, niche filtering,
and biotic interactions. Four theoretical metacommunity archetypes- neutral dynamics, patch dynamics, mass effects, and species
sorting- make different assumptions about the importance of these three processes in explaining patterns of abundance across
space. I investigated which metacommunity archetypes drive epifaunal invertebrate community structure in eelgrass (Zostera
marina L.) meadows in four bioregions within coastal British Columbia. Species ranges were inspected using "Elements of
Metacommunity Structure" analysis to determine whether dispersal is limiting. Variation partitioning was used to determine the
relative importance of spatial structure (influencing dispersal) and environmental variables (influencing niche filtering) in explaining
variation in species abundance across sites. I found that dispersal is not limiting, as several species representing multiple phyla and
dispersal modes were present at all sites from Haida Gwaii to the Southern Gulf Islands. Average nitrates, dissolved oxygen, sea
surface temperature and salinity best explained variation in community structure across all sites, however these variables were
spatially structured by region. The mass effects archetype- wherein high dispersal rates allow inferior competitors to maintain
populations at sites by migration- best fit my results, however stronger evidence on dispersal rates and interspecific interactions will
allow a more conclusive inference. My research provides insight on how diversity in BC eelgrass meadows is maintained, and how
community structure may respond to habitat fragmentation and climate change in the future.
 Freek de Haas (PhD Student, Otto Lab)
Interesting R- and K-selection dynamics in finite population sizes

R- and K-selection refers to a model of two competing types growing logistically and who differ in their intrinsic growth rate (R)
and/or carrying capacity (K). These types of selection are density dependent: R-selected types are favored in relatively small
populations while K-selected types are favored when the population size is close to carrying capacity. Consequently, K-selected
types dominate the equilibrium dynamics in deterministic models where population sizes are assumed to be large enough for drift
to be ignored. Here we examine a model of R- and K-selection in finite populations, where the benefit of an R-selected type in small
population sizes can be explored. We use numeric simulations and a spatially explicit individual based simulation to explore R- and
K-selection in finite populations and observe interesting dynamics at the wave front of these spreading populations. Our model
comes to drastically different conclusions with respect tot the fate of the R-selected type compared to previously studied models
that assume large population sizes and no drift.

Dr. Matthew Pennell

Rethinking comparative methods
 Emily Lim (Honours Student, ​Harley​ Lab)
Caprellid amphipods (Caprella spp.) are vulnerable to both physiological and habitat-mediated effects of ocean acidification

Ocean acidification (OA) is one of the most significant threats to marine life, and is predicted to drive important changes in marine
communities. Although OA impacts will be the sum of direct effects mediated by alterations of physiological rates and indirect
effects mediated by shifts in species interactions and biogenic habitat provision, direct and indirect effects are rarely considered
together for any given species. Here, we assess the potential direct and indirect effects of OA on a ubiquitous group of crustaceans:
caprellid amphipods (Caprella laeviuscula and Caprella mutica). Direct physiological effects were assessed by measuring caprellid
heart rate in response to acidification in the laboratory. Indirect effects were explored by quantifying caprellid habitat dependence
on the hydroid Obelia dichotoma, which has been shown to be less abundant under experimental acidification. We found that OA
resulted in elevated caprellid heart rates, suggestive of increased metabolic demand. We also found a strong, positive association
between caprellid population size and the availability of OA-vulnerable O. dichotoma, suggesting that future losses of biogenic
habitat may be an important indirect effect of OA on caprellids. For species such as caprellid amphipods, which have strong
associations with biogenic habitat, a consideration of only direct or indirect effects could potentially misestimate the full impact of
ocean acidification.
 Rhea Storlund (MSc Student, Trites Lab)
Steller hearts: An investigation into marine mammal cardiac function and anatomy

Marine mammal hearts deviate from the typical terrestrial mammalian pattern in that their hearts are dorsoventrally flattened,
tend towards a bifid apex, and have an enlarged aorta. Given the extensive cardiovascular responses to diving, consisting of
bradycardia and vasoconstriction, these morphological differences are likely related to cardiac function. However, this premise
relies on two unsubstantiated claims: first that marine mammal hearts function differently than their terrestrial counterparts, and
second that the observed anatomical differences have a functional significance. To address these knowledge gaps, I compare
marine mammal cardiac anatomy and function to terrestrial mammals. Previous examinations of marine mammal cardiac function
have used invasive techniques and have taken measurements during forced dives. Both of these conditions are known to affect the
parameters being measured, thereby making it difficult to interpret the results in terms of normal cardiac function. Instead, I use
the non-invasive techniques of electrocardiography (ECG) and echocardiography to study cardiac function and anatomy in
anaesthetized Steller sea lions (n=8). I measure cardiac dimensions, fractional shortening, blood flow, stroke volume, heart rate, and
characterize the ECG waveform. To identify differences in cardiac function and anatomy that exist in relation to diving, I compare
these parameters with pre-existing values for other species of both marine and terrestrial mammals from the literature. To my
knowledge, this is the first complete cardiac study of a marine mammal using these non-invasive techniques together, and the first
documented echocardiographic study of Steller sea lions. The results will be valuable in assessing the health of stranded sea lions as
well as for health checks in zoos, aquariums, and rehabilitation facilities. This baseline assessment will provide measurements of
normal cardiac function that can be related to cardiac anatomy and may provide insight into diving cardiovascular physiology that
would warrant further exploration.
 Le Tam Nhan Nguyen​ (MSc Student, Pante Lab)
Characterization of the nucleoprotein’s nuclear localization signal function during influenza A virus infection

Replication of the genome of influenza A virus, which consists of eight viral ribonucleoprotein complexes (vRNPs), occurs in the
nucleus of infected cells. The nuclear import of vRNPs is mediated by nuclear localization signals (NLSs) on the viral nucleoprotein
(NP) of the vRNPs. NP contains two NLSs, a well-studied NLS1 at the N-terminus and a newly identified NLS2 in the middle of the
protein. ​Through site-directed mutagenesis and functional analysis, we found that NLS2 is a non-classical monopartite ​with the
sequence 212-GRKTR-216 in NP from seasonal influenza A virus strains. However, as with other NLSs, there may be variation in the
basic residues of NLS2, which may affect the nuclear import of vRNPs and infection​. Using bioinformatic analysis, ​we found that all
variants of the NLS2 are present in NP and other proteins from different influenza virus strains and even in other non-influenza viral
proteins​. ​To further study the contribution of the NLSs to the nuclear import of the viral genome, we have successfully generated
recombinant viruses containing mutations within NLS1 or NLS2 using reverse genetics. To investigate how these mutations inhibit
viral infection, tissue culture cells were infected with the wild-type or NLS mutant viruses and the infection was followed using
confocal microscopy after immunostaining of viral NP over a time course. ​Reverse genetic-derived NLSs mutant viruses ​were still
able to establish infection, but at a slow rate in comparison to wild-type viruses. NP was transported into the nucleus of the mutant
viruses-infected cells during the early stage of the viral life cycle, but the assembly or release of progeny virus from the cell was
impeded at 18 h and 24 h post infection. Based on these results, we propose that mutations of NLS1 or NLS2 could potentially
inhibit vRNP assembly; thereby, also inhibiting the nuclear export of vRNPs.
 Jacelyn Shu​ (MSc Student, Brauner Lab)
Enhanced hemoglobin-O2 unloading in teleosts is sustained after catecholamine removal

Recent findings indicate that teleost fishes may be able to greatly enhance hemoglobin-oxygen (Hb-O​2​) unloading to tissues under
stress. With the release of circulating catecholamines, red blood cell (RBC) b-adrenergic Na​+​/H​+​ exchangers (b-NHE) are activated,
protecting RBC pH and building a H​+​ gradient across the RBC membrane. At the tissues, plasma-accessible carbonic anhydrase (CA)
short-circuits this H​+​ gradient, reducing RBC pH and enhancing Hb-O​2​ unloading. Previous studies show that in rainbow trout, Hb-O​2
unloading may double with no change in perfusion compared to a system without short-circuiting. However, catecholamines in vivo
may be removed within minutes of release, potentially restricting the benefits of enhanced Hb-O​2​ unloading to a narrow time
frame. Here, we show that this system remains active following experimentally removing catecholamines from the blood: adding CA
to b-adrenergically stimulated RBCs in vitro caused a large increase in PO​2​ that may mirror enhanced Hb-O​2​ unloading to tissues in
vivo. Even an hour after catecholamine removal, 50% of the maximal ∆PO​2​remained, indicating that the system can be sustained in
the absence of circulating catecholamines. This system may therefore play an important role in supplying tissues with O​2​ not only
during short stressful periods, but also in the time following.

Luis Camacho (PhD Student, ​Avilés ​Lab)

Ants shape predation rate patterns along elevational gradients in the tropical Andes

Macroecological studies suggest reduced predation rates at higher elevations and latitudes, but the mechanisms behind these
patterns remain unaddressed. We focused on the equatorial Andes to test if changes in predation across elevations correlate with
changes in the contribution of different predators, their abundances, and efficiencies. We sampled 4000-meter elevational
gradients on the western and eastern slopes and measured predation rates using live baits. We found predation rates decreased
with elevation. This was mainly driven by reductions in ant predation whose relative importance also decreased. Predator density
and efficiency appeared to be important mechanisms driving predation rates. Our results agree with known general patterns of
predation, and suggest temperature and net primary productivity as their potential primary drivers.
 Shuang Yang (PhD Candidate, Pante Lab)
The underlying mechanism of a unique nuclear entry pathway used by the parvovirus minute virus of mice

Parvoviruses are small, non-enveloped, single-stranded DNA viruses that replicate in the nucleus of the host cell. Rodent
parvoviruses are not only able to preferentially target rapidly dividing cancer cells, but also possess oncolytic properties. They have
thus been evaluated as potential anti-cancer agents, either as unmodified infectious virus, attenuated mutants, or vectors. For the
development of such parvoviruses-based therapy, we need to better characterize their life cycle, particularly their nuclear entry
pathway. Our laboratory has previously found that the parvovirus minute virus of mice (MVM) and other parvoviruses enter the
nucleus by a unique mechanism that involves small, transient disruptions of the nuclear envelope (NE) and partial dismantling of the
underlying nuclear lamina. We are now investigating the molecular mechanism by which MVM disrupts the NE. Since
N-myristoylation of different viral proteins has been reported to play roles in virus-cellular membranes interaction for many other
non-enveloped viruses, and capsid proteins of MVM have putative N-myristoylation sites, we have been investigating the
involvement of protein N-myristoylation in the nuclear entry mechanism of MVM. We found that the protein N-myristoylation
inhibitor 2-Hydroxymyristic acid (HMA) significantly prevented viral gene expression in MVM-infected mouse fibroblast cells. More
importantly, when added at different time point post-infection, the inhibitory effects of HMA on viral gene expression coincided
with the time of the nuclear entry of MVM. To further illustrate the role of protein N-myristoylation during this process, we used
semi-permeablized HeLa cells in which the influx of TRITC-dextran into the nucleus is an indicator for MVM-induced NE
disruption. With this system we further found that when protein N-myristoylation was blocked by HMA, MVM-induced NE
disruption was also inhibited. Therefore, we propose that protein N-myristoylation plays an important role in the mechanism of
MVM-induced NE disruption during the nuclear entry of MVM.

Dr. Stilian Louca

Unraveling bacterial diversification through geological time with massive phylogenies
 Jordan Hamden (PhD Candidate, Soma Lab)
11-dehydrocorticosterone, a corticosterone metabolite, in mice, rats, and birds

Glucocorticoids (GCs) are steroid hormones secreted by the adrenal glands into the blood to coordinate multiple organ systems. In
addition, lymphoid tissues, such as bone marrow, thymus, and spleen, locally produce GCs to regulate lymphocyte selection and
immune responsiveness. Corticosterone is metabolized to the inactive 11-dehydrocorticosterone (DHC) within tissues by the
enzyme 11β-HSD2, and corticosterone is regenerated from DHC by 11β-HSD1. Recent data demonstrate that regeneration of
corticosterone from DHC plays a major role in local elevation of corticosterone levels in mouse lymphoid tissues. However, there
are very little data on DHC levels in blood or tissues of mice. Furthermore, it is well documented that GCs increase in response to
stressors, but changes in DHC levels after stressors have never been measured in any animal. We developed an immunoassay to
measure DHC levels in the circulation and tissues. In mice, we measured corticosterone and DHC levels in whole blood, bone
marrow, thymus, and spleen at postnatal day 5 (PND5), PND23, and PND90. Overall, corticosterone levels were greater than DHC
levels. At PND5, corticosterone was locally elevated in bone marrow, thymus, and spleen, and DHC was locally elevated in bone
marrow, but locally reduced in thymus. In the bone marrow, the local elevation of both corticosterone and DHC might be the result
of local de novo synthesis from cholesterol or sequestration of DHC from the blood. In the thymus, the local elevation of
corticosterone and the local reduction of DHC may indicate that DHC is being regenerated into corticosterone. For both PND23 and
PND90, corticosterone was higher in blood
than all tissues. DHC was locally elevated in the bone marrow and thymus at PND23, and in the bone marrow at PND90.
Interestingly, blood corticosterone was lowest at PND5, while blood DHC was highest at PND5. We also measured corticosterone
and DHC in rats and birds before and after restraint stress. Circulating corticosterone and DHC increased after restraint stress, but
corticosterone had a greater increase than DHC. Additionally, corticosterone levels were higher than DHC levels for both rats and
birds before and after restraint. Future studies will determine the sources of circulating and local DHC. These data are a crucial first
step for understanding how various tissues locally regenerate GCs from metabolites.
 Matt Osmond (PhD Student, Otto Lab)
Predicting the types of mutations that rescue populations from extinction

Populations exposed to harsh environmental changes (such as drug treatment, herbicides, or pollutants) often go extinct. However,
resistance sometimes evolves fast enough to rescue populations and promote persistence. There is now over 20 years of theory
investigating how the probability of evolutionary rescue depends on the characteristics of available genotypes, but we still don't
know how evolutionary rescue affects the characteristics of genotypes that contribute to persistence. Meanwhile, there is a large
literature devoted to understanding the characteristics of genotypes that contribute to adaptation in the absence of demography,
the distribution of fitness effects (DFE) along an adaptive walk being of central importance. In this talk I will explain how we have
added demography to Fisher's geometric model to predict the DFE of rescued populations, i.e., looking at only those adaptive walks
that end in persistence. This approach suggests the genetic pathways by which evolutionary rescue will occur, illustrates the
importance of sex and recombination in adaptation to changing environments, and begins a line of inquiry into the genetic
signatures of evolutionary rescue that may be identified in the genomes of natural populations.
 Sarah Fortune (PhD Candidate, Trites Lab)
Drones, time-depth recorders and prey sampling reveal complex foraging behaviour in bowhead whales.

As zooplanktivorous predators, bowhead whales (​Eubalaena mysticetus​) must routinely locate prey patches of sufficient energetic
density to meet their metabolic needs. However, little is known about Eastern Canada-West Greenland bowhead whale foraging
behaviour and the quality and quantity of their prey resource in Canadian waters. We used a new approach to study bowhead
feeding ecology that included: 1) multi-scale biologging to record underwater feeding behaviour using long-term SPLASH tags
equipped with satellite telemetry-linked time-depth recorders (TDRs) (n=10) and short-term fine-scale archival TDRs (n=5); 2)
Unmanned Aerial Systems (i.e., drones) to document behaviours not previously seen during focal follows; and 3) vertical prey
collection at discrete depths using an Optical Plankton Counter (OPC), multi-frequency echosounder (n=74) and conical mesh nets
(n=28). Tag attachments ranged from 3.19 – 12.7 hours for fine-scale TDRs and for several days for long-term SPLASH tags. Analysis
of video showed that the whales were not feeding at the surface during daytime (0-20 m), while the TDR data indicated that they
were conducting principally (~68%) long and deep square-shaped foraging dives (136.66 m ± 135.72 SD). Interestingly, the whales
were also routinely conducting shorter and shallower, u-shaped dives (~26%) suggesting that they were also exploiting shallowly
aggregated prey (67.83 m ± 84.81 SD). Vertical prey sampling revealed two prey layers comprised of almost exclusively calanoid
copepods at depths that correspond to bowhead whale dive depths. Zooplankton biomass was dominated by lipid rich, Arctic
copepods (e.g., ​Calanus hyperboreus ​and ​C. glacialis​). Our finding contrasts with previous feeding ecology studies that found whales
consistently dove to depth to feed and did not alternate between deep and shallow feeding strategies. Combining drones with TDRs
and prey sampling showed a more complex feeding ecology then previously understood and provides a means to evaluate the
energetic balance of individuals under current environmental conditions.
 Poster Session
Aaron Purdy (MSc Student, Trites Lab)
Rethinking the aerobic dive limit

The diving capacity of marine mammals is defined by their aerobic dive limit (ADL), which is the duration of time they may remain
submerged using only their on-board oxygen stores to create energy. This cornerstone concept of diving physiology is often
presented as a distinct switch between aerobic and anaerobic metabolic pathways at a given threshold, and is presumed to be
indicated through sudden lactate emergence and proportional increases in recovery time for gas homeostasis. However, few studies
have empirically tested this hypothesis in freely diving marine mammals. Our study investigated the pattern of lactate emergence as
well as oxygen and carbon dioxide recovery times in a group of freely-diving trained Steller sea lions (​Eumetopias jubatus​). We
collected voluntary blood samples post-dive from 4 adult females after dives to 40m for varying duration (1-7 min) to determine
blood lactate levels. We also measured the time for oxygen and carbon dioxide to reach pre-dive levels using flow-through
respirometry. Contrary to expectations, lactate levels demonstrated a progressive increase with dive time both before and after the
calculated ADL (cADL), with significant levels evident even following very short dives. Oxygen and carbon dioxide recovery increased
at a steady rate in proportion to increased dive time, with no apparent inflexion point—with CO​2​ recovery rates proving the limiting
factor. With a steady increase in blood lactate levels and no distinct change in recovery time at the cADL, our results suggest that
anaerobic metabolism is occurring long before the body’s total oxygen stores have been consumed. This lends support to the idea
that there is not a distinct switch between aerobic and anaerobic pathways at a given threshold. Our findings related to how marine
mammals manage oxygen should be considered—particularly when incorporating physiological costs and constraints into models of
diving physiology and optimal foraging.
 Ailene MacPherson​ (PhD Student, Otto Lab)
Density-dependent selection in finite populations

Density-dependent selection, for example "r" and "K" selection, is of long standing interest in light of the evolution of life history
strategies. Numerous mathematical models of have explored when "r" selected or "K" selected alleles should be favoured by
natural selection, how allele frequency dynamics are influenced by the genetic architecture of the "r" and "K" traits, and how
environmental variability effects the preferred life-history strategy. To date, however, these models have relied on the assumption
that populations sizes are large enough that the effect of genetic drift is minimal. Relaxing this assumption we explore the effect of
finite population size and drift on the evolution of life history strategy within a population. Extending these single population
results to meta-populations we show that density-dependent selection can have important impacts on migration and expansion
load as well as the probability of "mutational meltdown".

Francisco Henao Diaz (PhD Student, Pennell Lab)

Young clades appear to be diversifying faster: meta-analysis of speciation rates reveals ubiquitous age-dependent effects

Variation in speciation rates is frequently invoked to explain disparity in species richness across clades, space, ecological gradients,
and time. Consequently, many studies have estimated rates of speciation from both molecular phylogenies and the fossil record.
However, there has not been a comprehensive examination of the actual rates of diversification that have been estimated; such a
meta-analysis would be useful because it would provide a baseline for comparison to rates computed from specific groups (e.g.,
“rapid radiations”) and rates estimated from other types of data.. To fill this gap, we estimated many empirical previously published
phylogenies and then examined whether these rates were correlated to attributes of the clade. In addition to estimating the
distribution of speciation rates, we found a persistent pattern that younger clades always appear to be diversifying faster (i.e., rates
of speciation were negatively age-dependent). We used simulations and subsampling of empirical datasets to investigate the causes
and consequences of this anomaly. Through this analysis we uncovered a previously unrecognized artifact that may have
implications for how we interpret many macroevolutionary patterns.
Heat​her Reid (Directed Studies Student, ​Harley Lab)
Cold tolerance and winter microhabitat use of an intertidal snail, ​Littorina scutulata

Climate change is often considered synonymous with extreme high temperatures, and most research has focused on species’ upper
thermal limits. However, changes at the cold end of the temperature range will be equally important as species move towards the
poles, up slope, and to deeper depths. Intertidal marine species already live near the edge of their thermal limits, in large part due
to daily exposure to air. The cold tolerance of intertidal species is a relatively understudied area, as is the use of microhabitats in
winter. Many microhabitats are available to intertidal invertebrates, including rock bottoms and underneath algae, and these
microhabitats may play an important role in invertebrate survival by providing thermally buffered refugia. We are examining the
cold tolerance of an intertidal snail (​Littorina scutulata​) through exposure trials. Early results indicate that activity of ​L. scutulata ​is
high at air temperatures down to -3°C, but movement is restricted past 3 hours at -6°C and any time at -9°C. Temperature loggers
were deployed to determine the temperature differences between microhabitats and their surroundings, while lab experiments
have been conducted to determine the microhabitat use of ​L. scutulata ​at different temperatures. Early results indicate that below
0°C, snails more likely to be found under algae or underneath a rock than they are at 0°C or 10°C, and snails are more active after
exposure to low temperatures when under ​Fucus​. This research is expected to provide important information on microhabitat use
and the effects of cold exposure on intertidal invertebrates in winter.
 Kaleigh Davis (PhD Student, ​O’Connor Lab)
Metabolic scaling of biological nitrogen fixation

Metabolic scaling theory (MST) describes how the temperature dependence of highly conserved metabolic processes, i.e.
photosynthesis and respiration, drives the rates of higher order biological processes. While MST has done a good job of predicting
multi-scale patterns using temperature, it has largely ignored the role of temperature-resource interactions in regulating these
metabolic processes. Here, we investigate the temperature dependence of biological nitrogen fixation (BNF), a third highly
conserved metabolic process that provides essential raw materials for photosynthesis. Using a meta-analysis, we test the hypothesis
that the temperature-dependent nitrogenase pathway that underlies BNF (E​a​ = ~1.5eV) confers a general temperature dependence
on BNF that is different from that of respiration and photosynthesis. We will then use a mesocosm experiment to test the
hypothesis that this thermal asymmetry influences community composition and productivity over temperature gradients.
Quantifying the temperature dependence of nitrogen fixation within the framework of MST will allow us to better understand the
biochemical underpinnings of community-level processes and predict how they might change with climatic warming.
 Silu Wang (PhD Candidate, Irwin Lab)
Admixing aggression genes through a moving warbler hybrid zone

The rapid speciation of Setophaga warblers is an interesting puzzle for understanding divergence and speciation. We can gain
insights of such rapid speciation from hybrid zones between young Setophaga sister pairs. One of the hybrid zones between S.
townsendi and S. occidentalis in the Pacific Northwest was thought to be moving over time, based on the shift of plumage hybrid
indices at the same sites over time. Such movement might lead to the fusion of gene pools between species in such rapidly moving
and extensively hybridizing taxa. We found genome-wide Fst between allopatric parental populations indeed decayed over time. A
genome-wide association study (GWAS) was conducted to detect genetic regions underlying plumage variations in the genome. We
found that although the aggression and quality-associated genetic regions demonstrated more differentiation between parental
populations than the rest of the genome, such differentiation decayed over time. The quality-associated genetic regions decayed
more than the rest of the genome. This study sheds light on population genetic cause and consequences of a moving hybrid zone,
which could influence divergence in this young clade.