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Geology topics

Julie A. Savidge

Publications and source records attributed to Julie A. Savidge.

At least 19 recordsLinked to original sources

Demographic responses to experimental manipulation of an enclosed population of invasive brown treesnakes (Boiga irregularis)

Understanding how invasive species populations respond to perturbations can give insight into effective management. However, opportunities to test theoretical population responses are rare, as experimental perturbations often run counter to management goals (e.g., suppression). On the island of Guam, a 5-ha outdoor enclosure (with no immigration or emigration) constructed in 2003 serves as a unique population-scale study area for invasive brown treesnakes ( Boiga irregularis ). Capture-mark-recapture studies (CMR) and several experiments have taken place since 2004. Here, we analyze a subset of the CMR data collected between 2008 and 2012, a time period encompassing two experimental perturbations: (1) supplemental feeding to determine if the population is food limited and (2) targeted removal of adult and large-subadult cohorts to quantify the timing and magnitude of the population's demographic response to a simulated control effort. We used the CMR data to fit a Pradel model in a Bayesian framework to compare the population growth rate, survival probability, and recruitment rate prior to, during, and post perturbations. We found that increasing food resources increased population growth and recruitment rates, but that survival remained relatively constant. The removal perturbation resulted in a decrease in population growth rate and survival immediately after the removal, but population growth rate and recruitment began increasing 1 year after the removal perturbation. Our results suggest that brown treesnakes in the enclosed population are prey limited, likely constraining population growth, and that the suppression of population growth after removal of adults and large sub-adults is short-lived (i.e., less than a year).

Guam

Reproductive condition of an invasive snake in urban, savanna, and forest habitats

Predation by Brown Treesnakes ( Boiga irregularis ) has caused the decline or extinction of all native forest vertebrates on Guam, including birds, lizards, and bats. Loss of the highest-quality prey has caused shifts in Brown Treesnake size and life history. We sought to understand how reproductive condition varies among habitats island-wide, particularly comparing urban and savanna habitats (where some primarily non-native large prey remain) with prey-depauperate forest habitats. We dissected 100 snakes from each of 18 sites stratified by 6 habitat types and calculated a body condition index, relative follicle size (for females; N = 817) and relative testes size (for males; N = 992) for each. Low detection of reproductive females continues to hamper research on their biology, as only 11% of females had secondary vitellogenic follicles (N = 89). Although variation among sites within habitats was high, above-average reproductive development of both males and females was most frequently observed in urban and savanna habitats, emphasizing the potential for prey subsidies in these habitats to fuel population growth and affect invasive snake management objectives. Furthermore, the trade-off between body condition and reproductive condition appeared to be more severe in forested habitats and less severe in urban habitats for both males and females. There were few obvious seasonal patterns, although there was a slightly lower probability of females having secondary vitellogenic follicles in the wet season compared to the dry season.

NeoBiota

Growth rate variation in Brown Treesnakes (Boiga irregularis): An invasive species of conservation concern

Somatic growth rate is a fundamental trait that influences metabolism, lifespan and reproductive maturity and is critical for understanding population dynamics and informing management actions. Brown Treesnakes ( Boiga irregularis ) introduced to Guam are highly invasive and can reproduce year-round without discrete cohorts. We compared snake size trajectories described by the conventionally used von Bertalanffy growth function versus the Gompertz model. Using quantile regression with a regularized effect for individual snakes we modeled growth rates of 270 marked, wild snakes as a function of size. The Gompertz model explained more of the variation in growth and rendered more realistic predictions of asymptotic sizes than did the von Bertalanffy model. With the Gompertz model, growth rates were 1.05–1.16× faster in males than in females. Females reached asymptotic sizes at shorter snout-vent lengths than males. Growth rate was positively correlated with amount of precipitation, and modeling wet-dry seasonality on Guam as a sinusoidal function identified a growth peak in September—October. Effects of seasonality and precipitation, however, were minor compared to individual and sex related differences in size-adjusted growth rates. We estimated that the 50th (and 5th, 95th) growth-rate percentile males in our study population become sexually mature at an age of 33 (∞, 15) months, while females mature at 41 (∞, 18) months, where ∞ indicates that the slowest growing snakes never reach maturity. However, 50% of the snakes mature at a size below the median, and age at maturity may be as low as 10.4 (males) and 13.7 (females) months for average-sized hatchlings that grow fast. Our results have implications for the timing of management options for this species and our approach can be broadly applied to animals where repeated growth data are obtained and age is unknown.

Ecology and Evolution

Trait heritability and its implications for the management of an invasive vertebrate

Control methods that target specific traits of an invasive species can produce results contrary to the aims of management. If targeted phenotypes exhibit heritability, then it follows that the invasive species could evolve greater resistance to the applied control measures over time. Additional complications emerge if those traits targeted by control are also inversely related to reproductive success. Given this, prudent considerations for invasive species management are to quantify the heritability of traits selected through control measures and gauge their relationship with reproductive success. Herein we provide a case study utilizing long-term field data and a multi-generational pedigree of an experimentally-closed population of brown treesnakes (N = 426; Boiga irregularis ) on Guam. We employed an “animal model” to estimate the narrow-sense heritability ( h 2 ) for annual body condition, a trait related to both susceptibility to a primary tool used for brown treesnake control (i.e., live-lure traps) and annual reproductive success. Annual body condition displayed significant heritability [ h 2 = 0.149 (95% highest posterior density interval: 0.059–0.220)]. Considering a negative effect of body condition on susceptibility to trap capture but positive effect on reproductive success, significant heritability of body condition suggests the potential for live-lure traps to lose efficacy over time while also eliciting an undesirable effect on brown treesnake fecundity. Our results highlight the potential for negative repercussions that can stem from management actions, while also serving to underscore the evolutionary implications that are often overlooked but subsumed within invasive species control.

Biological Invasions

Genomic pedigree reconstruction identifies predictors of mating and reproductive success in an invasive vertebrate

The persistence of an invasive species is influenced by its reproductive ecology, and a successful control program must operate on this premise. However, the reproductive ecology of invasive species may be enigmatic due to factors that also limit their management, such as cryptic coloration and behavior. We explored the mating and reproductive ecology of the invasive Brown Treesnake (BTS : Boiga irregularis ) by reconstructing a multigenerational genomic pedigree based on 654 single nucleotide polymorphisms for a geographically closed population established in 2004 on Guam ( N = 426). The pedigree allowed annual estimates of individual mating and reproductive success to be inferred for snakes in the study population over a 14‐year period. We then employed generalized linear mixed models to gauge how well phenotypic and genomic data could predict sex‐specific annual mating and reproductive success. Average snout–vent length (SVL), average body condition index (BCI), and trappability were significantly related to annual mating success for males, with average SVL also related to annual mating success for females. Male and female annual reproductive success was positively affected by SVL, BCI, and trappability. Surprisingly, the degree to which individuals were inbred had no effect on annual mating or reproductive success. When juxtaposed with current control methods, these results indicate that baited traps, a common interdiction tool, may target fecund BTS in some regards but not others. Our study emphasizes the importance of reproductive ecology as a focus for improving BTS control and promotes genomic pedigree reconstruction for such an endeavor in this invasive species and others.

Ecology and Evolution

Contact rates with nesting birds before and after invasive snake removal: Estimating the effects of trap-based control

Invasive predators are responsible for almost 60% of all vertebrate extinctions worldwide with the most vulnerable faunas occurring on islands. The brown treesnake ( Boiga irregularis ) is a notorious invasive predator that caused the extirpation or extinction of most native forest birds on Guam. The success of avian reintroduction efforts on Guam will depend on whether snake-control techniques sufficiently reduce contact rates between brown treesnakes and reintroduced birds. Mouse-lure traps can successfully reduce brown treesnake populations at local scales. Over a 22-week period both with and without active snake removal, we evaluated snake-trap contact rates for mouse- and bird-lure traps. Bird-lure traps served as a proxy for reintroduced nesting birds. Overall, mouse-lure traps caught more snakes per trap night than did bird-lure traps. However, cameras revealed that bird-lure traps had a snake contact rate almost 15 times greater than the number of successfully captured snakes. Snakes that entered bird-lure traps tended to be larger and in better body condition and were mostly captured in bird-lure traps, despite numerous adjacent mouse-lure traps. Traps placed along grid edges caught more snakes than interior traps, suggesting continuous immigration into the trapping grid within which bird-lure traps were located. Contact between snakes and bird-lure traps was equivalent before and after snake removal, suggesting mouse-lure traps did not adequately reduce the density of snakes that posed a risk to birds, at least at the timescale of this project. This study provides evidence that some snakes exhibit prey selectivity for live birds over live mouse lures. Reliance on a single control tool and lure may be inadequate for support of avian reintroductions and could lead to unintended harvest-driven trait changes of this invasive predator.

NeoBiota

Do observer fatigue and taxon-bias compromise visual encounter surveys for small vertebrates?

Context. Visual encounter surveying is a standard animal inventory method, modifications of which (e.g. distance sampling and repeated count surveys) are used for modelling population density. However, a variety of factors may bias visual survey counts. Aims. The aim of the present study was to evaluate three observer-related biases: (1) whether fatigue compromises detection rate as a survey occasion progresses; (2) whether long-term fatigue or boredom compromise detection rates over the course of a survey period; and (3) whether observers exhibit biases in detection rates of different animal taxa. Methods. We analysed >2.3 × 10 4 observations of lizards and small mammals from nocturnal pedestrian visual encounter surveys, each 4 h in duration, conducted by a pool of 29 observers, each of whom surveyed for up to 31 nights. Key results. Detections of sleeping (diurnal) emerald tree skinks ( Lamprolepis smaragdina ) exhibited a small but statistically verified decline as the evening progressed, whereas detections of sleeping (diurnal) green anoles ( Anolis carolinensis ) increased as the evening progressed. Detections of nocturnal geckos (several species pooled) showed a weak and non-significant declining trend. Small mammal sightings (rats, shrews and mice pooled) declined strongly over the course of an evening. The participants saw greater or equal numbers of animals the more nights they surveyed. Most participants exhibited statistically significant, and often strong, taxonomic detection bias compared with the pool of peer observers. The skills of some observers appeared to be consistently above average; others consistently below average. Conclusions. Data on sleeping lizards suggest that neither short-term nor long-term observer fatigue is of much concern for 4-h visual searches. On the contrary, differences among observers in taxonomic bias and overall detection skills pose a problem for data interpretation. Implications. By comparing temporal detection patterns of immobile (e.g. sleeping) with actively moving animal taxa, sampling biases attributable to searcher fatigue versus the animals’ circadian rhythm can be disentangled and, if need be, statistically corrected for. Observer skill differences and observer-specific taxonomic biases may hamper efforts to statistically evaluate survey results, unless explicitly included as covariates in population models.

Wildlife Research

Optimizing walking pace to maximize snake detection rate: A visual encounter survey experiment

Visual encounter survey efforts can be defined and constrained by duration, distance, or both duration and distance simultaneously. This study examines the optimal walking pace that will maximize the number of animal detections within a limited time frame. We predicted that animal sighting rate per unit of distance would decline with increasing pace, but that maximal sighting rate per unit of time would occur at some intermediate pace because walking faster means covering more ground and passing more animals that could potentially be detected. We conducted a controlled experiment during which we searched for Brown Treesnakes ( Boiga irregularis ) while walking 220-m-long transects at three different speeds. For transects inside the forest, detection rate per unit distance decreased (nonlinearly) with increasing pace. When considering catch per unit time, however, we found 5% more snakes at the medium pace compared with the slow pace, and 63% more snakes at the fast pace compared with the slow pace. The pattern appeared different for surveys along forest edges with higher vegetation density, in which snakes might be more difficult to spot. Surprisingly, pace had no detectable effect on the body sizes or perch heights of snakes successfully detected. Finding the optimal search pace for a particular study organism in a focal habitat has the potential to substantially increase survey cost-efficacy.

Herpetologica

Passive restoration following ungulate removal in a highly disturbed tropical wet forest devoid of native seed dispersers

Overabundant ungulate populations can alter forests. Concurrently, global declines of seed dispersers may threaten native forest structure and function. On an island largely devoid of native vertebrate seed dispersers, we monitored forest succession for 7 years following ungulate exclusion from a 5-ha area and adjacent plots with ungulates still present. We observed succession from open scrub to forest and understory cover by non-native plants declined. Two trees, native Hibiscus tiliaceus and non-native Leucaena leucocephala , accounted for most forest regeneration, with the latter dominant. Neither species is dependent on animal dispersers nor was there strong evidence that plants dependent on dispersers migrated into the 5-ha study area. Passive restoration following ungulate removal may facilitate restoration, but did not show promise for fully restoring native forest on Guam. Restoration of native forest plants in bird depopulated areas will likely require active outplanting of native seedlings, control of factors resulting in bird loss, and reintroduction of seed dispersers.

Guam

Quantile regression of microgeographic variation in population characteristics of an invasive vertebrate predator

Localized ecological conditions have the potential to induce variation in population characteristics such as size distributions and body conditions. The ability to generalize the influence of ecological characteristics on such population traits may be particularly meaningful when those traits influence prospects for successful management interventions. To characterize variability in invasive Brown Treesnake population attributes within and among habitat types, we conducted systematic and seasonally-balanced surveys, collecting 100 snakes from each of 18 sites: three replicates within each of six major habitat types comprising 95% of Guam’s geographic expanse. Our study constitutes one of the most comprehensive and controlled samplings of any published snake study. Quantile regression on snake size and body condition indicated significant ecological heterogeneity, with a general trend of relative consistency of size classes and body conditions within and among scrub and Leucaena forest habitat types and more heterogeneity among ravine forest, savanna, and urban residential sites. Larger and more robust snakes were found within some savanna and urban habitat replicates, likely due to relative availability of larger prey. Compared to more homogeneous samples in the wet season, variability in size distributions and body conditions was greater during the dry season. Although there is evidence of habitat influencing Brown Treesnake populations at localized scales (e.g., the higher prevalence of larger snakes—particularly males—in savanna and urban sites), the level of variability among sites within habitat types indicates little ability to make meaningful predictions about these traits at unsampled locations. Seasonal variability within sites and habitats indicates that localized population characterization should include sampling in both wet and dry seasons. Extreme values at single replicates occasionally influenced overall habitat patterns, while pooling replicates masked variability among sites. A full understanding of population characteristics should include an assessment of variability both at the site and habitat level.

PLoS ONE

Experimental landscape reduction of wild rodents increases movements in the invasive brown treesnake (Boiga irregularis)

Experimental studies evaluating the effects of food availability on the movement of free-ranging animals generally involve food supplementation rather than suppression. Both approaches can yield similar insights, but we were interested in the potential for using food suppression for the management and control of invasive predators, in particular, the brown treesnake ( Boiga irregularis ) on Guam. However, understanding a species’ response to food resources is critical before employing such a strategy. We studied the movements of 24 radio-tagged B. irregularis initially caught within four 4-ha unfenced plots in rodent-abundant (control) and rodent-suppressed (treatment) grassland habitats over a 40-day period. Because monitoring duration differed among snakes, we also analyzed short-term (16-day) activity areas. Over the 16 days, snakes associated with rodent-suppressed plots had 86% larger activity areas (ha), 94% greater dispersal distances (m), and 43% greater movement rates (m/h) than snakes associated with control plots. Boiga irregularis moved extensively, but these movements were not always reflected in the size of the snake’s total activity area. Movement rates did not differ between sexes, but snakes in above-average body condition moved greater distances per hour than those in below-average condition irrespective of treatment. Our study indicates that a relatively small prey suppression effort can cause almost immediate and significant changes in B. irregularis movement. On Guam, prey suppression might enhance control efforts by either increasing trap capture success or discouraging snakes from entering areas of conservation or management concern. However, the outcome of using prey suppression as a control tool in areas threatened with the accidental introduction of the brown treesnake is more difficult to predict and might have negative consequences such as elevated dispersal rates.

Management of Biological Invasions

Ontogenetic and ecological variation in invasion risk of Brown Treesnakes (Boiga irregularis) on Guam

Size structure within populations of invasive species may have consequences for relative risk at all stages of the invasion process, with implications for management interventions such as interdiction, suppression, and eradication. To assess relative distributions of invasive Brown Treesnakes among demographic categories of management interest, we undertook the most comprehensive and controlled sampling in >25 years of research into this ecologically and economically destructive introduced predator. We collected a seasonally-balanced sample of 100 snakes from each of 18 sites, stratified by six habitat types, encompassing the species’ entire extralimital range. Samples indicated significant differences in distributions of female and male snakes among management classes (juvenile, transitional, and mature) by site and habitat. We found substantial heterogeneity in localized population characteristics over relatively small geographic distances, only modest influence of habitat type, higher prevalence of reproductively mature snakes in savanna and urban habitats, and an alarmingly high proportion of snakes that are too small to be effectively targeted by current rodent-baited control tools (mean = 38.2%, range = 19 to 72%). Failure to account for such variability in high risk demographic fractions may hinder successful interventions.

Guam

To cross or not to cross: modeling wildlife road crossings as a binary response variable with contextual predictors

Roads are significant barriers to landscape-scale movements of individuals or populations of many wildlife taxa. The decision by an animal near a road to either cross or not cross may be influenced by characteristics of the road, environmental conditions, traits of the individual animal, and other aspects of the context within which the decision is made. We considered such factors in a mixed-effects logistic regression model describing the nightly road crossing probabilities of invasive nocturnal Brown Treesnakes ( Boiga irregularis ) through short-term radiotracking of 691 snakes within close proximity to 50 road segments across the island of Guam. All measures of road magnitude (traffic volume, gap width, surface type, etc.) were significantly negatively correlated with crossing probabilities. Snake body size was the only intrinsic factor associated with crossing rates, with larger snakes crossing roads more frequently. Humidity was the only environmental variable affecting crossing rate. The distance of the snake from the road at the start of nightly movement trials was the most significant predictor of crossings. The presence of snake traps with live mouse lures during a portion of the trials indicated that localized prey cues reduced the probability of a snake crossing the road away from the traps, suggesting that a snake's decision to cross roads is influenced by local foraging opportunities. Per capita road crossing rates of Brown Treesnakes were very low, and comparisons to historical records suggest that crossing rates have declined in the 60+ yr since introduction to Guam. We report a simplified model that will allow managers to predict road crossing rates based on snake, road, and contextual characteristics. Road crossing simulations based on actual snake size distributions demonstrate that populations with size distributions skewed toward larger snakes will result in a higher number of road crossings. Our method of modeling per capita road crossing probabilities as a binary response variable, influenced by contextual factors, may be useful for describing or predicting road crossings by individuals of other taxa provided that appropriate spatial and temporal resolution can be achieved and that potentially influential covariate data can be obtained.

Ecosphere

Detection rates of geckos in visual surveys: Turning confounding variables into useful knowledge

Transect surveys without some means of estimating detection probabilities generate population size indices prone to bias because survey conditions differ in time and space. Knowing what causes such bias can help guide the collection of relevant survey covariates, correct the survey data, anticipate situations where bias might be unacceptably large, and elucidate the ecology of target species. We used negative binomial regression to evaluate confounding variables for gecko (primarily Hemidactylus frenatus and Lepidodactylus lugubris) counts on 220-m-long transects surveyed at night, primarily for snakes, on 9,475 occasions. Searchers differed in gecko detection rates by up to a factor of six. The worst and best headlamps differed by a factor of at least two. Strong winds had a negative effect potentially as large as those of searchers or headlamps. More geckos were seen during wet weather conditions, but the effect size was small. Compared with a detection nadir during waxing gibbous (nearly full) moons above the horizon, we saw 28% more geckos during waning crescent moons below the horizon. A sine function suggested that we saw 24% more geckos at the end of the wet season than at the end of the dry season. Fluctuations on a longer timescale also were verified. Disturbingly, corrected data exhibited strong short-term fluctuations that covariates apparently failed to capture. Although some biases can be addressed with measured covariates, others will be difficult to eliminate as a significant source of error in longterm monitoring programs.

Andersen Air Force Base

Movements and activity of juvenile Brown Treesnakes ( Boiga irregularis )

Understanding the spatial ecology and foraging strategy of invasive animals is essential for success in control or eradication. We studied movements and activity in juvenile Brown Treesnakes on Guam, as this population segment has proven particularly difficult to control. Distance between daytime refugia (from telemetry of 18 juveniles, 423-800 mm snout-vent length) ranged from 0-118 m ( n = 86), with a grand mean of 43 m. There were tendencies for shorter snake movements on nights directly following a full moon and on dry nights, but variation among snakes was of a larger magnitude and would greatly reduce chances to detect moon or rain effects unless corrected for. Snake activity was estimated from audio recordings of signals from “tipping” radio transmitters, analyzed for pulse period and amplitude. Activity was highest in the hours immediately after sunset, and gradually declined throughout the night before dropping abruptly in conjunction with sunrise. Snake activity was higher on rainy nights, and tended to be highest during waning moons and when the moon was below the horizon. We conclude that small Brown Treesnakes forage actively and appear to move far enough to regularly encounter the traps and bait used on Guam for control purposes, suggesting that alternative explanations are required for their low capture rates with these control tools.

Copeia

Selective predation by feral cats on a native skink on Guam

Two species of skinks (Fig. 1) occur in a 5-ha plot on Guam where we have been conducting intensive research on Brown Treesnake (Boiga irregularis) population biology for nearly a decade (Rodda et al. 2007). The Pacific Blue-tailed Skink (Emoia caeruleocauda [de Vis 1892]) is native to Guam, whereas the Curious Skink (Carlia ailanpalai Zug 2004) is invasive. On the evening of 27 November 2012, PML and MSP discovered a pile of vomited skinks (Fig. 2) inside the plot (UTM 55P: 269125 m E, 1508930 m N).

Reptiles & Amphibians

Canine detection of free-ranging brown treesnakes on Guam

We investigated canine teams (dogs and their handlers) on Guam as a potential tool for finding invasive brown treesnakes (Boiga irregularis) in the wild. Canine teams searched a 40 × 40 m forested area for a snake that had consumed a dead mouse containing a radio-transmitter. To avoid tainting the target or target area with human scent, no snake was handled or closely approached prior to searches. Trials were conducted during the morning when these nocturnal snakes were usually hidden in refugia. A radiotracker knew the snake's location, but dog handlers and search navigators did not. Of 85 trials conducted over four months, the two canine teams had an average success rate of 35% of correctly defining an area ≤ 5 × 5 m that contained the transmittered snake; the team with more experience prior to the trials had a success rate of 44% compared with 26% for the less experienced team. Canine teams also found 11 shed skins from wild snakes. Although dogs alerted outside the vicinity of transmittered snakes, only one wild, non-transmittered snake was found during the trials, possibly reflecting the difficulty humans have in locating non-transmittered brown treesnakes in refugia. We evaluated success at finding snakes as a function of canine team, number of prior trials (i.e. experience gained during the trials), recent canine success at finding a target snake, various environmental conditions, snake perch height, and snake characteristics (snout-vent length and sex). Success rate increased over the course of the trials. Canine team success also increased with increasing average humidity and decreased with increasing average wind speed. Our results suggest dogs could be useful at detecting brown treesnakes in refugia, particularly when compared to daytime visual searches by humans, but techniques are needed to help humans find and extract snakes once a dog has alerted.

New Zealand Journal of Ecology