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Bjorn Lardner

Publications and source records attributed to Bjorn Lardner.

16 recordsLinked to original sources
USGS2025

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↗
USGS2021

Tools for increasing visual encounter probabilities for invasive species removal: A case study of brown treesnakes

Early detection and rapid response (EDRR) are essential to identifying and decisively responding to the introduction or spread of an invasive species, thus avoiding population establishment and improving the probability of achieving eradication. However, detection can be challenging at the onset of a species invasion as low population densities can reduce detection probability and conceal the true extent of the situation until the species is well established. This is doubly challenging if the invading species displays cryptic behavior or is nocturnal, thus further limiting opportunities for its discovery. Survey methods that maximize a searcher’s ability to detect an incipient population are therefore critical for successful EDRR. Brown treesnakes (Boiga irregularis) on Guåhan are a classic cautionary example of the dangers of not detecting an invasion early on, and the risk of their introduction to other islands within the Marianas, Hawai’i and beyond remains. Nocturnal visual surveys are known to detect brown treesnakes of all sizes and are the primary detection tool used by the Brown Treesnake Rapid Response Team, but detection probability remains low in complex forest habitats. As such, we investigated the use of two potential enhancements to nocturnal visual surveys – a live mouse lure and spray scent attractant – that may create hotspots of increased detection probability during surveys. We found that, while brown treesnake detection probabilities were low for all surveys, visual surveys conducted on transects with live mouse lures resulted in detection probabilities that were 1.3 times higher than on transects without live mouse lures. Conversely, the spray scent attractant did not increase the probability of detecting brown treesnakes compared to transects without scent, and in fact had detection probabilities that were 0.66 times lower, though the reasons for this phenomenon are unclear. Unlike scent attractants, live mouse lures likely provide both visual and olfactory cues that attract brown treesnakes to transects and thus provide more opportunities to detect and capture them. These enhancements were trialed on Guåhan, where prey populations are depressed. It remains unclear whether live mouse lures will be as effective for EDRR applications in prey-rich settings.

Guam↗
USGS2021

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↗
USGS2019

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↗
USGS2019

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↗
USGS2019

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↗
USGS2019

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↗
USGS2018

Inferring the absence of an incipient population during a rapid response for an invasive species

Successful eradication of invasives is facilitated by early detection and prompt onset of control. However, realizing or verifying that a colonization has occurred is difficult for cryptic species especially at low population densities. Responding to the capture or unconfirmed sighting of a cryptic invasive species, and the associated effort to determine if it indicates an incipient (small, localized) population or merely a lone colonizer, is costly and cannot continue indefinitely. However, insufficient detection effort risks erroneously concluding the species is not present, allowing the population to increase in size and expand its range. Evidence for an incipient population requires detection of ≥1 individual; its absence, on the other hand, must be inferred probabilistically. We use an actual rapid response incident and species-specific detection estimates tied to a known density to calculate the amount of effort (with non-sequential detections) necessary to assert, with a pre-defined confidence, that invasive brown treesnakes are absent from the search area under a wide range of hypothetical population densities. We illustrate that the amount of effort necessary to declare that a species is absent is substantial and increases with decreased individual detection probability, decreased density, and increased level of desired confidence about its absence. Such survey investment would be justified where the cost savings due to early detection are large. Our Poisson-based model application will allow managers to make informed decisions about how long to continue detection efforts, should no additional detections occur, and suggests that effort to do so is significantly higher than previously thought. While our model application informs how long to search to infer absence of an incipient population of brown treesnakes, the approach is sufficiently general to apply to other invasive species if density-dependent detection estimates are known or reliable surrogate estimates are available.

PLoS ONE↗
USGS2018

Evaluating airsoft electric guns for control of invasive brown treesnakes

Firearms are often used in lethal control of invasive vertebrates, but safety and regulatory aspects limit the circumstances under which they can be used. During August 2016 at the Brown Treesnake Project laboratory on Guam, we evaluated hobby‐grade Airsoft Electric Guns (AEGs)—a lower powered, less‐hazardous, and less‐regulated alternative to firearms—for capture and control of small animals, with specific emphasis on invasive brown treesnakes ( Boiga irregularis ). Tests of AEGs differing in power with ammunition (plastic pellets) masses ranging from 0.20 to 0.39 g, fired at gelatin blocks from distances of 4, 8, and 12 m, showed that heavy ammunition is of overriding importance for maximizing lethality: 0.39‐g pellets penetrated more deeply at 12 m than did 0.20‐g pellets at 4 m. Inspection of tissue damage in brown treesnake carcasses subjected to fire with the 0.39‐g ammunition from the same distances suggested that injuries sustained by a direct hit from 12 m would often be lethal, and snakes would be unlikely to survive multiple hits from automatic fire discharged at approximately 17/s. Limited trials with live snakes helped us to understand behavioral responses in a snake hit by ≥1 pellets, including distance traveled over time. Based on these factors, we assessed the risk that a snake injured by pellet fire might evade subsequent capture by rapid responders in the proximity. We also discuss ethical considerations and regulatory advantages of using AEGs. Published 2018. This article is a U.S. Government work and is in the public domain in the USA.

Wildlife Society Bulletin↗
USGS2016

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↗
USGS2015

Stability of detectability over 17 years at a single site and other lizard detection comparisons from Guam

To obtain quantitative information about population dynamics from counts of animals, the per capita detectabilities of each species must remain constant over the course of monitoring. We characterized lizard detection constancy for four species over 17 yr from a single site in northern Guam, a relatively benign situation because detection was relatively easy and we were able to hold constant the site, habitat type, species, season, and sampling method. We monitored two species of diurnal terrestrial skinks ( Carlia ailanpalai [Curious Skink], Emoia caeruleocauda [Pacific Bluetailed Skink]) using glueboards placed on the ground in the shade for 3 h on rainless mornings, yielding 10,286 skink captures. We additionally monitored two species of nocturnal arboreal geckos ( Hemidactylus frenatus [Common House Gecko]; Lepidodactylus lugubris [Mourning Gecko]) on the basis of 15,212 sightings. We compared these count samples to a series of complete censuses we conducted from four or more total removal plots (everything removed to mineral soil) totaling 400 m 2 (about 1% of study site) in each of the years 1995, 1999, and 2012, providing time-stamped quantification of detectability for each species. Unfortunately, the actual population trajectories taken by the four species were masked by unexplained variation in detectability. This observation of debilitating latent variability in lizard detectability under nearly ideal conditions undercuts our trust in population estimation techniques that fail to quantify venue-specific detectability, rely on pooled detection probability estimates, or assume that modulation in predefined environmental covariates suffices for estimating detectability.

Journal of Herpetology↗
USGS2015

Factors affecting defensive strike behavior in Brown Treesnakes ( Boiga irregularis ) provoked by humans

Striking is a typical antipredator defense exhibited by many species of snakes. While trapping Brown Treesnakes (Boiga irregularis) on Guam, we observed that snakes most frequently struck at an approaching person at a site where snakes had been trapped, marked, and handled in the past. Using a combination of between-sites and within-site comparisons, we assessed if the propensity to strike was correlated with capture histories (both recent and long-term), snake size, body condition (a proxy to nutritional stress), sex, or tail condition (broken or intact), while controlling for confounding variables. We confirmed that propensity to strike was higher at the site where we had been conducting capture-mark-recapture for several years. However, we were unable to demonstrate a correlation between striking tendencies and individual recent or long-term capture histories. The only morphological covariate that had an effect on strike propensity was sex, with females striking more often than males. After removing the site effect from our model, we found that snakes missing parts of their tails were more likely to strike than snakes with intact tails. We have yet to identify the factor(s) that cause the pronounced difference across sites in snake propensity to strike, and data from additional sites might help elucidate any geographical patterns.

Herpetological Conservation and Biology↗
USGS2014

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↗
USGS2009

Prey preferences and prey acceptance in juvenile Brown Treesnakes (Boiga irregularis)

On the Pacific island of Guam, control of the invasive Brown Treesnake (Boiga irregularis) relies largely on methods that use mice as bait. Juvenile B. irregularis feed primarily on lizards and their eggs, but little is known about their prey preference. We conducted an experiment to investigate preferences for, and acceptance of, dead geckos, skinks, and neonatal mice, in juvenile B. irregularis ranging from 290 mm to ca. 700 mm snout-vent length (SVL). Snakes of all sizes showed a preference for geckos over skinks and neonatal mice. Geckos were the first prey chosen in 87% of 224 initial trials (56 snakes subjected to four trials each; 33% would be expected from a random choice). The smallest snakes had the most pronounced preference. Although many of the snakes accepted neonatal mice and/or skinks, some snakes of all sizes were reluctant to feed on anything but geckos, especially when well fed. We also addressed the hypothesis that repeated encounters with a particular prey type increase a snake's preference for that prey. Our study does not support this hypothesis. Our results suggest that control methods relying solely on rodent bait may be inefficient for targeting snakes < 700 mm SVL and that individual heterogeneity in prey preference may cause a significant part of this juvenile cohort to be completely refractory to capture with rodent bait, even if the bait is dead and small enough to be readily swallowed.

Herpetological Conservation and Biology↗