Catriona Harris

Invasive species control: Incorporating demographic data and seed dispersal into a management model for Rhododendron ponticum

Abstract Rhododendron ponticum is a serious invasive alien plant in the British Isles and is of significant conservation and economic concern. Here, we integrate information on both the demographics and spatial dynamics of this species within an individual-based, spatially-explicit model and investigate the effectiveness of different control strategies. Importantly, we simulate seed movement and dispersal using a mechanistic seed dispersal model. We investigate the effectiveness of initiating control at the edge versus the core of the infestation, with and without returning each year to remove seedlings. We compare these results to an age-dependent strategy whereby the oldest plants are removed each year. Age-dependent control, in which the oldest plants were removed first, was the most effective strategy investigated, both in terms of the probability of successful eradication and the number of years taken to control. We demonstrate that this is because the older (and taller) plants towards the core produce more seeds that, on average, travel further. Indeed, our results suggest that the expansion of the invading front is actually driven as much by seeds that disperse long distances from these larger plants as by the seed rain from recently matured plants located much closer to the front. Finally, we investigate the potential use of ‘quarantine lines’ — corridors of unsuitable habitat that are sufficiently wide to contain an infestation, preventing spread to vulnerable areas. This study has provided generic insights into best practice for management based on the current understanding of the biology and ecology of this pernicious, invasive plant.

Severity of Expert-Identified Behavioural Responses of Humpback Whale, Minke Whale, and Northern Bottlenose Whale to Naval Sonar

Controlled exposure experiments using 1 to2 kHz sonar signals were conducted with 11 humpback whales (Megaptera novaeangliae), one minke whale (Balaenoptera acutorostrata), and one northern bottlenose whale (Hyperoodon ampullatus) during three field trials from 2011 to 2013. Ship approaches without sonar transmis-sions, playbacks of killer whale vocalizations, and broadband noise were conducted as controls. Behavioural parameters such as horizontal movement, diving, social interactions, and vocalizations were recorded by animal-attached tags and by visual and acoustic tracking. Based on these data, two expert panels independently scored the severity of behavioural changes that were judged likely to be responses to the experimental stimuli, using a severity scale ranging from no effect (0) to high potential to affect vital rates (9) if exposed repeatedly. After scoring, consensus was reached with a third-party moderator. In humpback whales, killer whale playbacks induced more severe responses than sonar exposure, and both sonar exposures and killer whale playbacks induced more responses and responses of higher severity than the no-sonar ship approaches and broadband noise playbacks. The most common response during sonar exposures in all three species was avoidance of the sound source. The most severe responses to sonar (severity 8) were progressive high-speed avoidance by the minke whale and long-term area avoidance by the bottlenose whale. Other severe responses included prolonged avoidance and cessation of feeding (severity 7). The minke whale and bottlenose whale started avoiding the source at a received sound pressure level (SPL) of 146 and 130 dB re 1 μPa, respectively. Humpback whales generally had less severe responses that were triggered at higher received levels. The probability of severity scores with the potential to affect vital rates increased with increasing sound exposure level (SEL). The single experiments with minke and bottlenose whales suggest they have greater susceptibility to sonar disturbance than humpback whales, but additional studies are needed to confirm this result.

Dose response severity functions for acoustic disturbance in cetaceans using recurrent event survival analysis

Behavioral response studies (BRSs) aim to enhance our understanding of the behavior changes made by animals in response to specific exposure levels of different stimuli, often presented in an increasing dosage. Here, we focus on BRSs that aim to understand behavioral responses of free-ranging whales and dolphins to manmade acoustic signals (although the methods are applicable more generally). One desired outcome of these studies is dose-response functions relevant to different species, signals and contexts. We adapted and applied recurrent event survival analysis (Cox proportional hazard models) to data from the 3S BRS project, where multiple behavioral responses of different severities had been observed per experimental exposure and per individual based upon expert scoring. We included species, signal type, exposure number and behavioral state prior to exposure as potential covariates. The best model included all main effect terms, with the exception of exposure number, as well as two interaction terms. The interactions between signal and behavioral state, and between species and behavioral state highlighted that the sensitivity of animals to different signal types (a 6-7 kHz upsweep sonar signal (MFAS) or a 1-2 kHz upsweep sonar signal (LFAS)) depended on their behavioral state (feeding or non- feeding), and this differed across species. Of the three species included in this analysis (sperm whale (Physeter macrocephalus), killer whale (Orcinus orca) and long-finned pilot whale (Globicephala melas)), killer whales were consistently the most likely to exhibit behavioral responses to naval sonar exposure. We conclude that recurrent event survival analysis provides an effective framework for fitting dose-response severity functions to data from behavioral response studies. It can provide outputs that can help government and industry to evaluate the potential impacts of anthropogenic sound production in the ocean.

The Challenges of Analyzing Behavioral Response Study Data: An Overview of the MOCHA (Multi-study OCean Acoustics Human Effects Analysis) Project.

This paper describes the MOCHA project which aims to develop novel approaches for the analysis of data collected during Behavioral Response Studies (BRSs). BRSs are experiments aimed at directly quantifying the effects of controlled dosages of natural or anthropogenic stimuli (typically sound) on marine mammal behavior. These experiments typically result in low sample size, relative to variability, and so we are looking at a number of studies in combination to maximize the gain from each one. We describe a suite of analytical tools applied to BRS data on beaked whales, including a simulation study aimed at informing future experimental design.

A Simulation-Based Method for Quantifying and Mitigating the Effects of Anthropogenic Sound on Marine Mammals

SAFESIMM (Statistical Algorithms For Estimating the Sonar Influence on Marine Megafauna) is a simulation-based framework for calculating the probable numbers of animals affected by underwater sounds; such as active sonar, which has been implicated in marine mammal fatalities. SAFESIMM is the culmination of a large, multi-year collaborative project between BAE Systems and the University of St Andrews. The resulting software tool uses the latest research on the effects of sound on marine mammals together with data on the distribution, abundance and hearing characteristics of these species. The end result is a simulation-based statistical model that quantifies the probability of physical effects and behavioural responses, along with the expected numbers of occurrences and associated uncertainty in the predictions. The parameters used in each simulation are easily altered, allowing rapid comparison of alternative scenarios. This paper describes: 1. The problem in general terms; 2. The logical high-level solut...

Marine mammals and sonar: Dose‐response studies, the risk‐disturbance hypothesis and the role of exposure context

This manuscript was written following the Behavioral Response Research Evaluation Workshop (BRREW), jointly sponsored by the US Office of Naval Research, US Navy Living Marine Resources, and US National Oceanic and Atmospheric Administration - National Marine Fisheries Service. PLT acknowledges funding from the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland). MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.

A simulation approach to assessing environmental risk of sound exposure to marine mammals

Abstract Intense underwater sounds caused by military sonar, seismic surveys, and pile driving can harm acoustically sensitive marine mammals. Many jurisdictions require such activities to undergo marine mammal impact assessments to guide mitigation. However, the ability to assess impacts in a rigorous, quantitative way is hindered by large knowledge gaps concerning hearing ability, sensitivity, and behavioral responses to noise exposure. We describe a simulation‐based framework, called SAFESIMM (Statistical Algorithms For Estimating the Sonar Influence on Marine Megafauna), that can be used to calculate the numbers of agents (animals) likely to be affected by intense underwater sounds. We illustrate the simulation framework using two species that are likely to be affected by marine renewable energy developments in UK waters: gray seal (Halichoerus grypus) and harbor porpoise (Phocoena phocoena). We investigate three sources of uncertainty: How sound energy is perceived by agents with differing hearing abilities; how agents move in response to noise (i.e., the strength and directionality of their evasive movements); and the way in which these responses may interact with longer term constraints on agent movement. The estimate of received sound exposure level (SEL) is influenced most strongly by the weighting function used to account for the species presumed hearing ability. Strongly directional movement away from the sound source can cause modest reductions (~5 dB) in SEL over the short term (periods of less than 10 days). Beyond 10 days, the way in which agents respond to noise exposure has little or no effect on SEL, unless their movements are constrained by natural boundaries. Most experimental studies of noise impacts have been short‐term. However, data are needed on long‐term effects because uncertainty about predicted SELs accumulates over time. Synthesis and applications. Simulation frameworks offer a powerful way to explore, understand, and estimate effects of cumulative sound exposure on marine mammals and to quantify associated levels of uncertainty. However, they can often require subjective decisions that have important consequences for management recommendations, and the basis for these decisions must be clearly described.

Measuring whale and dolphin call rates as a function of behavioral, social, and environmental context

Cetacean sound-production rates are highly variable and patchy in time, depending upon individual behavior, social context, and environmental context. Better quantification of the drivers of this variability should allow more realistic estimates of expected call rates, improving our ability to convert between call counts and animal density, and also facilitating detection of sound-production changes due to acoustic disturbance. Here, we analyze digital acoustic tag (DTAG) records and visual observations collected during behavioral response studies (BRSs), which aim to assess normal cetacean behavior and measure changes in response to acoustic disturbance; data sources include SOCAL BRS, the 3S project, and Bahamas BRS, with statistical contributions from the MOCHA project (http://www.creem.st-and.ac.uk/mocha/links). We illustrate use of generalized linear models (and their extensions) as a flexible framework for sound-production-rate analysis. In the context of acoustic disturbance, we also detail use of two-dimensional spatially adaptive surfaces to jointly model effects of sound-source proximity and sound intensity. Specifically, we quantify variability in pilot whale group sound production rates in relation to behavior and environment, and individual fin whale call rates in relation to social and environmental context and dive behavior; with and without acoustic disturbance.

Statistical analysis of data from acoustic tags: Methods for combining data streams and modeling animal behavior

Statistical analysis of data from multi-sensor acoustic tags presents several characteristic challenges. Datasets generally include time-series of many measurements on a small number of individuals; different data streams often have distinct temporal resolutions and precisions. The MOCHA project (Multi-study Ocean acoustics Human effects Analysis) is a three-year effort focused on developing innovative statistical methods for such data. Here, we present several approaches for appropriate, effective statistical analysis of such datasets, with an emphasis on quantitative assessment of changes in marine mammal behavior in response to acoustic disturbance. Issues to be addressed will include: combining data streams from multi-sensor tags (and also concurrent visual observation data) for statistical analysis; statistical methods to characterize or summarize normal behavior and detect departures from normal; methods for analysis of call-production-rate data from acoustic tags; and methods for combining analysis...