USGS ScienceSearch

Geology topics

Frank J. Mazzotti

Publications and source records attributed to Frank J. Mazzotti.

At least 37 records · Page 2Linked to original sources

Novel ecological and climatic conditions drive rapid adaptation in invasive Florida Burmese pythons

Invasive species provide powerful in situ experimental systems for studying evolution in response to selective pressures in novel habitats. While research has shown that phenotypic evolution can occur rapidly in nature, few examples exist of genome‐wide adaptation on short ‘ecological’ timescales. Burmese pythons ( Python molurus bivittatus ) have become a successful and impactful invasive species in Florida over the last 30 years despite major freeze events that caused high python mortality. We sampled Florida Burmese pythons before and after a major freeze event in 2010 and found evidence for directional selection in genomic regions enriched for genes associated with thermosensation, behavior, and physiology. Several of these genes are linked to regenerative organ growth, an adaptive response that modulates organ size and function with feeding and fasting in pythons. Independent histological and functional genomic datasets provide additional layers of support for a contemporary shift in invasive Burmese python physiology. In the Florida population, a shift towards maintaining an active digestive system may be driven by the fitness benefits of maintaining higher metabolic rates and body temperature during freeze events. Our results suggest that a synergistic interaction between ecological and climatic selection pressures have driven adaptation in Florida Burmese pythons, demonstrating the often‐overlooked potential of rapid adaptation to influence the success of invasive species.

Florida

Evaluating GPS biologging technology for studying spatial ecology of large constricting snakes

Background : GPS telemetry has revolutionized the study of animal spatial ecology in the last two decades. Until recently, it has mainly been deployed on large mammals and birds, but the technology is rapidly becoming miniaturized, and applications in diverse taxa are becoming possible. Large constricting snakes are top predators in their ecosystems, and accordingly they are often a management priority, whether their populations are threatened or invasive. Fine-scale GPS tracking datasets could greatly improve our ability to understand and manage these snakes, but the ability of this new technology to deliver high-quality data in this system is unproven. In order to evaluate GPS technology in large constrictors, we GPS-tagged 13 Burmese pythons ( Python bivittatu s) in Everglades National Park and deployed an additional 7 GPS tags on stationary platforms to evaluate habitat-driven biases in GPS locations. Both python and test platform GPS tags were programmed to attempt a GPS fix every 90 min. Results : While overall fix rates for the tagged pythons were low (18.1%), we were still able to obtain an average of 14.5 locations/animal/week, a large improvement over once-weekly VHF tracking. We found overall accuracy and precision to be very good (mean accuracy = 7.3 m, mean precision = 12.9 m), but a very few imprecise locations were still recorded (0.2% of locations with precision > 1.0 km). We found that dense vegetation did decrease fix rate, but we concluded that the low observed fix rate was also due to python microhabitat selection underground or underwater. Half of our recovered pythons were either missing their tag or the tag had malfunctioned, resulting in no data being recovered. Conclusions : GPS biologging technology is a promising tool for obtaining frequent, accurate, and precise locations of large constricting snakes. We recommend future studies couple GPS telemetry with frequent VHF locations in order to reduce bias and limit the impact of catastrophic failures on data collection, and we recommend improvements to GPS tag design to lessen the frequency of these failures.

Florida

Everglades Invasive Reptile and Amphibian Monitoring Program

The Everglades Invasive Reptile and Amphibian Monitoring Program (EIRAMP) provides a scientific framework for integrated EDRR, outreach, and monitoring for invasive wildlife in South Florida. EIRAMP also monitors native reptiles, amphibians and mammals to assess impacts of invasive species. South Florida has more nonnative species of reptiles and amphibians than anywhere else in the world. Some of these species become invasive and harm the environment, economy, and/or public health. Many are predators that can severely impact native species and ecosystems. Controlling and monitoring these invaders are essential to the success of multi-billion dollar Everglades restoration efforts. Prevention and eradication are the most cost-effective ways to limit impacts of invasive species (see Figure 1). Once populations are widely established, management becomes expensive, long-term, and often ineffective. Early detection and rapid response (EDRR) offers the best chance to get ahead of the invasion curve and successfully contain or eradicate populations while they are still localized.

Florida

Parasite spillover: Indirect effects of invasive Burmese pythons

Identification of the origin of parasites of nonindigenous species (NIS) can be complex. NIS may introduce parasites from their native range and acquire parasites from within their invaded range. Determination of whether parasites are non-native or native can be complicated when parasite genera occur within both the NIS’ native range and its introduced range. We explored potential for spillover and spillback of lung parasites infecting Burmese pythons ( Python bivittatus ) in their invasive range (Florida). We collected 498 indigenous snakes of 26 species and 805 Burmese pythons during 2004–2016 and examined them for lung parasites. We used morphology to identify three genera of pentastome parasites, Raillietiella , a cosmopolitan form, and Porocephalus and Kiricephalus , both New World forms. We sequenced these parasites at one mitochondrial and one nuclear locus and showed that each genus is represented by a single species, R. orientalis , P. crotali , and K. coarctatus . Pythons are host to R. orientalis and P. crotali , but not K. coarctatus ; native snakes are host to all three species. Sequence data show that pythons introduced R. orientalis to North America, where this parasite now infects native snakes. Additionally, our data suggest that pythons are competent hosts to P. crotali , a widespread parasite native to North and South America that was previously hypothesized to infect only viperid snakes. Our results indicate invasive Burmese pythons have affected parasite-host dynamics of native snakes in ways that are consistent with parasite spillover and demonstrate the potential for indirect effects during invasions. Additionally, we show that pythons have acquired a parasite native to their introduced range, which is the initial condition necessary for parasite spillback.

Florida

Optimal control of an invasive species using a reaction-diffusion model and linear programming

Managing an invasive species is particularly challenging as little is generally known about the species’ biological characteristics in its new habitat. In practice, removal of individuals often starts before the species is studied to provide the information that will later improve control. Therefore, the locations and the amount of control have to be determined in the face of great uncertainty about the species characteristics and with a limited amount of resources. We propose framing spatial control as a linear programming optimization problem. This formulation, paired with a discrete reaction-diffusion model, permits calculation of an optimal control strategy that minimizes the remaining number of invaders for a fixed cost or that minimizes the control cost for containment or protecting specific areas from invasion. We propose computing the optimal strategy for a range of possible model parameters, representing current uncertainty on the possible invasion scenarios. Then, a best strategy can be identified depending on the risk attitude of the decision-maker. We use this framework to study the spatial control of the Argentine black and white tegus ( Salvator merianae ) in South Florida. There is uncertainty about tegu demography and we considered several combinations of model parameters, exhibiting various dynamics of invasion. For a fixed one-year budget, we show that the risk-averse strategy, which optimizes the worst-case scenario of tegus’ dynamics, and the risk-neutral strategy, which optimizes the expected scenario, both concentrated control close to the point of introduction. A risk-seeking strategy, which optimizes the best-case scenario, focuses more on models where eradication of the species in a cell is possible and consists of spreading control as much as possible. For the establishment of a containment area, assuming an exponential growth we show that with current control methods it might not be possible to implement such a strategy for some of the models that we considered. Including different possible models allows an examination of how the strategy is expected to perform in different scenarios. Then, a strategy that accounts for the risk attitude of the decision-maker can be designed.

Florida

Life histories and conservation of long-lived reptiles, an illustration with the American crocodile (Crocodylus acutus)

Successful species conservation is dependent on adequate estimates of population dynamics, but age-specific demographics are generally lacking for many long-lived iteroparous species such as large reptiles. Accurate demographic information allows estimation of population growth rate, as well as projection of future population sizes and quantitative analyses of fitness trade-offs involved in the evolution of life-history strategies. Here, a long-term capture–recapture study was conducted from 1978 to 2014 on the American crocodile ( Crocodylus acutus ) in southern Florida. Over the study period, 7,427 hatchlings were marked and 380 individuals were recaptured for as many as 25 years. We estimated survival to be strongly age dependent with hatchlings having the lowest survival rates (16%) but increasing to nearly 90% at adulthood based on mark–recapture models. More than 5% of the female population were predicted to be reproductive by age 8 years; the age-specific proportion of reproductive females steadily increased until age 18 when more than 95% of females were predicted to be reproductive. Population growth rate, estimated from a Leslie–Lefkovitch stage-class model, showed a positive annual growth rate of 4% over the study period. Using a prospective sensitivity analysis, we revealed that the adult stage, as expected, was the most critical stage for population growth rate; however, the survival of younger crocodiles before they became reproductive also had a surprisingly high elasticity. We found that variation in age-specific fecundity has very limited impact on population growth rate in American crocodiles. We used a comparative approach to show that the original life-history strategy of American crocodiles is actually shared by other large, long-lived reptiles: while adult survival rates always have a large impact on population growth, this decreases with declining increasing growth rates, in favour of a higher elasticity of the juvenile stage. Crocodiles, as a long-lived and highly fecund species, deviate from the usual association of life histories of “slow” species. Current management practices are focused on nests and hatchling survival; however, protection efforts that extend to juvenile crocodiles would be most effective for conservation of the species, especially in an ever-developing landscape.

Journal of Animal Ecology

Conservation genetics of American crocodile, Crocodylus acutus , populations in Pacific Costa Rica

Maintaining genetic diversity is crucial for the survival and management of threatened and endangered species. In this study, we analyzed genetic diversity and population genetic structure at neutral loci in American crocodiles, Crocodylus acutus, from several areas (Parque Nacional Marino Las Baulas, Parque Nacional Santa Rosa, Parque Nacional Palo Verde, Rio Tarcoles, and Osa Conservation Area) in Pacific Costa Rica. We genotyped 184 individuals at nine microsatellite loci to describe the genetic diversity and conservation genetics between and among populations. No population was at Hardy-Weinberg Equilibrium (HWE) over all loci tested and a small to moderate amount of inbreeding was present. Populations along the Pacific coast had an average heterozygosity of 0.572 across all loci. All populations were significantly differentiated from each other with both FST and RST measures of population differentiation with a greater degree of molecular variance (81%) found within populations. Our results suggest C. acutus populations in Pacific Costa Rica were not panmictic with moderate levels of genetic diversity. An effective management plan that maintains the connectivity between clusters is critical to the success of C. acutus in Pacific Costa Rica.

Nature Conservation

Expert elicitation, uncertainty, and the value of information in controlling invasive species

We illustrate the utility of expert elicitation, explicit recognition of uncertainty, and the value of information for directing management and research efforts for invasive species, using tegu lizards (Salvator merianae) in southern Florida as a case study. We posited a post-birth pulse, matrix model in which four age classes of tegus are recognized: hatchlings, 1 year-old, 2 year-olds, and 3 + year-olds. This matrix model was parameterized using a 3-point process to elicit estimates of tegu demographic rates in southern Florida from 10 herpetology experts. We fit statistical distributions for each parameter and for each expert, then drew and pooled a large number of replicate samples from these to form a distribution for each demographic parameter. Using these distributions, as well as the observed correlations among elicited values, we generated a large sample of matrix population models to infer how the tegu population would respond to control efforts. We used the concepts of Pareto efficiency and stochastic dominance to conclude that targeting older age classes at relatively high rates appears to have the best chance of minimizing tegu abundance and control costs. We conclude that expert opinion combined with an explicit consideration of uncertainty can be valuable in conducting an initial assessment of what control strategy, effort, and monetary resources are needed to reduce and eventually eliminate the invader. Scientists, in turn, can use the value of information to focus research in a way that not only increases the efficacy of control, but minimizes costs as well.

Ecological Economics

Comparison of climate envelope models developed using expert-selected variables versus statistical selection

Climate envelope models are widely used to describe potential future distribution of species under different climate change scenarios. It is broadly recognized that there are both strengths and limitations to using climate envelope models and that outcomes are sensitive to initial assumptions, inputs, and modeling methods Selection of predictor variables, a central step in modeling, is one of the areas where different techniques can yield varying results. Selection of climate variables to use as predictors is often done using statistical approaches that develop correlations between occurrences and climate data. These approaches have received criticism in that they rely on the statistical properties of the data rather than directly incorporating biological information about species responses to temperature and precipitation. We evaluated and compared models and prediction maps for 15 threatened or endangered species in Florida based on two variable selection techniques: expert opinion and a statistical method. We compared model performance between these two approaches for contemporary predictions, and the spatial correlation, spatial overlap and area predicted for contemporary and future climate predictions. In general, experts identified more variables as being important than the statistical method and there was low overlap in the variable sets (<40%) between the two methods Despite these differences in variable sets (expert versus statistical), models had high performance metrics (>0.9 for area under the curve (AUC) and >0.7 for true skill statistic (TSS). Spatial overlap, which compares the spatial configuration between maps constructed using the different variable selection techniques, was only moderate overall (about 60%), with a great deal of variability across species. Difference in spatial overlap was even greater under future climate projections, indicating additional divergence of model outputs from different variable selection techniques. Our work is in agreement with other studies which have found that for broad-scale species distribution modeling, using statistical methods of variable selection is a useful first step, especially when there is a need to model a large number of species or expert knowledge of the species is limited. Expert input can then be used to refine models that seem unrealistic or for species that experts believe are particularly sensitive to change. It also emphasizes the importance of using multiple models to reduce uncertainty and improve map outputs for conservation planning. Where outputs overlap or show the same direction of change there is greater certainty in the predictions. Areas of disagreement can be used for learning by asking why the models do not agree, and may highlight areas where additional on-the-ground data collection could improve the models.

Ecological Modelling

The gold tegu, Tupinambis teguixin (Linnaeus, 1758) sensu lato (Squamata: Teiidae): evidence for an established population in Florida

Gold tegus, Tupinambis teguixin (Linnaeus, 1758), are generalist predators from South America and are ecologically similar to Argentine black and white tegus ( Salvator merianae ), a successful invader in Florida. We trapped gold tegus in Broward and Miami-Dade counties, Florida, USA. In Miami-Dade County, collection occurred from 2008 through 2016. We combined new trapping records with previous literature records. Ten gold tegus of both sexes and multiple size classes over a nine year period indicate a reproducing population in Miami-Dade County. Tupinambis teguixin is the sixth established non-native teiid species in Florida. Additionally, we report Tupinambis teguixin from Sarasota and Lee counties, Florida, USA. Determining population status in those counties requires further study. It is critical to differentiate this species from Salvator merianae during removal efforts. The possibility of eradicating Tupinambis teguixin exists if proper identification enables reliable monitoring of the populations.

Florida

Large reptiles and cold temperatures: Do extreme cold spells set distributional limits for tropical reptiles in Florida?

Distributional limits of many tropical species in Florida are ultimately determined by tolerance to low temperature. An unprecedented cold spell during 2–11 January 2010, in South Florida provided an opportunity to compare the responses of tropical American crocodiles with warm-temperate American alligators and to compare the responses of nonnative Burmese pythons with native warm-temperate snakes exposed to prolonged cold temperatures. After the January 2010 cold spell, a record number of American crocodiles ( n = 151) and Burmese pythons ( n = 36) were found dead. In contrast, no American alligators and no native snakes were found dead. American alligators and American crocodiles behaved differently during the cold spell. American alligators stopped basking and retreated to warmer water. American crocodiles apparently continued to bask during extreme cold temperatures resulting in lethal body temperatures. The mortality of Burmese pythons compared to the absence of mortality for native snakes suggests that the current population of Burmese pythons in the Everglades is less tolerant of cold temperatures than native snakes. Burmese pythons introduced from other parts of their native range may be more tolerant of cold temperatures. We documented the direct effects of cold temperatures on crocodiles and pythons; however, evidence of long-term effects of cold temperature on their populations within their established ranges remains lacking. Mortality of crocodiles and pythons outside of their current established range may be more important in setting distributional limits.

Ecosphere

Betrayal: radio-tagged Burmese pythons reveal locations of conspecifics in Everglades National Park

The &ldquo;Judas&rdquo; technique is based on the idea that a radio-tagged individual can be used to &ldquo;betray&rdquo; conspecifics during the course of its routine social behavior. The Burmese python ( Python bivittatus ) is an invasive constrictor in southern Florida, and few methods are available for its control. Pythons are normally solitary, but from December&ndash;April in southern Florida, they form breeding aggregations containing up to 8 individuals, providing an opportunity to apply the technique. We radio-tracked 25 individual adult pythons of both sexes during the breeding season from 2007&ndash;2012. Our goals were to (1) characterize python movements and determine habitat selection for betrayal events, (2) quantify betrayal rates of Judas pythons, and (3) compare the efficacy of this tool with current tools for capturing pythons, both in terms of cost per python removed (CPP) and catch per unit effort (CPUE). In a total of 33 python-seasons, we had 8 betrayal events (24 %) in which a Judas python led us to new pythons. Betrayal events occurred more frequently in lowland forest (including tree islands) than would be expected by chance alone. These 8 events resulted in the capture of 14 new individuals (1&ndash;4 new pythons per event). Our effort comparison shows that while the Judas technique is more costly than road cruising surveys per python removed, the Judas technique yields more large, reproductive females and is effective at a time of year that road cruising is not, making it a potential complement to the status quo removal effort.

Florida

Spatial and temporal variability in estuary habitat use by American alligators

Estuarine habitat occupied by Alligator mississippiensis , a primarily freshwater species, is spatially and temporally heterogeneous largely due to a salinity gradient that fluctuates. Using long-term night light survey data, we examined seasonal patterns in alligators&rsquo; habitat use by size classes in midstream and downstream estuary zones of Shark River, Everglades National Park, in southern Florida. We observed predominantly large-sized alligators (total length&thinsp;&ge;&thinsp;1.75 m); observations of alligators in the small size classes (0.5 m&thinsp;&le;&thinsp;total length&thinsp;<&thinsp;1.25 m) were rare especially in the higher-salinity downstream zone. The density of alligators in the downstream zone was lower than that of the midstream zone during the dry season when salinity increases due to reduced precipitation. Conversely, the density of the large size alligators was higher in the downstream zone than in the midstream zone during the wet season, likely because of reduced salinity. We also found a significant declining trend over time in the number of alligators in the dry season, which coincides with the reported decline in alligator relative density in southern Florida freshwater wetlands. Our results indicated high adaptability of alligators to the fluctuating habitat conditions. Use of estuaries by alligators is likely driven in part by physiology and possibly by reproductive cycle, and our results supported their opportunistic use of estuary habitat and ontogenetic niche shifts.

Florida

Fluctuating water depths affect American alligator ( Alligator mississippiensis ) body condition in the Everglades, Florida, USA

Successful restoration of wetland ecosystems requires knowledge of wetland hydrologic patterns and an understanding of how those patterns affect wetland plant and animal populations.Within the Everglades, Florida, USA restoration, an applied science strategy including conceptual ecological models linking drivers to indicators is being used to organize current scientific understanding to support restoration efforts. A key driver of the ecosystem affecting the distribution and abundance of organisms is the timing, distribution, and volume of water flows that result in water depth patterns across the landscape. American alligators (Alligator mississippiensis) are one of the ecological indicators being used to assess Everglades restoration because they are a keystone species and integrate biological impacts of hydrological operations through all life stages. Alligator body condition (the relative fatness of an animal) is one of the metrics being used and targets have been set to allow us to track progress. We examined trends in alligator body condition using Fulton&rsquo;s K over a 15 year period (2000&ndash;2014) at seven different wetland areas within the Everglades ecosystem, assessed patterns and trends relative to restoration targets, and related those trends to hydrologic variables. We developed a series of 17 a priori hypotheses that we tested with an information theoretic approach to identify which hydrologic factors affect alligator body condition. Alligator body condition was highest throughout the Everglades during the early 2000s and is approximately 5&ndash;10% lower now (2014). Values have varied by year, area, and hydrology. Body condition was positively correlated with range in water depth and fall water depth. Our top model was the &ldquo;Current&rdquo; model and included variables that describe current year hydrology (spring depth, fall depth, hydroperiod, range, interaction of range and fall depth, interaction of range and hydroperiod). Across all models, interaction between range and fall water depth was the most important variable (relative weight of 1.0) followed by spring and fall water depths (0.99), range (0.96), hydroperiod (0.95) and interaction between range and hydroperiod (0.95). Our work provides additional evidence that restoring a greater range in annual water depths is important for improvement of alligator body condition and ecosystem function. This information can be incorporated into both planning and operations to assist in reaching Everglades restoration goals.

Florida

Considerations for building climate-based species distribution models

Climate plays an important role in the distribution of species. A given species may adjust to new conditions in-place, move to new areas with suitable climates, or go extinct. Scientists and conservation practitioners use mathematical models to predict the effects of future climate change on wildlife and plan for a biodiverse future. This 8-page fact sheet written by David N. Bucklin, Mathieu Basille, Stephanie S. Romañach, Laura A. Brandt, Frank J. Mazzotti, and James I. Watling and published by the Department of Wildlife Ecology and Conservation explains how, with a better understanding of species distribution models, we can predict how species may respond to climate change. The models alone cannot tell us how a certain species will actually respond to changes in climate, but they can inform conservation planning that aims to allow species to both adapt in place and (for those that are able to) move to newly suitable areas. Such planning will likely minimize loss of biodiversity due to climate change.

Report

Dry years decrease abundance of American alligators in the Florida Everglades

The Everglades has been greatly reduced and is threatened by land use change and altered hydrology. The Comprehensive Everglades Restoration Plan calls for monitoring and assessment of key ecosystem attributes, one of which is abundance of American alligators. We examined 10 years of alligator night spotlight counts from Arthur R. Marshall Loxahatchee National Wildlife Refuge along two canals and in the interior marsh to determine trends and how dry years affect alligator abundance. Alligators showed population response to hydrologic conditions. In particular, there were declines in abundance after dry years followed by an apparent recovery in abundance in subsequent years. Increases in abundance were lower in the marsh than L-40 Canal. In addition, there was evidence that intensity of dry events affected population dynamics with greater declines observed in years with drier conditions. Results revealed that overall population of alligators increased from 2004 to 2013, but that increases varied by survey route. These results demonstrate that dry years cause a decline in alligator abundance proportional to the intensity of the dry event, and that it is important to make a distinction between canals and marsh when measuring alligator response to hydrology.

Florida

Home range, habitat use, and movement patterns of non-native Burmese pythons in Everglades National Park, Florida, USA

Background Studies on the spatial ecology of invasive species provide critical information for conservation managers such as habitat preferences and identification of native species at risk of predation. To understand the spatial ecology of non-native Burmese pythons ( Python molurus bivittatus ), now well-established in Everglades National Park and much of South Florida USA, we radio-tracked 19 wild-caught adult pythons, 16 with VHF tags during 2006 through 2009 and 3 by GPS tags between 2010 and 2011. Our goal was to identify individual core-use areas and quantify home ranges, as well as to explore correlations of python movements with environmental parameters such as the presence of surface water. Results Radio-tracking periods ranged from 87 to 697 days for snakes with VHF tags, with a total of 5,119 tracking days (mean&thinsp;&plusmn;&thinsp;1 SD&thinsp;=&thinsp;319.9&thinsp;&plusmn;&thinsp;184.3 days); GPS tracking periods ranged from 12 to 93 days, with a total of 146 tracking days (mean&thinsp;&plusmn;&thinsp;1 SD&thinsp;=&thinsp;48.7&thinsp;&plusmn;&thinsp;40.7 days). We observed mean individual radio-tracked python home ranges of 22.5 km 2 (2250 ha) with overall low site fidelity; all home ranges were within the park boundary. Python core-use areas included slough and coastal habitat types, and we delineated 18 common-use areas (that is, areas where individual core-use areas spatially overlapped). Tree islands were a principal feature of common-use areas, even if they were not the predominant habitat type. Multiple common-use areas were in proximity to roads. The longest movements of individual pythons correlated well with presence of surface water, and occurred during both wet and dry seasons. Conclusions High-use areas determined from python habitat-use and movement data may be optimal locations for targeted control efforts and further studies on impacts to native fauna.

Florida

Factors affecting individual foraging specialization and temporal diet stability across the range of a large “generalist” apex predator

Individual niche specialization (INS) is increasingly recognized as an important component of ecological and evolutionary dynamics. However, most studies that have investigated INS have focused on the effects of niche width and inter- and intraspecific competition on INS in small-bodied species for short time periods, with less attention paid to INS in large-bodied reptilian predators and the effects of available prey types on INS. We investigated the prevalence, causes, and consequences of INS in foraging behaviors across different populations of American alligators ( Alligator mississippiensis ), the dominant aquatic apex predator across the southeast US, using stomach contents and stable isotopes. Gut contents revealed that, over the short term, although alligator populations occupied wide ranges of the INS spectrum, general patterns were apparent. Alligator populations inhabiting lakes exhibited lower INS than coastal populations, likely driven by variation in habitat type and available prey types. Stable isotopes revealed that over longer time spans alligators exhibited remarkably consistent use of variable mixtures of carbon pools (e.g., marine and freshwater food webs). We conclude that INS in large-bodied reptilian predator populations is likely affected by variation in available prey types and habitat heterogeneity, and that INS should be incorporated into management strategies to efficiently meet intended goals. Also, ecological models, which typically do not consider behavioral variability, should include INS to increase model realism and applicability.

Oecologia