Application of Spatial and Closed Capture-Recapture Models on Known Population of the Western Derby Eland (Taurotragus derbianus derbianus) in Senegal

Camera trapping with capture-recapture analyses has provided estimates of the abundances of elusive species over the last two decades. Closed capture-recapture models (CR) based on the recognition of individuals and incorporating natural heterogeneity in capture probabilities are considered robust tools; however, closure assumption is often questionable and the use of an M_h jackknife estimator may fail in estimations of real abundance when the heterogeneity is high and data is sparse. A novel, spatially explicit capture-recapture (SECR) approach based on the location-specific capture histories of individuals overcomes the limitations of closed models. We applied both methods on a closed population of 16 critically endangered Western Derby elands in the fenced 1,060-ha Fathala reserve, Senegal. We analyzed the data from 30 cameras operating during a 66-day sampling period deployed in two densities in grid and line arrays. We captured and identified all 16 individuals in 962 trap-days. Abundances were estimated in the programs CAPTURE (models M₀, M_h and M_h Chao) and R, package secr (basic Null and Finite mixture models), and compared with the true population size. We specified 66 days as a threshold in which SECR provides an accurate estimate in all trapping designs within the 7-times divergent density from 0.004 to 0.028 camera trap/ha. Both SECR models showed uniform tendency to overestimate abundance when sampling lasted shorter with no major differences between their outputs. Unlike the closed models, SECR performed well in the line patterns, which indicates promising potential for linear sampling of properly defined habitats of non-territorial and identifiable herbivores in dense wooded savanna conditions. The CR models provided reliable estimates in the grid and we confirmed the advantage of M_h Chao estimator over M_h jackknife when data appeared sparse. We also demonstrated the pooling of trapping occasions with an increase in the capture probabilities, avoiding violation of results.     

Edge Effect on Density Estimates of a Radiotracked Population of Yellow-Necked Mice

ABSTRACT Density estimates for small-mammal populations from capture-mark-recapture (CMR) data have played an important role in many studies of theoretical and applied ecology. Defining effective trapping area (ETA) is one of the main issues affecting accuracy of density estimates. Our objective was to assess sensitivity of CMR density estimates to correctors based on movement parameters calculated from trapping and radiotelemetry data. From May to November 2005, we conducted monthly CMR trapping in a beech (Fagus sylvaticus) forest of the province of Trento, northern Italy. In conjunction with CMR, we radio-marked 32 yellow-necked mice (Apodemus flavicollis) captured from July to October and located them daily using radiotelemetry. We estimated population size (N) by model averaging with Program MARK. We calculated ETA using several definitions of the boundary strip, including full and half mean maximum distance moved (MMDM) from capture-recapture and telemetry data and mean radius of mean monthly home ranges. The boundary strip (W) increased with the amount of behavioral information embodied in the estimates. The largest W and lowest density values were based on radius of mean home ranges followed by MMDM calculated from telemetry data. The ETA based on movement distances increased more than proportionally when N decreased, suggesting that low population density combined with scarce resources results in rodents moving more in search of food, thus leading to overestimated ETA and underestimated density values. Although robust behavioral information would certainly improve density estimates, we suggest caution in relating ranging movements to capture probability and hence in using correctors based on movement distances to infer density values.     

How Does Spatial Study Design Influence Density Estimates from Spatial Capture-Recapture Models?

When estimating population density from data collected on non-invasive detector arrays, recently developed spatial capture-recapture (SCR) models present an advance over non-spatial models by accounting for individual movement. While these models should be more robust to changes in trapping designs, they have not been well tested. Here we investigate how the spatial arrangement and size of the trapping array influence parameter estimates for SCR models. We analysed black bear data collected with 123 hair snares with an SCR model accounting for differences in detection and movement between sexes and across the trapping occasions. To see how the size of the trap array and trap dispersion influence parameter estimates, we repeated analysis for data from subsets of traps: 50% chosen at random, 50% in the centre of the array and 20% in the South of the array. Additionally, we simulated and analysed data under a suite of trap designs and home range sizes. In the black bear study, we found that results were similar across trap arrays, except when only 20% of the array was used. Black bear density was approximately 10 individuals per 100 km(2). Our simulation study showed that SCR models performed well as long as the extent of the trap array was similar to or larger than the extent of individual movement during the study period, and movement was at least half the distance between traps. SCR models performed well across a range of spatial trap setups and animal movements. Contrary to non-spatial capture-recapture models, they do not require the trapping grid to cover an area several times the average home range of the studied species. This renders SCR models more appropriate for the study of wide-ranging mammals and more flexible to design studies targeting multiple species.     

Nests and nesting sites of Brazilian forest rodents (Nectomys squamipes andOryzomys intermedius) as revealed by a spool-and-line device

Patterns of nest construction and nest site selection of two murid rodents of the Brazilian Atlantic rainforest were described. The animals were captured in an area of about 1.5 ha of primary forest, marked, equipped with a spool-and-line device and released at the capture point. The next day we followed the thread to find their nests. In total 11 nests ofNectomys squamipes (Brants, 1827) and 13 ofOryzomys intermedius (Leche, 1886) were found. They were all egg-shaped, about 15 cm long and 10 cm wide, without any apparent entrance and consisted of dry leaves and grass. The two species differed in the selection of nest sites. The nests ofN. squamipes were close to streams and built inside decomposed fallen logs or using natural entrances between the soil and tangled roots or rocky crevices. Nests ofO. intermedius were built under or inside fallen logs and inside aerial root systems of palms. The spool-and-line technique used here is useful to provide detailed information on several aspects of the ecology and natural history of small mammals.     

Agouti reintroduction recovers seed dispersal of a large-seeded tropical tree

The aim of animal reintroductions has mainly been species recovery; only few reintroduction initiatives focus on ecosystem restoration. Therefore, reintroduction consequences on ecological interactions are seldom assessed. We used the interaction between a reintroduced population of agoutis (Dasyprocta leporina) and a vulnerable tropical endemic tree (Joannesia princeps) to examine reintroduction effects on seed dispersal and seedling establishment. To test the outcomes of this interaction, we tracked seeds of J. princeps in two adjacent forest areas with and without reintroduced agoutis. We also assessed if dispersal distances affected seedling survival. To determine seed fate and dispersal distance, we used spool-and-line tracking, together with camera traps to identify dispersers. Agoutis were the only species removing J. princeps seeds, thus dispersal only occurred where agoutis had been reintroduced; in the area without agoutis, all seeds remained intact on the soil, even one year after the experiment's beginning. At the reintroduction area, most seeds were preyed upon by agoutis but 7% remained dispersed and 2% germinated after ten months. Only seeds buried by agoutis were able to germinate. Most dispersed seeds were dispersed 15 m or farther and longer dispersal distances benefited J. princeps, since seedlings farther from a conspecific adult tree had greater survival probability. Agoutis were also seen burying seeds of two other plant species; these mammals have the potential to benefit dozens of large-seeded species in our study system. Agouti reintroduction thus exemplifies the value of trophic rewilding programs to re-establish ecological interactions and restore ecosystem functioning.     

Estimating Black Bear Density Using DNA Data From Hair Snares

ABSTRACT DNA-based mark-recapture has become a methodological cornerstone of research focused on bear species. The objective of such studies is often to estimate population size; however, doing so is frequently complicated by movement of individual bears. Movement affects the probability of detection and the assumption of closure of the population required in most models. To mitigate the bias caused by movement of individuals, population size and density estimates are often adjusted using ad hoc methods, including buffering the minimum polygon of the trapping array. We used a hierarchical, spatial capture-recapture model that contains explicit components for the spatial-point process that governs the distribution of individuals and their exposure to (via movement), and detection by, traps. We modeled detection probability as a function of each individual's distance to the trap and an indicator variable for previous capture to account for possible behavioral responses. We applied our model to a 2006 hair-snare study of a black bear (Ursus americanus) population in northern New York, USA. Based on the microsatellite marker analysis of collected hair samples, 47 individuals were identified. We estimated mean density at 0.20 bears/km². A positive estimate of the indicator variable suggests that bears are attracted to baited sites; therefore, including a trap-dependence covariate is important when using bait to attract individuals. Bayesian analysis of the model was implemented in WinBUGS, and we provide the model specification. The model can be applied to any spatially organized trapping array (hair snares, camera traps, mist nests, etc.) to estimate density and can also account for heterogeneity and covariate information at the trap or individual level.     

Dispersal and population structure of the rufous bettong,Aepyprymnus rufescens (Marsupialia: Potoroidae)

Abstract Many species of herbivorous mammals declined to extinction following European settlement of inland Australia. The rufous bettong, Aepyprymnus rufescens (a macropodoid marsupial), is ecologically similar to many of these species. We used analysis of microsatellite markers to determine dispersal patterns and mating system characteristics in a cluster of local populations of A. rufescens, with the aim of gaining a better understanding of regional population dynamics in such species. Particularly, we asked whether the rufous bettong showed source‐sink dynamics, as Morton (1990) hypothesized that many mammals may have been made vulnerable to extinction through such processes. We compared populations separated by distances of up to 12 km, and detected significant genetic differentiation among local populations (F_ST = 0.016). Females displayed greater genetic structuring than males, suggesting that females dispersed over shorter distances or less frequently than males. Geographic distance was weakly related to genetic distance between populations suggesting some gene flow at this scale, and paternity assignment indicated that dispersal can occur over distances of up to 6.5 km. Our study populations varied widely in density, but density did not explain the pattern of genetic differentiation observed. These findings of significant structure among populations, some influence of distance on genetic divergence and that density explains little of the divergence among populations, suggested that source‐sink dynamics did not play a large role among these populations. Variance in male mating success was low (maximum assigned paternity for an individual male was 14% of offspring). While data on multiple maternity were limited, roughly half of repeat maternity was assigned to the same male, suggesting that the mating system of the rufous bettong is not purely promiscuous.     

How Small is too Small? Camera Trap Survey Areas and Density Estimates for Ocelots in the Bolivian Chaco

Studies on carnivores, which are generally difficult to observe directly because they are elusive and nocturnal, are carried out through indirect methods, e.g. , camera trapping and radiotracking. The first method has been used to estimate population densities of species that can be differentiated as individuals using unique pelage patterns. However, the use of capture–recapture methodology has raised doubts regarding the estimation of the sampling area around the camera traps, which is obtained using maximum distances traveled by individuals photographed at two or more different locations. In this paper, the results from camera traps are compared with a radiotracking study carried out simultaneously with ocelots ( Leopardus pardalis ) to confirm whether maximum distances observed in camera traps coincide with ranging patterns determined from radio telemetry, and in turn whether the sampling areas estimated from camera traps are appropriate for estimating density. Mean maximum distance moved was 2880 m according to camera trap records during a 60-d survey period while, with radiotracking, the maximum distance moved was 3176 m during the same period. The difference is not significant, and the sampling areas estimated with camera traps to assess ocelot density are reliable. However, if the area covered by cameras is reduced to less than three to four times average home range for the target species, then density estimates from camera trapping are exaggerated because of the reduced observed distances and the fact that multiple individuals can overlap in relatively small areas.     

Bayes and Empirical Bayes Estimators of Abundance and Density from Spatial Capture-Recapture Data

In capture-recapture and mark-resight surveys, movements of individuals both within and between sampling periods can alter the susceptibility of individuals to detection over the region of sampling. In these circumstances spatially explicit capture-recapture (SECR) models, which incorporate the observed locations of individuals, allow population density and abundance to be estimated while accounting for differences in detectability of individuals. In this paper I propose two Bayesian SECR models, one for the analysis of recaptures observed in trapping arrays and another for the analysis of recaptures observed in area searches. In formulating these models I used distinct submodels to specify the distribution of individual home-range centers and the observable recaptures associated with these individuals. This separation of ecological and observational processes allowed me to derive a formal connection between Bayes and empirical Bayes estimators of population abundance that has not been established previously. I showed that this connection applies to every Poisson point-process model of SECR data and provides theoretical support for a previously proposed estimator of abundance based on recaptures in trapping arrays. To illustrate results of both classical and Bayesian methods of analysis, I compared Bayes and empirical Bayes esimates of abundance and density using recaptures from simulated and real populations of animals. Real populations included two iconic datasets: recaptures of tigers detected in camera-trap surveys and recaptures of lizards detected in area-search surveys. In the datasets I analyzed, classical and Bayesian methods provided similar – and often identical – inferences, which is not surprising given the sample sizes and the noninformative priors used in the analyses.     

MICROGEOGRAPHIC VARIATION IN MITOCHONDRIAL DNA OF MEADOW VOLES (MICROTUS PENNSYLVANICUS) IN RELATION TO POPULATION DENSITY

We examined mitochondrial-DNA (mtDNA) sequence heterogeneity on four adjacent trapping grids in an island population of meadow voles (Microtus pennsylvanicus) at two different population densities. Four restriction endonucleases revealed 20 different mtDNA composite phenotypes in samples totaling 198 meadow voles. There were significant heterogeneities in the distribution of four common mtDNA composite phenotypes among the four trapping grids, suggesting that there is population subdivision on a fine scale. Genetic distances between grids, mtDNA diversity within grids, and G_ST also varied during the study period. We found a decrease in genetic distance and an increase in diversity when the population density was high and vice versa when the population density was low. When population density was high, the coefficient of gene differentiation was smaller than the same coefficient observed when the population density was low. These changes in population subdivision and diversity are consistent with theoretical expectations of population structure in which effective female population size and dispersal are the critical variables. The data also support the hypothesis of maintenance of mtDNA diversity by population subdivision, rapid population growth rate, and dispersal.     

cameratrapR: An R package for estimating animal density using camera trapping data

• 1.Camera trapping plays an important role in wildlife surveys, and provides valuable information for estimation of population density. While mark-recapture techniques can estimate population density for species that can be individually recognized or marked, there are no robust methods to estimate density of species that cannot be individually identified.
• 2.We developed a new approach to estimate population density based on the simulation of individual movement within the camera grid. Simulated animals followed a correlated random walk with the movement parameters of segment length, angular deflection, movement distance and home-range size derived from empirical movement paths. Movement was simulated under a series of population densities. We used the Random Forest algorithm to determine the population density with the highest likelihood of matching the camera trap data. We developed an R package, cameratrapR, to conduct simulations and estimate population density.
• 3.Compared with line transect surveys and the random encounter model, cameratrapR provides more reliable estimates of wildlife density with narrower confidence intervals. Functions are provided to visualize movement paths, derive movement parameters, and plot camera trapping results.
• 4.The package allows researchers to estimate population sizes/densities of animals that cannot be individually identified and cameras are deployed in a grid pattern.

     

Microhabitat selection and daily movements of two rodents (Necromys lasiurus and Oryzomys scotti) in Brazilian Cerrado, as revealed by a spool-and-line device

Studies on patterns of habitat use by mammals are necessary for understanding the mechanisms involved in their distribution and abundance. In this study, we used the spool-and-line method to investigate habitat utilization by two sigmodontine rodents from Brazilian Cerrado, Necromys lasiurus and Oryzomys scotti. We conducted the study in a Cerrado area in central Brazil (15°56′S and e 47°56′W) where the animals were caught in an area of 7.68 ha of Cerrado sensu stricto. Captured individuals were marked, equipped with a spool-and-line device, and released at the same capture point. The next day we followed the thread to record their daily movements and find their nests. To investigate microhabitat selection we compared habitat characteristics along trails of each studied species with general habitat characteristics of the study area. Although the mean 24-h distance was greater for N. lasiurus (mean ± SE: 41.9 ± 42.2 m, N=3) than for O. scotti (28.7 ± 14.2 m, N=6) this difference was not significant (Mann–Whitney test, U=26, P>0.6). We detected significant differences among observed microhabitats variables of both species and available microhabitat characteristics as determined by discriminant analysis (Wilks′s lambda F=3.001; df=14, 116; P<0.001). Both species were associated to microhabitat characteristics whose values differed markedly from the overall available habitat. Along the first canonical discriminant function of the DFA both them were associated with greater grass height than the mean height available and along the second axis N. lasiurus selected areas with higher fruit availability and more shelters than those selected by O. scotti. For stronger inferences regarding differential patterns of habitat utilization by Cerrado rodents we suggest the simultaneous use of both spool-and-line and standard trapping methods.     

Information-Theoretic Model Selection and Model Averaging for Closed-Population Capture-Recapture Studies

Specification of an appropriate model is critical to valid statistical inference. Given the “true model” for the data is unknown, the goal of model selection is to select a plausible approximating model that balances model bias and sampling variance. Model selection based on information criteria such as AIC or its variant AIC_c, or criteria like CAIC, has proven useful in a variety of contexts including the analysis of open-population capture-recapture data. These criteria have not been intensively evaluated for closed-population capture-recapture models, which are integer parameter models used to estimate population size (N), and there is concern that they will not perform well. To address this concern, we evaluated AIC, AIC_c, and CAIC model selection for closed-population capture-recapture models by empirically assessing the quality of inference for the population size parameter N. We found that AIC-, AIC_c-, and CAIC-selected models had smaller relative mean squared errors than randomly selected models, but that confidence interval coverage on N was poor unless unconditional variance estimates (which incorporate model uncertainty) were used to compute confidence intervals. Overall, AIC and AIC_c outperformed CAIC, and are preferred to CAIC for selection among the closed-population capture-recapture models we investigated. A model averaging approach to estimation, using AIC, AIC_c, or CAIC to estimate weights, was also investigated and proved superior to estimation using AIC-, AIC_c-, or CAIC-selected models. Our results suggested that, for model averaging, AIC or AIC_c should be favored over CAIC for estimating weights.     

The Sustainability of Wolverine Trapping Mortality in Southern Canada

Range declines, habitat connectivity, and trapping have created conservation concern for wolverines throughout their range in North America. Previous researchers used population models and observed estimates of survival and reproduction to infer that current trapping rates limit population growth, except perhaps in the far north where trapping rates are lower. Assessing the sustainability of trapping requires demographic and abundance data that are expensive to acquire and are therefore usually only achievable for small populations, which makes generalization risky. We surveyed wolverines over a large area of southern British Columbia and Alberta, Canada, used spatial capture-recapture models to estimate density, and calculated trapping kill rates using provincial fur harvest data. Wolverine density averaged 2 wolverines/1,000 km² and was positively related to spring snow cover and negatively related to road density. Observed annual trapping mortality was >8.4%/year. This level of mortality is unlikely to be sustainable except in rare cases where movement rates are high among sub-populations and sizable un-trapped refuges exist. Our results suggest wolverine trapping is not sustainable because our study area was fragmented by human and natural barriers and few large refuges existed. We recommend future wolverine trapping mortality be reduced by ≥50% throughout southern British Columbia and Alberta to promote population recovery. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Adult philopatry and dispersal in the field vol Microtus agrestis

Summary Using mark-recapture data, we related the movements of adult field voles to population density, sex ratio and population growth. Dispersal movements (defined as distances larger than 1 home range diameter) were few in both sexes; 4 out of 197 (2.0%) in males and 8 of 316 (2.5%) in females. The distance moved between sequential trapping periods was similar for males and females; the mean being 10.2 m and 9.0 m respectively. Both males and females moved larger distances during the breeding season than during the nonbreeding period. The distance moved between sequential trapping periods showed a strong negative relation to density, i.e. both sexes moved shorter distances at higher densities, but there were no differences between periods of increasing and declining population densities. These results contradict the dispersal predictions of all major hypotheses proposed to explain population fluctuations in small mammals. The dispersal patterns fit a geometric distribution, suggesting that competition is the primary factor determining the dispersal characteristics of this population.     

Assessment of the range of attraction of pheromone traps to Agriotes lineatus and Agriotes obscurus

• 1 The range of attraction of YATLOR pheromone traps was studied to gain information on the number of traps needed for mass trapping of males of two Agriotes species.
• 2 Male click beetles of the species Agriotes lineatus (L.) and Agriotes obscurus (L.) (25–30 individuals per release point) were marked and released at a distance of 2, 5, 10, 15, 20 and 60 m from a pheromone trap both along and opposite to the known prevailing wind direction. Traps were regularly inspected over approximately 1 month. The percentage of recaptured beetles was calculated and analyzed using analysis of variance. Maximum sampling ranges and effective sampling areas were calculated.
• 3 Averaged over all five trials and distances, approximately 40% of the released beetles (A. lineatus and A. obscurus) were recaptured. The percentage recapture of male adults was significantly affected by release distance, whereas no differences were found for species and release direction.
• 4 Males were recaptured from all release points and the percentage recapture decreased (in part significantly) with increasing distance from 76% (2 m) to 35% (15 m) and 9% (60 m), respectively. Most of the beetles were recaptured within the first 3 days after release, independent of the distance, except 60 m. The effective sampling area for A. lineatus was 1089 m² after 12 days and increased to 1735 m² after 30 days. Corresponding values for A. obscurus were considerably higher: 1518 m² for 12 days and 2633 m² for 30 days.
• 5 We conclude that the range of attraction of the pheromone traps for A. lineatus and A. obscurus is comparatively low, providing high percentage recapture only for release distances up to 10 m. Accordingly, any approach targeted on preventing mating by male mass trapping would require a dense network of pheromone traps.

     

Habitat selection by two species of small mammals in the Atlantic Forest,Brazil: Comparing results from live trapping and spool-and-line tracking

Habitat selection by small mammals is usually evaluated using data from live trapping, which provides little information about the movements of individuals. Few studies used movement data or compared the results of different sampling methods to study habitat selection by these animals. We evaluated habitat selection by the rodent Nectomys squamipes and the marsupial Micoureus paraguayanus in the Atlantic Forest of Brazil using the spool-and-line technique. We also determined if percentage of captures reflected the amount of movements in each habitat. Habitat selection was determined comparing use and availability of five habitat types at two spatial scales (movement paths and movement areas) using compositional analysis, which allowed ranking of habitats according to their relative use by animals. The use of available habitat types was non-random for both species at both spatial scales. The two species had contrasting habitat affinities directly related to their particular habits, with N. squamipes using predominantly the stream habitat, and M. paraguayanus using mainly the restinga forest habitat. Patterns of habitat selection were similar at both spatial scales probably due to the small size of movement areas, which may not represent habitat use at a broader scale. For both species, live trapping and movement data provided the same ranking in habitat use, demonstrating that simple capture indices may be used to study habitat selection by these species across different habitat types.     

The decay of similarity with geographical distance in parasite communities of vertebrate hosts

Aim The rate at which similarity in species composition decays with increasing distance was investigated among communities of parasitic helminths in different populations of the same host species. Rates of distance decay in similarity of parasite communities were compared between populations of fish and mammal hosts, which differ with respect to their vagility and potential to disperse parasite species over large distances. Location Data on helminth communities were compiled for several populations of three mammalian host species (Ondatra zibethicus, Procyon lotor and Canis latrans) and three fish host species (Perca flavescens, Catostomus commersoni and Esox lucius) from continental North America. Methods Distances between localities and similarity in the composition of helminth communities, the latter computed using the Jaccard index, were calculated for all possible pairs of host populations within each host species. Similarity values were then regressed against distance to see if they decayed at exponential rates, as reported for plant communities; the significance of the regressions was assessed using randomization tests. Results The number of hosts examined per population did not correlate with the number of helminth species found per population, and thus sampling effort is unlikely to have confounded the results. In four (two mammals and two fish) of the six host species, similarity in helminth communities decayed exponentially with distance. When the log of similarity is plotted against untransformed distance, the slopes obtained for the two fish species are lower than those obtained for the two mammalian host species. Main conclusions Similarity in the composition of parasite communities appears to decay exponentially with increasing distance in some host species, but not in all host species. The rate of decay is not necessarily associated with the vagility of the host. Although distance decay of similarity is generally occurring, it seems that other ecological processes, related either to the host or its habitat, can obscure it.     

Differential responses of three Agriotes click beetle species to pheromone traps

1 Previous work had suggested that adult male click beetles (Agriotes spp.) show differential responses to species specific pheromone traps. This hypothesis was tested using mark–release–recapture methods to estimate the maximum sampling range and the effective sampling area of traps for three species. 2 Captured beetles of the species Agriotes lineatus, Agriotes obscurus and Agriotes sputator were marked to show the direction of release, the distance of the release point from the trap and the replicate. Analysis of variance showed that there were significant differences in recapture rates between species and release distances. There were no significant differences between release direction and replicates. 3 Calculated linear speeds suggested differences in movement rates in the order: A. lineatus > A. obscurus > A. sputator. There were also substantial differences between the species in the maximum sampling ranges and effective sampling areas of the traps. These placed the species in the same order. 4 The results are used to estimate the minimum cost of mass trapping programmes to prevent males from mating, giving values of €165/ha/year (A. lineatus), €247.5/ha/year (A. obscurus) and €2343/ha/year (A. sputator). 5 Implications for the use of pheromone traps in wireworm pest management are discussed. It is concluded that current U.K. recommendations based on the cumulative total catch of the three species over a sampling season can be improved by considering the spatial relationships between the adult trapping system and larval distribution. The current constraint to this is the general inability to separate wireworms into species.