A field study of the effect of prebaiting on censusing by the capture-recapture method in a vole population

Populations of the vole, Clethrionomys rufocanus, in a lowland woodlot of Hokkaido were studied for the presence of effects of prebaiting on censusing by the capture-recapture method. A grid of 121 live-traps, spaced 5 m apart, was laid out on each of two plots, one of which alone was prebaited three days long. Owing to very high densities and great trap-efficiency, sufficient and favourable samples could be available for statistical analysis, except the trend of delayed catch for young in sampling. The population on either plot, however, proved to be markedly variable in catchability of unmarked animals in the course of trapping; while the probability of recapture was counted as invariable on the average from day to day, the recapture frequency was different between juveniles, subadults and adults. Needless to say, the catchability was distinctly greater for marked voles than for unmarked ones, whether prebaited or not, through the trapping period, except that the first-day catchability for unmarked ones on the prebaited plot seems not to be significantly lower than that for marked ones. Consequently, it turns out that the prebaiting has almost never helped to eliminate the important bias induced by differential trap-response of marked and unmarked animals; its contribution is only that the catchability for unmarked ones is slightly higher on the part of the prebaited plot on earlier days of the period. In accordance with the heterogeneous catchability, the Γ-form distribution analogous to the geometric could be applied with thorough fitness to the capture frequency in order to estimate the whole populations. The fact that the estimates are reliable, being not at any rate underestimates, was further confirmed by the result of a follow-up work conducted by means of the removal method with wider trap-spacing which brought forth distinct underestimation chiefly referable to unexposure to traps of the partial populations. The subject of unexposure was discussed by laying stress on the relation between minimum range length and trap-spacing.     

Adrenal analysis for critique of the social stress theory in natural populations of a montane vole

Summary The vole,Eothenomys smithi, in highlands of Shikoku was studied to disclose ups and downs of adrenal activities under the influence of the breeding level in natural populations. It has proved that the vole's adrenal in both sexes may be much more strongly by rise and fall of the fertility as well as by shift of the age structure in populations than the rat's. Therefore, as for perhaps all kinds of small mammals, we consider necessary to follow a pertinent procedure to avoid thoroughly these sources of error in studies to draw any conclusion for or against theChristian's theory. The present data analysed by such a method failed to afford any confirmative proofs; the fact that the pregnancy rate was increased in parallell to the density is against his view. However, never the whole content of the theory has been invalidated by these results.     

On the problem of trap-response types of small mammal populations

1. So far as the population data thus far amassed go, commensal rats in residential habitats are of type II; house mice may be of type I or III in fields but might show type II in residences, and fieral species or subspecies may range from type I to III. Altogether alloresponsive populations seem to be much commoner than the isoresponsive.
2. It is as yet little disclosed to what degree the differentiation of the type is correlated with speciation or subspeciation, even if only a proof that a subspecific population would hold type III as its fixed feature is given. The response types are supposed to be associated with the levels of population shyness. Hence some characters reflected in those may possibly be connected with sexual isolation provoking speciation.
3. Some awkward respects due to the heterogeneous trappability, depending upon the trap-response type, among a mixed population of marked and unmarked animals have been confirmed regarding the methodology for estimating parameters.

     

An approach to the edge effect in proof of the validity ofDice's assessment lines in small-mammal censusing

A second fieldwork for the sake of solving the edge-effect subject was carried out in almost the same ways and plans in summer of 1971 in succession to that of 1970. It was found out that the population density was reduced to two-thirds, but the data were sufficient to form the subsequent conclusions:

1. The initial population estimated by the removal census method turned out to be well compatible with the numbers of survivals and emigrants of marked voles calculated byJolly 's method.
2. The natural, entire home ranges of both sexes reached by determining observed range length and width with 24 voles, selected on rigid conditions, agreed with those of the range-conservative type in 1970 in both acreage and elliptic shape.
3. In the majority of the select specimens, considered to reveal the natural ranges, the random distribution in number of captures per station within range was proved by means of I_σ so far as such capture density as treated here is concerned. On the ground of the empirical evidence, the above range size approached by the routine observational method could be made surprisingly accordant with the range size in terms of r-values calculated byWierzbowska 's method.
4. From these proofs, the validity ofDice 's assessment lines makes evident, and besides, it could be further substantiated by use ofMarten 's notion of mouse-equivalents through the mediation ofWierzbowska 's method.
5. Consequently, I have largely come to the conclusion thatDice 's assessment lines stand on justifiable basis from theoretical and empirical angles.

     

Inquiry into effects of prebaiting on removal census in a vole population

A field work with two plots of grid, a snap trap being set on each station spaced 5 m apart, was executed in the summer of 1968 to evaluate prebaiting in census trapping by comparing the result in one plot, prebaited for three days, with that in the other not prebaited. Since the population was as high as some 230 per acre on the average in density and formed of the vole, Microtus montebelli, alone, sufficient samples were gathered irrespective of the plot size as small as 50×50 m. Owning to the circumstances, multiple collisions inflicted so intense influence on sampling especially in the prebaited plot that z-equation for census adjusted to the effect was well applicable to the data in either plot. In sampling, the fact that small voles are apt to be caught later than large voles was statistically evidenced in either sex, and yet any proof that males tend to be caught prior to females was not offered. It was ascertained in either plot that the daily catch was realized according to the same rule through the whole period of trapping in both external belt and internal square within the plot; hence it follows that no considerable immigration occurred. One of the beneficial effects of prebaiting is sure to be that the probability of capture was markedly enhanced in the prebaited plot, and a second is supposed, though inconclusively, to be that a good sampling could be executed consistently through the census period giving rise to no inordinate catches perhaps due to heterogeneous sampling as was seen in earlier days in the not prebaited plot. The supposition has derived from the condition that most of the whole population is trappable, which is established by interrelation among population density, size of home range and trap spacing. It was suggested that the effect of prebaiting should be evaluated from the view-point of the interrelation, because the basic utility of prebaiting consists in that it may help to our utlimate purpose to estimate the whole population.     

Estimating effective population size using RADseq: Effects of SNP selection and sample size

Effective population size (N_e) is a key parameter of population genetics. However, N_e remains challenging to estimate for natural populations as several factors are likely to bias estimates. These factors include sampling design, sequencing method, and data filtering. One issue inherent to the restriction site‐associated DNA sequencing (RADseq) protocol is missing data and SNP selection criteria (e.g., minimum minor allele frequency, number of SNPs). To evaluate the potential impact of SNP selection criteria on N_e estimates (Linkage Disequilibrium method) we used RADseq data for a nonmodel species, the thornback ray. In this data set, the inbreeding coefficient F_IS was positively correlated with the amount of missing data, implying data were missing nonrandomly. The precision of N_eestimates decreased with the number of SNPs. Mean N_e estimates (averaged across 50 random data sets with2000 SNPs) ranged between 237 and 1784. Increasing the percentage of missing data from 25% to 50% increased N_e estimates between 82% and 120%, while increasing the minor allele frequency (MAF) threshold from 0.01 to 0.1 decreased estimates between 71% and 75%. Considering these effects is important when interpreting RADseq data‐derived estimates of effective population size in empirical studies.     

A method for the estimation of parameters for natural stage-structured populations

A relatively simple method is proposed for the estimation of parameters of stage-structured populations from sample data for situation where (a) unit time survival rates may vary with time, and (b) the distribution of entry times to stage 1 is too complicated to be fitted with a simple parametric model such as a normal or gamma distribution. The key aspects of this model are that the entry time distribution is approximated by an exponential function withp parameters, the unit time survival rates in stages are approximated by anr parameter exponential polynomial in the stage number, and the durations of stages are assumed to be the same for all individuals. The new method is applied to four Zooplankton data sets, with parametric bootstrapping used to assess the bias and variation in estimates. It is concluded that good estimates of demographic parameters from stagefrequency data from natural populations will usually only be possible if extra information such as the durations of stages is known.     

Shifting trends: detecting environmentally mediated regulation in long-lived marine vertebrates using time-series data

Assessing the status and trends in animal populations is essential for effective species conservation and management practices. However, unless time-series abundance data demonstrate rapid and reliable fluctuations, objective appraisal of directionality of trends is problematic. We adopted a multiple-working hypotheses approach based on information-theoretic and Bayesian multi-model inference to examine the population trends and form of intrinsic regulation demonstrated by a long-lived species, the southern elephant seal. We also determined the evidence for density dependence in 11 other well-studied marine mammal species. (1) We tested the type of population regulation for elephant seals from Marion Island (1986–2004) and from 11 other marine mammal species, and (2) we described the trends and behavior of the 19-year population time series at Marion Island to identify changes in population trends. We contrasted five plausible trend models using information-theoretic and Bayesian-inference estimates of model parsimony. Our analyses identified two distinct phases of population growth for this population with the inflexion occurring in 1998. Thus, the population decreased between 1986 and 1997 (−3.7% per annum) and increased between 1997 and 2004 (1.9% per annum). An index of environmental stochasticity, the Southern Oscillation Index, explained some of the variance in r and N. We determined analytically that there was good evidence for density dependence in the Marion Island population and that density dependence was widespread among marine mammal species (67% of species showed evidence for population regulation). This approach demonstrates the potential functionality of a relatively simple technique that can be applied to short time series to identify the type of regulation, and the uncertainty associated with the phenomenon, operating in populations of large mammals.     

These are not the trends you are looking for: poorly calibrated single-pass electrofishing data can bias estimates of trends in fish abundance

There is increasing interest in the potential of single-pass and timed electrofishing to assess status and trends in fish populations. However, where capture probability varies over time, there is a risk that uncalibrated electrofishing data could fail to detect, or provide biased estimates of trends. This study analysed a long-term electrofishing dataset collected over 50 years in an intensively studied catchment where egg deposition and emigrant production declined by c. 82% and 35% over the same time. The electrofishing data were used to illustrate the effects of changing capture probability on estimated trends in juvenile Atlantic salmon Salmo salar abundance. Temporal variability in capture probability was modelled. Trends in abundance were then estimated from uncalibrated single-pass electrofishing count data and compared with estimates from data calibrated for capture probability. The calibrated data revealed significant declines in S. salar fry (age 0) and parr (age ≥ 1) abundance. However, the trend estimates from the uncalibrated data were positively biased and not significant. Exploration of alternative (realistic) scenarios with different trends in true abundance and capture probability suggests that uncalibrated electrofishing data can provide very misleading estimates of trends. The problem is exacerbated in data where capture probability is low. It is recommended that single-pass and timed electrofishing methods should not be used to assess trends in fish populations without regular (annual) calibration.     

Evaluation of three methods to estimate density and detectability from roadside point counts

Roadside point counts are often used to estimate trends of bird populations. The use of aural counts of birds without adjustment for detection probability, however, can lead to incorrect population trend estimates. We compared precision of estimates of density and detectability of whistling northern bobwhites (Colinus virginianus) using distance sampling, independent double-observer, and removal methods from roadside surveys. Two observers independently recorded each whistling bird heard, distance from the observer, and time of first detection at 362 call-count stops in Ohio. We examined models that included covariates for year and observer effects for each method and distance from observer effects for the double-observer and removal methods using Akaike's Information Criterion (AIC). The best model of detectability from distance sampling included observer and year effects. The best models from the removal and double-observer techniques included observer and distance effects. All 3 methods provided precise estimates of detection probability (CV = 2.4–4.4%) with a range of detectability of 0.44–0.95 for a 6-min survey. Density estimates from double-observer surveys had the lowest coefficient of variation (2005 = 3.2%, 2006 = 1.7%), but the removal method also provided precise estimates of density (2005 CV = 3.4%, 2006 CV = 4.8%), and density estimates from distance sampling were less precise (2005 CV = 9.6%, 2006 CV = 7.9%). Assumptions of distance sampling were violated in our study because probability of detecting bobwhites near the observer was <1 or the roadside survey points were not randomly distributed with respect to the birds. Distances also were not consistently recorded by individual members of observer pairs. Although double-observer surveys provided more precise estimates, we recommend using the removal method to estimate detectability and abundance of bobwhites. The removal method provided precise estimates of density and detection probability and requires half the personnel time as double-observer surveys. Furthermore, the likelihood of meeting model assumptions is higher for the removal survey than with independent double-observers. © 2011 The Wildlife Society.     

Do island plant populations really have lower genetic variation than mainland populations? Effects of selection and distribution range on genetic diversity estimates

Ecological and evolutionary studies largely assume that island populations display low levels of neutral genetic variation. However, this notion has only been formally tested in a few cases involving plant taxa, and the confounding effect of selection on genetic diversity (GD) estimates based on putatively neutral markers has typically been overlooked. Here, we generated nuclear microsatellite and plastid DNA sequence data in Periploca laevigata, a plant taxon with an island–mainland distribution area, to (i) investigate whether selection affects GD estimates of populations across contrasting habitats; and (ii) test the long‐standing idea that island populations have lower GD than their mainland counterparts. Plastid data showed that colonization of the Canary Islands promoted strong lineage divergence within P. laevigata, which was accompanied by selective sweeps at several nuclear microsatellite loci. Inclusion of loci affected by strong divergent selection produced a significant downward bias in the GD estimates of the mainland lineage, but such underestimates were substantial (>14%) only when more than one loci under selection were included in the computations. When loci affected by selection were removed, we did not find evidence that insular Periploca populations have less GD than their mainland counterparts. The analysis of data obtained from a comprehensive literature survey reinforced this result, as overall comparisons of GD estimates between island and mainland populations were not significant across plant taxa (N = 66), with the only exception of island endemics with narrow distributions. This study suggests that identification and removal of markers potentially affected by selection should be routinely implemented in estimates of GD, particularly if different lineages are compared. Furthermore, it provides compelling evidence that the expectation of low GD cannot be generalized to island plant populations.     

Size-frequency estimates of secondary production byMysis relicta in Lakes Michigan and Huron

Data from five Great Lakes studies ofMysis relicta populations were reanalyzed to calculate secondary production estimates using the size-frequency method. Production estimates (P) ranged from 0.25 to 3.2 g dry weight m^–2 yr^–1. Average annual biomass {xxB} and mean annual density (xxD) were 0.11–1.11 g dry weight/ m² and 25–434 animals/ m², respectively. P:{xxB} ratios varied only between 2.2 and 3.3. Maximum and minimum biomass values within a study varied by a factor of 519 for one study but by less than 17 for the others. Highest estimates of P, {xxB} and {xxD} were calculated for collections from a 50-m station in Lake Michigan despite the larger populations suspected to be present at greater depths sampled in the other studies. These conservative estimates provide a basis for scaling trophic interactions involvingM. relicta and emphasize findings by previous workers that night-time sampling with vertical net hauls is the best available technique for quantitative studies ofM. relicta populations in the Great Lakes.     

A computer simulation study on the number of loci and trees required to estimate genetic variability in cacao (Theobroma cacao L.)

Current methods for measures of genetic diversity of populations and germplasm collections are often based on statistics calculated from molecular markers. The objective of this study was to investigate the precision and accuracy of the most common estimators of genetic variability and population structure, as calculated from simple sequence repeat (SSR) marker data from cacao (Theobroma cacao L.). Computer simulated genomes of replicate populations were generated from initial allele frequencies estimated using SSR data from cacao accessions in a collection. The simulated genomes consisted of ten linkage groups of 100 cM in length each. Heterozygosity, gene diversity and the F statistics were studied as a function of number of loci and trees sampled. The results showed that relatively small random samples of trees were needed to achieve consistency in the observed estimations. In contrast, very large random samples of loci per linkage group were required to enable reliable inferences on the whole genome. Precision of estimates was increased by more than 50% with an increase in sample size from one to five loci per linkage group or 50 per genome, and up to 70% with ten loci per linkage group, or equivalently, 100 loci per genome. The use of fewer, highly polymorphic loci to analyze genetic variability led to estimates with substantially smaller variance but with an upward bias. Nevertheless, the relative differences of estimates among populations were generally consistent for the different levels of polymorphism considered.     

Impact of removal on occupancy patterns of the invasive rainbow lorikeet (Trichoglossus moluccanus) in Tasmania

Introduced species may threaten both biodiversity and agriculture, necessitating an understanding on the factors that influence their distribution, and the efficacy of control measures. In Tasmania, Australia, the introduced rainbow lorikeet (Trichoglossus moluccanus) may be widespread, but data on where they occur and the efficacy of control methods are limited. We used an occupancy modelling framework (presence–absence data) to undertake a survey of two populations of invasive rainbow lorikeets to: (i) understand their distribution across the north and south of the island, and (ii) evaluate the impact of removing birds from the southern population by quantifying occupancy before (2016) and after (2018) removal. The best model explaining occupancy in both populations included a negative relationship with distance from central urban areas. We found no change in site occupancy or detectability in the southern population after removal of 208 birds (potentially comprising >50% of their original population size). This result may be explained by one of three possibilities: (i) the population is larger than previously thought, (ii) the population recovered quickly after reduction, or (iii) removal of birds reduced population density but not area of occupancy. We highlight the importance of urban habitats for the invasive rainbow lorikeet and suggest that alternative methods (e.g. abundance/density-based monitoring) may better detect impacts of removal.     

A model for analyzing insect stage-frequency data when mortality varies with time

A model is developed for the analysis of insect stage-frequency data which may be applied to populations with age-dependent mortality. The analysis of stage-frequency data is divided into two steps. In the first step, the number of different mortality rates and their values are estimated. The second step provides estimates of developmental rates and variances for each developmental stage and in addition provides estimates of the number of recruits to each stage. The model may be used both in analysis and prediction of insect stage frequencies. Hence, in addition to estimating developmental and mortality rates from stage-frequency data, it may also be used as a simulation model for an insect population. The model is applied to two populations of Hemileuca oliviaeCockerell , a lepidopterous pest of New Mexico grasslands. The model identifies, in the two populations, different mortality rates that are related to plant productivity.     

Estimating population size using single‐nucleotide polymorphism‐based pedigree data

Reliable population estimates are an important aspect of sustainable wildlife management and conservation but can be difficult to obtain for rare and elusive species. Here, we test a new census method based on pedigree reconstruction recently developed by Creel and Rosenblatt (2013). Using a panel of 96 single‐nucleotide polymorphisms (SNPs), we genotyped fecal samples from two Swedish brown bear populations for pedigree reconstruction. Based on 433 genotypes from central Sweden (CS) and 265 from northern Sweden (NS), the population estimates (N = 630 for CS, N = 408 for NS) fell within the 95% CI of the official estimates. The precision and accuracy improved with increasing sampling intensity. Like genetic capture–mark–recapture methods, this method can be applied to data from a single sampling session. Pedigree reconstruction combined with noninvasive genetic sampling may thus augment population estimates, particularly for rare and elusive species for which sampling may be challenging.     

Estimating regional landbird populations from enhanced North American Breeding Bird Surveys

Estimating the size of bird populations is central to effective conservation planning and prudent management. I updated estimated regional bird populations for the East Gulf Coastal Plain of Mississippi using data from 275 North American Breeding Bird Surveys from 2009 to 2013. However, regional bird populations estimated from count surveys of breeding birds may be biased due to lack of empirical knowledge of the distance at which a species is effectively detected and the probability of detecting a species if it is present. I used data recorded within two distance classes (0–50 m and >50–400 m) and three 1‐min time intervals on 130 Breeding Bird Surveys to estimate detection probability and effective detection distance for 77 species. Incorporating these empirical estimates of detection probability and detection distance resulted in estimated regional populations for these species that were markedly greater than regional populations estimated without species‐specific estimates of detection parameters. Using the same Breeding Bird Survey data, I also estimated probability of site occupancy for 66 species and extrapolated this to the proportion of area occupied in the East Gulf Coastal Plain of Mississippi. I combined the area occupied with the reported range of breeding territory size for 54 species to obtain independent estimates of regional bird populations. Although the true population of these species is unknown, estimated populations that incorporated empirical estimates of detection probability and detection distance were more likely to be within the range of independently estimated, occupancy‐based, regional population estimates than were population estimates that lacked empirical detection and distance information.     

Effects of Transect Selection and Seasonality on Lemur Density Estimates in Southeastern Madagascar

I investigated how transect type (trails vs. cut transects) and seasonality influenced density estimates for 5 lemur taxa (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus) in the Vohibola III Classified Forest in SE Madagascar. I surveyed tree height and diameter and lemur populations from June 1 to December 28, 2004 along 2 1250-m trails local people used and 2 1250-m transects cut parallel to the trails in primary rain forest. Despite dendrometric variations within and between trails and transects, only density estimates of Hapalemur griseus griseus differed significantly by transect type. The spatial variation may be a result of removal by local people of giant bamboo, which is the main food for Hapalemur griseus griseus, along trails. Conversely, seasonality influenced density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus. The temporal variations may be related to seasonal torpor for Cheirogaleus major and increased detection probabilities during periods of fruit exploitation for Eulemur rubriventer, Microcebus rufus, and Hapalemur griseus griseus. Transect type and seasonality did not affect density estimates for Avahi laniger, which may be related to the highly folivorous and low-energy diet of the nocturnal lemur. Researchers surveying lemurs along line transects should be aware that transect selection may influence density estimates for Hapalemur griseus griseus and that seasonality may influence density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus.     

Evaluating sample size to estimate genetic management metrics in the genomics era

Inbreeding and relationship metrics among and within populations are useful measures for genetic management of wild populations, but accuracy and precision of estimates can be influenced by the number of individual genotypes analysed. Biologists are confronted with varied advice regarding the sample size necessary for reliable estimates when using genomic tools. We developed a simulation framework to identify the optimal sample size for three widely used metrics to enable quantification of expected variance and relative bias of estimates and a comparison of results among populations. We applied this approach to analyse empirical genomic data for 30 individuals from each of four different free‐ranging Rocky Mountain bighorn sheep (Ovis canadensis canadensis) populations in Montana and Wyoming, USA, through cross‐species application of an Ovine array and analysis of approximately 14,000 single nucleotide polymorphisms (SNPs) after filtering. We examined intra‐ and interpopulation relationships using kinship and identity by state metrics, as well as F_ST between populations. By evaluating our simulation results, we concluded that a sample size of 25 was adequate for assessing these metrics using the Ovine array to genotype Rocky Mountain bighorn sheep herds. However, we conclude that a universal sample size rule may not be able to sufficiently address the complexities that impact genomic kinship and inbreeding estimates. Thus, we recommend that a pilot study and sample size simulation using R code we developed that includes empirical genotypes from a subset of populations of interest would be an effective approach to ensure rigour in estimating genomic kinship and population differentiation.     

Investigation into the edge effect by use of capture-recapture data in a vole population

A substantial explication of the edge effect has been attempted by use of capture-recapture data for a vole population (Microtus montebelli), gathered intwo plots of 100×100 m or less during 12 days, cheked twice daily, in August 1970; the sample was quite sufficient for the aim. The edge effect as guessed by increased catch per trap is usually suspected to ensue from range-settlers in the outside boundary strip of a plot and immigrants. But by a theoretical analysis I could attain a tentative conclusion that no increased catch per trap will occur unless any invasion takes place. Then it follows that, apart from the effect of invasion, the role of the adjoining outside settlers in the edge effect is essentially required to be studied in the light of knowledge on the truth of size and shift in home range. The variation in range behavior for 183 adult voles, captured 6 times or more, could be grouped into eight types, of which the range-conservative type possessed 52% of the sample and the group of the type was justly utilized for giving averages of range size. Besides, it was seen from the observed frequency of types that a considerable number of immigrants onto the census plot were induced perhaps being allured by trap baits, but the majority of them proved to be assigned to the voles that have their ranges inside the assessment line ofDice ; the rest referable to effective immigrants was only a few (7%). I could perceive no reason such as disproves the idea ofDice 's additional boundary strip. Viewed from maps of ingress shift of ranges, the effect of ingress must have been greater in the outer trap rows than in the inner within the plot, so that it might well be called edge effect in general; such effect, however, is seen gradually diminishing toward the center, and hence it is almost unlikely that one should find any clear-cut intra-plot assessment lines demarcating such an inner square as quite free from edge effects. Averages of observed range length and width (ORL and ORW), as reliable measures for the true range size, were determined from the above group of specimens; as a result, the remarkable concept of elliptic range shape was established by regarding ORL as long axis and ORW as short one, and, directly from these averages, the mean range sizes worked out at 0.04 for females and 0.09 for males in acreage which proved to be surprisingly well agreeable with those of isotope-revealed ranges for voles given byGodfrey (1954) andAmbrose (1969). The catchability for marked voles () was estimated by the maximum likelihood method by use ofJolly 's formulae (1965), but that for unmarked ones () was made by the regression census formula; as a result it was shown that the population was clearly of π>p type and that the trap-experience that voles underwent one month or more ago can make them retain as high catchability as π.