The Laxmann’s shrew Sorex caecutiens. Step towards "sociality"

The Laxmann’s shrew Sorex caecutiens. 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Step towards "sociality" Nikolay A. Shchipanov, Alexey A. Kalinin, Tatiana Demidova This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6810348/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Sorex are commonly considered highly aggressive territorial animals. Our research shows that species in this genus can significantly differ in the extent of home ranges overlap and their inclination to contact with conspecifics. We assay space usage and propensity to engage in social contacts in the Laxmann’s shrew, S. caecutiens in comparison with the common shrew, Sorex araneus . Although both studied species shared area under increased population density, the Laxmann’s shrew reveal significantly larger shared area, and the proportion of shared area in this species significantly exceeded that was expected for random co-location of animals. In the common shrew shared area corresponded to expectation for random co-location of animals. In a preference test with a conspecific in wire-mesh container we found, that whereas the common shrew was rather indifferent, the Laxmann’s shrew reveal propensity to contact with conspecific. When estimated a suit of variables obtained in trials with respect to pace-of-life continuum, we found that Laxmann's shrew corresponded "slow species"; it was more shy, less active in general, more thorough explorer, with greater propensity to engage in social contacts. We assume that the Laxmann's shrew made at least a step towards more complex social structure. shrews home range preference test social contacts sociality Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The studying of social evolution suggests understanding how “previously selfish, free-living individuals join together to form a group resembling an individual in its own right”, passing thee stages – social group formation, social group maintenance, and social group transformation (Bourke 2011 ). There are a number of theoretical models regarded partner choice, direct and indirect reciprocity, etc. (eg. Novak 2006, 2012; Fuentes 2025 ; Chassy et al., 2025 , and other), and in all scenarios, evolution begins when individuals of the same species can remain in contact outside the mating season. In any case the first step to sociality is tolerance of members of one's own species, and hence the presence of a common space and some tendency towards social contact. This suggests that such a move might manifest itself in as significant extend of shared area and a greater propensity for social contact. Among placental mammals, shrews are of particular interest for such a step. These insectivorous mammals can be considered as the life form of the first placental mammals (Luo et al. 2011 ). Since the majority of shrews considered solitary living (Churchfield 1990 , Rychlik 1998 ), it is reasonable to assume that that the ancestor of placentals was likely solitary, even though Eulipotyphla are not more basal than other mammalian orders (Lukas, Clutton-Brock, 2013 ). It is of interest to study more shrew species in wild with respect to the “biodiversity of social systems to be able to perform meaningful comparative studies about the evolution of sociality” (Valomy et al. 2015 ). On the other hand, it is the third most diverse order of mammals after rodents and bats, with more than 370 species family shrews, but we have very few knowledge of the social organization of most species. The common belief that all shrews are equally asocial animals may be the result of insufficient study of their behaviour and space use (Valomy et al. 2015 ). At least, some shrews of the subfamily Crocidurinae are more tolerant, and often shared area, whereas subfamily Soricinae is mainly represented by solitary, aggressive species (Rychlik 1998 ). More detailed studies show differences in communication and space use between closely related species. For example, Eurasian water shrew, Neomys fodiens , is known as highly aggressive strictly solitary, territorial animal, while Iberian water shrew, N. anomalus exhibited communal using of space and amicable behaviours (Krushinska, Rychlik 1993 ; Krushinska et al. 1994 ; Rychlik, Zwolak, 2005 , 2006 ). Of all the shrews, red-toothed shrews, Sorex , were regarded as the most aggressive and least social animals (rev. Churchfield 1990 ; rev. Rychlik 1998 ). It should be noted, however, that our understanding of social behaviour in this rather diverse genus, which contains at least 142 known species, is based chiefly on the study of the common shrew (Crowcroft 1955 , 1957 ). Due to technical difficulties in handling shrews, the extreme susceptibility of animals to traps and captivity, both the space use, and behaviour of most species remain poorly understood. However, interactions in paired contact tests with animals from wild indicate that at least four Sorex species found in central Russia differ in their aggressiveness towards conspecifics (Shchipanov et al. 1998 ). Of those species the Laxmann’s shrew S. caecutiens was found the most “social”: the species exhibited rather diverse patterns of integrative interactions, which accounted for more than 33% of all contacts. The amicable interactions were rather diverse and included such patterns as climbing on the partner, crawling under the partner, sitting side by side in tactile contact, which were absent in common shrews (Shchipanov et al. 1998 ). It was of interest to obtain more quantitative characteristics of behaviour to compare S. araneus and S. caecutiens . The Laxmann’s shrew is a numerous species with a wide range, inhabiting from the north-eastern Europe to Far-East, however its behaviour and space use are barely studied. In Central Russia, the preferred habitat of S. caecutiens is mossy coniferous forests, where Laxmann’s shrews of their first year of life established dense groups with shared home ranges (Shchipanov et al. 2001 , 2005 ). We monitor small mammals using capture-mark-recapture (CMR) technique in various habitats, and in particular in mossy coniferous forest, which permit us to study space use in this species in a dense local population. Behaviour of this species, which was earlier studied in contact test gives only general picture of interactions of individuals (Shchipanov et al. 1998 ). Now we for the first-time assay sociability of the Laxmann’s shrew from wild in Crowley’s (2000, 2004) preference test. Previously, when studying sociability in this test in the common shrew, we assessed personality traits such as activity, boldness, exploratory activity, and propensity for social contacts (Shchipanov, Demidova 2022 ). The test allows us to quantify behavioural traits, and is applicable for comparative studies of behaviour for both individuals and species. Social behaviours that benefit survival can be associated with life histories in terms of the pace-of-life syndrome (hereafter POLS) (Silk, Hodgson 2021 ). Note that the above mentioned behavioral traits are important for both interspecific and intraspecific differentiation in terms of POLS schematic behavioral profile (Réale et al. 2010 ). In this study, we intend 1) quantitatively compare shared area in the common and the Laxmann's shrews, 2) report the results of the first study of S. caecutiens in a preference test, and 3) compare the results of preference tests in S. caecutiens and S. araneus in terms of POLS. Materials and methods Study area and animal trapping We monitor small mammals in five habitats in the central part of European Russia in the vicinity of the Bakanovo village, the Staritsa Region of Tver Oblast’, N 56°18′; E 34°53′ (Shchipanov, Kalinin 2024b ). The Laxmann's shrew is more numerous in mixed coniferous (spruce and pine) mossy old-growth forest, and is also common, but not numerous, in the adjacent forest area that was cleared in 1998 and is now overgrown with young birch forest. In other habitats surveyed, in a young forest on an abandoned field, on a former pasture and on the territory of an abandoned village, S. caecutiens was sparse, with only single individuals observed in some years. The common shrew contrary was rare in the mossy forest and more numerous in grassland habitats (Shchipanov, Kalinin 2024 a). The traps in all the habitats except of village area was installed as lines of 50 traps with 7.5 metres between traps, and as grid in the village area because of the configuration of plot available to monitoring. On the grid the traps were placed as four lines of 18 traps and one line of 15 traps with 7.5 m interval between traps in the lines, and 10 m distance between the lines. We used special trapping protocol, which was tailored to shrews (Shchipanov et al. 2000 , 2005 ). This imply using original live-trap, with wire treadle (the chart was given in above-mentioned articles), oat flakes with unrefined sunflower oil as bait, and daily exposition of operative traps shortened to 1.5 h after daily activation. While checking traps animals inspected, marked, if necessary, weighed, their position was recorded, and they were released at the place of capture. Animals were marked by toe-clipping. Unfortunately, this remained the only reliable way of mass individual lifelong marking of shrews. Note, we minimise harm to individual using numbers which require minimal number of toes to be clipped. This way of marking did not influenced survival of shrews (Shchipanov et al. 2005 ; Shchipanov et al. 2022 ). This regimen of inspection minimizes mortality in traps. No animal died in a trap in this study. After inspection, the traps remained at their position to be available for visits but were not operational. Thus, animals could move over 90% of daytime for free. This allows us to consider each capture of an individual as independent and reflective of its actual presence on the spot. For the behaviour study, we used common shrews from the village area and Laxmann’s shrews from the forest. Animals for behavioral testing were removed from traps during regular trap inspections at 8:00–9:30 AM, placed in cages with abundant food (mealworms, grasshoppers, fly larvae, and chicken hearts), subjected to the test, and released at the catch sites no later than 11 AM in case of S. araneus , and 12–13 in case of S. caecutiens . The greater time in the latter shrew resulted from a more distant sampling area. After the tests animals were released at a place of capture. Animals All the data in this study are given for the first-year shrews, since a complete population turnover in shrews takes one year, and first-year animals make up the vast majority of the population by mid-summer (Churchfield, 1990 ; Kalinin et al., 2008). Space sharing in both the common and the Laxmann's shrews was estimated using data collected in coniferous mossy forest and adjoined young birch forest at the cleared area, in 2014–2024. A total of 223 S. caecutiens individuals were captured 614 times, and 438 S. araneus individuals were captured 984 times. Of those 124 S. caecutiens and 252 S. araneus were captured more than twice and were regarded to be resident. Behaviour of Laxmann’s shrew was studied in 2024, in July. Fourteen Laxmann’s shrews from the forest plot were used in trials. In common shrews we used data for the first trial of 20 animals from the village area, since we conducted ongoing annual surveillance of the common shrew behaviour in this habitat. The trials with common shrews, which were used for comparison, were conducted during the same time period as the Laxmann’s shrew, in July 2024. The numbers, which was estimated as number of residents per 100 traps (Shchipanov, Kalinin 2024 b), in the period, when animals were taken for tests, made 22 ind./100 traps (11 on line of 50 traps) in the Lxmann’s shrew in the forest, and 23 ind./100 traps (on average 4.14 on line of 18 traps on grid) in the common shrew. Behavioral tests To study behaviour in common shrew, we used modified Crawley’s preference test. It implies 5 min trials in a single 50 × 50 cm chamber with specified virtual zones. The test has been described in details in our previous publications (Shchipanov, Demidova 2020 ; 2022 ). Briefly, two wire-mesh containers (cylindrical, 10 cm high and 8 cm in diameter) with a non-transparent cover were placed at a fixed position on the diagonal of the arena. One of the containers was with a stimulating animal, and the other remained empty. The focal animal could have olfactory, acoustic and visual communication with the stimulator through the mesh wall of the container. The study was conducted in an isolated room with artificial lighting. The brightness of the light was approximately equal to natural light outdoors. The focal animal was placed in the centre of arena in plastic glass, and released manually. The operator left the room after the focal animal was released, and the trial was conducted in the operator's absence. The orientation of the arena position towards the cardinal points was altered by us randomly between trials. The stimulating animal, unfamiliar to a focal individual, was placed in a container immediately prior to a trial. Because we were limited by the number of animals available for testing, we used both focal and stimulating animals from the same habitat. We considered animals with distance between the activity centres above median value of the distance between border captures in the entire sample (in case of both species it made approximately 60 m) as "unfamiliar". In this project we used for analysis only first (for both focal and stimulating animal) trials. Trials were video recorded by means of a Sony Handycam in MPEG-2 format, and the videos were analysed in the EthoVision XT (Noldus) software. Distance travelled, time spent in an area and mean velocity were determined for the border zone (BZ) (a 3-cm band along the walls), the zone of possible contact with the stimulating animal (around the container within a strip 3 cm from the edge) (AZ), the free zone similarly sized near the empty container (FZ), the tops (covers) of the containers with an animal (AC), and the free container (FC), and the area of the arena that excluded the above-mentioned zones, the central zone (CZ) (Fig. 1 ). Analyses Normal distribution was studied in Shapiro-Wilk test for p > 0.15. Variability of normally distributed data were shown as ± SE unless otherwise stated. Shared space Shared space was estimated as the proportion ( P ) of traps in an individual's home range that were visited by other neighbouring resident shrews. It was calculated as proportion of traps ( P ) visited by two and more resident shrews (animals with two and more than captures) – shared traps ( T s ), to the total number of traps ( T T ) visited by residents: P = T s / T T . (1) It has been found in common shrews, that shared area in yearling conspecifics increased with the number of shrews lived at the observed area due to random overlap (Shchipanov, 2021 ). Therefore, to avoid bias related to the different population abundance, we compared proportions of shared traps plotted it against number of resident shrews (animals with two and more captures) revealed in a habitat (on a line of 50 traps, 375 m). The study was performed as analysis of regressions. To understand if the proportions P resulted from the random coincidence of visited spots, we tested the null hypothesis about random distribution of visits in traps. To test the hypothesis, we simulated the expected proportion of shared traps, P e . The simulation assumes that each trap has an equal probability of receiving one of the 50 trap numbers in a trial. The total number of trials ( t T ) equaled the sum of traps (trap numbers), visited by all residents on a line of 50 traps in a given period. It could be calculated as number of residents, N , multiplied by mean number of traps, m t , visited by resident individual: t T = m t N . After receiving n random traps numbered from 1 to 50 in n trials, ( n = t T ), we counted the total number of trap-numbers obtained, as well as the number of traps-numbers that appeared two or more times. Calculations were performed using EXCEL's built-in randomization. After 1000 iterations we got average number of visited traps, n t , and average number of traps visited by two and more individuals n J , and calculated P e as P e = n J / n t (2) The expected proportion was calculated for number of animals, N , ranged from 5 to 31 (min and max N revealed on trapping lines) with taking into account average number of traps visited by mean individual per trapping session. In S. araneus the average number of traps (trap-numbers), m t , visited during a trapping session made 429 trap-numbers (sum for all trapping sessions) divided by 252 residents (revealed for all trapping sessions), or 1.70 traps visited by mean individual per a trapping session. In S. caecutiens m t made 254/124 = 2.05 traps visited by mean individual per a trapping session. Note that the different number of traps on a home range of residents, m t , influenced the number of trials in simulation, thus expected P e in S. araneus and S. caecutiens are not equal for equal N . Actual and expected proportions were compared in sign test. To assess whether trap sharing differed significantly under similar population size, we compared the proportion of traps shared between the medium (5–10 individuals per line) and high (over 10 individuals) abundance classes. The data sets (proportions of shared traps) in the cases were found normally distributed in Shapiro-Wilk test. Therefore, we studied differences in proportion of shared traps with application t -test for independent variables (groups). Since the proportions of shared traps and captures in shared traps were normally distributed, we assessed their goodness of fit using Pearson correlation, assuming that if there was no significant disproportion, we would find a significant correlation. These analyses were performed in STATISTICA 8 (StatSoft, USA). Behaviours We studied the full set of variables obtained during the trials: distances, velocity and duration of stay in the specified above virtual zones. To assay whether it was some preference of contact to social versus non-social object we calculated indexes of preference in distance I d and in time I T as: I X =X AZ / (X AZ + X FZ ) – 0.5, (3) where X is variable in question, either distance – d, or time – T, in the zone of container with conspecific (AZ) or in the zone of empty container (FZ). The index ranges from − 0.5 to 0.5. It tends to zero when the presence of the stimulus animal in the container has no significance for the focal individual; it is positive when the focal individual shows an interest in the partner, and negative when it avoids interact with a partner. Comparing indexes with null was performed in sign test. Normally distributed distances travelled within zones were compared between zones of the same species using Student's t-tests for dependent variables, as overall activity differed from trial to trial. Duration of stay in various zones of the arena was compared in one-way ANOVA, with p -value obtained in Tukey post-hock test. Residuals were found normally distributed. To understand whether there were significant differences in behavioral variables between species, we compared the suit of studied characteristics using Kruskal-Wallis ANOVA since a number of variables were not normally distributed. All statistical tests were performed in Statistica 8 (StatSoft, USA). To reveal tendency in bias in behavioral patterns in the studied species we used weighted deviation from general median value ( M G ) of a variable calculated as: d XS = ( M XS – M XG ) / M XG , (4) where d XS is deviation of variable X in species in question (either S. araneus or S. caecutiens ), M XS – median value of variable X in species (S) in question, and M XG is median value of variable X in generalized (G) sample of both species. Positive deviation indicates a predominance in the compared variable, and a negative value indicates a loss in the variable compared with the median value in generalized sample. Results Shared space In both Laxmann's and common shrews, the proportion of shared traps, P (Eq. 1) was positively related to population size, i.e., to the number of animals found on the observed lines as residents. The regression line was steeper for S. caecutiens (B = 0.0281 ± 0.0073) compared to S. araneus (B = 0.0134 ± 0.0027) (Fig. 2 ). The average proportion of shared traps, P , at population density below 10 resident individuals per line made 0.20 ± 0.03 for S. caecutiens and was significantly ( p < 0.013) higher than in S. araneus : 0.08 ± 0.03. This proportion, with population size above 10 resident individuals per line of 50 traps, increased to 0.45 ± 0.05 in S. caecutiens and it was significantly higher ( p < 0.04) than the proportion in S. araneus , the latter was P = 0.27 ± 0.06. Comparing with the proportion expected for random co-location in the Laxmann’s shrew showed that 12 of 14 (85.7%) actual P values exceeded P e level, which is sufficient to consider the difference as non-random in the sign test (z = 2.4, p < 0.016). In the common shrew the actual P values were distributed almost perfectly evenly with respect to the expected values, i.e., 8 above and 9 below the P e line (see Fig. 2 ). According to sign test it is ideal correspondence to the value expected random coincidence. Resident S. araneus has 4.4 ± 0.3 captures, and as shown above, the animal visited on average 1.7 trap numbers, so on average there were 2.3 captures per trap; and for S. caecutiens − 6.3 ± 1.7 captures per 2.1 trap numbers, or 3 captures per trap. One could assume that neighbours will more frequently visited common traps in the case of seeking for contacts, or individual traps, in case of avoiding of contacts, thus we expected to find some disproportion between the piece of common traps and proportion of the common activity. In fact, we found almost complete correspondence between the proportion of visits to common traps and the proportion of common traps in S. araneus ( r = 0.93, p < 0.0001), similarly in S. caecutiens ( r = 0.88, p < 0.0001). Behaviour The Laxmann’s shrew in preference test The average distance that was covered in a trial was 27.4 ± 3.2 m, and the longest travelled distance made 43.1 m. The distance travelled in the zones of containers was 3.4 ± 0.5 m in FZ versus 6.3 ± 0.8 m in AZ, so the distance travelled near container with conspecific was significantly longer ( p 0.28, t = 1.1, df = 19). Distance covered in the border zone in S. caecutiens 9.34 ± 1.9 m was not significantly ( p < 0.09, t = 1.9, df = 13) different from distance covered in central area 6.72 ± 0.9 m, whereas in S. araneus the track in BZ was significantly ( p < 0.001, t = 3.9, df = 19) longer (BZ it was 15.1 ± 1.5 m, and CZ = 10.4 ± 1.1 m). Velocities were not normally distributed. In S . caecutiens, median value of the average velocity in arena was 10.7 cm/s, and maximal speed was recorded in central area, it made about 27 cm/s. These values are quite similar to common shrews: median = 11.8, and max = 26 cm/s. A specific trait in the Laxmann’s shrew was very slow movements of near the container with conspecific compared with movements near empty container. These velocities were normally distributed (Shapiro-Wilk W = 0.958, p > 0.6 and W = 0.93, p > 0.38), and made 5.7 ± 0.6 cm/s and 10.4 ± 1.0 cm/s respectively. Thus, S. caecutiens moved near container with conspecific significantly slower ( p < 0.0001, t =-5.5, df = 13). Apparently, this was due to communication with a partner through the mesh wall of the container, as evidenced, in particular, by the dense tracks around the container (see Fig. 1 ). Another important trait in Laxmann’s shrew behaviour is significantly greater time spent in AZ compared with any other zone. In the common shrew the longest staying was observed in the border zone – BZ (Fig. 3 ). Indexes of preference indicated sustainable interest of S. caecutiens to a stimulating animal in container. Median value of indexes (Eq. 2) of distance I d , and time I T made 0.18 and 0.27 respectively, and both indexes were significantly ( p < 0.003) greater than zero. Comparison of the Laxmann’s and the common shrew Significant differences between these shrews were found for eight of the 22 characteristics obtained in preference test. Distances. S. araneus covered significantly greater distances in both the border zone (BZ) and the central zone (CZ), while the distance travelled in the conspecific’s container zone (AZ) was greater for S. caecutiens . The distances passed on the top of containers, both with stimulating animal (AC) and empty (FC), as well as near empty container (FZ) were larger, but insignificantly, in S . caecutiens . As a result of this disproportion, the total distance travelled during the test was not significantly different. Because distance is a valuable indicator of general activity, we additionally compared distances unrelated to social stimuli, i.e., movement outside the container with the conspecific animal - NC distance. This distance was significantly larger in S. araneus (Table 1 , distance). Table 1 Results of the comparison of the common shrew, S. araneus (SA), and the Laxmann’s shrew, S. caecutiens (SC), in the Kruskal-Wallis ANOVA, and bias of median values from the median value in generalised sample (G). Characteristics with p < 0.05 are highlighted in bold, with p < 0.1 are highlighted in bold italics; d X – weighted deviation of median value (formulae 4). Variable Zone Median value d X U Z p -level SA, N = 20 SC, N = 14 G, N = 34 SA SC Distance, cm BZ 1479.7 759.1 1274.1 0.161 -0.404 71.0 -2.414 0.016 CZ 1070.1 748.4 947.2 0.130 -0.210 71.0 -2.414 0.016 FC 75.4 91.8 84.9 -0.112 0.081 97.0 1.505 0.132 FZ 269.0 281.5 275.2 -0.022 0.023 125.0 0.525 0.600 AC 76.7 82.7 82.7 -0.072 0.000 121.0 -0.665 0.506 AZ 411.1 548.4 515.0 -0.202 0.065 87.0 1.855 0.064 NC 3094.2 2185.3 2583.7 0.022 -0.070 80.0 -2.100 0.0358 Total travelled 3488.3 3172.6 3411.8 0.198 -0.154 98.0 -1.470 0.142 Velocity, cm/s BZ 14.0 13.3 13.5 0.037 -0.013 116.0 -0.840 0.401 CZ 18.1 19.5 19.1 -0.054 0.020 112.0 0.980 0.327 FC 6.7 9.3 7.5 -0.107 0.251 54.0 3.009 0.003 FZ 11.4 10.6 11.0 0.034 -0.040 125.0 -0.525 0.600 AC 5.8 7.6 6.4 -0.097 0.185 117.0 0.805 0.421 AZ 7.2 5.6 6.7 0.079 -0.166 87.0 -1.855 0.064 Average 11.8 10.7 11.4 0.034 -0.062 99.0 -1.435 0.150 Duration of stay, s BZ 111.5 62.6 94.2 0.183 -0.336 93.0 -1.645 0.100 CZ 61.0 29.8 51.8 0.179 -0.425 52.0 -3.079 0.002 FC 9.6 9.8 9.8 -0.024 0.000 128.0 0.420 0.675 FZ 26.4 25.4 25.5 0.033 -0.006 119.5 0.717 0.473 AC 10.5 8.6 9.4 0.120 -0.077 111.5 -0.997 0.319 AZ 57.2 121.6 82.3 -0.305 0.477 71.0 2.414 0.016 Preference I d 0.04 0.18 0.09 -0.557 0.930 96.0 1.348 0.178 I T 0.16 0.27 0.19 -0.195 0.383 102.0 1.129 0.259 Velocities. Substantial difference in velocities was found only on the cover of empty container (FC), where S. caecutiens was significantly swifter, and zones of container with stimulating animal (AZ), where the Laxmann’s shrew was slower. We could note, that with the exception of container covers, and central zone S. caecutiens moved a bit slower. As a result, average velocity was smaller in the Laxmann’s shrew, however due to high fluctuations of the variable, the difference was not significant (Table 1 , velocity). Duration. Compared with S. araneus , S. caecutiens spent almost twice as much time in the container zone with a conspecific (AZ) and almost half as much time in the central (CZ) and in the border (BZ) zones. No difference between these species in time spent on the tops of containers (FC and AC), and in the zone of empty container (FZ) was found (Table 1 , duration). Preference. Although S. caecutiens demonstrated greater interest in the stimulating animal compared to S. araneus , we did not find significant difference in indexes of preference both in distance and in time. Please note, the indices only show the choice in favour of "social" versus “non-social” object, and in this study both species preferred to remain and move within the zone of conspecifics (AZ). We could only mention, that if the indexes in S. caecutiens were significantly above zero, the distance index, I d , in S. araneus was borderline significant ( p < 0.07) compared to zero, but index of time, I T , in this species was significantly ( p < 0.001) greater than null. Indeed, the median value of both indexes in S. caecutiens were substantially greater than those in S. araneus (Table 1 , preference), however due to very high fluctuations of the indexes in the common shrew the differences between the species were not significant. In the common shrew I d ranged from − 0.41 to 0.39, I T from − 0.37 to 0.34, and CV in I d was 250% and CV in I T comprised 113%. In S. caecutiens ; in the indexes were more homogeneous: I d ranged from 0.03 to 0.33 and in I T from 0.05 to 0.41; CV were 71% and 61% respectively. Discussion Space use The move toward sociality involves an increase in the strength and complexity of interactions between animals in a group (Cantor et al., 2021 ). Undoubtedly, the most reliable data for such studies are direct observations of animals, but in the case of secretive species, it is necessary to use indirect data, which in the case of small mammals are the results of CMR studies. CMR data adequately reflect social networks (Wanelik, Farine, 2022). Given that a more shared territory reflects a closer social bond, we decided compare the two studied shrew species. A pitfall in such studies is the differences in estimates of space use, which depend on the observation method. S. araneus is commonly known as territorial solitary species (rev. Churchfield, 1990 ; Ivanter et al., 1994 ; Ivanter, Makarov, 2001 ; rev. Rychlik, 1998 ). Only females of the second year of life could be considered as territorial, but home ranges of these females widely overlap ranges of year-old shrews, and old males widely overlap each other, old female’s and year old shrews ranges (Churchfield, 1990 ). The home ranges of yearling shrews also overlapped, at least peripherally (Sillito, 1963; Croin Michielsen, 1966 ). Using our capture protocol, which allows the animal to roam freely for more than 90% of the day, we found that young shrews' home ranges can overlap substantially (Shchipanov et al., 2001 , 2005 ). More detailed study reveal, that even old females could share area, and in general sharing area in the common shrew depends on local population density, and corresponded to probability of random co-location (Shchipanov, 2021 ). Space using in the Laxmann’s shrew is almost unstudied, and we do not now publications on this issue, except of our own. We found that S. caecutiens shape dense populations with widely overlapping home ranges in mossy coniferous forests (boreal type) and sparse populations with non-overlapping or even non-contiguous home ranges in disturbed grassland habitats (Shchipanov et al., 2005 ). Therefore, comparing space using we shall take into account population density. Our study showed that both species were able to survive in sparse groups in isolated home ranges, but that when population density increased, common areas emerged. Compared with common shrew, at the same population density, year-old Laxmann's shrews shared a significantly higher proportion of home ranges with their conspecifics. The proportion of shared activity, measured by the proportion of captures in shared traps, was consistent with the proportion of these traps, suggesting that the likelihood of social contact was also significantly higher in S. caecutiens . More importantly, in contrast to the common shrew, the proportion of shared territory in the Laxmann's shrew was significantly greater than expected from random co-location. As a result, the space use of the Laxmann's shrew indirectly indicates a greater propensity for social contacts in this species compared to the common shrew. “Sociability”. Under the lack of ability to direct observations one could estimate social behaviour of shrews in trials with animals taken from wild for a short time. Previously, the behaviour of shrews has been studied in tests involving direct contacts of shrews, i.e. in contact tests (rev. Rychlik 1998 ). These tests provided a good idea of the relationship between the different forms of interaction, but did not permit assess the motivation of an individual to engage in social contact because parameters of a test should be scored for the pair as a unit (File and Hyde, 1978 ; File and Seth, 2003 ). Crawley’s ( 2000 , 2004 ) preference test allow estimated motivation to social contacts (Moy et al., 2004 ; Millan, Bales, 2013 ). The test was suggested as a measure of sociability, that is, “motivation for social contact” and implies estimation time spend in the zone of social (container with animal stimulant) versus non-social object (Moy et al., 2004 ). We tailored the test to study shrews taken from a monitored site and that should be returned back as quick as possible (Shchipanov, Demidova, 2020 ). We supposed that sociability could be estimated by index of preferences either in time or in distance travelled in the zone adjoined (0.5 average body length) container with conspecific. Although we used the term “sociability”, we mean that since in shrews most contacts represent agonistic behaviour, the index only shows the propensity to engage social contacts. That's why we used the term sociability in quotation marks here. The indexes were constructed in such a way that a negative value indicated avoidance of a partner, and a positive value indicated a tendency to contact with an individual of the same species. In the common shrew, which has beeen studied for several years, we observed significant fluctuations of preference indexes (Shchipanov, Demidova, 2022 ). In two of four years, the index of distance in the common shrew was significantly below zero, and index of time was not different with zero. In two other years both indexes were significantly above zero. The inter-annual changes in sociability, as it was assessed by indexes, were closely associated with non-residents activity and the exploration behaviour. We hypothesize that inter-annual differences in the sociability of common shrews are a by-product of selection favouring either thorough or superficial exploration (Shchipanov, Demidova, 2022 ). It is not surprising than, that when compared the common shrew and the Laxmann’s shrew in preference test we did not find clear differences in the indexes, which in this study were significantly above zero in both species. Although the Laxmann's shrew had significantly higher distance and time indexes, due to the high variability of these indexes in the common shrew, we could not find significant differences. However, we should note, that we obtained only first data in the Laxmann’s shrew, and we need obtain multiannual data to assay the indexes carefully. When we studied common shrews with this test we found, that time, velocity and duration of stay in various zones of arena could become more salient characteristics than the indexes (Shchipanov, Demidova, 2022 ). In this study, we noted sharply different distributions of activity across arena zones in the compared shrews. Whereas the common shrew spent most of time in the border zone, the Laxmann’s shrew spent most of time in the zone of a conspecific. Therefore, we could assume, that the major motivation in the common shrew was escape from the arena, while in the Laxmann’s shrew to investigate a partner, i.e. to engage in social contact. Therefore, results of study in preference tests allowed us regard the Laxmann’s shrew as more “sociable”, compared with the common shrew, species. In terms of POLS. A suite of closely related behavioral traits shapes behavioral syndrome (Sih et al., 2004 ; Réale and Dingemanse, 2010 ; Biro and Stamps, 2008 ; Carter et al., 2013 ). It was assumed, that these traits are related to differences in suite of physiological (e.g. metabolic, hormonal, immunity) traits that have coevolved with the life-history traits of each species, as the pace-of-life syndrome – POLS (rev. Réale et al., 2010 ). In particular, POLS concept suggested that in behaviour of so called “slow living species” the lower aggressiveness, shyness, thorough exploration, lower activity and higher sociability will correlate, versus correlated boldness, superficial exploration, higher activity and lower sociability in the “fast living species” (Sih and Bell, 2008 ; Réale et al., 2010 ; Varma et al., 2020 ). The variables we analyse in this study have previously been used to study the common shrew in terms of POLS. The bolder individuals of the common shrew were found more “sociable”, and more superficial in exploration as compared to shier shrews (Shchipanov, Demidova, 2022 ). It was of interest to us compared the common and the Laxmann’s shrews in terms of POLS. We assay shift of the variables in the species from some neutral position, the median value of a variable in generalized sample (see Table 1 ). Considering the high fluctuations of almost all the variables studied, since in the trials we used animals just taken from wild, we include in the study all variables differing with probability value p < 0.15. We traditionally interpreted distance and time in central area as measure of boldness, distance travelled without contact with “social object” and average velocity as characteristic of general activity. Of all the characteristics of exploration of non-social novel object (empty container), we can use only activity on the top, since the distance in the zone differed insignificantly. However, it is quite indicative characteristic, since it is evidence in favour of more thorough study. Both velocity and distance in FC were taken with opposite sign, since the less are the value of the variables the more thorough was exploration. Activity in the zone of stimulating conspecific, the distance, time of stay and velocity served to characterise propensity to social contact. The latter variable was taken with opposite sign, since the more attentive was an animal to conspecifics, the slower should it moved. As a result of this study, we obtained a scheme that corresponds well to the continuum proposed for behaviour in Réale et al. ( 2010 p. 4053). Interestingly, all significant and near to significant deviations in S. caecutions were directed towards the position of “slow”, and in S. araneus , on the contrary towards “fast living” species (Fig. 4 ). Conclusion At low population density, yearlings of both species lived in individual areas, but as their numbers increased, both species began to share territory. The proportion of shared area in both species increased with population density, however, it was always significantly greater in the Laxmann's shrew, and if in the common shrew this proportion corresponded to expectation for random co-location, in the Laxmann’s shrew this proportion was significantly larger than expected for random co-location of individuals. Larger shared area was accompanied in Laxman's shrews by significantly greater attention to conspecifics in behavioral tests of animals from wild. Our current data showed, that compared with the common shrew, with respect to POLS continuum the behaviour, of the Laxmann's shrew corresponded "slow species", more shy, less active in general, more thorough explorer, with greater propensity to engage in social contacts. All this allow us to conclude that the Laxmann's shrew made at least a step to more complex social structure. Declarations Ethics All applicable international, national and/ or institutional guidelines for the care and use of animals were followed. All procedures performed in experiments involving animals were approved by the Bioethical Committee on Animal and Human Research at A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences (which are in accordance with recommendations of US and EU ethical committees) (permission No. 30 issued on February 27, 2019, with extension No. 30a issued April 10, 2023), following all relevant laws and regulations. This article does not contain any experiments on human subjects performed by any of the co-authors. The capture method used does not affect animal welfare. No animal died in traps in the course of the study. All authors have read, understood, and have complied as applicable with the statement on ‘Ethical responsibilities of Authors’ as found in the Instructions for Authors. Authors contribution Conceptualisation, data analysis and writing original draft: NS; field experiments, catching animals and conducting behavioral tests: NS, AK, TD; writing and editing NS, TD. All authors have read and agreed to publish the final version of the manuscript. Funding This work was supported by A.N. Severtsov Institute of Ecology and Evolution state assignment contract No. 0089‐2021‐0007. Conflict of interest. The authors have no conflict of interest to declare. Data availability . Not applicable. References Biro P, Stamps J (2008) Are animal personality traits linked to life-history productivity? Trends Ecol Evol 23:361–368. https://doi:10.1016/j. tree.2008.04.003 Bourke AF (2011) Principles of social evolution. Oxford University Press Cantor M, Maldonado-Chaparro AA, Beck KB et al (2021) The importance of individual-to-society feedbacks in animal ecology and evolution. 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Biol Lett 11:20150825. http://doi.org/10.1098/rsbl.2015.0825 Varma V, Vasoya H, Jain A, Binoy VV (2020) The bold are the sociable: personality traits and laterality in an indigenous megafish, the deccan mahseer (Tor khudree). Ichthyol Res 67:483–492. https://doi.org/10.1007/s10228-020-00744-8 Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6810348","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":475620147,"identity":"6da4b540-1e2b-4469-adab-e46034dd27f5","order_by":0,"name":"Nikolay A. Shchipanov","email":"","orcid":"","institution":"A.N.Severtsov Institute of Ecology and Evolution","correspondingAuthor":false,"prefix":"","firstName":"Nikolay","middleName":"A.","lastName":"Shchipanov","suffix":""},{"id":475620148,"identity":"c59b2e68-030e-4074-8a65-4947c626c7ef","order_by":1,"name":"Alexey A. Kalinin","email":"","orcid":"","institution":"A.N.Severtsov Institute of Ecology and Evolution","correspondingAuthor":false,"prefix":"","firstName":"Alexey","middleName":"A.","lastName":"Kalinin","suffix":""},{"id":475620149,"identity":"4398fe9a-2a71-405c-86e7-e72ea209a857","order_by":2,"name":"Tatiana Demidova","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA60lEQVRIiWNgGAWjYDCCA0DEA+dVMDCwsROrBaLtDFALMxFaGOBaGNuABCEtfMe7Ew+8qTgsb89+9pjEz3nb5PmYGdg+8+DRInnm7IaDc84cNuzhyUuT7N1227CNmYF5Nj4tBjdyNxzmbUtj7GHIMZPg3XabEaSFGa+W+2/BWux7+N+YSf6dc9uesJYbvCAtNok9Ejlm0rwNtxMJapE8kwvyi01yz403xtYyx24ntzEzNjPOwaOF7/jZzR/eVEjYtvfnGN58U3Pbdn5782GGN3i0IAMWCQjN2ECkBmAcfiBa6SgYBaNgFIwoAADnUU2PJJMRLAAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0001-7864-1893","institution":"A.N. Severtsov Institute of Ecology and Evolution","correspondingAuthor":true,"prefix":"","firstName":"Tatiana","middleName":"","lastName":"Demidova","suffix":""}],"badges":[],"createdAt":"2025-06-03 10:42:54","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6810348/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6810348/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":85478376,"identity":"234a8d89-0824-490a-a60c-5026091fbe0e","added_by":"auto","created_at":"2025-06-26 10:27:27","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":755355,"visible":true,"origin":"","legend":"\u003cp\u003eVirtual zones and position of the containers with stimulating animal (black) and empty (white). On the left chart positions of the containers with an animal and empty (free), and zones: BZ – border zone, CZ – central zone, AZ – vicinity of container with stimulating conspecific, AC – cover of container with stimulating shrew, FZ – vicinity of empty container, FC – cover of empty container. On the right chart we show typical digitized track for 5 min trial in Laxmann’s shrew is shown. The bold solid line borders the floor, the track (red line) behind the floor reflects the jumps onto the walls of the chamber. Various dashed lines show margins of various zones. Please note the close dense ring of tracks around the container with stimulating animal compared with the loose trace near empty container\u003c/p\u003e","description":"","filename":"image1.png","url":"https://assets-eu.researchsquare.com/files/rs-6810348/v1/61bc9e3d5b492321255a4a03.png"},{"id":85478373,"identity":"68218351-83ea-4a9a-9bd3-8177a6a64959","added_by":"auto","created_at":"2025-06-26 10:27:27","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":85418,"visible":true,"origin":"","legend":"\u003cp\u003eProportion of traps, \u003cem\u003eP\u003c/em\u003e (eq. 1) visited by two and more residents in \u003cem\u003eS. araneus\u003c/em\u003e (blue) and in \u003cem\u003eS. caecutiens\u003c/em\u003e (red) with respect to number, \u003cem\u003eN\u003c/em\u003e, of resident conspecific shrews on a line of 50 live-traps in 2014 - 2024. Markers show actual values of proportion under certain \u003cem\u003eN\u003c/em\u003e in one of the years, squares are for the Laxmann’s shrew, and rings are for the common shrew. The single solid line is the regression and the double line is the expected proportion \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e (eq. 2) for the Laxmann’s shrew; the dashed single line is the regression, and the double dashed line is the expected proportion, \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e, for the common shrew\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-6810348/v1/1d98de9fa774bb926553aec4.png"},{"id":85478371,"identity":"b17df0dc-c482-40ae-b5e7-99fe9cfc7379","added_by":"auto","created_at":"2025-06-26 10:27:27","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":76150,"visible":true,"origin":"","legend":"\u003cp\u003eResults of ANOVA for duration of stay in various zones of the arena in \u003cem\u003eS. caecutiens\u003c/em\u003e (red lines and marker) and \u003cem\u003eS. araneus\u003c/em\u003e(blue line and marker). \u003cem\u003ep\u003c/em\u003e-values obtained in Tukey post-hock test are given in the table under the chart, for \u003cem\u003eS. caecutiens\u003c/em\u003e above, and for \u003cem\u003eS. araneus\u003c/em\u003e - below the diagonal; \u003cem\u003ep\u003c/em\u003e\u0026lt;0.05 highlighted bold, in red for \u003cem\u003eS. caecutiens\u003c/em\u003e, and in blue for \u003cem\u003eS. araneus\u003c/em\u003e. BZ – border zone, CZ – central zone, FC – cover of the empty container, FZ – zone around empty container, AC – cover of the container with conspecific, AZ – zone around container with conspecific.\u003c/p\u003e","description":"","filename":"image3.png","url":"https://assets-eu.researchsquare.com/files/rs-6810348/v1/65b2597003772d2c91e70d86.png"},{"id":85478780,"identity":"652f402f-2533-4cdc-9796-76f2732a7170","added_by":"auto","created_at":"2025-06-26 10:35:27","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":87834,"visible":true,"origin":"","legend":"\u003cp\u003eWeighted deviation of median value in \u003cem\u003eS. araneus\u003c/em\u003e and \u003cem\u003eS. caecutiens\u003c/em\u003e from median value in generalised sample, \u003cem\u003ed\u003c/em\u003e\u003csub\u003eX\u003c/sub\u003e (eq. 4) of the studied variables (detailed in text). Left and right opposite columns of behavioral characteristics correspond Réal’s with coauthors schematic of the potential integration of different traits along a pace-of-life continuum (Réal et al., 2010 p. 4053). Positive value of \u003cem\u003ed\u003c/em\u003e\u003csub\u003eX\u003c/sub\u003e indicated shift towards increasing of a variable, and negative one show its diminishing, compared with median value in general sample (see Table 1). Signs of deviations in table 1 that marked movement towards the named group of behaviours are shown near the name of each block, and scale of deviation (proportion to median in generalized sample) is shown above the chart. Red colour is for Laxmann’s shrews; blue colour is for common shrews. Full colour bars display characteristics differed for \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05, pale colored for \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.1, and uncolored for \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.15 (see Table 1). D is for distance, T is for time, V is for velocities. BZ is for border zone, CZ is for central area, AZ is for zine around container with conspecific animal, NC is the area of arena unrelated to zones of social stimuli, and FC is for cover of empty container.\u003c/p\u003e","description":"","filename":"image4.png","url":"https://assets-eu.researchsquare.com/files/rs-6810348/v1/447c10fe99e38f41f7c23617.png"},{"id":89492657,"identity":"85f6fd03-6836-433c-9c5b-e4a34fc1d9ae","added_by":"auto","created_at":"2025-08-20 14:12:51","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1772935,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6810348/v1/2317ad78-a5d1-4ef0-a6d9-8ee530887502.pdf"}],"financialInterests":"","formattedTitle":"\u003cp\u003eThe Laxmann’s shrew \u003cem\u003eSorex caecutiens\u003c/em\u003e. Step towards \"sociality\"\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe studying of social evolution suggests understanding how \u0026ldquo;previously selfish, free-living individuals join together to form a group resembling an individual in its own right\u0026rdquo;, passing thee stages \u0026ndash; social group formation, social group maintenance, and social group transformation (Bourke \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). There are a number of theoretical models regarded partner choice, direct and indirect reciprocity, etc. (eg. Novak 2006, 2012; Fuentes \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Chassy et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2025\u003c/span\u003e, and other), and in all scenarios, evolution begins when individuals of the same species can remain in contact outside the mating season. In any case the first step to sociality is tolerance of members of one's own species, and hence the presence of a common space and some tendency towards social contact. This suggests that such a move might manifest itself in as significant extend of shared area and a greater propensity for social contact.\u003c/p\u003e \u003cp\u003eAmong placental mammals, shrews are of particular interest for such a step. These insectivorous mammals can be considered as the life form of the first placental mammals (Luo et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Since the majority of shrews considered solitary living (Churchfield \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1990\u003c/span\u003e, Rychlik \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), it is reasonable to assume that that the ancestor of placentals was likely solitary, even though \u003cem\u003eEulipotyphla\u003c/em\u003e are not more basal than other mammalian orders (Lukas, Clutton-Brock, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). It is of interest to study more shrew species in wild with respect to the \u0026ldquo;biodiversity of social systems to be able to perform meaningful comparative studies about the evolution of sociality\u0026rdquo; (Valomy et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). On the other hand, it is the third most diverse order of mammals after rodents and bats, with more than 370 species family shrews, but we have very few knowledge of the social organization of most species. The common belief that all shrews are equally asocial animals may be the result of insufficient study of their behaviour and space use (Valomy et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). At least, some shrews of the subfamily \u003cem\u003eCrocidurinae\u003c/em\u003e are more tolerant, and often shared area, whereas subfamily \u003cem\u003eSoricinae\u003c/em\u003e is mainly represented by solitary, aggressive species (Rychlik \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). More detailed studies show differences in communication and space use between closely related species. For example, Eurasian water shrew, \u003cem\u003eNeomys fodiens\u003c/em\u003e, is known as highly aggressive strictly solitary, territorial animal, while Iberian water shrew, \u003cem\u003eN. anomalus\u003c/em\u003e exhibited communal using of space and amicable behaviours (Krushinska, Rychlik \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Krushinska et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Rychlik, Zwolak, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOf all the shrews, red-toothed shrews, \u003cem\u003eSorex\u003c/em\u003e, were regarded as the most aggressive and least social animals (rev. Churchfield \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; rev. Rychlik \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). It should be noted, however, that our understanding of social behaviour in this rather diverse genus, which contains at least 142 known species, is based chiefly on the study of the common shrew (Crowcroft \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1955\u003c/span\u003e, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1957\u003c/span\u003e). Due to technical difficulties in handling shrews, the extreme susceptibility of animals to traps and captivity, both the space use, and behaviour of most species remain poorly understood. However, interactions in paired contact tests with animals from wild indicate that at least four \u003cem\u003eSorex\u003c/em\u003e species found in central Russia differ in their aggressiveness towards conspecifics (Shchipanov et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Of those species the Laxmann\u0026rsquo;s shrew \u003cem\u003eS. caecutiens\u003c/em\u003e was found the most \u0026ldquo;social\u0026rdquo;: the species exhibited rather diverse patterns of integrative interactions, which accounted for more than 33% of all contacts. The amicable interactions were rather diverse and included such patterns as climbing on the partner, crawling under the partner, sitting side by side in tactile contact, which were absent in common shrews (Shchipanov et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). It was of interest to obtain more quantitative characteristics of behaviour to compare \u003cem\u003eS. araneus\u003c/em\u003e and \u003cem\u003eS. caecutiens\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe Laxmann\u0026rsquo;s shrew is a numerous species with a wide range, inhabiting from the north-eastern Europe to Far-East, however its behaviour and space use are barely studied. In Central Russia, the preferred habitat of \u003cem\u003eS. caecutiens\u003c/em\u003e is mossy coniferous forests, where Laxmann\u0026rsquo;s shrews of their first year of life established dense groups with shared home ranges (Shchipanov et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). We monitor small mammals using capture-mark-recapture (CMR) technique in various habitats, and in particular in mossy coniferous forest, which permit us to study space use in this species in a dense local population. Behaviour of this species, which was earlier studied in contact test gives only general picture of interactions of individuals (Shchipanov et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Now we for the first-time assay sociability of the Laxmann\u0026rsquo;s shrew from wild in Crowley\u0026rsquo;s (2000, 2004) preference test. Previously, when studying sociability in this test in the common shrew, we assessed personality traits such as activity, boldness, exploratory activity, and propensity for social contacts (Shchipanov, Demidova \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The test allows us to quantify behavioural traits, and is applicable for comparative studies of behaviour for both individuals and species. Social behaviours that benefit survival can be associated with life histories in terms of the pace-of-life syndrome (hereafter POLS) (Silk, Hodgson \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Note that the above mentioned behavioral traits are important for both interspecific and intraspecific differentiation in terms of POLS schematic behavioral profile (R\u0026eacute;ale et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this study, we intend 1) quantitatively compare shared area in the common and the Laxmann's shrews, 2) report the results of the first study of \u003cem\u003eS. caecutiens\u003c/em\u003e in a preference test, and 3) compare the results of preference tests in \u003cem\u003eS. caecutiens\u003c/em\u003e and \u003cem\u003eS. araneus\u003c/em\u003e in terms of POLS.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area and animal trapping\u003c/h2\u003e \u003cp\u003eWe monitor small mammals in five habitats in the central part of European Russia in the vicinity of the Bakanovo village, the Staritsa Region of Tver Oblast\u0026rsquo;, N 56\u0026deg;18\u0026prime;; E 34\u0026deg;53\u0026prime; (Shchipanov, Kalinin \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e). The Laxmann's shrew is more numerous in mixed coniferous (spruce and pine) mossy old-growth forest, and is also common, but not numerous, in the adjacent forest area that was cleared in 1998 and is now overgrown with young birch forest. In other habitats surveyed, in a young forest on an abandoned field, on a former pasture and on the territory of an abandoned village, \u003cem\u003eS. caecutiens\u003c/em\u003e was sparse, with only single individuals observed in some years. The common shrew contrary was rare in the mossy forest and more numerous in grassland habitats (Shchipanov, Kalinin 2024 a). The traps in all the habitats except of village area was installed as lines of 50 traps with 7.5 metres between traps, and as grid in the village area because of the configuration of plot available to monitoring. On the grid the traps were placed as four lines of 18 traps and one line of 15 traps with 7.5 m interval between traps in the lines, and 10 m distance between the lines.\u003c/p\u003e \u003cp\u003eWe used special trapping protocol, which was tailored to shrews (Shchipanov et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). This imply using original live-trap, with wire treadle (the chart was given in above-mentioned articles), oat flakes with unrefined sunflower oil as bait, and daily exposition of operative traps shortened to 1.5 h after daily activation. While checking traps animals inspected, marked, if necessary, weighed, their position was recorded, and they were released at the place of capture. Animals were marked by toe-clipping. Unfortunately, this remained the only reliable way of mass individual lifelong marking of shrews. Note, we minimise harm to individual using numbers which require minimal number of toes to be clipped. This way of marking did not influenced survival of shrews (Shchipanov et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Shchipanov et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This regimen of inspection minimizes mortality in traps. No animal died in a trap in this study. After inspection, the traps remained at their position to be available for visits but were not operational. Thus, animals could move over 90% of daytime for free. This allows us to consider each capture of an individual as independent and reflective of its actual presence on the spot.\u003c/p\u003e \u003cp\u003eFor the behaviour study, we used common shrews from the village area and Laxmann\u0026rsquo;s shrews from the forest. Animals for behavioral testing were removed from traps during regular trap inspections at 8:00\u0026ndash;9:30 AM, placed in cages with abundant food (mealworms, grasshoppers, fly larvae, and chicken hearts), subjected to the test, and released at the catch sites no later than 11 AM in case of \u003cem\u003eS. araneus\u003c/em\u003e, and 12\u0026ndash;13 in case of \u003cem\u003eS. caecutiens\u003c/em\u003e. The greater time in the latter shrew resulted from a more distant sampling area. After the tests animals were released at a place of capture.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eAnimals\u003c/h3\u003e\n\u003cp\u003eAll the data in this study are given for the first-year shrews, since a complete population turnover in shrews takes one year, and first-year animals make up the vast majority of the population by mid-summer (Churchfield, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Kalinin et al., 2008). Space sharing in both the common and the Laxmann's shrews was estimated using data collected in coniferous mossy forest and adjoined young birch forest at the cleared area, in 2014\u0026ndash;2024. A total of 223 \u003cem\u003eS. caecutiens\u003c/em\u003e individuals were captured 614 times, and 438 \u003cem\u003eS. araneus\u003c/em\u003e individuals were captured 984 times. Of those 124 \u003cem\u003eS. caecutiens\u003c/em\u003e and 252 \u003cem\u003eS. araneus\u003c/em\u003e were captured more than twice and were regarded to be resident.\u003c/p\u003e \u003cp\u003eBehaviour of Laxmann\u0026rsquo;s shrew was studied in 2024, in July. Fourteen Laxmann\u0026rsquo;s shrews from the forest plot were used in trials. In common shrews we used data for the first trial of 20 animals from the village area, since we conducted ongoing annual surveillance of the common shrew behaviour in this habitat. The trials with common shrews, which were used for comparison, were conducted during the same time period as the Laxmann\u0026rsquo;s shrew, in July 2024. The numbers, which was estimated as number of residents per 100 traps (Shchipanov, Kalinin 2024 b), in the period, when animals were taken for tests, made 22 ind./100 traps (11 on line of 50 traps) in the Lxmann\u0026rsquo;s shrew in the forest, and 23 ind./100 traps (on average 4.14 on line of 18 traps on grid) in the common shrew.\u003c/p\u003e\n\u003ch3\u003eBehavioral tests\u003c/h3\u003e\n\u003cp\u003eTo study behaviour in common shrew, we used modified Crawley\u0026rsquo;s preference test. It implies 5 min trials in a single 50 \u0026times; 50 cm chamber with specified virtual zones. The test has been described in details in our previous publications (Shchipanov, Demidova \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Briefly, two wire-mesh containers (cylindrical, 10 cm high and 8 cm in diameter) with a non-transparent cover were placed at a fixed position on the diagonal of the arena. One of the containers was with a stimulating animal, and the other remained empty. The focal animal could have olfactory, acoustic and visual communication with the stimulator through the mesh wall of the container. The study was conducted in an isolated room with artificial lighting. The brightness of the light was approximately equal to natural light outdoors. The focal animal was placed in the centre of arena in plastic glass, and released manually. The operator left the room after the focal animal was released, and the trial was conducted in the operator's absence. The orientation of the arena position towards the cardinal points was altered by us randomly between trials. The stimulating animal, unfamiliar to a focal individual, was placed in a container immediately prior to a trial. Because we were limited by the number of animals available for testing, we used both focal and stimulating animals from the same habitat. We considered animals with distance between the activity centres above median value of the distance between border captures in the entire sample (in case of both species it made approximately 60 m) as \"unfamiliar\". In this project we used for analysis only first (for both focal and stimulating animal) trials.\u003c/p\u003e \u003cp\u003eTrials were video recorded by means of a Sony Handycam in MPEG-2 format, and the videos were analysed in the EthoVision XT (Noldus) software. Distance travelled, time spent in an area and mean velocity were determined for the border zone (BZ) (a 3-cm band along the walls), the zone of possible contact with the stimulating animal (around the container within a strip 3 cm from the edge) (AZ), the free zone similarly sized near the empty container (FZ), the tops (covers) of the containers with an animal (AC), and the free container (FC), and the area of the arena that excluded the above-mentioned zones, the central zone (CZ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eAnalyses\u003c/h3\u003e\n\u003cp\u003eNormal distribution was studied in Shapiro-Wilk test for \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.15. Variability of normally distributed data were shown as \u0026plusmn;\u0026thinsp;SE unless otherwise stated.\u003c/p\u003e\n\u003ch3\u003eShared space\u003c/h3\u003e\n\u003cp\u003eShared space was estimated as the proportion (\u003cem\u003eP\u003c/em\u003e) of traps in an individual's home range that were visited by other neighbouring resident shrews. It was calculated as proportion of traps (\u003cem\u003eP\u003c/em\u003e) visited by two and more resident shrews (animals with two and more than captures) \u0026ndash; shared traps (\u003cem\u003eT\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e), to the total number of traps (\u003cem\u003eT\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e) visited by residents:\u003c/p\u003e \u003cp\u003e \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u003cem\u003eT\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e / \u003cem\u003eT\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e. (1)\u003c/p\u003e \u003cp\u003eIt has been found in common shrews, that shared area in yearling conspecifics increased with the number of shrews lived at the observed area due to random overlap (Shchipanov, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Therefore, to avoid bias related to the different population abundance, we compared proportions of shared traps plotted it against number of resident shrews (animals with two and more captures) revealed in a habitat (on a line of 50 traps, 375 m). The study was performed as analysis of regressions.\u003c/p\u003e \u003cp\u003eTo understand if the proportions \u003cem\u003eP\u003c/em\u003e resulted from the random coincidence of visited spots, we tested the null hypothesis about random distribution of visits in traps. To test the hypothesis, we simulated the expected proportion of shared traps, \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e. The simulation assumes that each trap has an equal probability of receiving one of the 50 trap numbers in a trial. The total number of trials (\u003cem\u003et\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e) equaled the sum of traps (trap numbers), visited by all residents on a line of 50 traps in a given period. It could be calculated as number of residents, \u003cem\u003eN\u003c/em\u003e, multiplied by mean number of traps, \u003cem\u003em\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e, visited by resident individual: \u003cem\u003et\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e = \u003cem\u003em\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e\u003cem\u003eN\u003c/em\u003e. After receiving \u003cem\u003en\u003c/em\u003e random traps numbered from 1 to 50 in \u003cem\u003en\u003c/em\u003e trials, (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u003cem\u003et\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e), we counted the total number of trap-numbers obtained, as well as the number of traps-numbers that appeared two or more times. Calculations were performed using EXCEL's built-in randomization. After 1000 iterations we got average number of visited traps, \u003cem\u003en\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e, and average number of traps visited by two and more individuals \u003cem\u003en\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e, and calculated \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e as\u003c/p\u003e \u003cp\u003e \u003cem\u003eP\u003c/em\u003e \u003csub\u003ee\u003c/sub\u003e = \u003cem\u003en\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e / \u003cem\u003en\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e (2)\u003c/p\u003e \u003cp\u003eThe expected proportion was calculated for number of animals, \u003cem\u003eN\u003c/em\u003e, ranged from 5 to 31 (min and max \u003cem\u003eN\u003c/em\u003e revealed on trapping lines) with taking into account average number of traps visited by mean individual per trapping session. In \u003cem\u003eS. araneus\u003c/em\u003e the average number of traps (trap-numbers), \u003cem\u003em\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e, visited during a trapping session made 429 trap-numbers (sum for all trapping sessions) divided by 252 residents (revealed for all trapping sessions), or 1.70 traps visited by mean individual per a trapping session. In \u003cem\u003eS. caecutiens m\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e made 254/124\u0026thinsp;=\u0026thinsp;2.05 traps visited by mean individual per a trapping session. Note that the different number of traps on a home range of residents, \u003cem\u003em\u003c/em\u003e\u003csub\u003et\u003c/sub\u003e, influenced the number of trials in simulation, thus expected \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e in \u003cem\u003eS. araneus\u003c/em\u003e and \u003cem\u003eS. caecutiens\u003c/em\u003e are not equal for equal \u003cem\u003eN\u003c/em\u003e. Actual and expected proportions were compared in sign test.\u003c/p\u003e \u003cp\u003eTo assess whether trap sharing differed significantly under similar population size, we compared the proportion of traps shared between the medium (5\u0026ndash;10 individuals per line) and high (over 10 individuals) abundance classes. The data sets (proportions of shared traps) in the cases were found normally distributed in Shapiro-Wilk test. Therefore, we studied differences in proportion of shared traps with application \u003cem\u003et\u003c/em\u003e-test for independent variables (groups). Since the proportions of shared traps and captures in shared traps were normally distributed, we assessed their goodness of fit using Pearson correlation, assuming that if there was no significant disproportion, we would find a significant correlation.\u003c/p\u003e \u003cp\u003eThese analyses were performed in STATISTICA 8 (StatSoft, USA).\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eBehaviours\u003c/h2\u003e \u003cp\u003eWe studied the full set of variables obtained during the trials: distances, velocity and duration of stay in the specified above virtual zones. To assay whether it was some preference of contact to social \u003cem\u003eversus\u003c/em\u003e non-social object we calculated indexes of preference in distance \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e and in time \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e as:\u003c/p\u003e \u003cp\u003e \u003cem\u003eI\u003c/em\u003e \u003csub\u003eX\u003c/sub\u003e =X\u003csub\u003eAZ\u003c/sub\u003e / (X\u003csub\u003eAZ\u003c/sub\u003e + X\u003csub\u003eFZ\u003c/sub\u003e) \u0026ndash; 0.5, (3)\u003c/p\u003e \u003cp\u003ewhere X is variable in question, either distance \u0026ndash; d, or time \u0026ndash; T, in the zone of container with conspecific (AZ) or in the zone of empty container (FZ). The index ranges from \u0026minus;\u0026thinsp;0.5 to 0.5. It tends to zero when the presence of the stimulus animal in the container has no significance for the focal individual; it is positive when the focal individual shows an interest in the partner, and negative when it avoids interact with a partner. Comparing indexes with null was performed in sign test.\u003c/p\u003e \u003cp\u003eNormally distributed distances travelled within zones were compared between zones of the same species using Student's t-tests for dependent variables, as overall activity differed from trial to trial. Duration of stay in various zones of the arena was compared in one-way ANOVA, with \u003cem\u003ep\u003c/em\u003e-value obtained in Tukey post-hock test. Residuals were found normally distributed.\u003c/p\u003e \u003cp\u003eTo understand whether there were significant differences in behavioral variables between species, we compared the suit of studied characteristics using Kruskal-Wallis ANOVA since a number of variables were not normally distributed. All statistical tests were performed in Statistica 8 (StatSoft, USA).\u003c/p\u003e \u003cp\u003eTo reveal tendency in bias in behavioral patterns in the studied species we used weighted deviation from general median value (\u003cem\u003eM\u003c/em\u003e\u003csub\u003eG\u003c/sub\u003e) of a variable calculated as:\u003c/p\u003e \u003cp\u003e \u003cem\u003ed\u003c/em\u003e \u003csub\u003eXS\u003c/sub\u003e = (\u003cem\u003eM\u003c/em\u003e\u003csub\u003eXS\u003c/sub\u003e \u0026ndash; \u003cem\u003eM\u003c/em\u003e\u003csub\u003eXG\u003c/sub\u003e) / \u003cem\u003eM\u003c/em\u003e\u003csub\u003eXG\u003c/sub\u003e, (4)\u003c/p\u003e \u003cp\u003ewhere \u003cem\u003ed\u003c/em\u003e\u003csub\u003eXS\u003c/sub\u003e is deviation of variable X in species in question (either \u003cem\u003eS. araneus\u003c/em\u003e or \u003cem\u003eS. caecutiens\u003c/em\u003e), \u003cem\u003eM\u003c/em\u003e\u003csub\u003eXS\u003c/sub\u003e \u0026ndash; median value of variable X in species (S) in question, and \u003cem\u003eM\u003c/em\u003e\u003csub\u003eXG\u003c/sub\u003e is median value of variable X in generalized (G) sample of both species. Positive deviation indicates a predominance in the compared variable, and a negative value indicates a loss in the variable compared with the median value in generalized sample.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eShared space\u003c/h2\u003e \u003cp\u003eIn both Laxmann's and common shrews, the proportion of shared traps, \u003cem\u003eP\u003c/em\u003e (Eq.\u0026nbsp;1) was positively related to population size, i.e., to the number of animals found on the observed lines as residents. The regression line was steeper for \u003cem\u003eS. caecutiens\u003c/em\u003e (B\u0026thinsp;=\u0026thinsp;0.0281\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0073) compared to \u003cem\u003eS. araneus\u003c/em\u003e (B\u0026thinsp;=\u0026thinsp;0.0134\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0027) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe average proportion of shared traps, \u003cem\u003eP\u003c/em\u003e, at population density below 10 resident individuals per line made 0.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 for \u003cem\u003eS. caecutiens\u003c/em\u003e and was significantly (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.013) higher than in \u003cem\u003eS. araneus\u003c/em\u003e: 0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03. This proportion, with population size above 10 resident individuals per line of 50 traps, increased to 0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 in \u003cem\u003eS. caecutiens\u003c/em\u003e and it was significantly higher (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.04) than the proportion in \u003cem\u003eS. araneus\u003c/em\u003e, the latter was \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06.\u003c/p\u003e \u003cp\u003eComparing with the proportion expected for random co-location in the Laxmann\u0026rsquo;s shrew showed that 12 of 14 (85.7%) actual \u003cem\u003eP\u003c/em\u003e values exceeded \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e level, which is sufficient to consider the difference as non-random in the sign test (z\u0026thinsp;=\u0026thinsp;2.4, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.016). In the common shrew the actual \u003cem\u003eP\u003c/em\u003e values were distributed almost perfectly evenly with respect to the expected values, i.e., 8 above and 9 below the \u003cem\u003eP\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e line (see Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). According to sign test it is ideal correspondence to the value expected random coincidence.\u003c/p\u003e \u003cp\u003eResident \u003cem\u003eS. araneus\u003c/em\u003e has 4.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 captures, and as shown above, the animal visited on average 1.7 trap numbers, so on average there were 2.3 captures per trap; and for \u003cem\u003eS. caecutiens\u003c/em\u003e \u0026minus;\u0026thinsp;6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7 captures per 2.1 trap numbers, or 3 captures per trap. One could assume that neighbours will more frequently visited common traps in the case of seeking for contacts, or individual traps, in case of avoiding of contacts, thus we expected to find some disproportion between the piece of common traps and proportion of the common activity. In fact, we found almost complete correspondence between the proportion of visits to common traps and the proportion of common traps in \u003cem\u003eS. araneus\u003c/em\u003e (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.93, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001), similarly in \u003cem\u003eS. caecutiens\u003c/em\u003e (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.88, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eBehaviour\u003c/h2\u003e \u003cdiv id=\"Sec12\" class=\"Section3\"\u003e \u003ch2\u003eThe Laxmann\u0026rsquo;s shrew in preference test\u003c/h2\u003e \u003cp\u003eThe average distance that was covered in a trial was 27.4\u0026thinsp;\u0026plusmn;\u0026thinsp;3.2 m, and the longest travelled distance made 43.1 m. The distance travelled in the zones of containers was 3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 m in FZ \u003cem\u003eversus\u003c/em\u003e 6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8 m in AZ, so the distance travelled near container with conspecific was significantly longer (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.003, \u003cem\u003et\u003c/em\u003e=-3.6, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;13). In the common shrew sampled in the same period of time these distances (in AZ\u0026thinsp;=\u0026thinsp;3.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 m, and in FZ\u0026thinsp;=\u0026thinsp;4.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 m) were not different (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.28, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.1, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;19). Distance covered in the border zone in \u003cem\u003eS. caecutiens\u003c/em\u003e 9.34\u0026thinsp;\u0026plusmn;\u0026thinsp;1.9 m was not significantly (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.09, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.9, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;13) different from distance covered in central area 6.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 m, whereas in \u003cem\u003eS. araneus\u003c/em\u003e the track in BZ was significantly (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3.9, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;19) longer (BZ it was 15.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5 m, and CZ\u0026thinsp;=\u0026thinsp;10.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1 m).\u003c/p\u003e \u003cp\u003eVelocities were not normally distributed. In \u003cem\u003eS\u003c/em\u003e. caecutiens, median value of the average velocity in arena was 10.7 cm/s, and maximal speed was recorded in central area, it made about 27 cm/s. These values are quite similar to common shrews: median\u0026thinsp;=\u0026thinsp;11.8, and max\u0026thinsp;=\u0026thinsp;26 cm/s. A specific trait in the Laxmann\u0026rsquo;s shrew was very slow movements of near the container with conspecific compared with movements near empty container. These velocities were normally distributed (Shapiro-Wilk W\u0026thinsp;=\u0026thinsp;0.958, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.6 and W\u0026thinsp;=\u0026thinsp;0.93, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.38), and made 5.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6 cm/s and 10.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0 cm/s respectively. Thus, \u003cem\u003eS. caecutiens\u003c/em\u003e moved near container with conspecific significantly slower (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001, \u003cem\u003et\u003c/em\u003e=-5.5, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;13). Apparently, this was due to communication with a partner through the mesh wall of the container, as evidenced, in particular, by the dense tracks around the container (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAnother important trait in Laxmann\u0026rsquo;s shrew behaviour is significantly greater time spent in AZ compared with any other zone. In the common shrew the longest staying was observed in the border zone \u0026ndash; BZ (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIndexes of preference indicated sustainable interest of \u003cem\u003eS. caecutiens\u003c/em\u003e to a stimulating animal in container. Median value of indexes (Eq.\u0026nbsp;2) of distance \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e, and time \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e made 0.18 and 0.27 respectively, and both indexes were significantly (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.003) greater than zero.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eComparison of the Laxmann\u0026rsquo;s and the common shrew\u003c/h2\u003e \u003cp\u003eSignificant differences between these shrews were found for eight of the 22 characteristics obtained in preference test.\u003c/p\u003e \u003cp\u003eDistances. \u003cem\u003eS. araneus\u003c/em\u003e covered significantly greater distances in both the border zone (BZ) and the central zone (CZ), while the distance travelled in the conspecific\u0026rsquo;s container zone (AZ) was greater for \u003cem\u003eS. caecutiens\u003c/em\u003e. The distances passed on the top of containers, both with stimulating animal (AC) and empty (FC), as well as near empty container (FZ) were larger, but insignificantly, in \u003cem\u003eS\u003c/em\u003e. \u003cem\u003ecaecutiens\u003c/em\u003e. As a result of this disproportion, the total distance travelled during the test was not significantly different. Because distance is a valuable indicator of general activity, we additionally compared distances unrelated to social stimuli, i.e., movement outside the container with the conspecific animal - NC distance. This distance was significantly larger in \u003cem\u003eS. araneus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, distance).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eResults of the comparison of the common shrew, \u003cem\u003eS. araneus\u003c/em\u003e (SA), and the Laxmann\u0026rsquo;s shrew, \u003cem\u003eS. caecutiens\u003c/em\u003e (SC), in the Kruskal-Wallis ANOVA, and bias of median values from the median value in generalised sample (G). Characteristics with \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 are highlighted in bold, with \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.1 are highlighted in bold italics; \u003cem\u003ed\u003c/em\u003e\u003csub\u003eX\u003c/sub\u003e \u0026ndash; weighted deviation of median value (formulae 4).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eVariable\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eZone\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e \u003cp\u003eMedian value\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e\u003cem\u003ed\u003c/em\u003e\u003csub\u003eX\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eU\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eZ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003ep\u003c/em\u003e-level\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSA,\u003c/p\u003e \u003cp\u003eN\u0026thinsp;=\u0026thinsp;20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSC,\u003c/p\u003e \u003cp\u003eN\u0026thinsp;=\u0026thinsp;14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eG,\u003c/p\u003e \u003cp\u003eN\u0026thinsp;=\u0026thinsp;34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSC\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"7\" rowspan=\"8\"\u003e \u003cp\u003eDistance, cm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1479.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e759.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e1274.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.161\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.404\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e71.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-2.414\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.016\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eCZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1070.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e748.4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e947.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.130\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.210\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e71.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-2.414\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.016\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e75.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e91.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e84.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.112\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.081\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e97.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.505\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.132\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFZ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e269.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e281.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e275.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.022\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.023\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e125.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.525\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.600\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e76.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e82.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e82.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.072\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e121.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.665\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.506\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eAZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e411.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e548.4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e515.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e-0.202\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e0.065\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e87.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e1.855\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.064\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eNC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e3094.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e2185.3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e2583.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.022\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.070\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e80.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-2.100\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.0358\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal travelled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3488.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3172.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3411.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.198\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.154\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e98.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-1.470\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.142\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"6\" rowspan=\"7\"\u003e \u003cp\u003eVelocity, cm/s\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBZ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e13.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.037\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.013\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e116.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.840\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.401\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCZ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e19.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e19.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.054\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.020\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e112.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.980\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.327\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eFC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e6.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e9.3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e7.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e-0.107\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e0.251\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e 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align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.421\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eAZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e7.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e5.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e6.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.079\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.166\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e87.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-1.855\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.064\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAverage\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.034\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.062\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e99.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-1.435\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.150\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003eDuration of stay, s\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e111.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e62.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e94.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.183\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.336\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e93.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-1.645\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.100\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eCZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e61.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e29.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e51.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.179\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-0.425\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e52.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e-3.079\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.002\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.024\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e128.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.420\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.675\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFZ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e26.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e25.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.033\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.006\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e119.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.717\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.473\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.120\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.077\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e111.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.997\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.319\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eAZ\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e57.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e121.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e82.3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e-0.305\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e0.477\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e71.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e2.414\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.016\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003ePreference\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.557\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.930\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e96.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.348\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.178\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.195\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.383\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e102.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.129\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.259\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eVelocities. Substantial difference in velocities was found only on the cover of empty container (FC), where \u003cem\u003eS. caecutiens\u003c/em\u003e was significantly swifter, and zones of container with stimulating animal (AZ), where the Laxmann\u0026rsquo;s shrew was slower. We could note, that with the exception of container covers, and central zone \u003cem\u003eS. caecutiens\u003c/em\u003e moved a bit slower. As a result, average velocity was smaller in the Laxmann\u0026rsquo;s shrew, however due to high fluctuations of the variable, the difference was not significant (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, velocity).\u003c/p\u003e \u003cp\u003eDuration. Compared with \u003cem\u003eS. araneus\u003c/em\u003e, \u003cem\u003eS. caecutiens\u003c/em\u003e spent almost twice as much time in the container zone with a conspecific (AZ) and almost half as much time in the central (CZ) and in the border (BZ) zones. No difference between these species in time spent on the tops of containers (FC and AC), and in the zone of empty container (FZ) was found (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, duration).\u003c/p\u003e \u003cp\u003ePreference. Although \u003cem\u003eS. caecutiens\u003c/em\u003e demonstrated greater interest in the stimulating animal compared to \u003cem\u003eS. araneus\u003c/em\u003e, we did not find significant difference in indexes of preference both in distance and in time. Please note, the indices only show the choice in favour of \"social\" \u003cem\u003eversus\u003c/em\u003e \u0026ldquo;non-social\u0026rdquo; object, and in this study both species preferred to remain and move within the zone of conspecifics (AZ). We could only mention, that if the indexes in \u003cem\u003eS. caecutiens\u003c/em\u003e were significantly above zero, the distance index, \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e, in \u003cem\u003eS. araneus\u003c/em\u003e was borderline significant (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.07) compared to zero, but index of time, \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e, in this species was significantly (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) greater than null. Indeed, the median value of both indexes in \u003cem\u003eS. caecutiens\u003c/em\u003e were substantially greater than those in \u003cem\u003eS. araneus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, preference), however due to very high fluctuations of the indexes in the common shrew the differences between the species were not significant. In the common shrew \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e ranged from \u0026minus;\u0026thinsp;0.41 to 0.39, \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e from \u0026minus;\u0026thinsp;0.37 to 0.34, and CV in \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e was 250% and CV in \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e comprised 113%. In \u003cem\u003eS. caecutiens\u003c/em\u003e; in the indexes were more homogeneous: \u003cem\u003eI\u003c/em\u003e\u003csub\u003ed\u003c/sub\u003e ranged from 0.03 to 0.33 and in \u003cem\u003eI\u003c/em\u003e\u003csub\u003eT\u003c/sub\u003e from 0.05 to 0.41; CV were 71% and 61% respectively.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eSpace use\u003c/h2\u003e \u003cp\u003eThe move toward sociality involves an increase in the strength and complexity of interactions between animals in a group (Cantor et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Undoubtedly, the most reliable data for such studies are direct observations of animals, but in the case of secretive species, it is necessary to use indirect data, which in the case of small mammals are the results of CMR studies. CMR data adequately reflect social networks (Wanelik, Farine, 2022). Given that a more shared territory reflects a closer social bond, we decided compare the two studied shrew species. A pitfall in such studies is the differences in estimates of space use, which depend on the observation method.\u003c/p\u003e \u003cp\u003e \u003cem\u003eS. araneus\u003c/em\u003e is commonly known as territorial solitary species (rev. Churchfield, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Ivanter et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Ivanter, Makarov, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; rev. Rychlik, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Only females of the second year of life could be considered as territorial, but home ranges of these females widely overlap ranges of year-old shrews, and old males widely overlap each other, old female\u0026rsquo;s and year old shrews ranges (Churchfield, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). The home ranges of yearling shrews also overlapped, at least peripherally (Sillito, 1963; Croin Michielsen, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1966\u003c/span\u003e). Using our capture protocol, which allows the animal to roam freely for more than 90% of the day, we found that young shrews' home ranges can overlap substantially (Shchipanov et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). More detailed study reveal, that even old females could share area, and in general sharing area in the common shrew depends on local population density, and corresponded to probability of random co-location (Shchipanov, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Space using in the Laxmann\u0026rsquo;s shrew is almost unstudied, and we do not now publications on this issue, except of our own. We found that \u003cem\u003eS. caecutiens\u003c/em\u003e shape dense populations with widely overlapping home ranges in mossy coniferous forests (boreal type) and sparse populations with non-overlapping or even non-contiguous home ranges in disturbed grassland habitats (Shchipanov et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Therefore, comparing space using we shall take into account population density.\u003c/p\u003e \u003cp\u003eOur study showed that both species were able to survive in sparse groups in isolated home ranges, but that when population density increased, common areas emerged. Compared with common shrew, at the same population density, year-old Laxmann's shrews shared a significantly higher proportion of home ranges with their conspecifics. The proportion of shared activity, measured by the proportion of captures in shared traps, was consistent with the proportion of these traps, suggesting that the likelihood of social contact was also significantly higher in \u003cem\u003eS. caecutiens\u003c/em\u003e. More importantly, in contrast to the common shrew, the proportion of shared territory in the Laxmann's shrew was significantly greater than expected from random co-location. As a result, the space use of the Laxmann's shrew indirectly indicates a greater propensity for social contacts in this species compared to the common shrew.\u003c/p\u003e \u003cp\u003e \u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026ldquo;Sociability\u0026rdquo;.\u003c/span\u003e \u003c/p\u003e \u003cp\u003eUnder the lack of ability to direct observations one could estimate social behaviour of shrews in trials with animals taken from wild for a short time. Previously, the behaviour of shrews has been studied in tests involving direct contacts of shrews, i.e. in contact tests (rev. Rychlik \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). These tests provided a good idea of the relationship between the different forms of interaction, but did not permit assess the motivation of an individual to engage in social contact because parameters of a test should be scored for the pair as a unit (File and Hyde, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1978\u003c/span\u003e; File and Seth, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Crawley\u0026rsquo;s (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) preference test allow estimated motivation to social contacts (Moy et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Millan, Bales, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The test was suggested as a measure of sociability, that is, \u0026ldquo;motivation for social contact\u0026rdquo; and implies estimation time spend in the zone of social (container with animal stimulant) \u003cem\u003eversus\u003c/em\u003e non-social object (Moy et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). We tailored the test to study shrews taken from a monitored site and that should be returned back as quick as possible (Shchipanov, Demidova, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). We supposed that sociability could be estimated by index of preferences either in time or in distance travelled in the zone adjoined (0.5 average body length) container with conspecific. Although we used the term \u0026ldquo;sociability\u0026rdquo;, we mean that since in shrews most contacts represent agonistic behaviour, the index only shows the propensity to engage social contacts. That's why we used the term sociability in quotation marks here.\u003c/p\u003e \u003cp\u003eThe indexes were constructed in such a way that a negative value indicated avoidance of a partner, and a positive value indicated a tendency to contact with an individual of the same species. In the common shrew, which has beeen studied for several years, we observed significant fluctuations of preference indexes (Shchipanov, Demidova, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In two of four years, the index of distance in the common shrew was significantly below zero, and index of time was not different with zero. In two other years both indexes were significantly above zero. The inter-annual changes in sociability, as it was assessed by indexes, were closely associated with non-residents activity and the exploration behaviour. We hypothesize that inter-annual differences in the sociability of common shrews are a by-product of selection favouring either thorough or superficial exploration (Shchipanov, Demidova, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). It is not surprising than, that when compared the common shrew and the Laxmann\u0026rsquo;s shrew in preference test we did not find clear differences in the indexes, which in this study were significantly above zero in both species. Although the Laxmann's shrew had significantly higher distance and time indexes, due to the high variability of these indexes in the common shrew, we could not find significant differences. However, we should note, that we obtained only first data in the Laxmann\u0026rsquo;s shrew, and we need obtain multiannual data to assay the indexes carefully.\u003c/p\u003e \u003cp\u003eWhen we studied common shrews with this test we found, that time, velocity and duration of stay in various zones of arena could become more salient characteristics than the indexes (Shchipanov, Demidova, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In this study, we noted sharply different distributions of activity across arena zones in the compared shrews. Whereas the common shrew spent most of time in the border zone, the Laxmann\u0026rsquo;s shrew spent most of time in the zone of a conspecific. Therefore, we could assume, that the major motivation in the common shrew was escape from the arena, while in the Laxmann\u0026rsquo;s shrew to investigate a partner, i.e. to engage in social contact.\u003c/p\u003e \u003cp\u003eTherefore, results of study in preference tests allowed us regard the Laxmann\u0026rsquo;s shrew as more \u0026ldquo;sociable\u0026rdquo;, compared with the common shrew, species.\u003c/p\u003e \u003cp\u003e \u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eIn terms of POLS.\u003c/span\u003e \u003c/p\u003e \u003cp\u003eA suite of closely related behavioral traits shapes behavioral syndrome (Sih et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; R\u0026eacute;ale and Dingemanse, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Biro and Stamps, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Carter et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). It was assumed, that these traits are related to differences in suite of physiological (e.g. metabolic, hormonal, immunity) traits that have coevolved with the life-history traits of each species, as the pace-of-life syndrome \u0026ndash; POLS (rev. R\u0026eacute;ale et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In particular, POLS concept suggested that in behaviour of so called \u0026ldquo;slow living species\u0026rdquo; the lower aggressiveness, shyness, thorough exploration, lower activity and higher sociability will correlate, \u003cem\u003eversus\u003c/em\u003e correlated boldness, superficial exploration, higher activity and lower sociability in the \u0026ldquo;fast living species\u0026rdquo; (Sih and Bell, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; R\u0026eacute;ale et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Varma et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The variables we analyse in this study have previously been used to study the common shrew in terms of POLS. The bolder individuals of the common shrew were found more \u0026ldquo;sociable\u0026rdquo;, and more superficial in exploration as compared to shier shrews (Shchipanov, Demidova, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). It was of interest to us compared the common and the Laxmann\u0026rsquo;s shrews in terms of POLS. We assay shift of the variables in the species from some neutral position, the median value of a variable in generalized sample (see Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Considering the high fluctuations of almost all the variables studied, since in the trials we used animals just taken from wild, we include in the study all variables differing with probability value \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.15. We traditionally interpreted distance and time in central area as measure of boldness, distance travelled without contact with \u0026ldquo;social object\u0026rdquo; and average velocity as characteristic of general activity. Of all the characteristics of exploration of non-social novel object (empty container), we can use only activity on the top, since the distance in the zone differed insignificantly. However, it is quite indicative characteristic, since it is evidence in favour of more thorough study. Both velocity and distance in FC were taken with opposite sign, since the less are the value of the variables the more thorough was exploration. Activity in the zone of stimulating conspecific, the distance, time of stay and velocity served to characterise propensity to social contact. The latter variable was taken with opposite sign, since the more attentive was an animal to conspecifics, the slower should it moved.\u003c/p\u003e \u003cp\u003eAs a result of this study, we obtained a scheme that corresponds well to the continuum proposed for behaviour in R\u0026eacute;ale et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e p. 4053). Interestingly, all significant and near to significant deviations in \u003cem\u003eS. caecutions\u003c/em\u003e were directed towards the position of \u0026ldquo;slow\u0026rdquo;, and in \u003cem\u003eS. araneus\u003c/em\u003e, on the contrary towards \u0026ldquo;fast living\u0026rdquo; species (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eAt low population density, yearlings of both species lived in individual areas, but as their numbers increased, both species began to share territory. The proportion of shared area in both species increased with population density, however, it was always significantly greater in the Laxmann's shrew, and if in the common shrew this proportion corresponded to expectation for random co-location, in the Laxmann\u0026rsquo;s shrew this proportion was significantly larger than expected for random co-location of individuals. Larger shared area was accompanied in Laxman's shrews by significantly greater attention to conspecifics in behavioral tests of animals from wild. Our current data showed, that compared with the common shrew, with respect to POLS continuum the behaviour, of the Laxmann's shrew corresponded \"slow species\", more shy, less active in general, more thorough explorer, with greater propensity to engage in social contacts. All this allow us to conclude that the Laxmann's shrew made at least a step to more complex social structure.\u003c/p\u003e "},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll applicable international, national and/ or institutional guidelines for the care and use of animals were followed. All procedures performed in experiments involving animals were approved by the Bioethical Committee on Animal and Human Research at A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences (which are in accordance with recommendations of US and EU ethical committees) (permission No. 30 issued on February 27, 2019, with extension No. 30a issued April 10, 2023), following all relevant laws and regulations. This article does not contain any experiments on human subjects performed by any of the co-authors. The capture method used does not affect animal welfare. No animal died in traps in the course of the study. All authors have read, understood, and have complied as applicable with the statement on \u0026lsquo;Ethical responsibilities of Authors\u0026rsquo; as found in the Instructions for Authors.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors contribution\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConceptualisation, data analysis and writing original draft: NS; field experiments, catching animals and conducting behavioral tests: NS, AK, TD; writing and editing NS, TD. All authors have read and agreed to publish the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by A.N. Severtsov Institute of Ecology and Evolution state assignment contract No. 0089‐2021‐0007.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest.\u003c/strong\u003e The authors have no conflict of interest to declare.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e. Not applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eBiro P, Stamps J (2008) Are animal personality traits linked to life-history productivity? Trends Ecol Evol 23:361\u0026ndash;368. https://doi:10.1016/j. tree.2008.04.003\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBourke AF (2011) Principles of social evolution. Oxford University Press\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCantor M, Maldonado-Chaparro AA, Beck KB et al (2021) The importance of individual-to-society feedbacks in animal ecology and evolution. 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Ichthyol Res 67:483\u0026ndash;492. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10228-020-00744-8\u003c/span\u003e\u003cspan address=\"10.1007/s10228-020-00744-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"shrews, home range, preference test, social contacts, sociality","lastPublishedDoi":"10.21203/rs.3.rs-6810348/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6810348/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSorex are commonly considered highly aggressive territorial animals. Our research shows that species in this genus can significantly differ in the extent of home ranges overlap and their inclination to contact with conspecifics. We assay space usage and propensity to engage in social contacts in the Laxmann\u0026rsquo;s shrew, \u003cem\u003eS. caecutiens\u003c/em\u003e in comparison with the common shrew, \u003cem\u003eSorex araneus\u003c/em\u003e. Although both studied species shared area under increased population density, the Laxmann\u0026rsquo;s shrew reveal significantly larger shared area, and the proportion of shared area in this species significantly exceeded that was expected for random co-location of animals. In the common shrew shared area corresponded to expectation for random co-location of animals. In a preference test with a conspecific in wire-mesh container we found, that whereas the common shrew was rather indifferent, the Laxmann\u0026rsquo;s shrew reveal propensity to contact with conspecific. When estimated a suit of variables obtained in trials with respect to pace-of-life continuum, we found that Laxmann's shrew corresponded \"slow species\"; it was more shy, less active in general, more thorough explorer, with greater propensity to engage in social contacts. We assume that the Laxmann's shrew made at least a step towards more complex social structure.\u003c/p\u003e","manuscriptTitle":"The Laxmann’s shrew Sorex caecutiens. Step towards \"sociality\"","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-26 10:27:23","doi":"10.21203/rs.3.rs-6810348/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"cfa62509-a7d1-4a52-9266-c638d6e12107","owner":[],"postedDate":"June 26th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-08-20T14:04:43+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-26 10:27:23","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6810348","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6810348","identity":"rs-6810348","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}