INTRODUCTION
Dispersal is a key process connecting wildlife populations across landscapes. By enabling individuals to leave their natal or established ranges, disperse to new areas, and reproduce elsewhere, it maintains demographic connectivity and gene flow among spatially separated populations (Clobert et al., 2012;Baguette et al., 2013). Yet dispersal also requires animals to negotiate unfamiliar and potentially hazardous landscape features. Depending on the interaction between landscape structure and species-specific movement behavior, such features may restrict or redirect animal movement, thereby shaping functional connectivity (Baguette and Van Dyck, 2007). At different spatial and temporal scales, these movement restrictions can limit access to resources, restrict demographic exchange, or impede gene flow (Cosgrove et al., 2018). Identifying the features that redirect movement is therefore important for understanding functional connectivity and for designing effective wildlife-management and conservation strategies.
Rivers are prominent landscape features that may restrict the dispersal of terrestrial mammals and gene flow between populations on opposite banks (Patton et al., 1994;Eriksson et al., 2004). Their effects, however, are not uniform. Wider rivers have been associated with reduced dispersal or stronger genetic differentiation in some mammals such as European badgers (Meles meles) and European ground squirrels (Spermophilus citellus), whereas smaller rivers showed no such effect (Frantz et al., 2010;Ćosić et al., 2013). Similarly, large Amazonian rivers can delimit the ranges of larger-bodied primates, whereas smaller upstream reaches may be permeable (Wallace, 1852;Ayres and Clutton-Brock, 1992). Barrier effects may also vary between taxa, sexes and even individuals, as illustrated by contrasting responses among Bornean small mammals, sex-biased dispersal in brown bears and occasional crossings by individual wildcats between genetically structured populations (Hartmann et al., 2013;Brunke et al., 2019;Spitzer et al., 2026). Whether a river acts as a barrier therefore depends on both species and individual traits as well as the circumstances of crossing.
River resistance can also vary among sites and seasons. Water depth and flow, for example, may influence river crossings, while islands and bridges can provide local crossing opportunities (Bond et al., 2012). Seasonal ice may have the same effect: caribou (Rangifer tarandus) avoided open water and selected ice when moving across large water bodies (Leblond et al., 2016). Yet most evidence for river effects comes from population genetic patterns or from general movement responses, and the routes and conditions of individual cross-river movements during dispersal remain poorly documented.
Raccoon dogs (Nyctereutes procyonoides) are capable of long-distance movements and rapid spread into newly available areas (Diao et al., 2022;Deshpande et al., 2025). GPS-based studies of invasive populations in northern Europe found no overall evidence that water bodies constrained raccoon dog dispersal, and even suggested crossings of the large Tornionjoki River between Finland and Sweden (Deshpande et al., 2025). In coastal Finland, ice cover further facilitated movements between islands that might otherwise be inaccessible by land (Toivonen et al., 2025). In Korea, population genetic evidence likewise suggests that major rivers, including the Han River, may not completely segregate raccoon dog populations (Hong et al., 2018). Nevertheless, these studies do not reveal how individual river-crossing events unfold during dispersal, what crossing modes may be used, or whether such events are associated with altered activity.
Here, we report river-crossing events and infer plausible modes of crossing during long-distance dispersal by GPS-tracked raccoon dogs in South Korea. We characterize individual spatial trajectories, including changes in net squared displacement across pre-dispersal, dispersal, and settlement phases, and compare ODBA-derived activity levels among phases. By linking individual crossing events and inferred crossing modes to movement and activity dynamics, we examine how locally and seasonally available opportunities can make rivers permeable during dispersal. This perspective frames functional connectivity as a dynamic, rather than fixed, property of riverine landscapes and has implications for wildlife management in rapidly changing environments.
MATERIALS AND METHODS
1. Study Site
The GPS tracking data used in this study were derived from a broader raccoon dog movement study conducted across the Seoul Metropolitan Area (Choi, 2026). The focal area in the study comprised the reaches of the Imjin and Han Rivers where river-crossing events were recorded: Yeoncheon County and Paju City on the Imjin River, and the cities of Hanam and Namyangju on the Han River (Figure 1). Land cover ranges from forest and paddy agriculture in the north to high-density urban development in the south (Ministry of Environment, Republic of Korea, 2023). Channel width was measured from the water classes of the same land-cover map along the relevant main-stem reaches, extending approximately 40 ㎞ along the Imjin River and 23 ㎞ along the Han River. Median (maximum) channel widths were 213 (492) m for the Imjin River and 320 (1,856) m for the Han River in the focal study area.
2. Methods
1) Animal capture, collaring, and tracking
Raccoon dogs were captured in medium-sized live traps (Tomahawk Live Traps, USA; Woojin Global, South Korea) baited with minced pork or duck meat. Traps were checked on site or by remote video cameras at least twice within 24 h. Captured individuals were guided into a denim handling bag, measured for body mass, hind foot length, age class and sex. Further details are given in Choi (2026) and Choi et al. (2026). Of the 65 individuals tracked between August 2022 and May 2026, the three individuals in which cross-river movements were found constitute the case series described here. These three were live-trapped and released at the capture site (ID: DKC, PTH) or, after rehabilitation at a wildlife rescue center, translocated within the same city (ID: SNG). Each was fitted with a GPS collar (Druid Tech, China; approximately 145 g, <4% of body mass) that also recorded tri-axial acceleration continuously at 25 Hz, from which overall dynamic body acceleration (ODBA) was computed on board at 1-min intervals and stored as the mean of each 10-min interval (Wilson et al., 2006). ODBA is reported in the collar’s output units of 0.1 mg, so 1,000 units correspond to 0.1 g. Analyses used 1-h GPS fixes and 10-min mean ODBA from the dispersal period and the equally long periods immediately before and after it, excluding the period following capture or release.
All capture activities were conducted under permits issued by the respective local government offices. The animal capture, handling and tagging procedures were approved by the Seoul National University Institutional Animal Care and Use Committee (IACUC; approval no. SNU-230613-3-1, SNU-241101-2).
2) Identification and measurement of cross-river movements
A candidate cross-river movement was recorded whenever two consecutive GPS locations fell on opposite sides of a river channel. Candidates were examined visually on satellite imagery and web maps, and we retained those crossing a channel at least 50 m wide, whether the inferred route passed over open water or followed a road spanning the channel; movements associated with narrower tributaries, tidal creeks or approaches parallel to a channel were excluded. This threshold was set to focus on large rivers and to exclude small streams, seasonally dry channels and false crossings caused by GPS location error.
For each retained movement, we measured the length of a straight line between the two consecutive GPS locations intersecting mapped water. If the line crossed more than one water section separated by a mid-channel island, each section and their total were reported. Channel width at the site was measured as twice the distance from the channel centerline to the nearest non-water cell, that is the diameter of the largest circle fitting inside the channel, and the same measure was applied along the main stem of each river to describe the reaches.
Each site was then examined on Sentinel-2 true-color imagery from the Copernicus Data Space Ecosystem, using the cloud-free scene nearest in date to each crossing event, and on web maps. From these, we identified nearby bridges, islands and other structures that could have provided a crossing opportunity, measured the span of any structure that a movement followed, and recorded the surface condition of the channel.
3) Dispersal phases and activity
For each focal individual, we identified dispersal from the net squared displacement (NSD) time series and daily movement rates (Bunnefeld et al., 2011). Dispersal was defined as sustained directional movement away from the initial range without an immediate return, and ended when a new, spatially stable range was established. NSD was calculated relative to the median location during the pre-dispersal period. Daily mean ODBA was calculated for days with at least 60 ten-minute ODBA records. For each dispersal event, we compared ODBA during dispersal with the immediately preceding and subsequent periods of equal duration. For each crossing, mean 10-min ODBA was calculated between the two GPS locations bracketing it.
Spatial processing and statistical summaries were carried out in Python 3.10 (https://www.python.org), and maps were inspected in QGIS 3.34.2 (https://www.qgis.org). All spatial computation used EPSG:32652 (UTM zone 52N).
RESULTS AND DISCUSSION
1. River crossings during dispersal
Five river-crossing movements were identified in three of the 65 GPS-tracked raccoon dogs, all during dispersal and all between sunset and sunrise, at 19:00–23:00 or 03:00–06:00 KST (Table 1, Figure 2). The three individuals were female and differed in age, study area and timing of movement, but each crossed a major river while dispersing (Table 1).
Dispersal was attempted by 30 individuals (16 females and 14 males) and completed by 22 individuals (12 females and 10 males; Choi, 2026); river crossings were detected in three of these dispersers (10%) and in none of the individuals that remained resident, although the accessibility of major rivers from resident home ranges varied. Large rivers therefore may bound the routine movements that make up a resident home range while remaining permeable during dispersal.
A juvenile female (ID: DKC) dispersed between 19 and 22 October 2024, within the autumn period in which juvenile raccoon dogs usually leave their natal range (Sutor, 2008;Drygala et al., 2010;Figure 1). On the night of 20 October, the Imjin River was crossed northward in two steps intersecting 50 and 278 m of mapped water, and southward again across a 254 m section about 24 h later, by way of direct passage and use of a mid-channel island, with settlement on the south bank (Table 1). Both movements are consistent with passage through open water, the island reducing the longest continuous section.
Two adult females dispersed later than the natal dispersal season, between 15 and 18 February 2025 (ID: PTH) and between 31 January and 5 February 2026 (ID: SNG). On the night of 16 February 2025, PTH moved out onto the Imjin riverbed at 20:00 and was located within the channel at 21:00 and 22:00 without reaching the opposite bank; it then crossed to the north bank between 22:00 and 23:00, over 312 m of mapped water before settled in the north (Table 1;Figure 2). On 4 February 2026, SNG crossed the Han River northward at a disused road bridge about 540 m long and returned before dacwn without using any structure, crossing a reach about 500 m wide on an oblique course that intersected 1,729 m of water (Table 1; Figure 2A and B). Satellite imagery acquired four days earlier showed the reach to be extensively ice covered (Figure 2C). Ice cover is known to open movement routes across water for raccoon dogs in winter (Toivonen et al., 2025) and is the most plausible explanation here.
The river-crossing events were therefore consistent with four potential crossing modes: direct passage across open water, a mid-channel island, a disused road bridge and an ice-covered reach. Together, these cases show why channel width alone is an incomplete proxy for river resistance. Mid-channel islands, bridges and seasonal ice may each alter the routes available to a dispersing individual.
2. Movement and activity across dispersal phases
Net squared displacement rose rapidly during dispersal and leveled off once the animals reached their new home ranges (Figure 3). Dispersal lasted 4, 4 and 6 days in DKC, PTH and SNG, over cumulative paths of 26.4, 24.4 and 27.8 km. Daily mean ODBA during dispersal was 1.2 to 3.4 times that of the equally long period immediately before it, but the juvenile and the two adults reached this elevation in different ways (Figure 3). The juvenile (ID: DKC) was already active before departure (daily mean ODBA: 2,032), rose only modestly during dispersal (2,499) and returned to a similar level afterwards (1,988; Figure 3A). Each adult was far less active in the days before departure and dispersed at 2.9 and 3.4 times that level (909 to 2,603 in PTH and 438 to 1,504 in SNG); activity then declined but stayed well above the pre-dispersal level (2,134 and 987; Figure 3B, 3C). During river crossings, mean 10-min ODBA was 5,683 in DKC, 5,532 in PTH and 4,275 in SNG, 2.1–2.8 times their daily means during dispersal. These cases illustrate that dispersal can be preceded by contrasting activity patterns, a period of low activity before the winter dispersals of the adults, consistent with the reduced winter activity or road crossings reported for raccoon dogs (Kowalczyk and Zalewski, 2011;Choi et al., 2026), and sustained high activity around the autumn dispersal of the juvenile. However, age class and season are confounded in these three cases, and separating these effects will require more individual long-distance dispersals tracked across both seasons and age classes.
The present case series shows that raccoon dogs can move between opposite banks of major rivers during long-distance dispersal, a behavior we did not observe during home-range residency. These occasional crossings of barriers illustrate the high dispersal capacity of raccoon dogs (Deshpande et al., 2025). However, the three individuals do not permit inference about the probability of crossing, the effects of sex or age, or the relative importance of river characteristics. Although all crossing individuals were female, this pattern may reflect the small sample rather than sex-specific behavior. Our observations nevertheless suggest that the permeability of a river varies among locations and seasons according to the presence of islands, crossing structures, and ice cover, and quantifying these effects will require high-resolution tracking across more seasons, river types and life stages. More broadly, they suggest that how much a barrier restricts population connectivity cannot be read from its physical structure alone, but depends on the crossing opportunities available to an animal at a particular place and time.












