The effect of urbanization and behavioral factors on coyote net displacement and its implications for seed dispersal

doi:10.21203/rs.3.rs-2743672/v1

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The effect of urbanization and behavioral factors on coyote net displacement and its implications for seed dispersal

https://doi.org/10.21203/rs.3.rs-2743672/v1

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published 31 Oct, 2023

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Coyotes (Canis latrans) may be important seed dispersal vectors in urban areas, given their omnivorous diet and wide-ranging movement patterns potentially able to bypass fragmentation. However, fragmentation itself, anthropogenic food sources, and human activity can limit their natural movement patterns. Previous research has found urbanization limits movement range across mammals; however, it remains unclear the degree to which this may cascade into seed dispersal as seed retention time plays an important role in seed dispersal distance. Additionally, social and temporal changes in behavior influence coyote movement patterns, likely interacting with the effect of urbanization on net displacement and seed dispersal. We used GPS telemetry data to analyze the effect of urbanization on coyote net displacement (n = 94 individuals), in interaction with social and temporal factors, for a series of seed retention timeframes. We found that urbanization led to shorter net displacement in comparison to rural areas. The effect of urbanization increased with increasing seed retention timeframe and disproportionately affected long-distance seed dispersal. While seasonality did not influence the effect of urbanization, social status had a strong interaction effect, whereby the effect of urbanization was strong for transient and dispersing coyotes but negligible for resident coyotes, as territoriality was likely the main limiting factor for the latter, whereas the former, which were wider ranging, were most affected by the urban matrix. Interestingly, despite the urban effect, coyote net displacement varied across social statuses and seasons in both landscape types, suggesting coyotes can provide a heterogenous seed dispersal contribution within and across plant species. Future research on fine-scale movement patterns and scat analysis is needed to better understand the cascading effects of decreased long-distance net displacements on urban plant populations.

cityscape

net displacement

mesopredators

coyotes

zoochory

carnivore

seed dispersal

cities

Urban areas are characterized by isolated remnants of vegetation patches scattered across a developed matrix composed of varying degrees of built infrastructure and human activity. The presence of animals with large movement capabilities and the ability to bypass the inhospitable developed landscape found in urban areas is key for maintaining seed dispersal between remnant vegetation patches. While birds are effective seed dispersers in urban areas and can travel for long distances using small patches as stepping stones (Han and Keeffe 2019), their gape size and shape limits the range of fruits ingested (Mazer and Wheelwright 1993). In contrast, mammals can ingest larger seed species in greater quantities and retain them for longer periods of time, given their body size and digestive tracts (Willson 1993). Mammals can also disperse seeds that attach to their fur (Hovstad et al. 2009) and those acting as predators can also disperse seeds previously eaten by their prey (Hämäläinen et al. 2017). Hence, mammals are potential seed dispersers for a wider range of plant species within the urban environment, and may be key in determining the composition of plant communities in urban forests, as in other environments (Melo et al. 2010).

However, anthropogenic footprint and built infrastructure have been found to shorten movement ranges of mammals in urban landscapes in comparison to those in rural and forested landscapes. These shorter movement ranges caused by the anthropogenic footprint and built infrastructure have been suggested to have implications for seed dispersal distance (Tucker et al. 2018). However, seed dispersal distance is the combined result of animal movement and seed retention time (Nield et al. 2020). In addition, the movement range of mammals that are socially complex and subject to seasonality, may be further altered by behavior changes associated with social dynamics, breeding and pup-rearing requirements and resource availability (Franckowiak et al. 2019; Larson et al. 2020; Ellington et al. 2020). Therefore, a more in-depth analysis of how urbanization alters movement ranges of mammals considering their behavioral complexity, and the retention time of the seeds they transport, is needed to further understand the ecological implications of urbanization.

In this study we used coyotes (Canis latrans) as a model species, given their wide movement ranges (Gehrt et al. 2009), and their remarkable adaptation to both urban and rural areas. Coyotes have been identified as an effective seed disperser for several plant species, via both primary ingestion of wild and farmed fruits (Rubalcava-Castillo et al. 2020) and secondary ingestion of seeds through prey (Sarasola et al. 2016mäläinen et al. 2017), and through epizoochory, including diaspores of grass and bushes (Quick et al. 2017). Furthermore, coyote movement patterns are strongly influenced by their social dynamics, whereby group-living resident coyotes partition the landscape through territoriality, and solitary transient and dispersing coyotes traverse the landscape between territories (Kamler and Gipson 2000; Mitchell et al. 2015). Temporal dynamics also influence coyote movement patterns, following breeding, pup-rearing, and dispersal activity, and climate-driven changes in resource and shelter availability (Sasmal et al. 2019).

We investigated differences in coyote net displacement over four seed retention timeframes, reflecting their diverse seed dispersal modes. To quantify net displacement, we used telemetry data from GPS-collared coyotes (n = 94 individuals) in urban and rural landscapes in southern Ontario, Canada. We integrated into the analysis the effect of social status (i.e., resident, transient, dispersing), biological season (i.e., breeding, pup-rearing, dispersal), and climate season (i.e., spring, summer, autumn, winter). We hypothesized that 1) reduced habitat availability, increased anthropogenic food availability, and increased encampment behavior (Ellington and Gehrt 2019) would increase movement tortuosity in urban areas, leading to shorter net displacement over time, affecting longer-retained seeds the most. 2) Non-territorial coyotes, given their larger movement ranges and predominantly linear movements (Webster et al. 2022), would be more affected by urbanization, leading to increased differences in net displacement between landscape types for all seed retention timeframes. 3) Seasonal changes resulting in biological constraints on coyote movement (i.e., pup-rearing season) (Kitchen et al. 1999) would decrease differences in net displacement between landscape types, and those biological seasons resulting in greater movement requirements (i.e., dispersal and breeding season) would increase differences in net displacement between landscape types. 4) Harsher climate seasons would lead to increased differences in net displacement between landscape types, given more stable resource and shelter availability in urban areas (Sugden et al. 2021).

Ultimately, this study aims to explore the effect of urbanization on the scale of seed dispersal by coyotes for seed species with different retention time ranges, considering their social and temporal behavioral complexity, and identify intraspecific differences in their seed dispersal contributions in both urban and rural landscapes.

2.1. Study area

Our study area included the Greater Toronto Area (GTA), a metropolitan region, and the surrounding rural areas within southern Ontario (Fig. 1). The largest urban landscape in the study area, the GTA, contains forested river valleys crossing transversally over high-density and medium-density residential areas. These forested river valleys are well-suited habitats for coyotes and connect the lakeshore with the green belt, a large protected natural area surrounding the GTA. Coyotes also utilize other green areas in the GTA, such as parks, cemeteries, golf courses, and backyards, similar to other mesopredators in the area (Rosatte and Allan 2009). The rural landscape in the study area is composed mainly of plantations, agricultural fields, orchards, densely wooded vegetation, and scattered small towns.

To distinguish between urban and rural landscapes, we defined urbanization degree following the Global Human Settlement project (Florczyk et al. 2019), whereby an urban area was characterized by contiguous grid cells containing a minimum of 1,500 people per km² or a minimum of 50% share of built-up land, totaling a minimum of 50,000 inhabitants. Following this definition, all areas outside urban centers were considered rural, which included the following landcover classes: agricultural land, small towns, and low-density human settlements.

2.2 Model species

Coyotes have adapted well to cities by fine-tuning their space-use to the presence of humans while making use of the large availability of resources within. In general, diurnally, coyotes encamp in forested or enclosed urban green areas (Ellington and Gehrt 2019), while nocturnally, they make use of residential areas (Thompson et al. 2021) and vegetated areas (Murray and St Clair 2017; Wurth et al. 2020), potentially moving daily between source areas of non-native plant species and forested land, where they deposit scats.

Coyotes have been found to have important contributions to seed dispersal of native species in temperate regions, e.g., Bearberries (Arctostaphylos glauca), a native species to the Canadian Pacific coast, and Prickly cactus (Opuntia spp.) (Silverstein 2005), potentially including the Eastern prickly-pear cactus (Opuntia humifusa), an endangered species of prickly pear endemic to southern Ontario. They can also be important in the spread of invasive species such as Rose hip (Rosa rugosa), for which seeds have been consistently found in their scats (Garbary et al. 2013). Nonetheless, with opportunistic omnivorous diets, and high germination rates of the seeds they deposit (Silverstein 2005), the plant species effectively dispersed by coyotes are possibly numerous.

2.3 Movement analysis

We used telemetry data from 94 coyotes that were captured, radio-collared, and released within southern Ontario (Fig. 1, Table S1). The coyotes were fitted with GPS collars by the Ontario Ministry of Natural Resources and Forestry and the Toronto Wildlife Centre between 2010 and 2021, following methods described by Wheeldon (2020) and Thompson et al. (2021). GPS collars (Wildcell SG, Lotek Wireless Inc., Newmarket, Ontario) recorded locations at variable fix rates, but location data were rarefied to a constant fix rate of 3 hours. We estimated net displacement at each timepoint by measuring the Euclidean distance between contiguous GPS datapoints (Fig. 2).

2.4 Behavioral factors

We included the interaction of factors influencing coyote behavior: social status, biological season, and climate season. (1) Social status distinguished coyotes following their movement patterns, i.e., (i) resident, if coyote used the same area continuously for ≥ 3 months; (ii) transient, if coyote did not use the same habitat fragments continuously; and (iii) dispersing, if the coyote was traveling in a unidirectional mode. Coyotes with different social statuses were evenly distributed across urban and rural landscapes. (2) Biological seasons included time periods when coyotes are known to breed (January to April), rear pups (May to August), or disperse (September to December); these may not be applicable for every coyote, but they serve to account for anticipated behavioral changes throughout the biological year that might affect net displacement (Sasmal et al. 2019). (3) Climate seasons included time periods defining temperature changes affecting fruit production, resource availability, and foliage acting as shelter: spring (April to June), summer (July to September), autumn (October to December), and winter (January to March). For this, we noted the characteristics at the start point of the movement trajectory and discarded any potential seed dispersal events with differing characteristics between the start point and endpoint.

2.5 Seed dispersal analysis

To understand the implications of changes in net displacement on seed dispersal, we analyzed coyote net displacement over time within four seed retention timeframes (Gelmi-Candusso et al., 2019). For this, we assumed that coyote net displacement within a seed retention timeframe is a potential seed dispersal event, with the start point of each step being a potential seed ingestion/attachment event, and the endpoint of each step being a potential seed deposition/release event (Fig. 2). To reduce autocorrelation between estimates obtained from one movement trajectory, we averaged the net displacements occurring within an equal timeframe for each trajectory.

The seed retention timeframes analyzed herein considered that coyotes can disperse seeds through both epizoochory, whereby seeds may stay attached to coyote fur for up to two days, and endozoochory, whereby seeds reside in the gastrointestinal tract for up to three days in canids (i.e., gut-passage time) (Graae et al. 2004; Hernot et al. 2005; Varela and Bucher 2006; Quick et al. 2017; Draper et al. 2021). Thus, the range of seed retention timeframes included (i) 0–6 hrs, (ii) 6–24 hrs, (iii) 24–48 hrs, and (iv) 48–72 hrs. Steps beyond the 72 hrs time range from a start point were not included in the analysis, as these go beyond seed retention time ranges found in the literature for carnivore seed dispersal.

2.6 Statistical analysis

We analyzed the effect of urbanization (i.e., landscape type) on net displacement within each seed retention timeframe, with individual variation as a random effect, using a generalized linear mixed model (GLMM). We log-transformed the response variable, and followed a gamma distribution using the glmer() function in the lme4 package (Bates et al. 2015).

We integrated the effect of factors influencing behavior (i.e., social status, biological season, and climate season) by analyzing net displacement differences within each factor level in interaction with landscape type using a GLMM. To avoid double interaction factors, we ran the models for each seed retention timeframe. The GLMMs followed a gamma regression with log-transformed data and accounted for individual variation as a random effect. Given the partial temporal overlap between biological and climate seasons, we compared both models using AIC to evaluate which seasonal scale better explained the variability seen in our data.

To understand the potential seed dispersal kernels (i.e., probability density of seed dispersal with increasing distance from source), including how far most seeds would be dispersed in the different landscapes and how far those rare long-distance seed dispersal events would reach, we quantified coyote net displacement in terms of quantiles (5%, 50%, 95%). We considered net displacements in the top 95% as potential long-distance seed dispersal events (Cain et al. 2000). We used the ggpredict() function, which computes predicted values for all possible levels and values from the models’ predictors, to predict net displacement across all our conditions analyzed and thus determine potential seed dispersal distance. While the relationship between net displacement and seed dispersal distance may also be influenced by the complex number of factors determining where a seed will be deposited after being retained, these predictions are meant for comparing the potential seed dispersal distance across all the conditions tested under a constant setting. Future research on effective seed dispersal distance using scat analysis and seed tracking should further analyze any potential seed aggregation patterns following coyote scent-marking behavior (Barrette and Messier 1980).

As expected, the difference in net displacement between urban and rural landscapes was significant for all seed retention timeframes and increased with increasing seed retention timeframe. Based on the GLMM output, coyotes in urban landscapes had a greater decrease in net displacement with increasing seed retention timeframe than coyotes in rural landscapes (Table S3). However, the main urban-rural difference across seed retention timeframes was seen for net displacements in the top 95%, i.e., the potential long-distance seed dispersal events. For these, the difference between urban and rural landscapes increased exponentially with increasing seed retention timeframe, in comparison to net displacements in the lower 50%, which increased linearly with increasing seed retention timeframe (Fig. 3).

The effect of urbanization on net displacement was strongly influenced by social status. Dispersing coyotes were the most affected by urbanization, whereby they exhibited a significant 56–69% decrease in net displacement in urban areas (Fig. 4, Table S2), and they were also the main drivers of potential long-distance seed dispersal events in both landscape types (Table 1). Transient coyotes were also affected by urbanization, whereby they exhibited a significant 15–19% decrease in net displacement in urban areas (Fig. 4, Table S2). The urban-rural difference in net displacement increased linearly with seed retention timeframe for transient coyotes and increased exponentially for dispersing coyotes (Fig. 4). As a result, dispersing coyotes had comparable net displacement to transient coyotes in urban areas, including for longer seed retention timeframes, leading to similar maximum potential seed dispersal distance. In contrast, resident coyotes did not have significant differences in net displacement between landscape types, and this was the case for all seed retention timeframes, whereby they exhibited a 0–8% decrease in net displacement in urban areas (Fig. 4, Table S2).

Table 1

Coyote net displacement with respect to social status while moving within urban or rural landscape. Descriptive statistics for movement ranges across seed retention timeframes are given: i) mean daily net displacement across individuals with 95% confidence interval (CI) range, ii) the maximum distance of 50% of their daily net displacements, iii) the maximum distance of 95% of their daily net displacements, and iv) the number of individuals included in each category.
	Landscape	Seed retention timeframe	Mean ± CI (km)	50% (km)	95% (km)	N
Resident coyotes	URBAN	0-6hrs	0.65 ± 0.10	0.56	1.43	23
		6-24hrs	1.17 ± 0.29	0.98	3.39	23
		24-48hrs	1.37 ± 0.37	1.11	4.09	23
		48-72hrs	1.39 ± 0.34	1.11	3.98	23
	RURAL	0-6hrs	0.84 ± 0.81	0.87	1.75	42
		6-24hrs	1.41 ± 0.14	1.44	3.01	43
		24-48hrs	1.63 ± 0.20	1.56	3.88	42
		48-72hrs	1.67 ± 0.22	1.58	4.31	41
Transient coyotes	URBAN	0-6hrs	0.70 ± 0.23	0.52	1.79	15
		6-24hrs	1.70 ± 0.73	1.10	4.16	16
		24-48hrs	2.73 ± 1.51	1.64	5.92	16
		48-72hrs	3.53 ± 2.10	1.86	7.56	16
	RURAL	0-6hrs	1.07 ± 0.29	0.77	2.24	43
		6-24hrs	2.33 ± 0.42	1.74	5.49	41
		24-48hrs	3.86 ± 0.86	2.53	9.27	40
		48-72hrs	4.82 ± 1.25	3.05	11.40	40
Dispersing coyotes	URBAN	0-6hrs	0.72 ± 0.46	0.43	1.36	4
		6-24hrs	1.75 ± 1.17	0.97	3.70	4
		24-48hrs	3.35 ± 2.80	1.61	7.10	4
		48-72hrs	4.46 ± 3.97	1.89	10.77	4
	RURAL	0-6hrs	1.87 ± 1.43	1.03	4.87	4
		6-24hrs	5.41 ± 4.52	2.54	12.39	4
		24-48hrs	10.65 ± 9.67	4.57	26.09	4
		48-72hrs	15.09 ± 13.71	6.47	35.55	4

Biological seasons influenced net displacement but did not influence the difference in net displacement between urban and rural landscapes, as the decrease in urban areas ranged between 24–39%, comparable to that found in the landscape-only model. The largest and smallest differences in net displacement between urban and rural landscapes were seen during breeding season (38–39% decrease in urban areas) and pup-rearing season (24–29% decrease in urban areas), respectively (Table S2). As with the landscape-only model, the difference in net displacement between urban and rural landscapes increased with increasing seed retention timeframe for all biological seasons, albeit at a lower rate for pup-rearing season (Fig. 4, Table S3).

Climate seasons influenced to a limited extent the effect of urbanization on net displacement. The largest and smallest differences in net displacement between urban and rural landscapes were seen during winter (39–42% decrease in urban areas) and summer (18–26% decrease in urban areas), respectively (Table S2). The difference in net displacement between urban and rural landscapes increased with increasing seed retention timeframe, especially for autumn, winter, and spring (Fig. 4, Table S3). Biological and climate seasons partially overlapped temporally, and both had a strong effect on net displacement across seed retention timeframes; however, AIC differences between the models suggest climate seasonality better explained the variability in our data for all seed retention timeframes.

We found a reduction in the net displacement of coyotes in urban areas, whereby this reduction became stronger with increasing time scale, supporting previous global findings on terrestrial mammal movement (Tucker et al. 2018). In addition, our results suggested that urbanization affects long-distance movements most and more with increasing seed retention timeframe. Including socials status in the model explained why, as urbanization disproportionately affected non-territorial coyotes, which were the main drivers of long-distance movements. In contrast, while biological and climate seasons influenced net displacement, the difference between urban and rural landscapes remained constant across seasons, with a slight decrease during summer and an increase during winter, supporting our initial hypothesis regarding the effect of climate harshness.

Decreased forest cover and increased spatial aggregation of vegetation created by fragmentation has led to shorter mean seed dispersal distance by mesopredators in other landscapes (Herrera et al. 2016), and these are characteristics also found in urban areas. Furthermore, in urban landscapes, the availability of anthropogenic food sources and rodent populations in residential areas may also reduce the foraging travel distance to areas surrounding the habitat patches, potentially reducing the need to travel between habitat fragments. In non-urban fragmented landscapes, the presence of movement corridors can reduce the difference in seed dispersal distance between fragmented and continuous habitats (Herrmann et al. 2016), as they create a directed form of movement between green patches, bypassing landscape fragmentation. Similarly, improving coyote movement corridors within cities may help bypass the inhospitable landscape, thereby alleviating the effect of urbanization on coyote net displacement and its cascading effect on seed dispersal.

Interestingly, our analysis found that urbanization had little effect on the difference in net displacement of resident coyotes between urban and rural landscapes. These results suggest that, in our study area, the main limiting factor for resident coyotes’ net displacement was their territory, rather than landscape fragmentation. However, this might not be the case for resident coyotes living in cities with fewer or smaller green spaces, as in our study area the forested river valleys travelling across the urban matrix provide large areas where resident coyotes can thrive (Thompson et al. 2021), potentially reducing the difference in net displacement between urban and rural landscapes. In contrast to resident coyotes, urbanization had a strong effect on the net displacement of transient and dispersing coyotes (i.e., non-territorial coyotes), suggesting landscape configuration to be a main constraining factor with respect to their net displacement, as these coyotes have wider movement ranges and utilize more inhospitable land, such as smaller suboptimal urban green fragments and residential areas(Newsome et al. 2015; Mitchell et al. 2015; Thompson et al. 2021). This decrease in net displacement strongly reduces the long-distance seed dispersal potential of non-territorial coyotes in urban areas, which can lead to negative implications for plant adaptation to climate change and the restoration of anthropogenically disturbed areas (Cain et al. 2000; Nathan et al. 2008; Mokany et al. 2014).

We found temporal changes in coyote net displacement following biological and climate seasonality. In both landscape types, net displacement decreased during summer and pup-rearing seasons in comparison to the other seasons. Summer and pup-rearing seasons partially overlapped, therefore it is unclear whether this reduced net displacement was due to movement restrictions while caring for dependent pups or an abundance of available resources (Mueller et al. 2011); however, given the model comparison results, our findings suggest the latter might be a more important factor for coyote net displacement. Interestingly, while the overall urban-rural difference in coyote net displacement across seasons was considerably less than when considering coyote social status, the urban-rural difference was twice as large during the harshest season (i.e., winter), than during summer, net displacement was less affected by climate seasonality in urban areas than in rural areas. These findings reflect the more constant availability and widespread abundance of resources in urban areas, the ability of coyotes to exploit anthropogenic food sources, and a higher abundance and richness of prey species exploiting yard features, leading to a wider dietary breadth in urban coyotes (Murray et al. 2015; Hansen et al. 2020; Sugden et al. 2021).

Coyote heterogeneous contribution to seed dispersal

Our results showed differences in coyote net displacement across seasons and social statuses, in both landscape types, albeit to a lesser extent within urban areas. The seasonal differences in coyote net displacement increase the plant specificity in their seed dispersal contribution, providing different seed dispersal range to different plants following their specific phenology and seasonality in fruit production, suggesting future studies should focus on species-specific seed dispersal, especially in rural areas. Most importantly, the intraspecific and seasonal diversity in coyote movement patterns, habitat use (Kamler and Gipson 2000), and food consumption (Bartel and Orrock 2022) create functional differences in the seed dispersal kernels coyotes create. A heterogenous array of seed dispersal kernels provide plant species with different movement pathways and diverse microhabitats for deposition, and have been found to be key in habitat regeneration (Gonzalez-Varo et al. 2013; Escribano-Avila et al. 2014), suggesting an important role of coyotes and likely other wide-ranging omnivore predators in urban habitat renaturalization.

Urbanization limited the net displacement of non-territorial coyotes, had a stronger effect for seeds with longer retention times, and predominantly affected long-distance seed dispersal events. Maintaining animal movement between vegetated fragments by enhancing connectivity is a crucial factor in restoring seed dispersal by terrestrial wildlife in urban areas. Our study also suggests coyotes have a high likelihood of providing an intraspecific heterogeneous contribution to seed dispersal, given their complex social dynamics and biological/climate-driven seasonality in their movement patterns, indicating the potential species-specific effect of seed dispersal by coyotes. Future research identifying seed species dispersed in temperate climates through scat analysis, tracking seeds to quantify effective seed dispersal distance, and defining urban foraging areas through behavioral state analysis, is needed to further understand the ecological implications of seed dispersal by coyotes, in terms of spread of invasive species and their contribution to urban forest biodiversity.

Animal Ethics statement

The collaring of the animals used for this study was approved by the Trent University Animal Care Committee (protocol no. 10016) and the Ontario Ministry of Natural Resources Wildlife Animal Care Committee (protocol nos. 10-219, 11-219, 12-219, 75-12, 75-13, 75-14, 75-16, 75-17, 75-18, 75-19, 75-20, 75-21).

Author Contributions

T.A.G.C. conceived the study, designed the model and the computational framework, implemented the model, analyzed the data, drafted the manuscript, and designed the figures. T.J.W. and B.R.P. collected and curated the coyote tracking data and provided comments and edits on the manuscript. M-J.F. supervised the development and findings of this work and contributed to the writing of the manuscript. All authors provided critical feedback and helped shape the research, analysis, and manuscript.

Conflict of Interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Ethics approval

Ethics approval was obtained to conduct live-trapping and collaring.

Availability of data and code

The net displacement data will be deposited in Dryad upon acceptance of the manuscript.

Code availability

Code available on the corresponding author’s Github.

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The effect of urbanization and behavioral factors on coyote net displacement and its implications for seed dispersal

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Abstract

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1. Introduction

2. Methods

2.1. Study area

2.2 Model species

2.3 Movement analysis

2.4 Behavioral factors

2.5 Seed dispersal analysis

2.6 Statistical analysis

3. Results

4. Discussion

5. Conclusions

Declarations

Author Contributions

Conflict of Interest

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Availability of data and code

Code availability

References

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Supplementary Files

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