Brock R. McMillan

SPACE USE AND SOCIALITY OF RIVER OTTERS (LONTRA CANADENSIS) IN MINNESOTA

Abstract The river otter (Lontra canadensis) has a complex social system, which varies widely across its range. We examined patterns of space use and social interactions for a native population of river otters in southeastern Minnesota. We radiomarked 28 river otters and monitored annual home ranges and core areas, static and dynamic interactions, and site fidelity. We compared these characteristics and interactions between sexes and age classes. Annual home ranges of male river otters were 3.2 times greater than those of females (P = 0.042), and annual core areas of males were 2.9 times greater than those of females (P = 0.083). The static interactions among river otters were extensive, with 69% of the individuals exhibiting core-area overlap. Overall dynamic interactions were positive (i.e., animals were closer together than expected). Males used 74% of their year 1 home range during year 2, and females used 75%. Females on average used only 39% of their core area from year 1 during year 2, whereas males used 65%. In general, conspecifics were not excluded from home ranges or core areas and signs of cooperation were evident, suggesting that river otters in southeastern Minnesota were social rather than territorial.

Biotic resistance and disturbance: rodent consumers regulate post‐fire plant invasions and increase plant community diversity

Biotic resistance and disturbance are fundamental processes influencing plant invasion outcomes; however, the role of consumers in regulating the establishment and spread of plant invaders and how disturbance modifies biotic resistance by consumers is unclear. We document that fire in combination with experimental exclusion of rodent consumers shifted a native desert shrubland to a low-diversity, invasive annual grassland dominated by Bromus tectorum (cheatgrass). In contrast, burned plots with rodents present suppressed invasion by cheatgrass and developed into a more diverse forb community. Rodents created strong biotic resistance to the establishment of aggressive plant invaders likely through seed and seedling predation, which had cascading effects on plant competition and plant community diversity. Fire mediated its positive effects on plant invaders through native plant removal and by decreasing the abundance and diversity of the rodent community. The experimental disruption of plant and consumer-mediated biotic resistance of plant invaders using fire and rodent exclusion treatments provides strong evidence that native plants and rodents are important regulators of plant invasion dynamics and plant biodiversity in our study system. While rodents conferred strong resistance to invasion in our study system, fluctuations in rodent populations due to disturbance and climatic events may provide windows of opportunity for exotic plant species to escape biotic resistance by rodent consumers and initiate invasions.

Negative effects of an exotic grass invasion on small-mammal communities.

Exotic invasive species can directly and indirectly influence natural ecological communities. Cheatgrass (Bromus tectorum) is non-native to the western United States and has invaded large areas of the Great Basin. Changes to the structure and composition of plant communities invaded by cheatgrass likely have effects at higher trophic levels. As a keystone guild in North American deserts, granivorous small mammals drive and maintain plant diversity. Our objective was to assess potential effects of invasion by cheatgrass on small-mammal communities. We sampled small-mammal and plant communities at 70 sites (Great Basin, Utah). We assessed abundance and diversity of the small-mammal community, diversity of the plant community, and the percentage of cheatgrass cover and shrub species. Abundance and diversity of the small-mammal community decreased with increasing abundance of cheatgrass. Similarly, cover of cheatgrass remained a significant predictor of small-mammal abundance even after accounting for the loss of the shrub layer and plant diversity, suggesting that there are direct and indirect effects of cheatgrass. The change in the small-mammal communities associated with invasion of cheatgrass likely has effects through higher and lower trophic levels and has the potential to cause major changes in ecosystem structure and function.

Wallowing Behavior of American Bison (Bos bison) in Tallgrass Prairie: an Examination of Alternate Explanations

Abstract Wallowing is a common behavior of American bison (Bos bison). Past explanations and current hypotheses suggested for wallowing behavior include grooming behavior associated with shedding, male-male interaction (typically rutting behavior), social behavior for group cohesion, play behavior, relief from skin irritation due to biting insects, reduction of ectoparasite (tick and lice) load and thermoregulation. We monitored circannual and circadian patterns of wallowing frequency by American bison during 1996–1997 in the tallgrass prairie region of eastern Kansas. Wallowing activity increased from April to late June or July (during 1996 and 1997, respectively), decreased during midsummer, peaked again in September, decreased from September to October and then remained low from November to March. Diurnally, wallowing was low in early morning, increased to a peak in early afternoon and then decreased during mid afternoon and evening. Within the herd adult males wallowed more frequently than adult females and both adult males and females wallowed more frequently than yearlings. We observed behaviors that were consistent with all of the hypotheses previously suggested to explain wallowing behavior by bison. Based on our observations we suggest that the alternate explanations for wallowing behavior are not mutually exclusive. However, only the relief from biting insects hypotheses was consistent with both the circannual and circadian patterns of frequency of wallowing by American bison.

Vegetation Responses to an Animal-generated Disturbance (Bison Wallows) in Tallgrass Prairie

Abstract Bison (Bos bison) were a keystone species in the tallgrass prairie region of the Great Plains of North America. Cattle (Bos taurus) have been described as a functional equivalent to bison and have replaced bison in most of the grassland that remains intact. However, non-grazing behaviors influence grassland dynamics and are dissimilar between bison and cattle. Wallowing behavior (a non-grazing behavior by bison, but not cattle) creates disturbances (wallows) that were a common feature (may have numbered more than 100 million) of tallgrass prairie prior to extirpation of bison and conversion of most land to row-crop agriculture. We hypothesized that wallows are a unique disturbance that significantly influence both the structure and function of tallgrass prairie. We examined the response of plants to wallowing disturbances on Konza Prairie Biological Station, Kansas, where a herd of approximately 200 bison had year-round access to 1000 ha of native tallgrass prairie. We determined the influence of this disturbance type on aboveground net primary production (ANPP), plant species richness and diversity, and plant life form richness and diversity. The ANPP at the edge of wallowing disturbances was double the production in wallows and in adjacent prairie, which were not different from year to year. Plant species richness and diversity were significantly lower in wallows than at the edge of wallows and in adjacent prairie during all years of the study. However, composition of species were dissimilar among locations (e.g., 16% of all plant species samples were found only in the wallows); and, therefore, wallows increased the local and likely regional diversity of plant species. Likewise, wallowing disturbances increased local richness and diversity of plant life forms. It appears that these once common wallowing disturbances increase the diversity of tallgrass prairie beyond that of prairie that is grazed only (i.e., prairie with cattle).

Habitat enhancement and native fish conservation: can enhancement of channel complexity promote the coexistence of native and introduced fishes?

Native fishes worldwide have declined as a consequence of habitat loss and degradation and introduction of non-native species. In response to these declines, river restoration projects have been initiated to enhance habitat and remove introduced fishes; however, non-native fish removal is not always logistically feasible or socially acceptable. Consequently, managers often seek to enhance degraded habitat in such a way that native fishes can coexist with introduced species. We quantified dynamics of fish communities to three newly constructed side channels in the Provo River, Utah, USA, to determine if and how they promoted coexistence between native fishes (nine species) and non-native brown trout (Salmo trutta L.). Native and introduced fishes responded differently in each side channel as a function of the unique characteristics and histories of side channels. Beaver activity in two of the three side channels caused habitat differentiation or channel isolation that facilitated the establishment of native species. The third side channel had greater connectivity to and similar habitat as the main channel of the Provo River, resulting in a similar fish community to main channel habitats (i.e. dominated by brown trout with only a few native fish species). These results demonstrate the importance of understanding habitat preferences for each species in a community to guide habitat enhancement projects and the need to create refuge habitats for native fishes.

LONG-TERM STUDY OF ABUNDANCE OF THE HISPID COTTON RAT IN NATIVE TALLGRASS PRAIRIE

Abstract We studied patterns of and factors influencing abundance of hispid cotton rats (Sigmodon hispidus) in tallgrass prairie habitats from autumn 1981 to spring 2001 at the Konza Prairie Biological Station, Kansas. Abundance of cotton rats was low during autumn (X̄ = 0.57 individuals/trapline) and extremely low in spring (X̄ = 0.04 individuals/trapline). Autumn abundance varied widely in 20 years (range: 0.00–2.50 individuals/trapline) and among 14 sites (0.05–1.35 individuals/trapline). Spring prairie fires had a positive influence on cotton rats during the 1st (X̄ = 0.8 individuals/trapline) and 2nd autumns (X̄ = 1.0 individuals/trapline), but not during the 3rd or later autumns (X̄ = 0.2 individuals/trapline). Cotton rats were associated strongly with lowland prairie relative to breaks and upland prairie. Autumn abundance was correlated positively with previous winter (December–February) average maximum temperature, but was not correlated with previous winter precipitation, summer (June–August) average maximum temperature, summer precipitation, or aboveground net primary productivity. Although previous winter maximum temperature accounted for only 32% of interautumn variation in abundance, winter harshness appears to be the major factor driving temporal variation in autumn abundance of hispid cotton rats.

ENVIRONMENTAL FACTORS AFFECTING BIOMASS AND DISTRIBUTION OF STUCKENIA PECTINATA IN THE HERON LAKE SYSTEM, MINNESOTA, USA

The Heron Lake System historically has been an important resource for waterfowl in Southern Minnesota, USA. In the early 1900s, the system was a major nesting, feeding, and staging area for breeding and migrating waterfowl mainly due to the extensive growth of sago pondweed, Stuckenia pectinata. In recent years, the abundance of S. pectinata has decreased dramatically. We conducted a study from 2002 through 2003 to identify factors limiting the success of S. pectinata. Distribution and biomass of S. pectinata differed significantly over the years of this study. Biomass was related to environmental factors, including water transparency and water temperature early in the season. Water transparency during May (time of early growth) had a negative relationship with maximum biomass at each site over all years. Water temperature had a positive relationship with increases in seasonal biomass yield of S. pectinata in the Heron Lake System.

Water Developments and Canids in Two North American Deserts: A Test of the Indirect Effect of Water Hypothesis

Anthropogenic modifications to landscapes intended to benefit wildlife may negatively influence wildlife communities. Anthropogenic provisioning of free water (water developments) to enhance abundance and distribution of wildlife is a common management practice in arid regions where water is limiting. Despite the long-term and widespread use of water developments, little is known about how they influence native species. Water developments may negatively influence arid-adapted species (e.g., kit fox, Vulpes macrotis) by enabling water-dependent competitors (e.g., coyote, Canis latrans) to expand distribution in arid landscapes (i.e., indirect effect of water hypothesis). We tested the two predictions of the indirect effect of water hypothesis (i.e., coyotes will visit areas with free water more frequently and kit foxes will spatially and temporally avoid coyotes) and evaluated relative use of free water by canids in the Great Basin and Mojave Deserts from 2010 to 2012. We established scent stations in areas with (wet) and without (dry) free water and monitored visitation by canids to these sites and visitation to water sources using infrared-triggered cameras. There was no difference in the proportions of visits to scent stations in wet or dry areas by coyotes or kit foxes at either study area. We did not detect spatial (no negative correlation between visits to scent stations) or temporal (no difference between times when stations were visited) segregation between coyotes and kit foxes. Visitation to water sources was not different for coyotes between study areas, but kit foxes visited water sources more in Mojave than Great Basin. Our results did not support the indirect effect of water hypothesis in the Great Basin or Mojave Deserts for these two canids.

Site Characteristics of River Otter (Lontra canadensis) Natal Dens in Minnesota

Abstract Knowledge of the habitat and spatial characteristics of natal dens used by female river otters (Lontra canadensis) is limited to a few observations. Documentation of these characteristics in human-altered landscapes may improve management opportunities for this species. We monitored 8 adult (>2 y old) radio-marked female otters during the natal denning season (March–May) in southeastern Minnesota during 2003–2004 and quantified 10 micro- and 2 macro-habitat characteristics of dens. Females began denning in March, with a mean initiation date of 31 March, and used natal dens for a mean of 49 d (se = 3). Two females used man-made brush piles as dens, four used small limestone caves, one used a cavity in the roots of a big-toothed aspen (Populus grandidentata) and one used a beaver (Castor canadensis) bank-den. Dens were located a mean of 316 m (se = 79) from the nearest body of water and averaged 61 m (se = 15) of elevation higher than the nearest body of water. Seven of eight females placed dens outside of their normal activity areas, and all females appeared to select den sites that were protected from flood events. Our results suggest that to promote successful reproduction of river otter populations, managers should protect potential denning sites in upland habitats adjacent to bodies of water.