With many natural landscapes undergoing restoration efforts, there is a growing need for the optimization of direct seeding practices. Seeds planted on wildlands are often consumed by rodents leading to reduced plant establishment. Coating seeds in rodent aversive products may prevent seed predation. We tested ten seed-coating formulations containing products expected to deter rodents, namely: ghost and cayenne pepper powders; essential oils from bergamot, neem, and pine; methyl-nonyl-ketone, anthraquinone, activated carbon, beta-cyclodextrin and a blank coating containing no rodent deterrents to serve as a control treatment. Each treatment was applied to Pseudoroegneria spicata (bluebunch wheatgrass) seeds. These seeds germinated similarly to uncoated control seeds unless the coating contained methyl-nonyl-ketone which reduced germination. Seeds were offered to Ord's kangaroo rats (Dipodomys ordii) that strongly avoided the treatments in favor of uncoated control seeds. Notably, the blank coating, lacking active ingredients, still elicited 99% avoidance. However, these results indicated behavior when alternative food sources are readily available, a scenario rare in nature. To address this, a second feeding experiment was conducted to observe D. ordii's behavior under calorie-restricted conditions. D. ordii were subjected to a fast period and then offered only one treatment. Under these conditions, many subjects chose to consume coated seeds, but to a lesser degree than subjects offered control seeds. Seeds coated in ghost pepper, neem oil, and activated carbon reduced consumption by 47-50%. Given these lab results, we would expect these seed-coatings to increase the establishment of native seeds following the direct seeding of wildlands by deterring rodent seed-predation.

The landscapes of the Great Basin and Mojave Deserts are changing due to plant invasion. Highly flammable invasive grasses increase the size and frequency of fire causing a cascade of effects through the plant and animal communities. One of the most influential animal groups in desert systems is small mammals. We sought to learn how small mammals are impacted by fire and how their influence on the plant community differs between burned and unburned habitat. Small mammals did not have higher rates of mortality as a direct result of a controlled burn. In the Great Basin, there were short-term reductions in abundance, richness, and diversity of the small mammal community in burned plots. In the Mojave, species richness and diversity increased in burned plots shortly after fire and no abundance differences were detected. These results correspond with our prediction based on the dominant small mammal species at each site. Small mammals are primarily granivores; however, they also have strong impacts on the plant community via folivory. We tested for small mammal impacts on seedling survival in burned and unburned habitat. Small mammal access, burned vs. unburned habitat, and plant species were all important determinants of survival. Small mammals greatly reduced survival at both sites in burned and unburned habitat and often had a stronger impact in unburned than burned plots. Accounting for small mammal folivory may be a crucial step in successful post-fire rehabilitation. Finally, we used seed trays to test how small mammals influence the persistence of seed on the landscape. Small mammals reduced persistence of an invasive and native plant species in the Great Basin in 2012, yet a year later when small mammal abundance was reduced, no small mammal effect was observed. In the Mojave, persistence was reduced for the majority of species both years of the study. Small mammals did not appear to avoid seed of invasive plant species as we had predicted and may be important consumers reducing the reproductive potential of these invaders. If small mammals do prefer non-native seedlings over natives and are also consuming non-native seed, they may be greatly reducing the presence of non-natives both on the unburned landscape as well as after fire. Non-native consumption by small mammals could aid in the biotic resistance of these desert ecosystems. This research further enforces the important role that small mammals play as consumers, dispersers, and regulators of the plant community.

Granivores are important components of sagebrush communities in western North America. These same regions are being altered by the invasion of the exotic annual Bromus tectorum (cheatgrass) that alters physical and biological dynamics in ways that appear to promote its persistence. This research directly relates to the restoration of B. tectorum-dominated systems in two inter-related ways. First, because these landscapes have large quantities of seeds applied during restoration, it is important to determine the major granivore communities in intact sagebrush communities and in nearby cheatgrass-dominated communities. Second, it is important to develop an understanding of patterns of seed harvest by granivores. In addition to the data chapters there are two review chapters; Chapter 1 highlights factors contributing to seed removal and Chapter 7 provides ecologically based techniques that could minimize the negative consequences of granivores during ecological restoration. Common groups of ants showed increased abundances; uncommon species and functional groups were generally negatively impacted by cheatgrass (Chapter 2). Conversely, rodents were negatively impacted by conversion to cheatgrass (Chapter 4). Ant seed removal was highly context-dependent (Chapter 3), depending on the background vegetation (large-scale among-patch effects), foraging distance from the nest mound (small-scale among-patch effects), and the presence of other seed species in mixture (within-patch effects). In addition, cheatgrass provided associational resistance to native seeds in mixture, meaning the presence of cheatgrass increased native seed survival. In Chapter 5 a novel statistical technique in the ecological sciences showed that rodents have marked preferences for some seeds over others and that more seeds were removed in sagebrush compared to cheatgrass-dominated sites, although associational effects among seed mixtures were not detected. In Chapter 6 we show that the amount of seed harvested depended on both intraspecific and interspecific seed density. B. tectorum seeds had associational susceptibility (increased harvest) in the presence of native seeds. Although the reciprocal effect may occur, we did not find statistical support for it. These sets of studies are not only of basic ecological interests, but are also important for developing management strategies for restoration of these degraded lands.

The effect of shared enemies between invasive and native species has been argued to facilitate biological invasions (i.e., the apparent competition hypothesis or ACH). This study investigated a previously untested possibility: whether granivorous small mammals facilitate cheatgrass (Bromus tectorum) invasion by driving food-mediated apparent competition between cheatgrass and native grasses. Specifically, we tested three predictions that must be true if such apparent competition occurs. First, cheatgrass invasion augments total seeds available to granivorous small mammals. Second, density of granivorous small mammals increases in response to increased seed availability (simulated with experimental additions of cheatgrass seeds). Third, granivorous small mammals prefer seeds from native grasses over cheatgrass seeds. We tested these predictions in the Great Basin Desert of Utah, USA. Cheatgrass invasion augmented total yearly seed production. Granivorous small mammals preferred native seeds over cheatgrass seeds. However, neither abundance, richness, nor diversity of granivorous small mammals increased in response to experimental additions of cheatgrass seed. We therefore conclude that granivorous small mammals did not drive food-mediated apparent competition during the study period. The lack of support for the ACH in this study may suggest that the role of small mammal-driven apparent competition is either unimportant in the Great Basin, or that the appropriate indirect interactions between small mammals, cheatgrass, and native grasses have yet to be evaluated. Testing the third prediction required the separation of seeds from the soil matrix. We employed a chemical flotation methodology to recover target seeds from soil, and developed a novel method of decanting target material from the flotation solution. We compared the utility of the novel method to that of a traditional decantation method. Specifically, we compared effectiveness (the proportion of seeds recovered from a known sample), rapidity (the time required to decant that sample), efficiency (the number of seeds decanted per second), and recovery bias (the effect of relative density on seed recovery) between methods. Our proposed method was more effective, more rapid, more efficient, and less biased than the traditional method. Therefore, any future work relying on flotation to analyze seed banks should clearly describe how samples are decanted and should consider the proposed method as a potential means of enhancing the efficiency of chemical flotation.

Promouvoir le service de contrôle biologique des bio-agresseurs des cultures nécessite de comprendre le fonctionnement des populations d’ennemis naturels dans des mosaïques agricoles qui sont hétérogènes et dynamiques. Cette thèse porte sur l’étude des déterminants de l’abondance des carabes impliqués dans la régulation des adventices via la consommation de leurs graines. L’analyse des variations d’abondance des espèces à une échelle nationale nous a permis d’évaluer le rôle relatif des facteurs locaux et paysagers et de mettre en évidence le rôle important de la proportion de colza et de prairie dans le paysage sur l’abondance de carabes consommateurs de graines dans les parcelles cultivées. Le suivi expérimental de la composition des communautés de carabes ainsi que du statut nutritionnel et des mouvements de plusieurs espèces dans des dispositifs colza/blé et colza/bordure pérenne sur l’ensemble de la période d’activité des carabes nous a permis de mettre en évidence que (i) les carabes répondent l’hétérogénéité de cette mosaïque agricole (type de culture, qualité de l’habitat au sein d’un même type de culture), avec des réponses souvent spécifiques (ii) que le colza d’hiver joue un rôle important dans le fonctionnement des populations de carabes consommateurs de graines du fait de sa haute qualité d’habitat pour ces espèces, (iii) que la variation de la qualité d’habitat peut jouer sur le mouvement des carabes vers l’habitat adjacent et (iv) que la présence d’une bordure pérenne au voisinage d’une culture favorise le maintien et le fonctionnement de plusieurs espèces consommatrices de graines d’adventices dans la mosaïque agricole. / Promoting biological pest control in crops requires to understand the functioning of natural enemies in heterogeneous and dynamic agricultural mosaics. This thesis focuses on the study of the determinants that influence the abundance of ground beetles involved in the regulation of weeds via the consumption of their seeds. In an analysis of the variations of species abundance at a national scale we identified the respective role of local and landscape factors and showed that oilseed rape and grassland proportions in the landscape were key determinants of the abundance of carabid species in cultivated fields. We then studied carabid communities through combined measured of their abundance, nutritional status and between-habitat movements, during an entire cropping season, in pairs of habitats consisting in winter oilseed rape fields with either a crop (winter cereal fields) or a semi-natural habitat (grassy field margins) as adjacent habitat. We demonstrated that (i) ground beetles respond to the heterogeneity of arable mosaic (crop type, habitat quality within the same crop type), often with specific responses, (ii) winter oilseed rape plays an important role in the functioning of weed seed-eating carabid species due to its high habitat quality, (iii) variations in habitat quality can influence the movement of beetles to the adjacent habitat and (iv) the presence of a grassy field margin in the vicinity of a crop promotes the maintenance and functioning of several species in arable mosaics.

Crop rotation may be used to prevent the continuous selection of particular weed species adapted to one crop type. This might be useful for weed management, economy in herbicide applications and promoting biodiversity. Common simple crop sequences might be diversified by introducing perennial forage crops. Impacts of such perennial crops on weeds were studied with four approaches : 1) Large-scale weed surveys in 632 fields in western France showed that weed species composition differed most strongly between perennial alfalfa crops and annual crops. Comparisons of fields before, during and after perennial alfalfa suggested that community composition varies in a cyclic way during such crop rotations. Several weed species problematic in annual crops were suppressed during and after perennial crops, but the appearance of other species led to equal or even higher plant diversities. 2) A 3-year field experiment with contrasting crop management options allowed an investigation of the underlying mechanisms for this: The absence of soil tillage reduced weed emergence but increased the survival of established plants. The permanent vegetation cover and frequent hay cuttings reduced weed growth, plant survival and seed production. 3) Greenhouse experiments testing the regrowth ability of individual plants after cutting showed strong differences between species and functional groups. An two-factorial experiment suggested that the negative impacts of cutting and competition on weed growth were mainly additive. 4) Special measurements of weed seed predation in the field experiment showed positive correlations with vegetation cover, indicating that this ecosystem service may be particularly fostered by perennial crops. Consistent preferences of seed predators for certain weed species indicates that seed predation may be another cause of the observed weed community shifts.

La rotation de cultures peut être utilisée pour empêcher la sélection continue d’espèces adventices adaptées à un type de culture. Elle pourrait favoriser la gestion des adventices, l’économie d’herbicides et la biodiversité. Les successions de cultures simples d’aujourd’hui pourraient être diversifiées par des cultures fourragères pérennes. Les impacts des ces cultures sur les adventices ont été étudié utilisant quatre approches : 1) Des relevés d’adventices sur 632 champs dans l’ouest de la France ont montré que la composition spécifique varie le plus entre des cultures fourragères pérennes et des cultures annuelles. Une comparaison des champs avant, pendant, et après des cultures fourragères pérennes a suggéré que la composition des communautés varie d’une manière cyclique pendant ces rotations. Plusieurs espèces problématiques dans des cultures annuelles ont été supprimées pendant et après les cultures pérennes, mais l’apparition d’autres espèces a produit une diversité de plantes comparable, voire supérieure. 2) Une expérimentation au champ de trois ans avec des modes de gestion contrastés a permis d’étudier les mécanismes sous-jacents: L’absence de travail du sol a réduit la levée des adventices, mais a augmenté la survie des plantes adultes. Le couvert végétal permanent et les fauches fréquentes ont réduit la croissance, la survie des plantes et la production de graines. 3) Des expérimentations sous serre analysant la croissance poste fauche de plantes individuelles ont montré des différences importantes entre espèces et groupes fonctionnels. Une expérimentation à deux facteurs a suggéré que les impacts négatifs de la fauche et de la compétition sur la croissance des adventices ont été additifs. 4) Des mesures spéciales de prédation de graines d’adventices sur l’expérimentation au champ ont montré des corrélations positives avec le couvert végétal et la prédation, indiquant une importance particulière de ce service écosystémique dans des cultures pérennes. La préférence des graines de certaines espèces montre que la prédation de graines peut être une autre cause des changements de communautés d’adventices. / Crop rotation may be used to prevent the continuous selection of particular weed species adapted to one crop type. This might be useful for weed management, economy in herbicide applications and promoting biodiversity. Common simple crop sequences might be diversified by introducing perennial forage crops. Impacts of such perennial crops on weeds were studied with four approaches : 1) Large-scale weed surveys in 632 fields in western France showed that weed species composition differed most strongly between perennial alfalfa crops and annual crops. Comparisons of fields before, during and after perennial alfalfa suggested that community composition varies in a cyclic way during such crop rotations. Several weed species problematic in annual crops were suppressed during and after perennial crops, but the appearance of other species led to equal or even higher plant diversities. 2) A 3-year field experiment with contrasting crop management options allowed an investigation of the underlying mechanisms for this: The absence of soil tillage reduced weed emergence but increased the survival of established plants. The permanent vegetation cover and frequent hay cuttings reduced weed growth, plant survival and seed production. 3) Greenhouse experiments testing the regrowth ability of individual plants after cutting showed strong differences between species and functional groups. An two-factorial experiment suggested that the negative impacts of cutting and competition on weed growth were mainly additive. 4) Special measurements of weed seed predation in the field experiment showed positive correlations with vegetation cover, indicating that this ecosystem service may be particularly fostered by perennial crops. Consistent preferences of seed predators for certain weed species indicates that seed predation may be another cause of the observed weed community shifts. / Fruchtfolgen können dazu dienen, die kontinuierliche Selektion von Unkrautarten zu verhindern, die an eine bestimmte Kultur angepasst sind. Dies könnte dem Unkrautmanagement, der Einsparung von Herbiziden, und der Biodiversität dienen. Heutige, sehr einfache Furchtfolgen könnten durch mehrjährige Futterkulturen diversifiziert werden. Die Einflüsse solcher mehrjähriger Kulturen auf Unkräuter wurden in vier Ansätzen untersucht : 1) Vegetationsaufnahmen auf 632 Feldern in Westfrankreich zeigten, dass die Unkrautzusammensetzung zwischen mehrjährigen Futterkulturen und einjährigen Kulturen stark variiert. Der Vergleich von Feldern vor, während und nach mehrjährigen Futterkulturen legte nahe, dass die Pflanzengemeinschaft während solcher Fruchtfolgen zyklisch variiert. Mehrere problematische Unkrautarten wurden während und nach den mehrjährigen Kulturen zurückgedrängt. Das Auftauchen anderer Arten führte jedoch zu einer gleichbleibenden oder leicht erhöhten Pflanzenvielfalt. 2) Ein dreijähriger Feldversuch mit verschiedenen Bearbeitungsoptionen ermöglichte es, die zugrunde liegenden Mechanismen zu untersuchen: Die fehlende Bodenbearbeitung hat das Auflaufen der Unkräuter reduziert und das Überleben der adulten Pflanzen erhöht. Die permanente Vegetationsbedeckung und die häufigen Heuschnitte haben das Wachstum, das Überleben und die Samenproduktion vermindert. 3) Gewächshausexperimente zum Nachwachsen von Unkrautpflanzen nach Heuschnitten zeigten große Unterschiede zwischen verschiedenen Arten und funktionellen Gruppen. Ein Experiment mit zwei Faktoren lässt vermuten, dass die negativen Effekte der Schnitte und der Konkurrenz auf das Unkrautwachstum sich addieren. 4) Spezielle Messungen der Prädation von Unkrautsamen auf den untersuchten Feldern zeigten positive Korrelationen mit der Vegetationsbedeckung, was auf eine besondere Wichtigkeit dieser Ökosystemdienstleistung in ausdauernden Kulturen hindeutet. Die Präferenz von bestimmten Samenarten deutet darauf hin, dass Samenprädation ein weiterer Grund für die beobachteten Änderungen der Unkrautgemeinschaften sein kann.