Detection of Xylem Cavitation in Corn under Field Conditions

We report the detection of cavitation events in corn (Zea mays) plants growing under field conditions in Greeley, CO. To our knowledge this study reports the first successful attempt to monitor continuously for long periods the cavitation events of a crop plant using acoustic detection techniques. Cavitation events occur in corn plants using acoustic detection techniques. Cavitation events occur in corn plants irrigated daily when the xylem pressure potentials fall below about −1.0 megapascals. In unirrigated corn we estimate that approximately half of all vessels cavitate on any one day when xylem pressure potentials fall below about −1.8 megapascals. We postulate that root pressure developed every night in irrigated and unirrigated corn is adequate to rejoin cavitated water columns.     

How does synchrony with host plant affect the performance of an outbreaking insect defoliator?

Phenological mismatch has been proposed as a key mechanism by which climate change can increase the severity of insect outbreaks. Spruce budworm (Choristoneura fumiferana) is a serious defoliator of North American conifers that feeds on buds in the early spring. Black spruce (Picea mariana) has traditionally been considered a poor-quality host plant since its buds open later than those of the preferred host, balsam fir (Abies balsamea). We hypothesize that advancing black spruce budbreak phenology under a warmer climate would improve its phenological synchrony with budworm and hence increase both its suitability as a host plant and resulting defoliation damage. We evaluated the relationship between tree phenology and both budworm performance and tree defoliation by placing seven cohorts of budworm larvae on black spruce and balsam fir branches at different lags with tree budburst. Our results show that on both host plants, spruce budworm survival and pupal mass decrease sharply when budbreak occurs prior to larval emergence. By contrast, emergence before budbreak decreases survival, but does not negatively impact growth or reproductive output. We also document phytochemical changes that occur as needles mature and define a window of opportunity for the budworm. Finally, larvae that emerged in synchrony with budbreak had the greatest defoliating effect on black spruce. Our results suggest that in the event of advanced black spruce phenology due to climate warming, this host species will support better budworm survival and suffer increased defoliation.     

Extreme weather events and plant–plant interactions: shifts between competition and facilitation among grassland species in the face of drought and heavy rainfall

Biotic interactions play an important role in ecosystem function and structure in the face of global climate change. We tested how plant–plant interactions, namely competition and facilitation among grassland species, respond to extreme drought and heavy rainfall events. We also examined how the functional composition (grasses, forbs, legumes) of grassland communities influenced the competition intensity for grass species when facing extreme events. We exposed experimental grassland communities of different functional compositions to either an extreme single drought event or to a prolonged heavy rainfall event. Relative neighbour effect, relative crowding and interaction strength were calculated for five widespread European grassland species to quantify competition. Single climatic extremes caused species specific shifts in plant–plant interactions from facilitation to competition or vice versa but the nature of the shifts varied depending on the community composition. Facilitation by neighbouring plants was observed for Arrhenatherum elatius when subjected to drought. Contrarily, the facilitative effect of neighbours on Lotus corniculatus was transformed into competition. Heavy rainfall increased the competitive effect of neighbours on Holcus lanatus and Lotus corniculatus in communities composed of three functional groups. Competitive pressure on Geranium pratense and Plantago lanceolata was not affected by extreme weather events. Neither heavy rainfall nor extreme drought altered the overall productivity of the grassland communities. The complementary responses in competition intensity experienced by grassland species under drought suggest biotic interactions as one stabilizing mechanism for overall community performance. Understanding competitive dynamics under fluctuating resources is important for assessing plant community shifts and degree of stability of ecosystem functions.     

Clumped distribution of oak leaf miners between and within plants

Leaf miners typically show non-random distributions both between and within plants. We tested the hypothesis that leaf miners on two oak species were clumped on individual host trees and individual branches and addressed whether clumping was influenced by aspects of plant quality and how clumping and/or interactions with other oak herbivores affected leaf-miner survivorship. Null models were used to test whether oak herbivores and different herbivore guilds co-occur at the plant scale. Twenty individual Quercus geminata plants and 20 Quercus laevis plants were followed over the season for the appearance of leaf miners and other herbivores, and foliar nitrogen, tannin concentration, leaf toughness and leaf water content were evaluated monthly for each individual tree. The survivorship of the most common leaf miners was evaluated by following the fate of marked mines in several combinations that involved intra- and inter-specific associations. We observed that all leaf miners studied were clumped at the plant and branch scale, and the abundance of most leaf-miner species was influenced by plant quality traits. Mines that occurred singly on leaves exhibited significantly higher survivorship than double and triple mines and leaves that contained a mine or a leaf gall and a mine and damage by chewers exhibited lowest survivorship. Although leaf miners were clumped at individual host trees, null model analyses indicated that oak herbivores do not co-occur significantly less than expected by chance and there was no evidence for biological mechanisms such as inter-specific competition determining community structure at the plant scale. Thus, despite co-occurrence resulting in reduced survivorship at the leaf scale, such competition was not strong enough to structure separation of these oak herbivore communities.     

Root endophyte symbiosis in vitro between the ectomycorrhizal basidiomycete Tricholoma matsutake and the arbuscular mycorrhizal plant Prunus speciosa

We previously reported that Tricholoma matsutake and Tricholoma fulvocastaneum, ectomycorrhizal basidiomycetes that associate with Pinaceae and Fagaceae, respectively, in the Northern Hemisphere, could interact in vitro as a root endophyte of somatic plants of Cedrela odorata (Meliaceae), which naturally harbors arbuscular mycorrhizal fungi in South America, to form a characteristic rhizospheric colony or “shiro”. We questioned whether this phenomenon could have occurred because of plant–microbe interactions between geographically separated species that never encounter one another in nature. In the present study, we document that these fungi formed root endophyte interactions and shiro within 140 days of inoculation with somatic plants of Prunus speciosa (=Cerasus speciosa, Rosaceae), a wild cherry tree that naturally harbors arbuscular mycorrhizal fungi in Japan. Compared with C. odorata, infected P. speciosa plants had less mycelial sheath surrounding the exodermis, and the older the roots, especially main roots, the more hyphae penetrated. In addition, a large number of juvenile roots were not associated with hyphae. We concluded that such root endophyte interactions were not events isolated to the interactions between exotic plants and microbes but could occur generally in vitro. Our pure culture system with a somatic plant allowed these fungi to express symbiosis-related phenotypes that varied with the plant host; these traits are innately programmed but suppressed in nature and could be useful in genetic analyses of plant–fungal symbiosis.     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Benefit of Shading by Nurse Plant Does Not Change along a Stress Gradient in a Coastal Dune

The proximity of adult neighbors often increases the performance of woody seedlings under harsh environmental conditions but this nurse plant effect becomes less intense when abiotic stress is alleviated, as predicted by the stress gradient hypothesis (SGH). Although some studies have tested how the net nurse effect is changed by stress, few studies have tested how the mechanism that drives the facilitative effect of nurse responds to changes in stress. We conducted field experiments in a subtropical coastal dune to test if shading drives the known nurse effect of adults of the tree Guapira opposita on seedling performance of another tree species, Ternstroemia brasiliensis. We transplanted T. brasiliensis seedlings to three neighbor environments: under a G. opposita crown, under artificial shade and without neighbor as a control. Furthermore, assuming that proximity to the seashore correlates with stress intensity, we tested if the potential shade-driven facilitation became less intense as stress decreased. Regardless of the proximity to the seashore, after a year, the survival of T. brasiliensis seedlings was twice as high when the seedlings were under G. opposita or under artificial shade compared to the control, indicating that the nurse effect is driven by shade and that this facilitation mechanism is constant along the stress gradient. However, G. opposita and artificial shade had a negative effect on seedlings growth. Overall, our results showed that the facilitation mechanism behind the nurse effect did not wane as the stress was reduced. Furthermore, in spite of the potential costs in terms of biomass production, our study highlights the potential of nurse plants and artificial shade as techniques to improve the survival of transplanted seedlings used in the restoration of degraded shrubland coastal dunes.     

不同支持物对攀援植物--葎草雌雄株光合特性及生物量结构的影响

葎草[Humulus scandens(Lour.)Merr.]为草本雌雄异株攀援植物,采用野生种群人为控制的方法,设置分枝找到乔木支持物(高度(3±0.5)m)、灌木支持物(高度(1±0.5)m)和无支持物3种生长方式,测定雌(♀)、雄(♂)株分枝的叶面积参数、光合参数、叶绿素含量、可溶性糖含量及茎、叶、花在分枝、构件、单株水平下生物量分配比等指标,分析支持物对分枝光合特性和生物量结构的影响,探讨雌、雄分枝利用支持物的性别差异及响应支持物的生态适应差异。结果表明:(1)分枝的单叶面积、总叶面积、叶面积比率、比叶面积对支持物响应存在性别差异,♀株叶面积参数均大于♂株;支持物差异主要影响叶绿素a含量,对叶绿素b、叶绿素a/b影响不显著。(2)支持物差异对分枝叶片的光合参数均有显著的影响(P0.05),支持物对光合参数影响顺序为GsPnTrCi,性别差异影响顺序为GsTrPnCi;可溶性糖含量在性别间、支持物间均表现出极显著的差异(P0.01),♂株糖含量显著高于♀株。(3)在分枝水平下,♀株叶、茎及花生物量分配比都未受到支持物不同的影响(P≥0.05),而♂株在叶、茎的生物量分配比方面受影响显著(P0.05);在构件水平下,支持物差异显著影响了♂株分枝间的营养生长构件分配比,显著影响了♀株分枝间的生殖生长构件分配比;在单株水平下,叶分配比仅在支持物间差异显著(P0.05),而茎、花分配比在性别间、支持物间均有显著差异。(4)雌雄分枝的光合特性和生物量结构对不同支持物的响应差异明显,使葎草单株的生理整合性和适应性大幅度提高。研究以雌雄异株攀援植物为材料,从分枝水平分析支持物对雌雄株光合特性及生物量结构的影响,研究结果对探讨雌雄异株攀援植物的生态适应性具一定的价值。     

光照条件、植株冠层结构和枝条寿命的关系——以桂花和水杉为例

根据衰老理论的代谢率假说,生物寿命与其代谢率有关,个体大小相同的生物体,在质量较好的微生境中通常比较差生境中具有更高的代谢速率。因此,生物体在资源供给较差的生境中通常比资源供给较充足的生境中具有更长的寿命。枝条是木本植物植冠构建的基本单元之一,如果枝条遵循代谢率假说,则可推测在光照较好环境下的植物枝条或小枝将比其在遮荫环境下具有更短的寿命,即枝条寿命与光照条件成反比。以常绿物种桂花(Osmanthus fragrans)和落叶物种水杉(Metasequoiaglyptostroboides)为研究对象,通过测量不同光照环境下,植株大小(株高和胸径)、冠层深度、冠层轮廓(冠层深度/冠层宽度)、相对冠层宽度(冠层宽度/植株高度)以及植株凋落枝条寿命等性状,探讨了光照条件对成年植株冠层形态结构和植株枝条寿命的影响。调查发现:1)枝条的寿命在遮荫条件下显著高于全光照条件下,与理论预测吻合;2)随遮荫程度增加,植株冠层深度和冠层轮廓增加,相对冠层宽度减小;3)枝条的平均寿命与植株冠层深度和冠层轮廓成正比,与植株相对冠层宽度成反比。这表明光照条件可能通过改变植株冠层结构来影响枝条寿命。未来需要进一步研究枝条生物量分配、叶片光合能力和呼吸速率在不同生活型物种之间的差异,以便更全面的理解枝条寿命与生境质量之间的关系。     

Spatial structuring of bacterial communities within individual Ginkgo biloba trees

Plant‐associated microorganisms affect the health of their hosts in diverse ways, yet the distribution of these organisms within individual plants remains poorly understood. To address this knowledge gap, we assessed the spatial variability in bacterial community diversity and composition found on and in aboveground tissues of individual Ginkgo biloba trees. We sampled bacterial communities from > 100 locations per tree, including leaf, branch and trunk samples and used high‐throughput sequencing of the 16S rRNA gene to determine the diversity and composition of these communities. Bacterial community structure differed strongly between bark and leaf samples, with bark samples harbouring much greater bacterial diversity and a community composition distinct from leaves. Within sample types, we observed clear spatial patterns in bacterial diversity and community composition that corresponded to the samples' proximity to the exterior of the tree. The composition of the bacterial communities found on trees is highly variable, but this variability is predictable and dependent on sampling location. Moreover, this work highlights the importance of carefully considering plant spatial structure when characterizing the microbial communities associated with plants and their impacts on plant hosts.     

Biochemical analysis of ‘kerosene tree’ Hymenaea courbaril L. under heat stress

Hymenaea courbaril or jatoba is a tropical tree known for its medically important secondary metabolites production. Considering climate change, the goal of this study was to investigate differential expression of proteins and lipids produced by this tree under heat stress conditions. Total lipid was extracted from heat stressed plant leaves and various sesquiterpenes produced by the tree under heat stress were identified. Gas chromatographic and mass spectrometric analysis were used to study lipid and volatile compounds produced by the plant. Several volatiles, isoprene, 2-methyl butanenitrile, β ocimene and a numbers of sesquiterpenes differentially produced by the plant under heat stress were identified. We propose these compounds were produced by the tree to cope up with heat stress. A protein gel electrophoresis (2-D DIGE) was performed to study differential expression of proteins in heat stressed plants. Several proteins were found to be expressed many folds different in heat stressed plants compared to the control. These proteins included heat shock proteins, histone proteins, oxygen evolving complex, and photosynthetic proteins, which, we believe, played key roles in imparting thermotolerance in Hymenaea tree. To the best of our knowledge, this is the first report of extensive molecular physiological study of Hymenaea trees under heat stress. This work will open avenues of further research on effects of heat stress in Hymenaea and the findings can be applied to understand how global warming can affect physiology of other plants.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.     

Invasive feral pigs impact native tree ferns and woody seedlings in Hawaiian forest

Invasive mammals can fundamentally alter native plant communities, especially on isolated islands where plants evolved without them. The globally invasive feral pig (Sus scrofa) can be particularly destructive to native plant communities. Tree ferns are an important understory component in many forests facilitating the establishment of a variety of species. However, the extent and effects of feral pig damage to tree ferns, and associated impacts on plant community regeneration, are largely unknown. We quantified the effect that feral pig damage has on tree fern growth, survival, and epiphytic woody seedling abundance over 1 year on 438 randomly selected tree ferns of three endemic species (Cibotium chamissoi, Cibotium glaucum, and Cibotium menziesii) in a Hawaiian montane wet forest with high tree fern and feral pig densities. Across all tree fern species, feral pigs damaged 13 % of individuals over 1 year. Compared with undamaged tree ferns, moderately- to heavily-damaged individuals had decreases of 4 to 27 % in trunk length increment and lost tenfold more fronds. Tree fern angle (standing, leaning, prone, or semi-prone) and woody seedling abundance co-varied with feral pig damage. Specifically, damaged tree ferns were more often prone or semi-prone and supported more seedlings, but also had annual mortality up to 34 % higher than undamaged tree ferns. Overall, feral pig damage had substantial negative effects on tree ferns by reducing growth and survival. Given the importance of tree ferns as regeneration sites for a variety of native plants, feral pig damage to tree ferns will likely alter future forest composition and structure. Specifically, feral pig damage to tree ferns reduces potential establishment sites for species that either regenerate preferentially as epiphytes or are currently restricted to epiphytic establishment due to ground rooting by feral pigs.     

Tree species diversity alters plant defense investment in an experimental forest plantation in southern Mexico

How plant species diversity affects traits conferring herbivore resistance (e.g., chemical defenses), as well as the mechanisms underlying such effects, has received little attention. One potential mechanism for the effect of diversity on plant defenses is that increased plant growth at high diversity could lead to reduced investment in defenses via growth–defense trade‐offs. We measured tree growth (diameter at breast height) and collected leaves to quantify total phenolics in 2.5‐year‐old plants of six tropical tree species (N = 597 plants) in a young experimental plantation in southern Mexico. Selected plants were classified as monocultures or as polycultures represented by mixtures of four of the six species examined. Tree species diversity had a significant negative effect on total phenolics, where polycultures exhibited a 13 percent lower mean concentration than monocultures. However, there was marked variation in the effects of diversity on defenses among tree species, with some species exhibiting strong reductions in phenolic levels in mixtures, whereas others were unresponsive. In addition, tree species diversity had no effect on growth, nor was the negative effect of diversity on chemical defenses mediated by a growth–defense trade‐off. These results demonstrate that tree diversity can alter investment in chemical defenses in long‐lived tree species but that such effect may not always be under strong control by plant endogenous resource allocation trade‐offs. Regardless of the underlying mechanism, these findings have important implications for predicting effects on consumers and ecosystem function.     

乌拉山自然保护区白桦种群的年龄结构和点格局分析

白桦群落是乌拉山森林植被的主要类型之一,在高海拔阴坡、半阴坡以纯林形式分布.根据乌拉山自然保护区白桦林不同林龄结构设置3个典型样地,采用种群径级结构代替年龄结构、点格局分析(Ripley's K-Function)方法探讨了乌拉山白桦种群年龄结构、空间分布规律和种群动态.结果表明:(1)乌拉山自然保护区白桦种群径级结构呈典型的“金字塔”型,种群自然更新良好,属增长型种群;(2)由于种内不同个体间为争夺空间和资源,种群在第Ⅲ、Ⅳ径级死亡率较高,自疏作用明显;(3)白桦种群的存活曲线接近于Deevey Ⅰ型曲线;(4)在研究尺度内白桦种群以幼树、中龄树为主时呈聚集分布,而成龄树或老龄树占多数时呈随机分布,即随着种群年龄的增加,其分布格局逐渐由集群分布向随机分布转变.乌拉山白桦种群在小于1.5m的尺度呈聚集分布,即具有2株以上个体“丛生”现象.在环境条件相似的情况下,白桦种群自身的生物、生态学特性是影响其分布格局的最主要因素.     

Branching morphology as an indicator of environmental disturbance: Testing the vegetative fragmentation of Acanthophora spicifera and the turf morphology of Laurencia papillosa

A new method for interpreting branching morphology is introduced and demonstrated to be a practical means of assessing ecological events. The “turf” configuration of Laurencia papillosa (Forssk.) Grev. and the vegetative fragmentation of Acanthophora spicifera (Vahl) Børg. are examined on a fringing reef platform at Galeta Point, Panama. Predictions of branching morphology are tested by comparing plants in the fore-reef, that are exposed to intense wave action and potentially large numbers of herbivorous fish, with those in the back-reef, where wave action and grazing are minimal. Differences in plant size, number of branches and length of branch segments, together with the number and location of missing branches, are quantified. It is shown that in some instances branching morphology can be used to test ecological events.There is little difference in the branching structure of Laurencia papillosa in the foreand back-reef. The number and the length of branch segments were similar and few branches were lost to wave action or herbivory. Some grazing of determinate branches occurred on the reef flat, but this had no apparent effect upon branching morphology. Plants in the fore-reef, however, were significantly smaller than those in the back-reef. It follows that fore-reef plants are best characterized as dwarf plants as opposed to a “turf”.The branching morphology of Acanthophora spicifera is altered by wave action in the fore-reef. Plants were shorter and had proportionally fewer branches in the fore-reef than in the back-reef. There was, however, no difference in the length of branch segments between the two areas. Numerous branching scars, resulting from the loss of indeterminate branches, were found. In the fore-reef, the smaller-sized plants of A. spicifera are best explained by vegetative fragmentation.     

Drought-induced mortality of a foundation species (<Emphasis Type="Italic">Juniperus monosperma</Emphasis>) promotes positive afterlife effects in understory vegetation

Climate change-induced droughts have contributed to large-scale die-offs of dominant tree species throughout much of the southwestern United States. These mortality events provide ecologists with the opportunity to determine whether afterlife effects associated with the die-off occur and the potential implications for future ecosystem changes. We studied both the afterlife and interaction effects of condition (dead trees, living trees, and open areas) on understory vegetation in a Juniperus monosperma woodland of northern Arizona 7 years after a major mortality event. Five major findings resulted: (1) there was a positive afterlife effect on understory plants, in which vegetation under dead junipers contained almost double the amount of cover; (2) the competitive effect on understory plants was exemplified by a 1.3 times greater cover and 1.6 additional species in open areas compared to under living junipers; (3) plant community composition significantly differed by aspect and condition; (4) the highly invasive cheatgrass (Bromus tectorum) was 1.5 times greater under dead junipers compared to live junipers; and (5) litter depth and light availability were negatively and positively correlated with plant cover, respectively, but weakly correlated with afterlife effects. Our results indicate that mortality events can promote changes in understory vegetation through afterlife effects. In ecosystems where foundation species suffer high rates of mortality, changes in plant population dynamics and ecosystem function may promote an altered trajectory in community composition with the potential to increase the presence of invasive species. Continued species die-offs associated with climate change-induced drought may contribute to an increased occurrence and legacy of afterlife effects.     

Requirements for plant coexistence through pollination niche partitioning

Plant–pollinator interactions are often thought to have been a decisive factor in the diversification of flowering plants, but to be of little or no importance for the maintenance of existing plant diversity. In a recent opinion paper, Pauw (2013 Trends Ecol. Evol. 28, 30–37. (doi:10.1016/j.tree.2012.07.019)) challenged this view by proposing a mechanism of diversity maintenance based on pollination niche partitioning. In this article, I investigate under which conditions the mechanism suggested by Pauw can promote plant coexistence, using a mathematical model of plant and pollinator population dynamics. Numerical simulations show that this mechanism is most effective when the costs of searching for flowers are low, pollinator populations are strongly limited by resources other than pollen and nectar, and plant–pollinator interactions are sufficiently specialized. I review the empirical literature on these three requirements, discuss additional factors that may be important for diversity maintenance through pollination niche partitioning, and provide recommendations on how to detect this coexistence mechanism in natural plant communities.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.