Southern Ontario’s Ecoregions in Slow Motion: An Eight-Year Journey Along the Bruce Trail

Guest post by Daniel Stuart, MEnvSc Candidate in the Department of Physical & Environmental Science at the University of Toronto-Scarborough

During the final year of my undergraduate program the idea of hiking all 900-or-so kilometres of the Bruce Trail somehow lodged itself in my head. It was 2010 and I was twenty-one years old, immersed in the idealism of that age and on the doorstep of a career as an ecologist. At the time hiking from Queenston Heights along the Niagara River to the town of Tobermory at the northern tip of the Bruce Peninsula (Figure 1.) seemed an appropriate way to gain a more meaningful appreciation of my home province’s landscape. This would turn out to be true in part, but little did I know that the more valuable takeaway would be a practical education in the transitional ecosystems that define Southern Ontario’s landscape. For those without the time to hike it themselves, take a tour with me along the trail from south to north exploring its subtle but undeniable ecological shifts.

Figure 1: Bruce Trail Map (Bruce Trail Conservancy, 2020

As life sometimes goes, it was another two years before I finally purchased the Bruce Trail Reference guidebook and embarked on my first sojourn, a three day hike that would take me from the southern terminus of the trail at Queenston Heights back to Hamilton where I lived at the time. I hopped on a free shuttle bus heading for a casino in Niagara Falls and upon arriving was accosted by the bus driver when he spotted my backpack and water jug, realizing I had no intention of gambling that day.  It was September 2, 2012 and the first miles of the trail were peppered with sightings of uncommon shrubs and trees like Bladdernut (Staphylea trifolia), Sassafras (Sassafras albidum), Spicebush (Lindera benzoin), Pignut Hickory (Carya glabra), and Hill’s Oak (Quercus ellipsoidalis), many of which display full fruit in the late summer. These shrubs and trees share a common trait: in Canada they are confined to the Carolinan Ecoregion.

The Carolinan Ecoregion (defined as Ecoregion 7E in Ontario; Figure 2.) occupies the southernmost portions of Ontario, extending from the shores of Lake Erie to approximately Grand Bend in the west, London, Hamilton, and Toronto in the east. Named for the forests typical of the Atlantic Coast from Long Island to Georgia, this region is dominated by a large variety of deciduous (or, leafy) trees including those listed above that fail to thrive in cooler climates to the north or west (Colthurst & Waldron, 1993). In the Niagara Region the sheltering cliffs and slopes of the Niagara Escarpment offer a slightly warmer microclimate that encourages the region to “punch above its weight” in terms of plant diversity.

Figure 2: Ecoregions of Ontario (Crins et al., 2009)

My first journey from the Niagara River ended in utter failure when with painfully blistered soles, just 26 kilometres into my expedition I swallowed my pride and called a friend to pick me up at the Brock University campus in St. Catharines. I would eventually work up to 30- and even 40-kilometre days, but this would take years of training and a good deal of re-conditioning every spring to tighten up my legs that would seemingly turn to jelly each winter.

The “southern feel” of the Bruce Trail gradually diminishes as one hikes westward toward Hamilton, the conspicuously common open-grown oaks (Quercus spp.) gently replaced by the familiar Sugar Maple (Acer saccharum)-dominant woodlands that emblemize Canada. The extensive forested tracts of the Dundas Valley offer the final display of southern species before mounting the escarpment where suddenly one stands firmly in the Great Lakes-St. Lawrence Ecoregion (defined as Ecoregion 6E in Ontario; Figure 2.).  The abruptness of the transition surprised me. I recall spotting the northernmost stand of a southern tree, a population of Chinquapin Oak (Quercus muehlenbergii) perched below the escarpment brow next to Sydenham Road in Dundas. Although I understand that southern species are occasionally found north of the official boundaries of the Carolinian Ecoregion, along the Bruce Trail I encountered no other Carolinian-specialist plant. The sheltered valleys of the Hamilton area seem to provide a last bastion for southern plants that struggle to tolerate the exposed landscape above Burlington and beyond.

From the Burlington heights the Great Lakes-St. Lawrence forest extends northward all the way to the edge of the Canadian Shield, which itself transitions into the seemingly endless Boreal forest that blankets the northern part of our continent. Unlike the Carolinian region which comprises mostly deciduous trees, or the Boreal region which compromises mostly coniferous trees, the Great Lakes-St. Lawrence forest is a roughly equal mix of the two. This forest type features strong representation from leafy trees like Sugar Maple (Acer saccharum), American Beech (Fagus americana), and Black Cherry (Prunus serotina) along with their needled counterparts like Eastern White Pine (Pinus strobus), Eastern White Cedar (Thuja occidentalis), and Eastern Hemlock (Tsuga canadensis).

I hiked the central stretches of the Bruce Trail at a slower rate between 2014 and 2018, a section that traverses a hilly complex of woodlots, river valleys, and bucolic landscapes. I came across a Striped Maple (Acer pensylvanica) in the Caledon area and a small Jack Pine (Pinus banksiana) stand on a north-facing slope near the Hockley Valley, both typically northern trees. My first Northern Holly Fern (Polystichum lonchitis) was observed in Noisy River Provincial Park near the village of Creemore, a plant that in places coated the trailside by the time I reached Owen Sound. Similarly, I spotted a tiny American Hart’s Tongue Fern (Asplenium scolopendrium var. americanum) in the Beaver Valley, a globally uncommon species whose core range is concentrated around Owen Sound and the lower reaches of the Bruce Peninsula.

By May of 2019 I was hiking in earnest, setting aside many weekends to cover the approximately 210 kilometres from the west edge of the Beaver Valley near Kimberley, through Owen Sound and to the base of the Bruce Peninsula near Wiarton. The birding that spring was glorious, and I often hiked with binoculars somewhat annoyingly tugging against my neck. In the Beaver Valley I observed my first ever Louisiana Waterthrush (Parkesia motacilla) along the rushing banks of Bill’s Creek. A Philadelphia Vireo (Vireo philadelphicus) flitted between branches in a woodlot near Walter’s Falls, a Golden-winged Warbler (Vermivora chrysoptera) was spotted within a thicket at the Bighead River Overnight Rest Area, and a Green Heron (Butorides virescens) squawked at me near the Bognor Marsh.

In early September 2019 I began the big push up the Bruce Peninsula toward Tobermory, in a four-day period that would take me from the town of Wiarton to Crane Lake Road just before the southeast boundary of Bruce Peninsula National Park. Logistics were more complicated now and I was forced to consider packing lightweight provisions that were adequate but could still be carried on my back. There were also safety considerations specific to the Bruce Peninsula, like establishing a check-in system where cell reception was poor, and to keep aware of Black Bear (Ursus americanus) and the docile but not entirely unthreatening Massasauga (Sistrurus catenatus), Ontario’s only venomous snake. Bear scat was an intermittent sight along the length of the peninsula, first observed just 14 kilometres past Wiarton along Malcolm Bluff.

Although forests remained of mixed composition typical of the Great Lakes-St. Lawrence region, cool northern exposures and thin-soiled areas took on a palpable “northern feel”, often dense with Eastern White Cedar (Thuja occidentalis), pine (Pinus spp.) and Eastern Hemlock (Tsuga canadensis). Wind-beaten crags offered habitat for abundant Bearberry (Arctostaphylos uva-ursi), a northern species yet unseen on my journey so far, and Rattlesnake Plantains (Goodyera spp.) became commonplace. By the time I reached the edge of the National Park the Boreal woods felt much closer.

Sadly, poor weather and low spirits cut my hike short in September, with soggy feet and an approaching storm promising to result in a miserable finale. Despite this setback my goal to finish the Bruce Trail remains firm. At this moment I have booked a campsite in the National Park this May 2020 and (barring any disasters) myself, along with three companions, will finish the final 40 kilometres toward the trail’s northern terminus.

To walk the Bruce Trail is to walk a cross-section of Southern Ontario. For me it has offered an education in landscape ecology earned by traversing it first-hand. It has been a limit-testing and a character-building experience. Although I now hike with a different outlook than my 21-year-old self, I must credit him with having the guts to recognize the journey’s value and for accepting its challenge.

References

Bruce Trail Conservancy. 2020. Explore the Trail. Bruce Trail Conservancy. <https://brucetrail.org//trail-sections>. Retrieved 13 February 2020.

Colthurst, K., Waldron, G. 2013. “What is a Carolinian Forest?”. Essex Region Conservation Authority. Carolinian Canada. <https://caroliniancanada.ca/legacy/SpeciesHabitats_Forests.htm>. Retrieved 13 February 2020.

Crins, William J., Paul A. Gray, Peter W.C. Uhlig, and Monique C. Wester. 2009. The Ecosystems of Ontario, Part I: Ecozones and Ecoregions. Ontario Ministry of Natural Resources, Peterborough Ontario, Inventory, Monitoring and Assessment, SIB TER IMA TR- 01, 71pp.

How many traits make a plant? How dimensionality simplifies plant community ecology.

Daniel C. Laughlin. 2013. The intrinsic dimensionality of plant traits and its relevance to community assembly. Journal of Ecology. Accepted manuscript online: 4 NOV. DOI: 10.1111/1365-2745.12187

Community ecology is difficult in part because it is so multi-dimensional: communities include possibly hundreds of species present, and in addition the niches of each of those species are multi-dimensional. Functional or trait-based approaches to ecology in particular have been presented as a solution to this problem, since fewer traits (compared to the number of species) may be needed to capture or predict a community’s dynamics. But even functional ecology is multi-dimensional, and many traits are necessary to truly understand a given species or community. The question, when measuring traits to delineate a community is: how many traits are necessary to capture species’ responses to their biotic and/or abiotic environment? Too few and you limit your understanding, too many and your workload becomes unfeasible.

Plant communities in particular have been approached using a functional framework (they don't move, so trait measurements aren't so difficult), but the number and types of traits that are usually measured vary from study to study. Plant ecologists have defined functional groups for plants which are ecologically similar, identified particular (“functional”) traits as being important, including SLA, seed mass, or height, or taken a "more is more" approach to measurements. There are even approaches that capture several dimensions by identifying important axes (leaf-height-seed strategy, etc.). Which of these approaches is best is not clear. In a new review, Daniel Laughlin rather ambitiously attempts to answer how many (and which) traits plant ecologists should consider. He asks whether the multi-dimensional nature of ecological systems is a curse (there is too much complexity for us to ever capture), or a blessing (is there a limit on how much complexity actually matters for understanding these systems)? Can dimensionality help plant ecologists determine the number of traits they need to measure? 

From Laughlin 2013. The various trait axes (related to plant organs) important for plant function.

Laughlin suggests that an optimal approach to dimensionality should consider each plant organ (root, leaves, height, figure above). Many of the traits regularly measured are correlated (for example, specific leaf area, leaf dry matter content, lifespan, mass-based maximum rate of photosynthesis, dark respiration rates, leaf nitrogen concentration, leaf phosphorus concentration are all interrelated), and so potentially redundant sources of information. However, there are measurements in the same organ that may provide additional information – leaf surface area provides different information than measures of the leaf economic spectrum – and so the solution is not simply measuring fewer traits per organ. Despite redundancy in the traits plant ecologists measure, it is important to recognize that dimensionality is very high in plant communities. Statistical methods are useful for reducing dimensionality (for example, principle coordinate analysis), but even when applied, Laughlin implies that authors often over-reduce trait data by retaining to only a few axes of information.

Using 3 very large plant species-trait datasets (with 16-67(!) trait measures), Laughlin applies a variety of statistical methods to explore effective dimensionality reduction. He then estimates the intrinsic dimensionality (i.e. the number of dimensions necessary to capture the majority of the information in community structure) for the three datasets (figure below). The results were surprisingly consistent for each data set – even when 67 possible plant traits were available, the median intrinsic number of dimensions was only 4-6. While this is a reasonably low number, it's worth noting that the number of dimensions analyzed in the original papers using those datasets were too low (2-3 only).

From Laughlin 2013. The intrinsic number of traits/dimensions
necessary to capture variation in community structure.

For Laughlin, this result shows that dimensionality is a blessing, not a curse. After all, it should allow ecologists to limit the number of trait measures they need to make, provided they choose those traits wisely. Once the number of traits measured exceeds 8, there appears to be diminishing returns. The caveat is that the traits that are important to measure might differ between ecosystems – what matters in a desert is different than what matters in a rainforest. As always, knowing your system is incredibly important. Regardless, the review ends on a highly optimistic note – that complexity and multi-dimensionality of plant communities might not limit us as much as we fear. And perhaps less work is necessary for your next experiment.

The niche as a changeable entity: phenotypic plasticity in community ecology

Schiffers, K. et al. 2011. Root plasticity buffers competition among plants: theory meets experimental data. Ecology. 92:3.

Ashton et al. 2010. Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms. Ecology. 91:11.

Nearly all explanations for coexistence in communities focus on differences between species. The scale of these differences may occur over large temporal (e.g. evolutionary history, phylogenetic relationships) or spatial scales (e.g. environmental tolerances), or at the scale of the individual. In plants, interactions at the local scale are given particular attention, including interactions mediated by trait differences between species. At finer scales still, there has been recent focus on differences between individuals of the same species, whether they are driven by genotypic differences (link) or plastic changes in individual phenotypes.

From Ashton et al. 2010

Phenotypic plasticity can be defined as phenotypic differences among individuals of the same genotype that occur in response to an environmental cue. The ability of plant species to alter their usage of resources, for example, has clear relevance to resource partitioning among species, since a given individual could adaptively take advantage of alternate resources in response to their particular competitive environment. In such a case, an individual’s realized niche is a function of phenotypic changes in response to the biotic and abiotic environment and thus physiologically-determined. This is in contrast to the usual approach to species’ niches, where physiological constraints are considered to determine a species’ fundamental niche. Although the plant literature shows clear examples of phenotypic plasticity among plants, including in response to competition (for example, perception of light quality leading to changes in growth form), the topic usually receives only passing mention in the community ecology literature.

The number of papers addressing questions of coexistence and competition through the lens of phenotypic plasticity is slowly rising.

From Schiffer et al. 2011, Lithium uptake is
significantly higher on the non-competitor side

A couple of papers from the last few years provide tantalizing glimpses into the possible contribution of plasticity to coexistence. In Schiffers et al. (2011), the authors use experimental and modeling approaches to test whether root uptake can change in response to the proximity of competitors. In the experimental study, the authors looked at the uptake of lithium (a stable nutrient that will be taken up in the place of potassium) by Bromus hordeaceus. They planted pairs of B. hordeaceus  at varying distances apart and then injected lithium into the soil at different differences from the focal plant. They found that lithium uptake was significantly higher on the non-competitor side of the focal plant than on the competitor side, suggesting that plastic changes in resource uptake occurred in response to competitor proximity. Modelling results from the same study suggest that plasticity may allow individuals minimize competitive pressure by making changes in belowground architecture, thereby using available space more efficiently.

Ashton et al. (2010) take a similar approach, looking at how the uptake of nutrients (in this case three forms of nitrogen (N)) varies among species depending on their competitive environment. They explored the ways in which plasticity could lead to changes in the realized niche. In particular, they explored two hypotheses: that plants would exhibit niche preemption, where the inferior competitor switched to a different form of nitrogen in the presence of the superior competitor; or dominant plasticity, where plasticity actually enhances competitive ability.  The authors looked at 4 species, 3 common and 1 rare(r), in an alpine tundra community, isolating naturally occurring pairs of each combination of species. These ‘competitive arenas’ were isolated, and other species within the arena were removed. After a year, the authors added N¹⁵ tracers to each arena, in three forms (NH4+, NO3-, and glycine): these tracers would allow them to track the N once it was incorporated into the plant tissue. The plants were then harvested and the amount of each type of nitrogen in each was measured. Plant biomass was also recorded, and used to estimate the ‘competitive response’ (basically the ratio of biomass when grown with a competitor compared biomass to when grown solo). Their findings were rather neat: the 3 common plants experienced no negative effect on biomass from growing in competition with the rare plant, but the rare plant had much lower biomass when grown in the presence of any of the common plants. Further, while the common plants showed changes in the form of N they used when growing with the rare plant, the rare plant did not switch its N preference. The rare plant’s lack of plasticity in response to competition may relate to its lower biomass when grown with superior competitors, and ultimately its lower abundance.

Although limited, these studies hint at the role that phenotypic plasticity could play in interspecific interactions. Unfortunately plasticity may be difficult to measure in many contexts, particularly since variation within a species can be attributed to genetic differences or phenotypic plasticity, and these factors must be teased apart. Further, there is an issue of differentiating the effects of resource limitations from ‘adaptive’ plastic changes in growth. While plants are relatively tractable for these types of studies (they’re sessile, they use limited abiotic resources), other organisms are less explored for a reason.

What these studies can’t address is the question of ‘how important is phenotypic plasticity, really’? Reviews of coexistence mechanisms list numerous possible ways by which coexistence is facilitated among species. For plants especially, the limited number of resources required for survival has lead to great consideration of the possible niche axes over which species can differentiate themselves. Phenotypic plasticity's contribution to coexistence may be that it provides another way by which plants can partition resources at very fine scales. And if nothing else, such results provide further evidence that variation within species may be an important component of coexistence.

Thanks to Kelly Carscadden for discussions on the topic.

Seed dispersal: plant height seems to be more important than seed size!

I really like papers that teach me something that I didn’t know. But, I love papers that show me that what I learned is wrong. This is the case of a new paper by Fiona Thomson, Angela Moles, Tony Auld, and Richard Kingsford on seed dispersal that appears in the last issue of the Journal of Ecology. This group from Australia analyzed the effects of seed size and plant height on their dispersal abilities. They reviewed intensively the literature gathering data on 200 species from 148 studies around the world. Surprisingly to me, they found plant height was much better at predicting seed dispersal than seed size. This might not sound so surprising for many people (and after seeing the paper, kind of intuitive), but there was a lot of evidence that seed size was the best predictor of dispersal, with species with smaller seeds dispersing further than species with bigger seeds. For wind dispersed species, their results are more intuitive, but they found this pattern in a number dispersal syndromes analyzed (i.e. unassisted, wind, ballistic, ingestion, and ant dispersal). So, in your next study on seed dispersal consider adding plant height as an explanatory variable.

Thomson, F. J., A. T. Moles, T. D. Auld, and R. T. Kingsford. 2011. Seed dispersal distance is more strongly correlated with plant height than with seed mass. Journal of Ecology 99:1299-1307. DOI 10.1111/j.1365-2745.2011.01867.x

BES day 2: Plants, plants and way more plants

From Sept 13

I attended the Journal of Ecology Centenary symposium all morning, where the talks were broad overviews of select areas in plant ecology. They were quite good; I really do feel that I was informed about recent research advances.

In the first talk of the morning, Sandra Lavorel gave a tour de force about how plant functional traits scale up to ecosystem services. She recognizes that there are trade offs in services, where one service (say agricultural value) is in direct conflict with a noter service (say species richness). She very cleverly asks whether these services are constrained by ecological trade offs or traits. It is known that functional traits affect ecosystem functions and services, and it is also known that there are strong tradeoffs in plant traits such as explained by the world wide leaf economic spectrum. Where plants have these tradeoffs they affect productivity and litter decomposition. Height for example affects productivity and other trophic levels supported. Abiotic gradients affect traits like height or leaf N, and these traits affect ecosystem function such as biomass or litter. Multiple service such as agronomic value, pollination, cultural value, richness, etc. To understand how traits relate to tradeoffs in services.

Next was Angela Moles who talked about how the study of invasions has progressed and whether there were important future directions. The have been 10,000 studies on invasions over the last 30 years and she recognizes that the fact that species evolve in their new ranges to be a critical future research need. Specifically, she asked: do exotics evolve to be more similar or different tha natives? And, can differences be predicted by environmental differences between home and away range. Most interestingly she brought up the point that if on-going change produce new species, should they then be conserved as natives? She went on to say that broad generalizations about trait differences between natives and exotics have produced largely idiosyncratic results, and so other priorities such take the forefront. She went on to say that impact on natives is actually an understudied problem, which needs to be rectified. Finally, she showed us that there is a generally positive relationship between disturbance and invasion. But invasions are favored when there is a change in disturbance rate, since natives are likely adapted to historical disturbance regime. She showed some relatively weak evidence that change in disturbance better predictor of invader richness and abundance then the amount of disturbance, but more work is needed.

Yadvinder Malhi talked about how productivity and metabolism were related to biomass in tropical forests. He sowed us that a small proportion of primary production is turned into biomass. Thus small changes in various pathways could have large consequences. In exhaustive studies in the tropics, he showed that increases in GPP (gross primary production) occurred with soils nutrient quality, and decreases with elevation, likely because of temperature effects on photosynthesis. He also showed that carbon use efficiency is lower than thought, about 30% of carbon turned to biomass. Further, higher productivity is associated with lower residence times, and he hypothesized that rapid growth leads to earlier senescence or less defences if trade offs exist. Biomass appears to be increasing with climate change, but potentially greater mortality and turnover.

Then James Bullock talked about where we are at with understanding seed dispersal. There has been a long history of not understanding long distance dispersal, LDD. The main empirical approaches have grown rapidly lately: tracking seeds, molecular methods and marking seeds or to track dispersers. But at the same time spread models have appeared and advanced. However, Bullock really supports mechanistic models for wind-dispersed species, and these models seem to really provide insights. He then compared a handful of models for invasive and scarce natives, and did mechanistic modeling with climate change. Changes in future wind speeds may result in even larger changes in spread rates. Only Ailanthus appears to have dispersal rates at or faster than the rate of climate change, most species do not appear to be able to move fast enough (except animal dispersed species). Movement on shoes major dispersal vector.

Hans Jacquemyn was the final speaker, and talked about evolution and habitat fragmentation. He studies calcareous grasslands, forests and heathlands in Belgium that have increasingly become isolated and fragmented. Observed declines in genetic diversity in populations, as they get smaller in size. More recent fragmentations have less loss of genetic diversity compared to older fragments. Reduced seed output in small populations for self-incompatible species, results in reduced population growth rates. To counteract plants can increase floral displays or increase selfing rates, which they observed. Also, he has observed changes in timing of flowering and investing more in nectar.

*Some thoughts about the BES. It is a great Society, and a great meeting. It is relatively small and it is nice to see how many members know each other. For those of us in North America, I think it is a great experience to go to one of these meetings.

**I also participated in the BESdigital workshop on communicating science in a digital era. I will have a post about this tomorrow –I've been without internet connection at both the Sheffield dorm and various airports.

Ecological interactions and evolutionary relatedness: contrary effects of conserved niches

Over the past several years a multitude of papers linking patterns of evolutionary relatedness to community structure and species coexistence. Much of this work has looked at co-occurrence patterns and looked for non-random patterns of relatedness. The key explanations of patterns has been that communities comprised of more distantly-related species is thought to be structured by competitive interactions, excluding close relatives. Alternatively, communities comprised of species that are closely related, are thought to share some key feature that allows them to persist in a particular set of environmental conditions or stress. This whole area of research is completely predicated on close relatives having more similar niche requirements then two distant relatives. This predication is seldom tested.In a recent paper in the Proceedings of the National Academy of Science, Jean Burns and Sharon Strauss examine the ecological similarity among 32 plant species and tested if evolutionary relationships offered insight into these similarities. The ecological aspects they examined were germination and early survival rates as well as interaction strengths among species. To assess how these were influenced by evolutionary relatedness, they planted each species in the presence of one of four other species varying in time since divergence from a common ancestor, creating a gradient of relatedness for each species. They found that germination and early survival decreased with increasing evolutionary distance. This surprising result means that species germinating near close relatives do better early on then if they are near distant relatives. The explanation could be that they share many of their biotic and abiotic requirements, and these conserved traits influence early success.

Conversely, when they examined interaction strengths over a longer period (measured as relative individual biomass with and without a competitor), they found that negative interactions were stronger among close relatives.

These two results reveal how evolutionary history can offer insight into ecological interactions, and that the mutually exclusive models of competitive exclusion versus environmental filtering do not capture the full and subtle influence of conserved ecologies. Evolutionarily conserved traits can explain both correlated environmental responses and competitive interactions.

Burns, J., & Strauss, S. (2011). More closely related species are more ecologically similar in an experimental test Proceedings of the National Academy of Sciences, 108 (13), 5302-5307 DOI: 10.1073/pnas.1013003108

Grassland diversity increases stability across multiple functions

As ecological systems are altered with cascading changes in diversity, the oft-asked question is: does diversity matter for ecosystem function? This question has been tested a multitude of times, with the results often supporting the idea that more diverse assemblages provide greater functioning (such as productivity, nutrient cycling, supporting greater pollinator abundance, etc.). Besides greater functioning, scientists have hypothesized that more diverse systems are inherently more stable. That is, the functions communities provide remain more constant over time compared with less diverse systems, which may be less reliable.

While the relationship between diversity and stability has been tested for some functions, Proulx and colleagues examined the stability of 42 variables over 7 years across 82 experimental plots planted with either 1, 2, 4, 8, 16 or 60 plant species in Jena, Germany. They examined patterns of variation (and covariation) in the functions and found that many show lower variation over time in plots with more plant species. Greater stability was found at many different trophic levels including plant biomass production, the abundance and diversity of invertebrates and the abundance of parasitic wasps -which indicate more complex food webs. They also found greater stability in gas flux, such as carbon dioxide. Despite the greater stability in these measures of above-ground functions, below ground processes, such as earthworm abundance and soil nutrients, were not less variable in high diversity plots.

How ecosystems function is of great concern; these results show that more diverse plant communities function more stably and reliably than less diverse ones. The next step for this type of research should be to address what kind of diversity matters. A greater number of species means more different kinds of species, with differing traits and functions. What aspect of such functional differences determine stability of ecosystem function?

This is an exciting paper that continues to highlight the need to understand how community diversity drives ecosystem function.

Proulx, R., Wirth, C., Voigt, W., Weigelt, A., Roscher, C., Attinger, S., Baade, J., Barnard, R., Buchmann, N., Buscot, F., Eisenhauer, N., Fischer, M., Gleixner, G., Halle, S., Hildebrandt, A., Kowalski, E., Kuu, A., Lange, M., Milcu, A., Niklaus, P., Oelmann, Y., Rosenkranz, S., Sabais, A., Scherber, C., Scherer-Lorenzen, M., Scheu, S., Schulze, E., Schumacher, J., Schwichtenberg, G., Soussana, J., Temperton, V., Weisser, W., Wilcke, W., & Schmid, B. (2010). Diversity Promotes Temporal Stability across Levels of Ecosystem Organization in Experimental Grasslands PLoS ONE, 5 (10) DOI: 10.1371/journal.pone.0013382

Plant rarity: environmental or dispersal limited?

In order to promote the persistence and possible spread of extremely rare plant species, ecologists need to know why a species is rare in the first place. In 1986, Deborah Rabinowitz identified seven forms of rarity, where rarity could mean several things depending on range size, habitat specificity and population sizes. When considering rarity, it often feels intuitive to look for environmental causes for these different forms of rarity. Habitat alteration is an obvious environmental change that affects abundance and distribution, but are rare species generally limited by habitat or resource availability? The alternative cause of rarity could just be that sufficient habitat exists, but that the rare species is simply unable to find or disperse to other sites. An extreme example of this would be the Devil's Hole pupfish which exists at only a single pool. It can survive elsewhere (such as in artificial tanks) but natural dispersal is impossible as its pool is in a desert.

Photo taken by Kristian Peters and available through GNU free documentation license

In a recent paper by Birgit Seifert and Markus Fischer in Biological Conservation, they examine whether an endangered plant, Armeria maritima subsp. elongata, was limited because of a lack of habitats or if it was dispersal limited. They collected seeds from eight populations and experimentally added these seeds to their original populations and to uninhabited, but apparently appropriate sites. They found that seeds germinated equally well in inhabited and uninhabited sites and seedlings had similar survivorships. They found that variation in germination rates were likely caused by originating population size and that low genetic diversity and inbreeding reduce viability.

These results reinforce two things. First is that conserving species may only require specific activities, such as collect and distributing seeds. Here ideas like assisted migration seem like valuable conservation strategies. Secondly, we really need to be doing these simple experiments to better understand why species are rare. If we fail to understand the causes of rarity, we may be wasting valuable resources when try to protect rare species.

Seifert, B., & Fischer, M. (2010). Experimental establishment of a declining dry-grassland flagship species in relation to seed origin and target environment Biological Conservation DOI: 10.1016/j.biocon.2010.02.028

Competitive coexistence, it's all about individuals.

Understanding how species coexist has been the raison d'etre for many ecologists over the past 100 years. The quest to understand and explain why so many species coexist together has really been a journey of shifting narratives. The major road stops on this journey have included searching for niche differences among species -from single resources to multidimensional niches, elevating the role for non-equilibrial dynamics -namely disturbances, and assessing the possibility that species actually differ little and diversity patterns follow neutral process. Along this entire journey, researchers (especially theoreticians) have reminded the larger community that that coexistence is a product of the balance between interactions among species (interspecific) and interactions among individuals within species (intraspecific). Despite this occasional reminder, ecologists have largely searched for mechanisms dictating the strength of interspecific interactions.

Image used under Flickr creative commons license, taken by Tinken

In order for two species to coexist, intraspecific competition must be stronger than interspecific -so sayeth classic models of competition. While people have consistently looked for niche differences that reduce interspecific competition, no one has really assessed the strength of intraspecific competition. Until now that is. In a recent paper in Science, Jim Clark examines intra- vs interspecific interactions from data following individual tree performances, across multiple species, for up to 18 years. This data set included annual growth and reproduction, resulting in 226,000 observations across 22,000 trees in 33 species!

His question was actually quite simple -what is the strength of intraspecific interactions relative to interspecific ones? There are two alternatives. First, that intraspecific competition is higher, meaning that among species differences only need to be small for coexistence to occur; or secondly, that intraspecific competition is lower, requiring greater species niche differences for coexistence. To answer this he looked at correlations in growth and fecundity between individuals either belonging to the same or different species, living in proximity to one another. He took a strong positive correlation as evidence for strong competition and a negative or weak correlation as evidence for resource or temporal niche partitioning. What he found was that individuals within species were much more likely to show correlated responses to fluctuating environments, than individuals among species.

This paper represents persuasive evidence that within-species competition is generally extremely high, meaning that to satisfy the inequality leading to coexistence: intra > inter, subtle niche differences can be sufficient. These findings should spur a new era of theoretical predictions and empirical tests as our collective journey to understanding coexistence continues.

Clark, J. (2010). Individuals and the Variation Needed for High Species Diversity in Forest Trees Science, 327 (5969), 1129-1132 DOI: 10.1126/science.1183506

The evolution of a symbiont

The evolution of negative interactions seems like a logical consequence of natural selection. Organisms compete for resources or view one another as a resource, thus finding ways to more efficiently find and consume prey. However, to me, the natural selection of symbiotic or mutualistic interactions has never seemed as straight forward (expect maybe the case where one species provides protection for the other, such as in ant-plant mutualisms). A specific example is the rise of nitrogen-fixing plants, who supply nutrients to bacteria called rhizobia capable of converting atmospheric nitrogen into forms, such as ammonia, usable to the plant host. Not only has this symbiosis evolved, but has seemed to evolve in very evolutionarily distinct lineages. The question is, what are the mechanisms allowing for this?

In a recent paper, Marchetti and colleagues answer part of the question. They experimentally manipulate a pathogenic bacteria and observe it turning into a symbiont. They transferred a plasmid from the symbiotic nitrogen fixing Cupriavidus taiwanensis into Ralstonia solanacearum and infected Mimosa roots with it. Plasmid transfer among distinct bacteria species is common and referred to horizontal genetic transfer (as opposed to vertical, which is the transfer to daughter cells). The presence of the plasmid caused R. solanacearum to quickly evolve into a root-nodulating symbiont. Two regulatory genes lost function, and this caused R. solanacearum to form nodules and to impregnate Mimosa root cells.

This extremely novel experiment reveals how horizontal gene transfer can supply the impetus for rapid evolution from being a pathogen to a symbiont. More importantly it reveals that sometimes just a few steps are required for this transition and how distantly-related bacterial species can acquire symbiotic behaviors.

Marchetti, M., Capela, D., Glew, M., Cruveiller, S., Chane-Woon-Ming, B., Gris, C., Timmers, T., Poinsot, V., Gilbert, L., Heeb, P., Médigue, C., Batut, J., & Masson-Boivin, C. (2010). Experimental Evolution of a Plant Pathogen into a Legume Symbiont PLoS Biology, 8 (1) DOI: 10.1371/journal.pbio.1000280

Plant genotypic diversity supports pollinator diversity

Research over the past 20 years has shown that plant communities with greater diversity maintain higher productivity, greater stability and support more diverse arthropod assemblages. More recently, several experiments have shown that interspecific diversity (namely genotypic differences) also affects community functioning. Pollination is often considered an essential function, and does plant genotypic diversity affect pollinator diversity and frequency?

In a recent paper in PLoS ONE, Genung and colleagues test whether plant genotypic diversity affects pollinator visits. They use an experimental system set-up by Greg Crutsinger that combines multiple genotypes of the goldenrod, Solidago altissima, and record pollinator visits over two years. Experimental plots contained 1, 3, 6, or 12 genotypes of S. altissima. After accounting for differences in abundance, Genung et al. show that as genotypic diversity increases, both pollinator richness and number of visits to the plot significantly increase. This increase is greater than expectations of randomly simulated assemblages combining proportional pollinator visits from monocultures.

The previous research at the species-level has made a persuasive rationale to protect species diversity in order to maintain ecosystem functioning. Now, research like this is making a case that there are consequences for not explicitly considering genetic diversity in conservation planning and habitat restoration.

Genung, M., Lessard, J., Brown, C., Bunn, W., Cregger, M., Reynolds, W., Felker-Quinn, E., Stevenson, M., Hartley, A., Crutsinger, G., Schweitzer, J., & Bailey, J. (2010). Non-Additive Effects of Genotypic Diversity Increase Floral Abundance and Abundance of Floral Visitors PLoS ONE, 5 (1) DOI: 10.1371/journal.pone.0008711

Predicting invader success requires integrating ecological and land use patterns.

Disclaimer, this was modified from an editorial I wrote for the Journal of Applied Ecology.

In the quest to understand species invasions, we often try to link the abundance and distribution of invaders to underlying ecological processes. For example, oft-studied are the links between exotic diversity and native richness or environmental heterogeneity. Seemingly independently, research into how specific land use or management activities affect invasion dynamics is also fairly common. While both research strategies are of fundamental importance, not often recognized, or at least explicitly studied, is that both ecological patterns and management activities simultaneously affect invasion success. Thus a truly integrative approach to understanding invader success must take into account variation in ecological communities and abiotic resource avalibility as well as land use patterns at multiple spatial scales. Such an approach is necessary if ecologists wish to predict potential invader abundance, spread and impact.

Diez et al. Examine how environmental and management heterogeneity interact to influence patterns of Hieracium pilosella (Asteraceae) inasions in the South Island of New Zealand. The spread of H. Pilosella in New Zealand is threatening native habitats (tussock fields) and the livestock grazing industry. Diez et al. Asked how environmental and management regimes affect H. Pilosella abundance and distribution across six large farms on the South Island. This is an interesting and important question, not just because they are examining how human-caused and ecological variation interact to affect H. Pilosella dynamics, but also because these sources are heterogeneity are realized at different spatial scales.

Diez et al. show that the abundance and distribution of H. Pilosella was significantly affected by the interaction of habitat type (i.e., short vs. tall tussocks) and farm management strategies (i.e., fertilization and grazing rates). At larger scales, H. Pilosella was more abundant in tall tussock habitats and was unaffected by fertilization, while in short tussocks, it was less abundant in fertilized patches. At small scales, H. Pilosella was less likely to be found in short tussocks with high exotic grass cover and high productivity (measured as site soil moisture and solar radiation). Conversely, in tall tussocks, H. Pilosella was more likely to be found on sites with high natural productivity. Diez et al. were able to tease these complex causal mechanism apart by using Bayesian multilevel linear models, for which they included example R code in an online appendix.

While it is a truism in ecology to say that heterogeneity affects ecological patterns, this paper deserves mention because they convincingly show that the spread of noxious exotic plants in a complex landscape, can potentially predicted by understanding the invader success in different habitat types and land management strategies. In their case they show how human activities, which were not designed to affect H. Pilosella, can strongly affect abundance in different habitat types. This type of approach to understanding invader dynamics can potentially arm managers with the ability to use existing land use strategies to predict how and where further invader targeting would be most useful.


Diez, J., Buckley, H., Case, B., Harsch, M., Sciligo, A., Wangen, S., & Duncan, R. (2009). Interacting effects of management and environmental variability at multiple scales on invasive species distributions Journal of Applied Ecology DOI: 10.1111/j.1365-2664.2009.01725.x

Taking below-ground processes seriously: plant coexistence and soil depth

Some of the earliest ecologists, like Eugen Warming and Christen Raunkiaer, were enthralled with the minutia of the differences in plant life forms and how these differences determined where plants lived. They realized that differences in plant growth forms corresponded to how different plants made their way in the world. Since this early era, understanding the mechanisms of plant competition is one of the most widely-studied aspects of ecology. This is such an important aspect of ecology because understanding plant coexistence allows us to understand what controls productivity in the basal trophic level for most terrestrial food webs. There are a plethora of plausible mechanisms for how plants are able to coexist, and most involve above-ground partitioning strategies (such as different leaf shapes) or phenological differences (such as germination or bolting timing). Yet, below-ground interactions among plants as a way to understand competition and coexistence have been making a strong resurgence in the literature lately. This resurgence has been driven by new hypotheses and technologies.In what is probably the best hypothesis test of the role for below-ground niche partitioning, Mathew Dornbush and Brian Wilsey reveal how soil depth can affect coexistence. They seeded 36 tallgrass prairie species into plot that were either shallow, medium or deep soiled, and asked if species richness and diversity were affected after 3 years. They found that species richness significantly increased with increased soil depth, revealing that deeper soils likely had greater niche opportunities for species. Not only did deeper soils harbor greater richness, but compositions were non-random subsets. The species inhabiting shallow soils were a subset of medium soils, and medium a subset of deep. This means that increasing depth opened new niche opportunities, unique from the ones for shallow soils.

This study is the first field-based experiment of soil depth and coexistence, that I know of and the results are compelling. Plant species are segregating below-ground niches, and perhaps we look for other partitioning strategies for species that inhabit the same soil depth.

Dornbush, M., & Wilsey, B. (2009). Experimental manipulation of soil depth alters species richness and co-occurrence in restored tallgrass prairie Journal of Ecology DOI: 10.1111/j.1365-2745.2009.01605.x

Other notable recent papers on below-ground processes:

Bartelheimer, M., Gowing, D., & Silvertown, J. (2009). Explaining hydrological niches: the decisive role of below-ground competition in two closely related species Journal of Ecology DOI: 10.1111/j.1365-2745.2009.01598.x

Cramer, M., van Cauter, A., & Bond, W. (2009). Growth of N-fixing African savanna species is constrained by below-ground competition with grass Journal of Ecology DOI: 10.1111/j.1365-2745.2009.01594.x

Meier, C., Keyserling, K., & Bowman, W. (2009). Fine root inputs to soil reduce growth of a neighbouring plant via distinct mechanisms dependent on root carbon chemistry Journal of Ecology, 97 (5), 941-949 DOI: 10.1111/j.1365-2745.2009.01537.x