Showing posts with label plants. Show all posts
Showing posts with label plants. Show all posts

Thursday, November 14, 2013

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.

Friday, March 2, 2012

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



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.

Wednesday, October 12, 2011

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

Wednesday, September 14, 2011

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.

Friday, April 29, 2011

Ecological interactions and evolutionary relatedness: contrary effects of conserved niches

ResearchBlogging.orgOver 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

Sunday, October 17, 2010

Grassland diversity increases stability across multiple functions

ResearchBlogging.orgAs 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

Thursday, April 8, 2010

Plant rarity: environmental or dispersal limited?

ResearchBlogging.orgIn 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

Friday, March 5, 2010

Competitive coexistence, it's all about individuals.

ResearchBlogging.orgUnderstanding 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

Wednesday, February 3, 2010

The evolution of a symbiont

ResearchBlogging.orgThe 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

Thursday, January 14, 2010

Plant genotypic diversity supports pollinator diversity

ResearchBlogging.orgResearch 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

Tuesday, January 5, 2010

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.

ResearchBlogging.orgIn 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

Wednesday, November 25, 2009

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

ResearchBlogging.orgSome 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

Wednesday, October 21, 2009

Adaptation and dispersal = (mal)adapted

ResearchBlogging.orgEver since Darwin, we often think of organisms as being in a constant battle against other organisms and local environments. Thus natural selection and the resulting arms race results in organisms highly adapted to local conditions and against local antagonists. At the same time, and especially driven by theoretical advances in the 1990's, researchers began to ask how dispersal -that is, the flow of genetic material from elsewhere, can disrupt local adaptation. On the one hand it may provide genetic variation allowing for novel solutions to new difficulties. On the other hand, dispersal may reduce the prevalence of fitness-increasing genes within local populations.

In a simple but elegant experiment, Jill Anderson and Monica Geber performed a reciprocal transplant experiment, moving Elliott's Blueberry plants between two habitats. One population was from highland, dryer habitats and the other from moist lowlands. They further evaluated performance in greenhouse conditions. Their results, published in Evolution, show that these two populations have not specialized to local conditions. Rather, due to asymmetric gene transfer, lowland individuals actually performed better when planted in highlands than compared to their home habitat. Further, in the greenhouse trials, lowland species did not perform better under higher moisture conditions. While genetic or physiological constraints may also limit adaptation, Anderson and Geber present a fairly convincing case that gene flow is the culprit.

These results reveal that populations may actually be relatively mal-adapted to local conditions, which has numerous consequences. For example, we need to be cognizant of adaptations to particular conditions when selecting populations for use in habitat restoration and when trying to predict response to altered climatic or land-use conditions. Importantly what does this mean for multi-species coexistence? Dispersal seems to limit the ability to adapt, and thus, better use local resources or maximize fitness, making for a better competitor. At the same time, dispersal can offset high death rates, allowing for the persistence of a population that would otherwise go extinct. Understanding how these two consequences of dispersal shape populations and communities is an interesting question, and work like Anderson and Geber's provides a foundation for future studies.

Anderson, J., & Geber, M. (2009). DEMOGRAPHIC SOURCE-SINK DYNAMICS RESTRICT LOCAL ADAPTATION IN ELLIOTT'S BLUEBERRY (

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Evolution DOI: 10.1111/j.1558-5646.2009.00825.x

Mycorrhizal Networks: Socialists, capitalists or superorganisms?


ResearchBlogging.orgMycorrhizal networks – fungal mycelium that colonize and connect roots of one or more plant species – are a very intriguing type of fungal-plant association. There is evidence of substantial facilitation between plant individuals via these fungal networks. This can have drastic implications for our understanding of nature, given that the common perception is that other mechanisms, like competition, herbivory or dispersal, are the main drivers of plant community associations. This may be far from reality if the existence of “socialist” networks is widespread (e.g. the ability to connect and profit from a network may be more important than competitive abilities). In the last issue of the Journal of Ecology ( that has a very interesting special feature on facilitation in plant communities), Marcel Van der Heijden and Tom Horton conducted a review of the topic. They found a general positive effect of mycorrhizal networks on seedlings and large plants (i.e. plants tend to grow better if they are associated with a network). However, the reviewers also found some networks can have a neutral or even a negative effect on plants. The plant responses were highly variable depending on other variables including fungal species, nutrients availability, and plant identity. The positive effect of some fungal networks on seedlings growing nearby adult trees of its same species is somehow opposite to the predictions of the Janzen-Connell hypothesis. We need further studies to understand the overall importance of mycorrhizal networks in relation to other better understood mechanisms.

van der Heijden, M., & Horton, T. (2009). Socialism in soil? The importance of mycorrhizal fungal networks for facilitation in natural ecosystems Journal of Ecology, 97 (6), 1139-1150 DOI: 10.1111/j.1365-2745.2009.01570.x

Wednesday, October 7, 2009

Exotic plants integrate into plant-pollinator networks

ResearchBlogging.orgAt almost any spot on the globe, there are species present that are not native to that locale, having been transported by human activities. Whether and how exotic species impact communities is a multifaceted problem that requires understanding the multitude of direct and indirect species interactions that occur. In a paper published in the Proceedings of the Royal Society, B, Montserrat Vila and colleagues asked if exotic plants where integrated into plant-pollinator networks, and whether this integration had any observable impacts on these networks. This is an important question, as most ecological theory predicts that plant-pollinator networks are actually quite resilient to perturbations since most associations tend to be between generalists as opposed to the more susceptible specialists.

They studied invaded plant communities across Europe, observing pollinator visits to flowers in multiple 50 x 50 m plots. They calculated connectance as the number of interactions standardized by network size. They showed that exotics fully integrated into plant-pollinator networks. Exotic species accounted for 42% of all pollinator visits and 24% of all network connections -a testament to the overall abundance of exotics in many communities. However, these exotics did not affect overall changes in network connectedness, revealing that these networks are quite robust to invasions.

That said, researchers must now ask if this is true in networks that do contain high numbers of specialists (e.g., orchids) or if the relative few specialists in generalist-dominated systems are more susceptible to changes from exotics.

Vila, M., Bartomeus, I., Dietzsch, A., Petanidou, T., Steffan-Dewenter, I., Stout, J., & Tscheulin, T. (2009). Invasive plant integration into native plant-pollinator networks across Europe Proceedings of the Royal Society B: Biological Sciences, 276 (1674), 3887-3893 DOI: 10.1098/rspb.2009.1076

Monday, September 21, 2009

Everything but extinct: invasion impacts on native diversity

ResearchBlogging.orgThere has been a persistent debate in the plant invasions literature about whether exotic plant invasions are a major threat to native plant persistence. While there are clear examples of animal invasions resulting in large scale extinction -e.g., the brown tree snake or Nile perch, evidence has been ambiguous for plants. Most ecologists are not so sanguine as to actually conclude that plant invasions are not a threat, and I think most believe that plant invader effects are an issue of temporal and spatial scale and that the worst is yet to come.

In a forthcoming paper from Heinke Jäger and colleagues in the Journal of Ecology, Cinchona pubescens invasions on the Galápagos Islands were monitored in long-term plots for more than seven years. What they found was that there was a four-fold increase in Cinchona density as the invasion progressed and that this increase had measurable effects on native species abundance. While they did not observe any native extirpations in their plots, native densities decreased by at least 50%. Of the greatest concern was that Island endemics appear to the most susceptible to this invasion.

What these results show is that, while there were not any observed extinctions, there were serious deleterious changes to native diversity. Further, the native species, and especially the endemics, are now more susceptible to other invasions or disturbances due to their lower abundances. The impact of exotic invaders may not be readily apparent but may be a major contributor to increased extinction risk.

Jäger, H., Kowarik, I., & Tye, A. (2009). Destruction without extinction: long-term impacts of an invasive tree species on Galápagos highland vegetation Journal of Ecology DOI: 10.1111/j.1365-2745.2009.01578.x

Wednesday, September 9, 2009

Exploring ecology through GMOs

This year's Tansley Lecture at the BES meeting was a superb presentation given by Ian Baldwin from the Max Planck Institute for Chemical Ecology. He was enjoyable to watch as his folksy, mid-western American style disarmed the listener and leaving them unprepared for his ascorbic wit and, at times, controversial message. Prof. Baldwin* is a chemical ecologist who studies plant biochemical and physiological processes and their interaction with herbivores. Through his use of molecular tools and superb natural history, he has gained new insights into how and why plants respond to herbivory. He has discovered the pathways allowing wild tobacco, Nicotiana attenuata, to detect chemicals in tobacco hornworm spit and the resulting chemical defense response. More than this though, part of his talk was about the use of transformed plants to study this plant defense response. Using genetic tools, his group was able to knockout certain segments of these biochemical pathways in order to determine how various chemicals affected hornworms. He showed chemical responses involved signaling hornworm predators whereas other responses directly targeted wornworm's ability to digest plant material.

I think that ecologists are often wary of GMOs and his talk was a convincing case for their use in basic research, and he advocated for a more reasoned approach to their use.




*Note: He has run into trouble with German authorities over using the title 'Dr.' -see here.

Tuesday, August 25, 2009

March of the polyploids!

ResearchBlogging.orgSpeciation by polyploidy (see here for a general description of polyploidy) is one of the mechanisms of speciation and evolutionary diversification. We all learn about it in Bio 101, right after allopatry and sympatry. It is thought to be an especially important driver of speciation in plants, and anecdotal evidence, such as the origination of the invasive polyploid, Spartina anglica in the UK in the 1800's, reinforced that view. But how important has been unanswered until now.

In a new publication in PNAS by Wood et al. -from the Loren Rieseberg lab (one of the best lab homepages BTW) this questions has been answered. The authors go through all available chromosome counts on the Missouri Botanical Garden's Index to Plant Chromosome Numbers, and assess the proportion of polyploid species. They find that about 15% of all angiosperm speciation events coincided with an increase in chromosome number (and about 30% of fern species). Further, about 35% of all genera contain polyploids. Looking across the phylogeny of major plant groups, they find that all major lineages, except Gymnosperms, have significant proportions of polyploids (again with ferns have the greatest proportion). Polyploidy is a ubiquitous feature of plant diversity and a major driver of plant speciation. And now we can quantify just how important.

Wood, T., Takebayashi, N., Barker, M., Mayrose, I., Greenspoon, P., & Rieseberg, L. (2009). The frequency of polyploid speciation in vascular plants Proceedings of the National Academy of Sciences, 106 (33), 13875-13879 DOI: 10.1073/pnas.0811575106

Tuesday, August 18, 2009

Unifying invader success and impact

ResearchBlogging.orgSomething that has continuously bothered me about our collective narrative concerning invasions has been the conflicting processes determining invader success and impact. Numerous studies (including some of my own) show that invaders are successful often because they are different from residents. That is, they are thought to occupy some unique niche. However, occupying a unique niche means that competition is minimized and these successful invaders should have relatively low impact on residents. Conversely, species that have large impacts are thought to be superior competitors, but why are they able to be so successful?

In a new paper in the Journal of Ecology, Andrew MacDougall, Benjamin Gilbert and Jonathan Levin use Peter Chesson's framework where ability for two species to coexistence (or conversely the strength of competitive exclusion) is a process relative to two factors -the magnitude of fitness differences and the degree of resource use overlap. Here competitive exclusion is rapid if species have a large fitness difference and high resource overlap, and slow if fitness differences are low. Species that are successful because of reduced resource overlap likely have little impact unless there are large fitness inequalities.

If we then view the invasions process on a continuum (see figure), then by determining basic fitness and resource use, we can predict success and impact. This is an exciting development and I hope it inspires a new generation of experiments.

MacDougall, A., Gilbert, B., & Levine, J. (2009). Plant invasions and the niche Journal of Ecology, 97 (4), 609-615 DOI: 10.1111/j.1365-2745.2009.01514.x

Friday, May 1, 2009

Enrichment and diversity loss: a mechanism tested

ResearchBlogging.org
To paraphrase Thomas Henry Huxley: How stupid of us not to have thought of that!

In what has to be one of the most elegant and simple experiments I've seen in a long time, Yann Hautier, Pascal Niklaus and Andy Hector tested a basic mechanism of why nutrient enrichment results in species loss. This is a critically important issue as it has been repeatedly shown that while adding nitrogen to plant communities causes increases in productivity, species go locally extinct. We may bare witness to local diversity declines because human activity has greatly increased nutrient deposition. This pattern has been observed for a couple of decades, but the exact mechanism has never been adequately tested, with some camps believing that enrichment increases below-ground competition for other resources that become limiting, or above ground for light.

As reveled in the most recent issue of Science, Hautier et al. performed an exceedingly simple experiment; they added light to the understory of plant communities with or without nitrogen additions. They made two compelling observations. First, when communities were enriched without elevated light, they lost about 3 of the 6 initial species compared to the control, while light addition in the enriched communities maintained the 6 member community (as did a light only treatment). The second result was that the light plus nitrogen treatment obtained much higher biomass than either the nitrogen or light only treatments, and in fact the light only treatment did not significantly increase productivity, meaning that the communities are not normally light-limited. Further, they failed to detect any elevated belowground competition for other resources.

These results reveal that nutrient enrichment causes diversity loss because increased plant size increases light competition and plants that grow taller with elevated nitrogen are better light competitors. An old problem solved with the right experiment.

Hautier, Y., Niklaus, P., & Hector, A. (2009). Competition for Light Causes Plant Biodiversity Loss After Eutrophication Science, 324 (5927), 636-638 DOI: 10.1126/science.1169640