Insect herbivory in fragmented forests: it’s complicated

I’m excited to announce a recent paper to come out of the lab, by former PhD student Dorothy Maguire, and with Dr. Elena Bennett. In this work, we studied the amount of insect herbivory in forest patches in southern Quebec: the patches themselves varied by degree of fragmentation (ie, small versus large patches) and by connectivity (ie, isolated patches, or connected to other forest patches). We studied herbivory on sugar maple trees, both in the understory and canopy, and at the edges of the patches. Our research is framed in the context of “ecosystem services” given that leaf damage by insects is a key ecological process in deciduous forests, and can affect the broader services that forest patches provide, from supporting biodiversity through to aesthetic value. Dorothy’s research was part of a larger project about ecosystem services and management in the Montérégie region of Quebec.

 

Maguire_Canopy.JPG

Dorothy Maguire sampling insects in the tree canopy (Photo by Alex Tran)

The work was tremendously demanding, as Dorothy had to select sites, and within each site sample herbivory at multiple locations, including the forest canopy (done with the “single rope technique). Dorothy returned to sites many times over the entire summer to be able to assess trends over time. Herbivory itself was estimated as damage to leaves, so after the field season was completed, thousands of leaves were assessed for damage. The entire process was repeated over two years. Yup: doing a PhD requires a suite of skills in the field and lab, and there is no shortage of mind-numbing work… Dedication is key!

As with most research, we had high hopes that the results would be clear, convincing, and support our initial predictions – we certainly expected that forest fragmentation and isolation in our study landscape would have a strong effect on herbivory – after all, our study forests varied dramatically in size and isolation, and herbivory is a common and important ecological process, and insect herbivores are known (from the literature) to be affected by fragmentation.

 

QuebecLandscape

The landscape of southern Quebec. Lots of agriculture, some patches of forest.

 

However, as with so much of ecological research, the results were not straightforward! “It’s complicated” become part of the message: patterns in herbivory were not consistent across years, and there were interactions between some of the landscape features and location within each patch. For example, canopies showed lower levels of herbivory compared to the understory, but only in isolated patches, and only in one of the study years! We also found that edges had less herbivory in connected patches, but only in the first year of the study. Herbivory also increased as the season progressed, which certainly makes biological sense.

So yes, it’s complicated. At first glance, the results may appear somewhat underwhelming, and the lack of a strong signal could be viewed as disappointing. However, we see it differently: we see it as more evidence that “context matters” a great deal in ecology. It’s important not to generalize about insect herbivory based on sampling a single season, or in only one part of a forest fragment. The story of insect herbivory in forest fragments can only be told if researchers look up to the canopy and out to the edges; the story is incomplete when viewed over a narrow time window. In the broader context of forest management and ecosystem services, we certainly have evidence to support the notion that herbivory is affected by the configuration of the landscape. But, when thinking about spatial scale and ecosystem processes, careful attention to patterns these processes “within” forest patches is certainly required.

We hope this work will inspire others to think a little differently about insect herbivory in forest fragments. Dorothy’s hard work certainly paid off, and although the story is complicated, it’s also immensely informative and interesting, and sheds light on how big landscapes relate to small insects eating sugar maple leaves.

Reference:

Maguire et al. 2016: Within and among patch variability in patterns of insect herbivory across a fragmented forest landscape. PlosOne DOI: 10.1371/journal.pone.0150843

 

Landscape structure, insect herbivory, and ecosystem services

I’m pleased to announce a new publication to come out of the lab, with lead author Dorothy Maguire and co-authored by Elena Bennett and Patrick James. In this work, Dorothy ponders and writes about the broader implications of insect herbivory. More specifically, how insect herbivory is affected by landscape connectivity (i.e., the degree to which habitats are linked to each other), and how plant-feeding insects may relate to ecosystem services (i.e., the values and services that humans get from our natural systems).

Female (l) and male (r) Gypsy moth, caught in the act.

Important insects when, as caterpillars, eat a lot of foliate: Female (l) and male (r) Gypsy moth, caught in the act.

We certainly know that insects can do all kinds of damage to plants in ecosystems, but do insects in more (or less) connected habitats do more damage? To address this question Dorothy scoured the literature and got the relatively unsatisfactory answer of “sometimes”: 49% of the papers suggest increased connectivity relates to more insect herbivory and 28% of the papers show less herbivory in more connected patches. The lack of a clear answer actually makes quite a bit of sense since every context can be quite different, and not all insects are equal. It is hard to generalize since effects in forests will not be the same as in fields, and insects that are out-breaking (i.e., with major population explosions) may be affected differently than non out-breaking species. Dorothy certainly found these contexts were important. The results were important to illustrate how we need to adapt any management options with close attention to both landscape feature and their interaction with the life-history of the herbivore.

The second part of Dorothy’s work delved deeper into the literature to ask about the effects of out-breaking versus non out-breaking herbivore species on a select suite of forest ecosystem services: effects on timber production, aesthetics, soil formation and Carbon sequestration. There were some interesting results of this and again, any particular effect of herbivory on an ecosystem service was highly sensitive to the outbreak status of the herbivore. For example, the aesthetics of a forest can be positively affected by low levels of herbivory since this may help create pleasant conditions for light infiltration to the forest floor. However, an out-breaking species may defoliate a tree more completely, thus reducing the aesthetic value. Another example is that low levels of herbivory may positively affect timber production because trees may show “compensatory” growth after light feeding by an insect. In contrast, timber production will be negatively affected by high levels of defoliation as this may reduce a tree’s ability to grow. Although some of these results may seem rather logical, Dorothy’s work was unique as it showed how the scientific literature supports the connections between a herbivore’s life-history and key ecosystem services.

Screen Shot 2015-06-11 at 6.55.21 AM

Visual representations of the hypothesized relationships between insect herbivory and ecosystem services. Specifically (a) timber production, (b) aesthetic value of forests. Graphs are divided into four sections representing positive and negative effects of herbivory on ES, during non-outbreak (low) vs. outbreak (high) levels of herbivory. Quadrants are coloured differently based on the hypothesized strength of the effect of herbivory on ES: weak (light grey), moderate (dark grey) and strong (black). Proposed relationships are derived from synthesis of the available literature. From Maguire et al.

The last part of the work was focused on building a conceptual framework – a framework that ties together landscape structure, the process of herbivory, and ecosystem services. This is meant to be a road map for any stakeholders with an interest in any or all of those factors. For example, should a forest manager be tasked with understanding how to increase or support a particular ecosystem service, she or he needs also to recognize how that service is tied to important processes such as herbivory, and the related connections to the broader landscape.

Screen Shot 2015-06-11 at 7.05.34 AM

This work is novel and important because it links the well known process of insect herbivory to concepts of ecosystem services and to the discipline of landscape ecology. The marrying of these areas is critically important as we face increasing pressures on our natural systems, and the complexity of the systems can be overwhelming. We hope this work piques more interest in this topic, and that the framework Dorothy provides is useful to all the stakeholders.

Reference:

Maguire, DY, PMA James, CM Buddle & EM Bennett Landscape connectivity and insect herbivory: A framework for understanding tradeoffs among ecosystem services. Global Ecology and Conservation. doi:10.1016/j.gecco.2015.05.006

 

An ode to graduate students

Last week I saw two of my graduate students successfully defend their PhDs. This is wonderful and exciting, and I am delighted that they are both moving on to post-doctoral research positions in other places. I am also saddened by their departures: seeing good students leave the lab creates a vacuum. This has caused me to reflect about the effect graduate students have on their supervisors:

I write, teach, research.

I see classrooms, computers, forests and fields.

I use keyboards, iPads, PowerPoint, and pipettes.

I publish or perish.

LOIs, RFPs, IFs, and h-factors.

Grants, emails, to-do lists and budgets.

Learning?

Always.

Literature and libraries can start the process,

But books and blogs barely break the silence.

It’s the tangible human that makes the difference.

My colleagues, my friends:

You are the Academy.

Do you have the answers?

How to avoid wandering alone in ivory towers?

How to slow the withering on tenured vines?

How to grasp frail tendrils of discovery?

How to find that perfect chorus of voices, words, arguments and insights?

Search again.

Find hope and optimism in our laboratories.

Open the door to the greatest discovery of all:

It’s their keen intellect, smiles, kind words or questions.

It’s crafted by their company.

Caffeine-fuelled conversations critique, criticize, challenge.

(Coffee is never bitter with graduate students)

Embracing curiosity, creativity and collaboration.

Wrangling words together: perform, propose, predict.

Execute, explain, engage.

Fieldwork, funding, fellowship.

Null hypothesis, clear objectives, conceptual frameworks.

Significance and broader impact,

Contributions to knowledge.

Contributions for humanity.

I hope I did enough; I wish for more.

Fleeting moments are now warm memories:

Catching spiders on the tundra, or caterpillars in the canopy.

Thank you, students: you teach me.

We move beyond metrics and money.

We write, we study, we learn.

We discover.

We grow.

Crystal Ernst successfully defended her PhD on 23 Feb.

Crystal Ernst successfully defended her PhD on 23 Feb.

Dorothy Maguire (middle) successfully defended her PhD on 27

Dorothy Maguire (middle) successfully defended her PhD on 27 Feb. Elena Bennett was Dorothy’s co-supervisor.

Trophic cascades in fragmented forests

Many birds eat insects and spiders. Some of these insects and spider are themselves predators, feeding on critters lower down in the food web. Some of the insects that are fed upon by birds, or other predators, also play important roles in forest, such as munching upon the fresh, green leaves of young trees (here’s a reminder).

Munch, munch, munch. The hungry caterpillar. (photo by Sean McCann, reproduced here with permission)

Munch, munch, munch. The hungry caterpillar. (photo by Sean McCann, reproduced here with permission)

These interactions are ongoing, all the time, in forests around the world. These forests, however, are changing in important ways. Some of them are getting smaller and smaller as humans continue to encroach on the land, via urbanization or agriculture. This results in a ‘fragmented’ landscape. A landscape with small forest patches, perhaps no bigger than your back yard. A landscape with larger forests, perhaps one in which you could get lost in. These forests are themselves connected to each other –sometimes directly by a corridor or hedgerow.

This is the context for PhD student Dorothy Maguire’s research. Within that context, Dorothy tackled a fascinating project, one that was just recently published. In this work, Dorothy and co-authors (including me, an undergrad at that time, Thomas Nicole, and McGill Professor Elena Bennett) put cages around small trees in different types of forests SW of Montreal. The cages (made of chicken wire) were in place to test the effects of ‘predator exclusions’ on the insects and spiders occurring on saplings. The prediction is that if you exclude larger predators, such as birds, this may allow a ‘release’ of other insects and spiders. In turn, this release may have trickle-down effects on an important process occurring in young trees: herbivory. For example, if a predator is more common because it’s not being eaten by birds, perhaps it will eat more caterpillars, which may mean the leaves on trees will be eaten less frequently. In ecology this is dubbed a ‘trophic cascade’. Dorothy did this work in the context of fragmented forests, and she worked in forests that were either small and isolated from other forests, or in forests that were large and connected to other forests. This was done because there’s an expectation that these ecological effects will be different depending on the degree of fragmentation happening on the landscape. For example, insectivorous birds may decrease in abundance in small, isolated patches, which means their effects on insect prey (and perhaps herbivory) may be reduced relative to effects in larger patches of forest.

Dorothy Maguire, working in a forest fragment.

Dorothy Maguire, working in a forest fragment.

During one summer field season, Dorothy and Thomas wrapped up some small sugar maple trees in chicken wire, left some alone as controls, counted insects and spiders over the summer months, and measured herbivory on the trees themselves. As expected, the effects of the ‘cage’ was significant: when you put a cage around a tree, you end up with more arthropods living on those trees. This confirms other papers which report a similar effect: insectivorous birds (and perhaps other vertebrate predators) have a significant, and meaningful impact on the insects and spiders living on trees. Or, stated another way, birds eat critters living on trees, and without these birds, there would certainly be more arthropods around!

Dorothy did not uncover a strong effect on the process of insect herbivory: although more insects and spiders were living in the trees protected by chicken wire, the leaves themselves were not affected. This could be because more insect predators were around, and thus compensating for the lack of birds, and eating just as many herbivorious insects (e.g., caterpillars) as the birds might have eaten.

The lanscape of southern Quebec. Lots of agriculture, some patches of forest.

The lanscape of southern Quebec. Lots of agriculture, some patches of forest.

Scaling up to the landscape context, there were no overall significant effects of the cage treatments in relation to the forest type, nor was the level of herbivory dependent on the landscape context. The general results for large, connected patches were no different than for small, isolated patches. However, the magnitude of the effect was marginally affected by the landscape context for the cage exclusion: vertebrate predator may have a more significant impact in smaller, isolated patches.

As with all research projects, this work resulted with as many questions as answers, which is equally frustrating and fascinating. It’s clear that vertebrate predators are important in these systems, but more work is needed to fully assess whether these effects are truly affected by the degree of forest fragmentation on the landscape. The lack of effects on the process of herbivory itself was equally intriguing – there are clearly many complex interactions occurring on small maple trees. Some of these interactions involve top-down predation events, but there are likely a suite of ‘bottom-up’ effects that are also influencing the system.

Reference:

MAGUIRE, D. Y., NICOLE, T., BUDDLE, C. M. and BENNETT, E. M. (2014), Effect of fragmentation on predation pressure of insect herbivores in a north temperate deciduous forest ecosystem. Ecological Entomology. doi: 10.1111/een.12166

Congratulations to the lab

Last week my laboratory attended the Entomological Society of Canada’s annual meeting, held in Guelph. I was so proud of the whole lab – we had an impressive showing at the meeting, and I was especially impressed with the three undergraduate students who presented their research to Entomologists from across Canada. Wow – I don’t think I had that amount of confidence when I was an undergrad!

I am THRILLED to announce that three arthropod ecology students were recognized for their excellent presentations.

First, Master’s student Étienne Normandin was awarded first prize for his oral presentation in the Biodiversity section.  His talk was titled Biodiversity of wild bees in two urban settings: Montreal and Quebec city. He’s co-supervised by Valérie Fournier at Laval University. Here’s a photo of Etienne doing some field work:

Field work!

Field work!

Second, PhD student Dorothy Maguire was the runner up in the the same Biodiversity section. Her oral presentation was on Insect herbivory in fragmented forest landscapes: linking land use with changes in biodiversity and ecosystem function. Dorothy is co-supervised by Elena Bennett. Here’s a photo of Dory doing what she loves!

Tree climbing!

Tree climbing!

Finally, PhD student Raphaël Royauté was runner up in the student poster competition, for his work titled Does physiological state affect individual variation in boldness in a jumping spider?  Raphaël is co-supervised by Dr. Charles Vincent, from Agriculture and Agri-Food Canada.  Here’s an older photo of me and Raph, not long after he first came to the lab for a short internship.

Raphael (left) and Chris (right), working hard.

Raphael (left) and Chris (right), working hard.

Spider cakes!

My graduate students are a very talented bunch – they are intelligent, creative, and have a good sense of humour.  Some of our lab group celebrated birthdays recently, and in honour of this, we had two cakes earlier this week.  The first, made by MSc student Sarah Loboda, is the VERY BEST SPIDER CAKE I have ever seen (or eaten!).  Check this out:

Spider Cake!

Of course, let’s discuss how anatomically correct that cake is!  Two body parts, pedicel, eight legs (coming from the cephalothorax, of course), and a bunch of eyes.

Spider cake! (eyes0

As you may know, most spiders in Canada have eight eyes, but since some do have six, I find it quite acceptable that this spider has six eyes.  Furthermore, not all spider eyes are identical so it is appropriate to have two kinds represented on the cake.  Well done, Sarah.

And in case that STUNNING MASTERPIECE isn’t enough, another student (Dorothy Maguire)  made a cake that is a very good approximation for the female epigynum of wolf spiders in the genus Pardosa.

Pardosa epigynum

And not just any Pardosa:  this is diagnostically similar to one of the species that graduate student Katie Sim is working on!  Incredible!

….want some proof – look at this image, taken from Dondale & Redner’s text on the Lycosidae of Canada.  Enough said.

Pardosa concinna epigynum

Arthropods in the tree-tops: Canopy ecology in Quebec (Part 3)

This is the final post in a three part series about studying canopy arthropods in Quebec.  Part 1 was about canopy access and Part 2 was about patterns of diversity.  This post is about ecological interactions in the canopy. 

I had the pleasure of supervising a M.Sc. student, Kathleen Aikens, who was keen to work on a canopy project that looked deeper into some of the ecological interactions occurring in our deciduous forest canopies.  This was possible since we had, by this time, acquired a lot of base-line data on arthropods in many strata of the forest.  Kathleen’s work included using exclosure cages to see whether or not bird predation might affect arthropods in the Canopy differently than in the understorey.   This was exciting work, as it took our laboratory in a new direction, and lets us start to unravel some of the complexities of the food-webs in the tree-tops.  Her main result was that birds did have a strong top-down effect on arthropods, and that effect did differ as a function of height.  Using some bait trials, we also found that predation by arthropods on arthropods was also stratified.   This research suggests that arthropods living in trees in our region of the world are always under significant predation pressure, from both vertebrate and invertebrate predators.

A cage experiment, to assess the effect of predators on insects living in the forest canopy.

More recently, my laboratory has started to collaborate closely with another group at McGill studying “ecosystem services” – this is work done with Elena Bennett, another colleague at McGill University.  The research framework with this project is about how different ecosystem services are affected by the fragmented landscape that occurs in a large region just south of Montreal.  Elena and I co-supervise a PhD student Dorothy Maguire, who is looking at the ecosystem function of insect herbivory, and studying how herbivory varies as a function of forest size and degree of isolation (i.e., from a large contiguous forest), and she is studying herbivory in the understorey as well as the canopy.  Herbivory is closely linked to ecosystem services because of its effect on nutrient cycling, forest aesthetics, and more.  Although this project is currently underway, Dorothy is uncovering some interesting results, already.  For example, she is finding that levels of insect herbivory differ between the understorey and the canopy, and that forest fragmentation is affecting insect herbivory.

Summary

I have provided some highlights of some of the work that our laboratory has done in Quebec’s deciduous forests (and my apologies to the students who I didn’t mention!).  Although we have come a long way, and uncovered some interesting research results, I still feel that the work is just beginning.  For example, the bulk of our work has been on only two tree species (Sugar Maple and American Beech), and we have only studied a fraction of the arthropods that exist in the canopies of our forests.  I would like to expand the research to include other plant-feeding guilds, bees and wasps.  I’m also always curious about the piles of dead and decaying leaves that we find nestled between the crotches of high branches – these micro-habitats surely contain suspended soil (e.g., see Lindo & Winchester 2007), and within those “islands” there should be a host of arthropods.   Not surprisingly, the forest canopies in southern Quebec are home to a marvelous diversity of arthropods.  It’s a scientist’s model system, and a delightful system in which to work and play.

Me (Chris Buddle) above the canopy at Mont St Hilaire!