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.

 

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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.

 

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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.

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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.

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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