Changing Forest Structure

It’s been a while since I posted here about the forest modelling I’ve been working on here at MSU. Over the last couple of months I’ve been working on finalizing the regeneration modelling component, refining the timber harvest rules, linking simulations to the bird occupancy modelling I started this spring, and writing it all up for manuscripts.

Across our study area we’ve found that regeneration of juvenile trees following timber harvest varies greatly. For example, from our empirical data we find that sugar maple saplings were present in over 70% of northern forest gaps but were completely absent from 96% of gaps in southern areas. Megan Matonis suggested in her thesis that this variation is related to snow depth, deer density and soil nutrient conditions. To examine the potential long-term effects of these differences in regeneration on forest structure I’ve been running our simulation model with pre-set levels of regeneration that reflect our observations, ranging from the maximum possible (given the space available in a post-harvest gap) to a complete absence of regenerating juvenile trees.

These ‘gaps’ I’m talking about are created in northern hardwood forests when individual or small groups of trees are removed in an uneven-aged timber management approach. The removal of these trees creates openings (‘gaps’) in the forest canopy allowing light into lower levels for younger trees [gaps may also be created naturally but we’re focusing on those created by human activity which is the dominant driver in our study area]. When harvesting trees in this approach foresters aim to produce a forest structure with a ‘reverse-J’ distribution of tree sizes; high densities of small, young trees and low densities of larger, older trees (approximating a gamma-distribution like I found in our data previously). The idea is that through time an abundant supply of competing smaller trees will replace larger trees trees that are removed.

Representing this approach in our model (using FVS keywords [.pdf]) requires quite a bit of code, but working through the example provided by Don Vandendriesche [.pdf] helped. This approach requires the model user to specify a residual basal area (the area occupied by trees) and the ratio between the number of trees in successive size classes (the q-factor).

To examine my initial results (and to help debugging during the whole modelling process) I used R to plot size-class distributions for tree densities and basal area. As is the norm I used size-classes defined by the diameter-at-breast-height of the trees (5 cm or about 2 inches). Then I combined plots for simulated years into animated .gif files to see how the distributions changed through time for different regeneration levels. Here are a couple of examples (click for larger versions):

By the end of these 200-year simulations the same stand has a very different forest structure. In the top example regeneration is sufficient to replace trees removed during harvest, growing into larger size-classes as more resources (light and space) become available. But in the bottom example we see the consequences of when no new trees grow to replace the the removed trees – by the mid-21st century there are no trees in the smaller size-classes and timber harvesting has to become less frequent to meet timber removal goals (and remain viable).

I’m continuing to analyse the model output in a more quantitative manner and assessing the impacts of these potential changes in forest structure on bird habitat (specifically the probability that different species will be present in a forest stand). All together this should make a nice manuscript and provide some interesting information for the foresters working in these northern hardwood forests.

Leverhulme Early Career Fellowship

Around the time I wrote this blog about the National Assessment of UK Forestry and Climate Change Steering Group report I was thinking about writing a proposal to the Leverhulme Trust for an Early Career Fellowship. I found out recently that my proposal was successful and so from January 2011 I will be back at King’s College, London!

The Leverhulme Trust makes awards in support of research and education with special emphasis on original and significant research that aims to remove barriers between traditional disciplines. Their Early Career Fellowships are awarded across all disciplines and in 2010 approximately 70 were expected to be awarded to individuals to hold at universities in the UK. Given the emphasis on original, significant and cross-disciplinary research made by the Trust I looked for something that matched my research skills in coupled human and natural systems modelling but that pushed work in that area in a new direction. I thought back to the ideas about model narratives I have previously explored with David O’Sullivan and George Perry (but have not worked on since then) and Bill Cronon’s plenary address at the Royal Geographical Society in 2006 on the need for ‘sustainable narratives’. With that in mind, and given the UK Forestry and Climate change report I had been reading, I decided to make a pitch for a project that would explore how narratives from the use of models could help individuals identify how local actions transcend scales to mitigate global climate change in the context of the anticipated woodland planting that will be ongoing in the UK in future years. It proved to be a successful pitch!

I’m sure I will blog plenty more about the project in the future, so for now I will just leave you with the proposal rationale (below). I’m looking forward to getting to work on this when I get back to London, but before that there’s plenty more things to get done on the Michigan forest landscape ecological-economic modelling.

Model narratives for climate change mitigation
The abstract, vast, and systemic narratives that dominate the issue of global climate change do little to illustrate to individuals and groups how their actions might contribute to mitigate the effects of what is often framed as a global problem (Cronon 2006). Ways to improve the ability of individuals and groups to identify how their local actions transcend scales to mitigate global climate change are needed. In this research I will explore how narratives produced from computer simulation models that represent individuals’ actions can provide people with insights into how their behaviour affects system properties at a larger scale. Although the narrative properties of simulation models have been highlighted (O’Sullivan 2004), the use of models to develop localised narratives of climate change which emphasise individual agency has yet to be explored. Confronting individuals with these narratives will also help researchers reveal important underlying, and possibly implicitly held, assumptions that influence choices and behaviour.

This research will address the following general questions:

  • How can computer simulation models be better used to reveal to individuals how their local actions can contribute to global environmental issues such as Climate Change Mitigation (CCM)?
  • What are the narrative properties of simulation models and how can they be exploited to help individuals find meaning about their actions as they relate to global climate change?
  • By using simulation tools to spur reflection what can we learn about the factors influencing individuals’ choices and behaviour with regards CCM options?

Answering these questions will require a uniquely interdisciplinary research approach that spans the physical sciences, social sciences and humanities. Such ground-breaking, boundary-crossing work is necessary if we are to re-connect the physical sciences with the publics they intend to benefit and find solutions to large-scale and pressing environmental problems. For example, one of the key findings from a recent report by the National Assessment of UK Forestry and Climate Change Steering Group (Read et al. 2009) was that “[t]he extent to which the potential for additional [greenhouse gas] emissions abatement through tree planting is realized … will be determined in large part by economic forces and society’s attitudes rather than by scientific and technical issues alone” (p.xvii). The report also argued the need “to better understand and consider the role of different influences affecting choices and behaviour. Without the appropriate emotional, cultural or psychological disposition, information will make no difference.” (p.210). Narratives based on scientific understanding which portray how individuals can make a difference to large-scale, diffuse environmental issues will be important for fostering such a disposition. Simulation models – quantitative representations of reality which provide a means to logically examine how high-level and large-scale patterns are generated by lower-level and smaller-scale processes and events – have the potential to contribute to the construction of these narratives.

Bird Occupancy Modelling

Birds have been given short shrift in my posts blog posts about the Michigan UP ecological-economic modelling project. It’s not that we have forgotten about them, it’s just that before we got to incoporating them into our modelling there were other things to deal with first. Now that we’ve made progress on modelling deer distribution it’s time to turn our attention to how we can represent the potential impacts of forest management on bird habitat so that we might better understand the tradeoffs that will need to be negotiated to achieve both economic and ecological sustainability.

Ovenbird (Seiurus aurocapillus)
Ovenbird (Seiurus aurocapillus)

One of the things we want to do is link our bird-vegetation modelling with Laila Racevskis‘ assessment of the economic value of bird species she did during her PhD research. Laila assessed local residents’ willingess-to-pay for ensuring the conservation of several bird species of concern in our study area. If we can use our model to examine the effects of different timber management plans (each yielding different timber volumes) on the number of bird species present in an area we can use Laila’s data to examine the economic tradeoffs between different management approaches. The first thing we need to do to achieve this is be able to estimate how many bird species would be present in a given forest stand.

Right now the plan is to estimate the presence of songbird species of concern in forest stands by using the data Ed Laurent collected during his PhD research at MSU. To this end I’ve been doing some reading on the latest occupancy modelling approaches and reviewing the literature on its application to birds in managed forests. Probably the most popular current approach was developed recently by Darryl Mackenzie and colleagues – it allows the the estimation of whether a site is occupied by a given species or not when we know that our detection is imperfect (i.e. when we know we have false negative observations in our bird presence data). The publication of some nice overviews of this approach (e.g. Mackenzie 2006) plus the development of software to perform the analyses are likely to be at the root of this popularity.

The basic idea of the approach is that if we are able to make multiple observations at a site (and if we assume that bird populations and habitat do not change between these observations) we can use the probability of each bird observation history at a site across all the sites to form a model likelihood. This likelihood can then be used to estimate the parameters using any likelihood-based estimation procedure. Covariates can be used to model both the probability of observation and detection (i.e. we can account for factors that may have hindered bird observation such a wind strength or the time of day). I won’t go into further detail here because there’s an excellent online book that will lead you through the modelling process, and you can download the software and try it yourself.

Two recent papers have used this approach to investigate bird species presence given different forest conditions. DeWan et al. 2009 used Mackenzie’s occupancy modelling approach to examine impacts of urbanization on forest birds in New York State (they do a good job of explaining how they apply Mackenzie’s approach to their data and study area). DeWan considered landscape variables such as perimeter-area ratios of habitat patches and proximity to urban area to create occupancy models for 9 birds species at ~100 sites. They found that accounting for imperfect bird detection was important and that habitat patch “perimeter-area ratio had the most consistent influence on both detection probability and occupancy” (p989).

In a slightly different approach Smith et al. 2008 estimated site occupancy of the black-throated blue warbler (Dendroica caerulescens) and ovenbird (Seiurus aurocapillus) in 20 northern hardwood-conifer forest stands in Vermont. At each bird observation site they had also collected stand structure variables including basal area, understory density and tree diameters (in contrast to DeWan et al who only considered landscape-level variables). Smith et al. write their results “demonstrate that stand-level forest structure can be used to predict the occurrence of forest songbirds in northern hardwood-conifer forests” (p43) and “suggest that the role of stand-level vegetation may have been underestimated in the past” (p36).

Our approach will take the best aspects from both these studies; the large sample size of DeWan et al. with the consideration of stand-level variables like Smith et al. More on this again soon I expect.

Holiday Publications!

Update January 2010: This paper is now online with doi 10.1016/j.foreco.2009.12.020.

I received some good news this morning as I prepared to head back to the UK for the holidays. The paper I started writing back in January examining the white-tailed deer distribution in our managed forest landscape (the analysis for which inspired posts on Bayesian and ensemble modelling) has been accepted for publication and is ‘In Press’! I’ve copied the abstract below.

Another piece of publications news I received a while back is that the paper I co-authored with Raul Romero-Calcerrada and others modelling socioeconomic data to understand patterns of human-caused wildfire ignition risk has now officially been published in Ecological Modelling.

Happy Holidays everyone!

Effects of local and regional landscape characteristics on wildlife distribution across managed forests (In Press) Millington, Walters, Matonis, and Liu Forest Ecology and Management

Understanding impacts of local and regional landscape characteristics on spatial distributions of wildlife species is vital for achieving ecological and economic sustainability of forested landscapes. This understanding is important because wildlife species such as white-tailed deer (Odocoileus virginianus) have the potential to affect forest dynamics differently across space. Here, we quantify the effects of local and regional landscape characteristics on the spatial distribution of white-tailed deer, produce maps of estimated deer density using these quantified relationships, provide measures of uncertainty for these maps to aid interpretation, and show how this information can be used to guide co-management of deer and forests. Specifically, we use ordinary least squares and Bayesian regression methods to model the spatial distribution of white-tailed deer in northern hardwood stands during the winter in the managed hardwood-conifer forests of the central Upper Peninsula of Michigan, USA. Our results show that deer density is higher nearer lowland conifer stands and in areas where northern hardwood trees have small mean diameter-at-breast-height. Other factors related with deer density include mean northern hardwood basal area (negative relationship), proportion of lowland conifer forest cover (positive relationship), and mean daily snow depth (negative relationship).The modeling methods we present provide a means to identify locations in forest landscapes where wildlife and forest managers may most effectively co-ordinate their actions.

Keywords: wildlife distribution; landscape characteristics; managed forest; ungulate herbivory; northern hardwood; lowland conifer; white-tailed deer

Plant Now and Use Sustainably

In the climate change debate there’s been a lot of talk about current Amazonian rainforest deforestation, but I’ve heard much less about the role of the UK’s forests for carbon sequestration. Given the relative size of the UK to the Amazonian rainforest that’s not so surprising – The Nature Conservancy suggests the area of rainforest cut down each year (20 million hectares) is the same as the combined area of England, Scotland and Wales. The estimated 5000 years it took [.pdf] to go from 75% of the UK covered by forests and woodlands to the current 12% just doesn’t compare.

Recently, however, the case been made for increasing UK forest and woodland cover as a form of climate change mitigation. This summer the UK Low Carbon Transition Plan identified woodland creation as a cost-effective way of mitigating climate change and recognised the importance of supporting tree-planting initiatives. More recently, the National Assessment of UK Forestry and Climate Change Steering Group has provided its response to the IPCC Fourth Assessment Report and argues there is a clear need for more woodlands.

One of the main findings of this initial assessment was that an increase in woodland area of 23,000 ha per year over the next 40 years could abate 10% of UK 2050 greenhouse gas emissions. With echos of the recommendation from the Stern Review on the Economics of Climate Change to ‘Act now or pay later’, the key message from this assessment is ‘Plant now and use sustainably’. The long maturation times of forest systems means that it may take take 50–100 years for actions to pay off.

Being such a long-standing investment it’s vital that the benefits of planted woodlands and forests are not outweighed by negative impacts on biodiversity, food security, landscape and water supply. From this stand-point there is much to be done and the assessment recommends that “further scientific and socio-economic analysis is required to enable the UK to achieve the full [climate] adaptation and mitigation potential of forestry” and that “clear, robust, research programmes will be needed to underpin the changes of forestry policy and practice which are required to meet the new and challenging circumstances”.

A question that immediately springs to my mind is where these woodlands should be placed to maximise their carbon sequestration payoff while minimising negative impacts on other aspects of the landscape. For example, if arable agricultural land is to be converted, how will biodiversity be affected by the removal of hedgerows? What would this conversion of agricultural land mean for local economies? Which species will benefit in terms of habitat connectivity and which will lose out? Addressing questions like these will be important as forest policy moves toward returning UK forest cover area near levels seen elsewhere in Europe.

Initial Michigan Forest Simulation Output

It’s taken a while but finally the model that I came to Michigan State to develop is producing what seems to be sensible output. Just recently we’ve brought all the analyses on the data that were collected in the field into a coherent whole. We’ll use this integrated model to investigate best approaches for forest and wildlife management to ensure ecological and economic sustainability. This post is a quick overview of what we’ve got at the moment and where we might take it. The image below provides a simplified view of the relationship of the primary components the model considers (a more detailed diagram is here).

The main model components I’ve been working on are the deer distribution, forest gap regeneration and tree growth and harvest sub-models. Right now we’re still in the model testing and verification stage but soon we hope to be able start putting it to use. Here’s a flow chart representing the current sequence of model execution (click for larger image):

As I’ve posted several times about the deer distribution modelling (here, here, and here for example) and because the integration of FVS with our analyses is more a technical than scientific issue, I’ll focus on the forest gap regeneration sub-model.

Most of the forest gap regeneration analyses used the data Megan Matonis collected during her two summers in the field (i.e., forest). During her fieldwork Megan measured gap and tree regeneration attributes such as gap size, soil and moisture regime, time since harvest, deer density, and sapling heights, density and species composition. Megan is writing up her thesis right now but we’ve also managed to find time to do some extra analyses on her data for the gap regeneration sub-model. Here’s the flow chart representing the model sequence to estimate initial regeneration in gaps created by a selection harvest in a forest stand (click for larger image):

In our gap regeneration sub-model we take a probabilistic approach to estimate the number and species of the first trees to reach 7m (this is the height at which we pass the trees to FVS to grow). The interesting equations for this are Eqs. 6 – 9 as they are responsible for estimating regeneration stocking (i.e. number of trees that regenerate) and the species composition of the regenerating trees. Through time the effects of the results of these equations will drive future forest composition and structure and the amount of standing timber available for harvest.

The probability that any trees regenerate in a gap is modelled using a generalized linear mixed model with a stand-level random intercept drawn from a normal distribution. The probability is a function of canopy gap area and deer browse category (high or low; calculated as a function of deer density in the stand).

If there are some regenerating trees in the gap, we use a logistic regression to calculate the probability that the gap contains as many (or more) trees as could fit in the gap when all the trees are 7m (and is therefore ‘fully stocked’). The probability is a function of canopy openness (calculated as a function of canopy gap area), soil moisture and nutrient conditions and deer density. If the gap is not fully stocked we sample the number of trees using from a uniform distribution.

Finally, we assign each tree to a species by estimating the relative species composition of the gap. We do this by assuming there are four possible species mixes (derived from our empirical data) and we use a logistic regression to calculate the probability that the gap has each of these four mixes. The probability of each mix is a function of soil moisture and nutrient conditions, canopy gap area, and stand-level basal area of Sugar Maple Ironwood. Currently we have parameterised the model to represent five species (Sugar Maple, Red Maple, White Ash, Black Cherry and Ironwood).

As the flow chart suggests, there is a little more to it than these three equations alone but hopefully this gives you a general idea about how we’ve approached this and what the important variables are (look out for publications in the future with all the gory details). For example, at subsequent time-steps in the simulation model we grow the regenerating trees until they reach 7m and also represent the coalescence of the canopy gaps. I haven’t integrated the economic sub-model into the program yet but that’s the next step.

So what can we use the model for? One question we might use the model to address is, ‘how does change in the deer population influence northern hardwood regeneration, timber revenue and deer hunting value?’ For example, in one set of initial model runs I varied the deer population to test how it affects regeneration success (defined as the number of trees that regenerate as a percentage of the maximum possible). Here’s a plot that shows how regeneration success decreases with increasing deer population (as we would expect given the model structure):

Because we are linking the ecological sub-models with economic analyses we can look at how these differences will play out through time to examine potential tradeoffs between ecological and economic values. For example, because we know (from our analyses) how the spatial arrangement of forest characteristics influences deer distribution we can estimate how different forest management approaches in different locations influences regeneration through time. The idea is that if we can reduce deer numbers in a given area immediately after timber harvest we can give trees a chance to survive and grow above the reach of deer – moving deer spatially does not necessarily mean reducing the total population (which would reduce hunting opportunities, an important part of the local economy). The outcomes may look something like this:

We plan to use our model to examine scenarios like this quantitatively. But first, I need to finish testing the model…

Synthetic Trees

When testing and using simulation models we often need to use synthetic data. This might be because we want to examine the effects of different initial conditions on our model output, or simply because we have insufficient data to examine a system at the scale we would like to. The ecological-economic modelling project I’m currently working on is in both these situations, and over the last week or two I’ve been working on generating synthetic tree-level data so that we can initialize our model of forest stand change for testing and scenario development. Here’s a brief overview of how I’ve approached the task of producing a ‘treelist generator’ from the empirical data we have for over 60,000 trees in Northern Hardwood stands across Upper Michigan.

One of the key measures we can use to characterise forest stands is basal area (BA). We can assume that for each stand we generate a treelist for there is some ‘target BA’ that we are aiming to produce. As well as hitting a target BA, we also need to make sure that the tree diameter-at-breast-height (DBH) size-class distribution and species composition are representative of the stands in our empirical data. Therefore, our the first step is to look at the diameter size-class distribution of the stands we want to emulate. We can do this by plotting histograms of the frequency of trees of different diameter for each stand. In the empirical data we see two characteristic distributions (Fig 1).

Fig 1. Example stand tree count histograms

The distribution on the left has very many more trees in the smaller size classes as a result of stands self-thinning (as larger trees compete for finite resources). The second distribution, in which the smallest size classes are under-represented and larger size classes have relatively more trees, does not fit so well with the theoretical, self-thinning DBH size-class distribution. Stands with a distribution like this have probably been influenced by other factors (for example deer browse on the smaller trees). However, it turns out that both these DBH size-class distributions can be pretty well described by the gamma probability distribution (Fig 2).

Fig 2. Example stand gamma probability distributions for data in Fig 1

The gamma distribution has two parameters, a shape parameter we will call alpha and a scale parameter we will call beta. Interestingly, in the stands I examined (dominated by Sugar Maple and Ironwood) there are two different linear relationships between the parameters. The relationship between alpha and beta for 80% of stands represents the ‘self-thinning’ distribution, and the other 20% represent distributions in which small DBH classes are under-represented. We use these relationships – along with the fact that the range of values of alpha for all stands has a log-normal distribution – to generate characteristic DBH size-class distributions;

  1. sample a value of alpha from a normal distribution (subsequently reconvert using 10alpha),
  2. for the two different relationships use Bayesian linear regression to find mean and 95% credible intervals for the slope and intercept of a regression line between alpha and beta,
  3. use the value of alpha with the regression parameters to produce a value of beta.

So now for each stand we have a target basal area, and parameters for the DBH size class distribution. The next step is to add trees to the stand with diameters specified by the probability distribution. Each time we add a tree, basal area is added to the stand. The basal area for a tree is calculated by:

TreeBA = TreeDensity * (0.005454* diameter2)

[Tree density can be calculated for each tree because we know the sampling strategy used to collect empirical data on our timber cruise, whether on a fixed area plot, n-tree or with a prism].

Once we get within 1% of our target BA we stop adding trees to the stand [we’ll satisfy ourselves with a 1% accuracy because the size of tree that we allocate each time is sampled from a probability distribution and so we it is unlikely we will be able to hit our target exactly]. The trees in our (synthetic) stand should now (theoretically) have the appropriate DBH size-class distribution and basal area.

With a number of trees in now in our synthetic stand, each with a DBH value, the next step is to assign each tree to a species so that the stand has a representative species composition. For now, the two species we are primarily interested in are Sugar Maple and Ironwood. However, we will also allow trees in our stands to be Red Maple, White Ash, Black Cherry or ‘other’ (these are the next most common species in stands dominated by Sugar Maple and Ironwood). First we estimate the proportion of the trees in each species. In stands with Sugar Maple and Ironwood deer selectively browse Sugar Maple, allowing Ironwood a competitive advantage. Correspondingly, in the empirical data we observe a strong linear and inverse relationship between the abundance of Sugar Maple and Ironwood (Fig 3).

Fig 3. Relationship between stand Sugar Maple and Ironwood abundance

To assign species proportions we first estimate the proportion of Sugar Maple from the empirical data. Next, using the strong inverse relationship above we estimate the corresponding proportion of Ironwood (sampled using normal distribution with mean and standard deviation from from Bayesian linear regression). The remaining species proportions are assigned according to the frequency of their presence in the empirical data.

Now we use these proportions to assign a species to individual trees. Because physiology varies between species, the probability that a tree is of a given size also varies between species. For example, Ironwood very seldom reach DBH greater than 25 cm and the vast majority (almost 99% in our data) are smaller than 7.6 cm (3 inches) in diameter. Consequently, first we assign the appropriate number Ironwood to trees according to their empirical size-class distribution, before then assigning all other trees to the remaining species (using a uniform distribution).

The final step in generating our treelist is to assign each tree a height and a canopy ratio. We do this using empirical relationships between diameter and height for each species that are available in the literature (e.g. Pacala et al. 1994). And we’re done!

In the model I’m developing, these stands can be assigned a spatial location either using a pre-existing empirical map or using a synthetic land cover map with known characteristics (generated for example using the modified random clusters method, as the SIMMAP 2.0 software does). In either case we can now run the model multiple times to investigate the dynamics and consequences of different initial conditions. More on that in the future.

Disturbance and Landscape Dynamics in a Changing World

Experimentation can be tricky for landscape ecologists, especially if we’re considering landscapes at the human scale (it’s a bit easier at the beetle scale [pdf]). The logistic constraints of studies at large spatial and temporal scales mean we frequently use models and modelling. However, every-now-and-then certain events afford us the opportunity for a ‘natural experiment’ – situations that are not controlled by an experimenter but approximate controlled experimental conditions. In her opening plenary at ESA 2009, Prof. Monica Turner used one such natural experiment – the Yellowstone fires of 1988 – as an exemple to discuss how disturbance affects landscape dynamics and ecosystem processes. Although this is a great example for landscapes with limited human activity, it is not such a useful tool for considering human-dominated landscapes.

Landsat satellite image of the Yellowstone fires on 23rd August 1988. The image is approximately 50 miles (80 km) across and shows light from the green, short-wave infrared, and near infrared bands of the spectrum. The fires glow bright pink, recently burned land is dark red, and smoke is light blue.

Before getting into the details, one of the first things Turner did was to define disturbance (drawing largely on Pickett and White) and an idea that she views as critical to landscape dynamics – the shifting mosaic steady state. The shifting mosaic steady state, as described by Borman and Likens, is a product of the processes of vegetation disturbance and succession. Although these processes mean that vegetation will change through time at individual points, when measured over a larger area the proportion of the landscape in each seral stage (of succession) remains relatively constant. Consequently, over large areas and long time intervals the landscape can be considered to be in equilibrium (but this isn’t necessarily always the case).

Other key ideas Turner emphasised were:

  • disturbance is a key component in ecosystems across many scales,
  • disturbance regimes are changing rapidly but the effects are difficult to predict,
  • disturbance and heterogeneity have reciprocal effects.

Landscape Dynamics
In contrast to what you might expect, very large disturbances generally increase landscape heterogeneity. For example, the 1988 Yellowstone fires burned 1/3 of the park in all forest types and ages but burn severity varied spatially. Turner highlighted that environmental thresholds may determine whether landscape pattern constrains fire spread. For instance, in very dry years spatial pattern will likely have less effect than years where rainfall has produced greater spatial variation in fuel conditions.

Turner and her colleagues have also found that burn severity, patch size and geographic location affected early succession in the years following the Yellowstone fires. Lodgepole pine regeneration varied enormously across the burned landscape because of the spatial variation in serotiny and burn severity. Subsequently, the size, shape and configuration of disturbed patches influenced succession trajectories. Turner also highlighted that succession is generally more predictable in small patches, when disturbances are infrequent, and when disturbance severity/intensity is low (and vice versa).

Ecosystem Processes
One of the questions landscape ecologists have been using the Yellowstone fires to examine is; do post-disturbance patterns affect ecosystem processes? Net Primary Production varies a lot with tree density (e.g., density of lodgepole pine following fire) and the post-fire patterns of tree density have produced a landscape mosaic of ecosystem process rates. For example, Kashian and colleagues found spatial legacy effects of the post-fire mosaic can last for centuries. Furthermore, this spatial variation in ecosystem process rates is greater than temporal variation and the fires produced a mosaic of different functional trajectories (a ‘functional mosaic’).

Another point Turner was keen to make was that the Yellowstone fires were not the result of fire suppression as is commonly attributed, but instead they were driven by climate (particularly hot and dry conditions). Later in the presentation she used the ecosystem process examples above to argue that the Yellowstone fires were not an ecological disaster and that the ecosystem has proven resilient. However, she stressed that fire will continue to be an important disturbance and that the fire regimes is likely to change rapidly if climate does. For example, Turner highlighted the study by Westerling and colleagues that showed that increased fire activity in the western US in recent decades is a result of increasing temperatures, earlier spring snowmelt and subsequent increases in vegetation moisture deficit. If climate change projections of warming are realised, by 2100 the climate of 1988 (which was extreme) could become the norm and events like the Yellowstone fires will be much more frequent. For example, using a spatio-temporal state-space diagram (seebelow), Turner and colleagues [pdf] found that fires in Yellowstone during the 15 years previous to 1988 had relatively little impact on landscape dynamics (shown in green in the lower left of the diagram). However, the extent of the 1988 fires pushed the disturbance regime up into an area of the state-space not characteristic of a shifting-mosaic steady state (shown in red).

The spatio-temporal state-space diagram used by Turner and colleagues [pdf] to describe potential landscape disturbance dynamics. On the horizontal x-axis is the ratio of disturbance extent (area) to the landscape area and on the vertical y-axis is the ratio of disturbance interval (time) to recovery interval. Landscapes in the upper left of the diagram will appear to an observer as relatively constant in time with little disturbance impact; those in the lower right are dominated by disturbance.

Remaining Questions
Turner finished her presentation by highlighting what she sees as key questions for studying disturbance and landscape dynamics in a changing world:

  • How will disturbance interact with one another?
  • How will disturbances interact with other drivers?
  • What conditions will cause qualitative shifts in disturbance regimes (like that shown in the diagram above)?

It was comforting to hear that a leader in the field identified these points as important as many of them relate closely to what I’ve been working on thinking about. For example, the integrated ecological-economic forest modelling project I’m working on here in Michigan explicitly considers the interaction of two disturbances – human timber harvest and deer herbivory. The work I initiated during my PhD relates to the second question – how does human land use/cover change interact and drive changes in the wildfire regime of a landscape in central Spain? And recently, I reviewed a new book on threshold modelling in ecological restoration for Landscape Ecology.

Much of Turner’s presentation and discussion applied to American landscapes with limited human activity. This not surprising of course, given the context of the presentation (at the Ecological Society of America) and the location of her study areas (all in the USA). But although natural experiments like the 1988 Yellowstone fires may be useful as an analogue to understand processes and dynamics in similar systems, it is also interesting (and important) to think about how other systems potentially differ from this examplar. For example, the Yellowstone fires natural experiment has little to say about disturbance in human-dominated landscapes that are prevalent in many areas of the world (such as the Mediterranean Basin). In the future, research and models of landscape succession-disturbance dynamics will need to focus as much attention on human drivers of change as environmental drivers.

Turner concluded her plenary by emphasising that ecologists must increase their efforts to understand and anticipate the effects of changing disturbance regimes. This is important not only in the context of climate as driver of change, but also because of the influence of a growing human population.

Cedar Swamps and Deer

Right now I should be back in East Lansing after a week of fieldwork in our Michigan Upper Peninsula (the UP) study area. We’ve been in the UP this last week to finish up on our mesic conifer planting and white-tailed deer density fieldwork that I’ve written about previously. However, an incident with a deer has delayed us (see the bottom of this post) so I’m doing some data entry and writing in Marquette while our Jeep is repaired.

In previous posts about the fieldwork we’ve done in the UP, I have included photos from forest stands containing deciduous hardwood species such as Sugar Maple or American Beech. Generally, it’s understood that white-tailed deer browse juveniles trees in hardwood stands during the daytime in the winter, but shelter overnight in nearby lowland conifer stands. One of the aspects of our project is to identify some quantitative relationships for this behaviour, and so we’ve often had take measurements in the cedar swamps adjacent to northern hardwood stands.

As you can see from the picture above, the density of cedar swamps can make tree measurements a bit tricky. A standard measure of forest stand density (or stocking) is ‘stand basal area’ – a measure of the area occupied by tree stems (i.e. trunks) in a given area. The northern hardwood stands in our study area can have a stand basal area of anywhere between 60 and 100 square feet per acre. Cedar swamps are much more densely populated, with stand basal area values of 280 to 350 square feet per acre. An example of the transition between these stand types is shown in the picture below (click for a larger image).

The high density of the cedar swamps combined with continual cover provided by the evergreen canopy (generally) make winter snow depths lower and winter air temperatures higher compared with the deciduous hardwood stands. The soggy conditions underfoot make surveying cedar swamps even trickier – one has to hop from tree-root island to tree-root island over puddles whilst trying not to impale oneself on the lower branches. Even with care given enough time you’re guaranteed scratches and wet boots.

We’ve completed our fieldwork for now and are just waiting for our Jeep to be fixed after we hit a deer on our last day of work. With so many deer in the area and the high number of miles we drive around our study area, it was only a matter time before we hit one. We were on a major highway and the deer came out of nowhere. We’ve often spooked deer driving on tracks through the forest – it seems to me that when they’re startled they just bolt in whatever direction they happen to be facing at the time. Even if that means running across the road in front of your vehicle. As you can see below, it left quite a dent in the radiator. But Megan did a good job of keeping us on the road and thankfully the only casualty was the deer.

Snowy UP Forests

Cut logs waiting for collection in the snow
On Monday several other members of the EE model research team and I met with foresters from Plum Creek and AFM to give them an overview of what we’ve been working on over the past year or so. Megan (Forestry Master’s student) and I gave them the lowdown on what we’ve been doing with regards fieldwork and analysis of the resulting data, Susan (Natural Resources Master’s student) spoke briefly about her work looking at factors influencing the prices of timber sales, and Mike (Forestry Prof.) was on hand to help paint the overall picture.

The foresters we spoke with were interested in our progress to date and asked for more details on tree species-specific patterns we find in our regeneration data so that they might work to continue the sustainability of their forest stands. Megan and are I are likely taking a trip to the study area again in late April to revisit a few sites from last spring and summer, so we’ll visit again then.

To get from one meeting to the other we drove through our study area. We wanted to see if we could find evidence of winter deer browse and generally get a feel for how the forests (and our study stands) look during the winter. We didn’t catch any deer in the act of browsing but, as the top picture below shows, we did see tracks and there were plenty of stunted maple saplings poking just above the snow nearby.

Deer tracks in the snow

snow and shadows