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University of Copenhagen

Macroecology and Community Ecology

Four hours ITX exam

26 January 2023

All aids (computer etc.) may be used, but no outside communication or

internet use. Please type or sketch everything – do not copy-paste figures
or tables from study materials. You can sketch by hand and take a photo,
draw on the computer or use statistical software.

The questions must be answered in English

Problem 1 counts 33 %
Problem 2 counts 33 %
Problem 3 counts 33 %

The exam set comprises 5 pages

Robert Buitenwerf Michael Krabbe Borregaard

(external examiner / censor) (examiner)

Jonathan Shik Hans Henrik Bruun

(examiner) (examiner)
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Problem 1
Communities of blackflies (Diptera: Simuliidae) were investigated in streams in a tropical area.
While adult female blackflies are biting flies, feeding on blood from mammals, the blackfly larvae
are aquatic filter feeders. They live on hard bottom substrates in streams. The larvae are equipped
with so-called labral fans (Fig. 1.1.), which they use to filter food items from the passing water
current. The species differ in fan morphology and this is thought to determine the size of the food
items they can catch. The species are also known to differ in their preferred abiotic habitat
conditions, such as stream flow velocity, water pH and temperature. So, some species prefer fast-
running water, while others like the conditions of more slowly running stretches of the streams.

Figure 1.1. Illustration of one species of blackfly, with the labral fans shown in detail

Scientists measured labral fan traits (length, width and number of fan rays) of 35 blackfly species
occurring in a set of stream sites (in total, 150 communities). For each of the 150 communities, they
calculated two standardized indices of trait dispersion (Mean Pairwise trait Distance – MPD, and
Mean Nearest-neighbour Trait Distance – MNTD). The two trait dispersion indices were combined
into an index of Standardized Effect Size (SES), which takes negative values for trait clustering and
positive values for trait over-dispersion. The observed SES values of trait dispersion were evaluated
by comparison to random draws of species from the regional species pool. Their results are
summarized in Table 1.1.

Questions
1. If the observed communities were mainly structured by exploitation competition for food items,
what expectations would you have about the trait dispersion? And why?
2. Conversely, if the observed communities were mainly structured by environmental filtering by
the abiotic environment, what expectations would you have about the trait dispersion? And why?
3. In the studied blackfly communities, what can you infer about community assembly processes
from the empirically observed results?
4. Propose an observational and/or experimental study, in the field or in the laboratory, designed to
discriminate between competing hypotheses for the observed patterns.

Table 1.1. For each of the three fan ray traits (rows), the number of communities with positive and negative index
values, respectively, is shown, as well as the t value and p value from a one-sample t-test.

Positive (%) Negative (%) t test value p value

SES Fan ray width 27 123 −8.209 <0.001
SES Fan ray length 21 126 −11.119 <0.001
SES Number of fan rays 33 117 −9.087 <0.001
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Problem 2
Marine snails are widely distributed from the poles to the Equator.

1. Figure 2.1(a) shows the diversity of species and genera as it varies with latitude. What is the
pattern exhibited in Figure 2.1(a)? Briefly mention and describe the main hypotheses for the pattern
shown in the figure. Could the same hypotheses explain the relationship for genera?

2. Figure 2.1(b) shows the species-genus ratio as it varies with latitude. Some researchers have
argued that variation in the species-genus ratio could be used to test for an ecological process.
Which process? Why did they argue this process will affect the species-genus ratio?

3. Can you come up with an evolutionary (long-term) process that might also lead to a high species-
genus relationship in a region (ignore methodological problems for now and discuss them under
point 4 just below)?

4. Species-genus relationships have been criticized as metrics for ecological and evolutionary
processes. Why?

5. The figure has two dashed lines showing the "expected species-genus ratio" derived from a
randomization. Describe how you could go about creating this expected relationship. Why does the
expected relationship vary with latitude in the way it does?

6. What is the definition of a null model and what are they used for in ecology?

7. Figure 2.1(c) shows the relationship between species-genus relationships for all species, wide-
ranging species and small-ranging species separated. What could cause the difference we observe?
What does that tell us about the cause of the species-richness gradient (you reviewed this in
question 1).

Figure 2.1. (a) Richness for species (squares) and genera (circles). (b) Species–genus (S/G) ratios for all marine
bivalves. Dashed lines represent 95% confidence intervals on the expected S/G ratios. (c) S/G ratio for all species (line),
and separated into wide-ranging species (open squares) and small-ranging species (filled squares)
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Problem 3
Direct predictive modelling of macroecological responses to climate warming requires proper
physiological assumptions. Functional traits provide a means of doing this. Here we use thermal
performance of ectotherms as an entry point to explore these dynamics. All questions count
equally.

Figure 3.1 (left): Latitudinal trends in A) warming tolerance (CTmax - Thabitat) and B) thermal safety margin (Toptimum –
Thabitat) in insects, where Thabitat is the mean annual surface air temperature. The black lines in each figure represent the
projected magnitude of warming by year 2100. (From Deutsch et al. 2008).

Figure 3.2 (right): Maximum and minimum thermal tolerance limits and range of annual extreme air (Ta) and operative
body temperatures (Te) as a function of latitude. Warm and cool color points indicate upper and lower thermal-
tolerance limits, respectively, after correcting for different acclimation temperatures. Lines indicate relationships from
best-fit linear models of thermal tolerance, which take into account taxonomy and different metrics of cold tolerance.
The gray region shows the range of hourly air temperatures (Ta) across the year, and the light yellow region shows the
range of extreme operative temperatures (Te) across the year, based on local regressions of lowland temperature data as
a function of latitude. (From Sunday et al. 2014).

1. Based on the warming tolerance and thermal safety margin results presented in Figure 1,
how and why do Deutsch et al. (2008) argue that impacts of climate warming on terrestrial
ectotherms will vary with latitude? Provide pros and cons of using measures of critical
thermal limits (i.e. CTmax and CTmin) as functional traits. How might a focus on climatic
variability instead of mean annual temperatures influence predictions about how latitude
influences vulnerability to climate warming?

2. Based on the results presented in Figure 2, how does a focus on Te by Sunday et al. (2014)
change predictions for latitudinal impacts of climate warming relative to those predicted by
Deutsch et al. (2008)? What additional organismal traits are important to consider when
predicting risks in warming environments?
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3. Why does Dan Janzen predict that “mountain passes are higher in the tropics”? Outline a
study that could test this hypothesis by comparing beetle species in Colorado and Ecuador.
What dependent and independent variables would be important to measure? Sketch and
compare thermal performance curves predicted for a beetle species inhabiting 1,000 meters
above sea level in Colorado vs. Ecuador. Label the axes and relevant features of the thermal
performance curves.