Assignment 2 Resources

Background reading and instructions/guidelines.

Instructions/guidelines
This assignment is worth 20% of your final mark.
Your task in this assignment is to produce a manuscript in the format of a scientific paper,
with an abstract, introduction, methods, results and discussion sections, acknowledgements
and a reference list, each section with its appropriate heading. See the ‘Resources – marking
rubric for Assignment 2’ within the Assessment tile for more information on the marking
criteria that will be used for this assignment.
To complete Assignment 2 successfully, you will need to do some background reading to
understand the pollination system and the community you will be writing about - the
fascinating community ecology associated with pollination and fruiting in monoecious figs in
general and in the species we are studying, the Port Jackson Fig, Ficus rubiginosa, in
particular. This reading material is provided below, plus further reading suggestions are
provided in the list of ‘Useful References’ provided below. These references are all available
from the UNE library and/or Google Scholar. You are welcome to use any or none of these
references for your assignment but you will be expected to source additional, relevant, peer-
reviewed papers whether you use any of those provided or not.
Examine the journal articles in the ‘Useful References’ to get formatting ideas for your own
manuscript. I am also providing a separate, short tutorial on scientific writing in the resources
for this assignment.
Beware – people who start looking into the amazing world of figs often get drawn in and it
becomes a life-long interest!
At the risk of being drawn in, watch the documentary, The Queen of Trees, about the
community of organisms associated with the Sycomore Fig, Ficus sycomorus, on YouTube:
https://www.youtube.com/watch?v=xy86ak2fQJM
This documentary gives a very good overview of the community of organisms associated
with an African fig, Ficus sycomorus. The community associated with Ficus rubiginosa is
similar in many respects (although no elephants).
Background reading
Firstly, a short glossary:
Syconium: A fig ‘fruit’ is more correctly referred to as a syconium. Syconia are the fleshy,
infolded structures that contain the separate male and female flowers of fig trees, shrubs and
vines. Each pollinated female flower – and there are hundreds of female flowers within each
syconium – is capable of developing a single seed, and when the seeds are mature the
syconium ripens and often changes colour to attract frugivores to eat these ripe fruit and
disperse the seed.
Foundress: A pollinator fig wasp that enters a syconium and lays eggs within it is known as
a foundress. The next generation of fig wasps - the progeny of foundresses that develop
within fig syconia - are referred to as emergents after they emerge from their galled flowers
within ripening syconia. A percentage of these emergents are successful in going on to being
foundresses themselves.
Ovipositor: A tubular structure on the female wasp, usually concealed but sometimes
extending outside the abdomen, with which the female wasp deposits eggs.

Your introduction should present an overview of your study that places it within the context
of relevant or similar research that has been published in peer-reviewed literature. It should
introduce the reader to the field of study, with definitions of particular jargon if necessary,
and outline the particular issues being studied. It should describe what other researchers have
shown, what our current knowledge is, and what gaps there are in our knowledge and
understanding (this part of your introduction should demonstrate a knowledge of the current
literature through citation of appropriate research including the latest relevant findings). You
should define your own hypothesis/hypotheses that you will be testing and conclude with a
clearly stated and precisely worded aims/objectives of your study.
Much of your introduction can be gleaned from the following readings but I stress that you
will be expected to add to this with your own review of the literature.
There are over 750 species of Ficus globally, of which about half, including F. rubiginosa,
are monoecious (i.e., they have separate male and female flowers on the same plant). Figs
occur around the globe in tropical and in many subtropical and temperate ecosystems. Figs
are widely recognised as keystone species because of the support they provide to populations
of the many species of vertebrate frugivores that feed on their fruit (Shanahan et al. 2001) and
the many avian insectivores that feed on the fig wasps (Mackay et al. 2018). They are called
‘keystone’ species because this support is so important to these other species that any decline
or loss of the keystone species would lead to the decline or loss of many other species as
well, with many consequential changes to the local ecosystem.
In this community study you will be exploring a symbiotic relationship. The symbiotic
relationship you will be examining is a plant-pollinator mutualism between a fig tree, F.
rubiginosa, and its pollinator, a fig wasp, Pleistodontes imperialis. Pollination of figs is
performed by specialised, fig-species-specific fig wasps, which enter the syconium of the fig
through the minute entrance (the ‘ostiole’) at the tip of a fig fruit to lay their eggs and to
pollinate the fig’s flowers. Figure 1 shows a pollinator wasp entering a syconium of F.
rubiginosa.
 Figure 1. At the top left of this photo a female pollinator fig wasp, Pleistodontes imperialis, can be
seen entering the syconium of Ficus rubiginosa via the ostiole, the opening at the apical tip of the
syconium (this syconium is approximately 5 mm in diameter). The wings of another pollinator are
sticking out of the ostiole – the ostiole is extremely tight and the pollinators’ wings and antennae are
damaged or torn off completely as they squeeze in through the entrance of the syconium. At the lower
right of the photo a parasitoid of the pollinator, Philotrypesis sp. (actually, Philotrypesis may be an
inquiline rather than a parasitoid, or possibly both (look up these terms)), appears to be waiting for the
pollinator to lay her eggs before she too lays her own eggs with those of the pollinator. You can see
the long ovipositor of Philotripesis which she inserts through the wall of the fig to lay her eggs from the
outside, rather than having to enter the syconium. Thus she is able to lay her eggs in many fig
syconia and even several different trees, whereas the pollinator lays her eggs in only one syconium.

The pollination system in figs is an example of a ‘nursery’ pollination system, so-called

because the pollinator’s larvae develop within the same inflorescences/infructescences of the
plant species that they pollinate. See Fig. 2 for a generalised life-cycle diagram of a fig and
its pollinating fig wasp.

Figure 2. A generalized life-cycle diagram of a monoecious fig and its pollinator fig wasp. Image from
Britannica.com
BTW if you eat fresh Australian-grown fruit of the commercially-grown fig (Ficus carica)
they are unlikely to contain any dead wasps because the pollinator is not in Australia. If you
eat dried figs imported from overseas, though, they are likely to originate from areas where
the pollinator is around, so your figs are likely to contain at least some dead fig wasps – extra
protein! However, most fig wasps leave the fruit before it ripens.

As well as looking at the plant-pollinator interaction you will also be examining the impacts
on this mutualism by some of the other members of the wasp community associated with this
species of fig, the parasitoids and hyperparasitoids of the pollinator (parasitoids are parasites
that ultimately kill their host; hyperparasitoids are parasitoids that parasitise and kill the
parasitoids). Thus you will be looking at a predator(parasitoid)-prey interaction as well as a
mutualism interaction in this community.

Our study species for this assignment, the Port Jackson Fig, Ficus rubiginosa, is a
monoecious fig that can grow to 30 metres tall by 30 metres wide. This fig occurs naturally
from the south-east corner of NSW to Cape York in north Queensland and up to 400 km
inland in both states. Our data was collected from the New England region and western
slopes of NSW as far west as Mt Kaputar and the Warrumbungles, the western extreme of the
species’ range i.e., we collected our fig–mutualism data from the western side of the Great
Dividing Range in the drier part of F. rubiginosa’s distributional range. We monitored fig
trees for flowering and fruiting and collected data at 24 sites; 8 sites with large fig
populations of more than 50 trees, 8 sites with small populations of between 5 and 15 trees
and 8 sites with single, relatively isolated trees. Satellite images (Google Earth) of examples
of these three population categories are provided below in Fig. 3.

Data collection occurred from January 2014 to August 2016. Fig wasps and seed numbers
were determined by collecting samples of ripening fruit as they became available in
monitored trees, for later dissection and counting of the contents under a microscope. Note
that the asynchronicity of flowering and fruiting in F. rubiginosa meant that ripening fruit
became available on different trees at different times of the year, throughout the year.
Collecting fruit at just the right time, as they were ripening but before the fig wasps had
departed, required monthly monitoring of the trees at all 24 sites throughout the data-
collection period of this study. Determining ‘just the right time’ to collect fruit was a matter
of squeezing likely-looking (yellow) fruit to see if they were beginning to soften. They soften
because they swell and develop a hollow centre, or lumen, into which the wasps emerge from
their galled flowers and into which the male fig flowers open and release their pollen (see
Fig’s 2 and 4). Soft, yellow fruit were then examined with a hand lens to ensure no escape
tunnels had been dug for departing fig wasps by the male pollinator fig wasps (Fig. 5). Male
pollinator wasps are blind (or almost so) and wingless. After mating with the females they dig
tunnels through the wall of the fig syconium for the females to make their escape. Some
males also exit through these tunnels but most of them die inside the dark confines of the
ripening syconium).
Figure 3. Satellite images (Google Earth) of examples of the three population-size categories: single
trees, small populations and large populations. The yellow pins are marking F. rubiginosa trees.

Figure 4. Dissected fruit of Ficus rubiginosa: fruit a is at the mid stage of development (‘C’ stage (see
Fig. 8)). Fruit b is at early ‘D’ stage when a hollow centre is beginning to develop within the swelling
fruit. Fruit c is at mid D stage when the hollow centre has developed fully and wasps are emerging
into the hollow from their galled flowers lining the inner wall of the fruit. These images are all at the
same scale and fruit b is 12 mm in diameter.
Figure 5. A ripening Ficus rubiginosa syconium with an escape tunnel (marked with a red arrow)
which has been dug by a male pollinator wasp.

Fruit collected at just the right time, before escape tunnels had been dug, thus still contained
all the fig wasps that had developed within them. These fruit were measured (length and
width; Fig. 6), to determine their volumes, and then placed within individual vials. Vials of
fruit were frozen back in the lab for later dissection and counting of their seed and wasp
contents. A sample of these fruit was dissected and the contents (seeds, unpollinated flowers,
male flowers, and various species of fig wasps) scraped out, sorted and counted. Analysis of
early counts showed no statistical difference between mean numbers of wasps or seed of
three, five or ten fruits from each tree, so all future counts were made using samples of three
fruit per tree.

Figure 6. Immediately upon collection, the length and diameter of syconia were measured, using
callipers, and the data was entered into a field book.

Syconia in this species grow in pairs in leaf axils at the ends of branches, with each branch tip
usually carrying six to twelve fruit in the final three to six leaf axils (up to 20 pairs of fruit or
more in some instances) (Fig. 7).
Figure 7. Branch of Ficus rubiginosa with red, ripe and yellow, ripening syconia. Fig wasps emerge
from syconia at the yellow, ripening stage of syconia development. Note that this branch was from a
tree on the coast near Grafton where, usually, more fruit per branch were produced and pollinated
than in the western, drier region where this study was conducted.

Syconium development in monoecious figs goes through five recognised stages, named A to
E, described by Galil and Eisikowitch (1968). These five stages are shown and described
below in Fig. 8 for F. rubiginosa.

Figure 8. Five recognised stages of syconial development: A = bud stage; B = female-flowering

stage, which is when the female flowers open within the syconium and the ‘foundress’ pollinator
wasps enter the syconium through the ostiole, the minute opening at the tip of the syconium, to
pollinate as many female flowers as they can and lay as many eggs as they can; these foundress
wasps then die inside the syconium; C = the stage during which seeds develop within the pollinated
flowers and wasp larvae develop within the galled flowers, all within the growing syconium, which
remains green throughout this stage; D = male-flowering stage, when wasps emerge from their galls,
mate, collect pollen from the male flowers and then exit the syconium through tunnels dug by the
male pollinator wasps (which are blind and wingless and die soon afterwards, usually within the
syconium); E = ripe stage, when the syconium softens and becomes red (in this species) and sweet to
attract frugivores. Scale bars = 10 mm.
Syconium development usually occurs synchronously within trees with only minimal overlap
between adjoining developmental stages. Female flowering (B stage; Fig. 8) and male
flowering (D stage; Fig. 8) are thus temporally separated within trees as well as within
syconia, to reduce the chances of inbreeding.
There are large and diverse communities of fig wasps and other organisms, from nematodes
to birds and mammals, associated with figs of all species. We will concentrate on just a sub-
set of the communities of wasps associated with Ficus rubiginosa. First, let’s look at the
pollinator – the pollinators of figs are wasps in the family Agaonidae. These particular ‘fig
wasps’ (there are many other fig wasps than just the pollinator) are extremely specialised for
a mutualistic association with figs. The pollinator of F. rubiginosa is Pleistodontes
imperialis. Female P. imperialis wasps are about 1.5 mm long and live for less than 2 days
(Dunn et al. 2008). In this short period of a day or two they mate with males within the
confines of the fig syconium in which they developed, they collect pollen from the male
flowers of the fig (that open at the same time as the wasps are emerging from the galled
female flowers in which they developed) – they stuff this pollen into pockets on their
thoraxes, they escape from the fig syconium via tunnels dug through the wall of the syconium
by male P. imperialis wasps and then fly off in search of a fig tree of the same species that
has syconia at B stage (see Fig. 8) with open female flowers inside. Research has shown that
fig wasps are capable of flying extraordinary distances in their short adult life-times, of up to
160 km or more between trees (Ahmed et al 2009).
However, this is a very chancy business. Ficus rubiginosa trees flower and fruit
asynchronously throughout the year and the female-flowering ‘B’ stage can last for as little as
a week or two within a tree. This means that relatively few fig trees will be in female-
flowering stage at any one time. In order to cover as much ground as possible in their search
for a tree with female flowers the female fig wasps leave their maternal tree by flying up
above the canopy to catch any prevailing breeze which then carries them off in whatever
direction the wind is blowing (Compton et al., 2000). So the wasps are dependent on there
being a fig tree of the same species producing female flowers in the direction in which they
are travelling, within a flight distance achievable within their short life-times. When humidity
levels are low the wasps’ life times can be less than a day (Dunn et al. 2008). From up in the
air column where these wasps are being blown across the landscape they are able to sense
particular chemicals given off by F. rubiginosa trees with female flowers. When they pick up
these chemicals they drop out of the air towards the trees below them, navigate towards the
particular fig tree giving off the female-flower chemical signature, locate a B-stage syconium
in that tree and enter the syconium to deliver their pollen load and lay their eggs. They then
die within the syconium (their wings are usually torn off as they enter through the very tight
ostiole anyway, so even if they were able to get out of the syconium they wouldn’t be able to
fly away). During favourable weather conditions – warm, humid weather down on the
northern NSW coast for example – they have about a 4% success rate in making this journey
to pollinate a fig syconium and lay their eggs. In poorer conditions –during relatively dry
weather on the Western Slopes of northern NSW for example – they have only a 1% success
rate or less. One thing we want to find out from this study is whether fig population-size
impacts on this success rate.
Aims of the study
So, the aims of this study, that you are doing and writing up as a scientific paper, are to (a)
determine whether the size of F. rubiginosa populations has any impact on the ability of
pollinators to locate trees with female flowers to lay their eggs and pollinate the fig’s female
flowers i.e., does F. rubiginosa population size impact on pollinator visitation rates and, if so,
what impact does this have on pollinator and/or fig reproductive success? and (b) determine
whether the size of F. rubiginosa populations impacts on the ability of parasitoids to locate
trees with pollinated female flowers to parasitise pollinator larvae i.e., does F. rubiginosa
population size impact on the number of pollinator larvae killed and replaced by parasitoids
and, if so, what impact does this have on pollinator and/or fig reproductive success?
To answer these questions you will need to analyse the data provided in the Resources – wasp
and seed data.
DO NOT write your aims exactly as I have just done! Write them in your own words.
Turnitin is very good at picking up plagiarism and you will be penalised for it. This applies to
the whole assignment and all your work at UNE. Do not copy the work of others. A lot of
what you write will necessarily be flagged by Turnitin simply because you will be using
phrases and jargon associated with this study that also appear elsewhere in the literature.
Don’t flinch from using the appropriate jargon but just be sure to write the bulk of your
manuscript in your own words.
The above information on figs in general and Ficus rubiginosa in particular is not all relevant
for your own introduction. A lot of it is provided simply to help you understand the system
being studied, which you are now going to write about. Remember that I am available to
answer questions via the general forum for this unit or by email (kmackay5@une.edu.au).
Remember to conclude your introduction with clearly and precisely stated aims.

Your methods section should describe the species you looked at, where the study was
conducted, how you conducted the study and collected the data, and how you analysed the
data. You should be able to write up the data-collection methods for this manuscript based on
the information provided above in ‘background reading’ but, if in doubt, ask other students or
me. Remember to also write up how you analysed this data. The methods section must
describe what you did and how you did it, so that the reader would be able to replicate this
study exactly as you did it.

I expect you to conduct analyses of variance (ANOVAs) to determine if (1) fig population
size had any effect on the total number of pollinator fig wasps that emerged from syconia,
and (2) if fig population size had any effect on the number of parasitoids and hyperparasitoids
of the pollinator fig wasps that emerged from syconia (i.e., the number of fig wasp larvae that
were killed by parasitoids – each parasitoid or hyperparasitoid represents one dead
pollinator). I expect you to present the results of these analyses (means and standard errors)
in graphs. You can use whatever software you like for this but if you don’t already use R I
suggest you learn to do so. I will provide a separate tutorial on using R that will explain
clearly how to do these particular analyses and graph the results. You will also need to
analyse, through an ANOVA, whether fig population size impacts on the number of
pollinator foundresses per syconium but the results of this analysis can be presented in the
text rather than graphically.

I expect you to use R to fit a General Linear Model to the data for the number of pollinators
that develop to maturity within a syconium (the dependent variable), with the number of
foundress pollinators per syconium being the independent variable. I don’t expect the fitted
model to be presented graphically but I do expect you to give the model formula in the
results text. Again, refer to the tutorial in R in the Assessment tile.
For the results section I don’t want the datasets that I am sending to you presented in the
paper. These would normally not be presented in a scientific paper. I want your analyses of
the data presented in text and graphs.

You should begin your discussion section by stating whether your results support your
hypotheses or not i.e., you should state how your results relate to the aims of your study and
whether they answered your question/s. You should then interpret your results and discuss
their relevance in relation to appropriate research findings by other scientists in the published
literature. You should also discuss the implications of your findings in the context of what
influences may lead to small population fragments or isolated trees in the landscape e.g.,
anthropogenic influences such as vegetation clearing for development or agriculture can lead
to vegetation being reduced to smaller fragments with greater distances between such
patches; dispersal by F. rubiginosa may lead to expansion of the species’ range into new
areas by individual, isolated trees; but don’t dwell too much on conjecture and be sure to
make it clear that this is all it is, and use it in terms of describing the relevance of your
findings. The main discussion points you should concentrate on should be those that reveal
the meaning of your results and how they compare with the results in other published work,
why your own findings may agree with or differ from the findings in the literature (justify
your own interpretation if you have a different finding or interpretation). End your discussion
with a concluding paragraph.

Note: There have been many studies of interactions between the pollinators of Ficus species
and other wasps using the same Ficus syconia for nurseries, including the pollinators’
parasitoids. Correlations show a negative effect of parasitoids on pollinator numbers (e.g.,
Segar and Cook 2012), as one would expect because the parasitoids kill their hosts, but care
needs to be taken when interpreting the results of such studies. Many predator-prey studies
show a positive correlation between predator numbers and prey numbers, as was found in this
study (Mackay, unpublished data), but this isn’t unexpected either because as prey numbers
go up, so do predator numbers. Think carefully about the implications of your own findings.
Each parasitoid that develops within a syconium was responsible for the death of a single
pollinator-wasp larva.

The discussion section can be the most important part of a paper. Read some papers, like the
ones in the following ‘useful references’ list, and get some ideas about how you should write
your own discussion from these papers.

Lastly, write your abstract at the beginning of your paper - this should include the main
points from your introduction, methods, results and discussion, finishing with a statement that
clearly relates your results/conclusions to your stated aim/s. No references are cited in the
abstract. Often readers will only read the abstract of published papers and only read further if
the abstract draws them in.

See the ‘Resources – marking rubric for Assignment 2’ in the Assessment tile for more
details.

See the ‘Resources – analyses using R’ in the Assignment tile for help with using R.

In addition, I will write a short tutorial on scientific writing soon and add it to the
resources for this assignment.
The broader community of organisms associated with F. rubiginosa

Note that the dataset provided is a cut-down version of a much larger data set that included
data on other wasps in this community – we found 15 species of fig wasps who’s larvae
developed within the syconia of F. rubiginosa: the pollinator (Pleistodontes imperialis), two
species of Sycoscapter and two species of Philotrypesis (parasitoids (or inquilines) of the
pollinator), one species of Watshamiella (a hyperparasitoid of the pollinator), one species of
Eukobelea and one species of Pseudidarnes, P. minerva, (these species are all ovule gallers,
like the pollinator), one species of Herodotia and one (possibly two) species of Meselatus
(both of these wasps species make their galls in the wall tissue of the syconium), one species
each of Ormyrus, Eurytoma, Sycophila and Megastigmus (all probably parasitoids of the wall
gallers) and an unknown species belonging to an unknown trophic level or functional group.
All of the species other than the ones we are investigating occurred in very small numbers –
usually zero per syconium, sometimes one and rarely two or more per syconium. Only
Eukobelea and Pseudidarnes could potentially impact on the number of pollinators
developing within a syconium, and then only indirectly through competition for female
flowers in which to develop. Because the numbers of Eukobelea sp. and Pseudidarnes
minerva wasps were so low and because the numbers of female flowers far exceeded the
numbers of developing pollinating wasps, the impact of these other wasp species on
pollinator numbers was considered not to be significant or even detectable in this study, so
they have been left out of our analyses.

The community of organisms associated with F. rubiginosa is broader than just the wasps
that develop within the syconia. For example, there are frugivorous birds that feed on the ripe
fruit and potentially disperse the seed of this fig. We found 27 avian frugivore species in just
the relatively small area of 50,000 km2 where this study was conducted (Mackay et al. 2019;
there are likely to be considerably more species than this, keeping in mind that the range of F.
rubiginosa covers approximately 3,000 x 400 km including coastal and tropical regions not
examined in this study ). Other plant species that grow in dry rainforest patches along with F.
rubiginosa potentially rely on this fig, with its large crops of fleshy fruit, to attract these avian
frugivores to disperse their fruit as well. We also observed 55 species of avian insectivores
that fed on the fig wasps in these fig trees (Mackay et al. 2018; and again, there are likely to
be more insectivore species than this over the geographic range of F. rubiginosa). Other birds
used the dense canopies of these trees to hide their nests in or simply as places to hide from
predators, to rest and/or preen. Mammals including pigs and goats feed on the fruit of F.
rubiginosa, so this fig potentially helps to support populations of these feral pest species.
Many other insects were observed in the canopies of F. rubiginosa trees during this study.
The European Honeybee, Apis mellifera, for example, was often observed gathering resin
from wounds on the branches of the fig trees, presumably to be used in their hives for some
purpose. Sheep and cattle were observed to often use the trees for shelter; in turn, the trees
under which these animals sheltered benefitted from the extra nutrients provided in their
dung.

Useful references:
Ahmed, S., S. G. Compton, R. K. Butlin, and P. M. Gilmartin. 2009. Wind-borne insects
mediate directional pollen transfer between desert fig trees 160 kilometers apart. Proceedings
of the National Academy of Sciences of the United States of America 106: 20342-20347.

Compton, S. G., Ellwood, M. D., Davis, A. J., & Welch, K. (2000). The Flight Heights of
Chalcid Wasps (Hymenoptera, Chalcidoidea) in a Lowland Bornean Rain Forest: Fig Wasps
are the High Fliers 1. Biotropica, 32(3), 515-522.

Dunn, D. W., D. W. Yu, J. Ridley, and J. M. Cook. 2008. Longevity, early emergence and
body size in a pollinating fig wasp – implications for stability in a fig–pollinator mutualism.
Journal of Animal Ecology 77: 927-935.

Galil, J., and D. Eisikowitch. 1968. Flowering cycles and fruit types of Ficus sycomorus in
Israel. New Phytologist 67: 745-758.

Gates, D. J., and J. D. Nason. 2012. Flowering asynchrony and mating system effects on
reproductive assurance and mutualism persistence in fragmented fig-fig wasp populations.
American Journal of Botany 99: 757-768.

Janzen, D. H. (1979). How to be a fig. Annual review of ecology and systematics, 10(1), 13-
51.

Kong, Y., R. Wang, D.-R. Yang, R. Sreekar, Y.-Q. Peng, and S. G. Compton. 2016. Non-
pollinator fig wasp impact on the reproductive success of an invasive fig tree: why so little?
Biocontrol Science and Technology 26: 1432-1443.

Mackay, K. D., Gross, C. L., & Rossetto, M. (2018). Small populations of fig trees offer a
keystone food resource and conservation benefits for declining insectivorous birds. Global
ecology and conservation, 14, e00403.

Mackay, K. D., & Gross, C. L. (2019). Climate Change Threatens a Fig-Frugivore Mutualism
at its Drier, Western Range Margin. Proceedings of the Linnean Society of New South
Wales (Vol. 141).

Segar, S. T., and J. M. Cook. 2012. The dominant exploiters of the fig/pollinator mutualism
vary across continents, but their costs fall consistently on the male reproductive function of
figs. Ecological Entomology 37: 342-349.

Shanahan, M., S. So, S. G. Compton, and R. Corlett. 2001. Fig-eating by vertebrate

frugivores: a global review. Biological Reviews 76: 529-572.