The latest paper from Paolo Biella‘s PhD work, on which I collaborated and that I’ve discussed before on the blog, has just been published in the journal Functional Ecology. It’s entitled “An empirical attack tolerance test alters the structure and species richness of plant–pollinator networks“. The paper presents more of Paolo’s work showing how the experimental removal of the floral resources provided by the more generalised plants in a community can significantly (and negatively) affect the patterns of interaction between flowers and pollinators that we observe. It’s another piece of evidence that demonstrates how important it is to not neglect the common plants that attract a lot of flower visitors when considering how to manage a habitat.
If anyone has trouble accessing the PDF, drop me a line and I will send it to you.
Here’s the reference:
Biella, P., Akter, A., Ollerton, J., Nielsen, A. & Klecka, J. (2020) An empirical attack tolerance test alters the structure and species richness of plant-pollinator networks. Functional Ecology DOI: 10.1111/1365-2435.13642
Here’s the abstract:
Ecological network theory hypothesizes that the structuring of species interactions can convey stability to the system. Investigating how these structures react to species loss is fundamental for understanding network disassembly or their robustness. However, this topic has mainly been studied in‐silico so far.
Here, in an experimental manipulation, we sequentially removed four generalist plants from real plant–pollinator networks. We explored the effects on, and drivers of, species and interaction disappearance, network structure and interaction rewiring. First, we compared both the local extinctions of species and interactions and the observed network indices with those expected from three co‐extinction models. Second, we investigated the trends in network indices and rewiring rate after plant removal and the pollinator tendency at establishing novel links in relation to their proportional visitation to the removed plants. Furthermore, we explored the underlying drivers of network assembly with probability matrices based on ecological traits.
Our results indicate that the cumulative local extinctions of species and interactions increased faster with generalist plant loss than what was expected by co‐extinction models, which predicted the survival or disappearance of many species incorrectly, and the observed network indices were lowly correlated to those predicted by co‐extinction models. Furthermore, the real networks reacted in complex ways to plant removal. First, network nestedness decreased and modularity increased. Second, although species abundance was a main assembly rule, opportunistic random interactions and structural unpredictability emerged as plants were removed. Both these reactions could indicate network instability and fragility. Other results showed network reorganization, as rewiring rate was high and asymmetries between network levels emerged as plants increased their centrality. Moreover, the generalist pollinators that had frequently visited both the plants targeted of removal and the non‐target plants tended to establish novel links more than who either had only visited the removal plants or avoided to do so.
With the experimental manipulation of real networks, our study shows that despite their reorganizational ability, plant–pollinator networks changed towards a more fragile state when generalist plants are lost.
2 More than just bees: the diversity of pollinators
3 To be a flower
4 Fidelity and promiscuity in Darwin’s entangled bank
5 The evolution of pollination strategies
6 A matter of time: from daily cycles to climate change
7 Agricultural perspectives
8 Urban environments
9 The significance of gardens
10 The shifting fates of pollinators
11 New bees on the block
12 Managing, restoring and connecting habitats
13 The politics of pollination
14 Studying pollinators and pollination
As you can see it’s a very wide-ranging overview of the subject, and written to be accessible to both specialists and non-specialists alike. To quote what I wrote in the Preface:
“While the book is aimed at a very broad audience, and is intended to be comprehensible to anyone with an interest in science and the environment, and their intersection with human societies, I hope it will also be of interest to those dealing professionally with plants and pollinators. The subject is vast, and those working on bee or hoverfly biology, for example, or plant reproductive ecology, may learn something new about topics adjacent to their specialisms. I certainly learned a lot from writing the book.”
The book is about 100,000 words in length, lots of illustrations, and there will be an index. My copy editor reckons there’s 450 references cited, though I haven’t counted. I do know that they run to 28 pages in the manuscript, and that’s with 11pt text. All going well it will be published before Christmas.
The Wildlife Trust for Bedfordshire, Cambridgeshire and Northamptonshire has invited me to run my Introduction to Pollinators and Pollination workshop again this year, but of course it will all be online. Details for signing up are on the images, or you can follow this link.
Here’s a description of the workshop:
Pollination of flowers ensures the reproduction of most British wild plants and many of our agricultural crops. This session will provide an introduction to the natural history of pollinators and how they interact with the flowers that they pollinate. The main groups of pollinators will be introduced, with guidance on how to identify them, and their ecology and behaviour will be explored. The session will also consider why conserving these species is so important, followed by a Q and A discussion showing what individuals can do to help ensure their future diversity and abundance.
This year’s Missouri Botanical Garden/St Louis University John Dwyer Public Lecture in Biology will be given by Alan Mosswho researches Himalayan bumblebees and their interactions with flowers. The lecture is being live-streamed on YouTube – details are in the flyer above.
As I near completion of the copy-editing phase of my forthcoming book it’s frustrating to see all of the great research that’s been produced in recent weeks that I probably won’t be able to cite! Here’s a few things that caught my eye:
In the journal New Phytologist, Rhiannon Dalrymple and colleagues, including Angela Moles who hosted me during my recent stay in Australia, have a great study entitled Macroecological patterns in ﬂower colour are shaped by both biotic and abiotic factors. The title pretty much sums it up: in order to fully understand how flowers evolve we need to consider more than just their interactions with pollinators. It’s another demonstration of how we must look beyond simplistic ideas about pollination syndromes to fully understand the complexities of the relationship between flowering plants and pollinators…..
…..talking of which, again in New Phytologist, Agnes Dellinger asks: Pollinationsyndromes in the 21st century: where do we stand and where may we go? It’s an insightful and far-reaching review of a topic that has intrigued me for more than 25 years. There are still a lot of questions that need to be asked about a conceptual framework that, up until the 1990s, most people in ecology and biology accepted rather uncritically. One of the main unanswered questions for me is how further study of largely unexplored floras will reveal the existence of new pollination systems/syndromes. Which leads nicely to….
…..an amazing paper in Nature this week by Rodrigo Cámara-Leret etal. showing that New Guinea has the world’s richest island flora. The described flora includes 13,634 plant species, 68% of which are endemic to New Guinea! And the description of new species each year is not leveling off, there’s still more to be discovered. A commentary on the paper by Vojtech Novotny and Kenneth Molem sets some wider context to the work, and quite a number of media outlets have covered the story. Why is this relevant to pollinators and pollination? Well, we actually know very little about this critical aspect of the ecology of the island: there’s only a handful of published studies of plant-pollinator interactions from New Guinea, mostly focused on figs, bird-flower interactions, and a couple of crops. For such a biodiverse part of the world that’s a big gap in our understanding.
Finally, James Reilly, Rachael Winfree and colleagues have a paper in Proceedings of the Royal Society series B showing that: Crop production in the USA is frequently limited by a lack of pollinators. Most significant findings to me were that of the seven crops studied, five of them have their yields limited by lack of pollinators, and that even in areas of highly intensive farming, wild bees provided as much pollination service as honeybees.
That’s a few of the things that I spotted this week; what have you seen that’s excited or intrigued you? Feel free to comment.
In the next few months my new book Pollinators & Pollination: Nature and Society will be published. As you can imagine, I’m very excited! The book is currently available to pre-order: you can find full details here at the Pelagic Publishing website. If you do pre-order it you can claim a 30% discount by using the pre-publication offer code POLLINATOR.
As with my blog, the book is aimed at a very broad audience including the interested public, gardeners, conservationists, and scientists working in the various sub-fields of pollinator and pollination research. The chapter titles are as follows:
Preface and Acknowledgements
1. The importance of pollinators and pollination
2. More than just bees: the diversity of pollinators
3. To be a flower
4. Fidelity and promiscuity in Darwin’s entangled bank
5. The evolution of pollination strategies
6. A matter of time: from daily cycles to climate change
7. Agricultural perspectives
8. Urban environments
9. The significance of gardens
10. Shifting fates of pollinators
11. New bees on the block
12. Managing, restoring and connecting habitats
13. The politics of pollination
14. Studying pollinators and pollination
UPDATE: turns out the figure I cited for number of bee species is out of date so I’ve corrected it below. Thanks to John Ascher for pointing this out.
Publication of my book Pollinators & Pollination: Nature and Society by Pelagic Publishing has been pushed back until the end of this year or early in 2021. The current pandemic has created problems for the printing and distribution sectors, as it has for so many industries. Therefore, to celebrate World Bee Day, here’s a preview of the bee section from Chapter 2 which is entitled (ironically enough) “More than just bees – the diversity of pollinators”.
2.3 Bees, wasps and sawflies (Hymenoptera)
The bees and their relatives rank only third in terms of overall pollinator diversity. Within this taxonomic Order, bees are not especially species rich (17,000 or so described species, perhaps 20,000 in total) – over 20,400 (see: https://www.catalogueoflife.org/col/details/database/id/67) compared with the other 50,000 social and solitary wasps, sawflies, and so forth. But what they lack in diversity the bees make up for in importance as pollinators of both wild and agricultural plants, and in their cultural significance. The general notion of what a bee is, and how it behaves, looks to the honeybee (Apis mellifera) as a model: social, with a hierarchy, a queen, and a large nest (termed a hive for colonies in captivity). In fact, this view of bee-ness, though long embedded within our psyche, is far removed from the biology of the average bee: most of them have no social structure at all, and a fair proportion of those are parasitic. In Britain we have about 270 species of bees, give or take (Falk 2015) though there have been extinctions and additions to this fauna (see Chapters 10 and 11). These species provide a reasonable sample of the different lifestyles adopted by bees globally. They can be divided into four broad groups.
Honeybees include several highly social species and subspecies of Apis, of which the ubiquitous western honeybee (A. mellifera) is the most familiar. Most colonies are found in managed hives, though persistent feral colonies can be found in hollow trees, wall cavities, and other suitable spaces. They are widely introduced into parts of the world where they are not native (e.g. the Americas, Australia, New Zealand) and there is some debate as to whether they are truly native to Britain and northern Europe, with supporting evidence and arguments on both sides. Colonies can be enormous and contain thousands of individuals, mostly female workers, with a single queen. Unmated queens and males (drones) are produced by the colony later in the season.
Bumblebees (Bombus spp.) are typically also social, though their nests are much smaller (tens to hundreds of individuals). Depending upon the species these nests can be in long grass, rodent holes, or cavities in buildings and trees. Twenty-seven of the more than 250 species have been recorded in the UK, but six of these are not strictly social; they are parasitic and belong to the subgenus Psithyrus which will be described below.
The so-called solitary bees are by far the largest group in Britain (about 170 species) and worldwide (more than 90% of all species). In the UK they belong to 15 genera, including Andrena, Anthophora, Osmia, Megachile, etc. The females of most of these bees, once they have mated, construct nests that they alone provision with pollen for their developing young. Nesting sites can be genus- or species-specific, and include soil, cavities in stone or wood, and snail shells. Some species are not strictly solitary at all and may produce colonies with varying levels of social structure, though without a queen or a strict caste system; we term them “primitively eusocial”. In fact sociality has evolved and been lost numerous times in the bees and in the rest of the Hymenoptera (Danforth 2002, Hughes et al. 2008, Danforth et al. 2019). It’s also been lost in some groups that have reverted back to a solitary lifestyle, and even within a single genus it can vary; for example in the carpenter bee genus Ceratina (Apidae: Xylocopinae) tropical species are more often social than temperate species (Groom & Rehan 2018).
The final group is termed the cuckoo bees and, like their avian namesake, they parasitise the nests of both social and solitary bees (though never, interestingly, honeybees). There are about 70 species in 7 genera, including the bumblebee subgenus, Psithyrus. Other genera include Melecta, Nomada and Sphecodes. In some cases the parasitic species are closely related evolutionarily to their hosts and may resemble them, for example some Psithyrus species. In other cases they may be only distantly related and in fact look more like wasps, e.g. Nomada species. Some genera of cuckoo bees are restricted to parasitising only a single genus of bees, others are parasites of a range of genera (Figure 2.4).
Although we often think of bees, overall, as being the most important pollinators, in fact species vary hugely in their importance. Pollinating ability depends upon factors such as abundance, hairiness, behaviour, body size, and visitation rate to flowers (Figure 2.1). Size is especially important for three reasons. First of all, larger animals can pick up more pollen on their bodies, all other things being equal. Secondly, in order to bridge the gap between picking up pollen and depositing it, flower visitors must be at least as large as the distance between anthers and stigma, unless they visit the stigma for other reasons. Finally, larger bee species tend to forage over longer distances on average (Greenleaf et al. 2007) thus increasing the movement of pollen between plants. However, most of the world’s bees are relatively small as we can see from the analysis of British bees in Figure 2.5. Many species have a maximum forewing length of only 4 or 5 mm, and the majority of species are smaller than honeybees. Remember also that these are maximum sizes measured from a sample; individual bees can vary a lot within populations and even (in the case of Bombus spp.) within nests (Goulson et al. 2002). So the assumption that all bees are good pollinators needs to be tempered by an acknowledgement that some are much better than others.
Figure 2.5: The sizes of British bees. Forewing length is a good measure of overall body size and the data are maximum lengths recorded for species, except for the social bumblebees and honeybee I have used maximum size of workers (queens are often much larger). The blue line indicates the honeybee (Apis mellifera). The biggest bee in this data set is the Violet Carpenter Bee (Xylocopa violacea) which, whilst not generally considered a native species (yet), has bred in Britain in the past. Data taken from Falk (2015).
Last week, during one of my lockdown garden pollinator surveys, I spotted a bee visiting Germander Speedwell (Veronica chamaedrys) in the garden that I didn’t recognise. It initially confused me as it looked superficially like a Blood Bee in the genus Sphecodes. However the bee was clearly collecting pollen, which Sphecodes spp., being cleptoparasites, don’t do. A quick check in Steven Falk’s Field Guide to the Bees of GreatBritain and Ireland and a look at Steven’s Flickr site, suggested that it was almost certainly the Red-girdled Mining Bee (Andrena labiata), which is frequently associated with Germander Speedwell.
I posted this video on Twitter and Steven kindly confirmed my identification:
The Red-girdled Mining Bee is considered “Nationally Scarce” and it has a scattered and southerly distribution, as you can see from the map above, which is from the National Biodiversity Network Atlas account for the species. It’s only recorded from about half a dozen sites in Northamptonshire according to Ryan Clark, the County Bee Recorder. However Steven tells me that it’s being seen more and more frequently in gardens, and indeed just the other day Sarah Arnold, who is also carrying out surveys, emailed me to say that she had spotted it in her garden in Kent.
So this is a bee that’s definitely one to look out for, especially if you have Germander Speedwell growing.