Tag Archives: Science

How much do we really understand about pollination syndromes?

Ecologists and evolutionary biologists have, for many years, sought to document repeated patterns that they see in nature; to understand the processes that determine these patterns; and to make predictions about how and when they are going to be observed in the future or in other parts of the world. There are many examples of such patterns, including: cyclical population dynamics of species such as lemmings; the occurrence of specific types of plant communities (e.g. rainforest, grasslands) in areas with particular climates; and convergent evolution of unrelated species to similar ecological niches, such as large, predatory placental and marsupial mammals (e.g. the dog and wolf family compared to the Tasmanian “wolf”).

An example of convergent evolution that has fascinated botanists since the 19^th century is the idea of “pollination syndromes”, which are sets of flower characteristics that have repeatedly evolved in different plant families due to the convergent selection pressures applied by some groups of pollinators. Thus, red, scentless flowers producing lots of nectar are typical of many hummingbird pollinated plants in the New World, whilst white, night-scented flowers often signify moth pollination. Good examples of plant species possessing these archetypical flower traits are have been used as text book examples for decades, repeatedly used to illustrate the predictable and specialised nature of some plant-pollinator interactions.

The problem is that until recently the pollination syndromes have rarely been subjected to critical tests of their frequency and predictive value (Ollerton et al. 2009 and references therein). It’s been tacitly assumed that (after more than 150 years of study) we clearly know all there is to know about them, even though there have been criticisms levelled at the syndromes since their inception, a fact that has been subsequently ignored (Waser et al. 2011).

However in the last 20 years biologists have begun to seek answers to questions such as: How often do plant species conform to the expectations of the classical pollination syndromes? How good is our ability to predict the pollinators of a plant based just on its flower characteristics? What is the role played by flower visitors that do not conform to the predictions of the pollination syndromes? Similarly, what is the role of animals that steal nectar or pollen, or act as herbivores, in shaping flower traits? What new examples of convergent evolution of flower traits remain to be discovered?

Research conducted in many different parts of the world has addressed these questions, questions which some biologists had assumed were already answered or which were not worth asking in the first place. And the answers to them are proving to be both surprising and controversial.

For example, the most comprehensive test of the frequency and predictability of pollination syndromes that has been conducted to date (Ollerton et al. 2009) concluded that only a small proportion of the 352,000 species of flowering plants could be categorised into the pollination syndromes as classically described. Likewise, they estimated that the predictive power of the pollination syndromes was about 30%. Other studies have shown that “secondary” flower visitors can be just as, or more, effective pollinators than the “primary” pollinator predicted by the syndromes (e.g. Waser & Price 1981,1990, 1991); that floral antagonists can play an important a role in shaping flower traits (e.g. Junker and Parachnowitsch 2015 and references therein); and that there are still examples of convergent evolution to “unexpected” pollinators waiting to be discovered in less well researched parts of the world, which in fact is most of the world (Ollerton et al. 2003).

Recently the very prestigious journal Ecology Letters published a paper that has challenged the challengers. Rosas-Guerrero et al (2014), by using a statistical technique called meta-analysis underpinned by a review of the available literature, suggested that pollination syndromes are much more predictable than Ollerton et al. (2009) concluded, and perhaps as high as 75%. However some of my collaborators and I see problems with their approach to studying pollination syndromes that have biased the conclusions that they draw, and therefore undermined the robustness of those conclusions, which we set out in a response to their original paper (Ollerton et al. 2015). We originally tried to publish this in Ecology Letters but for some reason the journal was not interested; it’s therefore freely available from Journal of Pollination Ecology if you follow that link.

I won’t go into the detail of what we perceive as problems in Rosas-Guerrero et al.’s approach to testing the syndromes (you can read the paper for yourself) but in summary they relate to how the literature review was conducted (which failed to include all of the studies that could have provided data for their meta-analysis); the significant bias in the current literature because plant-pollinator interactions are not studied randomly (biologists are often drawn to large-flowered plants possessing those archetypical, classical flower traits associated with particular syndromes); the variation in how different researchers determine the effectiveness of the pollinators in their system, meaning that these studies are not always comparable; and issues around annual variation in pollinator identity and presentation of data.

Despite providing a focus and framework for understanding pollination biology for over 150 years, the pollination syndromes continue to surprise us and to provide a vital antidote to scientific hubris: we really do not understand nearly as much about them as we assume.

In an era when we are more and more concerned about loss of pollinator diversity, including extinction at both a species- and country-level, do these debates really matter or are they of purely academic concern, of interest to a few botanists and ecologists? As you might expect, I’d argue that they do matter: there are still some fundamental aspects of pollination ecology that we don’t completely understand, or have only recently been seriously addressing, some of which I’ve worked on myself and which I’ve highlighted in this blog. These include the number of flowering plants that require animal pollination, the diversity of pollinators at a global and regional level, the relative importance of different types of pollinators, and whether or not plants and pollinators are more specialised in tropical compared to temperate communities. Without some of this fundamental knowledge we are unable to make effective arguments, policies and strategies for conserving pollinators.

References

Junker RR, Parachnowitsch AL (2015) Working towards a holistic view on flower traits—how floral scents mediate plant–animal interactions in concert with other floral characters. Journal of the Indian Institute of Science 95:43–67.

Ollerton J, Johnson SD, Cranmer L, Kellie S (2003) The pollination ecology of an assemblage of grassland asclepiads in South Africa. Annals of Botany 92:807–834.

Ollerton J, Alarcón R, Waser NM, Price MV, Watts S, Cranmer L, Hingston A, Peter CI, Rotenberry J (2009) A global test of the pollination syndrome hypothesis. Annals of Botany 103:1471–1480.

Rosas-Guerrero V, Aguilar R, Marten-Rodriguez S, Ashworth L, Lopezaraiza-Mikel M, Bastida JM, Quesada M (2014) A quantitative review of pollination syndromes: do floral traits predict effective pollinators? Ecology Letters 17: 388–400.

Waser NM, Price MV (1981) Pollinator choice and stabilizing selection for flower color in Delphinium nelsonii. Evolution 35:376–390.

Waser NM, Price MV (1990) Pollination efficiency and effectiveness of bumble bees and hummingbirds visiting
Delphinium nelsonii. Collectanea Botanica (Barcelona) 19:9–20.

Waser NM, Price MV (1991) Outcrossing distance effects in Delphinium nelsonii: pollen loads, pollen tubes, and seed set.
Ecology 72:171–179.

Waser NM, Ollerton J, Erhardt A (2011) Typology in pollination biology: lessons from an historical critique. Journal of Pollination
Ecology 3:1–7.

By lifting the restriction on the use of neonicotinoid pesticides Defra throws a (bee) brick at its own National Pollinator Strategy

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Yesterday a brick arrived in the post. Not just any brick, but a Bee Brick, designed by the Green&Blue company in Cornwall as an architectural addition that can provide habitat for cavity nesting solitary species such as the Patchwork leaf-cutter bee that I discussed during Pollinator Awareness week. A representative of the company recently got in touch, after having read my blog, and asked if I’d like a sample to try out in the garden. In the absence of any planned wall building I’ve placed it a couple of metres up on the flat top of a south facing summer house window. It’s probably a bit late in the season to attract any nesting solitary bees this year, but we’ll see; expect a report back from me at some point.

I had actually encountered the Bee Brick earlier this year at the Chelsea Flower Show which Karin and I attended as a 50th year bucket-list day out. It was ok, I enjoyed it, the plants and (some of) the gardens were great. But it was too busy, too expensive and too full of ostentatiously wealthy people for my tastes.

As if to serve as a counter-point to all the good work being done during Pollinator Awareness Week and by companies such as Green&Blue, came the recent news that Defra has agreed to lift the restrictions on use of two neonicotinoid pesticides on oil seed rape across a “limited” area in the east of the country. It will apparently apply mainly to Suffolk, and cover an area of about 30,000 hectares. That’s 5% of the UK’s oil seed rape crop.

The decision was made at the behest of the National Farmers Union, and seems to make no farming sense whatsoever given that nationally yields of oil seed rape have not been affected by the restriction on neonicotinoids, with the harvest this year looking to be above average. Not surprisingly the decision has drawn furious fire from a range of environmental organisations including Buglife and the Wildlife Trusts. Meanwhile Friends of the Earth have threatened legal action, a move prompted by the fact that the Government has refused to allow its independent advisors to publish the details of the decision, including how it was made and what was discussed.

Aside from the lack of transparency, what particularly worries me is that this decision opens the door to further use of these restricted pesticides over the next 12 months, on a region by region basis, until we are back where we were prior to the restrictions being imposed. The two year restriction on use of neonicotinoid pesticides comes to an end in December, at which point no one outside (and possibly inside) of Defra knows what is going to happen.

The National Farmers Union is being very selective with their use of information about the scientific evidence base for the effects of these pesticides on pollinators. Dr Chris Hartfield, the NFU’s horticultural policy adviser and lead on bee health issues, was quoted as saying “The majority of the research that has fuelled this debate has been based on artificial dosing studies. The big question in this area is, does this accurately reflect what happens to bees foraging in and around neonicotinoid crops? We don’t know, but the field studies haven’t shown that they are causing population declines in pollinators”.

Dr Hartfield and the NFU know full well that all of the evidence so far published shows that even at very small (field realistic) doses, neonicotinoid pesticides have been demonstrated to have important, sub-lethal effects on pollinators that may ultimately affect populations of some species. Surely the wisest course of action is to further restrict their use until we have studied the situation.

This is not the only occasion when the NFU have been less than objective with their use of scientific evidence. In the past couple of weeks I’ve had a group email exchange with Dr Hartfield in which he talked about the study by Carvalheiro et al. (2013) that “shows these [pollinator] declines have slowed (or even reversed) in the last 2 decades”. I responded by pointing out that the current situation is not as straightforward as that. The recent paper that we published in the journal Science showed that the rate of extinctions of UK bees and flower-visiting wasps has in fact increased over the period when Carvalheiro et al. (2013) see a slow down in declines in abundance.

There are a number of reasons why our results may be in disagreement with those of Carvalheiro et al., which we discuss in the paper, including the large statistical confidence interval around the rate of extinction during this latter period. However as with all such data, one or two studies will not give a definitive answer. I provided Dr Hartfield with a link to our paper but I’m still waiting to receive a reply.

Initiatives such as the Bee Brick, reduced mowing on road verges, the RHS’s Perfect for Pollinators plant list, etc., etc. are important but they are tiny contributions compared to the role that must be played by British agriculture if we are to conserve pollinator diversity in the UK. Farming accounts for 70% of the land surface in this country and has by far the greatest part to play in reducing biodiversity loss.

Within 12 months of Defra launching the National Pollinator Strategy, the same Government department has decided to bow to pressure and allow some use of a group of pesticides that we know are causing problems, even if they are not the whole story. Defra is effectively hurling what may be the first of many bricks at itself, ultimately weakening the Strategy. From conversations with politicians I know that these large departments do not have good internal communication and dialogue, but this seems to be an outstanding example of Orwellian double-think on the part of Defra.

Carol Klein’s Plant Odysseys starts 29th July (and I put in an appearance in episode 1)

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Back in June last year I talked about taking part in a day of filming with Carol Klein for her new four-part series, made with Oxford Scientific Films, called Plant Odysseys. It’s an exploration of horticultural biodiversity, each episode focused on a particular group of plants.

The first episode, devoted to roses, is broadcast this Monday 29th July at 7pm on BBC2, and I’m looking forward to seeing what the production team did with our footage from Chester. The name may be misspelled in the publicity material but it’ll still be me….

Garden pollinators for PAW no. 7 – Tree bumblebee (Bombus hypnorum)

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For my final post for Pollinator Awareness Week I’ve chosen another bumblebee, one with a fascinating history and ecology. The Tree bumblebee (Bombus hypnorum) is a relatively new arrival on our shores. It was first discovered near Southampton by Dave Goulson in 2001; since then has spread out through the country, as far north (currently) as central Scotland, and has recently been recorded from Ireland. It arrived in Northamptonshire in 2006. The Bees, Wasps and Ants Recording Society (BWARS) has been tracking its spread through a recording scheme: it’s very distinctive being the only one of our 25 bumblebees to have a ginger thorax, black abdomen and white tail.

All the evidence suggests that this was a natural range expansion for the species rather than a deliberate or accidental introduction by people. It’s what species do, they move around and change their distribution over timescales of decades to hundreds of years (there are lots of bird examples of this, including the Collared dove in the UK). There’s no suggestion that this was due to climate change, however: Bombus hypnorum has long been present in colder parts of Europe and Scandinavia.

That said, there is probably a human influence to its spread as the species is closely associated with houses and gardens, often nesting in bird boxes or roof spaces (we had one in our roof for three years running). The natural nesting habit for this bee is tree holes (hence the common name) which is why they are usually found in cavities above the ground. However, like the Buff-tailed bumblebee, they can also be found in compost heaps, as a recent posting on the Bees Knees Facebook group showed (if you’re not a member of that group I can recommend it as it’s full of friendly, practical gardening advice for those interested in how their garden can be beautiful, productive and wildlife-friendly).

Although the Tree bumblebee will take nectar and pollen from an assortment of garden plants it seems to be particularly associated with members of the rose family (Rosaceae) and is a frequent pollinator of rosaceous soft fruit such as raspberries and blackberries. I tried and failed yesterday to photograph the bee on our raspberries, so here’s a photograph of the outcome of that pollination.

The Tree bumblebee is rapidly becoming one of the commonest garden bumblebees. Is this likely to cause problems for our other bumblebees, by out-competing them for nectar and pollen, or even nesting sites? It’s too early to tell but I’d be surprised if it does.

Phew! That’s it! It’s been a bit of marathon preparing these posts on top of writing a large grant application and a thousand other jobs, but it’s been a lot of fun. Thanks to everyone who has viewed my posts For Pollinator Awareness Week and commented on them, either on the site or on Facebook. Hopefully they have raised a broader awareness of our amazing native pollinators.

Garden pollinators for PAW no. 6 – Buff-tailed bumblebee (Bombus terrestris)

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It would be impossible to write a series of blog posts about garden pollinators for Pollinator Awareness Week without considering the bumblebees (genus Bombus) and I intend to devote the last two posts to that group of bees. The bumblebees are arguably the UK’s most important pollinators of both wild and crop plants, certainly later in the season when colony numbers have increased. Earlier in the season it’s the solitary bees such as the Orange-tailed mining bee that are predominant.

Although common and widespread in gardens, the Buff-tailed bumblebee (Bombus terrestris) belongs to a group of bees in which the workers are rather variable in appearance and can be very difficult to distinguish from those in the Bombus lucorum group, which includes two other species (B. cryptarum and B. magnus).

This is a truly social species with an annual nest comprising workers and a queen. Nests are founded by queens that have mated the previous year and hibernated. They usually choose old rodent nests in which to begin their colonies, which is why they are sometimes found in garden compost bins. An interesting question that I’ve not seen answered is whether the queens actively displace mice or voles from such nests: does anyone know? This association between bumblebees and mice led Charles Darwin and Thomas Huxley into some speculation as to the role of spinsters in the British Empire.

In my garden the Buff-tailed bumblebee pollinates a range of crops including strawberries, squashes, courgettes, blackberries, runner beans, french beans, tomatoes, and raspberries. As the photo above shows they also visit the flowers of passion fruit, where they seem to be more effective than the smaller honey bees and solitary bees.

Garden pollinators for PAW no. 5 – Orange-tailed mining bee (Andrena haemorrhoa)

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The Orange-tailed mining bee (Andrena haemorrhoa) is also referred to as the Early mining bee due to its habit of emerging from over-wintered nests as early in the year as March. In truth, however, many Andrena species put in an early appearance, making them important pollinators of orchard fruit such as apples, which you can see from the photograph above, taken in my urban garden earlier this year. So “Orange-tailed” is a more descriptive name.

Thanks to the Orange-tailed mining bee and other early bees, this unnamed apple variety (which Karin and I rescued from the bargain area of a local garden centre) has gone on to produce a heavy crop of eating apples (see below). There’s considerable interest in the role of wild bees such as these as pollinators of fruit in commercial orchards, not just in Europe but in the USA too, where other Andrena spp. also pollinate apples.

The epithet “Mining bees” refers to the fact that these solitary species of the genus Andrena usually make their nests in soil, excavating deep tunnels in which to construct individual cells. It’s another generalist, taking pollen and nectar from a wide variety of garden and wild flowers. Dandelions are particularly important early in the year – so don’t over-manage your lawn and allow some to flower!

Garden pollinators for PAW no. 4 – Gatekeeper butterfly (Pyronia tithonus)

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For my fourth contribution to Pollinator Awareness Week I’m going to highlight the Gatekeeper (Pyronia tithonus), a butterfly that I featured on this blog last year. As I noted in that post, it’s fairly rare to have Gatekeepers in an urban garden which indicates that the shrubs and hedges grown by myself and my neighbours are providing the right microclimate for the adults. In addition the overgrown lawns of some adjacent gardens give opportunities for egg laying as the caterpillars are grass feeders.

Adult butterflies are very well camouflaged when resting with their wings folded. They take nectar from a range of plants in my garden but particularly love the dark, heavily scented infloresences of the buddleia variety pictured here. They also visit the wild blackberries scrambling through the hedge that separates us from next door’s garden and probably pollinate those flowers. Although it’s often said that butterflies are poor pollinators compared to bees, due to their general un-hairiness and habit of holding themselves above the stamens and stigmas in a flower, it very much depends on the type of flower. We have an unpublished manuscript that I hope to submit to a journal later this year showing that butterflies are actually better pollinators of one grassland plant than bumblebees.

Garden pollinators for PAW no. 3 – Little flower bee (Anthophora bimaculata)

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There are only five Anthophora species in the UK and the Little flower bee (Anthophora bimaculata), as the name suggests, is one of the smallest. This is a fast moving little beast and my fairly bog standard cameras (and a generally impatient personality) struggle to capture it: Steve Falk’s images do it justice better than I ever could.

Once again, as with the Patchwork leaf-cutter bee, the flowers of Lamb’s ear are a real favourite in my garden. Also like that bee, this is a solitary species, though this is one that nests in dry, sandy soil. I’ve not been able to locate any nests in my own patch and I wonder whether it’s nesting in a nearby garden.

Another generalist species, I’ve seen this bee visit food crops with open flowers such as blackberry and raspberry in my garden. Despite its size it is likely to be quite a good pollinator of those fruit as it’s abundant, fast moving between flowers, and hairy, and can carry significant amounts of pollen.

Interestingly, Anthophora bimaculata was not recorded by Muzafar Sirohi during his surveys of bees in Northampton town centre, so it’s another species that we can add to that urban list.

Garden pollinators for PAW no. 2 – Marmalade hoverfly (Episyrphus balteatus)

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One of the most frequently encountered of hoverfly species in urban gardens is the beautifully named Marmalade hoverfly (Episyrphus balteatus). This insect is a “true fly” of the order Diptera that is sometimes confused with superficially similar wasps (order Hymenoptera), though (as the common name suggests) the species is translucent orange and black in colour rather than waspish yellow and black. It also has a very flat abdomen whereas wasps are rounded, and they certainly don’t sting.

Individual insects are relatively ineffective as pollinators – they are small and not very hairy, so carry little pollen compared to bumblebees for instance. However they can be extremely abundant and that abundance makes up for any individual ineffectiveness. It’s a real generalist, visiting lots of different types of flowers, and in my garden they visit radishes (as I noted last year) and raspberries.

I often see individuals patrolling crops such as runner beans, not visiting the flowers but laying eggs on leaves and stems: the larvae of the Marmalade hoverfly is carnivorous and feeds on aphids, so it plays an interesting dual role of both pollinator and pest controller. Definitely a gardener’s friend!

Pollinator Awareness Week – 13th – 19th July 2015

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Next week has been designated Pollinator Awareness Week (PAW) by Defra and there are events and profile-raising activities going on all over the country.

The motivation behind the PAW is (quote) “to bring attention to the essential needs of pollinators and the simple actions that we can all take to help pollinators survive and thrive”.

With that in mind, next week I intend to produce one blog post a day that highlights, with photographs, a pollinator (or group of pollinators) that I’ve found in my own urban garden in Northampton. The purpose is to illustrate the diversity of pollinators that even a town garden can support, something about their fascinating life histories, and the different ecological requirements of these pollinators that our gardens can provide. For some of them I’ll even discuss the garden crops that they pollinate. First post will be on Monday.

If you, or the group you work with, are doing something for Pollinator Awareness Week feel free to share it in the comments section below.