Monthly Archives: April 2014

“These things aren’t to study. They’re to turn up very loud and say, hey, once upon a time, everything was just as easy as this”

May June 2010 Garden, River of Flowers, Cambridge 011

The title of this brief post is a quote by Led Zeppelin frontman Robert Plant, from an interview that the BBC reported just this morning, regarding the forthcoming release of previously unheard Zeppelin material.  You can read the story and hear the interview here.

Regular readers of this blog will know that I’m not averse to sprinkling musical references into my posts, and this was a great quote that seemed to chime with something else I read this morning.  Over at the Small Pond Science blog, evolutionary ecologist Amy Parachnowitsch has a thought provoking post entitled “Save the bees, but maybe not this way“.  I’ll let you read it for yourself, but in a nutshell Amy is concerned about the scientific legitimacy of a “Save the Bees” campaign being crowd funded by online activist network

I share this concern and it worries me that whoever is organising the campaign is exploiting the genuine desire by people to “do something for the bees” without any regard for what exactly it is that’s “being done”.  It seems to me to be purely a campaign fund-raiser by people who don’t understand the issues or how science works, the message being: “These things aren’t to study. They’re to turn up very loud and say, hey, once upon a time, everything was just as easy as this”.

The organisers promise “the world’s first large scale, grass-roots supported, totally independent study of what’s killing our bees that decisively challenges the junk science of big pharma”.  As Amy notes, this is hugely offensive to independent scientists who are working on bee conservation issues (such as myself).  But without ever actually saying what they are going to do with the money, they’ve already had pledges of money from over 78,000 people!  If only raising funds for real research was that easy!

To reiterate what I said in the comments to Amy’s post, something that really worries me is that over-emphasis on pesticides and honey bees as single issues affecting “pollinator conservation” deflects attention from other factors which are at least as important, such as habitat loss. Colleagues and I have a manuscript in preparation at the moment showing that native bee and flower-visiting wasp extinctions in Britain began in the mid-19th century and reached their highest rate during the period 1929-1959, during a time of rapid agricultural intensification (but prior to the introduction of neonicotinoid pesticides that is currently exercising many people).  Loss of pollinator diversity is an issue that has deep roots.

In actual fact, although wild bee diversity is declining in the UK, overall abundance seems to be stable as some species are doing extremely well, including a new natural colonist, the Tree Bumblebee (Bombus hypnorum) which is spreading fast and is locally common.  But clearly greater diversity provides us with future insurance against losses of other species.

There are positive things that can be done for pollinator populations by every citizen, beyond giving money to crappy, pseudo-scientific campaigns, as I talked about in a recent post of mine.  So please don’t contribute to this request, and use the money you save to buy some wild flower seeds and/or the Led Zeppelin reissues.  It will make the world a better place.



Filed under Bees, Biodiversity, Ecosystem services, Gardens, Honey bees, Pollination

Book review: “Pollination and Floral Ecology” by Pat Willmer

2013-11-28 14.48.27

Review of “Pollination and Floral Ecology” by Pat Willmer (2011) Princeton University Press. £65. pp. 832.

Some backstory:  In early 2012 I was asked by the review editor of the journal Annals of Botany to review this book, and I jumped at the chance as it’s the first major single-author overview of the field of pollination ecology for a number of years, by a well respected academic in the field.  Unfortunately the review took a lot longer than I expected, in part because I was also coordinating my department’s Research Excellence Framework submission, on top of other teaching, research and admin duties, and it was taking up quite a bit of my time.

In addition I had mixed feelings about the book and wanted my review to do it justice, not be over-critical but at the same time highlight what I saw as flaws.   In the words of the  Fairport Convention song, Who Knows Where the Time Goes? – my review was only completed last Christmas and duly submitted. Turns out that the journal has a backlog of book reviews to publish and the editor asked that, given it’s been a couple of years since the book was published, would I mind if the review was posted on the Annals of Botany blog rather than in the printed journal.  I happily agreed as it’s likely to get more readers on the blog, and said I’d also post it on my own blog.  So here it is:


Any text book that tries to assess and summarise the whole of a multidisciplinary research field such as pollination ecology and floral biology is required to be four things:  (1) comprehensive in its scope; (2) up to date in its coverage of the literature; (3) accurate in its assessment of the current state of the field; and (4) authoritative in the conclusions it presents.

This volume by Professor Pat Willmer of the University of St Andrews certainly ticks the first box.  It’s a huge book, and covers everything relating to the evolution of flower attraction and reward systems, ecological interactions with pollinators, biochemistry, physiology, agriculture and conservation; all in 29 chapters split into three sections, with 87 pages of references.  The literature extends to 2010, which is impressive for a book published in 2011 (though see my comments below about completeness of the literature).   Specialist terms are highlighted in bold to direct the reader to the glossary at the back, a useful device even if there are a few inaccuracies, which I’ll mention later.

So far so good, and the author is to be congratulated on putting together such a comprehensive, not to mention timely, single-author book.  It’s clearly the summation of a career devoted to studying pollinators and flowers, and the author’s passion for her subject is apparent throughout.

However when we come to points 3 and 4, things are less straightforward.  There are some issues with accuracy that are troubling in a book aimed at newcomers to the field as well as established researchers.  To give just a few examples:

– on p.18 we are told that asclepiads have “one stamen” (they have five); on p.169 and in the glossary that asclepiad pollinia are the pollen grains from one anther (they are the contents of half an anther); and on p.170 that the pollinaria are “glued” to pollinators (they actually clip on).

– in the glossary, tree ferns are referred to as “cycads”, an error that is repeated on p.89.

– on p.88 there is a statement suggesting that tree fern spores were dispersed by “animal fur” 300 million years ago, long before the evolution of mammals, and that this (and dispersal of spores of fungi and mosses) is the equivalent of pollination: it is not, it equates to seed dispersal.

These are troubling errors of basic botany that are forgivable in an early draft of the book (everyone makes mistakes) but not in the final published version, after it’s been read, reviewed, checked and edited.  If the book goes to a second edition I hope that these (and other) mistakes will be fixed.  But they do hint at a fundamental problem with a book (and a field) as large and complex as this: a single author is arguably unlikely to be able to do justice to all of the subject matter.

There are parts of the book where it is unclear (to me at least) what the author is actually saying.  For example, on p.96 there is a graph which, it is suggested, demonstrates that pollination by animals is “technically uncommon when assessed in terms of the numbers of broad taxonomic groups that use it”, though the legend to the figure claims that “most orders of plants have no families” that possess wind pollination.  This is confusing: what is to be concluded by someone new to the field?  Is animal pollination common or rare?  Likewise, on p.91 we are told that the “first angiosperms…would probably have had their pollen moved mainly by wind…”, but then on p.92 that “an element of insect pollination could be regarded as almost ancestral”.  Which is correct?

There are other aspects to the book that are simply out of date; for example the linear, rather deterministic schemes set out in Figures 4.6 and 4.8 showing that Cretaceous flowers were open and radially symmetrical, and only later evolved into complex, bilateral flowers in the Tertiary, ignores fossil discoveries showing that orchids evolved in the Cretaceous (Ramírez et al., 2007).  Likewise, discussion of “counterproductive” crypsis in flowers (p.124) neglects recent findings of cryptic, wasp-pollinated plants in South Africa (e.g. Shuttleworth & Johnson, 2009).

There is a theme emerging here: some of the botany that the book presents is inaccurate, confused or out-dated.  Fortunately the zoological aspects of the book are much better, as one might hope from a Professor of Zoology.

The final criterion, that the book should be “authoritative in the conclusions it presents”, is however, in my view, the main weakness of this volume.  The author is unhappy with recent developments in the field, particularly as they relate to community-scale assessments of plant–pollinator interactions, in terms of network analyses and predictive utility of pollination syndromes.  Clearly Professor Willmer is on a mission to rebalance what she perceives as failings within some of the current trends in studying pollination.  A book review is not the place for a technical dissection of the author’s arguments, which is best left to the peer-reviewed literature (though I would argue that that’s also the place to present some of the criticisms the author introduces, rather than into a text book such as this).  I could focus the whole of this review on these topics because: (a) they take up a large proportion of the book, about one-third of the text pages; and (b) they are highlighted on the cover as being one of the main contributions of the book; specifically, that the author provides a critique of previous work that does not distinguish between “casual visitors and true pollinators” that can in turn result in “misleading conclusions about flower evolution and animal-flower mutualism”. Unfortunately her targets are straw men, and one – I believe quite telling – example will suffice.

On p.447 there is a criticism of the use by Waser et al. (1996) of Charles Robertson’s historical data set, and specifically that the analyses they present “…did not distinguish visitors from pollinators even though Robertson’s database did include information on this”.  However Waser et al. clearly state (p.1045 of their paper) that only pollinators were included in the analyses, not all flower visitors, and that “visitation is not a synonym for pollination…non-pollinating visitors are excluded (as in Robertson 1928)” (p.1048).

Why should Professor Willmer make a statement to the contrary?  Evidently she wishes to impress upon her readers that (in her opinion) there are fundamental problems in current approaches to studying pollination at a community level.  But even if that were the case (and I don’t believe it is) misrepresenting previous studies to suit an argument is poor scholarship at best.

Regardless of whether some of her criticism is well founded, the author does not seem to appreciate that plant–flower visitor interaction networks are ecologically important regardless of whether or not a flower visitor acts as a pollinator.  More fundamentally, true pollination networks possess similar attributes to flower visitor networks, for example a nested pattern of interactions, and arguments about level of generalisation of species are a matter of scale, not category (Ollerton et al., 2003).

At the end of her Preface, Professor Willmer reveals to us quite a lot about her personal attitude to research when she states that some readers might find her approach “too traditional” in an “era where ecological modelers [might be claimed to] have more to tell us than old-style field workers”.  What the author fails to appreciate is that this is a grossly false dichotomy and that most of the pollination ecologists who have embraced new analytical methodologies for understanding plant–pollinator interactions are also “old-style field workers” with considerable experience of studying the ecology of flowers and their pollinators beyond the computer screen.

In summary this is a book that, for all its good qualities of comprehensiveness and (mostly) up to date coverage, should be read with caution: parts of it are neither as accurate nor as authorative as the field of pollination and floral ecology deserves.



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.

Ramírez SR, Gravendeel B, Singer RB, Marshall CR,  Pierce NE. 2007. Dating the origin of the Orchidaceae from a fossil orchid with its pollinator. Nature 448: 1042-1045.

Shuttleworth A, Johnson SD. 2009. The importance of scent and nectar filters in a specialized wasp-pollination system. Functional Ecology 23: 931-940.

Waser NM, Chittka L, Price MV, Williams N, Ollerton J. 1996. Generalization in pollination systems, and why it matters. Ecology 77: 1043-1060.


Filed under Bees, Biodiversity, Book review, Evolution, Mutualism, Pollination

Budget cuts to Kew are cultural vandalism



The Royal Botanic Gardens, Kew, is a beautiful place to visit, a tourist destination for visitors to London, and a green island in an urban ocean.  That’s the public face of the Gardens.  What is less well appreciated to most of the casual strollers around the flower beds and glasshouses, is that Kew is arguably the most important centre for botanical research anywhere in the world.  During its long history it has produced, and continues to deliver, top rate science that informs international conservation strategies, agriculture and horticulture, as well as basic plant science in ecology and evolutionary biology.

It’s also a welcoming, inclusive place that embraces scientific visitors from all over the world, as I know from personal experience.  Although I’ve never had a formal relationship with staff at Kew, I’ve benefitted enormously from informal links, which have facilitated research and teaching, including annual trips to the Kew Herbarium for my final year undergraduate students.

I first visited Kew as a naive 20 year old to look at their living plant collection during research for what became my first ever publication: “Adaptations to arid environments in the Asclepiadaceae” (British Cactus and Succulent Journal 1986).  So started a long appreciation of Kew and what it freely offers teachers and researchers, which has included access to specimens prior to overseas research trips, to assess distribution and flowering times; identification of specimens we’ve collected on those trips; and primary data for our study of fly pollination in the genus Ceropegia.  I’ve also used their archives for my work on John Tweedie.  Kew is an incredible resource that, in any civilised and culturally aware country, would be cherished and supported. Unfortunately it appears that I do not live in such a country.

Rumours have been circulating for a while about an impending, massive budget cut at Kew, on top of financial savings that have already been made.  Now it appears that those proposed cuts are  much bigger than anyone had thought and 120 posts, mainly in science, are threatened.  I won’t repeat the depressing statistics underlying all of this – I’ll just urge you to visit the online campaign against these cuts, read the details, watch the David Attenborough video, sign the petition, and share it with friends and colleagues.

Please don’t let Kew wither away; it’s too important to UK science, conservation and education to allow it to be gutted without a fight.



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Filed under Biodiversity, Biodiversity and culture, Gardens, John Tweedie, University of Northampton

Disturbed birds? Results of a visitor access study to the Upper Nene Valley

Parrot from Coton Manor

Human activities can have significant impacts on wildlife in quite subtle ways that are not always appreciated by those of us who enjoy going out to look at nature.  For example, simply walking close to sensitive areas such as bird nesting or roosting sites has the potential to drive those birds from an area.  This was the theme of a workshop we attended yesterday afternoon, hosted by the local Wildlife Trust as part of the Nene Valley Nature Improvement Area (NIA) Project.

During the afternoon Colin Wilkinson from the RSPB presented the results of a survey that had been commissioned to assess the level of public access and usage of the Upper Nene Valley gravel pits.  These pits have Special Protected Area status due to the numbers of migratory over-wintering birds that use them.  They are also well used by the public but at the moment we have no idea what impact this is having on the birds, though there is anecdotal evidence that it is considerable at some sites.

The consultants who conducted the study used a combination of face-to-face interviews, site surveys, etc.  There’s too much in the report to go into all of the detail – you can access the full text here – but I’ve copied the highlights from the summary below:

  • The majority (98%) of visitors were on a short visit from their home
  • Group size for interviewed groups ranged from 1-8; 51% of interviewees were visiting on their own. Stanwick Lakes was notable in that group size tended to be larger here.
  • Half of the 939 interviewees had dogs with them (636 dogs in total).
  • Across all sites and survey periods, dog walking was the most common main activity (48% of interviewees).
  • During the winter, a higher proportion of people interviewed were dog walking (48% of interviews during the winter compared to 36% in the spring at the 6 locations surveyed in both seasons).
  • Over the winter, the main activities given by interviewees were: dog walking (53%), walking (26%), and wildlife watching (6%).
  • Most (77%) interviewees had arrived by car to the survey point
  • Most interviewees were frequent visitors (60% indicated that they visited at least once a week).
  • Most visits were short: 50% of visitors stated that they spent less than one hour on site and, in total, 88% spent less than two hours at the survey location.
  • The quality of the site was the most common reason for choice of site (61% interviewees), but was not the most common ‘main’ reason’; 32% interviewees gave proximity to home as the main factor underpinning their choice of site. Proximity to home seemed particularly important for dog walkers (44%) and those fishing (40%).
  • A total of 863 visitor routes were collected, either through lines on paper maps during the interview or via GPS units which were given out.
  • There were significant differences between sites in the lengths of routes taken by visitors. There were also differences between activities. The mean route length for dog walkers was 3.1km. For cyclists the average route was 7.3km while those fishing tended to have the shortest routes (0.6km average).
  • At three of the six sites that were surveyed in the winter and the spring/summer, the median route length increased in the spring/summer when compared to the winter, stayed the same at two and fell at one, suggesting no real pattern of people walking further in the summer .
  • A relatively high proportion (78% of interviewees) indicated that they were aware of the importance of the area for wintering birds. Around a quarter (24%) of all interviewees responded that they were aware that of the international importance of the area for nature conservation.
  • 908 postcodes were mapped reflecting the home postcodes of visitors. The two main settlements were Northampton (137 postcodes from the winter interviews fell within the settlement) and Wellingborough (88 postcodes from the winter interviews).
  • Dog walkers and joggers lived closest to the site at which they were visiting, with median values of 2.3 and 2.9km respectively
  • Visitor rates (visits per household) declined rapidly with distance such that a relatively small proportion of people visit from distances beyond 3km of the surveyed access points.

The challenge now will be to understand if and how these visitors are impacting on the abundance and diversity of birds in the Upper Nene Valley, and what can be done to minimise any disturbance.  Clearly there’s a balance to be struck between public recreation and wildlife protection, and this will be the theme of future work by the Nene Valley NIA Project.

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Filed under Biodiversity, Birds, Nene Valley NIA