Watch my recent interview on Earth To Be

Recently I enjoyed chatting with Dr Daniela Scaccabarozzi for the YouTube channel that she runs called Earth To Be. In a wide ranging interview we discussed my recent book, how it came about, some of the things that intrigued me during its research (including a cockroach-pollinated flower!), and the role of people and pollinators in the wider ecosystem. Thanks to Daniela for the invitation to chat! Here’s the link to the interview.

Do bumblebees make honey? Yes and no…and…maybe [UPDATED]

As kids, my friends and I did a lot of digging. We always seemed to be burrowing into slopes or excavating trenches, pretending to be archaeologists or treasure hunters. Indeed, there was a lot of ground treasure to be found in the part of Sunderland where I grew up. The area has a long history of pottery and glass making, and ship building, and the remnants of these industries could be uncovered every time we stuck a spade in the earth. Over time I developed my own small museum of interesting, unearthed fragments, including bits of hand-painted ceramics, glass bottles, and unidentifiable metal shards, alongside various animal bones I’d excavated. My parents quietly indulged this interest, and my muck-streaked face and clothes, even if they didn’t quite understand what I was doing.

Aged about 10, my first encounter with a bumblebee nest was during one such dig. On the waste ground behind a large advertising hoarding, we began digging into a low, grass-covered mound and accidentally excavated what was probably a small nest of Buff-tailed Bumblebees (Bombus terrestris). I can recall being fascinated by the waxy, odd shaped cells and by the sticky fluid that some of them were leaking. Being an adventurous sort of child I tasted the liquid: it was sweet and sticky, and that was my first encounter with bumblebee “honey”.

I’m going to leave those quotation marks in place because if you do an online search for “do bumblebees make honey?” you generally find that the answer is “no, only honey bees make honey”.

Now, defining honey as something made by honey bee strikes me as a circular argument at best. And it also neglects the “honey” made by meliponine bees that is central to the culture of stingless bee keeping by indigenous groups in Central and South America, and the long tradition pre-colonial tradition of honey hunting by Aboriginal Australians. So if we widen our definition of “honey” as being the nectar*-derived fluid stored in the nests of social bees, then Apis honey bees, stingless bees and bumblebees must all, by logic, make honey. And likewise there’s wasps in the genus Brachygastra from Central and South America that are referred to as “honey wasps” because, well, I’m sure you can work it out!

But this is where things become a little trickier, because turning nectar* into honey involves some complex evaporation and enzymatic activity, so that the resulting fluid is more concentrated and dominated by the sugars glucose and fructose. Although analysis of honey bee honey is commonplace, and there’s been some research conducted on the honey of stingless bees, I don’t know of any studies that have compared Bombus honey with that of other bees, or with what is stored in the nests of honey wasps**. If I’ve missed anything, please do comment and let me know, but this strikes me as an area of research demanding some attention.

So do bumblebees make honey? That very much depends on our definitions, but I’m happy to accept that they do because “honey” is not a single thing: it’s an insect-derived substance that can take a range of forms but serves the same broad purpose of feeding the colony. And although insects have probably been producing it for millions of years, I think I’ve known the answer to the question for almost 50 of them…

UPDATE: A couple of people have commented on social media that there are legal definitions of “honey” as a foodstuff. Here’s the definition according to UK law***:

“the natural sweet substance produced by Apis mellifera bees from the nectar of plants or from secretions of living parts of plants or excretions of plant-sucking insects on the living parts of plants which the bees collect, transform by combining with specific substances of their own, deposit, dehydrate, store and leave in honeycombs to ripen and mature”

So, legally, we can’t call anything that isn’t made by Apis mellifera “honey”, at least from a foodstuffs regulation perspective. But that’s clearly different to what we have been discussing above, which is about a biological definition of honey.

It’s also interesting to look at the compositional requirements of honey as a foodstuff (presented in Schedule one of that document, if you follow the link above). The lower limit for moisture content is 20%. Now if you consider that most nectar in flowers has a sugar content of between about 20% and 50%, clearly there’s been a lot of evaporative work done by the bees to reduce the amount of water in the honey. I would love to know how bumblebee (and other insect) “honey” compares to this: do they put the same kind of effort into evaporating the water from the stored nectar? Given that the purpose of reducing the water content is to prevent fermentation by yeasts when it’s stored for a long time, and that there are bumblebee species which have colonies that are active for more than one year, I imagine that at least some species in some parts of their range may employ similar tactics.

Thanks to everyone who has been commenting and discussing the topic. It never ceases to amaze me how much we still do not understand about some fundamental aspects of the natural history of familiar species!

*And honeydew to a greater or lesser extent.

**I’m going to ignore honey pot ants for now as this is complex enough as it is and they don’t store the “honey” in nest cells.

***From what I can gather definitions in other countries are similar.

Nature’s graffiti: lichens pattern clay tiles

Following on from my recent blog post about biological crusts, I was intrigued by the patterns formed by these lichens on the clay tiles capping the brick gate columns of our local cemetery. It looks as though they have been created by successive waves of growth, but I may be wrong about that. Any lichen experts out there who can tell me what’s going on?

I think the species is Xanthoria parietina, but again I’m happy to be corrected. Below is a cropped close-up from a slightly different angle.

What are the best books about bees and other pollinators?

Clearly that’s a very subjective question and everyone has their own view on which books about a particular subject they would recommend! So coming up with a list of just five for the Shepherd book recommendation site was not easy. My list features authors such as Brenda Z. Guiberson, Megan Lloyd, Steven Falk, Dave Goulson, Mike Shanahan and Stephen L. Buchmann, which will hopefully inspire you to read some of these books.

Here’s the link:

If you think that I’ve missed your favourite from the list, please do comment below. And if you’re an author, consider signing up for Shepherd and curating your own list, they’ve been really helpful and it’s a useful service for readers and authors.

A new study shows that even short-tubed flowers can specialise on hawkmoths as pollinators

Of all of the “classical” pollination syndromes, flowers that are hawkmoth pollinated have one of the highest levels of predictability. If a flower is pale in colour, opens at night, is highly scented, and possesses a long tube at the bottom of which is a supply of nectar, there’s a very high likelihood that it’s pollinated by long-tongued hawkmoths (Sphingidae).

Indeed, one of the foundational stories about the development of our understanding of how pollination systems evolve, relates to Charles Darwin, the long-tubed orchid Angraecum sesquipedale and the hawkmoth Xanthopan morganii praedicta.

Fast forward 160 years and we now know that pollination syndromes are more complex than 19th and early 20th century scientists imagined – see my recent book Pollinators & Pollination: Nature and Society for a discussion of this topic. That’s not surprising because, as I point out, we probably have data on the interactions between plants and their pollinators for only about 10% of the estimated 352,000 species of flowering plants. There’s still much to be discovered!

As an example of how our understanding of specialised flower-hawkmoth interactions is developing, consider this recent study that I’ve just published with my Brazilian colleague Felipe Amorim and other collaborators. In it we have shown that, contrary to expectations, a species of Apocynaceae (Schubertia grandiflora) with a relatively short floral tube can specialise on hawkmoths with much longer tongues than we might predict.

The full reference with a link to the study is shown below, followed by the abstract. If you would like a PDF, please drop me a line via my Contact page:

Amorim, F.W., Marin, S., Sanz-Viega, P.A., Ollerton, J. & Oliveira, P.E. (2022) Short flowers for long tongues: functional specialization in a nocturnal pollination network of an asclepiad in long-tongued hawkmoths. Biotropica


Since Darwin, very long and narrow floral tubes have been known to represent the main floral morphological feature for specialized long-tongued hawkmoth pollination. However, specialization may be driven by other contrivances instead of floral tube morphology. Asclepiads are plants with a complex floral morphology where primary hawkmoth pollination had never been described. We detailed here the intricate pollination mechanism of the South American asclepiad Schubertia grandiflora, where functional specialization on long-tongued hawkmoth pollinators occurs despite the short floral tube of this species. We studied two plant populations in the Brazilian Cerrado and recorded floral visitors using different approaches, such as light-trapped hawkmoths for pollen analysis, direct field observations, and IR motion-activated cameras. Finally, using a community-level approach we applied an ecological network analysis to identify the realized pollinator niche of S. grandiflora among the available niches in the pollinator community. Throughout a period of 17 years, long-tongued hawkmoths were consistently recorded as the main floral visitors and the only effective pollinators of S. grandiflora. Flowers rely on highly modified corona and gynostegium, and enlarged nectar chambers, to drive visitors and pollination mechanism. Despite its relative short-tube, network analysis placed S. grandiflora in the module including exclusively long-tongued hawkmoth pollinators and the most phenotypically specialized sphingophilous plants in the community. These results represent the first example of functional specialization in long-tongued hawkmoths in an asclepiad species. However, this specialization is uncoupled from the long floral tubes historically associated with the sphingophily syndrome.

The value of butterfly specimens for understanding species extinctions – a new study just published.

The Chequered Skipper Reintroduction Project has featured in several posts over the last few years – see here and here – and University of Northampton PhD researcher Jamie Wildman has been working hard to complete his thesis under the less-than-ideal conditions imposed by the COVID-19 pandemic. The first paper from the project has just been published and it deals with Jamie’s monumental efforts to bring together all of the scattered data relating to preserved Chequered Skipper specimens held in museums and private collections. An existing database contained just 266 records; Jamie’s efforts increased that by an order of magnitude, adding a further 3,533 new records that document where and when specimens were collected, and by whom.

This 1,328 % increase in data means that we now know much more about the historical distribution of this butterfly and how that changed over time.

The Chequered Skipper went extinct in England in 1976 and this enhanced database will allow us to understand why that extinction occurred. This initial paper documents the strategy used to find the additional records as a road map for how others might proceed in the future. The full reference with a link to the paper is here:

Wildman, J.P., Ollerton, J., Bourn, N.A.D., Brereton, T.M., Moore, J.L. & McCollin, D. (2022) The value of museum and other uncollated data in reconstructing the decline of the chequered skipper butterfly Carterocephalus palaemon (Pallas, 1771). Journal of Natural Science Collections 10: 31-44

This is the abstract:

The chequered skipper butterfly Carterocephalus palaemon (Pallas, 1771) was declared extinct in England in 1976 after suffering a precipitous decline in range and abundance during the 20th Century. By searching and collating museum and other records, we show how a deeper understanding of this decline can be achieved, thus furthering conservation objectives. A preexisting Butterflies for the New Millennium (BNM) database of United Kingdom butterfly species records, created by Butterfly Conservation in conjunction with the Biological Records Centre (BRC), contained 266 historic C. palaemon records from England. United Kingdom (UK) museums and natural history societies were contacted for specimen data, and these sources added 2175 new records to the BNM. Owners of private specimen collections were also contacted, and these collections accounted for a further 465 records. Specimens originating from UK museums, other institutions, and private collections represent 2640 (71%) of total new records. Other sources, such as personal accounts held in museums, published and unpublished texts produced an additional 894 records. A further 437 records from museums, private collections, and other sources were considered partial and omitted from the data due to limited or misleading date and/or locality information. In summary, data from UK museums and other sources has infilled English C. palaemon distribution prior to 1976, offering further insight into potential environmental and anthropogenic drivers of decline at key sites. The quality and quantity of data obtained using the method outlined in this study suggests similar work could be carried out for other extinct or declining butterfly species to improve our knowledge of habitat requirements and historical distribution via modelling, identify causes of decline, and provide valuable information for potential reintroductions.

Is the tropical epiphytic house plant Monolena primuliflora an “ant plant”?

I love going to botanic gardens and I keep a “life list” of those that I have visited. So on a visit to Lund University last week, to give a seminar and take part in an MSc defence, I was delighted to be able to add another one to that list. Lund University Botanical Garden is quite small, like many such urban gardens, and this is not the best time of the year to visit. But there was a good show of early spring plants in flowers, the sun was shining, and quite a number of people were enjoying the peace and calm in the middle of a city.

The glasshouses were especially busy, and they have a nice collection of cold-sensitive plants arranged by habitat and taxonomy, such as cacti and succulents, ferns, orchids, and so forth. One of the reasons why I enjoy botanic gardens so much is that I always, without exception, see plants that I have never previously encountered, often doing unexpected things.

Lund was no exception, and I was particularly intrigued by a plant called Monolena primuliflora which was being grown in a hanging basket, as is often the case with epiphytic plants. It’s a species of Melastomataceae, a family that I know well from tropical field work. But this one looked unlike any melastome that I’d ever seen. In particular, I was drawn to the large rhizome or caudex from which the leaves emerge:

This immediately reminded me of some of the epiphytic “ant plants” such as species of Myrmecodia and Hydophytum and especially ferns such as Lecanopteris. All of these myrmecophyte genera have evolved swollen stems or rhizomes which house colonies of ants. The ants in turn defend the plants against herbivores, in a mutualistically advantageous relationship.

Sure enough, when I searched online for information about Monolena primuliflora, it’s widely described in the house plant community as an “ant plant” – see here and here for example. After I tweeted about this, biologist Guillaume Chomicki (who has been researching these ant-plant interactions) was intrigued but asked about the evidence for it being a myrmecophyte:

That got me thinking, so I dug around in the botanical literature for the evidence and found…..nothing. The standard monograph on the genus by Warner (2002) doesn’t mention it and as far as I can tell (please someone will correct me if I am wrong) there’s no documented study of this species or genus having a myrmecophytic relationship with ants.

If I’m correct, how has the idea of Monolena primuliflora as an ant plant come about? This is a relatively new introduction to the houseplant trade and I suspect that plant sellers have made assumptions about the swollen rhizome (as I did!) to make the plant sound more interesting. There’s no doubt that the rhizome is fascinating and unusual in the family, but its function may be to store water (as found in many epiphytic orchids) rather than to house ants.

In my recent book Pollinators & Pollination: Nature and Society, and in this article last year in the magazine British Wildlife, I discussed how the world of plants (and pollinators) is full of myths and misunderstandings. This seems to be another one and by writing this blog post I’m hoping that we can clarify the situation with regard to Monolena primuliflora. So if you have any further information about it, please do comment below.

My thanks to everyone on Twitter who commented about the plant, especially Guillaume for asking the question!

The coltsfoot is flowering! But why is it so different to dandelion?

Yesterday Karin and I took to our bikes and rode south through some very nice, managed beech and oak woodland that runs parallel to the Isefjorden in this part of Odsherred. In was cold but sunny, birds were singing, and we saw the occasional insect on the wing. The kind of day that reminds you that spring is coming fast. On the way back we paused at a small housing development near the former psychiatric hospital at Annebergparken. In an area of disturbed ground I was delighted to see a patch of coltsfoot (Tussilago farfara) in full flower, the dandelion-like inflorescences a beacon to passing bees and flies.

Although it resembles a small dandelion, and belongs to the same family (Asteraceae), Tussilago is only distantly related to Taraxacum. Coltsfoot is really a type of groundsel (tribe Senecioneae) whereas dandelions are related to chicory (tribe Cichorieae).

Coltsfoot is unusual in that it produces its flowering stems long before the leaves that give it its common name, the plant’s reproduction powered by the energy that it stored up the previous year. Dandelions, like most herbaceous plants, produce their leaves first, then flower. That’s not the only difference to dandelions though.

The Database of Pollinator Interactions (DoPI) lists 9 species of insect that have been recorded as visiting coltsfoot for nectar and/or pollen. In contrast, the entry for Taraxacum officinale lists more than130 species as flower visitors. I thought initially that it might be due simply to under-recording, but this study of coltsfoot in Germany only recorded 16 insect species. So the greater attractiveness of dandelion is likely to be real. Why the big difference in pollinators?

One reason for it could be that dandelions have a very different flowering strategy; they can be in flower 12 months of the year, depending on local weather conditions, with a reproductive peak in May or June. They therefore have the opportunity to interact with many more insects than coltsfoot, which in contrast you generally only see in flower between March and May at the very latest.

Dandelions are also much more abundant than coltsfoot which is no doubt also a big factor in determining how often insects are observed on the flower heads. It’s not unusual to see whole fields full of dandelions in flower but I’ve never seen coltsfoot do that, perhaps because they prefer to grow on rather disturbed ground.

There may be some other factors at play here that I’m not aware of, for example a lower rate of nectar production in coltsfoot. Having said that, the fact that dandelions produce any nectar at all is a real conundrum. All of the literature claims that Taraxacum officinale is “apomictic“, a plant reproductive strategy in which seeds are produced without requiring ovules to be fertilised by pollen. In fact the online Ecological Flora of Britain and Ireland entry for dandelions lists the pollen vector as “none” for that very reason. But I’m sure that the real story is more complicated, otherwise why would these plants invest so much of their energy and resources in attracting and rewarding flower visitors? I’ve not delved deeply into the Taraxacum literature so perhaps one of my readers knows?

Our encounter with coltsfoot reminded me of the work that I did last year with the Stanwick Lakes nature reserve in Northamptonshire, advising on how best to enhance and manage the site (which is primarily a bird reserve) for pollinators. One of my recommendations was that they enlist their volunteers to collect seeds and root or stem cuttings from the small, isolated populations of early-flowering plants such as coltsfoot (pictured on the reserve below) and introduce them around the site in suitable spots. This would both increase the availability of nectar and pollen for the first flower visitors of spring, and also the ecological connectivity between different parts of the site as the pollinators are able to move around more effectively. So I was delighted to see this post on LinkedIn from Liz Williams who works at Stanwick, demonstrating that they have taken my advice on board and begun the hard work of planting for pollinators.

My work with Stanwick was an example of the advisory and consulting services that I offer. If you’d like some advice on how to improve an area for pollinators, or for biodiversity more broadly, please do get in touch via my Contact page.

Solace in nature: sunset over Hov Vig bird reserve

The invasion of Ukraine by hostile Russian forces is a humanitarian disaster the likes of which Europe has not seen for decades, and hoped never to see again. Like many people, Karin and I have been watching the news about the war with a sense of helplessness, bewilderment and alarm, wondering how such things can come to pass in the 21st century. We thought we were past the stage where aggressive, narrow-minded dictators could bully their way into adjacent countries.

Faced with 24 hour media coverage of such desperate events, it’s easy to lose touch with the world around us. Karin and I are fortunate to be able to bicycle to some beautiful local spots where we can reflect and try to find some solace in nature. That’s what we did yesterday with a late afternoon visit to the Hov Vig bird reserve. In addition to my photos, which I’ll let speak for themselves, Karin filmed a short video for her YouTube channel which includes a marvelous array of bird calls.

Tonight we are taking part in a fund-raising event at the local culture house. Please think about how you can help to support Ukraine, in however modest a way, but also don’t forget to connect with nature. It will always endure, despite the destructive efforts of humans.

Practical methods for assessing insect pollination services provided by sites – download our new study for free

In September 2016, along with 11 other pollinator & pollination scientists, I took part in a two-day two-day workshop held at the UNEP-World Conservation Monitoring Centre in Cambridge. The aim was to develop a range of simple, practical methods to enable the valuation of insect pollination services to agricultural crops that are provided by a nature reserves or other natural or semi-natural habitats, for TESSA – the Toolkit for Ecosystem Service Site-Based Assessments.

After a long gestation, caused not least by the COVID-19 pandemic, the paper outlining the methods that we developed has been published. It’s open-access and downloadable for free. Here’s the full reference with a link to the paper:

Ratto, F., Breeze, T. D., Cole, L. J., Garratt, M. P. D., Kleijn, D., Kunin, B., Michez, D., O’Connor, R., Ollerton, J., Paxton, R. J., Poppy, G. M., Potts, S. G., Senapathi, D., Shaw, R., Dicks, L. V., & Peh, K. S.-H. (2022) Rapid assessment of insect pollination services to inform decision-making. Conservation Biology 1–13

And here’s the Abstract:

Pollinator declines have prompted efforts to assess how land-use change affects insect pollinators and pollination services in agricultural landscapes. Yet many tools to measure insect pollination services require substantial landscape-scale data and technical expertise. In expert workshops, 3 straightforward methods (desk-based method, field survey, and empirical manipulation with exclusion experiments) for rapid insect pollination assessment at site scale were developed to provide an adaptable framework that is accessible to non-specialist with limited resources. These methods were designed for TESSA (Toolkit for Ecosystem Service Site-Based Assessment) and allow comparative assessment of pollination services at a site of conservation interest and in its most plausible alternative state (e.g., converted to agricultural land). We applied the methods at a nature reserve in the United Kingdom to estimate the value of insect pollination services provided by the reserve. The economic value of pollination services provided by the reserve ranged from US$6163 to US$11,546/year. The conversion of the reserve to arable land would provide no insect pollination services and a net annual benefit from insect-pollinated crop production of approximately $1542/year (US$24∙ha–1∙year–1). The methods had wide applicability and were readily adapted to different insect-pollinated crops: rape (Brassica napus) and beans (Vicia faba) crops. All methods were rapidly employed under a low budget. The relatively less robust methods that required fewer resources yielded higher estimates of annual insect pollination benefit.