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.
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.
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: https://shepherd.com/best-books/bees-and-other-pollinators
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.
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 https://doi.org/10.1111/btp.13090
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 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.
Join me this Thursday at a free online talk organised by Buglife where I’ll be giving an introduction to how flowers function and the ways in which their behaviour manipulates pollinators to ensure reproduction. I’ll be covering:
- What are flowers and where did they come from?
- How flowers function and reward pollinators.
- Some case studies from my own research on flower and pollinator behaviour.
- Why is it important that we understand floral biology?
Here’s the link to register for the event: https://www.buglife.org.uk/events/to-be-a-flower-with-professor-jeff-ollerton/
I look forward to seeing you there!
How we, as a society, value nature, and the tension between valuing (or appreciating) nature versus appreciating (or pouring money into) human cultural activities, have been consistent themes of this blog since I started it almost a decade ago; see for example my posts “How do we value nature? Costanza, Monbiot and the clash of concepts” and “Is the angry response of (some) environmentalists in the aftermath of the Notre Dame fire reasonable?“
Putting a monetary price on nature runs counter to the personal philosophies of many conservationists, which I completely understand: I have mixed feelings too. However there’s a whole field of research devoted to it called Ecological Economics and the valuation of natural capital and ecosystem services now plays a central role in the policies and strategies of both businesses and governments: see for instance the UK Government’s recent report on “The Economics of Biodiversity: The Dasgupta Review“. And whether we like it or not, the Earth’s ecosystems and the biodiversity that they contain support our global economy in very tangible ways, a point that I emphasise in my book Pollinators & Pollination: Nature and Society. If you’re reading this with a cup of coffee in your hand, you have to consider the ecological and financial impact of the billions of wild and managed bees that support the global coffee industry.
“What’s all of this got to do with guitars?” I hear you asking. Well, music, and especially guitars, are another constant theme of the blog, including my love of the songs of Crosby, Stills, Nash and Young, and my restoration of an old acoustic guitar back in 2020.
These themes converged together in a rather unexpected way just over a week ago. It was my birthday and as a present Karin had offered to buy me a new guitar. So off we went to Copenhagen for the day. One of the city’s best guitar shops is Akustikken and there I tried out several makes and models of acoustic guitar, of varying price and quality, before finally settling on an Epiphone Texan in an aged sunburst finish (see the image below). It plays very nicely, felt right in my hands, and was moderately priced despite its solid wood construction (cheap guitars often use laminated wood).
The guitar that really caught my eye, however, was the one in the photograph above: a Martin 045-S Stephen Stills Signature Model. Now, this is a serious, serious guitar. Based on a 1930s model owned by Stills himself, it was hand made in the USA in a limited edition of 91, of which this was number 48. The woods from which it’s made are rare and exceptional, including Adirondack spruce, Madagascar rosewood, and ebony, all species about which there are significant conservation concerns (see Richard Hobbs’s great blog The Nature of Music for more on this – highly recommended for anyone interested in the interface between human culture and ecological conservation).
The price tag for this guitar? A mere160,000 Danish kroner, about £18,000 or $20,000…..
That was WAY outside of our budget! But when the staff learned that it was my birthday they kindly took the Martin out of its humidity-controlled glass case and let me play it. I was a bit overwhelmed and very nervous if I’m honest, it was easily the most expensive guitar I have ever had in my hands! Karin took a short video of me strumming a few chords which I uploaded to Twitter:
Now, I’d played guitars up to around $2,000 in price that day, so a reasonable question is: did the $20,000 guitar sound ten times as good? Well, not in my hands it didn’t…. But in one sense it doesn’t matter, you’re not just paying for what it sounds like, you’re paying for the story, for the association with Stills, and the highly skilled crafting of the guitar – it is an exceptionally beautiful and fine-sounding instrument.
This brings us back to nature. We know from a lot of ecological experiments that have been conducted over the years that there’s a positive relationship between biodiversity (measured by the number of species in an ecosystem) and the way in which that ecosystem functions. So if you have more different kinds of plants in a grassland, for example, there tends to be greater carbon capture, more efficient use of water and uptake of nitrates from the soil, more resilience to events like drought and fire, and so forth. This is a strong and pervasive argument for conserving species within ecosystems: the more we have, the better the “health” of that ecosystem.
But, as with the sound of guitars, there’s probably an upper limit to this and ecosystems with ten times as many species probably do not function ten times as well. But they do function better. Having said that, this is a complex area of research with some competing ideas (and scientists) – this Wiki provides quite a good summary.
Regardless of the technical details, there’s no doubt that having more pollinators in an ecosystem, for example, increases the reproduction of a wider range of the plants that are present. Or that the presence of a greater diversity of dung beetles improves the rate of dung removal in grasslands.
But of course nature is more complicated than this. Just as a well made and high-value guitar is never going to sound good in the hands of a poor guitarist, likewise, species diversity in itself is insufficient. It is the interactions between those species that determines much of the way in which the ecosystem functions, and an ecosystem is never going to function well over the long term if it is inappropriately managed or if the processes that shape ecosystems, such as grazing by wild herbivores or natural fire regimes, are absent or have been altered.
Ecology is a hugely complex science but perhaps by exploring metaphors like this, some of that complexity can be made accessible to a wider range of people. Tell me what you think, does the metaphor work for you?
I was saddened to learn recently of the death of Professor Leonard B. Thien of Tulane University who passed away at the end of October after a long illness. Although I didn’t know Professor Thien personally, I knew of his work in floral ecology, pollination biology and plant evolution, topics on which he had worked for since obtaining his PhD in 1968. Over the course of his career he published more than 80 articles on a huge range of botanical subjects, including ground-breaking work on mosquito pollination of orchids (Thien 1969). The orchid species Alaticaulia thienii is named in his honour.
The studies Leonard Thien published that really inspired me when I was first starting out on my journey as a researcher, however, involved his work on “relictual” angiosperms, i.e. flowering plants that have very long evolutionary histories and deep phylogenetic roots back to the early Cretaceous period, for example Magnolia and Illicium. Papers with titles such as “Patterns of pollination in the primitive angiosperms” (Thien 1980) piqued my interest and motivated me to work on Australian Piperaceae for a short while following my PhD (Ollerton 1996). It was a topic that I struggled to gain further funding for, and later molecular systematic studies changed many of our ideas about what constitutes the most basal groups of extant flowering plants. But nonetheless, the questions that Leonard inspired in me, regarding the ecologies of these relictual taxa, and whether we can infer the reproductive ecology of the earliest flowering plants from studies of their surviving descendants, are ones that intrigue me to this day (van der Kooi and Ollerton 2020).
Leonard Thien kept up this interest even as new DNA technologies over turned old ideas, and he was the first to study the reproductive ecology of Amborella trichopoda on New Caledonia, a species now considered to be the earliest surviving clade of flowering plants (Thien et al. 2003). This is just one part of a legacy of work that current and future generations will build upon as we develop our understanding of the relationships between pollinators, plants, and evolutionary processes.
I’m grateful to Peter Bernhardt for prompting this post and for sending me a copy of the In Memoriam article that he and and David White will publish in the Plant Sciences Newsletter in March, and to Lorraine Thien for providing the photograph that accompanies this post.
Ollerton, J. (1996) Interactions between gall midges (Diptera: Cecidomyiidae) and inflorescences of Piper novae-hollandiae (Piperaceae) in Australia. The Entomologist 115: 181-184
Thien, L.B. 1969. Mosquito pollination of Habenaria obtusata (Orchidaceae). American Journal of Botany 56: 232-237.
Thien, L.B. 1980. Patterns of pollination in the primitive angiosperms. Biotropica 12: 1-14
Thien, L.B., Sage, T.L., Jaffre, T., Bernhardt, P., Pontieri, V., Wesston, P.H., Malloch, D., Azuma, H., Graham, S.W., McPherson, M.A., Hardeep, S.., Sage, R.S. & Dupre, J.-L. 2003. The population structure and floral biology of Amborella trichopoda (Amborellaceae). Annals of the Missouri Botanical Garden 90: 466-490
van der Kooi, C.J. & Ollerton, J. (2020) The origins of flowering plants and pollinators. Science 368: 1306-1308
Towards the end of our stay in Glastonbury, Karin and I took an omega-shaped circular walk that looped over the famous Tor, through the town, and back to our cottage. At one point the road we walked passed through a cutting in the Jurassic sandstone called Wick Hollow. Several very large oak and beech trees had anchored themselves into this stone, their roots finding cracks in the rock and no doubt widening them over time as they grew. The trees were spectacular and I took a few shots with my phone, though these really don’t do them justice.
The shade and structure created by the trees allowed a diversity of ferns, mosses, lichens and seed plants to grow. I’m always amazed by the power and adaptability of plants, even large trees, to find a foothold in the unlikeliest of places and by doing so, create microclimates that allow other species to flourish. Life supports life.
An interesting study published this week in the journal Science has provided more evidence that natural regrowth of forests is faster and more efficient than tree planting for restoring habitats. Here’s the Guardian‘s take on it:
Here’s a link to the original study in the journal:
And here’s a link to something that I wrote on this topic last year, arguing that pollinators and seed dispersers play a vital role in this process:
Tree planting has its place, of course, especially as a way to get local communities engaged in positive action for the environment. But it’s not the solution for large-scale habitat restoration: in order to do that we need to harness nature’s own regenerative abilities.