Category Archives: Evolution

A new review gives us a deeper understanding of the evolution of plant-pollinator interactions

If you’ve read my book Birds & Flowers: An Intimate 50 Million Year Relationship, you’ll know that I spend a few pages discussing the long-standing paradigm of how interactions between plants and their pollinators evolve and result in the formation of new plant species. This is referred to as the Stebbins (or Grant-Stebbins) Most Effective Pollinator Principle (MEPP). The MEPP is fairly straightforward and intuitive: flowers evolve their colour, shape, scent, rewards, and so forth as adaptations to the type of flower visitor that successfully moves the most pollen between flowers.

However, the MEPP is not the only Principle in town – there’s also Aigner’s Least Effective Pollinator Principle (LEPP) which is not so intuitive. In the LEPP, flowers can adapt to pollinators that are less successful at pollination, as long as those adaptations do no interfere with the pollination services provided by other flower visitors.

As I note in Birds & Flowers, we don’t know which of these Principles is more frequent in nature, because the LEPP has been much less intensively studied than the MEPP. That’s in part because it’s less well known, but also because the field work and experimental procedures required to test the LEPP are much more challenging.

Hopefully this is about to change with the publication of a brilliant critical review of the MEPP by pollination ecologists Kathleen Kay and Bruce Anderson published in the journal Annals of Botany, entitled: Beyond the Grant–Stebbins model: floral adaptive landscapes and plant speciation. The paper is open access – follow that link and you can download a copy.

Kathleen and Bruce discuss not just the MEPP v the LEPP, but also other ways in which flowers can evolve, framed around the idea of floral evolution as movement across an “adaptive landscape,” where plants are not shaped only by one pollinator but by the need to maximise overall reproductive success. This perspective allows us to explore how flowers evolve when influenced by multiple pollinators, how transitions between floral forms take place, and how speciation occurs through a combination of factors beyond pollination alone. It emphasises that pollinators are important drivers of floral change, but speciation is more likely when divergence happens across several aspects of a plant’s ecology, not just through its flowers.

It’s a great review and well worth your time reading in detail. Perhaps my favourite line in the paper comes from the abstract: “The Grant–Stebbins model, while inspiring decades of empirical studies, is a caricature of pollinator-driven speciation and explains only a limited range of adaptive outcomes.” This is something that many of us have been arguing for years: the natural world is extremely complex, so we should not expect these ecologically critical interactions between flowers and their pollinators to have simple origins or ecologies.

Evolutionary implications of a deep-time perspective on insect pollination – a new review just published

When we think of pollination, we often picture bees buzzing around flowers or butterflies flitting from bloom to bloom. This relationship between plants and pollinators is one of the most well-known interactions in nature. But insect pollination didn’t begin with the colorful flowers we see today. In fact, pollinators were at work millions of years before flowering plants (angiosperms) even existed. In a new review led by Spanish researchers David Peris and Ricardo Pérez-de la Fuente, to which I added a modern ecological perspective, we explored this topic and why it’s relevant to our current understanding of plant-pollinator relationships.

Despite centuries of research on pollination, the fossil record of pollinating insects has only gained serious attention in the past few decades. What palaeontologists have uncovered is reshaping our understanding of pollination’s origins. It turns out that insects were pollinating plants long before flowers evolved—playing a crucial role in the reproduction of ancient gymnosperms, the group of seed-producing plants that includes conifers, cycads, and ginkgos.

Most people assume that insect pollination began with flowering plants, but the evidence tells a different story. Fossilised insects with specialised body structures for carrying pollen—such as hairy bodies or mouthparts adapted for nectar-feeding—have been found in deposits dating back hundreds of millions of years. These early pollinators likely visited gymnosperms, helping them reproduce in a world that looked vastly different from today’s landscapes.

Ancient pollination was driven by a diverse range of insects, many of which are now extinct. The fossil record reveals that various insect groups—including beetles, flies, wasps, and even some long-lost relatives of modern lacewings—were already acting as pollinators long before the first flower bloomed. This means that pollination as an ecological process has far deeper evolutionary roots than many realise.

As plants evolved, so did their pollinators. The rise of flowering plants during the Cretaceous period (around 100 million years ago) transformed pollination systems, leading to the incredible diversity of plant-pollinator relationships we see today. Many of the insect groups that once dominated pollination in prehistoric times have since declined or disappeared, replaced by the bees, butterflies, and other familiar pollinators that thrive in modern ecosystems.

Understanding this long history is essential—not just for scientists, but for anyone interested in biodiversity and conservation. When we focus only on present-day pollinators and plants, we miss a crucial part of the story. The fossil record helps us see how pollination has changed over time, which in turn can offer insights into how today’s ecosystems might respond to environmental pressures such as climate change and habitat loss.

Recognising the ancient history of insect pollination isn’t just an academic exercise—it has real-world implications. If we understand how pollination evolved and adapted to past environmental changes, we can better predict how it might shift in the future. Conservation efforts that aim to protect pollinators today can benefit from a long-term perspective, ensuring that we’re not just responding to recent trends but also considering deep-time ecological processes.

So the next time you see a bee visiting a flower, remember—you’re witnessing the latest chapter in a story that began hundreds of millions of years ago. The relationship between plants and pollinators is far older, more complex, and more fascinating than we ever imagined.

Here’s the reference with a link to the paper. It should be open access, but if you have problems obtaining it, send me a message via my Contact page:

Peris, D., Ollerton, J., Sauquet, H., Hidalgo, O., Peñalver, E., Magrach, A., Álvarez-Parra, S., Peña-Kairath, C., Condamine, F.L., Delclòs, X. & Pérez-de la Fuente, R. (2025) Evolutionary implications of a deep-time perspective on insect pollination. Biological Reviews (in press)

What are the limits to pollinator diversity? A new article poses the question

The most globally significant groups of pollinators are well known and have been studied for a long time: bees and wasps, flies, butterflies and moths, birds, bats and beetles are all familiar to those of us with an interest in pollination ecology. However, every few years a new type of pollinator or a novel pollination system is described from nature or from the fossil record, or we add further examples of previously neglected pollinator groups such as cockroaches.

This begs the question: how much is there still to discover? How close are we to describing the full diversity of animals that act as pollen vectors? Can looking at the past help us to predict what we might find in the future? That’s the topic of a Perspective article that I was invited to write for the special issue of the Journal of Applied Entomology on the theme of  The Neglected Pollinators that I mentioned last month. It’s a subject that I’ve thought about a lot over the last few decades and it was great to get an opportunity to air some ideas and speculation.

The article is open access and you can download a copy by following the link in this reference:

Ollerton, J. (2024) What are the phylogenetic limits to pollinator diversity? Journal of Applied Entomology (in press)

Here’s the abstract:

Although huge progress has been made over the past 200 years in identifying the diversity of pollinators of angiosperms and other plants, new discoveries continue to be made each year, especially in tropical areas and in the fossil record. In this perspective article I address the following questions: Just how diverse are the pollinators and what are the phylogenetic limits to that diversity? Which other groups of animals, not currently known to regularly engage with flowers, might be found to be pollinators in the future? Can we predict, from the fossil record and from discoveries in under-researched parts of the world, which animal groups might turn out in the future to contain pollinators? I also discuss why adding to our knowledge of plant–pollinator interactions is important, but also stress that an incomplete knowledge may not be a bad thing if it means that remote, inaccessible and relatively pristine parts of the world remain that way.

Speaking at Oxford Ornithological Society – 11th September

Later this month I’ve been invited by the Oxford Ornithological Society to give a talk about my new book Birds & Flowers: An Intimate 50 Million Year Relationship. The talk will summarise the main themes from the book, particularly the sheer diversity of birds that can act as pollinators, what it means for the ecology and evolution of flowers, why the conservation of such interactions matters, and the cultural significance of bird-flower interactions. I’ll also deal with the question of why Europe is so odd when it comes to the question of birds as pollinators.

The talk is on Wednesday 11th September at Exeter Hall, Kidlington, starting at 7.45 pm; it’s free to society members, and non-members are invited to make a donation. Do come along if you’re in the area!

More details here: https://www.oos.org.uk/programme.php

I’ll bring a few copies of Birds & Flowers and Pollinators & Pollination: Nature and Society if anyone wants to buy a signed book.

Also in the diary are talks at South Leicester Birdwatchers (13th November) and Northamptonshire Bird Club (5th March).

If you represent a birding club or natural history society and wish to book me for a talk, please get in touch via my Contact page.

A doubly-parasitic orchid? – China Diary 5

Walking into Kunming Institute of Botany yesterday morning, I passed a young guy who was carrying what I initially thought was a species of Orobanchaceae. I’ve a long-standing interest in the pollination ecology of these intriguing parasitic plants, so I stopped to have a chat. Turns out they were in fact orchids! Specifically, they were specimens of Gastrodia elata, one of the “potato orchids“, so named because those fat tubers are edible. They are widely used in South China – where they are known as Tianma, 天麻 – both as a food and medicinally. The tubers are eaten before the flowers are produced, and originally they were collected from the wild. But in the 1960s a Chinese botanist named Xuan Zhou discovered how to cultivate them and they are now grown in specialist nurseries. A fascinating account of the life of Xuan Zhou – “The Father of Gastrodia” – was published in the journal Plant Diversity last year, shortly after he died.

These orchids do not produce green leaves or stems, therefore they cannot photosynthesise. Instead, they gain all of their energy from a parasitic symbiotic relationship with a fungus – they are what is termed “myco-heterotrophic“. Most myco-heterotrophic plants have evolved from ancestors that were involved in mutualistic mycorrhizal relationships with fungi, in which the plant provides sugars to the fungus in return for mineral nutrients and water. In the case of Gastrodia elata, the fungus concerned is the non-mycorrhizal, wood-rotting Armillaria mellea. In the west we know this as Honey Fungus, a disease of trees and shrubs and the bane of many a gardener. This is also edible, incidentally, but best dried before cooking (and some have an intolerance to it, so take care).

I tweeted the photograph in a short thread just after taking it, and Stewart Nicol pointed me to a study of the orchid’s floral biology and pollination ecology in Japan by Naoto Sugiura. Turns out that, at least in the population which Naoto studied, the plant produces no nectar and deceives its pollinators, which are small bees, into visiting the flowers.

That’s why I’ve used the phrase “doubly-parasitic*” in the title of this post – the plant, it appears, parasitically exploits both the fungus from which it gains energy and the pollinators that ensure its reproduction. It’s (almost, but not quite) the flip side of “double mutualism” in which species provide two benefits for one another, e.g. the same bird is both a pollinator and a seed disperser of a particular plant, a phenomenon that I discussed in my recent book Birds & Flowers: An Intimate 50 Million Year Relationship.

But note the question mark in the title of this post. There’s an enormous amount that we don’t know about these myco-heterotrophic interactions and how they remain stable over the evolutionary history of the plant and the fungus. In order to be considered a parasite, by definition, an organism must have a negative impact on the reproductive fitness of its host. Do these orchids negatively impact either the fungus or the bees that pollinate it? As yet we don’t know. And I was intrigued by this comment from a 2005 review of ‘The evolutionary ecology of myco-heterotrophy‘ by Martin Bidartondo:

“no successful plant lineage would be expected to cheat both mycorrhizal fungi (by failing to provide photosynthates) and deceive insect pollinators (by failing to provide nectar or other rewards) due to the evolutionary instability inherent to specializing on two lineages.”

At first glance it appears that Gastrodia elata is a plant lineage that has done just that, though I’d like to see more work carried out on this system. Specifically, are all populations of the orchid bee pollinated and are all rewardless? And does this orchid really provide no benefit to the fungus, perhaps by synthesising secondary compounds that protect the Armillaria from infection by bacteria or being eaten by invertebrates. So many questions to be answered about this fascinating species interaction!

*With thanks to my wife Karin Blak for inspiring that phrase.

A new review of ‘Birds & Flowers’ in the Journal of Pollination Ecology

The reviews of Birds & Flowers: An Intimate 50 Million Year Relationship are starting to appear in blogs, magazines and journals. The latest, by Diane Campbell, has just been published in the Journal of Pollination Ecology and I’m so pleased that it was positive! I’ve only met Diane a couple of times at conferences but I have a lot of respect for her work. The review is fair and balanced, and gratifyingly enthusiastic, for example:

In this delightful book, [Ollerton] describes the ways that birds and flowers interact. As in his previous book, Pollinators & Pollination: Nature and Society, [he] takes a deeply personal approach to the subject. He combines anecdotes from his research travels around the world, to mountains of Kenya and Tanzania, the Andes of Peru, Brazil, and Nepal, among other places, with his contributions to, and masterful knowledge of, the recent literature…

The review is free to read and download from Journal of Pollination Ecology. I’m so glad that people are enjoying the book – if you’ve bought or borrowed a copy, please do leave a comment and let me know what you think.

The flower that’s pollinated by birds, bees….and the wind!

In my new book Birds & Flowers: An Intimate 50 Million Year Relationship I spend a bit of time discussing the idea of the bird pollination syndrome that we refer to as ‘ornithophily’, its limitations, and the fact that it has two distinct meanings that are often conflated. One of the problems with ornithophily, and indeed all of the syndromes, is that historically it’s sometimes blinkered scientists to the extent that they only look at the flower visitors that are “right” for the syndrome, ignoring the rest or dismissing them as “secondary pollinators”, a term I dislike.

Why do I dislike that term? Because it fails to capture the complexity of flower-pollinator interactions and relegates an important component of plant reproduction to a subsidiary role. I could go on about this at some length, but if you’re interested in discovering more, look at pages 62-65 of Birds & Flowers. There I contrast the classical Most Effective Pollinator Principle with the equally valid (but much less well studied) Least Effective Pollinator Principle, with a segue into one of my favourite tracks from Led Zeppelin’s second album: What is and What Should Never Be.

But back to the real subject of this post – a flower that corresponds to the classical bird pollination syndrome BUT is also pollinated by bees and (very surprisingly) wind! It’s such an interesting paper by Brazilian ecologists Amanda Pacheco, Pedro Bergamo & Leandro Freitas – here’s the reference and a link to the study:

Pacheco, A., Bergamo, P.J. & Freitas, L. (2024) An unexpected case of wind pollination: ambophily in an ornithophilous tropical mountaintop Orobanchaceae. Plant Systematics and Evolution 310, 9. https://doi.org/10.1007/s00606-024-01890-6

For over 100 years the classical pollination syndromes have acted as a framework for understanding the ecology and evolution of plant-pollinator interactions. But we’ve long known that while they can be a useful shorthand, they do not fully reflect the complexity of how pollination systems evolve. That shouldn’t surprise us because, as I point out in my two recent books, we have data (of any quality) on no more than 10% of the 350,000 or so species of flowering plants!

In addition, those plants for which we do have good data are NOT a random subset of the flowering plants: they have been specifically chosen by researchers because they look to be good systems with which to address particular ecological or evolutionary questions.

Which is fine, but we MUST recognise that this imposes significant restrictions on our understanding of the biodiversity of plant-pollinator interactions. The authors of this paper expressed it very well when they wrote that assumptions about:

“predictability may cause researchers to take for granted that only birds pollinate ornithophilous flowers, hindering research on the contribution of other vectors.”

To which I’d add: it also hinders our understanding of how these interactions evolve over long time scales and across multiple populations.

An obvious question is: how frequent are these sorts of complex pollination systems, involving different pollen vectors of an apparently specialised flower? The answer is that we simply don’t know, because most researchers would have not gone into this level of detail. So a huge congratulations to the authors for a great study – I hope it stimulates others to look beyond the ‘expected’ pollinators of flowers.

Photos: Nathália Susin Streher from the original paper.

Read my author interview and get a 25% discount off ‘Birds & Flowers’, ‘Pollinators & Pollination’ and other books from Pelagic Publishing!

I recently did a short interview with Pelagic Publishing’s marketing person, Sarah Stott, which you can read here: https://pelagicpublishing.com/blogs/news/birds-and-flowers-author-interview.

On that page you can sign up to Pelagic’s newsletter (which I STRONGLY recommend, because they produce some great natural history and science books, and not just mine!) and by doing so you can receive a 25% discount on all orders.

What are you waiting for?

Read a preview of my new book ‘Birds & Flowers: An Intimate 50 Million Year Relationship’

In the run up to release of my new book Birds & Flowers: An Intimate 50 Million Year Relationship, my publisher, Pelagic, has updated the book description and provided a preview of some of the pages and colour plates – follow this link to view them. On that page you can also pre-order the book direct from Pelagic Publishing, who will ship worldwide, or it’s available from many of the online book sellers.

I’m really excited to be sharing this with the readers of my blog and can’t wait for publication day! Early in 2024 I hope to do some talks, online and in person, to promote the book – so watch this space.

With best wishes to you all and hopes for a peaceful, and more sustainable, New Year.

Recent research and seminars on pollinators and pollination that have caught my eye

There’s so much good science and so many great talks coming out of the (broad) field of pollinator and pollination research at the moment! Here’s a few things that have come up on my radar. Feel free to comment and add your own examples of things I may have missed.