Category Archives: Mutualism

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!

Life brings stability: biological crusts on sandy subsoil

A couple of weeks ago we visited Karin’s family in Jutland and went for a couple of long walks around the area. One of these took us through some very nice mixed pine, oak, and birch forest close to a river. The forest was anchored into a thin horizon of mulchy topsoil, beneath which was almost pure sand, a post-glacial legacy of the wider, wilder rivers that ran through the region at the end of the last Ice Age.

Where our path ran parallel to the river I noticed that the exposed vertical sections were far from lifeless: the sandy faces had been colonised by algae, lichens, fungi, cyanobacteria, and mosses. These biological crusts had stabilised the sand and prevented it from eroding further back into the bank. On a miniature scale they were doing what forests and other vegetation does in mountainous areas all over the world: preventing landslides.

Biological crusts in turn provide opportunities for ferns and seed plants to germinate and gain a foothold: they are often the starting point for further ecological succession.

Not only are these crusts acting as substrate stabilisers and seed beds, but all of the usual ecological processes of photosynthesis, nutrient acquisition, decomposition, carbon storage, symbiosis and competition are taking place in just a few millimetres of biodiversity. There’s a lot going on in these thin veneers of life.

The chapter titles for my book: Pollinators & Pollination: Nature and Society

A few people have asked me about what’s covered in my book which is being published by Pelagic and is currently in production. Here’s the chapter titles:


1         The importance of pollinators and pollination                               

2         More than just bees: the diversity of pollinators                           

3         To be a flower                                                                                               

4         Fidelity and promiscuity in Darwin’s entangled bank                 

5         The evolution of pollination strategies                                              

6         A matter of time: from daily cycles to climate change                 

7         Agricultural perspectives                                                                        

8         Urban environments                                                                                  

9         The significance of gardens                                                                    

10      The shifting fates of pollinators                                                            

11      New bees on the block                                                                              

12      Managing, restoring and connecting habitats                                 

13      The politics of pollination                                                                        

14      Studying pollinators and pollination                                                  

As you can see it’s a very wide-ranging overview of the subject, and written to be accessible to both specialists and non-specialists alike. To quote what I wrote in the Preface:

“While the book is aimed at a very broad audience, and is intended to be comprehensible to anyone with an interest in science and the environment, and their intersection with human societies, I hope it will also be of interest to those dealing professionally with plants and pollinators. The subject is vast, and those working on bee or hoverfly biology, for example, or plant reproductive ecology, may learn something new about topics adjacent to their specialisms. I certainly learned a lot from writing the book.”

The book is about 100,000 words in length, lots of illustrations, and there will be an index. My copy editor reckons there’s 450 references cited, though I haven’t counted. I do know that they run to 28 pages in the manuscript, and that’s with 11pt text. All going well it will be published before Christmas.

Get a 30% discount if you pre-order my new book Pollinators & Pollination: Nature and Society


In the next few months my new book Pollinators & Pollination: Nature and Society will be published.  As you can imagine, I’m very excited! The book is currently available to pre-order: you can find full details here at the Pelagic Publishing website.  If you do pre-order it you can claim a 30% discount by using the pre-publication offer code POLLINATOR.

As with my blog, the book is aimed at a very broad audience including the interested public, gardeners, conservationists, and scientists working in the various sub-fields of pollinator and pollination research. The chapter titles are as follows:

Preface and Acknowledgements
1. The importance of pollinators and pollination
2. More than just bees: the diversity of pollinators
3. To be a flower
4. Fidelity and promiscuity in Darwin’s entangled bank
5. The evolution of pollination strategies
6. A matter of time: from daily cycles to climate change
7. Agricultural perspectives
8. Urban environments
9. The significance of gardens
10. Shifting fates of pollinators
11. New bees on the block
12. Managing, restoring and connecting habitats
13. The politics of pollination
14. Studying pollinators and pollination



The other pollinators: some recent videos that don’t focus on bees

The review of the biodiversity of pollinators that I published in 2017 estimated that on average about 18% of animal-pollinated plants within natural communities are specialised on bees. Bees also contribute to the reproduction of many of the plants that have generalist pollination systems, which account for perhaps 50% of plant species on average. But that stills leaves a significant fraction (maybe one third) that are specialised on the “other” pollinators, including flies, beetles, birds, bats, and so forth. There is growing awareness of how important these pollinators are for wild plant and crop pollination, but bees still hog most of the pollinator-related media.

In the last couple of weeks I’ve been sent links to videos that focus on these other pollinators so I thought I’d compile a list that show us something of the true diversity of animals that act as pollen vectors. Please add your own suggestions in the comments:

Elephant shrews, lizards, cockroaches*, crustaceans, and biting midges are covered in this SciShow video (HT Steve Hawkins)

Opossum pollination of a Brazilian plant is featured in this video (HT Felipe Amorim)

Here’s a recorded webinar on bird pollination by Dan Scheiman from Audubon Arkansas

A few videos on bat pollination by Jim Wolfe can be found here and here and here, and this is a short one that’s a supplement to a recent Journal of Applied Ecology paper on cactus pollination by Constance J. Tremlett et al.

The fascinating ecology of skunk cabbage (Symplocarpus foetidus), including fly and possibly beetle pollination, is the topic of this video.

Fly pollination is also highlighted in this short piece by the Natural History Museum, and this one deals with drone flies as managed pollinators for agriculture in New Zealand.


*Watch out for my report on a newly discovered cockroach-pollinated plant….hopefully coming later this year…..

Recent reviews in pollination biology: an annotated list: UPDATED x 3

2019-02-09 13.47.49

As it’s my birthday today, I thought I’d reward myself by completing a blog post that I started just after Christmas and never got round to finishing.  Review articles that summarise recent developments in a field are an important contribution to the scientific literature that allow us to pause and reflect on where a topic has been and where it is headed.  Having recently (co)authored a couple of reviews I can attest that they are useful in this respect for both the writers and for the readers.

In the past couple of years quite a number of critical and timely reviews have been published which are proving very useful to me: I’m currently writing a book and these reviews have been invaluable in summarising aspects of a field that is currently publishing in excess of 1000 research papers per year. So I thought I’d bring them together into a single listing with a short commentary on each.  No doubt I have missed many other reviews so please feel free to point out any gaps and I will update the list as I go along.

Each review is hot linked to the source; a good proportion of the reviews are open access, notably those from the recent special issue of Annals of Botany devoted to the ecology and evolution of plant reproduction.  Some reviews are very focused, but most are quite broad.  Several of these complement one another.  I hope you find them interesting and useful.

Barrett, S. & Harder, L. (2017) The ecology of mating and its evolutionary consequences in seed plants. Annual Review of Ecology, Evolution and Systematics 48: 135-157

Mating systems, i.e. who breeds with whom, are just as complex in plants as they are in animals.  However some features of seed plants, such as the fact that they don’t move, that most species have both male and female functions, and that their growth is modular and often indeterminate, represent significant challenges that have been overcome in a bewildering variety of ways.


Braun, J. & Lortie, C.J. (2018)  Finding the bees knees: A conceptual framework and systematic review of the mechanisms of pollinator-mediated facilitation.  Perspectives in Plant Ecology, Evolution and Systematics 36: 33-40

In a community, if one plant species positively affects another, we term this “facilitation”.  It can occur at a variety of life stages, including reproduction whereby the presence of one species increase the likelihood of another species being pollinated.  This review shows that it occurs fairly frequently at a variety of spatial scales, but there are still significant gaps in our understanding of the phenomenon.


Fuster, F., Kaiser‐Bunbury, C., Olesen, J.M. & Traveset, A. (2018) Global patterns of the double mutualism phenomenon. Ecography

When species provide benefits to one another in two different ways, for example an animal is both a pollinator and a seed disperser of a plant species, we refer to it as a “double mutualism”.  As this fascinating review shows, double mutualisms are very uncommon, but they are widespread, and probably under-recorded.


Minnaar, C., Anderson, B., de Jager, M.L. & Karron, J.D. (2019) Plant–pollinator interactions along the pathway to paternity. Annals of Botany 123: 225-245 

The male aspect of plant reproduction, i.e. pollen donation, is often neglected when we consider how pollination systems evolve.  This review provides as up to date account of where we are in understanding how paternity influences floral characters such as shape and colour.


Ollerton, J. (2017) Pollinator diversity: distribution, ecological function, and conservation. Annual Review of Ecology, Evolution and Systematics 48: 353-376

A very broad over view of our current understanding of the biodiversity of pollinators, taking a deep time and a wide spatial perspective to put current concerns about loss of pollinators into a wider perspective.


Parachnowitsch, A.L., Manson, J.S. & Sletvold, N. (2019) Evolutionary ecology of nectar. Annals of Botany 123: 247–261 

We often take nectar for granted – it’s just sugar and water, isn’t it?  As this review shows, nectar is dynamic and complex, and affects a range of ecological functions beyond just providing pollinators with a reward.  However there’s still a huge amount we don’t understand about how nectar traits evolve.


Toledo-Hernández, M., Wangera, T.C. & Tscharntke, T. (2017) Neglected pollinators: Can enhanced pollination services improve cocoa yields? A review.  Agriculture, Ecosystems and Environment 247: 137-148

Chocolate is most people’s favourite confectionery and is famously pollinated only by small midges.  Or is it? As this review shows, lots of other insects visit cocoa flowers, but their role as pollinators has not been well studied.


Vizentin-Bugoni J, PKM Maruyama, CS Souza, J Ollerton, AR Rech, M Sazima. (2018) Plant-pollinator networks in the tropics: a review. pp 73-91 In Dáttilo W & V. Rico-Gray. Ecological networks in the Tropics. Springer.

This book chapter that I co-authored with some very energetic and creative young Brazilian researchers summarises what’s currently known about plant-pollinator interaction networks in tropical communities.  One of the conclusions is that they are really not so different to those in temperate and subtropical biomes.


Wright, G.A., Nicolson, S.W. & Shafir, S. (2018) Nutritional Physiology and Ecology of Honey Bees. Annual Review Entomology 63:327-344

A review of how bees use nectar and pollen at the level of both the individual and the colony, focused on the most widespread of pollinator species.


As expected, several people have told me about reviews I’d missed, and in some cases ones that I had read but forgotten about!  I’ll list them below, though without annotations:

Bennett, J. et al. (2018) A review of European studies on pollination networks and pollen limitation, and a case study designed to fill in a gap, AoB Plants 10:

Knight, T. et al. (2018) Reflections on, and visions for, the changing field of pollination ecology. Ecology Letters 21: 1282-1295

Vallejo-Marin, M. (2018) Buzz pollination: studying bee vibrations on flowers. New Phytologist



I had deliberately restricted the reviews to 2017 onwards, but via email David Inouye kindly sent a few older ones through which are equally useful:

Brosi, B. J. (2016) Pollinator specialization: from the individual to the community. New Phytologist: 210: 1190–1194

Hahn, M. and C. A. Brühl (2016) The secret pollinators: an overview of moth pollination with a focus on Europe and North America. Arthropod-Plant Interactions: 1-8

Inouye, D. W., et al. (2015) Flies and flowers III: Ecology of foraging and pollination. Journal of Pollination Ecology 16



A more recent addition to this set of reviews was sent to me by Anne-Laure Jacquemart.  Although it’s focused just on one (rather variable) crop, I think it will be really useful for anyone interested in the pollination biology of crop plants:

Ouvrard, P. & Jacquemart, A.-L. (2019) Review of methods to investigate pollinator dependency in oilseed rape (Brassica napus).  Field Crops Research 231: 18-29




The biological mutualisms at the heart of sourdough bread


During the road trip to Denmark that I mentioned in a post back in September – see “There ain’t no b(ee) in Starbucks” – my wife Karin received a special gift from her sister Pia.  It was a small jar containing a starter culture for sourdough bread, a culture that Pia has been using since she received it from a friend, who long ago received it from another friend.  I didn’t know much about sourdough bread and did some reading. That Wikipedia link is a good introduction but don’t be put off by the complexities of “refreshment” – we’ve kept the starter culture in the fridge since early September and it’s been fine.  Karin used the culture for the first time this morning and made the rye bread you see above.

But on to the biology.  In essence the sourdough culture is a mix of wild lactic acid bacteria and wild yeasts, plus flour and water.  When added to the bread mix (which in our case contained water, salt, seeds and molasses, as well as rye flour) the yeasts feed on some of the sugars within the mix and the lactic acid bacteria feed on other sugars that the yeast cannot metabolise.  During that bacterial fermentation, byproducts are also produced on which the yeasts feed.  The yeasts in turn produce carbon dioxide which serves to leaven the dough, and the bacteria produce lactic acid as another byproduct, which gives the bread its slightly sour flavour.  This lactic acid also lowers the pH of the environment and, together with the production of anti-fungal chemicals, the lactic acid bacteria prevent the growth of other bacteria and moulds.  The yeasts, however, can tolerate these conditions and they thrive.

At least six species of yeast and 25 species of lactic acid bacteria have been shown to be  involved in this process, often as multi-species mixtures.  The exact biodiversity of the culture is dependent upon its source: micro-organisms vary a lot across the world.  But the heart of the relationship between yeasts and bacteria is always the same: they each facilitate the growth and reproduction of the other, and so the relationship is mutualistic, much like (most) relationships between plants and pollinators, birds and berries, and sea anemones and clownfish.

Of course there is a third organism involved in this mutualism: Homo sapiens.  By producing the resources on which these organisms feed, and then distributing the starter culture, we are providing the right conditions for the yeast and lactic acid bacteria to increase their populations.  In turn the yeast and bacteria play an important role in producing food for us, and in fact this way of making bread is thousands of years old.  Microorganisms and people all benefit: what could be more mutualistic than that?  Indeed, these interactions could be classified as a rare example of a ménage à trois mutualism.

There’s also a social-cultural dimension to all of this as the passing of gifts such as the starter culture binds friendships.  If any of our local friends are reading we’d be happy to share the sourdough culture once we’ve bulked it up.  The bread that it makes is delicious and from now on we’re going to try to give up buying the shop-bought kind.

If you want to read more about all of this, and have a try at making your own starter culture from scratch, there’s some great information and links on the Microbial Menagerie blog.

Many thanks to Pia for sharing the starter culture, and to Karin for baking the bread!

Trait evolution, resource specialization and vulnerability to plant extinctions among Antillean hummingbirds – a new study just published

Hummingbird bowl from BM

Hummingbirds are fascinating creatures and important pollinators for a wide range of plants in the New World (and, historically, possibly in the Old World – see this post from 2014: There were hummingbirds over the White Cliffs of Dover).  During the last decade I have been involved in some hummingbird-related research with several colleagues, particularly Dr Bo Dalsgaard and Dr Stella Watts, and it’s generated some really interesting findings about the biogeography, macroecology, and interactions with plants of these most elegant of birds.

The latest installment of this work is a test of some ideas relating to the vulnerability of hummingbirds on islands to the extinction of their plant partners.  It’s just been published and the reference is:

Dalsgaard B., Kennedy J.D., Simmons B.I., Baquero A.C., Martín González A.M., Timmermann A., Maruyama P.K., McGuire J.A., Ollerton J., Sutherland W.J. & Rahbek C. (2018) Trait evolution, resource specialization and vulnerability to plant extinctions among Antillean hummingbirds. Proceedings of the Royal Society series B (in press)

Here’s the abstract:

Species traits are thought to predict feeding specialization and the vulnerability of a species to extinctions of interaction partners, but the context in which a species evolved and currently inhabits may also matter. Notably, the predictive power of traits may require that traits evolved to fit interaction partners. Furthermore, local abiotic and biotic conditions may be important. On islands, for instance, specialized and vulnerable species are predicted to be found mainly in mountains, whereas species in lowlands should be generalized and less vulnerable. We evaluated these predictions for hummingbirds and their nectar-food plants on Antillean islands. Our results suggest that the rates of hummingbird trait divergence were higher among ancestral mainland forms before the colonization of the Antilles. In correspondence with the limited trait evolution that occurred within the Antilles, local abiotic and biotic conditions—not species traits—correlate with hummingbird resource specialization and the vulnerability of hummingbirds to extinctions of their floral resources. Specifically, hummingbirds were more specialized and vulnerable in conditions with high topographical complexity, high rainfall, low temperatures and high floral resource richness, which characterize the Antillean Mountains. These findings show that resource specialization and species vulnerability to extinctions of interaction partners are highly context-dependent.

As always I’m happy to send a PDF to anyone who drops me an email.

Plant-pollinator networks in the tropics: a new review just published.


As an ecologist who has carried out field work in the temperate zone (UK), the subtropics (Tenerife and South Africa) and the tropics (parts of South America, Africa and Australia)  I’ve always found the idea that the study of ecology can be divided into “tropical” and “non-tropical” a bit odd.  It’s as if the way that the natural world works somehow changes at about 23 degrees north or south of the equator, making things “different” around the equator.  The tropics are a very special, diverse place, it’s true, but so are many places outside the tropics.

With this in mind I was pleased when I was asked by some of my Brazilian colleagues to contribute to a chapter in a new book entitled Ecological Networks in the Tropics. It was an opportunity to review what is known about plant-pollinator networks in the tropics and the ways in which they are very similar to such networks at lower latitudes. Here’s the details of the chapter, followed by the abstract.  If anyone wants a copy please drop me an email:

Vizentin-Bugoni J, PKM Maruyama, CS Souza, J Ollerton, AR Rech, M Sazima. (2018) Plant-pollinator networks in the tropics: a review. pp 73-91 In Dáttilo W & V. Rico-Gray. Ecological networks in the Tropics. Springer.


Most tropical plants rely on animals for pollination, thus engaging in complex interaction networks. Here, we present a global overview of pollination networks and point out research gaps and emerging differences between tropical and non-tropical areas. Our review highlights an uneven global distribution of studies biased towards non-tropical areas. Moreover, within the tropics, there is a bias towards the Neotropical region where partial networks represent 70.1% of the published studies. Additionally, most networks sampled so far (95.6%) were assembled by inferring interactions by surveying plants (a phytocentric approach). These biases may limit accurate global comparisons of the structure and dynamics of tropical and non-tropical pollination networks. Noteworthy differences of tropical networks (in comparison to the non-tropical ones) include higher species richness which, in turn, promotes lower connectance but higher modularity due to both the higher diversity as well as the integration of more vertebrate pollinators. These interaction patterns are influenced by several ecological, evolutionary, and historical processes, and also sampling artifacts. We propose a neutral–niche continuum model for interactions in pollination systems. This is, arguably, supported by evidence that a high diversity of functional traits promotes greater importance of niche-based processes (i.e., forbidden links caused by morphological mismatching and phenological non-overlap) in determining which interactions occur, rather than random chance of encounter based on abundances (neutrality). We conclude by discussing the possible existence and direction of a latitudinal gradient of specialization in pollination networks.

Local and regional specialization in plant–pollinator networks: a new study just published

Euphorbia canariensis pollinators 2016-04-29 17 58 00

A fundamental feature of the natural world is that no species exists in isolation: all organisms interact with other organisms during their lives. These interactions take many forms and the outcome varies with the type of interactions. For example predator-prey interactions are clearly negative for the prey species, but positive for the predator. Other interactions result in positive outcomes for both species, including relationships between pollinators such as bees, birds and flies, and the flowers that they pollinate. An important feature of such interactions is how specialized or generalized it is; that is, how many different pollinators are actually involved in pollinating a particular type of flower, or how many types of flower does a specific pollinator visits.

In a newly published study, I have collaborated with colleagues from Denmark and Brazil to assess how local specialization (within a community) relates to regional specialization (across communities) using two separate data sets from the Brazilian rupestrian grasslands and Canary Island/North African succulent scrub vegetation.

Here’s the citation with a link to the paper (drop me a line if you can’t access it and need a PDF):

Carstensen, D.W., Trøjelsgaard, K., Ollerton, J. and Morellato, L.P.C. (2017) Local and regional specialization in plant–pollinator networks. Oikos (in press) doi:10.1111/oik.04436

The abstract is as follows:

“Specialization of species is often studied in ecology but its quantification and meaning is disputed. More recently, ecological network analysis has been widely used as a tool to quantify specialization, but here its true meaning is also debated. However, irrespective of the tool used, the geographic scale at which specialization is measured remains central. Consequently, we use data sets of plant–pollinator networks from Brazil and the Canary Islands to explore specialization at local and regional scales. We ask how local specialization of a species is related to its regional specialization, and whether or not species tend to interact with a non-random set of partners in local communities. Local and regional specialization were strongly correlated around the 1:1 line, indicating that species conserve their specialization levels across spatial scales. Furthermore, most plants and pollinators also showed link conservatism repeatedly across local communities, and thus seem to be constrained in their fundamental niche. However, some species are more constrained than others, indicating true specialists. We argue that several geographically separated populations should be evaluated in order to provide a robust evaluation of species specialization.”

This is what those two different habitats look like:

If you would like more information on plant-pollinator networks, including details of an edible game for Christmas (!), follow this link to the standingoutinmyfield blog.