Category Archives: Biogeography

Monarchs and Milkweeds Workshop summary, Oak Spring, Virginia, June 2019

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As I recounted in my last post about a train ride through American climate change, my wife Karin and I have been in the USA for the past couple of weeks, visiting colleagues in the west and ultimately heading eastwards to Virginia for a workshop on monarch butterflies and their milkweed hosts.  The meeting was organised by Anurag Agrawal, professor at Cornell University and author of the recent book Monarchs and Milkweed, highly recommended to anyone interested in the natural history of plants and insects.  The monarch (Danaus plexippus) is an iconic migrating  species that travels from Mexico to Canada and back, over the course of a few generations.  This behaviour, and their vast over-wintering assemblages, have become the focus of intense efforts to understand their ecology and biology.  Their caterpillar host plants are mainly milkweeds (Asclepias spp.) and bringing together both plant and animal scientists is important for gaining a fuller over view of the issues facing the monarchs and the milkweeds, and how both can be conserved in a time of anthropogenic change.

The venue for the workshop was Oak Spring, Upperville, the former home of Paul and Rachel (“Bunny”) Mellon which has been turned into the base of operations for a philanthropic foundation specialising in plant science, horticulture, and botanical art.  The Oak Spring Garden Foundation (OSGF) is “dedicated to inspiring and facilitating scholarship and public dialogue on the history and future of plants, including the culture of gardens and landscapes and the importance of plants for human well-being”.  The OSGF generously funded the workshop, including accommodation and travel for participants.  This brought together a small group of scientists from the USA, the UK and Brazil, together with an artist, a milkweed horticulturalist, and two science writers.  Their brief was to discuss the latest developments in our understanding of monarch butterflies, their decline and conservation, and the taxonomy, evolution and ecology of milkweeds and the wider groups of Lepidoptera and the plant family Apocynaceae to which these organisms belong.  My invitation to take part was due to the research on the pollination ecology of this family I’ve conducted, spanning about twenty five years and culminating in a recently published assessment of the diversity of pollination systems in Apocynaceae.

First things first: Oak Spring is one of the most tranquil, beautiful, and inspiring places where it’s ever been my privilege to stay.  Here’s a few photographs, but they really do not do justice to the buildings and garden, their setting, nor to the unique atmosphere of Oak Spring.

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So to the science.  The workshop started with a set of short presentations on our recent research findings and the motivations for our interests in these organisms.  On the second day we then moved on to discussing ideas for future collaborations between the participants and how that work might be funded in the future.  Presentations and discussions were mainly held in the Basket House, named for obvious reasons:

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Each of us was also interviewed on camera to build an online archive of the work we do and why we do it.

The advantage of face-to-face meetings such as this, and why Skype and so forth can never fully replace them, is the free-flowing conversations that occur within the formal sessions and outside them.  Among the many things that I learned from presentations and discussions were:

  • The California monarch population has declined by almost 90% this year and there’s an urgent need to understand why this has happened.  Climate change has been implicated, especially in relation to the increased frequency of wildfires in this region.
  • Existing methods of nectar extraction from milkweed flowers may strongly underestimate the volume available to flower visitors, and overestimate the sugar concentration.  Using a small centrifuge to spin out the nectar seems to be the most effective method.
  • Asclepias arrived in the Americas (probably from Africa) some 10 million years ago (mya).  However Danaus only arrived about 3.7 mya, so there was a long period of time in which the plant was not co-evolving with one of its major herbivores.
  • There is strong evidence of migrations along the Andes by a close relative of the monarch, Danaus erippus.  Migrations in this group of butterflies therefore extends beyond the iconic D. plexippus.
  • Sonoran Desert Asclepias are sister group to the rest of the New World Asclepias spp.  The exact route by which the African ancestors made it to the Americas is unknown, it could be via Asia and the Bering Strait, or across the Atlantic by way of island stepping stones.  Either way, the phylogenetic position of the Sonoran milkweeds implies that a lot of Asclepias species have gone extinct over the past 10 million years.
  • Climate change seems to be resulting in more complex and unpredictable windows of opportunity for monarch egg laying and caterpillar development.  The monarchs are most successful in late spring and late summer, but not in all years.
  • Likewise, extreme precipitation of the kind I recently documented in the USA is also likely to have a negative impact on the monarchs and their host plants.
  • There is molecular evidence that monarch butterflies went through a huge genetic bottleneck in the 1960s-1970s, for reasons that are not altogether clear.

All of these findings, and more that there isn’t space to document, point to a need for further research to better understand these organisms if we wish to secure their futures.

By the end of the workshop we had made some concrete decisions on future steps:

  •  The African members of the genus Asclepias, plus about 20 other closely related genera, require more critical taxonomic and phylogenetic assessment in order to understand their systematic relationship to the North and South American Asclepias species.
  • A poster (or possibly series of posters) will be produced that explain the ecology of the monarch, its relationship with milkweeds, the patterns of migration, and the value of milkweeds as nectar sources for a diverse range of pollinators.
  • We will explore a multi-agency grant application to further develop the collaborations between participants.

The final day of the workshop involved a field trip around Virginia to see some of the local milkweed species, many of which live in woodland.  That surprised me: I always envision Asclepias spp. as grassland or desert plants.  The leader of the field trip, Mark Fishbein, had a hit list of 8 species that he wanted us to see and in the end we located all of them, including a rare hybrid population of A. syriaca x A. exaltata, plus the tropical milkweed Asclepias curassavica planted in the OSGF garden, plus the distant relative dogbane Apocynum cannabinum.  Here are some images from that day:

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Searching for milkweeds along Skyline Drive, Shenandoah National Park

 

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Poke milkweed – Asclepias exaltata

 

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Hunting that elusive hybrid milkweed!

 

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Caterpillar of the monarch butterfly feeding on a milkweed

 

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Alessandro Rapini intent on getting a good photo of the A. syriaca x A. exaltata hybrid

 

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A bumblebee and a butterfly visiting A. exaltata

 

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Purple milkweed – Asclepias purpurascens

 

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Common milkweed – Asclepias syriaca – with a visiting skipper butterfly

 

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Climbing milkvine – Matelea obliqua – a member of a largely fly-pollinated group of New World asclepiads

 

Thanks to my fellow workshoppers for such a stimulating and enjoyable meeting, and to all the staff at Oak Spring for making us feel so welcome.  Particular thanks go to Prof. Sir Peter Crane who, as President of the Oak Spring Garden Foundation, was hugely supportive of the workshop, and to Angie Ritterpusch, Head of Events and Guest Services, for logistical and organisational support.

 

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Filed under Biodiversity, Biogeography, Butterflies, Climate change, Evolution

Are these first photographs of a living specimen of a rare African butterfly?

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Earlier this year my colleague at the Royal Botanic Gardens Kew, David Goyder, tweeted a link to a new book about the biodiversity of Angola which you can download for free by following this link.  David’s an authority on Apocynaceae, the family of plants on which I’ve also worked for many years (see this recent post), and has been sorking in Angola in recent years on a large biodiversity project.  So I was interested to see what was in the chapter  he had co-authored called “The flora of Angola: Collectors, Richness and Endemism“.  I was immediately struck by one of the images in Figure 5.3 showing an unnamed butterfly feeding on the flower of a species of Apocynaceae (Raphionacme michelii).

I made a note to myself to talk to David about adding the record to our Pollinators of Apocynaceae Database. But before I had a chance to do that, another apocynologist colleague, Ulrich Meve in Bayreuth, forwarded the chapter with a similar idea in mind.

We emailed David about the image and he sent us originals, but confessed he didn’t know what the insect was.  So I uploaded it to an African Lepidoptera forum on Facebook.  At which point a wave of excitement broke, because after some discussion as to whether it might be a new species, it turned out that the most likely candidate was an exceptionally rare butterfly called Acraea mansya in the family Nymphalidae.

According to Dominique Bernaud, an authority on the group, this species is hardly known beyond a few collections and he has never seen a photograph of a living specimen: if you follow this link you will see that the known distribution of the species does not include Angola,  and indeed it is not listed in the chapter on butterflies in the Angola biodiversity book.  So this is a new country record and (we think) the first images of living insects: so a double first for a beautiful species.

Here’s links to collection information for the plant and to David’s checklist of plants from the region, which gives details of the vegetation and the habitat.

An unanswered question, of course, is whether the butterfly is a pollinator of this species of plants.  Raphionacme belongs to a subfamily of Apocynaceae that have hardly been studied from the perspective of pollination ecology, so we simply don’t know.  Hopefully someone in the future will visit this remote region of Africa and find out!

Thanks to David for sending the images (a complete set of which is below), the National Geographic Okavango Wilderness Project, and the Wild Bird Trust, Parktown, South Africa.

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Filed under Apocynaceae, Biodiversity, Biogeography, Butterflies, Pollination

Pollinator availability, mating system and variation in flower morphology in a tropical savanna tree – a new, open-access study

Curatella image by Pedro Lorenzo

Widespread plant species can encounter a variety of different pollinators across their distributional range.  This in turn can result in local adaptation of flowers to particular pollinators, or to an absence of pollinators that results in adaptations for more self pollination.   A newly published study by one of my former PhD students, André Rodrigo Rech in Brazil, has looked at this in the widespread South American savanna tree Curatella americana.  André studied 10 populations separated in space by thousands of kilometres, in cerrado vegetation, one of the most threatened habitat types in Brazil.  Here’s the abstract:

Widely distributed organisms face different ecological scenarios throughout their range, which can potentially lead to micro-evolutionary differentiation at specific localities. Mating systems of animal pollinated plants are supposed to evolve in response to the availability of local pollinators, with consequent changes in flower morphology. We tested the relationship among pollination , mating system, and flower morphology over a large spatial scale in Brazilian savannas using the tree Curatella americana (Dilleniaceae). We compared fruit set with and without pollinators in the field, and analyzed pollen tube growth from self- and cross-pollinated flowers in different populations. Populations with higher natural fruit set also had lower fruit set in bagged flowers, suggesting stronger barriers to self-fertilization. Furthermore, higher levels of autogamy in field experiments were associated with more pollen tubes reaching ovules in self-pollinated flowers. Morphometric studies of floral and leaf traits indicate closer-set reproductive organs, larger stigmas and smaller anthers in populations with more autogamy. We show that the spatial variation in mating system, flower morphology and pollination previously described for herbs also applies to long-lived, perennial tropical trees, thus reemphasizing that mating systems are a population-based attribute that vary according to the ecological scenario where the plants occur

Here’s the full citation with a link to the paper which is open access:

Rech, A.R., Ré Jorge, L., Ollerton, J. & Sazima, M. (2018) Pollinator availability, mating system and variation in flower morphology in a tropical savannah tree. Acta Botanica Brasilica (in press)

The illustration of Curatella americana  and its pollinators is by Pedro Lorenzo.

This paper is a contribution to a special issue of Acta Botanica Brasilica dedicated to floral biology and pollination biology in Brazil It’s all open access and if you follow that link you can download the papers.

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Filed under Bees, Biodiversity, Biogeography, Brazil, Evolution, Pollination

The evolution of pollination systems in one of the largest plant families: a new study just published – download it for free

Figure 1 JUNE revision

Interactions between flowering plants and the animals that pollinate them are known to be responsible for part of the tremendous diversity of the angiosperms, currently thought to number at least 350,000 species.  But the diversity of different types of pollination system (bird, bee, moth, fly, etc.) is unknown for most large, related groups of plants (what systematists term “clades”) such as families and subfamilies.  In addition we know little about how these interactions with pollinators have evolved over time and in different parts of the world.  Only a handful of groups of flowering plants have been studied with respect to questions such as:

How much do we currently know about the diversity of pollination systems in large clades?

How is that diversity partitioned between the smaller clades (e.g. subfamilies, tribes, genera) of a family, and what are the evolutionary transitions between the major groups of pollinators?

Do these pollination systems vary biogeographically across the clade’s range?

These sorts of questions have been addressed for the massive, globally distributed Apocynaceae (one of the top 10 or 11 largest angiosperm families with more than 5,300 species) in a study just published using a new database of pollinators of the family.  What’s more, the work is open access and anyone can download a copy for free.  Here’s the citation with a link to the paper:

Ollerton, J., Liede-Schumann, S., Endress, M E., Meve, U. et al. [75 authors in all] (2019) The diversity and evolution of pollination systems in large plant clades: Apocynaceae as a case study. Annals of Botany 123: 311–325

In this study we have shown that (among other things):

  • The family is characterised by an enormous diversity of pollination systems involving almost all of the major pollen vectors and some that are nearly unique to the Apocynaceae.
  • Earlier diverging clades have a narrower range of pollination systems than those that evolved later.
  • Transitions from one type of pollination system to another are evolutionarily constrained, and rarely or never occur, whereas others have taken place much more often, e.g. between wasp and beetle pollination.
  • There is significant convergent evolution of pollination systems, especially fly and moth pollination, by geographically and phylogenetically distinct clades.

You’ll notice that there are 75 (!) authors on this paper.  That’s because we’ve pulled together a huge amount of previously unpublished data and used some state of the art analyses to produce this work.  It was a monumental effort, especially considering that my colleague Sigrid Liede-Schumann and I only decided to push ahead with this project about a year ago when we chatted at the International Botanical Congress that I posted about at the timeIn truth however the origins of this paper go back over 20 years to 1997 when when Sigrid and I published a study of what was then known about pollination systems in the Asclepiadaceae (the asclepiads).

In that paper we said that the research “is intended to be ongoing…[we]…hope to re-review asclepiad pollination within the next decade”.  At the time I didn’t think it would actually take more than 20 years!  However over that period a lot has changed.  For one thing the Asclepiadaceae no longer exists, broken up and subsumed within a much larger Apocynaceae.  Also, I’ve done a lot of work in the field and in the herbarium on some of the smaller groups within the family, such as CeropegiaOthers, including many of my co-authors, have also been working on different groups in various parts of the world.  Finally the level of sophistication of the analyses we are now able to do has increased beyond recognition compared to what we could achieve in the mid-1990s.  All of this means that now is the right time to produce this study.

Having said all of that, this is still a work in progress.  Our Pollinators of Apocynaceae Database contains a sample of just over 10% of the species in the family.  So lots more data on plant-pollinator interactions needs to be collected before we say we fully understand how pollination systems have evolved in this most remarkable family.  I’d be happy to talk with anyone who is interested in the family and being involved in future data collection.

The database will be freely available to anyone who wants to use it – lots more can be done with this information and, once again, I’m happy to chat with potential collaborators.

I was recently interviewed about the study, and about plant-pollinator interactions and the Apocynaceae more generally, for the In Defense of Plants podcast – here’s a link to that interview.

Finally, I’d like to express my sincerest thanks to my co-authors on this study – I really couldn’t have done it without you guys!

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Filed under Apocynaceae, Biodiversity, Biogeography, Evolution, Pollination, Wasps

Speaking about plant-pollinator research and science blogging in Wageningen in May

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If any of you are near Wageningen University on 31st May I’m giving a talk about some of our recent research called “The macroecology and macroevolution of plant-pollinator interactions”.  It’s preceded by a workshop on the whys and hows of science blogging.  Details are in the poster.

Here are the abstracts for the talk and the workshop:

Macroecology and macroevolution of plant-pollinator interactions

Plant-pollinator relationships are an ecologically critical form of interaction that ensures the long-term survival of the majority of the world’s plants species, and contribute to a large fraction of global agricultural output.  In additiondiversity and abundance of biotically pollinated plant species can be an important determinant of the diversity of animals at higher trophic levels.

Despite that global significance, most studies of plant-pollinator interactions are done at a local level, involving populations and communities of species, over modest time scales.  The ways in which these local sets of interactions scale up to produce global macroecological and macroevolutionary patterns, and the processes underpinning them, will be explored using two case studies.

The first is a data set of 67 plant communities, ranging from 70ºN to 34ºS, with which we investigated the roles of biotic and abiotic factors as determinants of the global variation in animal versus wind pollination.  Factors such as habitat type, species richness, insularity, topographic heterogeneity, current climate and late-Quaternary climate change were investigated. The predictive effects of these factors on the proportion of wind- and animal-pollinated plants were examined (see: Rech et al. 2016 – Plant Ecology & Diversity 9: 253-262).

Since these results were published  we have increased the number of plant communities in our database to >90, and our findings seem to be robust to these additional data.  The dominant influence of contemporary climate on the relative importance of wind-pollinated species suggests that communities may be sensitive to future climate change.  Communities in areas that are predicted to become drier may in time contain more wind-pollinated plants which may in turn reduce the diversity of pollinator species that are present.  There may also be implications for the prevalence of human pollen allergies.  Future work will focus on these two areas.

The second case study uses a newly assembled database of pollinators of the family Apocynaceae (one of the ten largest families of flowering plants), supported by a molecular phylogeny of the major clades.  This database has been used to explore phylogenetic and biogeographic patterns of pollinator exploitation (Ollerton et al. in review).  The findings from this study challenge some long-held assumptions about convergent evolution, the role of rewards such as nectar, and the notion that some specialised pollination systems are evolutionary “dead ends”.  It also highlights the function of novel floral features in determining pollinator type and behaviour, such as the fused gynostegium and pollinia found in the subfamily Asclepiadoideae.  In summary, Apocynaceae is emerging as an important model family for understanding the ecology and evolution of plant-pollinator interactions.

 

Blogging for EEB: why bother?

A growing number of scientists in Ecology and Evolutionary Biology (EEB) have their own blogs or post as guests on others’ blogs.  In this workshop we will explore motivations and strategies for blogging, and its advantages for early career researchers.  Why blog?  What does it do for one’s career?  Is it a distraction from actually doing science?  How does one build a blog readership?  We will also focus on two aspects that are sometimes seen as mutually exclusive: blogging as science outreach to the general public (sci-communication), versus blogging with other professional scientists in mind (sci-community).  As preparation for the seminar please read Saunders et al. (2017) Bringing ecology blogging into the scientific fold: measuring reach and impact of science community blogs

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Filed under Biodiversity, Biogeography, Macroecology, Pollination

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

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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.

Abstract:

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.

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Filed under Bees, Biodiversity, Biogeography, Macroecology, Mutualism, Pollination

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.

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Filed under Bees, Biodiversity, Biogeography, Brazil, Macroecology, Mutualism, Pollination, Tenerife

Pollinator biodiversity and why it’s important: a new review just published – download it for free

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In a new review paper that’s just been published in the Annual Review of Ecology, Evolution and Systematics I have looked at the question of just how diverse the pollinators are, and why pollinator biodiversity is ecologically important and therefore worthy of conservation.  I’ve taken a deep time and wide space approach to this, starting with what the fossil record tells us about when animal pollination evolved and the types of organisms that acted as pollinators in the past (the answer may surprise you if you’re unfamiliar with the recent paleontological literature on this topic).  Some of the most prominent biogeographical patterns have been highlighted, and I have tried to estimate the global diversity of currently known pollinators.  A conclusion is that as many as 1 in 10 described animal species may act as pollen vectors.

As well as this descriptive part of the review I’ve summarised some recent literature on why pollinator diversity matters, and how losing that diversity can affect fruit and seed set in natural and agricultural contexts.  Extinction of pollinator species locally, regionally, and globally should concern us all.

Although I was initially a little worried that the review was too broad and unfocused, having re-read it I’m pleased that I decided to approach the topic in this way.  The research literature, public policy, and conservation efforts are currently moving at such a fast pace that I think it’s a good time to pause and look at the bigger picture of what “Saving the Pollinators” actually means and why it’s so important.  I hope you agree and I’d be happy to receive feedback.

You can download a PDF of the review entitled Pollinator Diversity: Distribution, Ecological Function, and Conservation by following that link.

Pollination ecologists should also note that in this same volume of Annual Review of Ecology, Evolution and Systematics there’s a review by Spencer Barrett and Lawrence Harder called The Ecology of Mating and Its Evolutionary Consequences in Seed Plants.  If you contact those authors I’m sure they’d let you have a copy.

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Filed under Apocynaceae, Bees, Biodiversity, Biogeography, Birds, Butterflies, Climate change, Ecosystem services, Evolution, Honey bees, Hoverflies, IPBES, Macroecology, Mammals, Moths, Mutualism, Neonicotinoids, Pollination, Urban biodiversity, Wasps

Who was the father of biogeography? Let poetry decide! UPDATED

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Over at the Dynamic Ecology blog yesterday, Jeremy Fox posted in the weekly Friday Links feature a piece about clerihews – four line poems about an eminent individual that follows a strict AA BB rhyming structure.  Jeremy’s challenge of “+1000 Internet Points for anyone who writes a clerihew about an ecologist in the comments”, of course, was like a proverbial red rag.  The clerihews came rolling in, including some great contributions, and some dodgy rhymes…  I contributed a couple:

Darwin’s natural selection
Was received with circumspection
But with development of society
Evolution replaced piety

and

Following the theories of Darwin
Science and religion were a-warrin’
But after natural selection
Came more balanced introspection

But then I suddenly found myself in a clerihew face-off  with Brazilian ecologist Rafael Pinheiro, which is too good not to preserve for posterity:

RP:

Look to this poor man called Wallace
He was not born and raised in a palace
But don’t get fooled by this misleading photography
The man is the father of biogeography

JO:

Von Humboldt travelled and mapped plants
When schoolboy Wallace wore short pants
So in a more accurate historiography
Von Humboldt’s the father of biogeography

RP:

Humboldt came first, I will not deny
But Wallace is the father and I’ll tell you why
He was not the first to study species distribution
But the one who explained it through evolution

JO:

Sure, Hooker embraced Darwin’s evolution
And came up with a very modern conclusion
But fatherhood is not about interpretation
It’s about the initial insemination

Jeremy award us 10,000 Internet Points and we agreed to call it a draw 🙂  Thanks to Jeremy for the initial challenge and to Rafael for being such a good sport.  It was a lot of fun.

UPDATE:

Jeremy has also highlighted the contributed clerihews with this post on Dynamic Ecology, to which Rafael has commented:

Jeff Ollerton studied pollinators and plants
When graduateboy me read his papers wearing short pants
So, I must admit, I am happy to be the one
Who faced him in the first clerihew slam

To which there’s only one possible response:

Rafael Pinheiro it’s been my pleasure
To trade these clerihews at leisure
But your last one, truth be told
Makes me feel old

 

 

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Filed under Biogeography

How many trees are there in Amazonia: two recent studies reached very different conclusions – UPDATED

The region of South America that we know as “Amazonia” has arguably the greatest biological diversity of any part of the planet, certainly as far as plants are concerned.  In some places the number of tree species per hectare exceeds 400, an order of magnitude greater than the number for the whole of the British Isles.  However estimating the total number of even the described plant species in this vast area has proven controversial, as two recent studies exemplify.  The first study was by ter Steege et al. (2016) and entitled “The discovery of the Amazonian tree flora with an updated checklist of all known tree taxa“, whilst the second is from just last month: Cardoso et al. (2017) “Amazon plant diversity revealed by a taxonomically verified species list“.  Both of them are open access so click on the links if you want to read the full studies.

One might expect that two such studies focused on Amazonia, both using vouchered herbarium records, would reach broadly similar conclusions as to the number of tree species in the region.  Not a bit of it: ter Steege et al. (2016) report 11,676 species, whilst Cardoso et al. (2017) say that the figure is 6,727.  That’s almost a two-fold difference!  Why the discrepancy?  Inspired by an initial tweet by University of Glasgow taxonomist Roderic Page, I downloaded the data from both studies and looked at it closely.

Here’s a scatter plot of the number of tree species per plant family reported by both studies:

Amazon tree diversity

 

The red line shows where we would expect the data points to lie if both studies had reported the same number of tree species per family.  Clearly few families lie on this line and most are above it as we might expect: as I’ve said, ter Steege et al. (2016) concluded that there were far more tree species overall and this is reflected at the family level.  Note that I’ve graphed this using a log scale and what might seem to be small differences are actually very large indeed.

Although the findings from two studies are highly correlated (diverse families are diverse in both studies, ditto families with low diversity) the actual level of that species richness is very different.  For example, in the Annonaceae, ter Steege et al. report  480 species, Cardoso et al. report 388; in the Clusiaceae the figures are 247 versus 135.  Other families are excluded from one data set or the other: ter Steege et al. reckon there 7 species of trees in the Dilleniaceae whereas Cardoso et al. cite zero.  Here’s a link to the data set if you want to explore further.  

So what’s going on here?  Why do two studies with similar aims, published about 12 months apart, come to such different conclusions.  As far as I can see there are three reasons for this.

First of all, the studies used slightly different taxonomies when it came to considering families and species.  So for example, Cardoso et al. recognise the family Peraceae which ter Steege et al. do not.  Although I haven’t done it, I’m sure that if one were to dig down to the species level there would be differences in which species were accepted and which were considered synonyms.

Secondly, the exact definition of what constitutes a “tree” varies between botanists, and the non-botanists who are no doubt responsible for some of the plant collections: some consider anything to be woody and tall-ish to be a “tree”, others have more strict definitions.  Notes about growth form taken in the field consequently get included in herbarium databases and may be inaccurate, especially for the uncommon species that have rarely been seen in the field.

The final reason, and the one that seems to be responsible for most of the discrepancy, is the definition of what constitutes “Amazonia”.  In the first study ter Steege et al. defined it as including the “forests and savannahs of the Amazon basin and Guiana Shield”.  In contrast Cardoso et al. considered only “lowland Amazon rain forests”.  That’s a big difference as there’s lot of savannah in this region, as well as other habitat types.  When we did field work in Guyana some years ago we could travel very quickly between savannah and rainforest.  It was clear to us that there is a range of trees that are restricted to one habitat or another, including species of Dilleniaceae (mentioned above) that are savannah specialists (hence the family’s exclusion from the Cardoso et al. study).

Now neither of these studies is “wrong” in the sense of being inaccurate or misguided: both are great studies involving a huge effort on the part of the authors.  But the limitations and definitions of geography and taxonomy that I’ve highlighted do mean that they need to be treated as rather different and not directly comparable.

So how many tree species are there in Amazonia?  If we consider just the rainforest then it’s 6,727 (Cardoso et al. 2017).  If we consider all habitats in the region, including rainforest plus savannah etc., then the figure is 11,676 species (ter Steege et al. 2016).  One of the implications of this is that the non-rainforest “Amazonian” habitats collectively contain 4949 tree species.  Thus a large proportion of the diversity of the region is in habitats, such as savannah, which are less of a focus for conservation efforts and not as well known to the general public, but are at least as threatened by agriculture and mining as rainforest.

Thanks to Roderic Page for initially highlighting this on Twitter, and Sandy Knapp for discussion.

UPDATE:  In retrospect my conclusion above regarding the proportion of trees in non-lowland rainforest habitats was much too high, as a couple of commenters have noted below.  It’s worth reading what they have to say, and my responses.  It’s likely that the taxonomic differences between the two studies are at least as great as the geographical ones, but then taxonomic opinions vary hugely.  Just serves to emphasise what a controversial and problematic question this is!

 

 

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Filed under Biodiversity, Biogeography