Category Archives: Biogeography

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

Wageningen poster.jpg

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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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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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

Norway from the air.jpeg

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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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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The macroecology of animal versus wind pollination – a new study just published

In collaboration with colleagues in Brazil, Denmark, and elsewhere in the UK, we’ve just published a new research paper which looks at the global spatial distribution of wind and animal pollinated plant species, and the underlying historical and contemporary ecological causes of that distribution.  It’s a study that builds on my “How many flowering plants are animal pollinated?” paper in Oikos, and has been a long time in its gestation.  We’re very excited by its findings and plan to develop this project in the future.

As a bonus we made the cover of the journal with the amazing image below!  Big thanks to Pedro Viana and Jesper Sonne for the photos.

Here’s the citation with a link to the publisher’s website; the abstract is below.  If anyone wants a PDF copy, please ask.

Rech AR, Dalsgaard B, Sandel B, Sonne J, Svenning J-C, Holmes N & Ollerton J (2016) The macroecology of animal versus wind pollination: ecological factors are more important than historical climate stability. Plant Ecology & Diversity 9: 253-262

 

Abstract:

Background: The relative frequency of wind- and animal-pollinated plants are non-randomly distributed across the globe and numerous hypotheses have been raised for the greater occurrence of wind pollination in some habitats and towards higher latitudes. To date, however, there has been no comprehensive global investigation of these hypotheses.

Aims: Investigating a range of hypotheses for the role of biotic and abiotic factors as determinants of the global variation in animal vs. wind pollination.

Methods: We analysed 67 plant communities ranging from 70º north to 34º south. For these we determined habitat type, species richness, insularity, topographic heterogeneity, current climate and late-Quaternary climate change. The predictive effects of these factors on the proportion of wind- and animal-pollinated plants were tested using correlations, ordinary least squares (OLS) and logistic regression analyses with information-theoretic model selection.

Results: The proportion of animal-pollinated plant species was positively associated with plant species richness and current temperature. Furthermore, in forest, animal pollination was positively related to precipitation. Historical climate was only weakly and idiosyncratically correlated with animal pollination.

Conclusion: Results were consistent with the hypothesised reduced chance for wind-transported pollen reaching conspecific flowers in species-rich communities, fewer constraints on nectar production in warm and wet habitats, and reduced relative effectiveness of wind dispersal in humid areas. There was little evidence of a legacy of historical climate change affecting these patterns.

andre-capa-1

 

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Filed under Biodiversity, Biogeography, Brazil, Climate change, Macroecology, Pollination

The integration of alien plants in mutualistic plant–hummingbird networks – a new study by Maruyama et al. (2016)

The collaborations with researchers in Brazil and Denmark in which I’ve been involved in recent years, focused particularly on hummingbirds and networks of plant pollinator interactions, have been very productive, most recently seen in a study of the effects of hummingbird feeders on diversity and abundance of the birds.

This collaboration continues with a new study that has just been published in the journal Diversity and Distributions which deals with the way in which non-native plant species are exploited by assemblages of hummingbirds in the New World.  Here’s the abstract:

 

Aim:  To investigate the role of alien plants in mutualistic plant–hummingbird networks, assessing the importance of species traits, floral abundance and insularity on alien plant integration.

Location: Mainland and insular Americas.

Methods: We used species-level network indices to assess the role of alien plants in 21 quantitative plant–hummingbird networks where alien plants occur. We then evaluated whether plant traits, including previous adaptations to bird pollination, and insularity predict these network roles. Additionally, for a subset of networks for which floral abundance data were available, we tested whether this relates to network roles. Finally, we tested the association between hummingbird traits and the probability of interaction with alien plants across the networks.

Results: Within the 21 networks, we identified 32 alien plant species and 352 native plant species. On average, alien plant species attracted more hummingbird species (i.e. aliens had a higher degree) and had a higher proportion of interactions across their hummingbird visitors than native plants (i.e. aliens had a higher species strength). At the same time, an average alien plant was visited more exclusively by certain hummingbird species (i.e. had a higher level of complementary specialization). Large alien plants and those occurring on islands had more evenly distributed interactions, thereby acting as connectors. Other evaluated plant traits and floral abundance were unimportant predictors of network roles. Short-billed hummingbirds had higher probability of including alien plants in their interactions than long-billed species.

Main conclusions: Once incorporated into plant-hummingbird networks, alien plants appear strongly integrated and, thus, may have a large influence on network dynamics. Plant traits and floral abundance were generally poor predictors of how well alien species are integrated. Short-billed hummingbirds, often characterized as functionally generalized pollinators, facilitate the integration of alien plants. Our results show that plant–hummingbird networks are open for invasion.

 

The full reference is: Maruyama, P.K. et al. (2016) The integration of alien plants in mutualistic plant–hummingbird networks across the Americas: the importance of species traits and insularity.  Diversity and Distributions (in press).

Happy to send a PDF to anyone who would like one.

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Tropical Zombies: Moles & Ollerton (2016) is now published

P1080615Back in March 2014 I reported about a guest blog that Angela Moles (University of New South Wales) and I had written for the Dynamic Ecology blog entitled “Are species interactions stronger and more specialized in the tropics?”  The post generated a lot of comments, not all of them supportive of what we were saying.  It also resulted in an invitation from the editor of the journal Biotropica to write up the post as a commentary.  This we did and duly submitted, it went through a couple of rounds of peer review, and has now finally been published.

The paper is currently open access on the Biotropica website as an early view item; here’s the reference hyperlinked to it:

Moles, A. & Ollerton, J. (2016) Is the notion that species interactions are stronger and more specialized in the tropics a zombie idea? Biotropica DOI: 10.1111/btp.12281 

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Sex and drugs and the source of the Nile: Sir Richard Francis Burton

Burton photo

They say that things often come in threes, and so it has appeared recently in relation to an individual I have long admired and been fascinated by: Captain Sir Richard Francis Burton.  As far as I’m aware there is no significant Burton-related anniversary in 2015 (other than it being 125 years since his death), but nevertheless he’s popped up in a couple of places of late.  First of all there was a Radio 4 Great Lives programme about the man; then yesterday there was an article in Nature by Professor Clare Pettitt and on Wednesday night, at a WIldlife Trust event in Cambridge, I found myself chatting to a man whose name badge stated “Richard Burton”.

Clearly the universe was trying to tell me something and it reminded me of a piece of writing that I produced in October 1990 (!) to mark the 100th anniversary of Burton’s death, and never published.  To put this in context, I was 25, about a year into my PhD research, and anticipating the birth of my first child in December.  Re-reading the piece has been less painful than I thought it would have been. Some of the writing is a little clumsy and there are other aspects that I’d now focus on, but it’s not too bad.  Having said that, Karin said it sent her to sleep and that my writing has improved a lot in 25 years, so there’s no pleasing everyone!

Anyway I thought I’d post this piece of writing (very lightly edited) as an indulgent missive from my 25 year old to my 50 year old self.  And it’s dedicated to my daughter Ellen in her 25th year.

———————————————————-

Often, simply striving for fame is not enough. No matter how daring your exploits or how much you publish, the contingencies of history conspire to obscure you, consigning your life and works to the realms of the scholar or to that nebulous coterie, the “enthusiasts”. Such has been the fate of one of the most exciting of the many outstanding lives of the Victorian age.

This week marks the one hundredth anniversary of the death of one of our most important, yet underappreciated, scholar-travelers, Captain Sir Richard Francis Burton. Such anniversaries always seem to necessitate a reassessment of the celebratee’s life and work, and this one is no exception; two major new biographies, an extensive “biobibliography” chronicling Burton’s literary output, and a film “Mountains of the Moon“. Yet for all this, Burton is still not a widely known figure; although his adventures far surpass, in daring and in accomplishment, his contemporaries Livingstone and Stanley, still he does not enjoy their household-name status. This is in spite of, between 1890 and 1989, the publication of at least eight biographies, a bibliography, and many articles and essays devoted to the man’s exploits. Add to this Burton’s own vast literary output, none of which is noted for any bashful self-deprecation on the part of the author, and one begins to wonder at the criteria we use to apportion recognition.

Richard Francis Burton was born in 1821, the son of an army officer, Colonel Joseph Burton. His early life was spent travelling Europe with his family, fueled by the incessant wanderlust of his father. This gypsy start to life, as well as being an obviously formative prelude to his later travels, seemed to encourage the rowdier, untamed, hell-raising aspects in the characters of Richard and his brother Edward. The despair of their parents, the pair were soon packed off to college in England; Edward to Cambridge, Richard to Oxford. College and academic life did not suit either of the boys, and both left (in Richard’s case, forcibly; he was sent down after attending a proscribed horse race) to pursue military careers. Over the next 50 years, Richard Burton devoted himself to restlessly wandering the world, roaming Africa and Asia, North and South America, and Europe. He was one of the first Europeans to visit the Islamic sacred cities of Medina and Mecca; he explored India, often on covert missions for the British government; travelled in Africa where he searched for the source of the Nile (and only missed discovering it through ill-luck and the machinations of others); he lived for a time in South America as consul at the port of Santos in Brazil and observed first-hand the war between Paraguay and the allied forces of Argentina, Brazil and Uruguay; and in all travelled and observed enough to satisfy several lifetimes.

During his wanderings Burton saw and experienced much, events which invariably he became curious about, investigating further, writing down his views. Whether it was local uses of medicinal and psychoactive plants; details of tribal ritual, or the niceties of local sexual practice; the grammatical fine points of local dialects; geological formations; curiosities of natural history; or simply the price of staple vegetables in a native market, Burton was interested. These details inevitably found their way into his many books, articles and learned papers, packing paragraphs of ethnological, geographical, archaeological and natural history minutiae into his accounts of travels and expeditions.

It is Burton’s polymath approach to scholarly work that is the man’s most interesting feature. Perhaps it was his limited formal education (travelling tutors, two terms at Oxford) that fostered this approach. Though in many ways laudable, conventional academia can lead to a blinkered approach to research, ivory-towerism at its worst. If Burton had limited himself to purely single-strand studies, for example oriental languages (or even language), as may have resulted from following an academic career, the world, and Richard Francis Burton, would have been far poorer. Had he only recorded the bare geographical necessities required of, for example, his travels in the Great Lakes region of Africa, what dry accounts they would have been, and what details we would have lost.

This is not to say that Burton’s work was not scholarly, far from it. His translation of the Arabian Nights, although not the first, is certainly the definitive version, rich in anecdotal footnotes from a man for whom the deserts of Arabia were perhaps his first real home (as a child and a young man he had hated Britain, especially its climate), and his research and translation of the works of the Portuguese poet-explorer Camoes shows Burton at his most academic.

Burton has perhaps been more misunderstood, loathed and ignored than any of his contemporaries, or any comparable figure before or since. This is in part due to the man’s interests during his lifetime: translations of obscure erotica such as the Kama Sutra; a more than passing interest in the ins-and-outs of male and female circumcision; undercover reconnaissance of Indian homosexual brothels (which invariably led to rumours about Burton himself) all added to his infamy. Perhaps more than anything else, this meant that most of Burton’s books were not widely read, a trend which continues today, aided by the inflated prices demanded by booksellers for even the most popular of his works.

As if our view of Burton were not obscured enough, his over-zealous wife Isabel sought to soften history’s account of her “Jemmy” by burning almost all of his private papers after his death; writings which may have cast light on this enigmatic man were consigned wholesale to the grate. Because of this, Burton’s biographers have tended to be hard on Isabel, dwelling on her attempts to instill Catholicism into her part Muslim, part Atheist husband, and, of course, on her literary pyromania. This may be because of frustration on their part; biographers and commentators have never really been able to reason Burton out, and large parts of his life remain veiled in secrecy and obfuscation. The task has not been aided by Isabel’s actions. Yet she was devoted to Burton, who was never the easiest of men to get along with, being often bad tempered or absent for months on end.

But Isabel is only a scapegoat. Mainly, the problem is that there never has been any other person to compare with Burton. How could any man hope to fulfill all that he aspired to? Why the incessant wandering in search of new experiences? Why was it that the man did not focus his energies, rather spreading himself across a continent of interest, his curiosity endless? It has been said that if Richard Burton had concentrated his mind in this way, he could have been one of the foremost intellects of his time, rivaling Darwin or Huxley, Edison or Swan. Yet this misinterprets the man. It is doubtful whether Burton could have disciplined himself enough to centre on a single area of research; Burton was a searcher, a shifter of interests. Burton’s writings have been criticised as being unstructured, cluttered and self-indulgent, almost as if he had not the time nor inclination to properly revise and edit, but simply wanted to get the current project out of the way in order to get on with the next. This is borne out by the fact that, towards the end of his life, he had eleven desks set up in the study of his home in Trieste, where he was consul; each desk was for a different project and, when he tired of one, he would move to another, as if restlessly seeking for something.

But none of this need be considered as faults in Burton’s character; he was probably no more flawed, neurotic or self-obsessed than any of the great men of his time. It seems impossible than an intellect as deep and all-encompassing as his, which mastered some twenty nine languages, produced fifty books (many of them comprising more than one volume) plus innumerable essays and articles plus all the work that Isabel burned, could ever hope to be completely stable and well adjusted. Eccentricities of writing and behaviour seem inevitable.

Now, one hundred years after his death, is as good a time as any to properly reappraise the life of Richard Francis Burton. As an explorer, anthropologist, geographer, linguist, orientalist, translator, diplomat, swordsman, writer (and a lot more besides) he stands unrivalled by the broad sweep of his experience and knowledge. Yet his private life seems consistently to get in the way of any objective assessment of the man and his accomplishments.

The scandalous view of Burton, prevalent in England during his life and long after his death, was as a man obsessed by sex, a delver into the sordid details of native life and custom, promoter (though not practitioner, Isabel would never have allowed it) of polygamy, an unpopular critic of certain governmental interventions abroad, a user of cannabis, opium and other, more exotic drugs, and an ill-tempered, frequently drunk, godless, misogynist racist. His reputation as a fighter, even a murderer, was often played up by Burton, though there has is only one well documented account of him ever killing anyone, and then in self defence. Yet the real Burton, so far as we can tell, does not deserve this misrepresentation. His interest in all things erotic was partly academic and partly out of concern for the then current view that women should not find pleasure in sex. Are these the motives of an over-sexed misogynist?   His use of drugs, including alcohol, is well known, but was not unusual amongst Victorians exposed to the influences of the Far East, or for whom port, wine and whiskey were often viewed as medicinal necessities. Finally, Burton was no more racist than most Europeans of his time, yet it was from an intellectual stand point, not an emotional or cultural one. Most of the great academics of the period believed that there was a progression of human development, with white Europeans at the pinnacle. But no one who deeply despised Arabs or Indians could live and worship amongst them the way Burton did. He may have severely criticised them, but then he criticised everyone.

All of this points to a man more liberal than many people have believed; Burton was in many ways a free thinker, particularly given his upper middle class military background. Finally, there is the matter of his atheism, if such it was, which would today raise few eyebrows. Yet the man lived and prayed for much of his life as a Muslim, had been initiated into an esoteric Sufi brotherhood, and before that into a Hindu sect. This is not the life of a godless man, in the accepted sense of the word, but of a man searching for truth, who was too intelligent to believe he had ever found it in the rosary beads of his wife’s Catholicism or in the calling chant of a muezzin.

The life of Richard Francis Burton was dogged by ill-luck and, certainly towards its end, ill health, and furthermore seemed cursed by the intransigence of government officials and individuals with grudges. A character such as his finds no difficulties in making enemies, yet they always seemed to be foes with influence, willing to block his attempts at organising expeditions or soliciting official help for schemes to further the British Empire, or its servant Burton. He never did find the source of the Nile; this single act, more than any other, would have ensured his position as the greatest of the Victorian explorers. Yet had he been successful, would the constant round of lecture tours, press interviews, official visits, and all have given him time to think and write about anything else? I believe it would, though whether it would have satisfied his roving curiosity and incessant wanderlust seems unlikely.

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