Monthly Archives: September 2017

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 Buzz Club: citizen scientists protecting pollinators

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This is a guest post by Charlie Dance who is Development Officer at The Buzz Club.


It’s hard to over-stress the importance of pollinators. Not only do they play an indispensable role in global food security, they’re also essential in maintaining the diversity of plant species in natural habitats, thus supporting nature as a whole. The UK is home to thousands of different pollinators including bees, wasps and hoverflies. However, while many of these species seem to be declining or disappearing, we know surprisingly little about the majority of them. Why are some disappearing, and how quickly is it happening? What can we do to help? How can we turn our gardens into pollinator havens? It was to help answer questions like these that the Buzz Club was founded in 2015.

Run by volunteers at the University of Sussex, The Buzz Club is a citizen-science charity using the power of the public to provide important data on pollinators. We run a variety of nationwide surveys and experiments suitable for all ages and ideal for wildlife and gardening enthusiasts. Furthermore, we provide information about how to make our urban landscapes more pollinator friendly.

For more information and for a list of current projects, please visit our website: http://thebuzzclub.uk/

As a membership-based organisation, we rely on the small donation of £2 per month from members, all of which goes directly towards running the charity. Not only do new members receive a complementary welcome pack containing a specially designed seed mix, bee identification chart, pollinator-friendly gardening guide, magnifying lens and stickers (see photo below), they also get to learn more about pollinators whilst helping to generate useful data that can be used in our projects.

We believe that with your help we can find out how best to conserve bees and other pollinators. Our ultimate goal is to ensure that we look after insects, giving them and us a future.

Join the Buzz Club here: https://alumni.sussex.ac.uk/buzzclub

Facebook: http://www.facebook.com/TheBuzzClubUK

Twitter: http://www.twitter.com/The_Buzz_Club


From Jeff:  if citizen science is your thing, don’t forget that the Ivy Pollinators project will run again this year: https://jeffollerton.wordpress.com/2016/10/11/ivy-pollinators-citizen-science-project/

 

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Fly pollination in the trap flower genus Ceropegia: a new study just published

Flies on C. arabica from Sage Reynolds

Pollination of flowers by flies (the insect order Diptera) has long fascinated me, in part because it often subverts the idea of what “normal” flowers should look like, but also because it is much less well studied, and appreciated, compared to bee pollination.  This is despite the fact that fly pollinated flowers are at least as frequent as bee pollinated flowers in many plant communities, as I show in a forthcoming review in the journal  Annual Review of Ecology, Evolution and Systematics (more on that in November when it’s published).

Over the past decade I’ve been studying the large plant genus Ceropegia in the family Apocynaceae (subfamily Asclepiadoideae).  The flowers of these species temporarily trap their fly pollinators, releasing them after a period, during which pollination takes place and/or pollen is picked up.  The latest study from this work has just been published in the journal Flora, in collaboration with colleagues from eight different countries.  The title is:

Diversity of Diptera families that pollinate Ceropegia (Apocynaceae) trap flowers: an update in light of new data and phylogenetic analyses

If you follow that link you can download the PDF for free for the next 50 days.

One of the main findings from this new study is that the diversity of fly families that pollinate Ceropegia spp. is much greater than we had previously realised.  The total now stands at 16 different families, including some that rarely, if ever, pollinate other plants (as far as we yet know).

Another important finding is that this clade, which may contain as many as 1000 species in total, seems to have diversified despite that fact that all species are apparently fly pollinated.  This is unusual: diversification of plant clades often involves shifts to very different groups of pollinators, e.g. bee to bird or bat pollination.

There’s still lots to discover about this group of plants and this is just the latest output from what is an ongoing project focused on Ceropegia and the Apocynaceae more generally.

Here’s the abstract:

“Pollination by flies (Diptera) has been important to the diversification and ecology of the flowering plants, but is poorly understood in contrast to pollination by other groups such as bees, butterflies and birds. Within the Apocynaceae the genera Ceropegia and Riocreuxia temporarily trap flies, releasing them after a fixed, species-specific period of time, during which pollination and/or pollen removal occurs. This “trap flower” pollination system shows convergent evolution with unrelated species in other families and fascinated Stefan Vogel for much of his career, leading to ground-breaking work on floral function in Ceropegia (Apocynaceae). In this new study we extend the work of the latest broad analysis published by some of the authors (Ollerton et al., 2009 − Annals of Botany). This incorporates previously unpublished data from India and Africa, as well as recently published information, on the diversity of pollinators exploited by Ceropegia. The analyses are based on a more accurate phylogenetic understanding of the relationships between the major groups, and significantly widens the biogeographic scope of our understanding of fly pollination within Ceropegia. Information about the pollinators of 69 taxa (species, subspecies and natural varieties) of Ceropegia is now available. Twenty five families of Diptera are known to visit the flowers of Ceropegia, of which sixteen are confirmed as pollinators. Most taxa are pollinated by species from a single family. Overall, there were no major biogeographic differences in the types of Diptera that were used in particular regions, though some subtle differences were apparent. Likewise there were no differences between the two major clades of Ceropegia, but clear differences when comparing the range of Diptera exploited by Ceropegia with that of the stapeliads. This clade, one of the largest in the Asclepiadoideae, is a fascinating example of a species radiation driven by an apparently relatively uniform set of pollinators.”

Photo credit: flies on flowers of Ceropegia arabica in cultivation by Sage Reynolds.

 

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