Fossil record supports evidence of impending mass extinction

The research team based at the University of York has, for the first time, discovered a close association between Earth climate and extinctions in a study that has examined the relationship over the past 520 million years — almost the entire fossil record available.

Matching data sets of marine and terrestrial diversity against temperature estimates, evidence shows that global biodiversity is relatively low during warm ‘greenhouse’ phases and extinctions relatively high, while the reverse is true in cooler ‘icehouse’ phases.

Moreover, future predicted temperatures are within the range of the warmest greenhouse phases that are associated with mass extinction events identified in the fossil record.

The research, published in the latest issue of Proceedings of the Royal Society B., was carried out by University of York student Gareth Jenkins, together with his supervisor, Dr Peter Mayhew, and University of Leeds Professor Tim Benton, both of whom are population ecologists.

Dr Mayhew says: "Our results provide the first clear evidence that global climate may explain substantial variation in the fossil record in a simple and consistent manner. If our results hold for current warming — the magnitude of which is comparable with the long-term fluctuations in Earth climate — they suggest that extinctions will increase."

Of the five mass extinction events, four — including the one that eliminated the dinosaurs 65 million years ago — are associated with greenhouse phases. The largest mass extinction event of all, the end-Permian, occurred during one of the warmest ever climatic phases and saw the estimated extinction of 95 per cent of animal and plant species.

"The long-term association has not been seen before, as previous studies have largely been confined to relatively short geological periods, limited geographical extents and few groups of organisms," says Professor Benton. "But the evidence is striking."


Article: Mayhew, P.J., Jenkins, G.B. & Benton, T.G. (2008) A long-term association between global temperature and biodiversity, origination and extinction in the fossil record. Proc. R. Soc. B 275, 47–53

Source of article: Univeristy of Your's Press Release

Volunteer to help IBS

Dear Biogeographers:

The International Biogeography Society has three goals:

1. Foster communication and collaboration among biogeographers in disparate academic fields
2. Promote the training and education of biogeographers
3. Increase awareness and interest in the contributions of biogeographers

Are you interested in biogeography and would you like to help the society accomplish its goals? To do this you should do three things.

First, if you are not a member, joint IBS - Membership is a bargain at $40.00 per year ($30.00 per year for students). In addition to supporting the society, becoming a member of IBS provides several personal advantages: 1) the opportunity to participate in international meetings of the Society, 2) discounts (up to 20%) on books published in association with IBS, 3) free online access to four of Blackwell Publishing's journals: Journal of Biogeography, Global Ecology & Biogeography, Diversity & Distributions, and Ecography, all journals covering aspects of biogeography and ecology, and 4) a 20% discount on access to two additional Blackwell Publishing journals: Journal of Avian Biology and Oikos.

Second, check out the website (http://www.biogeography.org/) and see what you might like to be involved with. Possibly helping with meetings, hosting an off-year meeting, serving on a committee, judging posters, recruiting new members, or even nominating someone for the Alfred Russel Wallace Award.

Finally, contact me or any member of the board and tell us what you are interested in doing for IBS or we can perhaps suggest something. We can always use someone to help stuff flyers in meeting packets!

Cheers, V. A. Funk, President IBS

Are current projections of climate change-impacts on biodiversity misleading?

A recent study, published in Ecography, has profound implications for the future study of diversity on Earth. The article challenges the current view that patterns of current climate are sufficient to explain and predict diversity, and proposes that historical climate patterns are also of strong importance to diversity prediction.

This is the urgent question arising from the study “Quaternary climate changes explain diversity among reptiles and amphibians”, published in the journal Ecography.

Why is life on Earth not evenly distributed? Geographic patterns of species diversity and their underlying processes have intrigued scientists for centuries, and continue to spur scientific debate. Studies carried out over the past 20 years have led to the conclusion that species diversity is best predicted by contemporary patterns of energy and water, the so-called “contemporary climate” hypothesis. Because current climate gradients are correlated with past climate variability, it has also been suggested that current climate acts as a surrogate for evolutionary processes that have been triggered by past climate variability, giving rise to the “historic climate” hypothesis. Now, new high-resolution data on historic climate has allowed Dr Araújo in collaboration with Dr Rahbek and other colleagues to finally directly test the “historic climate” versus “contemporary climate” hypotheses of biological diversity. Their illuminating results are published in a recent paper in Ecography. Contrary to the expectations of many scientists they found that historic climate variability was a better predictor of reptilian and amphibian diversity in Europe than contemporary climate.

“The lack of quantitative spatial data on variation in climate over historical time has prevented more rigorous testing of these diverging hypotheses”, says Dr. Miguel B. Araújo from the National Museum of Natural Sciences (CSIC) in Madrid. As a consequence, “the debate on the causes of diversity gradients has turned to some degree into a discussion of semantics”.
Recent developments in general climate models have finally facilitated high resolution predictions of past climates. In collaboration with leading climatologists working on paleoclimate modeling in the United Kingdom, Drs. Araújo, Rahbek and colleagues provide the first comparative test capable of differentiating between the contribution of contemporary and historical climate drivers of diversity gradients across a complete lineage of species at a continental scale.

“In recent years, analytical attempts to shed light on the role of history in determining today’s patterns of species richness have focused on the strong residual variation of models using contemporary climate”, explains Dr. Carsten Rahbek from the Center of Macroecology at the University of Copenhagen. “It has been argued that these residuals provide information about the role of historical rather than contemporary constraints. However, such an analytical approach assumes that contemporary climate is the main explanatory force. In other words, the contemporary and historical hypotheses are not tested simultaneously in a directly comparable manner, and historical hypotheses are only invoked to explain what is left to elucidate after the implementation of contemporary environmental processes”, says Dr. Rahbek.

“Our results are striking in that they contradict previous studies of large-scale patterns of species richness” affirms Dr. Rahbek. “They provide the first evidence, using a quantitative analytical approach, that historic climate can contribute to current patterns of richness independently of, and at least as much as contemporary climate”. This study has profound implications for the study of diversity on Earth, and challenges the current view that patterns of contemporary climate are sufficient to explain and predict diversity.

Differentiating between contemporary and historical hypotheses is important, not only for theoretical reasons: “an understanding of the mechanisms that generate and maintain diversity provides valuable insights for predicting the impacts of contemporary climate changes on biodiversity”, says Dr. Araújo. “If contemporary climate does drive species richness, then current climate variables could be used to accurately predict the effects of climate change on biodiversity. If, as shown in our study, the mechanisms underlying contemporary patterns of species richness are in fact strongly influenced by the history of climate, then current-climate predictions may be seriously misleading and alternative approaches to predict the effects of climate change on biodiversity must be developed”.

Article:
Miguel B. Araújo, David Nogués-Bravo, José Alexandre F. Diniz-Filho, Alan M. Haywood, Paul J. Valdes, Carsten Rahbek Quaternary climate changes explain diversity among reptiles and amphibians Ecography. doi:10.1111/j.2007.0906-7590.05318.x

Source of article: Blackwell's press release

Editor's note: This is the first article of the new section of the IBS Newsletter and blog: "highlights in the literature". Highlights in the literature publishes press releases on exciting papers in all fields of biogeography. With this new section the IBS seeks to encourage popular writing of scientific papers in biogeography and help promoting biogeographical research outside the academic realm. If you have published, or are about to publish, a paper in biogeography and have prepared a press release in a widely accessible language, we are willing to consider its publication in the IBS Newsletter and Blog. Press releases of biogeographical paper must be sent to ibs@mncn.csic.es for consideration by members of the editorial board.

Christen Raunkiær – one of several early island biogeographers

Henning Adsersen
Center for Macroecology, Institute of Biology, University of Copenhagen
e-mail: adser(at)bi.ku.dk


Islands, especially isolated oceanic islands, have always held a fascination for me, both emotionally and intellectually. Likewise they have fascinated travelers ever since the first oversea transportation means were invented and employed. This fascination and fantasy created early a fertile substrate for tales, epics and accounts on island life, as evidenced by Homer’s Odyssey, Sheherazade’ tales on Sindbad, and the Atlantis myths. The romantic novel was nourished in the same medium, as evidenced by Gulliver, Robinson Crusoe, and The Treasure Island.

In island literature exaggerations are almost mandatory. The scientific literature on islands, of course, does not allow exaggerations but there is still a marked tendency to focus on island organisms or phenomena that are peculiar in one way or another. No island birds are more commonly known as the monstrous Mauritius dodo and hardly any island plants as admired as the Seychelles giant coco or the silver sword of Hawaii. This trend to concentrate on the odd island organisms (the dodo approach, Adsersen 1995) is of course fascinating but it does not tell the whole truth on island biology. Neither does the focus on peculiar insular processes like evolutionary radiation as shown by Darwin finches or Hawaiian honeycreepers. No matter how seminal this approach (the finch approach) has proven to be for ecology and evolutionary biology it elucidates only part of the island scenario. The good stories on dodos and finches have the mark of anecdotes as long as they are not compared to something else. This “something” could be the rest of the organisms in the island, or the organisms of comparable areas on continents, or a general global pattern. In other words, it is necessary to consider the island biota as entities that may be quantitatively assessed and characterized.

MacArthur and Wilson’s classical works constitute a paradigmatic breakthrough of this approach, and by their work island biogeography was established as a discipline (MacArthur and Wilson 1967). Their equilibrium theory of island biogeography (ETIB) has been cyclically admired, criticized, rejected, reformulated, and modified ever since their modest-looking book was published, and this discourse will probably go on. But everyone participating in this debate acknowledges that island biotas may and should be assessed quantitatively and that island biotas are dynamical entities subject to qualitative and quantitative changes in time and space. Maybe this is the kernel of their message, and maybe its simplicity is the reason why it has had such a profound impact on evolutionary science, biogeography, and (macro)ecology. Now when the first forty years of island biogeography has been celebrated by a magnificent symposium at Harvard it seems apposite to ask whether Mac Arthur and Wilson had any predecessors in this quantitative approach. Marc Lomolino in his contribution to the symposium did so, focusing mainly on the American arena. Here I wish to draw attention to some important European contributors to early island biogeography.

Right after the publication of The Theory of Island Biogeography there was a marked tendency to try to support or even prove the theory by demonstrating mathematical relationships between species richness S and area A (species-area relationships, SPAR). Nice SPARs like the Arrhenius equation (log S = C + z logA) or the Gleason equation (S = K + p logA) (Gleason 1922) may be found by the quantitative approach and they may be result of ETIB; but ETIB could be explained by several other mechanisms, so there is no direct link between SPAR and ETIB. SPARs are important (“Ecology’s most general pattern”, Lomolino 2000) but not limited to island biogeography. Even Arrhenius’ pioneering work (Arrhenius 1921) was not an island biogeography work: it was indeed done on islands in the Stockholm archipelago but his S and A assessment were done in nested plots within the islands, so he could only obtain monotonously increasing functions, and there is no reference to total island S or A. Maybe Darlington’s rule of thumb (tenfold island area yields double species number, or S = c Alog 2 = c A0.301) is the first SPAR employed on true island entities (Darlington 1957). Numerous reviews and textbooks enlighten the history and relations between SPAR and ETIB.

SPARs are based on numbers of species S in limited, defined areas. Islands normally have hard borders and well-defined area, so that S may be assessed rather precisely.
It is therefore not surprising that some of the first quantitative data in biogeography are number of species on islands. J.D.Hooker studied Darwin’s plant collections from Galápagos and published in 1847 a remarkable treaty: On the Vegetation of the Galapagos Archipelago, as compared with that of other Tropical Islands and of the Continent of America. He set up three main questions to answer:

• How are the species distributed into major taxonomical groups?
• How many species are “peculiar” (endemic) and where do the species with wider distributions occur elsewhere?
• How are the species distributed within the archipelago?

Hooker not only tried to answer the questions but also managed to compare the patterns to that of other tropical archipelagos and to continental situations. His approach to question 3 is especially interesting. It is a numerical analysis of the number of plant species (the Florulæ) of the 4 islands that Darwin visited. In this analysis he repeatedly related the number of “peculiar” species to the total flora, and he even presented the data in a clear tabular form.

Table I. Original Hooker’s data on the florulæ of the Galápagos Islands.

His material was not complete, of course, even if he apparently managed to get access to all herbarium material in England and France. His estimate of total species number was 265, of which 17 were considered introduced by man. Today we assume that S ~ 600 and the number of naturalized plant species about the same. In the table beneath the modern figures (based on Lawesson et al 1987) are inserted. None of Hooker’s conclusions will hold but considering that only four of the major islands were included some of the relative figures (e.g. percent of plants confined to Galápagos) are quite accurate.

Table II. Hooker’s data on the florulæ of the Galápagos Islands. Recent data inserted. In the column “Absolutely peculiar to the islet.” the inserted column to the right left refer to the four islands mentioned, the one to the left refer to the entire archipelago.

Hooker’s approach is quite modern and for his time extraordinarily quantitative, although we today to each question would add “and why?” He maintained his interest in islands throughout his career, maybe best expressed in his lecture to the British Association in 1866 (reprinted and commented in Williamson 1984). In this lecture, which admittedly is much more verbal than numerical, he builds up both anecdotal and quantitative evidence that the pattern we may observe in island biogeography can be explained only if we assume evolution (“derivative origin of species”), and that the derivate origin of species became evident ”when Zoology and Botany became the subjects of exact scientific studies”! So certainly Hooker should be regarded as one of the progenitors of quantitative island biogeography.

Half a century after Hooker’s lecture a Finnish botanist, Alvar Palmgreen, meticulously censused the flora of islands in the Åland Archipelago of the Gulf of Bothnia, and remarkable papers followed: “The species richness as a plant geographical parameter” – “Chance as a plant geographical parameter” – “Remoteness as a plant geographical parameter”. The papers were in German and Swedish – they were read at the time but also forgotten. Haila and Järvinen, however, call to the attention that Palmgreen not only studied species-area relationships; species isolation relationships, and stochasticity in species dispersion but also came very close to formulation of ETIB (Haila and Järvinen 1982). It is, however, difficult to read Palmgren’s papers, even if you understand the language. His style is very wordy and even if he compares quantities like species number or isolation he does it verbally. There is hardly one graph or one mathematical formulation in his entire opus. Nevertheless, as Haila and Järvinen expressed it, his thoughts antedate MacArthur’s and Wilson’s by half a century.

Christen Raunkiaer in 1902

Christen Raunkiær from Denmark was contemporary with Palmgreen, but unlike Palmgreen he was not forgotten. His life forms, frequency analyses, and contemplations on species abundance distributions are still cited in most modern text books on ecology and biogeography. His life form system was outlined in Danish in 1904 (Raunkiær 1904). Next step was to introduce the system and its application in biogeography, which he did in French in a preliminary form in 1906. The final version came in 1907 in his most renowned publication Planterigets Livsformer og deres Betydning for Geografien. (The Life-Forms of plants and their bearing on geography) (Raunkiær 1907). The system was immediately seized by Scandinavian and continental botanists and ecologists. It became almost as paradigmatic in ecology as ETIB and it is quite apposite to celebrate its first century together with ETIBs 40 years. His works were after a long delay translated to English with the title: “The Plant Life Forms and Statistical Plant Geography” (Raunkiær 1934) and world wide acknowledged.

There are probably two reasons why Raunkiær’s life form system attained its high regard: The one is that the life forms are based on clear logical and biological reasoning and any plant species can without to much difficulty be categorized even by non-experts – and the second is his strict quantitative approach. With his own words: “-then Plant Geography as a botanical science gives place to Plant Geography as a geographical science” His combination of clear-cut concepts and exact biogeographical analyses on large scales made it possible for him to formulate precise definitions of biomes and to compare plant communities across biomes.The same combination is one of the reasons for MacArthur and Wilson’s success.

Even though the word “statistical” recurs in several of Raunkiær’s titles, he had no great knowledge on statistics as practiced today. Parametric distributions, hypothesis testing and significance were concepts unknown to him, as they where to Hooker, Palmgreen and Arrhenius. First after Fischer’s paradigmatic contributions to statistics these concepts became applicable to biogeographers and soon important contributions emerged. Within island biogeography Preston’s monumental papers on Commonness and Rarity of Species (Preston 1948, Preston 1962) seem to be best known but the equally monumental monograph Patterns in the Balance of Nature by C.B. Williams (1964) (a coworker of Fischer’s) deserve as much recognition. Both Preston and Williams took their approach from abundance distributions (lognormal and logseries, respectively) in communities and ended up by reflecting on SPARs – very much fertilizing the ground for ETIB.

Figure 1. The Raunkiaerian J

Abundance distribution was another of Raunkiær´s interests. He invented the frequency analysis in vegetation ecology and observed the recurring pattern known as the Raunkiærian J (Figure 1). He even postulated that mature homogeneous vegetation could be recognized by this pattern. Such a daring postulate would of course raise discussion. McIntosh (1962) reviewed the discussion and concluded that the Raunkiærian J is one of those “ideas that seems to be invulnerable to attack and persist although subjected to multiple executions” (another parallel to ETIB?). Raunkiær’s observation was a seminal inspiration for Preston (1948) and he demonstrated that the Raunkiaerian J may be derived from the lognormal abundance distribution, (which on its side will lead to the Arrhenius equation). Later on Williams (1964) showed that it may be derived also from the logseries distribution (which lead to Gleason equation). Even very recently the Raunkiærian J attains attention, now under the more general term “hollow species abundance distributions” (McGill et al 2007).

It is less known that one of Raunkiær’s last publications was on island biogeography: The Life-Form Spectrum of Some Atlantic Islands. The reason why it is almost forgotten is that it appeared in the series Botaniske Studier (published by himself) that comprises several of his emeritus works. It is an 80 page monograph in which he compares the life-form spectra of most Atlantic archipelagos from Svalbard in the north to South Georgia in the south. His questions were almost the same as Hooker’s first two questions above, except that, instead of taxonomical supraspecific groups, he considered ecological ones:

1. How are the species distributed into life-form groups?
2. How many species are “peculiar” (endemic) and does their life-form spectrums differ from that of species with wider distributions occur elsewhere?

He also added a “why” to his question – and found that his climatically derived life form biome types give good explanations. His discussion on this is lengthy because he had not the necessary tools to make up the relevant exact evaluations but he would probably have been pleased to see the ordination diagram in figure 2. It is based on his data, and however incomplete they may have been they give very good support to his ideas.

Figure 2. Nonmetric multidimensional scaling of Raunkiær’s data on life form spectra from Atlantic islands. The ordination was run in PC-ORD 5 with Euclidian Distance as dissimilarity measure. The diagram is highly significant (P<0.01,>

Raunkiær was not the first to use quantitative data in biogeographical analyses. Early biogeographers like Schouw, De Candolle, Hooker and Warming used quantitative expressions to support their idea. The novelty in his approach was that he demanded and devised exact methods to gather the data and from the emerging patterns he generated his general ideas.Thus, even if his missed the mathematical and statistical skill of today he must be regarded as a pioneer of exact quantitative analyses of biogeographical data. He was very keen on this approach. Already in his first paper on frequency analysis (Raunkiær 1909) he expresses his motto: Numbers are the poetic meters of science (Tal er videnskabens versefødder). In this light it is not surprising that he chose 1000 plant species selected at random from the Index Kewensis when he set up his “normal” life form spectrum, and that he based his species abundance distribution J on 1000 vegetation analysis from all over the world.

Christen Raunkiaer around 1935

According to the many anecdotes told about him at the University of Copenhagen his exactitude gave the impression that he was pedantic and, as he also was rather introverted, he was not liked by his students – they rather feared him. In his time as professor ordinarius of botany (1911-1923) he had residence in the Botanical Garden and he insisted that the students use the garden as a living book of botany. Part of his teaching was questioning the students whenever he met them and often rather brusquely so. But at one point he surprised all the students because instead of all his unpleasant questions he just asked them to fold their hands – he then observed them, nodded, thanked and left them puzzled. The numbers he gathered was how many of them had the right thumb on top. Many of us know this analysis to be a classic in elementary statistical textbooks. Raunkiær may very well have been the first to make an exact count (on 1000 persons??). So he was actually very broadminded in his pedantry. Anyway, he took up his demanding behavior towards the students again, so much that the students sent a timid delegate to the professor to ask politely for a change. His response was remarkable: he looked coldly at the shivering student and said: ”I will not change my methods or my demands, but if the students are not satisfied by my teaching I will resign!” - and so he did, at an age of 63. This secured him a rather long emeritus period where he had peace to pursue his whims.

The allusion to poetry in his motto is another sign of his wider view. He must have been a keen reader of poetry. After his retirement he started “botanizing” in Danish poetry and he applied his “frequency analysis” as a method to characterize poets and epochs. The sample units were not circular area samples but verse lines and the observed objects were plant names. He ploughed through millions of verse lines written by hundreds of poets – so there was a need to define what a true poet is: It is a person who has published at least 1000 verse lines or one volume of poems. He determined the plant species spectrum for each of these poets in terms of total species richness and abundance distribution (abundance measured by number of mentions/1000 verse lines). So if you are interested and dedicated (and able to read Danish) you may learn that the neoclassicistic period had a much lower species richness and density then the romantic period, that Hans Christian Andersen was only the second Danish poet to mention heather, and that it is possible to distinguish between poets from Jutland and Zeeland by their predominant species. The style and stringency in his three publications on such matters are brilliant – and fun to read.

Shortly before he died he took up his old interest in plant geography. In one paper he adresses the relationship between range, species-genus ratio, and life forms. He selected the 10 largest plant families with in total 2772 genera and 43594 species (his own count!). He was able to show that genera that comprise more than one life form have considerable higher species to genus rate, and a wider geographical range. This search on the global scale for relationships between geographical features, taxonomical features and functional traits/types is a focal approach in biogeography even now.

And now we return to his island biogeography paper. It is from the same period, and apart from its undisputed scientific relevance it also contains statements about attitudes and sentiments that are shared by island biogeographers today. For instance its first sentence: “Islands, especially isolated oceanic islands, have always held a fascination for me, both emotionally and intellectually” – which happens to be the first sentence also of this essay. Thus, even if Raunkiær never would have ventured to add exaggerations or fantasies to his scientific publications, he certainly did not refrain from showing his sentiments. He also expresses modesty and self critique. His motto: ”Numbers are the poetic meters of science” is the prologue paragraph of his “Investigations and statistics of plant formations” from 1909. The paper has also an epilogue: ”Numbers are the poetic meters of science. Verses may halt, and so also may the numbers of science. I hope the numbers given in this work will be found to halt no more than their human origin inevitably entails”. This paragraph has never been more apposite than now: Biogeographers must remember that their brilliant theories and sophisticated models build on numbers (field data) that are of human origin, and as such they will sometimes halt so much that the entire theoretical construction becomes highly tottering.

Additional information about Christen Raunkiær including a complete list of his works and some biografies can be found at http://www.macroecology.ku.dk/resources/default.asp?p=Raunkiaer

References
Adsersen, H., 1995: Research on Islands: Classic, Recent, and Prospec­tive Approaches. In: Vitousek, P.M., Loope, L.L. and Adsersen, H. (Eds.): Islands: Biological Diversity and Ecosystem Func­tion. Ecologi­cal Studies vol 115, Springer, Heidelberg, pp.8-21.
Arrhenius, O. 1921. Species and area. - J. Ecol. 9: 95-99.
Darlington, P.J. 1957. Zoogeography. Wiley
Gleason, H. A. 1922. On the relation between species and area. - Ecology 3: 158-162.
Haila, Y. and Järvinen , O. 1982 The role of theoretical concepts in understanding the ecological theatre: a case study in island biogeography. – In Saarinen, E. (Ed.): Conceptual Issues in Ecology: 261-278. D. Riedel Publishing Company.
Hooker, J. D. 1847. On the vegetation of the Galapagos Archipelago, as compared with that of some other tropical islands and of the continent of America.- Transactions from the Linnean Society of London 20: 235-262
Lawesson, J. E., Adsersen, H. and Bentley, P. 1987. An updated and annotated check list of the vascular plants of the Galápagos Islands. - Reports from the Botanical Institute, University of Aarhus, 16.
Lomolino. M.V., 2000. Ecology’s most general, yet protean pattern: The species-area relationship.- Journal of Biogeography. 27: 17-26.
MacArthur, R. H. and Wilson, E. O. 1967. The Theory of Island Biogeography. - Princeton University Press. Princeton
McGill B. J. et al (17 coauthors) 2007: Species abundance distributions: moving beyond single prediction theories to integration within an ecological framework. Ecology Letters 10: 995-1015
McIntosch,R.P. 1962. Raunkiaer’s “law of frequency”. Ecology 43: 533-535
Palmgreen, A. 1922: Über Artenzahl und Areal sowie über die Konstitution der Vegetation. Eine vegetationsstatistische Untersuchung. Acta Forestalia Fennica 22:1-136.
Palmgreen, A. 1921. Die Entfernung als Pflanzengeographischer Faktor. – Acta Societas pro Fauna et Flora Fennica 49: 1-113
Palmgreen, A 1925. Die Artenzahl als Pflanzengeographischer Charakter sowie Der Zufall und die Sekuläre Landhebung als Pflanzengeographischer Faktoren. - Fennia 46: 1-139
Preston, F.W. 1948: The commonness, and rarity, of species. Ecology 29: 254-283
Preston, F.W. 1962.The canonical distribution of commonness and rarity: Part I: Ecology 43: 185-215, Part 2: Ecology 43: 410-432
Raunkiær, C. 1904. Om biologiske Typer, med Hensyn til Planternes Tilpasning til at overleve ugunstige Aarstider. Botanisk Tidsskrift 26: 14
Raunkiær, C. 1905. Types biologiques pour la géographie botanique. Videnskabernes Selskabs Oversigter 1905: 347-438
Raunkiær, C. 1907. Planterigets Livsformer og deres Betydning for Geografien. Copenhagen.
Raunkiær, C. 1909. Formationsundersøgels og Formationsstatistik. Botanisk Tidsskrift 30: 20-132
Raunkiær, C. 1930: Hjemstavnsfloraen hos Hedens Sangere Blicher og Aakjær. Schultz forlag, København.
Raunkiær, C. 1934. The Life Forms of Plants and Statistical Plants Geography being the collected Papers of C. Raunkiaer. Clarendon press, Oxford.
Raunkiær, C. 1936. The Life-form Spectrum of some Atlantic Islands. Botaniske Studier 4: 240-328
Raunkiær, C. 1937. Life-form, genus area, and number of species. Botaniske Studier 5: 343-356
Williams, C.B. 1964: Patterns in the Balance of Nature. Academic Press, London
Williamson 1984. Sir Joseph Hooker’s Lecture on Island Floras. – Biological Journal of the Linnean Society 22: 55-77

Communicating biogeography

Robert J. Whittaker was sole editor and then editor-in-chief of Global Ecology and Biogeography from 1995 to April 2004. He has been editor-in-chief of Journal of Biogeography since October 2004.



Some months ago, Joaquín Hortal approached me to write a piece for this newsletter, with the remit of providing some pointers as to what we look for in papers submitted to Journal of Biogeography. In some respects this is easy to do in that the journal policy is to be inclusive of major traditions within the field and international in scope, within which we give preference to papers posing and answering clearly phrased questions or testing hypotheses of general interest. However, having been given the opportunity, I also wanted to make some comment pertaining to presentation and rigour of papers in the natural sciences generally (i.e. not just submissions to Journal of Biogeography), which is a rather more risky thing to do. Not least because I might see my own words thrown back at me by referees of my papers in the future with the admonishment that I cannot follow my own guidelines as to best practice! I say this because I think it is deceptively difficult to write scientific papers well.
The process of writing the paper should not be regarded as just the bit that comes at the end, after all the challenging scientific stuff of laboratory analysis, experimentation and sophisticated statistical analysis has been done; rather it is integral to shaping the interpretation and developing ideas and arguments, which rarely come to the page fully formed. It involves establishing the context of the work through making multiple connections to the existing literature, which continues to grow at a phenomenal rate. All of this presents significant challenges to the scientific author, requiring a systematic and thorough approach to the task of constructing a paper. The end result must be accurate, meaningful and clearly communicated so that it is understandable by the target audience.

So how well are we doing and are there particular problems facing those starting out in biogeography? If this were a school report, I think the answer to the first part of the question would probably be an enigmatic ‘could do better.’ In illustration of concerns that I share, Todd et al. (2007) provide some quantification of the extent of mis-citations in a sample of 306 ecological papers selected from 51 journals, focusing on specific assertions made in the sampled papers. They report that some 7% of citations did not support the original statement at all, 11% were ambiguous, 6% were ‘empty’ (i.e. citing an opinion from a secondary source), with the remaining three quarters being ‘good’ citations. Todd et al. (2007) provide some comparative data suggesting that ecology may not be particularly unusual in having a significant level of sloppy citations, but, pulling no punches, they go on to conclude (p. 1660) that “mis-citing of references is widespread in ecology and is currently part of the normative values [my italics] of workers within the discipline.” I have given this example because: a) an effort has been made to quantify one key indicator of the rigour of scientific papers in a closely-related (overlapping) discipline to biogeography, and b) because it is something that we should all be capable of getting right. Based on my experience as a reader, reviewer and editor, I suspect that an analysis using the term ‘biogeography’ in place of ‘ecology’ would yield similar findings.

The question arises: how do errors arise within papers and what steps can we all take to minimize them? Todd et al. (2007) focus on the second part of this question, arguing that journals should ramp up their efforts to detect failings such as poor citation practice, implying that editors and reviewers should do more, whilst conceding that “…it is as authors that we have the most potential to improve citation practices.” My own take on this is that the onus should rest firmly and squarely on the authors of paper and not just with respect to citations but in all aspects of paper preparation and presentation.

One of my most rewarding roles as editor-in-chief of Journal of Biogeography is to scrutinise the final revised versions of all papers prior to formal acceptance. Whilst I get to read many wonderful and fascinating papers, setting out analyses of complex data sets with great skill, it is surprising how many papers (supposedly in their final form) contain significant flaws, internal inconsistencies, failures to follow guidelines, and failures to describe accurately the results presented in the tables and figures. Moreover, returning to citations, few papers (my own submissions to other journals included!) manage to compile reference lists that accurately match the list of papers cited. Such failings are despite the very best efforts of reviewers and editors and, as Todd et al. (2007) point out, they frequently sneak through into published papers in a wide range of academic journals. I suspect there are many reasons for such failures. These may include:

1. Pressure to publish, get the next research grant, etc, leading to rushed work.
2. Lack of guidance from senior colleagues or supervisors for those starting out in academic scholarship.
3. On multi-authored papers, senior colleagues may leave it to a single, often junior, author to undertake the revision, failing to provide adequate input (or any input!) on subsequent versions submitted to a journal.
4. On multi-authored papers, errors and mismatches may be introduced through sequential edits by different authors as they circulate drafts around by email (indeed I suspect that the error rate in multi-authored papers is generally higher than in single-author papers).
5. Minor errors are introduced during revision as a by-product of responding to reviewer/editor requests to add or correct analysis, or to drop analyses, tables or figures. The complexity of content of the average paper today means that it is almost inevitable that authors fail to update all of the detail in the text of second or third versions of a paper.
6. Some authors reason that if a submission to journal X has a 4 in 5 chance of rejection then it is not worth putting too much effort into getting things exactly right at first submission until they have seen if the referees like it, thus normalising a rather laissez-faire attitude to accuracy.
7. The problems of writing in a foreign language (which applies to many workers in biogeography).
8. The autocorrect function in the word-processor, combined with an inability to type accurately in the first place, introduces all sorts of minor errors, especially into reference lists, where they are hard to spot. So, some advice on Autocorrect: turn it off and instead run a separate spell check.
9. The complexity of much analytical work in biogeography is such that it is only possible with modern computers and spatial statistics, meaning that it can be extremely difficult for authors, reviewers and editors to check whether results are reasonable. However, the frequency with which minor errors creep through to late stages of manuscript submission raises the suspicion that insufficient efforts are made by many authors to check that their results are correct by replicating/cross-checking their analyses.
10. Finally, if the paper does make it through to production by a journal, there is always the possibility of errors being introduced during copy-editing or typesetting. Unfortunately, because everything is supplied electronically these days, many authors seem to think there is hardly any need to check their proofs. Think again!

I am sure that the above is a far from exhaustive list of ways in which errors become entrained in manuscripts during the process of their development, and I am also certain that they are not specific to biogeography. However, biogeography is a particularly diverse discipline, with many technical terms, sub-disciplines, and schools of thought, and it is a truly international subject. Whilst we can celebrate this diversity, some have also despaired of the failure of different schools of thought to engage fully within one another. I suspect this has at least something to do with deficiencies in the way ideas are developed and expressed in many biogeographical papers. So, I see it as very important that papers published in Journal of Biogeography, in addition to being accurate in their content and focused on a clear question or hypothesis of general interest, are also clearly written and as accessible as possible to a wide audience. At present, as Ladle (2008) reports, biogeography as a discipline is largely cryptic in the public sphere of the news media and the blogosphere, and in the UK at least, in school curricula. So, once you have overcome the “deceptively difficult” task of completing a paper, there is a further task, often neglected, of making steps to disseminate the findings in other media (e.g. through press releases) to help build a wider base of public understanding and appreciation of biogeography.

In summary, I concur with Todd et al. (2007) that the dash to publish is part of a complex array of factors leading to a degree of slippage from the rigour necessary in scientific work. In my view, the onus falls on senior academics, whether as colleagues, mentors, co-authors, supervisors, or educators, to emphasise repeatedly the importance of theoretical and empirical rigour in all aspects of science, from conception of a study to the preparation of each draft of a paper submitted to a journal. These comments may seem to suggest that the problems stem largely from inexperienced authors, but in fact I have seen well constructed and badly constructed papers written by people of all career stages, suggesting that much of it comes down to aptitude (i.e. traits of the authors) and training at the outset of a scientific career. In biogeography, there is no doubt that we have an increasingly sophisticated global community engaged in our science, as readily seen from the diversity of authors publishing in journals such as Journal of Biogeography. This provides evidence of a healthy and vibrant field. So, my critical comments should be seen in this light: as a community of scholars we are ‘doing well but could do better’ (so, still enigmatic but with a tone of encouragement). Finally, in purely pragmatic terms, if the goal is to get your paper published, it is worth following the sentiment of the old saying that if you watch the pennies, the pounds look after themselves. If you can prepare a manuscript in which all the fine detail is correct and clear, it is more likely that you will convince your target audience (reviewers, editors, and subsequent readers) that your findings are valid and meaningful, or (let’s not be too hasty here!) at least worthy of serious consideration and debate.


Preparing a manuscript for submission – some suggestions

• Read and follow the journal’s instructions for authors
• Make and complete a checklist of all the components of the paper that require cross-checking prior to submission
• In attributing ideas, check primary sources
• Not read it? Don’t cite it
• Co-authors should read and check the final (or penultimate) draft of both original and revised manuscripts prior to submission
• Ensure tables, figures and appendices are cited in numerical order
• Check the Results and Discussion sections to ensure the content accurately tallies with the final content of tables and figures
• Make sure you have your word processor set to the right dictionary (e.g. UK English or US English)

References
Ladle, R.J. (2008) Catching fairies and the public representation of biogeography. Journal of Biogeography, in press.
Todd, P.A,, Yeo, D.C.J., Li, D. & Ladle, R.J. (2007) Citing practices in ecology: can we believe our own words? Oikos, 116, 1599-1601.

Editor’s note: With this first commentary from Robert J. Whittaker we start a new series for the IBS Newsletter. We hope to open a discussion within the biogeographical community, starting with views from the editors of key journals publishing biogeography papers about current trends in publishing biogeography, the kind of manuscripts they would like to receive to be considered for publication in their respective journals, and how to improve the quality of our work. Here, the editors will have the opportunity to write on the topics of their choice, with the general aim of improving how research in biogeography is communicated to the scientific community and the public.

IBS 2009 - First announcement

Your IBS Board and specifically Ella Vázquez-Domínguez and the Local Committee at the Instituto de Ecología, Universidad Autónoma de México, México D.F. invite you to plan ahead now to attend the 2009 biennial meetings of the IBS in Mérida, Yucatán, México from 8 to 12 January 2009.

Mérida is in the midst of cultural, natural, historical, and geological riches. The beautiful colonial “White City” was founded in January 1542 on the ruins of the Mayan city of T’ho, and is only 120 km from the archeological wonders of Chichén Itzá. The center of the Chicxulub Crater, formed from an asteroid impact 65 mya and implicated in the K-T extinctions, is located 20 km outside of Mérida (in the pueblo of Chicxulub). Several biosphere reserves are within easy distance of Mérida (e.g., Celestun and Ría Lagarto Biosphere Reserves). Check out these and other enchanting features of Mérida on the web.

Climate should be ideal in January, and we’re planning field trips to explore the natural and archeological wealth of the region.

Your enthusiastic response to our survey about our biennial meetings told us what you most liked about our meetings and what needed improvement. The respondents also gave us clear directives on how we might improve upon them. Specifically, the membership particularly valued the personal, intimate nature of our meetings that encourages interaction and cross-discipline discussion. At the same time, members wanted increased opportunities for oral presentations, particularly for up-and-coming researchers and advanced graduate students; increased, dedicated opportunities for group discussions; longer time and increased space to view posters; and increased involvement of paleontologists, geographers, and geologists. The survey results offered specific means to these ends, and we are most grateful for this input. As a result of your comments we have altered the format of the meeting; we hope you like the results.

Board considerations for the Mérida meetings fall into three general categories: 1) format of symposia; 2) topics of symposia; and 3) limits on meeting size. We’re planning to experiment with several different symposium formats that will include that of previous meeting symposia; one that includes selected contributed papers; and we’ve agreed (after extensive debate) to experiment with ½-day of no more than 3 concurrent sessions. By creatively selecting concurrent session topics, we hope to avoid the typical effect of segregating participants by discipline.

We’re looking for symposium topics that will continue and enhance our success of involving biogeographers from diverse disciplines, intrigue those underrepresented sciences (e.g., paleo, geography, and geology) and the general public, and also be most pertinent to the regional setting. We’re currently considering the following topics for four symposia: Environmental Change and Ecological Collapse; K-T Boundary extinctions; Diversification Across the Neotropical-Nearctic Transition Zone; Biogeography of Paleoboundaries and Paleo Hotspots; Caribbean Biogeography—Integrating Marine & Terrestrial Biogeography; the Great American Biotic Exchange; Human Biogeography and the Biogeography of Infectious Diseases; and Asian American Disjuncts.

Additionally, we are very pleased to announce that John C. Avise will be in attendance to receive the 2007 – 2009 Alfred Russel Wallace Award, joining our first two distinguished recipients, John C. Briggs (2003 – 2005) and Jared Diamond (2005 – 2007). And we’ve already scheduled one workshop: “Visualizing Evolution in Space and Time.” For background information about the workshop, see the article by David Kidd in the Summer 2007 IBS Newsletter 5(2):6-8 (available online at http://biogeography.blogspot.com/2007/07/geophylogenies-uniting-space-and-time.html).

While the growth of our Society has been beyond our expectations and demonstrates both the vital role and enthusiastic support of our organization, we share the members’ concern that we don’t destroy the most valued aspects of our meetings by “loving it to death” with overwhelming numbers. Thus we’re setting a cap of 550 on meeting participation, and of 250 posters (in three sessions). It looks like it will be first-come, first-served; so stay tuned (via the IBS web site and Newsletter) for registration dates.

We’re working hard to respond to the membership’s input, and to build on the success of our first three biennial meetings. The Board will be meeting in January to finalize the format and topics of the meetings. If you have further input or questions, please email the VP for Conferences (Dave Hafner) at david.hafner(at)state.nm.us. If you would like to suggest speakers or titles of talks for the above symposia, please send that information to Dave. 2011 IBS Meetings.—It’s never too early to plan on the next meetings! If you have suggestions for a meeting venue, or (even better) would be interested in hosting the 2011 meetings, let Dave Hafner know!

Dave Hafner, VP Conferences and Ella Vázquez-Domínguez, director-at-large
Check the latest news on the conference at http://www.biogeography.org/meetings.htm