Is there a clear division between Historial and Ecological Biogeography?

           Biogeography is the area of biology that studies the geographic distribution of organisms and the reasons why they occur (or don’t) on certain places of the Earth. Traditionally it has two approaches: the historical, focusing on larger temporal and spatial scales, relating the present distribution pattern with evolutionary processes (e.g.: speciation by vicariance), and the ecological biogeography, that studies how ecological processes, such as climate, altitude and other environmental agents affects the distribution, usually taking into account a short time interval and small areas. But can we assume that there is a clear division of the biogeography?

Resultado de imagem para biogeography

Ratite’s biogeographical distribuction explained by the continental drift.

          For a long time the climate was considered to be the best factor to explain the organisms’ actual distribution, but with a better understanding of biological interactions and processes, it became obvious that only one characteristic couldn’t explain the current pattern of species occurrence, causing some researchers to criticize the ecological biogeography. But it isn’t just because the climate isn’t a variable that explains everything that we can disregard the effect of other landscape components on the delimitation of the biological distribution, seeing the diversity of niches that we have in the present.

          As an example, some abiotic factors, like substrate characteristics, can define the distribution of gastropods in Britain and echinoderms in Colombia; it all depends on the scale that we are working with (maybe the substrate doesn’t mean anything when we are analyzing the global distribution of gastropods). And this is just one of many ecological factors that takes part on the delimitation of an organism’s area of life: even ecological interactions and behavioral differences can create an non-physical barrier that constrains the dispersal.

           Interacting with these ecological processes we have historical ones, like vicariance and the landscape changes (e.g.: orogeny and volcanism) that shaped the present habitat diversity, and created many ecological niches, making it hard to discern between these two approaches.

          In other words, ecological processes do define species spatial distribution, but many landscape characteristics resulted from long-time geological mechanisms. If we see biogeography like this, can’t we say that the ecological biogeography is kind of consequence of the historical? That could be true in the past, but now we have an important landscape modifying agent: the human being.

           The age we are in, the Anthropocene, habitat modifications that would take centuries to occur naturally can happen in less than a decade, and it may even cause speciation or hybridization in some cases, like with the crab-eating fox (Cerdocyon thous) and the hoary fox (Lycalopex vetulus). Those two canids most likely hybridized by induction of the deforestation of the Atlantic Forest (alteration of natural habitats), connecting both species. So we’re having a process usually related with the historical biogeography, occurring in a short time period, caused by an alteration of the landscape of an area.

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Lycalopex vetulus (left) and Cerdocyon thous (right).

           Seeing how the two divisions of biogeography are increasingly merging, we should expect more researches considering both the historical and ecological approaches at the same time, or even as if they were one.

Literature:

Garcez, F. S. 2015. Filogeografia e história populacional de Lycalopex vetulus (Carnivora, Canidae), incluindo sua hibridação com L. gymnocercus. Dissertação (Mestrado) – Curso de Zoologia, Pontifícia Universidade Católica do Rio Grande do Sul, Rio Grande do Sul, 2015. Available on: http://repositorio.pucrs.br/dspace/handle/10923/7711. Access on: 10 abr. 2017.

Haila, Y. 2002. A conceptual genealogy of fragmentation research: from island biogeography to landscape ecology. Ecological Applications, 12(2), 321–334.

Heads, M. 2015. The relationship between biogeography and ecology – envelopes, models, predictions. Biological Journal of the Linnean Society, 115, 456–468.

Monge-Nájera, J. 2008. Ecological biogeography: a review with emphasis on conservation and the neutral model. Gayana 72(1), 102-112.

Santos, C.M.D. & Amorim, D.S. 2007. Why biogeographical hypotheses need a well supported phylogenetic framework: a conceptual evaluation. Papéis Avulsos de Zoologia, 47(4): 63-73.

What about historical biogeography in the 21st century?

            As life and Earth evolve together, biogeography can be considered as the area of biology that pursuits to establish patterns of biotic distribution and connections between the biotas, both resulting from the evolutionary process, being closely related to the triad: form, space and time. In this context, biogeography seeks to understand such patterns by addressing a set of biological and geographic information in order to describe the distribution of organisms on the planet and to give explanations for the history that would have led to such spatial configurations. On its larger scale, biogeography provides the historical perspective required to understand the evolution of biota as well as the geological evolution of the planet.

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           Traditionally, biogeography has been divided into ecological and historical. Ecological biogeography studies how ecological processes (such as climate or other physical or environmental factors) interfere in the distribution of organisms in a short period of time, while history biogeography comprises the action of processes, based on causes that existed in the past, over a long period of time in this pattern. Historical biogeography is in the midst of a scientific revolution and that it is a rapidly evolving discipline. As a result of this a great number of approaches and methodologies arose in the most recent decades (second half of the 20th century and the first years of the 21st century).

So this drives to the question: What is the real present state of historical biogeography?

               According to a study made by Posadas et al. (2013), based on the papers published in the Journal of Biogeography (JB) during 1998–2010, there is a clear tendency to increase the number of contributions per volume, which seems to indicate that biogeography as a whole is an expanding research field. However, historical biogeographical contributions have shown a more marked increase. While the number of total contributions published in JB more than doubled from 1998 to 2010 (from 90 in 1998 to 200 in 2010), the historical biogeographical contributions increased almost 10 times during the same period (from 11 in 1998 to 105 in 2010).

1Number of articles published in each volume of the Journal of Biogeography during 1998–2010 (total = black bars; historical biogeography = gray bars) (Posadas et al., 2013).

            Out of a total of 2095 papers analyzed, 610 papers (29%) deal with historical biogeography, wich were written by 2018 authors, with a mean of 3.3 authors per paper. A trend toward more authors per paper is evident. This could be an indicator of increased cooperation/collaboration in historical biogeographical work, a pattern not exclusive to historical biogeography but found in most of the contemporary scientific journals. The increasing specialization of researchers, the emergence of more complex techniques to address problems, the presence of multidisciplinary teams in research institutions and  the significant impacts on productivity and citation rates could explain in part this growth.

3Percentage of papers authored by one (black bars), two (diagonal line bars), three (horizontal line bars), and four or more authors (gray bars) per year. Trend lines added for papers authored by a single author (black) and by four or more authors (gray) (Posadas et al., 2013).

               The analysis shows a strong trend favoring works devoted to terrestrial habitats (75% terrestrial, 12% freshwater, and 13% marine habitats). A possible explanation for this particular distribution of papers is that it is strongly biased by the degree of difficulty involving access to different habitats, as well as to the smaller average cost of accomplishing taxonomic works in continental environments (including freshwater) relative to marine environments. An alternative explanation for this pattern is related to the distribution of species diversity on continents (including freshwater habitats) and oceans (87% inhabit continental habitats and the remaining 13% inhabit marine). Thus, the study of different habitats seems to be more influenced by taxon diversity than by the surface covered by each habitat.

                  From a taxonomic point of view, at the kingdom level  it was observed that 59% of the papers dealt with animal taxa, 35% with plant taxa, and 4% with taxa from more than one kingdom; only 2% of the papers dealt with taxa from other kingdoms (Protoctista, Fungi). The percentage of papers devoted to higher taxonomic groups correlates better with ‘the inefficiently distributed’  labor force in taxonomy than with taxonomic diversity. The distribution of historical biogeographical papers does not reflect the real diversity of these phyla, the megadiverse animal groups, for example, such as arthropods (particularly insects), nematodes, and marine invertebrates are underrepresented.

                   In relation with approaches and techniques, five approaches were used in almost 72% of the papers considered: phylogeography (35%),   biota similarity and PAE approaches (13%), molecular biogeography (12%), and cladistic biogeography and event-based methods (6% each). The great surge in phylogeographical papers seems to lead the evolution of historical biogeography over the last years. The increase in the number of historical biogeographical contributions per year has accompanied the growth of phylogeographical papers . The fact that phylogeography is the main force which is driving historical biogeographical research in recent years is reflected, for example, in the increase in papers dealing with infraspecific taxa and in those that use molecular clocks. Thus, the taxonomic scale is showing increasing representation of those papers focused on a single species or a few closely related species.

2Distribution of papers applying the five most used approaches per year (Posadas et al., 2013).

               Historical biogeography is a research field that has increased its representation throughout the analyzed period. The emergence of new approaches, the combination of techniques, and the increase in questions indicates that historical biogeography is an active discipline. However there are many challenges in the future to understand with more completeness the the infinite variables that influence the existing patterns and processes. Some of them are:(1) to increase the study of those taxa underrepresented according to the part of the biodiversity they represent; (2) to balance the amount of work devoted to different biogeographical regions; (3) to increase biogeographical knowledge of marine and freshwater habitats; (4) to maintain the diversity of approaches, preventing the reduction of time, spatial, and taxonomic scales addressed by the discipline; and (5) to continue integrating historical biogeography along with other sources of information from other disciplines (ecology, paleontology, geology) into a richer context for explaining past, present, and future patterns of biodiversity on Earth.

Literature:

Almeida, E.A.B. & Santos, C.M.D. 2011. Lógica da biogeografia de vicariância. In: Carvalho, C.J.B. & Almeida, E.A.B. (ed.), Biogeografia América do Sul:Padrões e Processos. São Paulo: Roca, pp. 52-62.

Crisci, J.V., Katinas, L. & Posadas, P. 2003. Historical Biogeography: an introduction. Cambridge: Harvard University Press.

Posadas, P; Grossi, M.A. &  Ortiz-Jaureguizar, E. 2013. Where is historical biogeography going? The evolution of the discipline in the first decade of the 21st century? Progress in Physical Geography, 37(3): 377–396.

Santos, C.M.D. & Amorim, D.S. 2007. Why biogeographical hypotheses need a well supported phylogenetic framework: a conceptual evaluation. Papéis Avulsos de Zoologia, 47(4): 63-73.