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Biogeography
is the science that studies the relationships among the distributions of
organisms at present and in the past (Westhoff et al. 1970). On the basis of the
recognition that certain plant families only occur in certain parts of
the world, the world has been split up into five floral kingdoms
, each with a set of characteristic plant families. With increasingly fine
mechanisms of distinction, biogeographers have further subdivided those areas,
using genera for the next level down, the regions and species
for the provinces and lower subdivisions in the hierarchy. By bundling
apparent coincidence of distributions among certain taxonomic groups,
biogeography provides special information for identifying partially different
assemblages of species with a tendency to share distribution ranges, without
having to know them all. Hence, biogeographical units may be used as identifiers
of distribution by proxy (Vreugdenhil 2002) or surrogates (Faith et al. 2001).
With each further subdivision in the system, the number of distinguishing
species decreases while the number of overlapping species increases, thus
decreasing the degree of overall distinction. Udvardy
(1975) divided the world into eight realms, which were each subdivided into
different biomes. Van der Hammen and Cleef (1986) and Van der Hammen and
Hooghiemstra (1996, 2001) very convincingly present, respectively review, the
latest insights on the origin of the phytogeographical distribution of vascular
plant genera in the neotropical Andean forests on the basis of paleo-ecological
analysis. It is fascinating to read how paleoecologists can trace back the
spreading of families, genera and sometimes even species across the globe,
spanning millions of years, using such inconspicuous clues as pollen, shells,
imprints of plants, soil composition and nowadays, genetic composition.
Paleoecological analysis in mountainous regions works much faster than in
lowlands, as it is easier to interpret ecosystem changes along temperature
gradients (Cleef, 1979, 1980). Lowland analysis requires a much broader net of
systematically organised soil profiles and sediment sections (A.M. Cleef 2003,
pers. com.). Particularly in the humid tropics, the distribution of species is
much more difficult to define and consequently when staking out biogeographical
units one is still likely to see less reliability and geographical precision in
present-day distributions in tropical lowlands than in mountain regions (A.M.
Cleef and T. van der Hammen, pers. com.). Prance
(1982 in 1989) proposed that during Pleistocene glacial advances, when the
climate of the region became drier and cooler, forest became fragmented into
“Pleistocene lowland forest refugia” in Central and Northern South America.
Independent speciation within those refugia would have lead to a large amount of
local endemism, often referred to as “centres of endemism”. These proposed
refugia are mainly based on the distribution of four woody plant families,
Caryocaraceae, Chrysobalanaceae, Dichapetalaceae and Lecythidaceae. Davis et al.
(1996), somewhat seem to question this “popular theory”, stating “Whether
or not they were refugia, the “fact” that centres of endemism1
exist for a large number of different organisms has been well established”.
Yet, there have been suggestions, that these apparent centres of richness are
merely well-collected areas”. The latter warning must be taken seriously given
such findings on endemic species (See 2.3.3) as presented for instance by House
et al. (2002). The latter advised extreme caution considering how difficult it
is to make any quantitative comparisons on species richness in a botanically
relatively well-sampled2
country like Honduras. If the knowledge about the distribution of all species -
including endemic species - is so highly biased by road access and centres of
investigation in such a small country, sampling bias is very likely to occur in
the vastness of the Amazon as well. Vreugdenhil (e.g. 1992, 1997, 2002) has
organised or participated in the organisation of the collection of data in most
countries in Central America and has seen how lopsided and often fragmentary
data sets can be. House et al. (2002) suspect there to be good reason to even
doubt the status of a part of the less-conspicuous endemic or limited
distribution species, as their distribution ranges just have not yet been
discovered. To go from such sketchy information to identify “centres of
richness” and “centres of endemism” requires great scientific caution.
Other modifiers are likely to capture such situation. The
CBD approaches biodiversity selection on a country-by-country basis. Within a
worldwide strategy to seek the greatest possible representation of ecosystems
and species, this has a hidden advantage: not only does it deal with the fact
that conservation must have a solid legal and management basis, which primarily
is dependent on national legislation and national management organisations, but
in most cases, the world’s division into national territories automatically
leads to plural representation of the world’s recognised biogeographical
regions, including the WWF’s “Global 200”. Only in some very large
countries, like Brasil, Congo, India, Russia, Anglo-America would it be possible
to develop a PA system with the omission of one or more entire biogeographical
regions. For a national protected areas system composition analysis of medium
sized to small countries, biogeographical divisions are not strictly necessary,
as detailed ecosystem maps and the selection criteria elaborated later in this
document lead to much more fine-tuned area composition than possible with
biogeographical regionalisation while the country-based approach de facto
serves as a course proxy for biogeographical regions. This is particularly the
case if PA systems are spread across a country’s entire territory as
recommended in Chapter IV. This method is also much more precise than the
approach suggested by Prance or the EBAs of BirdLife International. Furthermore,
Davis’ suggestion that concentrations of endemism and high species diversity
would go together is not necessarily the case: in the humid tropics, high
endemism is particularly expected at higher elevations of isolated mountains,
where biodiversity is much lower than in the surrounding humid tropical
lowlands. Dinerstein
at al. (1995) developed a hierarchical classification scheme that divides Latin
America and the Caribbean (LAC) into 8 Bioregions, 5 Major Ecosystem
Types (METs), 11 Major Habitat Types (MHTs), and 191 ecoregions.
A
bioregion is defined as a geographically related assemblage of ecoregions
that share a similar biogeographical history and thus have strong affinities at
higher taxonomic levels (e.g. genera, families). A
MET is a set of ecoregions that: 1.
share comparable ecosystem dynamics(3*);
2.
have similar response characteristics to disturbance (*); 3.
exhibit similar degrees of beta diversity (dependent on vast data sets)
and 4.
require an ecosystem specific conservation approach (*). The following classes have been identified: Tropical Broadleaf Forests,
Conifer/Temperate Broadleaved Forests, Grasslands/Savannahs/shrublands; Xeric
Formations; Mangroves. A MHT is a set of ecoregions that: 1.
experience comparable climatic regimes; 2.
have similar vegetation structure; 3.
display similar spatial patterns of biodiversity (*); and 4.
contain flora and fauna with similar guild structures and life
histories(*). The following MHTs have been distinguished for the study region.
Tropical Moist Broadleaf Forests; Tropical Dry Broadleaf Forests; Temperate
Forests; Tropical and Subtropical Coniferous Forests; Grassland Savannahs and
Shrublands; Flooded Grasslands; Montane Grasslands; Mediterranean Scrub; Deserts
and Xeric Shrublands, Restingas and Mangroves. An ecoregion is a geographically distinct assemblage of natural
communities that 1.
Share a large majority of their species and ecological dynamics [requires
large data sets; 2.
Share similar environmental conditions; and 3.
Interact ecologically in ways that are critical for their long-term
persistence (*). Ecoregions within the same MHT can be similar in their structure and
ecological processes, but they share few species. When evaluating the suitability of the eco-region classification system
of Dinerstein for biodiversity mapping purposes, the following observations
should be made. In the definition, the system pretends leans heavily on
biogeographical history and, in its classification levels use distinct
physiognomic criteria. However, the system is not organised systematically and
it uses modifiers that can’t be clearly be identified in the field. At the
level of the MET, none of the four characteristics as formulated in the
definition can be measured objectively and require expert consensus building
through workshops. As such, they can’t be mapped from remotely sensed imagery
and as result, the main ecoregions map in Dinerstein (1995
) must be considered highly speculative. Of the MHT, only the first two criteria
may be defined and identified objectively, but one would need a set of properly
defined selection criteria. Again, the MHTs can’t be mapped from remotely
sensed imagery, as half of its modifiers are unidentifiable. The primary characteristics of the ecoregions as they appear in the
definition cannot be measured objectively and analysis of these regions suggest
that they are geographically distinct versions of the MHT, which are more than
anything else, coarsely separated physiognomic climatic classes. The method is
intrinsically weak as it builds on a combination of a set of modifiers
consisting of consensus derived versus mappable criteria and a number of poorly
defined modifiers. Therefore, it cannot be reproduced or complemented by
independent researchers; with poorly defined modifiers, users cannot know the
criteria of distinction of the species assemblages. The ecoregions approach has
been designed for continental applications at a scale of 1:10,000,000 (D.J.
Graham, pers. com). Yet, the map has been very useful for giving a first quick
impression to analyse where major gaps occurred at a continental basis, using
the available techniques of its time, but for national protected areas system
analysis, it is too unspecific. Each one of the levels of Dinerstein’s method has elements that can not
be used as a classifier as they are either not defined and/or not readily
identifiable in the field or from remotely sensed data. The method must be
considered a one-time product that cannot be reproduced. The biogeographical focus for demarcating species regionalisation
patterns would be the strength of this mapping concept, but the scientific basis
for it is weak. Biogeographical distinction should not be derived from consensus
building workshops but from field-data showing patterns of distribution
variation, and much more work needs to be done to consolidate the foundation of
species regionalisation patterns. Usually workshops tend to strive for
consensus, but the results are not always objective and reproducible. They are
important for agreeing on objectives, methods, rules and criteria, verification
and for joint evaluation of the results. Actual classification and delineation,
however, should result from objectively identifiable and reproducible modifiers.
To summarise: as a distinct modifier biogeographical regionalisation may
significantly contribute to an ecosystems classification system. However, the
level of feasible detail is subject to the knowledge about the distribution of
indicator families, genera or species for such regionalisation. A system
primarily focussing on biogeographical patterns is too coarse for the design of
national protected areas systems, but it may serve to pre-analyse worldwide
representation of coarse sets of species, as seems to be applied by the
Worldwide Fund for Nature, (WWF, http://www.panda.org/resources/programmes/global200/pages/home.htm
2002). 1 Endenism in the sense of limited
distribution species (see 2.3.4) 2 This qualification must be seen in the
context; obviously, the sampling of Honduras remains much to be desired. 3 * marked criteria are not clearly defined
or identifiable. This page is part of our web-book on Biodiversity Conservation. For organized reading go to our on-line Table of Content, or download our book in pdf format. |
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