The Multi-Level Approach
(self-organisation of higher order biodiversity)
John Couwenberg
1. Terminology
1.1 Biodiversity: diversity related to living entities.
1.1.1 Diversity: difference in space and space-time; i.e. differences in character, number and distribution.
1.1.2 Entities: all that, which can evolve. Entities are all patterns (spatial and spatio-temporal: elements, relationships, processes) which show variability and reproduction (continuation of information) and which are thus prone to selection, on all possible levels (evolution is a multilevel process).
1.2 Ecosystem: system among the elements of which there are
living entities.
2. Introduction:
the concept of biodiversity on the ecosystem level
2.1 Biodiversity on the ecosystem level is one of the three main 'levels' of biodiversity, stressed on the 1992 UNCED biodiversity convention.
2.2 The three main levels of biodiversity: a) genetic (lower than species level); b) species; c) ecosystem (higher than species level).
2.3.1 Biodiversity on the ecosystem level was originally conceived as biotope diversity: based on species diversity, with the notion of a species in its natural environment being 'worth' more than in an artificial one. This is the classical species-centred viewpoint.
2.3.2 Within this species-centred frame, mires represent an environment/biotope species occurring in them. Arguments for conservation of biodiversity on the higher order ecosystem level may be based upon this; i.e. conservation of the diversity of Biotopes in which species occur.
2.3.3 The UNCED division is based on a mixture of evolutionary and ecological ordering: a species is not a pure ecological entity (one cannot go 'out there' and see a species), it is an eco-evolutionary entity, and the concept of 'species' (and 'genes') is primarily based on evolutionary considerations
2.4 An ecosystem can be viewed both as an ecological (static) as well as an evolutionary (evolving) entity, dependent on timescale. In the species-centred viewpoint, ecosystems are viewed as ecological (static) entities: they represent the 'home' of a group of organisms. (This group of organisms is however not a group of species.)
2.5 Higher order biodiversity is interpreted here as diversity of ecosystems.
2.6 This is the ecosystem-centred viewpoint: diversity on the ecosystem
level is not found merely in species-biotope diversity, but in diversity
of the ecosystem as such.
3.1 Self-organisation is based on building and maintaining differences in information (structural and dynamical). Building means ordering: some differences are strengthened and others are weakened; 'some differences and others' implies variance, 'maintaining' implies continuation and 'strengthening and weakening' implies selection. Self-organisation leads to less instability (i.e. adaptation).
3.2 As the term 'building' implies, self-organisation leads to new levels of higher order. Higher order levels represent semi-autonomous entities, which display characteristics, that the lower order constituing entities do not. Example: a brick does not provide shelter, a house (organised collection of bricks) does.
3.3 Nature is spontaneity (in organisation). The terms 'natural',
'spontaneous' and 'self-organising' are interchangeable: nature defines
itself.
4.1 A feedback loop is any kind of information that stems from a higher order entity, and has its influence on a lower order entity.
4.2 This influence can be positive, enhancing; or negative, diminishing.
4.3 Self-organisation can, based on this, be split up into two distinct parts: building the difference and sustaining difference, or: making and maintaining.
4.4 Making (building, enhancing) difference is primarily based on positive feedback loops. Examples: Some Sphagnum species grow to form a hummock, the hummock favours these specific Sphagnum species; acidification by Sphagnum, which favours growth of Sphagnumover other plants.
4.5 Managing (maintaining, diminishing) difference is primarily based on negative feedback loops. Example: if a hummock grows too high, it becomes too dry, which is unfavourable for the hummock forming Sphagnum species.
4.6 Positive and negative feedback loops are continuously having their
influence in different forms and on different spatial and spatio-temporal
scales simultaneously (different intensity and frequency).
5.1 Several forms of self-organisation can be recognised within mires. Most important seems to be self-organisation based on water (see also 'the concept of peat' and 'hydro-genetic mire types').
5.2 Water is everywhere in (most) mires and because it flows, it can relay information from one place to another. (cf. boats on and chemical waste in rivers.)
5.3 The most important 'message' carried by water in the self-organising context of a mire is that of stagnating/non-stagnating parts of the mire:
5.4 Stagnating parts are those parts that hamper the flow of water (e.g. 'catotelm, hummock), non-stagnating parts are those parts that do not hamper the flow of water as much as the stagnating ones (e.g. 'acrotelm', hollow).
5.5 Stagnation is dependent on time scale (see examples above). Actually everything is non-stagnating, one just needs to wait long enough. When talking of stagnating and non-stagnating elements, there is always a specific time scale involved.
5.6 Within a mire, stagnating/non-stagnating elements can be discerned
on various spatial and spatio-temporal scales.
6.1 On a low level (zero-level) one can find stagnating and non-stagnating elements in presence and absence of plant-material; i.e. plant tissue or not-plant tissue.
6.2 On a somewhat higher level (first-level), stagnating/non-stagnating is determined by a single Sphagnum plant and follows from the ratio of zero-scale stagnating/non-stagnating elements. (Clonal and populational level of Masing 1998)
6.3 On a second-level, stagnating/non-stagnating can be found on the spatial scale of the mire feature or microform (Masing, 1972; Microstructural, Masing 1998): hummocks, hollows pools, tussocks, etc...
6.4 On a third-level, stagnating/non-stagnating can be found on the spatial scale of the mire-site (Masing 1972; Microtope or coenocomplex, Masing 1998), complex (Osvald 1923): hummock-hollow patterns, large pools (Fig 3, from Sakaguchi 1980).
6.5 On a fourth level, stagnating/non-stagnating can be found on the spatial scale of the mire complex (Masing 1972; Mesotope or mesostructural, Masing 1998), the mire as a whole: stagnating bogs in a sea of non-stagnating fens e.g. in Minnesota (Wright et al. 1992).
6.6 On a fifth level, stagnating/non-stagnating can be found on the
spatial scale of the mire system (Masing 1972;
Macrotope or macrostructural and Regional, Masing
1998), landscapes with mire and non-mire areas (Fig. 4, from:
Romanova 1985)
7.1 Orderedness is the result of (self-)organisation.
7.2 There are many different 'governing-principles' in self-organisation: physical, chemical, biological (see 9.2).
7.3 Each governing principle creates a different set of scale levels (see e.g. 6). This we call a scale series.
7.4 Each governing principle has a spatio-temporal 'frequency': the 'size' of the 'scale-jumps' that are made in its scale-series (see e.g. Masing 1998).
7.5 Each governing principle has an 'intensity': some governing principles may be more dominant than others. This dominancy can however differ from scale level to scale level.
7.6 Since different governing principles exist, observation of orderedness is a question of knowing how to look (what to look for) (cf. 9.2).
7.7 Since governing principles have different spatio-temporal frequencies, observation of orderedness is a question of knowing where (and how long) to look.
7.8 When governing principles of a comparable intensity but with different frequency coincide, the system will appear less ordered (cf. Fig. 5, from Swanson & Grigal 1988)
7.9 When among different governing principles of different frequency, one is dominant, the system appears ordered (cf. Fig. 5, from Swanson & Grigal 1988).
7.10 In some ecosystems orderedness is hard to see (e.g.: tropical rain forest), in others it is easy to see (e.g.: 'kermi bogs'). This difference of 'observability' of orderedness should be treated as a form of biodiversity on the meta-level of different main-types of ecosystems. (As should the 'small' number of species that occur in mires!!). ­ see 9.
7.11 'knowing how/where to look' implies that the scales on which higher order levels can be observed is based on subjective choice. Because this choice is however shared by majority of humanity, it can be viewed as 'pseudo-objective' and be treated as such. Mires can be seen as a main-type of ecosystem in which water governs the self-organisation
7.12 Within a main-type of ecosystem (e.g. mires), the different levels
of self-organisation, i.e. the levels over which the feedback loops 'run',
i.e. the levels on which semi-autonomous entities can be discerned, should
provide the basis for defining higher order biodiversity.
8.1 Masing 1972, 1998 provides a structural classification on the different self-organisational (feedback) levels (2nd-5th) mentioned above (see also Zobel & Masing 1987).
8.2 Zobel & Masing 1987 provide a spatio-temporal classification of community dynamics on the different levels (2nd-5th) mentioned above.
8.3 Masing 1984 provides a classification based on the developmental stage on the 4th level (Fig 6, from Masing 1984).
8.4 Sakaguchi 1980 provides a classification
on the 3rd level: based on the pattern hummock-hollow
complexes shown (from the air: Fig. 6; see also Sakaguchi
1982).
9.1 Higher order biodiversity can be approached from several ways (we're talking multi-level here!!); Information (observables) can be found in:
9.2 Differences in the character of governing principles: physical (e.g. stagnating/non- stagnating, sunlight reflecting/non-reflecting), chemical (e.g. ions hemming apical dominance), biological (e.g. growth of Sphagnum species as dependent on water table).
9.2.1 Different organisation governing processes need different kinds of observation / description (see examples 9.2).
9.2.2 Different organisation governing processes are of a different 'strength' or 'dominance' and may result in stronger or weaker organisation (see 7.8, 7.9).
9.3 Differences in the number of governing principles: a tropical rainforest is probably more complex in this respect than a mire (this may primarily be caused by the number of different 1st level elements (species) and their mutual relationships; i.e. first order complexity).
9.4 Difference in the orderedness of a system see 8.
9.5 Differences in number of organisational levels: number of feedback-levels that can be discerned in a system, number of scales in a scale-series.
9.5.1 Spatially larger systems usually show a larger complexity in this sense
9.5.2 Temporally older systems usually are spatially larger; they have had more time to develop complexity. They carry more time as information (cf. palynological 'archive'-function of mires).
9.5.3 When the higher order levels are present, the lower order levels are so as well. In presence of systems incorporating the higher order entities, systems which do not incorporate them become an additional form of diversity. Example: a temporary hiatus in a peat deposit is additional information only if a complete time series is also available.
9.6 Differences in the entities on levels: number of different entities on a certain level.
9.6.1 Elements with different characters can be discerned on one level: e.g. hummock, hollow, lawn, carpet...
9.6.2 On a lower observational level on the same organisational level, one may also discriminate different types of the same sort of entity: deep, shallow, dish-shaped, symmetrical, assymetrical (etc...) hollows.
9.6.3 i.e. Biotope diversity (cf. 2.3.2).
10.1 Transformative value (Norton 1984) is an information function (cf. 'Mire classification for nature conservation,' discussion document by Joosten), and refers to the impulse to alter or transform existing preferences.
10.2 Current nature conservation is species-centred (2.3.1) Mires, because of their orderedness provide a paradigm system for learning to 'think multilevel' (ecosystem-centred).
10.3 Mires herewith provide a positive-feedback mechanism for finding
new ways of looking at nature and its conservation.
consequences for nature conservation
workshop challenge
Masing, V. (1972) Typological approach in mire landscape study (with a brief multilingual vocabulary of mire landscape structure). In: Estonia - Geographical studies, pp. 61-84. Tallinn: Eston. Geogr. Soc.
Masing, V. (1984) Estonian bogs: plant cover, succession and classification. In: European Mires (ed. by P.D. Moore), pp. 119-148. London: Academic Press.
Masing, V. (1998) Multilevel approach in mire mapping, research and classification. Contribution to this workshop
Norton, B.G. (1984) Environmental ethics and weak anthropocentrism. Environm. Ethics 6: 131-148.
Osvald, H. (1923) Die Vegetation des Hochmoores Komosse. Svenk. Växtsoc. Sällsk. Handl. 1: 1-436.
Romanova, E.A. (1985) Rastitel'nost' bolot. In: Il'ina, I.S., Lapsina, E.I., Lavrenko, N.N. i dr.: Rastitel'nyj pokrov Zapadno-Sibirskoj raviniy, pp. 138-140. Nauka, Novosi-birsk.
Sakaguchi, Y. (1980) On the genesis of banks and hollows in peat bogs - an explanation by a thatch line theory. Bull. Dep. Geogr. Univ. Tokyo, 12: 35-58.
Sakaguchi, Y. (1982) Characteristics of bank and hollow complexes in the Ozegahara moor. Sc. Res. of the Highmoor in C. Japan 31-46
Swanson, D.K. & Grigal, D.F. (1988) A simulation of mire patterning. Oikos 53: 309-314.
UNCED (1992) Convention on biological diversity. Rio de Janeiro.
Wright, H.E., Coffin, B.A. & Aaseng, N.E. (eds.) The Patterned Peatlands of Minnesota, 327 p. Minneapolis: Univ. of Minnes. Press.
Zobel, M. & Masing, V. (1987) Bogs changing in time and space. Arch.
Hydrobiol. Beih. Ergebn. Limnol. 27: 41-55.
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