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An Approach to the Agrarian History of Upland Country: Ecology and Habitat*

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An Approach to the Agrarian History of Upland Country: Ecology and Habitat* An approach to the Agrarian History of Upland Country: Ecology and Habitat* By ADRIAN H COWELL HE agrarian history of upland country the dynamic processes operating or to has not been widely studied and one of describe numerically the vegetational pat- T the reasons could be, as Estyn Evans terns in order to generate explanations for explains, that 'one of the attributes of a what is discerned. Because man's economic society which is essentially pastoral is that activities are a biological factor in the much of the documentation is lacking'.' The competitive interactions between orga- consequent lack of information about the nisms, they are of interest to the ecologist so social and economic implications of changes that the distinctions between historical in husbandry techniques may be overcome ecologists and ecological historians depend by the application of quantitative ecological rather more on the application of the study methods for, as Sir George Stapledon has than upon the methods employed. As argued, 'vegetation accurately read is a Stapledon has commented 'There are no remorseless and objective historian'.-" disciplines, only problems'.3 It is argued here that the deficiencies of Recondite discussion of the mathematical documentation so lamented by Estyn Evans and statistical theories involved in the may be made good by the type of analysis analysis of vegetation does not seem favoured by Stapledon. In this review of appropriate here and interested readers are ecological methods and studies of the referred to accounts by Kershaw and Greig- components of upland systems the object Smith. 4 It should be emphasized, however, will be to establish their relevance to the that all techniques used in the analysis of elucidation of the agrarian history of the pattern should be subordinated to biological British uplands. It should be said here that the considerations. Some explanation of the term 'upland' is applied sensu latu so as to techniques is required. Numerical studies include both upland and hill farming, rather since the war have contributed to the idea than restricted to definitions of altitude or that vegetative pattern is not generally vegetational status which would here create explicable in terms of chance, s The physiol- confusion. ogy, reproductive strategy, and dispersal Ecology is concerned with the biological mechanisms, the anatomy, morphology, relationships between organisms in a habi- and habit (i. e. whether it grows horizontally tat. Investigations take account of the (creeps) or vertically (climbs), whether it is physical environment to the extent that tussocky or clumped), as well as its competi- physical factors such as climate, geology, or tive efficiency, all impose pattern. 6 Clearly soil chemistry modify such species interac- the scales at which these attributes are tions. Ecology is a quantitative science in so influential differ so that at large scales pattern far as it is concerned to analyse and evaluate may be the result of gross environmental * A glossary and list of plant names is appended to this article. •~Waller, op cir. 41~ Greig-Smith, 'Pattern in Vegetation',J0ur qfEcolq~),, 67, 3, 1979; ' E Estyn Evans, The Personalit), qflreland: Habitat, Heritm;'e, History, Quantitative Plant Ecolml),, 1983. 1972. ~Grdg-Smith, op cit. :R Waller, Prophet of the New Age: the L(fe ami Times qf Sir George ~'K A Kershaw, Quantitative and Dynamic Ecology, 1973; J L Harper, Stapledon, z972. Population Biology of Plants, 1978. 63 64 THE AGRICULTURAL HISTORY REVIEW factors whilst at the smallest scale it may be porating a divisive polythetic classification explained by spatial exclusion. Plant as a first stage which allows additional sites to 141 assemblages may often provide an index of be introduced into the analysis without environment and it was observations of requiring a totally new computation. 9 :!i: species sociology at any type of site which A study by Basset of pastures in the advanced the techniques of classification. 7 Camargue provides an elegant example of Essentially the species composition at any how the technique may be used and inte- one site is recorded and analysed so as to grated into a most effective analysis of derive a form of 'information' statistic by habitat. ~o He was able to identify differences noting presence and absence between sites. in habitat which could be explained by Such a process may suggest particular species differences of management. Similar studies as 'environmental indicators'. For reasons of of upland environments by Bunce et al, and computer time a divisive classification is by Hill and Jones, ~I in which the concern was preferred, whilst for ecological reasons an to identify the appropriate management assemblage of species (polythesis) is an strategies for economic and conservation advantage. Classification according to a purposes could point to variations in habitat divisive polythetic scheme has permitted which were explicable in terms of past ecologists to develop a diagnostic view of utilization and were documented in maps species (aut-ecology) or of an assembly of and estate documents. A similar study of species (synecology) as well as to identify the limestone upland in the south Pennines, environmental parameters operating. using a different technique, revealed a Investigation of pattern at large scales pattern caused by human activity. 1"- In all required that a reliable means should be cases cited above the vegetative analysis has found in order to place a species or a site in a been complemented by extant documents graded series (ordinated). To do this, Prin- which suggests the potential of such an cipal Components Analysis (using P. C. A. as approach for historical purposes. a generic term for a whole range of similar One of the quantitative techniques used to techniques, which is not strictly correct) has investigate pattern at smaller scales is 'Pat- been employed, and the information content tern Analysis', in which species are recorded is calculated in multi-dimensional space in by contact along a transect and the data which each e!ement is computed orthogo- processed by a nested block analysis of nally to the preceding element. It is a variance. The data is then plotted as a graph technique which has been of considerable use of mean square or covariance against block since it permits the components of variation size which indicates both the scale of pattern to be identified. There are many similar as well as the intensity according to the size of techniques and computer programmes the peak. A most appropriate example of available according to requirement but a such a technique is provided by Chadwick's scheme which uses a process called by its study ofNardus stricta (Mat Grass) on upland author 'reciprocal averaging' is widely used. in North Wales. 13 Here a pattern was This method allows the components of discerned which could not be explained by variation between sites and species to be VM O Hill, R G H Bunce and M W Shaw, 'Indicator Species mutually informative simultaneously. 8 A Analysis', Jour of Ecology, 63, 2, 1975, p 597. useful extension of this method, known as ,op Bassett, 'The Vegetation of a Camargue Pasture', J0ur of Ecology, 66, 3, 1978, p 807. Indicator Species Analysis, has enhanced the " Hill, Bunce and Shaw, op cit; M O Hill and E WJones, 'Changes flexibility of reciprocal averaging by incor- in a Welsh Forest',Jour of Ecolog),, 66, 2, 1978, p 427 . ': D J Anderson, 'Studies on structure in plant communities I. An analysis of limestone grassland in Monks Dale, Derbyshire', Jour ~J Braun-Blanquet, Plant Sociolocy, New York, 1932. qfEcolot.,y, 53, pp 97-1o7. 'M O Hill, 'Reciprocal Averaging. An eigen vector method',Jour qf ,3 M Chadwick, Studies ofNardus striaa in North Wah's, 1959, unpubl Ecolq¢y, 61, I, 1973, p 273. PhD thesis, Univ Coil Bangor. !?: I: ?,;!! ",! UPLAND COUNTRY: ECOLOGY AND HABITAT 65 environmental variation or independent One of the central concerns of ecology is species characteristics. When additional evi- 'climax'. The concept, grossly simplified, dence from economic and social history was postulates that the interaction of species, introduced an hypothesis could be advanced habitat, and environment is directional so to explain the pattern. A change in the meat that at any one stage of development there are consumption pattern consequent upon the typical assemblages which indicate the state First World War resulted in smaller joints. of a community towards climax. That is to Where hitherto upland and hill flocks were say that an assemblage of plants on recently composed ofwethers, smallerjoints resulted colonized ground will create the conditions in a switch to 'fat lamb', which meant that for different plants to flourish. Each assemb- grazing was even more selective with a lage succeeds its predecessor according to the consequent neglect of N. stricta. When the rate of change created by the competitive rate of cloning by rhizomes was calculated interactions of all species present. The the scale of the pattern coincided with the implications of'stability' are that the compo- process of propagation of Nardus since the nents of a stable habitat are in a state of switch in the composition of flocks. A study competitive equilibrium which militates of less relevance to upland studies is never- against substantial further change. The point theless interesting. Harper and Sagar studied of equilibrium may be designated a the distribution of Ranunculus spp (butter- 'climax'. I7 Clearly the more quantitative cups, in this instance) along a transect of data which becomes available from studies of ridge and furrow in grassland of the English any stage or process the better are the chances midlands. ,4 The three species of buttercup of constructing a mathematical model by showed preference for different types of site which future and past states may be extrapo- on the ridge and furrow so that R.
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