PROJECT OUTLINE FOR A DOCTORAL DEGREE IN ECOLOGY. by Thabisisani Ndhlovu 3488 Mkoba 16 Gweru Zimbabwe Email: Thabisisani@yahoo.co.

uk

Title:

ENVIRONMENTAL CORRELATES OF MONTANE TREE SPECIES DISTRIBUTION IN THE NGANDA-DOMWE RAIN-SHADOW AREA OF THE NYIKA PLATEAU, MALAWI
Table of contents
Title: ...........................................................................................................................................................- 1 Table of contents ........................................................................................................................................- 1 1. INTRODUCTION..................................................................................................................................- 2 1.1 Background.......................................................................................................................................- 2 1.2 Objectives.........................................................................................................................................- 2 1.3 Research Questions...........................................................................................................................- 3 1.4 Study area .........................................................................................................................................- 3 2 MATERIALS, METHODS AND DATA ANALYSIS ...........................................................................- 5 2.1 Sampling design................................................................................................................................- 5 2.1.1 Entitation ...................................................................................................................................- 5 2.1.2 Selection of representative forest patches..................................................................................- 6 2.1.3 Sampling plot placement ...........................................................................................................- 6 2.2 Species measurements ......................................................................................................................- 6 2.3 Environmental measurements ...........................................................................................................- 7 2.3.2 Soil sampling .............................................................................................................................- 8 2.4.1 Floristic and environmental datasets..........................................................................................- 8 2.4.2 Classification of the tree species into compositional units.........................................................- 8 2.4.3 Determination of important patterns of tree species distribution ...............................................- 9 2.2.4 Determination and characterisation of correlation between major tree species compositional and distributional patterns and environmental variation..........................................................................- 10 3. References ............................................................................................................................................- 11 -

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1. INTRODUCTION 1.1 Background The Nyika Plateau is an extensive 1320 square kilometre high altitude plateau occurring between 2000m and 2600m atop a massive block of granite and granitic gneiss forming the western margin of the East African Rift Valley system (Meadows 1984; Meadows & Linder 1993). It sits astride the Malawi/Zambia border to the northwest of Lake Malawi with most of it in Malawi except a small seventy square kilometre western section (Dowsett-Lemaire 1985). The Plateau is covered by the largest montane complex in south central Africa and consequently represents an environmental resource of considerable value. However, although the montane zone of the Nyika plateau has been botanised intensively (DowsettLemaire 1985; Willis et al. 2001) not much work has been done on its fundamental autecology and synecology. Dowsett-Lemaire (1985) provided detailed information on the composition, structure, phenology, distribution and extent of montane forest vegetation but did not provide corresponding environmental information. As a result there is limited knowledge about montane forest vegetation-environment relationships. This study proposes to quantitatively discover and describe the major pattern of variation in montane tree species distribution in a high-altitude and low-rainfall area of the Nyika Plateau and interpret it in terms of underlying environmental variation. It is hoped this research will provide inductive insight into montane forest vegetationenvironment relationships on the plateau.

1.2 Objectives The objectives of this study are: 1. to identify and characterise the major units of tree species composition in the study area. 2. to identify and describe the major gradients of tree species compositional change in the study area. 3. to seek and describe correlations between the discovered tree species distributional pattern and the variation in a set of environmental factors.

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4. to quantify the importance of each environmental variable, as well as groups of topographical (slope angle, aspect & terrain form), edaphic (soil depth, insitu soil moisture, drainage, soil texture & soil nutrients), climatic (elevation), biological competition (canopy cover)and disturbance (fire intensity & severity) variables in explaining the species distributional patterns of montane trees in the study area. 5. to utilise the resultant descriptive model of vegetation-environment relations to provide an ecologically based explanation of montane tree species distribution in the study area.

1.3 Research Questions The following questions will be addressed in the study: 1. Do montane trees in the study area occur together in repeating groups of associated species? 2. What are the compositional characteristics of these species associations? 3. What environmental conditions are related to the identified species associations? 4. What are the important patterns of variation in montane tree species distribution in the study area? 5. Are these distributional patterns related to any variation in the selected environmental variables? 6. How significant is/are this/these relationship(s)? 7. What are the characteristics and features of this/these relationships?

1.4 Study area The study will be conducted in the northern part of the Nyika plateau located south of Mondwe valley and lying in the shadow of the high peaks of Nganda and Domwe. It will cover an area of 84.339 square kilometres and extend 10.726 kilometres south to north (10o24`-10o29`S) and 7.863 kilometres east to west (33o44`-33o49`).

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The climate is temperate with direct and diffuse radiation approximating 450 cal/cm2 per day - comparable with northern England and the extreme north-eastern United States (Lemon 1968). Temperatures range from below 0C in winter (June/July) to over 21C in summer (October/November) giving mean annual temperatures of the order of 13-15oC(Lemon 1968). There is a low annual average of frost - one night in May, two in June, three in July and one in August (Dowsett-Lemaire 1985). As a result of its location in the rain-shadow of the high peaks of Nganda and Domwe, the area experiences the lowest rainfall on the plateau (Dowsett-Lemaire 1985). Ninety percent of this rain usually averaging 898 to 988 mm per annum - falls from December through April, and is followed by a dry period of about six to seven months (Lemon 1968; Dowsett-Lemaire 1989). Soil moisture storage, however, is very good such that the streams run all year (Lemon 1968). The seasonal distribution and abundance of rainfall varies greatly from year to year (Dowsett-Lemaire 1989). The area is underlain by the intermediate metamorphic Chambo gneiss on which have developed deep well drained, reddish brown to red, dominantly fine textured soils of low chemical fertility. The soils fall within the dystric-fe group of FAOs Haplic Acriso soil class. Montane forest, constituting less than 5 percent of the landscape, occurs in the study area as widely scattered patches (none larger than 8 ha) with low and broken up canopies (dominated by Hagenia and Myrica), few large woody creepers, covering a densely herbaceous ground (Dowsett-Lemaire 1989). This patchy distribution is commonly attributed to the high frequency and extent of range fires on the exposed drier high plateau (Dowsett-Lemaire 1985). These range fires are thought to have, over time, eroded the margins of once widespread tracts of montane forests to produce the extant mosaic of small sharply defined patches confined to fire-sheltered depressions on valley sides and watercourses (Dowsett-Lemaire 1989). Above 2250 m the edges of the forest patches are usually sharply cut and secondary growth is never extensive (Dowsett-Lemaire 1985). Secondary growth gradually forms larger patches around the forest patches with decrease in altitude until at 2000-2100 m it covers vast areas of many hectares (Dowsett-Lemaire 1985).

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2 MATERIALS, METHODS AND DATA ANALYSIS 2.1 Sampling design 1. The sampling procedure is designed to optimise the amount of information gained in proportion to the time and effort to be spent during the vegetation survey. Care will be taken to ensure that the data sets are representative of the full range of variability in montane tree species distributional patterns. This will be achieved through a structured random design that will include every forest patch in the study area. 2. Individual forest patches will be regarded as the basic vegetation segments within which the sampling of tree species will be carried out. More detailed entitation is precluded by the small size of most forest patches in the study area which would yield vegetation segments that cannot be suitably delineated using 1: 250 000 aerial photographs. The entitation process is necessitated by the absence of spatially explicit information (e.g. maps, GIS layers) on montane forest patches in the study area.

2.1.1 Entitation 1. Air photo processing and image stratification will be carried out using the `Integrated Land and Watershed Management Information System` (ILWIS) package. ILWIS is a geographical information systems (GIS) software package developed by the International Institute for Aerospace Survey and Earth Sciences (ITC) (Valenzuela 1988). 2. Aerial photos covering the study area will be scanned, georeferenced, georectified and stitched to form a continuous mosaic. 3. This digital air photo mosaic will be interpreted using patterns of tone, texture, and contrast to identify montane forest patches. 4. The outlines of the montane forest patches will be digitised onscreen and stored in the GIS database as a theme. Included in this database will be data on location

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(centre coordinates), size, altitude, aspect and area of secondary growth for every forest patch in the study area. 2.1.2 Selection of representative forest patches 1. The continuous data on forest patch size, altitude, aspect and area of secondary growth will be categorised. 2. Cluster analysis will be used to place all the montane forest patches in the study area into a reasonable number of groups using categorized data on size, altitude, aspect and area of secondary growth. 3. Representative forest patches will be chosen randomly from each class such their combined area will be above 15% of the total area of montane forest patches in the class.

2.1.3 Sampling plot placement. 1. Fieldwork will be restricted to the selected representative forest patches. 2. Two parallel transects will be subjectively selected to cover the variation in each selected forest patch. 3. The lengths of these transects will vary with patch size. 4. Potential sample plots of 20m*10m will be located at 20 metre intervals along each transect.

2.2 Species measurements 1. Every woody plant rooted within the sample plot with basal stem diameter >/= 4cm and >/= 1.5m in height will be included in the sample. 2. The selected plants will be taxonomically identified to species level. 3. The stem diameter will be measured above the basal whorl. 4. The height of each plant will be estimated using a ranging rod. 5. Each plant will be classed as to being an (1) emergent (2) canopy (3) under-story tree. 6. Each plant will be examined for fire damage.

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2.3 Environmental measurements Table 1 Environmental variables collected in each sampling plot Variable Slope angle Elevation Aspect Terrain form Comments

determined from DEM determined from DEM determined from DEM 6-point scale: 0 - valley bottom 1 = concave valleyside, 2 = plane valleyside, 3 = convex valleyside, 4 = ridge, 5 = hilltop Soil depth 4-point scale based on observation of the surface relief in plot.1 = extensive rock outcrops, 2 = local rock outcrops, 3 = no rock outcrops, terrain uneven, 4 = no rock outcrops, even surface. Insitu soil volumetric soil moisture determined by weighing fresh samples, air moisture drying them, and reweighing. Sampling will be done on the same day for each group of close lying sites in order to ensure that moisture differences between samples are meaningful. Drainage assessed visually on a scale 1 (permanently waterlogged) to 5 (excessively drained) for each plot Soil texture determined from percent fractions of sand, silt and loam. Soil nutrients Organic content, total soluble salts (TSS in mS cm-1), pH, concentration of exchangeable cations (Ca, Mg, Na, K, P, N), concentration of micronutrients (Fe, Zn, Cu, Mn). determined from visual estimation of percent canopy cover determined from evidence of fire front within the forest profile (modal height of fire scars in plot) 3-point scale, based on observations of fire induced mortality: 1. Low severity = light fires that have minimal impacts on forest overstories, but may kill small shrubs. 2. Moderate severity = partial stand replacement fires that include areas of both low and high severity. Some overstory trees are killed or heavily damaged in high severity patches. 3. high severity = most trees, including overstory trees, are killed

Canopy cover Fire intensity Fire severity

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2.3.2 Soil sampling 1. In each sample plot five soil sub samples from profiles 0-30 cm below the organic material layer will be collected from the four corners and centre. This is within the rooting depth of most trees. 2. The five sub samples will be pooled together by thorough mixing to form a composite soil sample. During the mixing visible undecomposed organic material will be removed and discarded. 3. After mixing, the composite soil samples will be immediately air dried by being spread out on a warm, dry area for five days after which they will be bagged.

2.4 Data analysis 2.4.1 Floristic and environmental datasets Two datasets will be developed: 1) The floristic dataset (species*sampling plot) containing data on the abundance of montane tree species in a series of sampling plots. The relative basal area per sampling plot for each tree species will be used as the abundance value. 2) The environmental dataset (environmental variable*sampling plot) containing data on a number of environmental variables measured at the same sampling plots. See Table 1 for the list of environmental variables to be measured. Environmental variables will be log-transformed in order to approximate normal or random (uniform) distributions.

2.4.2 Classification of the tree species into compositional units 1. The floristic dataset will be analysed using two-way indicator species analysis (TWINSPAN) to produce a classification of tree species associations in the study area. TWINSPAN is a divisive hierarchical programme that uses indicator species to characterize and separate species association classes

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(Okland 1990). This clustering exercise is a data reduction process intended to simplify the interpretation of the complex species dataset. The simplification is achieved by summarising the species distribution patterns of the entire species complement to those of groups of co-occurring tree species. 2. The non-parametric Mann-Whitney test (Zar 1996) will be used to test whether there are significant variations between the means of all environmental factors for the two groups separated at each split in the classification.

2.4.3 Determination of important patterns of tree species distribution 1. The important patterns of variation in tree species composition and distribution in the study area will be extracted by one of two ordination techniques, Principal Component Analysis (PCA) or Detrended

Correspondence Analysis (DCA) (Jongman et. al 1987). 2. PCA and DCA are indirect gradient analysis techniques that detect dominant compositional gradients from species*plots matrices of species abundance values independently of site variables. These techniques provide an expression of pure compositional gradients. 3. PCA will be applied if the distribution of tree species along the dominant floristic gradient is linear. DCA, on the other hand will be applied if tree species are unimodally distributed along this gradient. 4. Preliminary multivariate analysis using DCA will be used to determine whether the tree species are unimodally or linearly distributed along the dominant floristic gradient. DCA provides a non-linear rescaling of the ordination axes in units of mean standard deviation of species turnover (SD units) (Odd et. al 1990). 5. A gradient length of >/= 4SD units will represent unimodal distribution and < 4SD units, linear distribution. 4SD units indicate a nearly complete turnover of species composition along a gradient.

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2.2.4 Determination and characterisation of correlation between major tree species compositional and distributional patterns and environmental variation 1. The fraction of variation in species composition and distribution explained by single environmental variables will be assessed by hybrid CCA, using each environmental variable in turn as the only constraining variable (Borcard et. al 1992). Randomly generated dummy environmental variables will keep the number of environmental variables constant during each separate run. 2. Variation explained will be the ratio of the eigenvalue of the first constrained CCA axis and the total inertia (total variation; the sum of all unconstrained eigenvalues of DCA) (Borcard et. al 1992). 3. The hypothesis of non-significant deviation of variation explained by a variable from that explained by a random variable will be tested with the Monte Carlo test (99 unrestricted permutations of the constraining variable)(ter Braak 1990). 4. Comparison between the DCA output and the outputs of the significant CCA runs will be carried out to detect and characterise relations between DCA axes and CCA axes. Those measured environmental variables which relate strongly to the first few DCA ordination axes will be taken as accounting for the main part of the variation in the species distribution. Environmental variables that do not relate strongly to the first few DCA axes will be considered as not accounting for the main part of the variation. They will, however, also be considered as they may still account for some of the remaining variation which can be substantial. 5. A measure of the -diversity associated with a particular explanatory variable will be derived from the gradient length of the first CCA axis (with one constraining variable, using non-linear rescaling in SD units) (Odd et. al 1990).

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6. The variation explained by different sets of environmental variables will be added up and thus partitioned (Borcard et. al 1992).

3. References Borcard, D., Legendre, P. & Drapeau, P. 1992. Partialling out the spatial component of ecological variation. Ecology, 73(3):1045-1055.

Dowsett-Lemaire, F. 1985. The forest vegetation of the Nyika Plateau (Malawi-Zambia): ecological and phenological studies. Bull. Nat. Plantentuin Belg. 55: 301-392.

Dowsett-Lemaire, F. 1989. The flora and phytogeography of the evergreen forests of Malawi I: Afromontane and mid-altitude forests. Bull. Nat. Plantentuin Belg. 59: 3-131.

Jongman, R.H., ter Braak, C.J.F. & van Tongeren, O.F.G. 1987. Data analysis in community and landscape ecology. Pudoc, Wageningen.

Lemon, P.C. 1968. Effects of fire on African plateau grassland. Ecology, 49, 316-322.

Meadows, M.E. & Linder, H.P. 1993. A palaeoecological perspective on the origins of Afromontane grasslands. Journal of Biogeography, 20 (4), 345-355.

Meadows, M.E. 1984. Late Quaternary vegetation history of the Nyika Plateau, Malawi. Journal of Biogeography, 11(3): 209-222.

Odd, E., Okland, R.H., Okland, T. & Pedersen, O. 1990. Data manipulation and gradient length estimation in DCA ordination. Journal of Vegetation Science 1: 261-270.

Okland, R.H. 1990. Vegetation ecology: theory, methods and applications with reference to Fennoscandia. Sommerfeltia supplement, 1: 1-216.
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Ter Braak, C.J.F. 1990. Update notes: Canoco, version 3.1. Agricultural Mathematics Group. Wageningen.

Valenzuela, C.R. 1988. ILWIS overview. ITC Journal 1988-1: 3-14.

Willis, C.K., Burrows, J.E., Fish, L., Phiri, P.S.M., Chikuni, A.C. & Golding, J. 2001. Developing a greater understanding of the flora of the Nyika. Syst. Geogr. Pl. 71:9931008.

Zar, J.H. 1996. Biostatistical analysis. Prentice Hall, New Jersey.

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