Modelling the Distribution of Cecropia Species Using GIS based Techniques

Implications for Tropical Forest Conservation Koulla Pallaris September 1998

This dissertation is submitted as a part of an MSc degree in Geography at Kings College London

Resumen El objetivo de este estudio fue probar y evaluar el xito de modelar la distribucin potencial de la especies Cecropia sp., basado en variables biofsicas mediante el uso de sistemas de informacin geogrfica SIG. Se utliz el modelo DOMAIN, desarrollado por Carpenter et al (1990), para predecir la distribucin de Cecropia, mientras que otras aplicaciones SIG se usaron para modelar y hacer mapas de las variables biofsicas con las cuales DOMAIN procesara los datos. El modelar y mapear distribuciones de

especies tiene implicaciones significtivas como herramientas de conservacin de la biodiversidad. En el pasado, las encuestas florsticas solamente se proponan

documentar la distribucin de las especies, pero stas no siempre se georeferenciaban en forma correcta. Las ventajas de georeferenciar y mapear los patrones de

distribucin de las especies, permite identificar importantes lugares ecologicos y por consiguiente, ayuda en la implementacin de estrategias administrativas apropiadas. Adems, puede ser muy importante para predecir posibles migraciones o invasin de especies, a niveles diferents, en respuesta a futuros cambios en el clima. La aplicacin de tales tcnicas se ha concentrado mayormente en identificar distribucin de animales, con base en la determinacin del hbitat mediante identificacin de comunidades vegetales con mayor probabilidad de ser habitades. Este modela la distribucin potencial de Cecropia sp. identificando reas de condiciones ambientales similares a aquellas en las cuales se encuentra la especie.

Acknowledgements
I would like to thank Jorge Rubiano, Mauricio Rincon, Matthew Letts for all their help in the field. I am grateful to Mauricio Rincon for supplying the original maps from which subsequent analysis was derived. Thank you also to everyone at CIAT, for the use of their facilities while writing up this dissertation. Finally, I would especially like to thank my supervisor, Dr Mark Mulligan for all his help, guidance and support.

I would like to dedicate this work to the memory of Dr Alvaro Jose Negret, who took the time to introduce me to and supply me with considerable information on the species that I have modelled.

CONTENTS

SECTION 1: INTRODUCTION.......................................................................1-22 CHAPTER 1..........................................................................................................1-7 1.1 Abstract.......................................................................................................1 1.2 The Study Area ............................................................................................2 CHAPTER 2........................................................................................................8-11 2.1 The Threat to Biodiversity ............................................................................8 2.1.2 Climate Change: Implications on Future Species Distributions. ..............12 CHAPTER 3......................................................................................................12-22 3.1. Environmental Constraints to Species Survival..........................................12 3.1.1 Climate as a Determinant of Species Distribution....................................13 3.1.2 Environmental Complexity and Species Interactions as Determinants of their Distribution..............................................................................................15 3.3 Factors Affecting the Distribution of Cecropia ...........................................16 3.4 GIS Approaches for Modelling Potential Species Distributions...................20 SECTION 2: SAMPLING AND METHODOLOGIES .................................23-35 CHAPTER 4......................................................................................................23-35 4.1 Defining a Sampling Strategy .....................................................................23 4.2 Determining Biophysical Variables for Cecropia........................................29 4.3 Locating Cecropia in the Field ...................................................................30 4.4 Field Validation of Modelled Climatic Variables ........................................32 SECTION 3: MODELLING CECROPIA DISTRIBUTION .........................36-43 CHAPTER 5......................................................................................................36-43 5.1 Modelling the Biophysical Variables Important for Cecropia Distribution..36 5.2 Modelling Plant Distribution using Domain Model ....................................40 SECTION 4: MODEL RESULTS AND DISCUSSION.................................44-58 CHAPTER 6......................................................................................................44-58 6.1 DOMAIN Output .......................................................................................44 6.2 Validation of Model Results........................................................................47

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6.3 Environmental limits of Cecropia ..............................................................50 6.4 Sensitivity Analysis of DOMAIN .................................................................54 6.5 Discussion of Results..................................................................................56 6.6 Conclusions................................................................................................58 REFERENCES ................................................................................................59-62 BOOKS................................................................................................................ 61 INTERNET RESOURCES ........................................................................................... 62 SOFTWARE ............................................................................................................ 62

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LIST OF TABLES
TABLE 4.1.1. CLASSES USED IN CLASSIFYING THE CATCHMENT. .............................. 24 TABLE 4.3.1. PRESENCE-ABSENCE CLASSIFICATION ................................................ 31

LIST OF FIGURES
FIGURE 1.2.1. THE LOCATION OF STUDY SITE AND BIOPHYSICAL REGIONS OF COLOMBIA .......................................................................................................................... 5 FIGURE 1.2.2. MONTHLY RAINFALL FOR TAMBITO. ................................................... 6 FIGURE 1.2. 3. AIR TEMPERATURE AND RELATIVE HUMIDITY. .................................. 6 FIGURE 4.1.1. CLASSIFIED SAMPLING MAP. ............................................................ 26 FIGURE 4.1.2. LOCATION OF TRANSECTS SHOWN ON A HILLSHADE TERRAIN MAP. ..... 27 FIGURE 4.1.3. PRIMARY FOREST TRANSECT ............................................................. 28 FIGURE 4.1.4. PALO VERDE TRANSECT .................................................................... 28 FIGURE 4.1.5. EXIT PATH TRANSECT ....................................................................... 28 FIGURE 4.1.7. ASOMADERO TRANSECT .................................................................... 29 FIGURE 4.1.6. PALM FOREST TRANSECT. ................................................................. 29 FIGURE 4.1.7. RIVER STATION TRANSECT ................................................................ 29 FIGURE 4.4.1. MEASURED MIN AND MAX TEMPERATURE ........................................ 33 FIGURE 4.4.2. THROUGHFALL TROUGH TO MEASURE THROUGHFALL. ...................... 35 FIGURE 4.4.3. MEASURED THROUGHFALL OVER AN ALTITUDINAL GRADIENT .......... 36 FIGURE 5.1.4. REGRESSION BETWEEN MEAN ANNUAL MAXIMUM TEMPERATURE AND
ALTITUDE ......................................................................................................... 38

FIGURE 5.1.5. REGRESSION BETWEEN MEAN ANNUAL MINIMUM TEMPERATURE AND

ALTITUDE ......................................................................................................... 38

FIGURE 5.2.1. SCHEMATIC DIAGRAM ILLUSTRATING THE DOMAIN PROCEDURE ...... 41 FIGURE 6.1.1. MODELLED DOMAIN OUTPUT. SIMILARITY MEASURE ...................... 46 FIGURE 6.1.2. HISTOGRAM OF FREQUENCY DISTRIBUTION OF SIMILARITY AREAS..... 47 FIGURE 6.2.1. THE CONGRUENCE OF VALIDATION DATA TO MODEL OUTPUT. .......... 48 FIGURE 6.2.2. CONGRUENCE OF CECROPIA CELLS WITH DOMAIN OUTPUT .............. 49 FIGURE 6.3.1. MAXIMUM TEMPERATURE LIMITS OF CECROPIA ................................. 51 FIGURE 6.3.2. MINIMUM TEMPERATURE LIMITS OF CECROPIA .................................. 52 FIGURE 6.4.1. DOMAIN OUTPUT, WITH 3 DATA LAYERS ......................................... 55 FIGURE 6.4.2. DOMAIN OUTPUT WITH ONE DATA LAYER ........................................ 55

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LIST OF PLATES
PLATE 1. DISTURBED REGION OF TAMBITO CATCHMENT............................................ 7 PLATE 2. CECROPIA TREE ON AN EXPOSED RIVER BANK............................................ 17 PLATE 3. CECROPIA LEAF ATTACKED BY HERBIVORY................................................ 18

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Section 1: Introduction
Chapter 1 1.1 Abstract
The objective of this study was to try and evaluate the success of modelling the potential distribution of a plant species, (Cecropia), based on simple biophysical variables through GIS techniques. Principally the DOMAIN model, developed by Carpenter et al (1990) was applied to predict the distribution of Cecropia, while other GIS packages were used for modelling and displaying the biophysical variables used by DOMAIN to compute the potential distribution and for carrying out validation analysis. Modelling and Mapping species distributions has significant implications as a tool in biodiversity conservation. In the past, floristic surveys have endeavoured to document species distribution but these were not always accurately georeferenced. Georeferencing and mapping the distribution patterns of species enables important ecological sights to be identified. Furthermore, it can facilitate the application of appropriate conservation and management decisions. It can also be used for predicting possible migration or invasion of species in response to future climate change. The application of such techniques has mainly concentrated on identifying animal distributions, based on habitat determination. This study takes a tropical plant species and models its potential distribution based on identifying areas similar in environmental conditions to those in which the species is found. GIS is also used to identify and analyse the environmental tolerance limits of Cecropia The success of the approach is discussed. This in part, will be determined by how well the model performs at predicting the distribution of Cecropia. The choice of biophysical variables used for the analysis is also evaluated.

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1.2 The Study Area

The study was carried out in a Colombian Tropical Montane Cloud Forest (TMCF), in the nature reserve of Tambito Cauca watershed (3000 ha). Colombia is characterised by five main biogeographical regions, which comprise of lowland tropical forests, 43%, Tropical montane cloud forests, 3.0%, lowland savannahs (llanos), 2.9% and mangroves, 0.4% (Place, 1993). Tambito is situated in the topographical region of the Pacific coast (figure 1.2.1). This region is the most variable in terms of vegetational community types. The coastal regions are characterised by mangrove forests, while further inland a gradient with altitude results in a cover of lowland moist to montane forests on the slopes of the Cordillera Occidental mountain range.

More specifically, the vegetation of Tambito is classified as Subandian, with dominant vegetation families consisting of Palmae, Lauraceas, Sapotaceas, Moraceas, Rubiaceas and Flacourtiaces, (Negret, 1996a). A high diversity of invertebrate faunal species such as butterflies and insects have been recorded as well as a significant number of vertebrates, for example, 411 species of birds, 46 species of reptiles and 51 mammal species. This makes Tambito an important ecological site for carrying out studies of the kind undertaken here.

Tambito is characterised by altitudes of between 1300 m to about 2700 m (Negret 1996a). A mean annual rainfall of 4000 mm (figure 1.2.2) means that the atmospheric environment is always very humid (> 60%) (figure 1.2.3). Seasonality in rainfall is observed, with the driest months between June and September. Diurnal temperatures oscillate between 13 C - 26C (figure 1.2.3). Persistent and frequent

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cloud cover at or near the vegetation canopy (plate 1) reduces the solar radiation received by the vegetation and the high humidity results in a general suppression of evapotranspiration.

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CARIBBEAN SEA

PACIFIC SEA

Tambito - Cauca

Pacific Region Caribbean Region Amazon Region Orinoco Region Andean Region

Figure 1.2.1. The Location of Study Site in Relation to the Biophysical Regions of Colombia

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Monthly Rainfall for Tambito - 1996 & 1997

800 700 600 Rainfall (mm) 500 400 300 200 100 0
May Aug Sep Nov Feb Mar Jan Dec Jun Apr Jul Oct

1996 1997

Month

Figure 1.2.2. Monthly Rainfall for Tambito. (Source: Weather station at Tambito).

Hourly Air Temperature and Relative Humidity from 28/07/98 - 07/08/98 30 25 Temperature (C) 20 60 15 40 Air Temperature (C) 10 5 11:00 00:00 13:00 02:00 15:00 04:00 17:00 06:00 19:00 08:00 21:00 10:00 23:00 12:00 01:00 14:00 03:00 16:00 05:00 Relative Humidity (%) 20 0 120 100 Relative Humidity (%) 80

Hours

Figure 1.2. 3. Air Temperature and Relative Humidity. (Source: Cleared Plot Weather Station.)

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Detail on the geology and soils of the area are not known, but the literature suggests that much of the Andes consist mainly of Red-Yellow coloured Latosols, of volcanic origin (Fittkau et al, 1993). Three broad land cover types can be found in Tambito (figure 1.2.4). Primary forest covers most of the area (61.8%) while Secondary forests cover a less, but significant area (35.9%). Two main types of Secondary forests were observed; mixed evergreen and palm forests (Plate 1). The Secondary forest is the result of abandoned cultivated farmland and remnants of the Lulo crop can still be found there. A small area of pasture remains (2.3 %) which is maintained by grazing animals.

Primary Forest Secondary Forest Pasture Land

Figure 1.2.4. Vegetation Zones of Tambito. (Source: Reclassified image of Mapa de Cobertura Vegetal, Negret 1996)

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Persistent Cloud Over Canopy

Palm Forest

Pasture Area

Plate 1. Disturbed Region of the Tambito Catchment.

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Chapter 2 2.1 The Threat to Biodiversity

All species, whether presently endangered or not are deemed to be under threat as a result of modifications to their environment. Indeed, such changes need not be drastic man-induced alterations in land use, as often perceived, but may be the result of evolutionary processes (Drake et al, 1996). Natural changes, observed in the geological time scale, have shown how climates and geological events have resulted in genetic change and mass extinctions in the past. Even intervention by man is not a new phenomenon. Evidence suggests that man has been present in the humid tropics from about 5000 years BP (Fittkau et al, 1993).

However, the alarming rates of tropical forest deforestation in recent decades have escalated concern over the preservation of species diversity. Myers (1983, in Whitmore and Sayer, 1991) expresses that 25-30% of all plant and animal species are likely to become extinct by AD 2000. Whether exaggerated or not, it is true to say that the threat to the survival of biodiversity as a result of land use change and/or climate change is immense.

At present our knowledge on the distribution of endangered species is limited, often due to their occurrence in isolated patches in the landscape. If their distribution can be modelled successfully, based only on the unique biophysical conditions in which they are found, then the task of surveying their occurrence in the field can be dramatically reduced. Such techniques, as adopted here, can expand our knowledge on their location and highlight whether conservation measures need to be taken. 8

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2.1.1 Deforestation: Its Implications on Biodiversity Loss

Colombias complex geoclimatic and ecological history has meant that it has evolved into one the most species rich areas of the Neotropical regions, with the richest area concentrated in the Choc region of the Pacific cost. It has been estimated to have 50,000 species (Gentry 1978), making up 10% of the worlds biota, (Hernndez and Snchez, 1988).

A significant threat to this species richness in Colombia is agricultural expansion, which remains an important economic base. Colombia remains the worlds second largest coffee exporter after Brazil. In addition, beans, rice, and cassava remain important national food crops. Further, forests have been lost to large commercial plantations as well as for the production of illicit crops, (Cavelier and Etter, 1995).

With constant pressure to bring more land into production, shifting cultivation and human settlements are responsible for 76.3 % of deforestation, where increasingly cultivated land is spreading to the upper, steeper forested slopes of the Andes (Fittkau et al, 1993). Estimation of annual deforestation rates in Colombia range from 1.8 % (FAO 1981-1988, reported in Whitmore and Sayer, 1991) to 2.3 % (Myers 1989, Whitmore and Sayer, 1991), while current rates are reported at 600,000 ha per annum (Cavelier, 1995). This has reduced the TMCF extent to 27% (Cavelier, 1995).

Often, post assessment of the impacts of deforestation on species diversity is made difficult if the initial conditions and composition of species are not known. Predicting

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biodiversity change and loss is further complicated by successional dynamics following disturbance. These are related to resilience, a measure of how quickly an area will return to equilibrium and elasticity, a measure of the degree to which an environment will return to pre-disturbed conditions following perturbation (Collins and Benning, 1996). In addition, individual species within a community adapt to new conditions in divergent ways.

Thus, if the distribution of endangered species is known then conservatory measures can be taken to avoid their loss. Secondly, if disturbance, natural or man induced did occur, a more accurate, quantified assessment of biodiversity loss could be made instead of estimates which express nothing more than alarmist rhetoric.

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2.1.2 Climate Change: Implications on Future Species Distributions.

Climate change has occurred throughout geological history. Studies documenting the Quaternary history of Tropical America reveal that alternate phases of dry and more humid climates during the last two million years had resulted in vegetational shifts in the Tropics. The expansion of glaciation in the Northern Hemisphere during the Quaternary had especially affected the climates of the equatorial montane regions.

The climate in the Colombian Andes is believed to have been very dry during the peak of the last glaciation (20,000-13,000 BP). A reduction in temperatures during this period is presumed to have been as much as 11C at elevations above 2000 m. Regional variation of annual rainfall would also have varied throughout the continued uplift of the Andes in the Tertiary period. These climatic fluctuations had resulted in altitudinal movements of montane vegetation belts (Fittkau et al, 1993).

Concern over recent predictions in future climate change and its effects on species biodiversity has been expressed (Frankel et al, 1995). Endangered and endemic species are more likely to be within their environmental tolerance limits and therefore any shift in base conditions will more likely result in their extinction. To be able to predict the climatic displacement of species on both a global scale (where they cross latitudinal phytic zones) and local scale (crossing altitudinal zones) it is important to identify the unique climatic and biophysical conditions which demarcate their distribution.

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Chapter 3 3.1. Environmental Constraints to Species Survival

The environment sets the constraints within which organisms survive, develop and evolve. If their adaptive capability is inadequate to cope with change then their continued survival will be curbed (Gaston and Williams, 1996). The niche concept of Hutchinson, (1967) is often used to explain the spatial patterns of species distributions. The habitat niche, which comprises the physical, chemical and biotic elements that favour the plants vegetative development is namely used in this study to explain the association of a species to its environment in space.

Box (1981), states that despite the complexity of ecological systems and of plantenvironment relations, results suggest that the basic features of plant form and vegetation structure, in the absence of over riding non climatic complications, are determined primarily by the general levels and mean seasonal patterns of climatic variables. Therefore, even though other factors such as competition can affect species dispersal, a general assessment of their distribution can be derived from biophysical variables alone. One of the objectives of this study is to evaluate the veracity of this approach.

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3.1.1 Climate as a Determinant of Species Distribution

Attempts at classifying species distributions have concentrated mainly at the global scale, where arbitrary boundaries of vegetation types have been distinguished along global climatic zones. One of the earliest attempts illustrating this, is that of Holdridge (1967) where areas had been divided into phytogeographic regions. However, these are not immutably fixed, and historical records have shown how these can fluctuate greatly in time and space.

Modern approaches such as that of Box (1981, 1996) also maintain the link between macroclimatic variables and plant life form distributions. Box (1981) calculates the distribution of plant life forms by means of defining climatic envelopes from correlation between distribution limits and regional climate conditions. However, Woodward & Williams (1987) argue that these approaches only provide very broad explanations for species distributions and that it is necessary to establish the fundamental physiological basis for these climate-species relationships.

Climate control on the organisation of vegetation structure and composition can indeed be much more intricate than suggested by such broad scale studies. This has been manifested by the continuum theory (Austin and Gaywood, 1994) which emphasises how vegetation organises itself along smaller scale environmental gradients, for example, in response to altitudinal changes or aspect on a hillside.

At present the greatest difficulty that hampers such a probabilistic representation of species dispersal patterns is the lack of scientific understanding of the underlying factors that influence the occurrence of species, populations and individuals, (Miller,

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1994). This study attempts to simulate the environmental gradients as they change with altitude within the Tambito catchment.

The magnitude and frequency of climate variability over time is also important in affecting species composition and may differ in importance at different spatial scales and for different species within a community. Climate patterns change with season, from year to year and with climate phenomena such as the El Nio. Further, microclimate conditions can change dramatically over time due to the fast productivity of the TMCF ecosystem. For example, a gap can close rapidly due to the fast productivity of pioneer species that occupy it and therefore change, amongst other factors, the radiation received by the herb layer.

However, consideration of these short-term changes is only important if one is interested in the productivity of a species. The distribution of a species is influenced by those changes in the environment that occur over time intervals equal to or greater than the organisms response time (Miller, 1994). Therefore, for a tree species, such a Cecropia, it is adequate to consider only the mean, long-term changes, as these are the conditions that would have resulted in its long-standing presence within a community.

Mapping the abundance of a species in space may indicate whether it is within its environmental tolerance limits. A high abundance would suggest that present conditions are favourable to its development while conversely, a small number of very old individuals may mean that its niche has been invaded by other competing species. If such patterns are observed, it may be possible to assess the degree of environmental change and time period over which it has occurred.

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3.1.2 Environmental Complexity and Species Interactions as Determinants of their Distribution

TMCF are structurally very complex. This is attributed to the unique environment, which varies significantly with altitude. These altitudinal variations result in different vegetation association (Negret, 1996b). Their occurrence on a global scale is within a wide range of annual and seasonal rainfall regimes (500-10,000 mm/year). The atmospheric environment is characterised by persistent cloud, especially on the higher slopes. An important characteristic of TMCF is their ability to absorb moisture from the clouds (cloud stripping). At these higher altitudes a change in species composition and structure is observed. The density of species increases, with trees being smaller in stature having smaller, thicker leaves (Hamilton et al, 1995). Epiphyte biomass is also more abundant with increasing altitude.

In addition to the environmental heterogeneity other ecological concepts, related to competition theory, evolutionary theory and population dynamics attempt to explain spatial patterns of species occurrence. Often the potential niche of a species, that is the space it could potentially inhabit in the absence of competitors, is limited to a realisation niche in the presence of competitors and predators (Hutchinson, 1967). Temporal changes in species distribution can be explained by the adaptability potential of a species. A populations extinction rate is expected to be negatively correlated with its size, where small endemic populations are more likely to become extinct compared to large ones, (MacArthur and Wilson, 1967). In addition, a species best adapted to its environment will flourish and may consequently migrate away to less favourable areas (MacArther, 1984). These species are less likely to be displaced by shifting environmental conditions.

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3.3 Factors Affecting the Distribution of Cecropia

The knowledge presently available to us on the distribution and dynamics of biological diversity is extremely inadequate (Gadbil, 1996). Gathering this knowledge is an immense challenge. However, our ignorance on the distribution of biodiversity, can be alleviated to a certain extent if we can concentrate on methods to try and model their spatial occurrence. This can be done on two levels; by modelling the distribution of areas with high species richness, in order to direct conservation priorities in the general sense or target species on an individual basis. Focusing on the latter will help distinguish threatened species or species with a restricted geographical range.

It could be argued that such a species-based approach to managing all of the Earths biodiversity is not a practical proposition (Gadbil, 1996), while the former poses difficulties in accurately predicting geographical shifts given the complex interactions of individuals to their environment. However, if relatively simple, inexpensive methods can be used to achieve these objectives, practicality may no longer be an overriding issue impeding knowledge acquisition.

This study concentrates on a single tree species. The aim is to assess whether it is possible to accurately predict its geographical range. Given the time constraints, a tree species was chosen over herbaceous plants, as it was easier to identify. Further, the aim was to select a species with some significance in the area. Cecropia (plate 2) was selected after learning about its ecological importance (Negret, personal communication) and medicinal properties, (Fuentes et al, 1986). Cecropia is an important Neotropical genus, belonging to the Moraceas family and comprises 60

16

species altogether (Franco-Rosselli and Berg, 1997). The species of Cecropia believed to be found in Tambito are C. burriada, C. eximia, and C. virgusa (Negret, 1996b).

Plate 2. Cecropia Tree on an exposed river bank.

Its medicinal uses are varied. For example, the sap of C. palmata and C. peltata is astringent and is used against warts, calluses, dysentery, ulcers and venereal diseases. The leaves are analgesic, with wound healing properties and can also be used against infections of the liver. In Colombia they consider it effective against Parkinson Disease. It is even eaten, where the young buds are cooked as a vegetable (Raintree Group, 1998).

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Cecropia are an important species as a habitat and food source to many faunal species. In particular, their symbiotic associations with Azteca ants species, which live in their hollow stems is an example of the important associations between plant and animal species that exist in Tropical forests (Vasconcelos and Casimiro, 1997). These ants feed on glycogen rich food bodies found on a special organs at the base of the petiole and in return for food and nesting, Azteca ants are known to protect their host against encroaching vines and herbivores such as leaf cutting ants, which macerate their leaves (plate 3).

Plate 3. Cecropia leaf attacked by herbivory

Cecropia also play an important role in tropical forest dynamics. For example, Franco-Rosselli and Berg, (1997) state that Cecropia are an important pioneer, light demanding species in natural gaps of primary forest in the Andean regions as well as in the tropical regions of the Pacific and Amazonian forests. They are more abundant

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in secondary forests however, especially in areas disturbed and exposed by landslides or riverbanks and along roadsides and paths.

Their presence in primary forests is the result of gap formation, and gap dynamics. Canopy opening may be caused by any number of natural occurrences, such as bole, breach or crown breakages, fall of liana tangles, lightning, landslides, and the death of the standing tree (Hartshorn, 1990). It is suggested that the microclimate of smaller gaps may be inimitable to Cecropia and therefore they will only colonise gaps at least 400 m2 in area (Mabberley, 1992). Cecropia can grow to 25 m tall (mesophanerophyte) and may survive from ten to forty years or more, therefore long after the original conditions which favoured its colonisation have changed due to gap closure (Mabberley, 1992).

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3.4

GIS

Approaches

for

Modelling

Potential

Species

Distributions
Traditional methods of analysing and displaying the distribution of organisms in community ecology have used the classic approaches of ordination, classification and direct gradient analysis. A fuller description of these methods can be found in traditional statistical ecology books (Gauch, 1982, Digby & Kirby, 1987). However, briefly ordination and direct gradient analysis have traditionally involved the ordering of species along an environmental gradient or gradients. Multiple dimensions of variables have been used to approximate some pattern of response of a set of species. The general objectives of such studies in the past have been primarily to formulate hypotheses about the specific relationships between species composition at a site and the underlying variables. Classification has involved the grouping of species in relation to an environmental gradient.

Modern approaches focus a different objective although these traditional concepts still underlie the analysis. The aim is not primarily to find the causal relationship between a species and its environment, but rather to predict the distribution given that we already assume knowledge of their environmental tolerance limits. GIS can be used to model the spatial distribution of species based on the niche concept. Different GIS map layers describing the environmental conditions of a target area can be used to extract the variables that determine their distribution. Subsequently, by relating these to a map layer of the actual distribution one is able to derive a relationship between the two. By defining this relationship it can then be used to predict a species potential spread over a wider area.

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Carpenter et al (1993) divide such distribution models into two main classes; those that simulate detailed interactive processes between organisms and their environment using explicit performance parameters and those which use pattern analysis techniques to exploit the correlation between environmental variables and the sampled distribution of target taxa. The former require detailed knowledge of organism response to a given range of environmental factors over time. However, they argue that while these models provide important biological and ecological insights they do not necessarily satisfy the immediate needs of conservation where adequate process information is usually lacking.

The advantages of GIS based techniques for such studies are immense. Traditional methods of documenting species diversity involved the collection of floristic inventories. However, such methods require a large amount of field expertise. These studies are further made difficult due to the complexity and inaccessibility of TMCF. This has been the main reason why knowledge of species diversity is patchy and very poorly known in TMCF (Gentry, 1990). GIS techniques can provide a valuable resource for mapping species distributions, meeting the urgency needed for their conservation given the threat of deforestation.

The application of GIS systems to explore the biodiversity of crop species have successfully been implemented, with the result of producing maps indicating areas of favourable climates for their growth (Jones et al, 1997). The combination of both GIS and satellite imagery constitutes another method of furthering our knowledge of diversity distribution (Corbley, 1996).

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A problem with trying to map past vegetation distributions, for comparison with the present, is that past data sets are seldom geographically well referenced. However, the potential now exits to map present species distributions with Global Positioning Systems (GPS). GIS methods, in additions to their simplicity, provide a visual geographical representation, which past surveys could not.

At present Colombias protected areas make up 8.7% of the country (Place, 1993). Extension and development of these protected areas is being planed (Negret, 1996b). This is one area where knowing the distribution of certain species can help the decision making progress in the layout of corridors joining different national parks together. In addition, mapping both disturbance and distribution of species, areas where mass extinctions are likely to occur may be distinguished and therefore help to highlight areas of conservation priority (Smith et al, 1997). Furthermore, species extinction rates will often depend upon the distribution of deforestation and the extent of disturbance or degradation of those areas that remain forested. Therefore GIS can be used in areas where extinctions are likely to be prominent.

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Section 2: Sampling and Methodologies

Chapter 4 4.1 Defining a Sampling Strategy
Neldner et al (1995) stress that biased or inadequately sampled data sets may result in misleading predictions and therefore lead to poor management techniques being implemented. GIS can be used for defining the entire range of environmental variability so that it can be adequately sampled (Margules & Stein, 1989). Thus, the initial step was to classify the area into classes of similarity. This was carried out in IDRISI for windows. Based on analysis of the Digital Elevation Model (DEM), slope distribution and aspect where derived. These were classed as shown in table 4.1.1. The vegetation distribution map (figure1.1.4, section 1) was used as another data layer in the classification. All these themes were overlaid to produce a classified image with 25 classes (figure 4.1.1).

The resulting sampling map identified 6 significant areas of variability. Transects were laid out within these areas to determine how these different combinations of biophysical characteristics influence the occurrence of Cecropia (figure 4.1.2). Due to difficulty in accessibility of the area, it was not always possible to access all these zones. Nevertheless, 6 transects being 1) Primary forest, 2) Palo Verde, 3) Exit Path, 4) Asomadero, 5) Palm forest and 6) River station were located at or near the forest paths and correspond to classes 6, 3, 5, 17, 19, and 12 respectively. Figures 4.1.34.1.7 illustrate the altitudinal cross-sections of the different transects which range from between 1400 m to 1800 m. These transects were mapped with a GPS (see 23

section 4.3) which was set on track mode to record their geographical position at 1 minute intervals along their entire length.

Table 4.1.1. Classes Used in Classifying the Catchment.

Class Class 1 Class 2 Class 3 0 - 15 15 - 35 > 35

Slope

Class 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Cover 1 2 3 1 2 3 1 2 1 2 3 1 2 3 1 2 3 1 2 3 1 2 3 1 2

Aspect 4 4 4 5 5 5 5 5 4 4 4 5 5 5 5 5 5 4 4 4 5 5 5 5 5

Slope 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8

Aspect Class 4 Class 5 North (0 - 45 & 225 - 360) South (45 - 225) Land Cover Type Class 6 Class 7 Class 8 Primary Secondary Pasture

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.9 91

.9 73

99 9

98 2

7. 00 9

.0 18

6.

6.

-7 6

-7 7

-7

-7

-7 6

2.537

2.528

2.519

2.510

Classes 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

2.501

2.492

2.483

2.474

0.8

-7

0.8

1.6 Kilometers

Figure 4.1.1. Classified Sampling Map.

-7

6.

69 8

25

# # #

# ## # #

Exit Path

# # # # # # # # # ## # ## # # # ## # # ## ## # ## ## ## # # # ## # # # # ## # # # # # # ## # # #

# ## # # ### # ## ## # # # ## # # # #### # # ## # # ## ## ## # ## # # # ## ## # # # ## # # ### # # ## # # # # # # ## # # # # # # ### # # ## ## # # # # # ## # # #

Primary Forest Path

# # # ## # ##

# # # # # # # ## # # # ## # # #### ## # ## # ## # # # # # # ## # # # # ## # # # # # # # # # # ## ## # ## # # ### # # # # # # # # # # # # ## # # # ## # # ## # # # # ## # # # ## ### # #

Palo Verde Path

Palm Forest Path

# # # # # ## # # ## ##### ###### ### #### # # # ## # ## # # ## # ## # ## # # % # ## % % # ## % %% # #% # # # % % % # % % %#%## # % % % ### # % % # % # % % % #% ## % % # # # % #% % % %% # ### % ## % # # % # # # %# %%# #### # # # # %% # # ## # # % # ## # ## # # % # # ## # % # ## ### # # # # %% % ## # # %% %% %% % % # ## ### ### # # # %% %% %% % # % # # # # ## %% % % % # % %% % # # ## ## # % % %% # ## # # # %% # # ## # % # # % % # # # # %

River Station Path

% %

% % % %

% % %% % % %% %% %

% %% % %

%% % % %

% % %

% % % % %

%%

% %

% % % % % % % % % % % % % %% % % % %% % % % % %% %% %% % % % %%% % % %% % %% % % %% % %% % % % % % % % % %% % % % % % % %% %% % % % %%%% %% % %% % % % % % %%% % % %% % % % %% % % % %% % % %%% %% % % % %%% % % % % % %% % % % %% % % % % % % % % % % % %% % % %% % % % % % % %% % % % % % % % % % %% % %% % % % %% % % % %%

Asomadero P ath

% % % % % % % % % % % %

Rivst_utm Exit_utm Prim2_utm Palo_utm Palm_utm Asom_utm Riosmtr Hilldtm 226 - 254 198 - 225 170 - 197 142 - 169 113 - 141 85 - 112 57 - 84 29 - 56 0 - 28 No Data
# # # # #
%

400

400

800 Meters

Figure 4.1.2. Location of Transects shown on a hillshade terrain map.

26

Primary Forest Transect 1700 Altitude (m) 1650 1600 1550 1500 1450 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Figure 4.1.3. Primary Forest Transect

Palo Verde Transect 1540 Altitude (m) 1520 1500 1480 1460 1440 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

Figure 4.1.4. Palo Verde Transect

Exit Path Transect 1420 1410 1400 1390 1380 1370 1360 1 2 3 4 5 6 7 8 9 10 11

Figure 4.1.5. Exit Path Transect

Altitude (m)

27

Asomadero Transect
1900 1850 1800 1750 1700 1650 1600 1550 1500 1450 10 13 16 19 22 25 28 31 34 37 40 43
5

Altitude (m)

Figure 4.1.7. Asomadero Transect

Palm Forest Transect
1570 1550

Altitude (m)

1530 1510 1490 1470 1450 1 2 3 4

Figure 4.1.6. Palm Forest Transect.

River Station Transect 1460 1450 1440 1430 1420 1410 1400 1390 1380 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Figure 4.1.7. River Station Transect

Altitude (m)

46

28

4.2 Determining Biophysical Variables for Cecropia

If the distribution of a species is to be predicted accurately it is important to understand the factors which affect the types and numbers of individuals within a community (Divenvoorden, 1995). The first step therefore, was to try and determine the factors affecting the distribution of Cecropia from the literature (Franco Rosselli and Berg, 1997) and from field observations. These biophysical variables where then modelled and distributed over the catchment in PCRASTER, a GIS modelling program, to produce different data layer maps (see section 3, chapter 5).

In TMCF, altitude appears to be the overriding gradient influencing the types of species found. Therefore, factors that change with altitude like minimum and maximum annual temperature are also likely to be important to species tolerance limits. Other factors, which appeared to be influencing the distribution of Cecropia, were their proximity to gaps, and paths. Cecropia are light demanding species and therefore these exposed areas are more likely to receive solar radiation. Thus, aspect was used as a surrogate for this variable. Slope was chosen to be important in determining the formation of gaps; the steeper the slope the more likely is the occurrence of gaps due to treefall and landsliding. Finally, their proximity to rivers was another apparent important factor for two possible reasons; either because the riverbanks are exposed to incoming solar radiation or due to the greater moisture availability along these routes. Therefore a further two map layers were produced, one showing the distance to rivers and the other showing moisture availability. A more detailed description of how these were derived is given in chapter 5, section 5.1.

29

4.3 Locating Cecropia in the Field

Mapping of the presents or absence of Cecropia was determined using a GPS, (Sutherland 1996). Simply, every time an individual was encountered along a transect, its position was located by coordinates. A GPS receiver uses 24 satellites in orbit to determine the geographical position on the ground. A minimum of three satellites with good reception is recommended in order to obtain 3d navigation. This was not always possible in the field, especially in the lower altitudes and depressions of the river valley. A differential global positioning system (DGPS) was applied and post-processing differential was carried out to obtain a higher degree of accuracy. A GPS signal contains a random error between 300 m 400 m. By taking the GPS output and comparing it to the coordinates of a control point where the coordinates are known to be correct, the error can be calculated and applied to the data. Thus, the error is reduced to about 30 - 40 m.

The data obtained was divided into two parts. Half was used to define the control points used by DOMAIN to calculate their potential distribution (see chapter 5, section 5.2). The data from the GPS was then loaded into a program (source: Dr Mark Mulligan) designed to produce an ASCII grid file defining 1 for Cecropia presence and 0 for its absence. The ASCII file was then imported into PCRASTER and a map depicting the recorded distribution of Cecropia was produced. This was used as the training layer in DOMAIN.

The other half was used to build a point coverage in ARC/INFO showing their presence or absence within each 100m grid cell along each transect. This coverage

30

would then be compared by overlay, with the DOMAIN output in order to validate the model. In some cases more than one species was located within each grid cell. Their abundance could then be calculated and compared to the biophysical variables, in order to determine why Cecropia are more abundant in some grid cells and not in others. The frequency of occurrence could also be derived and classified by assigning a presence class (table 4.3.1), (Smith, 1980).

Table 4.3.1. Presence-Absence Classification Presence Class Stands of one community type studied in which species occur % 1 2 3 4 5 1-20 21-40 41-60 61-80 81-100

Presence is determined by dividing the total number of grid cells in which the species is found by the total number of grid cells investigated. Species that have a high percentage of presence are often regarded as more or less characteristic of that community, and their presence there is probably a result of their adaptability to the underlying environment.

Originally the aim was to differentiate the distribution of the male and female tree but discerning the different organs was very difficult, as the canopy was often too high to distinguish the organelles.

31

4.4 Field Validation of Modelled Climatic Variables

Measuring temperature at different altitudes with a min-max thermometer validated the modelled temperature output. It was placed in 3 altitudinal positions along the primary forest transect and kept there over a day in order to record the minimum and maximum range of diurnal fluctuations. The results show that temperatures decrease in an almost linear fashion with altitude (figure 4.4.1). However, measurements at higher interval frequencies would have made this relationship clearer.

Minim m&M u T p u axim m em erature Distribution w A ith ltitude

25

Temperature ( C)

20 15 10 M umT perature inim em 5 0 1400 M imum Temperature ax

A ltitude (m )

1500

1800

Figure 4.4.1. Measured Min and Max Temperature

Rainfall could not be modelled and distributed over the catchment, as there was no apparent empirical relationship between rainfall data and altitude or cloud cover.

32

However, a validation of modelled moisture availability was attempted through measurements of throughfall at 5 different altitudes. Throughfall troughs were constructed which were suspending from the tree trunks and funnelled into an 8 litre capacity bottle (figure 4.4.2). This procedure assumed that throughfall is a proportion of rainfall and therefore of moisture availability over space.

33

Figure 4.4.2. Throughfall Trough to Measure Throughfall.

34

The results show no apparent pattern in rainfall distribution with altitude and therefore a relationship, which could be applied to validate the modelled moisture availability, could not be derived (figure 4.4.3). In addition problems with the design of the throughfall troughs were encountered during very heavy storms. Storms larger than 15 mm resulted in the collection bottles over-spilling and therefore voiding the data. In order to overcome this, the area of the trough exposed to the atmosphere should be made smaller. Further, the original assumption may not have much credibility given that throughfall can be enhanced beyond rainfall contribution (see table 1, appendix) as a result of direct canopy interception from clouds.

Distribution of Throughfall W Altitude ith

20.00 Throughfall (mm) 15.00 10.00 5.00 0.00 1476 1567 1604 Altitude (m ) 1677 1689

Figure 4.4.3. Measured Throughfall Over an Altitudinal Gradient

35

Section 3: Modelling Cecropia Distribution

Chapter 5

5.1 Modelling the Biophysical Variables Important for Cecropia Distribution

The distribution of climatic and other biophysical variables over the whole catchment were determined by simple analysis of 3 basic maps (source: Mauricio Rincon): the DEM, the river networks and a map classifying vegetation into primary, secondary and pasture (figures 5.1.1 5.1.3). PCRASTER was used to model and distribute these variables. Given that accuracy of the GPS the DEM and subsequent data layers were resampled to a resolution of 100 m.

The basic climate variables of minimum and maximum temperature were distributed by deriving an empirical relationship between altitude and temperature data obtained from national climate stations (source: CIAT database). A strong relationship, r2 = 0.8 was derived from the regression analysis of the mean annual minimum and maximum temperatures of different stations between latitudes of 2-3 and longitudes of 76 with altitudes between 1300 m 2900 m , to reflect the range found in Tambito (figures 5.1.4 &.5.1.5).

36

Derived variables, calculated from the DEM, included slope and aspect. The topographic wetness index calculated by Ln(/tan), where the logarithm of accumulated upslope area divided by the tangent of the slope, was computed to indicate the moisture gradient over the catchment (Kirkby and Beven 1986). Other factors perceived to be important to the distribution of Cecropia was their proximity to the river network. A map layer showing the distance from rivers was computed by calculating the Euclidean distances to produce a map of equidistant buffers radiating out from the main river networks. The map layers for each of these secondary attributes are illustrated in figures 5.1.6 5.1.11. These were then exported as ASCII grids and imported into DOMAIN for the analysis.

41

5.2 Modelling Plant Distribution using Domain Model

The DOMAIN model was developed at CSIRO by Carpenter et al (1993), to map potential distribution of plants and animals. DOMAIN takes a known distribution of points, for example, a record of species presents data defined by the latitudelongitude coordinate system and constructs an environmental habitat envelope or domain. Essentially this domain is the area where the species is most likely to be found. It uses the known distribution as a set of training points to compare with map layers of different environmental factors within the same location. By evaluating the similarity between grid cells where the species is found with all the environmental layers, a grid of continuously varying similarity values is generated (figure 5.2.1).

The model uses range standardisation based on the Gower metric algorithm (Gower 1971) to equalise the contribution from every climatic attribute within each grid cell to the presence of a single species. This is based on the principles of monothetic classification (Gauch, 1982) and is preferred over variance standardisation techniques because it is less susceptible to bias arising from dense clusters of sample points (Carpenter et al, 1993).

Two quantification measures can be computed; the distance measure, constrained between 0 and 1, where 0 is nearer to the computed domain of the species; and the similarity measure which defines similarity between points as a percentage, so that 100 percent lies within the domain of the species.

41

Another facility available in DOMAIN allows the investigator to explore how survey transects account for environmental variability within an area. The same transects can be used to examine the degree to which they intercept the domain of specified species.

The simplicity of the DOMAIN approach serves as an advantage as it functions on a limited number of input variables compared to other process-based spatial models (Carpenter et al, 1993). While the integration of system dynamics into distribution models are considered more accurate in terms of the way they explain spatial change with time (Booth, 1991) they require a detailed knowledge of species response. Moreover, environmental data is needed at much finer spatial scales and frequencies. The capability of the model in predicting the distribution of Cecropia is evaluated, as is its robustness in analysing the resulting output distribution map.

44

Section 4: Model Results and Discussion

Chapter 6 6.1 DOMAIN Output

The projected distribution of Cecropia produced by DOMAIN is shown in figure 6.1.1. This map shows similarity classes, were a value of S=100 would indicate a high probability that the species is found there. This result was derived by taking into account all the variables discussed in chapter 5. A frequency graph of similarity classes (figure 6.1.2) illustrates that the resulting distribution shows how similarity areas with over 65% classification confidence encompass a comparatively large area, 52%. It is necessary however, to define the threshold similarity measure for Cecropia in order to determine whether the model is over predicting its distribution.

Carpenter et al (1993) state that the DOMAIN model in comparison with other distribution models, produced distribution patterns that were most consistent with the known ecology of the animal species they were modelling. This conclusion was based on knowledge of the species. If knowledge is limited it is necessary to validate the modelled output by comparing the correspondence of observed Cecropia presence to modelled distribution.

44

97

-7 7. 02

7. 03

7. 00

6.

6.

-7

2.54

2.52

2.51

2.49

2.48

-7

-7

-7

Rivers Domain Similarity Values -10 - 2 3 - 14 15 - 26 27 - 38 39 - 51 52 - 63 64 - 75 76 - 87 88 - 100 Grid30ll

2 Kilometers

Figure 6.1.1. Modelled DOMAIN output. Similarity Measure

-7

6. 95

98

45

Histogram showing the distribution of modelled Similarity Classes

Figure 6.1.2. Histogram of Frequency Distribution of Similarity Areas

6.2 Validation of Model Results

The point coverage was rasterised and spatial analysis between the model output and measured data was carried out. The validation data set appears to overlay with areas of high similarity (figure 6.2.1). However, one could argue that the data set is concentrated within the small area covered and that it is necessary to sample much wider area in order to validate grid cells further away. However, taking the data available it was possible to determine the success of the model.

46

-7 7. 0 0

6. 98

2.54

2.52

-7 7. 03

# ## # ## # # # # # # # ## # # # #

# #

2.51

# # # # #

# # # # ##

# ##

# # # # ## # ## # # # # #

2.49

2.48

-7

-7

6.

-7

Cvprim2 Cvpalo2 Cvexit2 Cvpalm Cvasom Imgrdshift -10 - 2 3 - 14 15 - 26 27 - 38 39 - 51 52 - 63 64 - 75 76 - 87 88 - 100 No Data Grid30ll
# # # # #

0.5

0 0.5

1 Kilometers

Figure 6.2.1. The Congruence of Validation Data to Model Output.

-7 6. 95

7 .0 2

97

47

By overlaying the rasterised Cecropia cells with the DOMAIN grid, it was possible to determine which Cecropia cells corresponded with which similarity zones. Query analysis was performed in IDRISI to determine the frequency distribution of conforming cells (figure 6.2.2).

T eFrequencyDistrib tionofC iaC w the p ictedD m in h u ecrop ells ithin red oa 10 9 8 7 6 5 4 3 2 1 0 66 69 72 74 75 76 78 79 80 81 82 83 84 85 86 100 Sim ilarity>65% No of Cecropia cells in each similarity Class

Figure 6.2.2. Congruence of Cecropia cells with DOMAIN output

All Cecropia cells corresponded to similarity areas >66%. 22.7 % of the points where found within the 100% similarity zone (S=100) predicted by the model, while 61.4% where found within S>80.

48

6.3 Environmental limits of Cecropia

The variability of environmental conditions within the predicted zones of distribution could not be assessed because of problems with overlay of data layers when they were converted to IDRISI. However, it was proposed to extract the range of biophysical variables within the cells computed as having a high degree of similarity to see whether the tolerance limits of Cecropia vary significantly. If they are found within a narrow band of varying climatic conditions then it could indicate their susceptibility to future climate change.

Data query analysis of Cecropia cells with the data layers was possible and the variability of conditions within these cells could be analysed. By overlaying the river network map with the modelled output, it evident that maximum congruence corresponds to the areas near the main rivers (figure 6.1.1).

The maximum temperature range defining Cecropia distribution appeared to be between 24C - 28C, (figure 6.3.1) which corresponded to the upper range for the whole area, while their minimum temperature limits were between 13C - 16C (figure 6.3.2).

Analysis of aspect showed that more Cecropia were found on North facing slopes (63.2%) compared to South facing (36.8%). Cecropia do appear to have an altitudinal

49

gradient. They appear to be found between 1350 m to 1900 m, with their maximum range between 1400 m to 1600 m (figure 6.3.3).

M u T axim m emperatureRangedefining Cecropia Presence

16 14 12 10 8 6 4 2 0 24 25 26 27 28 M u T axim m emperature(C)

Figure 6.3.1. Maximum Temperature Limits of Cecropia

No of Cecropia Cells

50

M u T p inim m emeratureRange d efiningCecropia Presence

No of Cecropia cells

15 10 5 0 13 14 15 16 T p emerature(C )

Figure 6.3.2. Minimum Temperature Limits of Cecropia

Altitudinal Range of Cecropia Presence

12 10 Frequency of Occurence 8 6 4 2 0 <1400 1400-1500 1500-1600 1600-1700 1700-1800 >1800 Altitude Zones (m)

Figure 6.3.3. Altitude Limits of Cecropia

51

Frequency analysis could not be performed due to problems with data layers not matching. However, it was intended to extract the number of cells with Cecropia presence along each transect to see if any difference in environmental gradients along these reflected the abundance of Cecropia.

52

6.4 Sensitivity Analysis of DOMAIN

Further analysis was carried out in order to evaluate the models performance when run with less environmental data layers. It appears that the modelled output is less conservative. Figure 6.4.1 shows the predicted output based on 3 data layers; Proximity to rivers, slope and the DEM, those perceived most important to their distribution. It is even more general in its prediction when only one layer is use, that is proximity to rivers (figure 6.4.2).

It is important to assess when evaluating an area for conservation purposes, which scale is needed for determining species distributions. If only a general guide to species distributions is required then the output of DOMAIN is adequate for yielding this information. However, for rare species, with limited areal extents, it may be necessary to know the distribution at much finer scales. This can partly be achieved by carrying out the analysis with higher resolution (both temporal and spatial) biophysical data.

53

60-80

40-60

>80

Figure 6.4.1. DOMAIN output, with 3 data layers

40-60

>80
60-80

Figure 6.4.2. DOMAIN output with one data layer

54

6.5 Discussion of Results

The application of the DOMAIN model for predicting the Cecropia species can generally be perceived as achieving what it sets out to do. In order to be more confident of its performance, much more field validation is necessary. Its computational abilities had been compared in other studies, to other distribution models (Neldner et al, 1995) but in terms of its application to defining the adequacy of a sampling survey. Its initial application was to predict an animal species. It performance here in predicting the distribution of a vegetation species appears to identify areas most likely to have Cecropia. Therefore, it is an easy and simple method that can be applied in an initial assessment of an area, especially if knowledge and data of the area are very limited.

Its performance has been shown to be sensitive to the number of data layers used and therefore it is important to determine with accuracy which biophysical variables are important to a species survival. In this case this was determined from field observation and literature reviews. However, if not very much was known of the species niche requirements, ordination analysis of the biophysical variables, such as Principle Component Analysis could be carried out to ascertain which ones are the most significant in determining the distribution of a species, before carrying out the DOMAIN analysis.

55

DOMAINS limitations as a modelling package restrict analysis beyond initial computation of modelled distribution. If such models are to became a desirable means of modelling species distributions in may be useful to integrate spatial analysis operators in order to asses and display results. The use of different GIS packages in this study for the analysis and validation of the output posed a lot of problems when converting from one program to another, often with the result of shifting the latitudinal and longitudinal positions of the grids, requiring corrections to be applied.

Because such approaches require data acquisition using a GPS, the accuracy of its prediction will depend on the accuracy of geocoding. Under different satellite regimes, the signal received will vary. This may result in miss coding of ground data. In marginal areas and for species of limited range this may cause problems in the accuracy of their predicted distribution. Therefore the continued improvement of these systems is essential for high accuracy mapping of endangered and endemic species.

An alternative method that could have been used to validate the model could have been the use of an aerial photograph. The areal photograph available for the area was too crude for this purpose. However, if a 25 meter resolution image (or finer) could be obtained, then areas of Cecropia presence could be depicted and digitised and then warped onto the map produced by DOMAIN to see whether the areas of actual Cecropia and modelled Cecropia match. The advantage of an areal photograph is that it covers a comparatively larger area than that possible on the ground by foot.

56

6.6 Conclusions
This approach is important for two reasons; not only does have the potential to mark out possible hot spots of biodiversity but at the same time it allows us to interactively map out climatic gradients. Both these components mark out the base level conditions on which the impact of future climate change can be assessed. In addition, when planning areas for conservation, it is important to encompass long gradients of climatic conditions, as it is easier for species to migrate along larger gradients. Conversely, when barriers exit, any future climate change will result in mass extinctions (Van der Hammen 1995).

The DOMAIN modelled output appears to have predicted the distribution of Cecropia with a confident degree of accuracy. Thus the algorithm used by the model appears sound. However, Namkoog (1994) suggests that even though such approaches tell us how present conditions are affecting species distributions using this basis for predicting their future location may be crude, given that it does not tell us nothing about the evolutionary dynamics of a species.

Schemske et al (1994), highlight the need to assemble the necessary demographic information to determine if the numbers of individuals and populations of a species are increasing, decreasing or are stable, and which life history stages have the greatest effect on population growth and species persistence. Such demographic parameters may have a greater bearing on how they would respond in space to changing underlying conditions

57

If distribution models like DOMAIN could integrate further analytical tools for post assessing their performance and for interactively querying sites of species abundance, it may be useful in determining important areas at grid scale resolution. Individual grid cells up to 100-meter resolution appear to yield important information about the underlying conditions and their effect on the distribution of a species. This could be much more useful than existing global maps of species distribution, as these tend to be very general, with little information on specific species. This approach does hold the potential to scale up without loss of detail if the resolution of the input information reflects the robustness and abundance of the species. Nevertheless, this simple approach provides the starting point for assessing some of the most endangered ecosystems. GIS as a whole, provide an important interface for adequately addressing the issue biodiversity conservation.

58

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MacArthur R. H (1984). Geographical Ecology. Patterns in the Distribution of Species. Priceton University Press, New Jersey. Magurran A E (1988). Ecological Diversity and its Measurement. Croom-Helm, London. Miller R I (1994). Mapping the diversity in Nature. Chapman and Hall, London Namkoong G (1994). What do measures of Biodiversity tell us? in Boyle T J B and Boontawee B (eds.). Measuring and Monitoring Biodiversity in Tropical and Temperature Forests, CIFOR, proceeding of a IUFRO symposium. Place S E (1993). Latin America. Nature and society. Smith R L (1980). Ecology and Field Biology 3rd Edition. Harper and Row, New York. Sutherland W J (1996). Ecological Census Tecchniques; a handbook. Cambridge University Press, Cambridge. Van der Hammen T (1995) Global change, Biodiversity and Conservation of Neotropical Montane Forests. In Churchhill S P, Balslev H and Forero E and Luteyn J L (eds). Biodiversity and conservation of Neotropical Montane Forest. Proceeding of the Neotropical Montane forest Biodiversity and conservation symposium. The New York Botanical Gardens 21-26 June 1993. Whitmore C and Sayer J A (1992). Tropical deforestion and Species Extinction. Chapman and Hall, London

Internet Resources

The raintree Group, Inc, Austin, Texas 78756.@ http://www.rain-tree.com/cecropia.htm

Software

DOMAIN model, @ http://www.cgiar.org/cifor/research/intro_d.html IDRISI for Windows, version 2. Clarks University. @ http://www.idrisi.clarku-edu PCRASTER, Utrecht University, Netherlands @ http://www.geog.u.nl/pcraster.html ARC/INFO, CIAT, Colombia MAPCUT, Source Dr Mark Mulligan, Kings College London

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Appendix
Table 1. Throughfall Data.
Date Total Daily Rainfall (mm) Site Altitude Av Width of Angle of Area (m) Trough (cm) Inclination Exposed To (degrees) Atmosphere (m2) 1476 1567 1604 1677 1689 1476 1567 1604 1677 1689 1476 1567 1604 1677 1689 1476 1567 1604 1677 1689 20 15 20 18.79 20 20 15 20 18.79 20 20 15 20 18.79 20 20 15 20 18.79 20 10.5 26 37 16 24 10.5 26 37 16 24 10.5 26 37 16 24 10.5 26 37 16 24 0.42279961 0.28986108 0.343413269 0.388335306 0.392824547 0.42279961 0.28986108 0.343413269 0.388335306 0.392824547 0.42279961 0.28986108 0.343413269 0.388335306 0.392824547 0.42279961 0.28986108 0.343413269 0.388335306 0.392824547 Amount of Actual Throughfall Throughfall Collected (mm) (ml/1000=mm) 1.607 1.179 1.843 0.816 0.992 8 8 8 8 8 4.53 7.98 5.985 5.105 6.06 4.1 3.38 4.68 2.97 5.58 3.80085497 4.067465699 5.36671167 2.101276878 2.525300438 18.92149332 27.59942798 23.29554713 20.60075371 20.36532611 10.71429559 27.53042941 17.4279812 13.14585596 15.42673453 9.697265325 11.66075832 13.62789507 7.648029814 14.20481496

5th-Aug-98

6th Aug 98

8th Aug 98

9th Aug 98 ?

4.1 A B C D E 15.6 A B C D E 16.5 A B C D E A B C D E

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