Response of Invertebrates to Alien and Indigenous Vegetation Characteristics in Nduli and Luchaba Nature Reserves, Eastern Cape, South Africa

RESPONSE OF INVERTEBRATES TO ALIEN AND INDIGENOUS VEGETATION CHARACTERISTICS IN NDULI AND LUCHABA NATURE RESERVES, EASTERN CAPE, SOUTH AFRICA

INAM YEKWAYO

A dissertation submitted in fulfillment of the requirements for the degree of:

MASTER OF SCIENCE (MSc)

In Zoology

at WALTER SISULU UNIVERSITY

SUPERVISOR: Dr A.S. NIBA

September 2012

ABSTRACT

Most invertebrate species are becoming extinct due to habitat loss and alien plant invasions. Hence this study aimed at determining the response of invertebrates to alien and indigenous vegetation within protected areas in the King Sabatha Dalindyebo (KSD) Local Municipality, Eastern Cape, South Africa. Invertebrates were collected using pitfall traps, during 12 sampling occasions from May 2010 to April 2011 numbers of sampling sites. Although the sampling method was adapted to collecting ground dwelling invertebrates, opportunistic flying invertebrates were also collected. A total of 7 flying invertebrate orders, 25 families, 34 species and 248 individuals were attracted to traps while 5 orders, 19 families, 50 species and 1976 individuals of soil surface- dwelling invertebrates were collected. ANOVA test showed no significant differences (p > 0.05) in species richness and abundance across sites for soil surface-dwelling invertebrates. Bray-Curtis similarity measures in PRIMER and correspondence analysis (CA) in CANOCO showed that sampling units with alien invasive plants shared most soil surface-dwelling invertebrate species at ± 75% level of similarity. Sampling unit A from the Mix alien (MA) site shared most species with indigenous vegetation sites. Sampling units from indigenous vegetation sites shared most species at ± 65% level of similarity. Multivariate analysis using CANOCO indicated that certain site variables such litter depth influenced the distribution of soil surface-dwelling invertebrates across sites. The study provided preliminary data and information for promoting invertebrate biodiversity conservation within protected areas (Nduli and Luchaba Nature Reserves) of the KSD Local Municipality.

Key words: invertebrates, indigenous vegetation, alien vegetation, Lantana camara, Acacia mearnsii, and Eucalyptus.

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ACKNOWLEDGEMENTS

If God was not with me I would not have done this study, special thanks to Him for the strength He gave me to face all challenges. Thank you to my supervisor, Dr Niba for his assistance in making the completion of this study possible and also building me to be an independent scientist. National Research Foundation (NRF) is acknowledged for funding this project. Thanks to the staff of Nduli and Luchaba Nature Reserves for allowing the project to occur. Special thanks go to the following: Dr Ansie Deppenaar, Ashely Kirk- Spriggs and the Iziko Museum taxon specialists for identifying invertebrate species. The Mthatha dam analytical service is acknowledged for analyzing soil samples for this project. Thanks to Mr. Forbanka D.N for helping me during data analysis phase of the project. Lastly I want to thank my family, friends and colleagues especially Miss Nomfundo Nkabi for making it possible for me to do this project.

TABLE OF CONTENTS

ABSTRACT…………………………………………………………………………………………………..¡

DECLARATION.…………………………………………………………………………………………….¡¡

DECLARATION ON PLAGIARISM……………………………………………………………………..ііі

ACKNOWLEDGEMENTS…………………………………………………………………………………..iv

TABLE OF CONTENTS…………………………………………………………………………………….v

LIST OF TABLES……………………………………………………………………………………...... vi¡

LIST OF FIGURES…………………………………………………………………………………………v¡i¡

LIST OF APPENDICES……………………………………………………………………………………x

CHAPTER 1: INTRODUCTION

1.1. INTRODUCTION …………………………..………………………………………………………..1

1.2. RATIONALE AND SPECIFIC OBJECTIVES……………………………………………………12

CHAPTER 2: METHODOLOGY

2.1. STUDY SITE……………………………………………………………………………………………14

2.1. INVERTEBRATE SPECIES SAMPLING………………………………………………………..17

2.3. DATA ANALYSIS……………………………………………………………………………………..18

CHAPTER 3: RESULTS

3.1. RESULTS……………………………………………………………………………………………….20

CHAPTER 4: DISCUSSION

4.1. EFFECT OF SAMPLING METHOD ON SPECIES DISTRIBUTION……………………………………………………………………………………………..38

4.2. SPECIES DISTRIBUTION TRENDS ACROSS SITES…………………………………………………………………………………………………………..38

4.3. SPECIES DISTRIBUTION TRENDS ACROSS MONTHS…………………………………40

4.4. RELATIONSHIP BETWEEN SPECIES, SITES AND ENVIRONMENTAL VARIABLES…………………………………………………………………………………………………….41

4.5. INDICATOR POTENTIAL OF INVERTEBRATE TAXA SAMPLED DURING THE STUDY PERIOD……………………………………………………………44

4.6. CONCLUSION………………………………………………………………………………………….46

4.7. CONSERVATION RECOMMENDATIONS………………………………………………………47

4.9. REFERENCES………………………………………………………………………………………….49

4.8. APPENDICES…………………………………………………………………………………………..65

LIST OF TABLES

Table No & Title Page No

1: Number of specimens from each order expressed as a percentage 24

2: 2-way ANOVA for species richness of soil 25

surface-dwelling invertebrates

3: 2-way ANOVA for species abundance of soil surface-dwelling invertebrate 25

4: 2-way ANOVA for species richness of flying invertebrates 26

5: 2-way ANOVA for species abundance of flying invertebrates 26

8: Eigenvalues, cumulative percentage variance of species-environmental 36

relation (CCA) of axes 1 and 2 9: Intra-set correlation for both alien and indigenous vegetation sites 37 combined

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LIST OF FIGURES

Figure No & Title Page No

1: Map of study area: Nduli and Luchaba Nature Reserves 16

2a: Spatial trends for soil surface-dwelling invertebrate abundance 27

and richness across sites during sampling period

2b: Spatial trends for flying invertebrate abundance and richness 27

across sites during sampling period

3a: Temporal trends for soil surface-dwelling invertebrate abundance 28

and richness during the sampling period

3b: Temporal trends for flying invertebrate abundance and richness 28

during the sampling period

4a: Cumulative no of taxa caught for all soil surface-dwelling orders 29

combined, as well as dominant orders (Hymenoptera: Fomicidae,

Araneae, Coleoptera)

4b: Cumulative no of invertebrate species caught for all four sites 29

and for alien and indigenous vegetation sites separately

5: Cumulative species rank dominance curves for 30

a) Eucalyptus and b) Mix alien sites

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6: Cumulative species rank dominance curves for 30

a) Forest and b) Grassland indigenous sites

7: Cumulative species rank dominance curves for all sites 31

8a and b: a) Dendogram and b) NMDS ordinations 32

for alien sites (M- Mix alien, E-Eucalyptus )

8c: CA biplot for alien vegetation sites (E-Eucalyptus 32

and M-Mix alien)

9a and b: a) Dendogram and b) NMDS ordinations 33

for indigenous sites (F-Natural forest, G-Grassland)

9c: CA biplot for indigenous vegetation sites (F-Natural Forest 33

and G-Grassland)

10a: Dendogram showing percentage similarity of all site sampling units 34

10b: NMDS ordinations based on all site sampling units 34

10c: CA bi-plot for alien and indigenous vegetation sites 34

11: CCA tri-plot for all four sites, depicting relationship 35

between sites, species and environmental variables

LIST OF APPPENDICES

Appendix Page No

1: Full names of orders, families, species and acronyms of 65

soil surface-dwelling invertebrates

2: Captured flying invertebrate orders, families and species names 69

3: Data for environmental variables 72

CHAPTER 1

INTRODUCTION

Invertebrates as important components of biodiversity

Invertebrates are important components of biodiversity, inhabiting all areas from soil and litter layers to herb, understory and canopy layers (Oxbrough, et al., 2010). Invertebrates (non-arthropods, non-insect arthropods and insects) comprise about 97% of all animal species, and insects alone constitute the largest class of organisms, accounting for over 75% of all animal species (Martin & Major, 2001). Furthermore, the diversity of invertebrates is related to plant species diversity, structural vegetation diversity, spatial heterogeneity, micro-habitat patch size and density (Smith, et al., 1985). Invertebrate remain major energy conduits and agents of nutrients and material recycling in marine, freshwater and terrestrial ecosystems throughout the world. Sustainability of the earth’s life support systems depend on their well-being as invertebrates remain a critically important component of biodiversity in terms of species numbers and biomass, occurring in diverse ecosystems (New, 1998).

This faunal group also plays a significant role in the processes of pollination, nutrient cycling, seed dispersal, soil formation and fertility, plant productivity, organic decomposition and regulation of populations of other organisms through predation and parasitism (Keesing & Wratten, 1998; Lovell, et al., 2007; Martin & Major, 2001; Ward & Lariviere, 2004). Oxbrough, et al. (2010) noted that ground-dwelling invertebrates are relatively easily captured and identified, and their ecology as well as behaviour is relatively well known compared to other invertebrate taxa.

Invertebrates, especially brightly colored insects are important indicators of environmental changes, and exhibit a wide range of body sizes, growth rates, life history strategies and ecological preferences. These can be linked to specific environmental variables, thereby providing a greater understanding of their responses

to these variables and interaction into predictive models for various ecosystems (Andersen, et al., 2004; Lewinson, et al., 2005; Ward & Lariviere 2004).

 Invertebrates as bio-indicators

A bio-indicator refers to a species or group of species that readily reflects the abiotic or biotic state of an environment. This information is useful in managing ecological relationships, between species and their distribution trends (McGeoch, et al., 2002; Stewart, et al., 2007). McGeoch (1998) postulates three categories of bio-indicators: environmental, ecological and biodiversity indicators. Environmental indicators detect and monitor changes in environmental state, while ecological indicators demonstrate the impact of a stressor on biota, and also monitor long term stressor induced changes in biota. Biodiversity indicators are used to monitor and estimate the diversity of taxa in a specified area where measurable parameters or variables of biodiversity such as species richness and endemism are needed for conservation planning (McGeoch, et al., 2002; Stewart, et al., 2007).

Carignan and Villard (2002) noted that indicator species include species which occupy a broad range of habitats, have wide range of requirements in terms of habitat patch size, habitat structure and configuration. Indicator species depend on certain ecological processes where they are expected to display a wide range of sensitivities to habitat modification and disturbance of natural processes, and their protection likely to maintain ecosystem function (Carignan & Villard, 2002; Pearce & Venier, 2006).

Insects have been flagged as promising bio-indicators for over two and half decades because of their significant contribution to global species richness, biomass and ecological function as well as their responsiveness, life history and behavioural diversity (Stewart, et al., 2007). As a result, during selection and design of nature reserves, insects are used as bio-indicators to highlight areas of maximum collective diversity and are expected to be indicative of total biological diversity (McGeoch, et al., 2002).

Andersen (1997) further indicates that invertebrates in general, have a dominant biomass, are diverse, and occupy very narrow niches that remain suitable for a short period of time, making them suitable for use as bio-indicators. Several invertebrate groups are fundamentally important in ecosystem function and are too sedentary to colonize new patches even within relatively short distance which makes them more sensitive to fragmentation than other more mobile taxa (Andersen, 1997; Maes, 2004). Furthermore, several invertebrates react more rapidly to environmental changes than long living animals (vertebrates) because they usually complete their life cycle every year (Maes, 2004).

Underwood and Fisher (2006) indicated that ants for example, are useful tools to monitor ecosystem conditions because they are abundant and ubiquitous in both intact and disturbed habitats, as they are sensitive and rapid responders to environmental variables. Ants are important functionally at many different trophic levels and play important ecological roles in soil turnover and structuring, nutrient cycling, plant protection, seed protection and dispersal (Lewinson, et al., 2005). Samways (2005) further noted that brightly colored insects have the potential to be used as flagship groups in conservation planning and implementation programs. Invertebrates also remain the principal food source for other invertebrates and vertebrates such as fish which consume terrestrial invertebrates especially during summer months when aquatic invertebrates are limited. As a result, land use practices that influence the input of terrestrial invertebrates into streams are predicted to have consequences for fish production (Edwards & Huryn, 1996; Martin & Major, 2001; Oxbrough, et al., 2010; Sweka & Hartman, 2008;; Ward & Lariviere, 2004). Invertebrate groups such as termites, ants and earthworms have been found to be excellent indicators for soil fertility (Nadel, et al., 2006).

Martin and Major (2001) found that spiders are likely to play an important role in shaping terrestrial arthropod communities since spiders are at the top of invertebrate trophic levels. Ground-dwelling invertebrates such as spiders and carabid beetles are

frequently used to assess habitat quality in various forested ecosystems (Oxbrough, et al., 2010). Soil and leaf litter invertebrates generally have poor dispersal abilities, being often restricted to their specific microhabitats. Millipedes and snails have also been found to be effective as ecological and biodiversity indicators in South Africa (Uys, et al., 2009).

The importance of habitat quality in conserving invertebrates

Terrestrial and freshwater invertebrates largely depend on native vegetation for their survival, with many of these faunal groups facing high extinction rates as a result of increasing habitat loss, degradation and invasion by alien plants and animals to their environment (Gordon, 1998; Hamer & Slotow, 2009). Invertebrate biodiversity conservation can be promoted by habitat preservation and management rather than single species initiatives (Lewinson, et al., 2005). Hence, Hills, et al. (2008) has suggested that invertebrates that live in cave ecosystems depend largely on allochtonous leaf litter for their energy. They further stated that changes in the quality and quantity of forest floor leaf litter due to exotic plants lead to changes in litter dwelling invertebrate assemblages because exotic leaf litter can have different chemical and physical properties when compared to the native litter. Exotic leaf litter may provide structurally different habitat that differ in its rate of litter break down and release of nutrient compared from native litter (Harris et al., 2004).

Protected areas (native reserves) in South Africa serve as refugia, for providing high quality habitat patches for invertebrate biodiversity conservation but challenges resulting from their size and numbers do arise. Even within these reserves, alien invasive plants impact invertebrate species composition and distribution differently at micro-habitat and landscape levels (Samways, 2005).

Effects of alien plant invasions on ecosystem processes

Alien plants are species with an evolutionary history that occurred elsewhere i.e. outside their home range, and disperse rapidly by wind, water and animal transport to a 4

foreign environment. These plant species have become a worldwide threat to ecosystems and their biodiversity because they negatively affect all components of biodiversity from genes to ecosystem processes, by displacing native plants and consumer assemblages (Harvey, et al., 2010; Higgins, et al., 1999; Houlahan & Findlay, 2004; McGeoch, et al., 2006 ; Mgobozi, et al., 2008; Morales & Traveset, 2009; Moron, et al., 2009; Reinhart & VandeVoort, 2006; Strayer, et al., 2003).This scenario has resulted in a reduction of native biodiversity which in turn interferes with the delivery of a range of important ecosystem services such as nutrient cycling, increase carbon assimilation, increase flammability, biological control of plant or insect pests and pollination (Gooden, et al., 2009; Harvey, et al., 2010; Higgins, et al., 1999; Mgobozi, et al., 2008). Houlahan and Findlay (2004) noted that exotic species can cause fundamental changes in ecosystem processes and community structures that may have disastrous economic consequences which include loss of crops, forests, fisheries and grazing capacity. Plant invaders of natural ecosystems (environmental weeds) threaten ecosystem processes and species diversity at a local and global scale. These plant invaders inhibit recruitment of resident native species by preventing seedling establishment and growth as well as modification of plant pollinator interactions (Gooden, et al., 2009). Morales, et al. (2009) found that alien flowering plants compete with native plants for pollination and as a result, the presence of alien flowering plants leads to a decline in pollinator visitation and reproductive success of native plants.

Reinhart and VandeVoort (2006) noted that exotic plant species not only affect native tree species but also affect the ecosystem function and macro invertebrate communities. Standish (2004) has demonstrated that where invasive exotic plant species have replaced native vegetation, there is also an impact on native invertebrate assemblages. These impacts vary with the degree of change of the vegetation diversity and structure e.g. a comparison of litter invertebrate assemblages of exotic Chrysanthemoides monilifera shrub land and native heath land of similar structure showed minimal impacts while a comparison of native grassland invaded by exotic shrubs and trees with non-invaded native grasslands indicated significant differences in 5

composition of the invertebrate species assemblages (Standish, 2004). This study will determine the response of invertebrate assemblages to Lantana camara, Acacia mearnsii, and Eucalyptus invasions within protected areas of the KSD Local Municipality.

Brief taxonomic profile and description of common invasive plants identified at sites used in this study

Lantana camara

Lantana camara Linnaeus which is commonly known as lantana belongs to the family Verbenacea, it is a hardy, evergreen, struggling shrub with characteristic odour and can grow to 3 m height and occurs in clumps which flowers throughout the year and it is also a nectar producer (Dua, et al., 2010). L. camara is a noxious weed that grows in many tropical and subtropical parts of the world (Pasha, et al., 2007; Sharma, et al., 2007). It is native to South and Central America. Even though L. camara has been naturalised in the warm, moist subtropical and temperate regions of Limpopo, Mpumalanga, Kwazulu-Natal and Eastern Cape in South Africa, it has also became abundant in the more extreme central highlands of Gauteng province (Phenye & Simelane, 2005).

L. camara is considered one of the world’s worst alien species because in natural ecosystems it is implicated in widespread loss of native plant species diversity via recruitment limitation of native species and altered ecosystem structure and function (Babu, et al., 2009; Gooden, et al., 2009; Vivian-Smith & Panetta, 2009). Lantana has imposed a great threat to land productivity, grazing livestock, biodiversity and to the overall ecology (Pasha, et al., 2007). Goulson and Derwent (2003) found that L. camara invades pasture, crops and native ecosystems therefore causing substantial economic losses and environmental degradation. It forms a dense and impenetrable thicket that suffocates indigenous vegetation (Duggin & Gentle, 1998). It is an important weed of agriculture and forestry since it encroaches on plantations, orchards and on pastures where it forms dense thickets so that livestock cannot penetrate. The leaves of this

plant are toxic when ingested by livestock although toxicity varies between the strains (Goulson & Dewent, 2003).

L. camara does not only have negative impacts but also have positive impacts since all parts of Lantana are used traditionally for several ailments throughout the world (Barreto, et al., 2010). Leaves of this plant are used as an antitumeral, antibacterial and antihypertensive agent. Roots are used for treatment of malaria, rheumatism and skin rashes. Lantana is used as carminative, antispasmodic, antiemetic and to treat respiratory infections such as cough, cold, asthma and bronchitis (Barreto, et al., 2010). It is used for the development of furniture products, baskets, mulch, compost, drugs and other biological active agents (Pasha, et al., 2007).

Acacia mearnsii

Black wattle (Acacia mearnsii) belongs to the family Mimosaceae. It is a fast-growing leguminous tree that grows to 20m tall. It was first introduced in South Africa in 1864 through seeds collected from southern Australia as a shade tree for livestock, wind breaks and as a source of fuel wood on farms (Beck, et al., 2003; Govender, 2007). It is an extremely versatile and useful tree, not only to the forestry industry as a source of high quality raw material for pulp production and as a source of vegetable tannin, but also for fire wood and building purposes by the rural communities and local farmers (Beck, et al., 2003). Black wattle is one of the widespread and significant invasive alien trees in South Africa and great concern is frequently expressed over its potential effect on loss of biodiversity, reduced catchment water yields, increased soil erosion and by exacerbating the intensity of fires (Impson, et al., 2008). This species forms dense stands and maintains a high green leaf area throughout the year and frequently replaces seasonally dormant grasslands and fynbos (Dye & Jarmain, 2004). The success of it as an invasive plant in the landscape is largely attributed to the annual production of enormous seed crops which accumulate in the soil for many years (Impson, et al., 2008).

Eucalyptus

Gum tree (Eucalyptus) belongs to the family Myrtaceae. It is a fast-growing tree that can grow up to over 60m tall and is native to Australia (Grattapaglia & Sederoff, 1994). Bernhard-Reversat, et al. (2003) noted that Eucalyptus is extensively grown in the tropics either as farmer forestry or as industrial plantations mainly for paper pulp production. In Africa, Bernhard-Reversat, et al. (2003) said that, gum tree is grown on sandy soils which are usually nutrient poor so as to support tree growth while Weinberg, et al. (2010) noted that natural regeneration of Eucalyptus in south-eastern Australia depends on the proximal seed source, appropriate soil moisture and temperature, the degree of harvesting by seed predators and appropriate seedbed. Eucalyptus leaf litter contains large amounts of phenolic which are antibiotics and anti- feeding agent for invertebrate and vertebrate fauna and it also contains low nitrogen content which could account for low density of soil macro-fauna especially of earthworms (Bernhard-Reversat, et al., 2003; Mboukou-Kimbatsa, et al., 2007). It is a fast-growing tree that is desiccation tolerant, drought tolerant, adaptable and of high economic benefits. It is now known as one of the world’s three major fast-growing tree species which is widespread in tropical and subtropical regions (Castillo, et al., 2010; Liu & Li, 2010).

Impact of invasive alien plants on invertebrate faunal communities

Humans have been identified as a major vector in the dispersal of exotic species throughout the world (Houlahan & Findlay, 2004; Strayer, et al., 2003; Tallamy, 2004). Alien species have large impacts on invertebrate populations due to their unpalatable taste to most native phytophagous insects, and as a result they reduce invertebrate diversity (Chown, et al., 2008; Tallamy, 2004), since native insects share little or no evolutionary history with alien plants (Mgobozi, et al., 2008). Alien invasive plants have been implicated in the extinction of about 58 native plant species in the Cape Floral Kingdom and have also contributed to the endangered status of 3 435 other species of Southern African plants (Macdonald, et al., 2003). These plant species also have 8

significant impact on native vegetation and their associated food webs. It is estimated that about 75% of all animal species are phytophagous insects that also serve as food to a number of species such as reptiles, amphibians, birds and mammals. (Gerber, et al., 2008; Tallamy, 2004).

Robson, et al. (2009) noted that invertebrates are particularly sensitive to the replacement of native habitat and associated plant species since such replacement reveals idiosyncratic responses by a range of invertebrate taxa in different environments. As a result of such replacement, the diversity of several invertebrate groups such as coleopteran, spiders and ants has been found to be lower in native shrub land sites infested with exotic Chrysanthemoides monilifera than in non-weedy sites (Robson, et al., 2009).

Pryke and Samways (2009) found that alien pine plantations in the Table Mountain, Western Cape of South Africa are home to some invertebrate species even though the low species richness and abundance noted in their study showed that alien pine plantations have little or no invertebrate conservation value. However, disturbance caused by pine removal and replacement of fynbos increased invertebrate diversity (Pryke & Samways 2009).

Gerber, et al. (2008) assessed whether plant species richness and invertebrate assemblages in European riparian invaded by exotic knotweeds (Fallopia) differed from those that are found in native grassland or bush-dominated habitats. Their study showed that the riparian habitats invaded by knotweeds support lower numbers of plant species and lower overall abundance and morph-species richness of invertebrates as compared to native grassland-dominated and bush-dominated habitat. Robson, et al. (2009) compared invertebrate assemblages of edges exposed to the intrusion of large amounts of pine leaf litter with plantations and native Eucalyptus in the south-east region of Australia and found that pine plantations had significantly lower invertebrate species richness when compared with native Eucalyptus woodland.

Lowe, et al. (2008) noted that invertebrate assemblages are sensitive to invasion from bass and wattle after investigating the impact of invasive small-mouth bass (Micropterus dolomieu) and black wattle (Acacia mearnsii) on assemblages of macro invertebrates and fish in a Cobble-bed foothill river, in the Rondegat River and in the Cedarburg Mountains of South Africa. Similarly (Albelho & Graca, 2004; Larranga, et al., 2009) found that streams flowing through Eucalyptus plantations support lower taxon richness, density and biomass of invertebrates than streams flowing through native oak forests. Plant invaders Acacia longifolia, Acacia mearnsii, Lantana camara, Solanum mauritanum¸Eucalypus grandis and Pinus patula were used to assess the distribution of epigaeic fauna and the results showed lower species richness and diversity of invertebrates in exotic vegetation when compared to indigenous vegetations (Samways, et al., 1996).

Other studies have also found that exotic plantations support much lower biodiversity when compared to indigenous vegetation (Clark & Samways, 1997; Ratsirarson, et al., 2002; Samways 2005). Reinhart & VandeVoort (2006) further illustrated that replacement of native riparian trees with exotic trees did not only affect the most common family of macro-invertebrates but also affected the common family of predatory macro-invertebrates, which may automatically affect the detrital food web.

Response of invertebrates to vegetation characteristics

Several authors have investigated the response of invertebrates to alien and indigenous vegetation characteristics (Derraik, et al., 2005; Fork, 2010; Harris, et al., 2004; Heleno, 2008; Lovich, et al., 2009; McDonald, 2007; Pryke & Samways, 2009; Strayer, et al. 2003). It has been further illustrated that invertebrate richness and abundance in caves, can be influenced by the nature of the leaf litter introduced (Hills, et al. 2008). Yeates and Barmuta (1999) tested the effect of leaf species (Salix fragilis and Eucalyptus viminalis) and leaf state (senescent or green) on feeding selectivity and growth rates of Notalina sp. (Trichoptera), Koorrnonga sp. (Ephemeroptera) and Physastra gibbosa (Mollusca). They reported that green willow material was a better 10

food source because of the noticeably thicker biofilm that it supported and this willow may also retain high levels of nutrient abscised leaves. Although willow leaves may provide preferred source of food but that food will be available for a shorter period compared to Eucalyptus detritus. Majer, et al. (2003) compared invertebrate abundance and biomass at ordinal level on the trunks of four major species of Eucalyptus in Western Australia and they reported that invertebrate abundance and activity on the bark depended on seasonal moisture and the nature (rough or smooth) of the bark. Some authors have illustrated that the impact of invasive alien plants on invertebrates is not always negative e.g. Oliver, et al. (2006) compared terrestrial invertebrates found under paddock containing alien trees with those found in the surrounding grazed native pastures and found more native invertebrate species under paddock trees when compared with the surrounding pastures. This was because of the increased total litter, percentage of the tree litter, percentage organic carbon, percentage of total nitrogen and extracted phosphorus.

Strayer, et al. (2003) compared macro-invertebrate faunas of alien macrophyte (Trapa natans) with those of native macrophyte (Vallisneria americana) and noted that replacement of Vallisneria by Trapa increased system wide biodiversity and food for fish, even though macro-invertebrates in Trapa beds may not be readily available due to the reduced concentration of oxygen. Woinarski, et al. (2002) assessed how two invertebrate groups i. e. ants and terrestrial spiders respond to the effect of pastoralism and military training in the tropical savanna of north-eastern Queensland. Their study reported very few differences in species richness between undisturbed land and the one which is managed for military use, for both ants and terrestrial spiders. They further noted that invertebrate faunas do not only vary with habitat management regimes but also with natural underlying patterns of environmental variability at local scales. Major environmental gradients structuring invertebrate fauna are landscape position, and topographic variations.

Pryke and Samways (2009) investigated invertebrates response to alien pine plantations, natural vegetation (indigenous forests) and botanical gardens (reserves) around Cape Town, South Africa which is a biodiversity hotspot. Their findings supported the removal of alien pine as pine accounted for low invertebrate species richness and abundance. They further emphasized the conservation value of urban botanical gardens for preserving invertebrate fauna.

Much attention has been paid to negative effects of alien species on resident communities and functioning of invaded ecosystems (Hejda, et al., 2009). However, the presence of these alien species can increase the diversity of invertebrate pollinators since they become preferentially attracted to these alien plants (Proche, et al., 2008).

Rationale and specific objectives of the study

Most invertebrate species are becoming extinct as a result of increasing anthropogenic factors such as habitat loss and alien plant invasions. Therefore understanding the ecological interaction and associations that exist between faunal communities and vegetation characteristics at local habitat patch level remains critically important for biodiversity conservation planning and monitoring within protected areas of the wider O. R. Tambo District Municipality where baseline data is lacking. This preliminary study therefore aims at determining the response of invertebrate assemblages to alien and indigenous vegetation cover in two nature reserves situated within the KSD Local Municipality.

Specific objectives:

 To generate preliminary data on invertebrate assemblage composition and distribution patterns at apriori-selected sites in Nduli and Luchaba Nature Reserves.  To determine the effects of measured environmental variables on invertebrate species distribution patterns

 To propose management recommendations for conserving the invertebrate fauna within protected areas of the KSD Local Municipality based on findings of the study.

CHAPTER 2

MATERIALS AND METHODS

The study site

The study was carried out in Luchaba Nature Reserve (31̊30’S, 28̊42’E) and Nduli Nature Reserve (31̊35’S, 28̊45’E) in the KSD District Municipality, Eastern Cape, South Africa. Nduli Nature Reserve is a valley which is about 3 km south of Mthatha, and lies next to the N2 highway (Fig. 1). Nduli Nature Reserve was originally established in 1951 and re-proclaimed on the 15th of February 1972 in terms of the Cape Nature Conservation Ordinance of 1965. This reserve is now approximately 170 ha. Luchaba Nature Reserve is situated on a State land, adjacent to the Mthatha Dam next to water sports area (Fig. 1). The distance between the two reserves is approximately 10 km. The main reasons for establishment of these reserves were to protect local biodiversity, provide recreational facilities for visitors or tourists and create awareness (through education) of the need to conserve indigenous flora and fauna.

Nduli and Luchaba Nature Reserves have mean annual temperature of 17.6°C and the lowest average monthly temperatures occur from June to August while the highest average monthly temperatures occur during October and between December and February. These reserves have average monthly rainfall of 60 mm and average annual rainfall of 654 mm. In both reserves, the wind direction is South Westerly during all months of the year. The geology of Nduli and Luchaba Nature Reserves comprises predominantly shales and sandstones of the Beaufort series of the Karoo system and these land forms are interlaced with dolerite dykes.

These reserves fall within the Mthatha moist grassland in the grassland biome in Eastern Cape. The primary invasive alien plant species present in these reserves include the black wattle and Lantana, bugweed and inkberry and gum tree. In both reserves, medium sized herbivores are present and these include Blesbok, Black Wildebeest, Blue Wildebeest, Burchell’s Zebra, Fallow Deer, Impala, Mountain Reedbuck, Red Hartebeest 14

and Springbok. Both reserves are also home to a variety of wildlife, a series of wetlands and grasslands that support a wide variety of bird and antelope species. Faunal species that were historically present in these reserves include Oribi, Cape buffalo, Lion, Leopard, Eland, Common Reedbuck, Mountain Reedbuck and Red Hartebeest. These reserves are managed together and they fall within the King Sabata Dalinyebo local Municipality, which in turn falls within the jurisdiction of O.R. Tambo District Municipality (Anon., 2010).

Fig.1. Map showing study locations at Nduli and Luchaba Nature Reserves

Sampling site stratification

There were 2 sites (Fig. 1) selected from each reserve based on the variation in vegetation type. In Luchaba Nature Reserve, 2 alien invasive vegetation patches i.e. the Eucalyptus patch and the mix alien patch comprising of (Acacia auriculiformis and Lantana camara) were selected. In Nduli Nature Reserve 2 sites comprising of indigenous forest patch and indigenous grassland patch were selected. All 4 study sites measuring 20 m² were each stratified into 4 sampling units measuring 5 m² separated from each other by at least 5 m. Sampling units from Luchaba nature Reserve were labeled as, EA, EB, EC, ED whereby E stands for Eucalyptus site, and also labeled as, MA, MB, MC and MD with M stands for Mix alien site. While sampling units from Nduli Nature Reserve were labeled as FA, FB, FC, FD with F stands for Forest site and also as GA, GB, GC and GD with G stands for grassland site.

Invertebrate species sampling

Pitfall trap sampling method was used in sampling invertebrates. This method is one of the most widely employed methods for evaluating terrestrial invertebrate assemblages and it is cheap and easily replicated (Cheli & Corley, 2010). Within each sampling unit, four pitfall traps (plastic cups) with 7.5 cm in diameter and 9.5 cm deep, were sunk into the ground such that the open end of the cup was flushed with the ground surface. These pitfall traps were left open on the ground and half filled with soapy water and then allowed to capture invertebrates for the period of 24 hours. Invertebrate species caught in traps and were sorted, preserved in 70% alcohol and transported to the laboratory for identification, as voucher specimens. Unidentified specimens were sent to taxon experts or Museums of Natural history e.g. Iziko museum in Cape Town and Bloemfontein museum for identification.

Measurement of environmental variables

Corresponding environmental variables were measured e.g. soil variables (soil pH, soil phosphorus, soil potassium and soil zinc), litter depth, grass height, grazing intensity, percentage of alien plants and percentage insolation. Soil variables were measured by analyzing soil samples (top soil) collected at sites, sent to the Mthatha soil analytical services and the Department of Agricultural soil analysis division in Dohne, Eastern Cape.

Environmental variables were: 1) litter depth (lit dep) which was quantified by measuring the distance from the litter surface to the soil surface using a standardized ruler. 2) Grass height (gra hei) was determined by height intervals as short (grass which was foot height =1), medium (grass between the ankle and knee height =2) and tall (grass above the knee height =3). 3) Grazing intensity (gra int) was measured by classifying available dung and degree of trampling as low =1, medium =2 and high =3. 4) Presence of alien plants (%ali veg) was estimated by determining the percentage of total area of surface covered by these plants within each sampling unit. 5) Percentage insolation (% ins) was also estimated by the amount of sunlight that penetrated the sampling unit during the sampling interval. These environmental variables were collected so as to establish the relationships between them and invertebrates caught. Data on soil characteristics were collected once in each of the four seasons of the year while data numbered 1-5 were collected for a period of twelve months from May 2010 to April 2011.

Data analysis

Invertebrate assemblage patterns within sampling units were determined using total number of specimens (individuals) and species present. The data was tested for normality before applying analysis of variance (ANOVA), in STATISTICA. Bray-Curtis similarity measures in PRIMER were used to determine sites and sampling units that share common species. This uses a hierarchical agglomerative technique usually takes a

similarity matrix as their starting point, and successively fuses the samples into groups and the groups into clusters, starting with the highest mutual similarities then gradually lowering the similarity level at which groups are formed and ends with a single cluster containing all samples. Correspondence analysis (CA) in CANOCO (Canonical correspondence analysis) was used to establish relationships (if present) between species and sites, since CA is a multivariate treatment of data through simultaneous consideration of multiple categorical variables (ter Braak, 1986). Canonical correspondence analysis (CCA) in CANOCO was used to explore relationships (if any) between species, sites and environmental variables.

CHAPTER 3

RESULTS

 Overrall taxon richness across sampling sites

The pitfall trap sampling method used in this study was implemented for collecting soil surface dwelling invertebrates, even though opportunistic flying invertebrates were also collected. A total of 2224 invertebrate specimens were collected, and the order Hymenoptera contributed the highest percentage of all specimens collected at 60% (58% Formicidae) while the orders Trichoptera and Neuroptera had the lowest specimens count, at 0.04% (Table 1). Out of the total number of specimens collected, 1976 soil surface-dwelling individuals were sorted into 5 orders, 19 families and 50 species, while 248 flying individuals were sorted into 7 orders, 25 families and 34 species. When considering only soil surface-dwelling invertebrates, the Araneae was the richest order with 23 species in 10 families followed by the Coleoptera with 4 families and 15 species, while the order Stylommatophora was represented by 1 family and 1 species. A total of 585 specimens could not be identified and therefore separated into morpho-species belonging to 15 different orders and these species were excluded in the analysis.

 Spatio-temporal trend in invertebrates distribution across sites

Invertebrate species collected varied across months and sites. Only four invertebrate species: Langona warchalowskii, Pardosa crassipalpis (Araneae), and Camponotus sp., Technomyrmex sp. (Hymenoptera) were collected throughout the year and from all sites except for Langona warchalowskii (Araneae) that was not found in the mix alien site. Invertebrate species collected only from indigenous vegetation sites (forest and grassland) were Streblognathus aethiopicus, Tetraponera sp. (Hymenoptera), Bantua sp. (Blattodea), Anachalcos convexus, Aphodius, Sisyphus sp., Sagrinae sp. and Psammodes bertolonii (Coleoptera) and Cyclosa sp. (Araneae). Three invertebrate species: Messor capensis (Hymenoptera), Hopliini sp. (Coleoptera) and Afropisaura sp. 20

(Araneae) were unique to the Eucalyptus site while five species: Polyrhachis gagates (Hymenoptera), Kheper nigroaenus, Anisonyx ditus, Plagiodera coffra and Sonchia sternalis (Coleoptera) were sampled only at the mix alien site. Some species occurred across all sites e.g. Camponotus sp, Pheidole sp., Technomyrmex sp. (Hymenoptera), Deropeltis erythrocephala (Blattodea) and Pardosa sp., Dysdera crocata, Hyllus argyrotoxus, Xysticus sp., Pardosa crassipalpis (Araneae).

 Effect of site sampling units on invertebrate species richness and abundance

There were no statistically significant differences (p = 0.091482, p = 0.168024, respectively) in species richness and abundance observed across sites for soil surface dwelling invertebrate (Table 2 & 3). However, trends showed that the Eucalyptus site had the highest specimen count while the grassland vegetation site had the least (Fig. 2a). The forest vegetation site had the highest species and specimen count for flying invertebrate followed by Eucalyptus vegetation site (Fig. 2b), even though this trend was not statistically significant (p = 0.540425 richness, p = 0.048663 abundance) (Table 4 & 5).

 Effect of sampling months on species richness and abundance

There were also no significant differences (p > 0.05) observed across months for soil surface dwelling invertebrate specimen abundance, even though the highest specimen counts occurred in January while the lowest counts occurred in June (Fig. 3a). However, there were significant differences (p < 0.05) in species richness across months, with the highest number of species recorded in February while the least species occurred in September (Table 2 & 3 and Fig. 3a). Also, there were statistically significant differences (p <0.05) across months for flying invertebrate species richness and specimen count, (Table 4 & 5). The highest species richness occurred in June while the lowest species richness was in August and April. The highest specimen counts were recorded in July while the least was collected in November (Fig. 3b). Cumulative number of all taxa (order) representatives caught for soil surface dwelling

invertebrates increased sharply from December until February, while a gradual increase was observed for Araneae and Coleoptera. The Hymenoptera (Formicidae) reached an asymptote (levelly off) from January to April (Fig. 4a). A sharp increase from December to March in the cumulative number of all invertebrate species caught from all sites and species from alien as well as species from indigenous vegetation sites, were observed (Fig. 4b). While a gradual increase for invertebrate species caught from alien vegetation sites occured from December to March (Fig. 4b).

 Dominance plots

Cumulative species dominance plots performed for individual sites and across sites showed that Eucalyptus sampling unit (Su) ED from Eucalyptus showed highest species evenness while Su EC showed highest species dominance (Fig.5a). The mix alien site indicated Sus MB and MC to have the highest species evenness and the highest species dominance, respectively (Fig.5b). Su FC was found to have high species evenness while Su FD had the highest species dominance (Fig.6a). The indigenous grassland site showed that Su GD had highest species evenness (Fig.6b). Overall cumulative species dominance plots across all sites indicated that the indigenous grass site had the highest species evenness while alien Eucalyptus site had the highest species dominance of Pardosa crassipalpis (Araneae) and Pheidole sp. (Hymenoptera) (Fig.7).

 Similarity at alien and indigenous vegetation site sampling units

Similar invertebrate distribution patterns were observed using NMDS and CA ordinations. Two clusters were observed at approximately 70 percent level of similarity for alien vegetation sites where Sus EA and EC shared Langona warchalowskii (Araneae) (Fig. 8a,b,c). The second cluster at 60 percent level of similarity extracted similarity amongst Sus EB, ED, MB, MC and MD, with Evarcha sp. (Araneae) and Anoplolepsis custodiens (Hymenoptera) being the dominant invertebrate species. The mix alien site, Sus MA was found to be an outlier with Polyrhachis gagates

(Hymenoptera), Kheper nigroaenus, Plagiodera coffra (Coleoptera) and Theridion sp. (Araneae) found only in this Su.

Indigenous vegetation Sus , showed two similar clusters at 60 percent of similarity (Fig. 9a,b,c). The first cluster comprised of FA, FB, FC and GA that were dominated by Thalassiu sp. (Araneae) and Pachyphaleria capensis (Coleoptera). The second cluster comprised of three Sus (GB, GC and GD) from the same site, sharing species like Deropeltis erythrocephala (Blattodea) and Langona warchalowskii (Araneae). The indigenous natural forest Su (FD) was an outlier with Bantua sp (Blattodea), Carebara vidua (Hymenoptera) and Anachalcos convexus (Coleoptera) found only at this Su. Results for all sampling units (both alien and indigenous vegetation sites combined) extracted three main clusters (Fig. 10a,b,c), with Su from two sites (MA and GB) sharing species such as Dysdera crocata (Araneae) and Deropeltis erythrocephala (Blattodea) at approximately 65 percent level of similarity. The second cluster comprised of Sus EA, EB, EC, ED, MB, MC, MD and GA from three different sites (Eucalyptus, mix alien and grassland). These Sus shared Anoplolepsis custodiens (Hymenoptera), Vallonia sp. (Stylommatophora) and Hydrophilus sp. (Coleoptera) at approximately 62 percent level of similarity . The third cluster at 60 percent level of similarity, comprised of sampling units (FA, FB and FC) from the same site, domminated by Hyllus orgyrotoxus (Araneae).

 CCA for alien and indigenous vegetation sites combined

Litter deposition, grazing intensity and percentage of alien vegetation significantly (p < 0.05) accounted for species distribution trends across alien vegetation sampling units (EA, EB, EC, ED and MB), while soil pH, soil potassium, grass height, percentage shade as well as soil phosphorus significantly (p < 0.05) accounted for species distribution across indigenous vegetation sampling units (Fig. 11). Eigenvalues and cumulative percentage variance of species-environment relationship for both axes 1 and 2 as well as total inertia data sets (Table 8).

Table 1. Number of specimens from each order expressed as a percentage of total caught

Order Total percentage

Hymenoptera 60%

Araneae 26.6%

Diptera 6.7%

Coleoptera 2.7%

Lepidoptera 2%

Blattodea 0.8%

Orthoptera 0.4%

Stylommatophora 0.17%

Hemiptera 0.09%

Trichoptera 0.04%

Neuroptera 0.04%

Table 2. A 2-way ANOVA for species richness of soil surface-dwelling invertebrates, at 5% level of probability

Source of SS df MS F p

variation

Intercept 1111.688 1 1111.688 348.2130 0.000000

Months 109.563 11 9.960 3.1198 <0. 005556***

Sites 22.396 3 7.465 2.3383 0.091482

Error 105.354 33 3.193

Table 3. A 2-way ANOVA for abundance of soil surface-dwelling invertebrate, at 5% level of probability

Source of SS df MS F p

variation

Intercept 79788.52 1 79788.52 54.18815 0.000000

Months 22009.73 11 2000.88 1.35889 0.238058

Sites 7912.40 3 2637.47 1.79123 0.168024

Error 48590.35 33 1472.43

Table 4. A 2-way ANOVA for species richness of flying invertebrate, at 5% level of probability

Source of SS df MS F p

Variation

Intercept 238.5208 1 238.5208 191.8827 0.000000

Months 58.7292 11 5.3390 4.2951 <0.000544***

Sites 2.7292 3 0.9097 0.7318 0.540425

Error 41.0208 33 1.2431

Table 5. A 2-way ANOVA for abundance of flying invertebrates, at 5% level of probability

Source of SS df MS F p

Variation

Intercept 1323.333 1 1323.000 94.50000 0.000000

Months 544.500 11 49.500 3.53571 <0.002375***

Sites 122.500 3 40.833 2.91667 0.048663

Error 462.000 33 14.000

Fig 2a. Spatial trends: Soil surface-dwelling invertebrate abundance and richness across sites during the sampling period (May 2010-April 2011)

Fig 2b. Spatial trends: Flying invertebrate abundance and richness across sites during sampling period (May 2010-April 2011)

Fig 3a. Temporal trends: Soil surface-dwelling invertebrate abundance and richness during the sampling period (May 2010-April 2011)

Fig 3b. Temporal trends: Flying invertebrate abundance and richness during sampling period (May 2010-April 2011)

Fig 4a. Cumulative no of taxa caught for all soil surface-dwelling orders combined, as well as dominant orders (Hymenoptera: Fomicidae, Araneae, Coleoptera)

Fig 4b. Cumulative no of invertebrate species caught for all four sites, and for alien and indigenous vegetation sites separately during sampling from August-April 29

100 EA 100 MA EB MB EC MC ED MD

80 80 %

% e

e c

c n

n a

n i

i m

m o

D 60

D 60 e

v i

e t

i a

l u

l m

u u

m C 40

C 40

20 20 1 10 100 1 10 100 Species rank Species rank a) b)

Fig 5. Cumulative species rank dominance curves for a) Eucalyptus site and b) Mix alien sites

100 GA 100 FA GB FB GC FC GD

90 FD 80

e c

% n

e a

n i

n m

i 80 o

D 60

D e

v i

e t

i a

t 70 l

a u

m u

u C 40

C 60

50 20 1 10 100 1 10 100 Species rank a) Species rank b)

Fig 6. Cumulative species rank dominance curves for a) Forest site and b) Grass indigenous sites

100 E M F G 80

D 60

C 40

20 1 10 100 Species rank

Fig 7. Cumulative species rank dominance curves for all sites (Eucalyptus-E, Mix alien- M, forest-F and grass-G)

Transform: Fourth root Resemblance: S17 Bray Curtis similarity 50 Transform: Fourth root Resemblance: S17 Bray Curtis similarity

2D Stress: 0.06 60

t i MB

r a MC

m 70 i MD

t n 80 ED

r e EB

P EC 90 MA

EA 100

M Sampling sites

a) b)

KhePol nig gag The sp. 1.0 Plag cof MA

Dip gag Der ery Hyd MB Hyl arg Dys cro Ano cus Tha Eva sp. Cam MD Son ste MCAni dit

Par sp. Par cra Val Tec Phe Xys sp. ED Pac cap Zel uqu Sis EB Eva dot Hop Car vid Che law EA Thy aur Lan war Mes capECAfr

-0.6 -0.8 1.0

Fig 8. a) Dendogram, b) NMDS ordinations and c) CA biplot (o-Sampling units and ∆- Species) for alien sites (M- Mix alien, E-Eucalyptus )

Transform: Fourth root Resemblance: S17 Bray Curtis similarity Transform: Fourth root Resemblance: S17 Bray Curtis similarity 2D Stress: 0.06 40 FA

GAFC y GC

i r GB a 60

e FB

c r 80 GD

100 FD

G Sampleing sites

a) b)

1.0 Ban FD Ana con Car vid

Che law

Hog sp. Thy Thy juv Sag Thy aur GD CycFB

Cam Pac cap Tec Eva sp. Dre sp. Phe Tha Par cra Lan war TetFC Xys sp. Zel uqu Der ery FA GB Psa ber Hyl arg Xer cru Par sp. Clu sp. GA Hyd Aph Dys cro Che fur

GC Str aet Gym Sis

-1.0 -0.6 0.8

Fig 9. a) Dendogram, b) NMDS ordinations and c) CA biplot (o-Sampling units and ∆- Species) for indigenous sites (F-Natural forest, G-Grassland)

Transform: Fourth root Resemblance: S17 Bray Curtis similarity

i r

a 60

e c

r 80

e P

100

D C D A B A C B A C D B D B A C

F F F F

E E E E

G G G G

M M M M a) Sampling sites

Transform: Fourth root Resemblance: S17 Bray Curtis similarity

2D Stress: 0.19 FB FD

GA FC MA MD MB MC

GB ED

EB FA EA GD EC

GC b)

FD Ban 0.8 Ana con

Cyc FB Hog sp. Tet FC Dre sp. Aph GA Xer cru Tha Khe nig FA Car vid Pol gag Che law Sag Plag cof MA Clu sp. Hyl argThe sp. Thy GD Hyd Psa ber Thy juv Cam GB Dys cro Tec Eva sp. Der ery Lan war Zel uqu Dip gag MB Phe Che fur Pac cap Par sp. Son ste Par cra Ani dit MDMC Xys sp. Ano cus Val EA Thy aur Eva dot Sis ED Hop EC Str aet Mes cap Gym GC Afr EB

-0.6 -0.6 1.0 c)

Fig 10. a) Dendogram, b) NMDS ordinations and c) CA-biplot (o-Sampling units and ∆- Species) for alien and indigenous vegetation sites (M-Mix alien, E-Eucalypus, F-Natural forest, G-Grassland) 34

Ban FB Cyc

0.8 FD Ana con Zn Ph TetDre sp. Hog sp. FC Clu sp. .sha Car vid Tha FAAph MA Xer cru GA Gym Hyl arg Pol gag GCChe fur Khe nig Plag cof Str aet Son ste MD The sp. GraK hei Ani dit Cam Che law MC Tec Hyd Pac cap Phe Val Zel uqu Sis Dys cro MB Dip gag Par sp. Eva sp. Lit dep Par cra P Der ery Eva dotEA Xys sp. GB Ano cus Lan war Psa ber Gra int ED Sag Hop EC Thy aur Mes cap Thy juv GD Afr EB Thy .Ali veg

-0.6 -1.0 1.0

Fig 11. CCA tri-plot for all four sites (E-Eucalyptus, M-Mix alien, F-Natural forest, G- Grassland), depicting relationship between sites, species and environmental variables, o-Sampling units, ∆-Species and ↗ -Environmental variables

Table 8. Eigenvalues, cumulative percentage variance of species-environmental relation (CCA) for axes 1 and 2 for both alien and indigenous vegetation sites combined Axes 1 2

Eigenvalues 0.151 0.135 Species-environment correlations 0.968 0.976 Cumulative percentage variance of species data 13.6 25.8 Cumulative percentage variance of species-environment relation 20.9 39.5 Total inertia 1.109

Test of significance of first canonical axis: eigenvalue = 0.151 F-ratio = 0.944 P-value = 0.3780

Test of significance of all canonical axes: Trace = 0.722 F-ratio = 1.247 P-value = 0.0420

Table 9. Intra-set correlation for both alien and indigenous vegetation sites combined

Environmental Axis 1 Axis 2

Variables

Litter deposition (Lit dep) -0.4714 -0.1542

Grass height (Gra hei) 0.6274 0.1004

Grazing intensity (Gra int) -0.3772 -0.2897

pH 0.3033 0.5466

Potassium (K) 0.7138 0.1158

Phosphorus (P) 0.1283 -0.1402

Zinc (Zn) -0.0668 0.5955

Percentage alien vegetation (.Ali veg) -0.6341 -0.4955

Percentage insolation (.Sha) 0.728 0.3685

CHAPTER 4

Discussion

 Effects of sampling method on species distribution

Pitfall trapping is frequently used for sampling active ground dwelling invertebrates because of its simplicity, efficacy and low cost (Cheli & Corley, 2010; Nunes, et al., 2011; Sabu & Shiju, 2009). This method is most effective in open habitats with reduced structural complexity such as savannas, dry forests; grassland and scrub vegetation since capturing value can be affected by vegetation complexity (Nunes, et al., 2011; Prasifka, et al., 2007; Sabu & Shiju, 2009). The use of pitfall traps in this study generally enhanced the collection of data on representative taxa of soil surface-dwelling invertebrates across sampling sites.

However, inefficiency in capturing either bottom dwellers or flying invertebrates (since it measures activity abundance rather than density estimates), meant that this sampling method did not provide an absolute measure of invertebrate populations collected across sites used for this study. Furthermore, the resting and evasive behavior of many taxa was not considered, leading to an under representation of such taxa (Nunes, et al., 2011; Sabu & Shiju, 2009). However, because the sampling method was standardized throughout the sampling period with uniform sampling efforts, data collected for this study using pitfall traps could be considered as very representative of the soil surface dwelling invertebrate assemblage across study sites.

Species distribution trends across study sites

Although no statistically significant differences were observed across sites, graphical trends showed that invertebrate species richness and population counts varied across sites. High invertebrate abundance at the Eucalyptus and the mix alien sites may be due to foraging behavior at these resource-poor sites where individuals probably spent

more time searching for food resources as a result of frequent disruptions in food web interactions and energy flow (Gerber, et al., 2008).

Invasive plant communities may not be functionally equivalent to the native plant communities in attracting local faunal assemblages (Mgobozi, et al., 2008). Also, alien plants do not always have a negative impact on native biodiversity as illustrated by Pryke and Samways (2009) who observed that alien pine plantations along the Table Mountain biotope in the Western Cape are home to some invertebrate species even though the low species turnover at this biotope showed that they have little or no invertebrate conservation value. Bonham et al. (2002) found that plantation development on cleared farmland can improve the conservation status of native invertebrates in areas where these plantations adjoin remnant patches of native vegetation as invertebrates in such areas can expand into exotic forest thereby lowering their risk of local extinction. The indigenous Acacia and grassland patches located adjacent to the alien vegetation sites used in this study probably accounted for increased migration for a few opportunistic invertebrate species into alien vegetation patches, resulting in high invertebrate abundance at the alien sites. Invertebrate population can also expand into indigenous grassland thereby increasing their survival rates (Wiezik, et al., 2007). High invertebrate abundance at the Eucalyptus site could also probably be as result of the proximity of this site to the edge of the Mthatha dam with its moist biotope characteristics. Ahrens and Kraus (2007) observed high densities of spiders living near freshwater-terrestrial interface.

Invertebrate species diversity across indigenous vegetation sites can be attributed to the complex habitat structure, higher plant diversity; increased food web interactions and energy flow at these vegetation sites (Mgobozi, et al., 2008; Wiezik, et al., 2007). Supporting this, Gamez-Virues et al. (2010) noted that higher plant diversity contributes to greater herbivore diversity because each plant species has a specific suite of consumers, and high biodiversity is associated with habitat heterogeneity (Myung-Pyo,

et al., 2008). High species richness may increase ecosystem stability, thereby sustaining and maintaining local biodiversity (Wiezik, et al., 2007).

 Species distribution trends across months

Statistically significant differences were observed across months for soil surface dwelling invertebrate richness probably due to species-specific habitat requirements during summer season, while the reverse is true during winter. Some invertebrate taxa (e.g. certain beetles, ants) are influenced by seasonality (rainfall) and temperature (Davis, 2002; Hahn & Wheeler, 2002). Furthermore, Davis (2002) observed maximum local diversity of dung beetles after rainfall that decreased as the soil surface becomes drier. Most beetles occur in terrestrial habitats which are warmer and moist (Davis, et al., 2004; Edward & Aschenborn, 1988). In this study, the beetle assemblages collected comprised mostly of members of the family Scarabaeidae (Anachalcos convexus, Aphodius sp., Hopliini sp., Diplognatha gagates, Kheper nigroaenus and Sisyphus sp.) captured mostly during summer when resource availability was at optimal levels.

Foraging activity of most ants has been found to be positively related with moisture and nutrient availability (Hahn & Wheeler, 2002). Some ant species under the family Formicidae (Camponotus maculates, Carebara vidua, Polyrhachis gagates, Tetraponera sp.) were collected at high population levels during the wet summer season. This trend was also observed by Hahn and Wheeler (2002).

Ahrens and Kraus (2007) noted that some spider species that desiccate more easily are constrained to certain moist microhabitats and thus their movements are limited to those habitats. In the current study, some spider species e.g. Pardosa crassipalpis, Hogna sp. (Lycosidae), and members of the family Salticidae e.g. Thyenula juvenca, Thyenula aurantiaca, Evarcha dotata, Thyene, Heliophanu were collected during the summer period when habitat conditions were wet and warm enough to avoid desiccation. Dippenaar-Schoeman (2002) using pitfall traps at study sites in the Richards Bay area of Kwazulu Natal observed more Lycosidae species from young and

more open forest than from mature (100 year old) forest because physical characters such as the amount of sunshine on the forest floor can be more important than the dominant plant species present. Lycosidae are able to colonise disturbed habitats very quickly because disturbed habitats are usually more open with less diverse vegetation cover (Dippenaar-Schoeman, 2002). This could probably be the reason for high population counts of P. Crassipalpis and Hogna sp. at alien vegetation sites in this study. Spiders regulate their body temperatures during winter season by using leaf litter as insulation from cold temperature (Cramer & Maywright, 2008; Kraus & Morse, 2005). In this study spider species caught during winter months included Afropisaura sp., Thalassium sp. (Pisauridae) and Cyclosa sp. (Araneidae). Lower temperatures conditions have been shown to constrain the number and distribution of many spider species living in cold conditions (Cheli, et al., 2010).

The study illustrated that the summer month of January was the most reliable period for sampling soil surface dwelling invertebrates in Nduli and Luchaba Nature Reserves. Nchai (2008) also used observed high ground-dwelling invertebrate richness and abundance in summer and the reverse during winter in the Western Cape of South Africa using pitfall traps. Hahn and Wheeler (2002) observed a similar trend in Panama using the baiting method, with high insect (ant) diversity during the wet summer season.

 Relationship between species, sites and environmental variables

 Effect of litter depth

Litter depth was found to be an important variable influencing the distribution of invertebrates across the Eucalyptus and Natural forest sites. High abundance of invertebrates at these sites could probably be due to leaf litter depth used by some species as growth substrate for egg-laying and shelter from predators and desiccation (Kazemi, et al., 2009; Magura, et al., 2004). Wiezik, et al., (2007) observed an increase in the diversity of ground-dwelling beetles due to high habitat complexity through

gradual accumulation of litter and deadwood at various levels of decomposition as well as increased canopy complexity leading to deep litter layers. Furthermore, these litter layers offer increased resources and structural complexity, microclimate diversity, refugia for protection from predators, increasing prey availability and also providing a buffering effect against temperature fluctuations (Magura, et al., 2004; Oliver, et al., 2006; Oxbrough, et al., 2010; Robson, et al., 2009). Halaj et al. (2008) noted that some invertebrate species such as molluscs are typically adapted to survive in cooler and moister habitats, and this phenomenon may explain why Vallonia sp. was found only in the forest site in this study. Wenninger and Inovye (2008) also found that moisture may influence the distribution of some invertebrate species. Hills et al. (2008) noted that the richness and abundance of invertebrate assemblages can be influenced by the nature of leaf litter. Litter quality probably also accounted for variation in faunal richness and abundance trends observed in this study.

 Effect of grazing intensity

Grazing intensity influenced the distribution of invertebrate species either positively or negatively depending on grazing pressure. Invertebrate distribution across alien vegetation sites was found to be influenced by grazing intensity. Grazing at very high densities by livestock or human activity can reduce plant diversity leading to the reduction of faunal diversity due to exposure to predators, thereby affecting their distribution ranges in such habitats (Allombert, et al., 2005; Cheli, et al., 2010; Souminan & Olofsson, 2000). Allombert et al. (2005) noted that gastropod diversity and abundance can be reduced by deer grazing. This form of grazing reduces biomass of herbaceous vegetation, thereby reducing the food resource available to gastropods. However, abundance of invertebrates across alien vegetation sites can probably also be as a result of animal grazing. Since, twigs and leaves browsed by mammalian herbivores increase herbivorous insects on twigs; due to changes in the chemistry, morphology and growth rate of the plant tissue (Cheli, et al., 2010).

 Effect of alien vegetation

High percentage of alien vegetation influenced invertebrate assemblage composition and distribution trends as noted by low species diversity observed in the Eucalyptus site. Robson et al. (2009) noted that alien pine plantation sites are simple and less diverse than non-plantation sites and tend to harbour lower invertebrate assemblages than in non-plantation sites. The low species richness at the Eucalyptus site compared to the natural forest site, observed in this study, can also be due to the fact that invasive trees are pest free in introduced areas since they are waxy, robust and have fibrous leaves which are physically resistant to herbivores. They have low nutritional value, and in most cases tend to have leaves that support mutualistic and entomopathic fungal associations as well as foster antibiotic responses (Holmoquist, et al., 2011). Additionally, low species richness in alien vegetation sites can probably be due to the fact that indigenous insect herbivores tend not to feed extensively on invasive plants because those plants contain flavonoids that may deter generalist feeders (Standish, 2004). Distribution of food resources at varying quality is responsible for structuring insect communities (Wenninger & Inouye, 2008). Standish (2004) noted that the invasive plant, Tradescantia has a negative impact on active epigaeic invertebrates resulting in decreased abundance and richness of invertebrates on this plant.

Wiezik et al. (2007) using the dry sieve method, found high species richness of ground- dwelling invertebrates in native forest ecosystems than semi-natural managed forests and plantations because of high habitat complexity which resulted in increased availability of micro-habitat, food, and shelter. Mgobozi et al. (2008), using pitfall trap and vegetation beating methods, observed lower abundance and species richness in sites invaded by Chromolaena odorata due to simpler habitat structure and lower plant diversity comparing to sites that are not invaded as well as sites that are newly invaded. However, the impact of alien vegetation plants on native biodiversity does not necessary have to be negative (Harris, et al., 2004). Furthermore, Parr et al. (2010) found that an invasive grass (Andropogon gayanus) had no effect on richness and

abundance of ant, spider and other invertebrates. Statistically, no significant impact was observed on soil surface dwelling invertebrate richness across alien and indigenous vegetation sites, even though this trend may change for individual taxa or for all taxa sampled at different spatial and temporal scales.

Afrin et al. (2010), Brunel et al. (2010) and Heleno et al. (2009) observed that replacement of native plants by alien plants is likely to affect other trophic levels, particularly phytophagous insects. Furthermore, invasive alien plants significantly affect native herbivores by outcompeting native plants and therefore causing a reduction in the resources available to native herbivores and higher trophic levels which feed on herbivores (Heleno, 2008; Maerz, et al., 2005). This can re-structure communities and also lead to evolutionary changes (Rodriguez, 2006). Alien plants such as Acacia and Eucalyptus reduce the amount of nutrients and water available to native species; alter soil chemistry and soil moisture-holding capacity (Afrin, et al., 2010). Also, Derraik et al. (2005) noted that replacement of a native shrub (Olearina bullata) cover by invasive plants lead to local extinction of some ground-dwelling invertebrate species.

 Effects of soil characteristics

This study further showed that soil constituent characteristics (pH, zinc, phosphorus and potassium), percentage insolation and grass height influenced invertebrate distribution across indigenous sites. Kazemi et al. (2009) noted that different invertebrate species associate with specific soil pH ranges due to their degree of vulnerability and resistance to acidity or alkalinity of the soil and to the availability of their required nutrients. High soil acidification in pine forest has been observed to have negative effect on invertebrate diversity. Acidic soils have less nutrients available thereby providing less suitable environments for invertebrates (Kazemi, et al., 2009).

Indicator potential of invertebrate taxa sampled during the study period

Several soil surface invertebrate species were found to be potentially sensitive to sites from which they were collected. The Araneae followed by the Coleoptera constituted 44

the highest number of soil surface-dwelling invertebrate species collected during the study period. Spiders have been used extensively as ecological indicators both in South Africa and other parts of the World because they are easily collected using pitfall traps, are abundant in most habitat types, with large numbers of species, and are important components of healthy terrestrial ecosystems (Cardoso 2009; Dippenaar-Schoeman, 2006; Myung-Pyo, et al., 2008; Pearce & Venier, 2006).

In South Africa, Lowes et al. (2005) noted that beetles can be used as ecological indicators because they play an important role in maintaining ecosystem processes such as nutrient cycling and are sensitive to environmental changes as they rely on the resources provided by organic leaf litter. The Coleoptera comprised about 40% of all insect species and have been used as indicators of disturbance and habitat quality, particularly in the tropics and subtropics (Clark and Samways, 1997; Stewart, et al., 2007).

The study showed that snails (Stylommatophora) had the lowest number of species. The activity of snails depends on weather conditions and other environmental variables (Matilda, et al., 2011). These variables were probably not optimal at sites used for this study, therefore accounting for the rarity of invertebrate group.

In general, invertebrate species collected during the study varied across months and sites. Only four invertebrate species were collected throughout the year and from all sites except for Langona warchalowskii (Araneae) that was not found in the mix alien site. Nine invertebrate species were collected only from indigenous vegetation sites (forest and grassland) while three and five species were unique to the Eucalyptus site and mix alien site respectively.

Some species occurred across all sites and therefore have good potential for use as bio- indicators e.g. Camponotus sp, Pheidole sp., Technomyrmex sp. (Hymenoptera), Deropeltis erythrocephala (Blattodea) and Pardosa sp., Dysdera crocata, Hyllus argyrotoxus, Xysticus sp., Pardosa crassipalpis (Araneae).

 Conclusion

Soil surface dwelling invertebrate species richness and distribution trends were influenced by seasonality as well as habitat, complexity and heterogeneity across sites. Species-rich taxa belonged to the orders Araneae, Coleoptera and Hymenoptera (Formicidae). Some measured site variables were important in determining species distribution patterns across sites e.g. litter depth, percentage of alien vegetation, grazing intensity, grass height, percentage isolation, soil characteristics (pH, zinc, phosphorus, potassium). Invertebrate species richness at the indigenous vegetation site (Natural forest) was probably due to its complex habitat structure. Despite the low habitat quality of the Eucalyptus alien vegetation site, high invertebrate abundance was observed from this site for a few taxa e.g. Pardosa crassipalpis (Araneae) probably because of increased migration of representative invertebrate species populations from adjacent indigenous Acacia and grassland patches. The response of invertebrate species to alien and indigenous vegetation sites used for the study was species or taxon specific. Certain rare invertebrate species (e.g. Vallonia sp. (Stylommatophora) Gryllus bimaculatus (Orthoptera), Bantua sp. (Blattodea), Proctarrelabis capensis (Neuroptera), Cimex lectulerius (Hemiptera), Dipseudopsis sp. (Trichoptera)) were found only at the indigenous natural forest site probably because of the cool and moist habitat characteristics associated with this site.

The different invertebrate individuals associated with alien and indigenous vegetation cover in this study suggested that some individuals were good indicators of exotic or indigenous vegetation characteristics (Samways, et al., 1996).

 Conservation recommendations

Given that many protected areas in South Africa (e.g. Kruger National Park and Camdeboo National Park) are invaded by invasive plants (Foxcroft, et al., 2011; Masubelele, et al., 2009; Pauchard & Alaback, 2004) with adverse consequences on their invertebrate faunal composition and distribution. This study therefore:

 Provides preliminary baseline information for raising community awareness through education about invasive alien plants within protected areas of the KSD Local Municipality, their effects on invertebrate assemblages and the need to eradicate them.

 Recommend that a variety of indigenous natural vegetation types in Luchaba and Nduli Nature Reserves be preserved, to provide high quality habitat patches and microhabitat conditions for various growth stages of invertebrate taxa. This can be achieved in various ways including replacing invasive with indigenous plant species as well as reducing the rate of spread introduction of invasive plant species into these reserves from reserve edges (Pauchard & Alaback, 2004).

 Recommend implementation of the Coarse and fine filter approaches for conserving species-rich taxa (e.g. members of the orders Araneae, Coleoptera, Hymenoptera (Formicidae) and rare species (e.g. Vallonia sp. (Stylommatophora) Gryllus bimaculatus (Orthoptera), Bantua sp. (Blattodea), Proctarrelabis capensis (Neuroptera), Cimex lectulerius (Hemiptera), Dipseudopsis sp. (Trichoptera)) respectively in Nduli and Luchaba Nature Reserves.

 Hunter (2005), however, noted that the coarse filter approach protects a representative array of natural ecosystems and their constituent processes, structure and species, even though not all species are conserved in this approach. Hence the fine filter can also be used to cater for rare and highly exploited invertebrate species.

 Further recommends the use of higher invertebrate taxonomic levels (e.g. order, family) as surrogates for conservation planning given the current taxonomic impediment of identifying most invertebrate specimens to species level. Also historical voucher depositions can be established to standardize the use of morpho-species in future invertebrate inventory studies (New, 1999).

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Appendix 1. Full names of orders, families, species and acronyms of soil surface- dwelling invertebrates collected

Orders Families Species Acronyms Hymenoptera Formicidae Anoplolepsis custodiens Ano cus

Hymenoptera Formicidae Carebara vidua Car vid

Hymenoptera Formicidae Technomyrmex sp. Tec

Hymenoptera Formicidae Messor capensis Mes cap

Hymenoptera Formicidae Pheidole sp. Phe

Hymenoptera Formicidae Polyrhachis gagates Pol gag

Hymenoptera Formicidae Streblognathus aethiopicus Str aet

Hymenoptera Formicidae Tetraponera sp. Tet

Hymenoptera Formicidae Camponotus sp. Cam

Blattodea Blaberidae Bantua sp. Ban

Blattodea Blattidae Deropeltis erythrocephala Der ery

Coleoptera Scarabidae Anachalcos convexus Ana con

Coleoptera Scarabaeidae Aphodius sp. Aph

Coleoptera Scarabaeidae Hopliini sp. Hop

Coleoptera Scarabaeidae Diplognatha gagates Dip gag

Coleoptera Scarabaeidae Kheper nigroaenus Khe nig

Coleoptera Scarabaeidae Anisonyx ditus Ani dit

Coleoptera Scarabaeidae Sisyphus sp. Sis

Coleoptera Scarabaeidae Gymnopleurus sp. Gym

Coleoptera Hydrophilidae Hydrophilus sp. Hyd

Coleoptera Chrysomelidae Plagiodera coffra Pla cof

Coleoptera Chrysomelidae Sagra sp. Sag

Coleoptera Chrysomelidae Sonchia sternalis Son ste

Coleoptera Tenebrionidae Psammodes bertolonii Psa ber

Coleoptera Tenebrionidae Pachyphaleria capensis Pac cap

Araneae Pisauridae Cyclosa sp. Cyc

Araneae Pisauridae Afropisaura sp. Afro

Araneae Pisauridae Thalassius sp. Tha

Araneae Miturgidae Cheiracanthium lawrencei Che law

Araneae Miturgidae Cheiracanthium furculatum Che fur

Araneae Cornnidae Dysdera crocata Dys cro

Araneae Salticidae Hyllus argyrotoxus Hyl arg

Araneae Salticidae Thyene sp. Thy

Araneae Salticidae Langona warchalowskii Lan war

Araneae Salticidae Evarcha dotata Eva dot

Araneae Salticidae Evarcha sp. Eva

Araneae Salticidae Thyenula juvenca Thy juv

Araneae Salticidae Thyenula aurantiaca Thy aur

Araneae Lycosidae Pardosa crassipalpis Par cra

Araneae Lycosidae Pardosa sp. Par

Araneae Lycosidae Hogna sp. 1 Hog sp. 1

Araneae Gnaphosidae Zelotes uquathus Zel uqu

Araneae Gnaphosidae Xerophaeus crustosus Xer cru

Araneae Clubionidae Clubiona sp. Clu

Araneae Theridiidae Steatoda capensis Ste cap

Araneae Theridiidae Theridion sp. The

Araneae Eresidae Dresserus sp. Dre

Araneae Thomisidae Xysticus sp. Xys

Stylommatophora Valloniidae Vallonia sp. Val

Collected morph-species belong in these groups:

Order: Neuroptera

Order: Archeognatha

Order: Hemiptera

Order: Dermaptera

Order: Isoptera

Order: Trichoptera

Order: Lepidoptera

Order: Orthoptera

Order: Diptera

Subphylum: Myriapoda

Subclass: Collembola

Appendix 2. Captured flying invertebrate orders, families and species names

Orders Families Species

Diptera Syrphidae Microdon testaceus

Diptera Syrphidae Allograpta fuscotibialis

Diptera Asilidae Damalis heterocera

Diptera Ephydridae Ochthera

Diptera Lauxaniidae Homoneura

Diptera Calliphoridae Lucillia

Diptera Muscidae Stomoxy calcitrans

Diptera Rhiniidae Rhyncomyia

Lepidoptera Nymphalidae Acraea horta

Lepidoptera Hesperridae Leucochitonea levubu

Lepidoptera Hesperridae Ancylotrypha

Lepidoptera Lycaenidae Cacyreus marshali

Lepidoptera Lycaenidae Chrysoritis

Lepidoptera Lycaenidae Ornipholidotos peucetia

Lepidoptera Geometridae Chiasmia simplicilinea

Lepidoptera Geometridae Zerenopsis lepida

Lepidoptera Arctiidae Thyretes coffra

Orthoptera Acrididae Oedaleus

Orthoptera Acrididae Nomadacris

Orthoptera Acrididae Eyprepocnemis plorans

Orthoptera Grylloidae Brachytrupes membranaceus

Neuroptera Ascalaphidae Proctarrelabis capensis

Hemiptera Cimicidae Cimex lectulerius

Hemiptera Pentatomidae Coridius nubilis

Trichoptera Dipseudopsidae Dipseudopsis

Hymenoptera Braconidae Bathyaulax

Hymenoptera Braconidae Hylaeus heraldicus

Hymenoptera Crabronidae Bembix

Hymenoptera Crabronidae Philanthus triangulum

Hymenoptera Crabronidae Dasyproctus bipunctatus

Hymenoptera Megachalidae Megachile chrysorrhoea

Hymenoptera Apidae Thyreus delumbatus

Hymenoptera Sphecidae Ammophila ferruginapes

Hymenoptera Sphecidae Chalybion tibiale

Hymenoptera Sphecidae Chalybion sp.

Appendix 3. Data for environmental variables

SUs Lit dep Gra hei Gra int pH K Zn % Alie veg % Ins

EA 3.1 1 1 4.88 186.75 19.5 54 50

EB 3.3 1 2 3.15 168.5 27 54 33

EC 5 1 1 3.91 194.25 14.75 80 72

ED 4.1 1 1 4.01 161.5 17 71 68

MA 0.6 2 1 3.5 231 34.25 76 70

MB 0.4 2 2 3.12 166.5 22.75 70 62

MC 0.5 2 2 3.15 179 24 68 63

MD 0.5 2 1 2.87 163.75 21.5 72 67

FA 1.8 1 1 4.76 232.75 12 5 85

FB 3.8 1 1 5.55 194.25 21.75 0 80

FC 2 2 1 6.45 231.5 20.75 0 67

FD 2.4 2 1 5.93 217.25 18.5 0 71

GA 0 3 1 4.29 347.75 17.5 0 87

GB 0 3 1 4.65 391.75 21.5 0 89

GC 0 3 1 4.53 381.25 15.5 0 85

GD 0 3 1 4.53 273.25 29.75 0 89