Bracken (Pteridium aquilinum) is a cosmopolitan species of fern which possesses extrafloral nectaries. A survey of the arthropod fauna associated with the plant throughout its range in South Africa identified the most widespread and damaging bracken herbivores and those species most likely to be influenced by ants visiting the extrafloral nectaries. Seventeen herbivorous arthropods were found to be definitely feeding on bracken and a further thirteen species with a less certain status were recorded. In addition, several species of ants were observed on the fronds and feeding at the extrafloral nectarie. On the basis of their widespread distribution and abundance, four bracken herbivores emerged as being particularly detrimental to the plant in South Africa. They were an eriophyid gall mite, a leafhopper and two moths. The biology of the two lepidopterans, Appana cinisigna and Panotima sp., suggested that they were potentially vulnerable to ants at various stages of their life histories. Laboratory experiments were undertaken to observe ant-lepidopteran interactions under controlled conditions. The ant Crematogaster peringueyi rapidly removed exposed A. cini igna eggs, but did not appear to regard Panotima eggs as food items. Small instar larvae of both species proved vulnerable, but the larger larvae appeared capable of escaping ant predation. Ant exclusion experiments in the field, using natural and artificially augmented ant densities, were carried out at two sites near Grahamstown. The dominant ant species was C. peringueyi. Neither the numbers of lepidopterans nor levels of herbivory were significantly reduced by the presence of ants . Despite the implications of the laboratory tests, the results of these field experiments did not support the hypothesis that ants which visit bracken extrafloral nectaries benefit the plant. Ant-bracken field studies on other continents also reported no significant ant-related effects, though marginal reductions in the abundance of certain herbivores have been noted at one site in the U.K. Since ant protection does not appear to be an inevitable consequence of having extrafloral nectaries, their value to bracken is in some doubt. The most likely situation where effective protection will occur is when high densities of vulnerable herbivores are preyed upon by large numbers of aggressive ants. However, even under these conditions, enhancement of plant fitness is not inevitable. This raises the question of why extrafloral nectaries have been retained in a plant that is as successful and widespread as bracken.

Several ant species may be associated with the same honeydew producing insect in citrus orchards. Recently antagonism and competition between two such ants were reported from Letaba. This report stimulated the present study, the purpose of which is to ascertain which ant species are most numerous in citrus orchards ln the Eastern Cape Province; the reasons for these ants being able to occupy orchards; and, finally, their relationships. A survey of the Sundays River Valley and the Fish River Valley revealed that three species - Anoplolepis custodiens Smith, Anoplolepis steingroeveri Forel, and Pheidole megacephala Fabricius - were commonly found in citrus orchards in association with Soft Brown Scale insects. The territorial relationships of these ants were determined by plotting their distribution at intervals for over a year. It was found that this distribution could be correlated with seasonal climatic conditions, the effects of farming practices, brood rearing requirements and behavioural differences between the three species. The ant, A.steingroeveri, had not been previously studied, and since foraging is an integral part of territorial behaviour, some aspects of its biology, and more especially its foraging activity, were investigated to compare it with the known biology of the other two species. Observations on the foraging activity of A. steingroeveri were made at monthly lntervals for almost a year, and correlated with climatic conditions prevailing at the time. Some seasonal and dally variations in foraging behaviour could not be readily explained in terms of the effects of temperature and humidity on the ants. Other factors, includ1ng trophallactic stimulation and overcast skies, were found to influence ant activity. Owing to the apparent lack of information in the literature and possibility of periodic honeydew production by Soft Brown Scale insects controlling ant activity was investigated in the laboratory. The results of this study showed that it was not likely that honeydew excretion restricted the activity of the ants. Several investigators have also found that many interrelated climatic factors correlated with ant foraging behaviour, but have expressed dissatisfaction at not being able to reach the crux of the matter. To clarify the issue a constant temperature chamber was constructed in which a colony of Crematogaster ants was housed and the ants were subjected to a series of temperatures at different humidities, all other factors were eliminated or kept constant. The effects of these two factors on the thermokinetics and foraging activity of the ants were registered. This thesis contains several sections and at the end of each the results are discussed in the light of the available literature and comparisons are drawn between the species under investigation. The main results are given in the summary. After the summary two appendices are included. One is an attempt to compare the effect on ant size of the honeydew diet of orchard colonies with the proteinaceoue diet of veld ants. The other appendix is an experiment to control citrus ants with an insecticide.

MEnvSC / Department of Zoology

PhDENV / Department of Ecology and Resource Management / See the attached abstract below

Thesis (MSc)--Stellenbosch University, 2011. / ENGLISH ABSTRACT: Since its introduction in 1898 into South Africa, the Argentine ant, Linepithema humile [Mayr 1868 (Hymenoptera: Formicidae)], has invaded human-occupied areas (i.e. urban and agricultural areas) and natural areas characterised by few, if any, anthropogenic disturbances. However, compared to other countries in which the Argentine ant has been recorded, and until the past few decades, very little research had been done on this invasive ant in South Africa. Consequently, several issues concerning its ecological and social effects are still under-researched. The first of these issues concerns the lack of knowledge about the distribution of the Argentine ant in the natural areas, particularly the protected areas (PAs), of South Africa. In order to determine how many PAs are occupied by this invasive ant, a study was conducted in the Western Cape Province (WCP). It was found that, of the 614 PAs documented for WCP, ten have a known presence and nine known absence records of the Argentine ant. The remainder of the PAs have no known occupancy records for this ant. A second issue concerns the seasonal bait preference of the Argentine ant in a fynbos habitat. Six bait treatments (two carbohydrate and protein baits, a mixture of the carbohydrate and protein treatments, and a control) were applied in a fynbos habitat across a sampling grid in four different Latin Square designs, i.e. once for every season. Based on these experiments, it was determined that the Argentine ant prefers the mixture of carbohydrate and protein treatments, and that this preference does not change according to season. Furthermore, previous studies conducted in Jonkershoek Nature Reserve [JNR (in WCP)] determined the existence of a distribution boundary of Argentine ants in an area known as Swartboschkloof. Therefore, the third issue concerned the exact location of the distribution boundary and possible reasons for its establishment. This distribution boundary of the Argentine ant was found to be present 450 m from Swartboschkloof hiking trail. A combination of several explanatory variables may contribute to the maintenance of this boundary, i.e. a change in the horizontal and vertical vegetation distribution, as well as in the slope and aspect across the distribution boundary. With these explanatory variables, the increasing presence of an indigenous ant species, Anoplolepis custodiens, from this boundary may also have contributed to the distribution boundary. In addition, the short-term effect of a fire that swept through this area was also assessed, which revealed that Argentine ants are severely negatively affected by fire (at least over the short-term), i.e. their abundance decreased considerably after the fire and their local distribution range contracted. The final issue concerns the public perceptions of invasive alien species (IAS) in general and the Argentine ant specifically, at JNR. This study revealed that the majority of visitors to JNR were aware of the presence of IAS in South Africa and in its PAs, while very few visitors knew about the Argentine ant. This study also revealed that future research concerning South Africans perceptions of IAS might play a strong contributing role in conservation. / AFRIKAANSE OPSOMMING: Sedert die Argentynse mier, Linepithema humile [Mayr 1868 (Hymenoptera: Formicidae)], in 1898 in Suid-Afrika ingevoer is, het dié mier mens-bewoonde gebiede (soos stedelike en landbou gebiede) en natuurlike gebiede, gekenmerk deur min, indien enige, antropogeniese versteuringe, binnegedring. Nietemin, in vergelyking met ander lande waar die Argentynse mier opgeteken is, en tot die onlangse paar dekades, is min navorsing oor hierdie indringermier in Suid-Afrika onderneem. Gevolglik is daar verskeie kwessies rondom die mier se ekologiese en sosiale uitwerkinge wat nog nie nagevors is nie. Die eerste kwessie het betrekking op die gebrek aan kennis oor die Argentynse mier se verspreiding in die natuurlike gebiede, veral die beskermde gebiede (BG), van Suid-Afrika. Om te bepaal hoeveel BG deur hierdie indringermier beset word, is ʼn ondersoek in die Wes-Kaapprovinsie (WKP) uitgevoer. Daar is bevind dat, van die 614 BG gedokumenteerd in die WKP, het tien bevestigde aanwesigheid- en nege bevestigde afwesigheidrekords van hierdie mier. Die oorblywende BG het geen bekende besettingsrekords van hierdie mier nie. ʼn Tweede kwessie het betrekking op die seisoenale lokaasvoorkeur van die Argentynse mier in ʼn fynbos habitat. Ses lokaas-behandelings (twee koolhidraat en proteïen lokaas, ʼn mengsel van die koolhidraat en proteïen behandelings, en ʼn kontrole) is aangewend in ʼn fynbos habitat, oor ʼn steekproefruitgebied, in vier verskillende Latyns-kwadraatpatrone (“Latin Sqaure designs”), d.i. een vir elke seisoen. Op grond van hierdie eksperimente is vasgestel dat die Argentynse mier die mengsel van koolhidrate en proteïne verkies, en dat hierdie voorkeur nie seisoenaal verander nie. Boonop, vorige ondersoeke wat in die Jonkershoek Natuurreservaat [JNR (in die WKP)] uitgevoer is, het ʼn verspreidings-grens van Argentynse miere ontdek in ʼn gebied bekend as Swartboschkloof. Gevolglik het die derde kwessie betrekking op die presiese ligging van hierdie grens en moontlike redes waarom dit gevestig het. Dié verspreidings-grens van die Argentynse mier is 450 m vanaf die Swartboschkloof voetslaanpad gevind. ʼn Kombinasie van verskeie verklarende veranderlikes kon tot hierdie grens bygedra het, d.i. ʼn verandering in die horisontale en vertikale plantegroeiverspreiding, sowel as in die helling en ligging oor die verspreidings-grens van die Argentynse mier. Tesame met hierdie verklarende veranderlikes, kon die toenemende teenwoordigheid van ʼn inheemse mier, Anoplolepis custodiens, vanaf hierdie grens ook tot die verspredings-grens bygedra het. Daarbenewens is die korttermyn-effek van ʼn vuur wat deur die area beweeg het, ook bestudeer. Die ondersoek het getoon dat die Argentynse mier (ten minste oor die korttermyn) erg negatief deur vuur beïnvloed is, d.i. hul volopheid het ná die vuur aansienlik verminder en hul plaaslike verspreidings-grens het gekrimp. Die finale kwessie het betrekking op openbare persepsie van uitheemse indringerspesies (UIS) oor die algemeen en spesifiek die Argentynse mier, by JNR. Hierdie ondersoek het aan die lig gebring dat die meerderheid van besoekers aan JNR bewus was van die teenwoordigheid van UIS in Suid-Afrika en in dié se BG, terwyl baie min egter van die Argentynse mier geweet het. Hierdie ondersoek het ook aan die lig gebring dat toekomstige navorsing rakende Suid-Afrikaners se persepsie van UIS ʼn sterk bydra tot bewaring kan maak.

Two ant species, Pheidole megacephala (Fabricius) and Anoplolepis custodiens (Smith) reach pest status in citrus orchards through precipitating outbreaks of homopterous pests. However, predacious ants, including these two ant species, play an important role in pest suppression in agroecosystems and are therefore beneficial to these systems. If A. custodiens and P. megacephala are important natural control agents in citrus, using ant bands to break the mutualism between the ants and the Homoptera as a method of ant control is preferable to poisoning. Ant communities were sampled by pitfall trapping in three experimental subunits of 2-, 4-, 15- and 30-year-old citrus orchards, in the Grahamstown area of the Eastern Cape. In one subunit in each orchard, populations of P. megacephala and A. custodiens were suppressed by poison applications. In a second subunit, trees were banded with trunk barriers so that ants were prevented from foraging in the trees and a third subunit served as the untreated control. Bait pupae of bollworm, false codling moth and fruit fly were planted in bait trays in all of the subunits to investigate predation on these citrus pests in the relative absence of predacious ants and where they were excluded from the trees. Pheidole megacephala dominated exclusively in all of the plots. Community composition did not change dramatically with increasing age of the trees, but species diversity and species abundance did. Rank-abundance curves showed that community diversity was highest in the 2-year-old plots and lowest in the 30-year-old plots. The Simpson and Shannon-Wiener diversity indices and their evenness measures indicated that diversity and equitability were highest in the poisoned subunits and lowest in the banded subunits. Principle component analysis revealed that the poisoned subunits were similar and distinct in species composition, that there was significant monthly variation in species composition and that community stability increases with an increase in orchard age. The presence of P. megacephala was significantly negatively correlated (rs = -0.293; p < 0.001) with pest pupal survival. Pupal survival was significantly higher for bollworm (p < 0.001), FCM (p < 0.001) and fruit fly (p < 0.001) in the poisoned subunits, than in the banded and control subunits. There was a general trend for survivorship to increase with an increase in the age of the trees. A significant difference (p < 0.001) was found between the months in which the trials were carried out. Pupal survival was significantly lower (p < 0.001) for FCM than for bollworm and fruit fly. In citrus orchards, ant communities are organised by ecological processes and interactions and are influenced by methods of ant control. Ant bands are preferable to poisoning as a method of ant control, so that beneficial species are left on the ground to prey on pests that pupate in the soil. Maintaining high ant species diversity in citrus orchards would be beneficial as predation on the pupae was more effective where ant species diversity was higher.

MSc (Zoology) / Department of Zoology / See the attached abstract below

The structuring of ant assemblages in a Southern African savanna was investigated using data from the only long-term, large-scale savanna fire experiment in Africa. A comprehensive survey of three habitats in the Kruger National Park (KNP) revealed a total of 169 ant species from 41 genera. The sampling efficiency and consistency of pitfall traps and Winkler samples for inventory, bioindicator and ecological studies in savanna habitats was compared using ants. Pitfall traps were more efficient and productive than Winkler sampling for epigaiec ants, with a greater total species richness and higher abundance of ants recorded. Suggestions were made to improve Winkler sampling output, and to allow quantitative data to be collected. With the structuring of local assemblages, competition was the most significant local factor tested. The relationship between ant dominance and ant species richness was consistent across three continents. A model developed to test mechanisms that could be responsible for the form of this relationship supported the hypothesis that competitive exclusion by dominant ants at least partially reduces species richness. Stress was only partially responsible for low dominance and low species richness, while scatter in the data points is related to patchiness of ants at baits. These findings contrast strongly with previous claims regarding the relationship between richness and dominance. Habitat complexity was not found to play an important role in determining ant assemblage body size in this savanna system. The size-grain hypothesis (Kaspari&Weiser 1999) which predicts that environmental rugosity results in positive allometric scaling of leg length on body length because of changes in locomotion costs, was tested by comparing the body sizes of ants from areas of contrasting habitat complexity. No support for the hypothesis was found. Phylogenetic independent contrast methods did however support the allometric relationship found by Kaspari and Weiser (1999). Ant assemblages in KNP exhibited a remarkable degree of resistance, and in some cases resilience, to burning. Species richness or abundance did not vary with different burning treatments, although ant assemblage composition was sensitive to burning treatment. This difference, however, was only pronounced between burnt and unburnt plots, not between burning treatments. The degree of response of ant assemblages is likely to be related to two main contributory factors: mean annual rainfall and changes in vegetation structure with burning, and the assemblage's history of association with fire. An overview of published research on the effects of fire on fauna in Southern Africa was undertaken. Few studies have examined the effects of fire on amphibians or reptiles and few experimental studies have been undertaken using an experimental fire regime applied over appropriately long time intervals. Most studies provided no information on the scale of the study. Replication was often not reported, and was generally inadequate. Information on the effects of fire on fauna in Southern Africa is fragmentary, and consequently informed management decisions regarding the consequences of burning policies on the conservation of biodiversity both within and outside protected areas are problematic. Recommendations and suggestions for improving fire research are given. / Thesis (DPhil (Zoology))--University of Pretoria, 2005. / Zoology and Entomology / unrestricted

MSc (Zoology) / Department of Zoology / See the attached abstract below

MSc (Zoology) / Department of Zoology / The concept of taxonomic diversity has been widely used to investigate diversity patterns and the mechanism underlying community assembly. However, functional and trait diversity can further explain the factors driving community assembly because they capture different aspects of species ecological roles such as habitat requirements and resource use. To investigate the factors shaping community assembly along the elevational Soutpansberg transect, ants were used as a model study organism because they are widely distributed geographically, ubiquitous and play several important roles in ecosystems. Functional and taxonomic diversity patterns along the elevation, their drivers (seasonality, soil properties, temperature and habitat structure) were examined using a long-term dataset (8 years) collected seasonally. Morphological and physiological traits at a community level are quantified and their relationship to temperature, soil properties and habitat structure modelled. Traits were used to test the size-grain hypothesis, Janzen’s rule and Brett’s rule. Functional and taxonomic diversity had a humped-shaped pattern on the northern and a decreasing pattern on the southern aspect. However, taxonomic diversity did not decrease monotonically on the southern aspect. Functional diversity was mainly related to habitat structure and temperature while taxonomic diversity was explained by seasonality, soil properties and temperature. Functional and taxonomic diversity were positively related at a larger scale but habitat specific at a smaller scale. Taxonomic diversity explained more variation in functional diversity than habitat structure and their interactions. The results supported the size-grain hypothesis and Janzen’s rule but not the Brett’s rule. Patterns in taxonomic and functional diversity mirrored each other but were the result of different mechanisms. Temperature was however important for both measures. Incorporating functional diversity analysis into taxonomic diversity contributed significantly in understanding the different mechanisms underlying community assembly along the Soutpansberg transect. This together with trait responses to environmental variables such as a decrease in size with increased temperature could be used to model the responses of ant communities to global change drivers such as climate and land use change / NRF