Potential impact and host range of Pereskiophaga brasiliensis Anderson (Curculionidae): a new candidate biological control agent for the control of Pereskia aculeata Miller (Cactaceae) in South Africa
- Authors: Mdodana, Lumka Anita
- Date: 2018
- Subjects: Curculionidae -- South Africa , Cactus -- South Africa , Biological pest control agents , Alien plants-- South Africa , Pereskiophaga brasiliensis Anderson (Curculionidae) , Pereskia aculeata Miller (Cactaceae)
- Language: English
- Type: text , Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/62350 , vital:28157
- Description: Pereskia aculeata Miller (Cactaceae) is a damaging invasive alien plant in South Africa that has negative impacts to indigenous biodiversity and ecosystem functioning. Mechanical and chemical control are not effective against P. aculeata so biological control is considered the only viable option. Two biological control agents, the leaf-feeding beetle Phenrica guerini Bechyne (Chrysomelidae) and the stem-wilting bug Catorhintha schaffneri (Coreidae), have been released in South Africa thus far. Post-release evaluations have indicated that P. guerini will not reduce P. aculeata densities to acceptable levels alone, while C. schaffneri was released very recently, so it is too soon to determine how effective that agent will be. Even if C. schaffneri is extremely damaging, it is likely that further agents will be required to reduce the densities of P. aculeata to acceptable levels within a reasonable time-scale. Additional agents should target the woody stems of P. aculeata which are not impacted by the damage of either of the released agents. Pereskiophaga brasiliensis Anderson (Curculionidae) is a promising potential candidate agent that feeds on the thick woody stems of the plant in the larval stage. Climatic matching, genetic matching and field based host specificity observations all indicated that P. brasiliensis was a promising candidate. In this study, the impact of P. brasiliensis to the target weed, P. aculeata, was quantified under quarantine conditions to determine whether it was sufficiently damaging to warrant release. This was followed by host specificity testing to determine whether P. brasiliensis was suitably host specific for release in South Africa. Impact studies indicated that P. brasiliensis was damaging to P. aculeata at insect densities that would be expected in the field. Pereskiophaga brasiliensis reduced the number of leaves of P. aculeata to a greater extent than it reduced shoot lengths, but both plant parameters were significantly reduced due to the feeding damage from the insect. This suggests that the damage from P. brasiliensis may be compatible with that of C. schaffneri which reduces shoot length to a greater degree than the number of leaves. Pereskiophaga brasiliensis is therefore sufficiently damaging to warrant release, and although interaction studies with the other agents would be required, it is expected that it should complement other existing agents. Although P. brasiliensis is sufficiently damaging, at present the host specificity data indicates that it is not suitably specific for release in South Africa because oviposition and larval development to the adult stage was recorded on both indigenous and alien plant species within the families Cactaceae and Basellaceae. This non-target feeding was recorded during no-choice tests, which are very conservative, but significant non-target damage and development to the adult stage was recorded on an indigenous plant from a different family to the target weed. Further host specificity testing, including paired and multiple choice tests, are required to confirm the broad host range of P. brasiliensis. Other biological control agents that damage the woody stems of P. aculeata should be considered. The stem-borer, Acanthodoxus machacalis (Cerambycidae) is considered the most promising of the other candidate agents as it can be sourced from a climatically matched region where genetically suitable P. aculeata plants are found, it is sufficiently damaging to the woody stems of P. aculeata and there is no evidence that the species has a broad host range. Acanthodoxus machacalis should be sourced from Rio de Janeiro, Brazil, and imported into quarantine in South Africa for host specificity testing.
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Interactions between two biological control agents released on Pereskia aculeata Miller (Cactaceae), in South Africa
- Authors: Mnqeta, Zezethu
- Date: 2017
- Subjects: Pereskia aculeata -- Biological control , Cactus -- Biological control -- South Africa , Alien plants -- Biological control -- South Africa , Flea beetles -- South Africa , Coreidae -- South Africa , Insects as biological pest control agents -- South Africa
- Language: English
- Type: text , Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/7949 , vital:21327
- Description: Pereskia aculeata Miller (Cactaceae) is an alien invasive plant introduced into South Africa from Brazil, which has negative impacts on native biodiversity in South Africa. Phenrica guerini Bechyne (Chrysomelidae) and Catorhintha schaffneri Brailovsky & Garcia (Coreidae) are two biological control agents released against P. aculeata in South Africa. Phenrica guerini was first released against P. aculeata, in 1991, followed by C. schaffneri in 2014. The overall aim of this study was to improve the biological control programme against P. aculeata in order to decrease its density to a level where it does not threaten the native biodiversity of South Africa. The first part of this study evaluated the efficacy of P. guerini on P. aculeata at Port Alfred (Eastern Cape) to better understand the role of P. guerini in the biological control of P. aculeata. An insecticide exclusion experiment was conducted over 100 days. Plots with P. guerini had a mean of 187 (SE ± 62) fewer leaves/m2 than plots without P. guerini. The agent reduced percentage cover in plots with P. guerini, with a mean of 19.42% (SE ± 3.15) lower cover than plots without P. guerini. Although P. guerini had an impact on P. aculeata at Port Alfred previous studies have indicated that a reduction to below 50% cover is required for native biodiversity to recover and the agent only reduced cover to 62% at Port Alfred. Phenrica guerini has therefore not reduced percentage cover sufficiently to completely control the weed. The data collected from Port Alfred was compared to the performance of the agent nationwide. Although P. guerini was found at far more sites than previously recorded, there were very few sites with comparable levels of damage to Port Alfred. This evidence suggests that P. guerini is not sufficiently damaging to reduce P. aculeata to acceptable levels and other biological control agents should be considered.Interactions between two biological control agents can have complex and unexpected impacts for a biological control programme. The second part of this study was to investigate interactions between C. schaffneri and P. guerini under laboratory conditions to test whether the two agents, individually or jointly, enhanced or reduced their impact on P. aculeata. Potted P. aculeata plants were exposed to one of four treatments: control (no agents), P. guerini only, C. schaffneri only and both species in combination. Four stocking densities, ranging from 2 to 12 insects per plant were used. Catorhintha schaffneri alone at high densities was more damaging than all other treatments with a significantly greater reduction in the mean number of leaves, 11.7 (SE ± 1.29), and shoot lengths, 2.17cm (SE ± O. 75). Even at lower density treatments, the combination of the two agents was not significantly more damaging than C. schaffneri alone and C. schaffneri was always more damaging than P. guerini alone. Mortality of P. guerini was significantly higher than C. schaffneri at the highest stocking density when in combination. Phenrica guerini contributes towards the biological control of P. aculeata at some sites in South Africa but not enough to completely control the weed. The antagonistic interaction between P. guerini and C. schaffneri suggests that these agents should not be released together because this would impact negatively on the overall biocontrol programme against P. aculeata. Catorhintha schaffneri should be released at sites were P. guerini is not present and evaluations of the success of this agent in the field should be conducted. Extrapolation of laboratory-based studies into the field is often challenging so mass-rearing of P. guerini should continue until there is convincing proof that C. schaffneri alone is more effective than P. guerini in the field.
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Complex interactions involving the Cape fig, Ficus sur Forsskål, and its associated insects
- Authors: Zachariades, Costas
- Date: 1995
- Subjects: Insect-plant relationships Mutualism (Biology) Fig -- South Africa Fig wasp -- South Africa Ants Homoptera
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5655 , http://hdl.handle.net/10962/d1005338
- Description: The inadequacy of arbitrarily classifying interactions between species as antagonistic, neutral or mutualistic has become clear in recent years. Both direct and indirect interactions between species can vary between mutualism and antagonism, depending on the intrinsic and extrinsic contexts of the interaction. This study investigated the characteristics of an ant-plant-homopteran interaction in southern Africa. The polyphagous homopteran Hilda patruelis (Tettigometridae) feeds primarily on the trunk-borne fruiting branches and figs of the Cape fig tree, Ficus sur, and produces honeydew which attracts tending ants. Ten of the sixteen ant species/species groups present on F. sur tended H. patruelis, with Pheidole megacephala the most frequent attendant. Ants attracted to F. sur by H. patruelis honeydew or other liquid food sources also preyed on insects on the tree, including adults of the small agaonid fig wasps whose larvae feed on the ovules in the developing figs. One fig wasp species (Ceratosolen capensis) is also the tree's only pollinator. No benefits to H. patruelis from being tended by ants were detected, either in terms of reduced parasitism, or predation by a lycaenid caterpillar. A P. megacephala colony foraging on a F. sur tree was found to receive a high proportion of its likely energy requirements from the tree, mainly in the form of H. patruelis honeydew, during periods when it was bearing fruit. It is probable that the H. patruelis-P. megacephala interaction constitutes a direct mutualism at times, but that benefits to the homopteran are intermittent or weak. Both H. patruelis and ants benefitted from F. sur, directly or indirectly, through the provision of food (and for some ants, nesting sites). The removal of phloem sap by H. patruelis did not detectably reduce the trees' reproductive output, either in terms of pollinator or viable seed production. The indirect effects of ant and H. patruelis presence on the F. sur trees were on average positive, as ants preyed disproportionately heavily on fig wasp species parasitic on or competing with the pollinator, thus increasing pollinator production. Effects of ant presence on seed production were not investigated, but have been demonstrated as beneficial elsewhere. However, there is great varatlon both in the composition of the wasp fauna arriving to oviposit at different crops, and in ant densities per fig, on several temporal and spatial scales. This results in high variability in the effects of ants on the pollinator and, through it, the tree, from positive to zero and potentially even negative. Despite this conditionality of beneficial outcomes for the tree, the mean effect of ants on the F. sur population studied was to increase pollinator production by up to nearly 20%. This study is among the few to have demonstrated an overall benefit to a plant of having homopteran-tending ants present on it.
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The phytophagous insect community on the Veld Fig, Ficus Burtt-Davyi Hutch
- Authors: Ross, Sally Jane
- Date: 1994
- Subjects: Phytophagous insects -- South Africa , Phytophagous insects , Insect-plant relationships
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5739 , http://hdl.handle.net/10962/d1005425 , Phytophagous insects -- South Africa , Phytophagous insects , Insect-plant relationships
- Description: The quest for patterns in community organisation is a daunting task which may be made easier by concentrating on communities associated with a restricted range of resources and therefore of relative simplicity. Here, the phytophagous insect community on the African fig tree Ficus burtt-davyi Hutch. was studied in an attempt to gain some insight into the factors which influence the composition of insect herbivore communities at a very local level, on individual plants of one host species at a single location. The tree's phenological patterns were detennined, due to their relevance to herbivores, particularly those which are host-specific feeders. The trees exhibited inter-tree asynchrony and intra-tree synchrony in fruit crop initiation, whereas leaf production was synchronous both within and between trees. Sixteen frequently occurring phytophagous insect species fed on the 123 F. burtt-davyi trees in the study area over a period of one year. Factors with the potential to influence the composition of this community were investigated at levels of the whole community (species richness), the guild, and the individual species. At each 'level' the effects of the measured factors on fluctuations in community composition were investigated, both over time (i.e. temporally) and spatially from tree to tree. During the year the phytophage community was influenced largely by temperature, although rainfall and tree phenological changes did exert varying influences on the abundances of guilds and individual species. Tree to tree variation in species richness (and thus commensurately, in the frequencies of occurrence of guilds and individual species) was influenced primarily by tree architectural complexity. Architecturally more complex trees hosted a greater number of species, a relationship largely attributable to effects of passive sampling and within-tree microhabitat heterogeneity and/or the availability of living space. The distributions of the leaf and stem piercing species were strongly associated with the presence of ants and this relationship manifested itself within the community as a whole. The degree of isolation of trees had consequences for individual species and for overall species richness, with the numbers of species present decreasing as trees became more isolated. A detailed analysis of guild distributional patterns revealed that the most important influential factors were those also evident at the level of the whole community and that species within guilds were, on the whole, no more similar to one another with respect to their habitat preferences than species from different guilds. The grouping of species into functional units therefore threw no additional light on the way in which this community is organised. An analysis of possible interspecific interactions between all of the phytophagous species in the community revealed only positive associations, both between species within guilds and between those in different guilds. These were doubtless attributable to autocorrelation as a result of similar habitat preferences. Competition was therefore rejected as an organising force within the community.
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Interactions between figs (Ficus spp., Moraceae) and fig wasps (Chalcidoidea, Agaonidae)
- Authors: Ware, Anthony Brian
- Date: 1993
- Subjects: Fig -- South Africa Fig wasp -- South Africa Pollination Agaonidae Chalcid wasps
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5719 , http://hdl.handle.net/10962/d1005405
- Description: Fig trees (Ficus spp., Moraceae) and fig wasps (Chalcidoidea, Agaonidae) are uniquely associated. In one fig wasp group, the pollinators (Agaoninae), each species is generally host species-specific. The relationship is one of obligate mutualism where the wasps provide pollination services and in return utilises some of the ovules for larval development. Non-pollinating fig wasps (generally belonging to subfamilies other than the Agaoninae) may be gallers or parasitoids, and can also be host species-specific. In the accompanying studies we examined the factors governing the interactions between fig wasps and their host trees. Surveys of fig trees and their associated pollinating fig wasps conducted in southern Africa, Madagascar and The Comores generally confirmed their specific relationships. An examination of F. sycomorlls in Madagascar resulted in the reclassification of F. sakalavarum as a distinct species with its own specific pollinator species. Biological and chemical evidence is presented demonstrating that the pollinators were able to distinguish their hosts through volatiles which emanated from the figs when they were ready to be pollinated. Environmental factors were found to influence wasp behaviour. Ambient temperature governed the timing of wasp emergence from their natal figs. When dispersing from their natal figs, the fig wasps flew upwards and then were blown downwind. Once nearing trees bearing figs ready to be pollinated, the wasps lost height and flew upwind towards the trees. E. baijnathi females apparently avoided figs which already contained a conspecific foundress. Scanning electron microscope studies of pollinating female fig wasp antennae showed that while all the species possessed multiporous plate sensilla, in only a few species were these sensilla elongated. Multiporous plate sensilla elongation is rare or absent among other female chalcids and may have evolved within the Agaoninae in order to facilitate their location on receptive host figs. Pollinator choice specificity appears to break down in a number of cases. In the first case examined, two pollinator species were recorded from the figs of African F. sycomorus. One. C. arabicus, pollinates the figs while the other, C. galili, acts as a 'cuckoo' by utilising some of ovules for oviposition without providing pollen. In the second case three pollinating fig wasp species were recorded from the rigs of F. lutea. Two were found to be incidental visitors and were not specifically attracted to the tree. The hybrid seeds from these crosses were successfully germinated but the seedlings did not grow passed the cotyledon stage of their development. In the concluding study the consequences of Ficus phenology and the structure of the fig's unusual inflorescence on the nonpollinating fig wasp community were examined. Various factors affecting the population levels and species richness were also examined. Future possible research directions were discussed.
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The systematics and phylogenetics of the Sycoecinae (Agaonidae, Chalcidoidea, Hymenoptera)
- Authors: Noort, Simon van
- Date: 1993
- Subjects: Agaonidae Chalcid wasps Fig wasp -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5784 , http://hdl.handle.net/10962/d1005472
- Description: The Sycoecinae are a distinct and well-defined subfamily of old world fig wasps (Agaonidae, Chalcidoidea 1 Hymenoptera) , exclusively associated with the figs of Ficus species (Moraceae). The most likely sister group of the Sycoecinae was determined to be the Sycoryctini (Sycoryctinae) based largely on synapomorphies of the underside of the head. 67 sycoecine species and 3 subspecies were recognised and included in a phylogenetic analysis of the subfamily. This analysis clearly delimited six genera (four African and two extra-African), although the phylogenetic relationships between the genera were not strongly supported and remain flexible. Comparisons of the phylogeny of the Sycoecinae with the classifications of the Agaoninae and their host fig trees (Ficus, Moraceae) suggest a degree of cospeciation sensu lato. Numerous homoplasies were detected within the Sycoecinae, some of which were shared with another group of fig wasps that also enter the fig to oviposit, the Agaoninae. The anatomy of the figs apparently provides strong selection pressures that have resulted in both parallelisms and convergences within and between the two subfamilies. Among the 67 species and 3 subspecies that were recognised, 43 species and 2 subspecies are described as new. The males of three previously recognised species are also described for the first time. One generic and two specific synonyms are established together with five new combinations. Keys are provided to the genera and species, for both sexes.
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Polymorphism and fighting in male fig wasps
- Authors: Vincent, Stephanie Louise
- Date: 1992
- Subjects: Fig wasp Chalcid wasps -- Morphology Wasps
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5740 , http://hdl.handle.net/10962/d1005426
- Description: Male fig wasps (Hymenoptera: Chalcidoidea) exhibit a fascinating range of morphology and behaviour. A cluster analysis, based on descriptions of the males of several hundred species, distinguished six major morphological groups. Behaviourial observations suggest that male morphology is related to the levels of inter-male aggression. Three behaviourial groupings were identified. Fighting species generally mated in the fig cavity, pacifist species mated in the females' galls or outside the figs. Mating sites are thus the primary determinants of male morphology and behaviour. In fighting species males were larger than their females, whereas pacifists and aggressors were equal in size or smaller than conspecific females. The large males in fighting fig wasps appear to be a consequence of sexual selection because larger males tended to win fights. Within a species there were no differences in the size of the galls that produced males and females, even in species where sexual size differences were present, suggesting that there is a heritable component to wasp size. No alternative advantages for smaller males were detected. Although fights were sometimes fatal, damage was not always a consequence of fighting behaviour and was recorded in both fighting and pacifist species. Sex ratios in several species were more female biased at higher population densities. Sex ratios of species with 'internally' ovipositing species were heavily female biased, but approached 1:1 in more outbred species with 'externally' ovipositing females. Levels of matedness, among females ranged from 73% to 99%. No evidence for sperm exhaustion was obtained. Species of Philotrypesis with both winged and flightless males were present only in southern African Ficus species from subsections Platyphyllae and Chlamydodorae. No species had only winged males. The flightless males of some Philotrypesis species were themselves polymorphic. In one polymorphic Philotrypesis species, winged males were found to be rare at high densities, but common at low densities. Digitata and religiosa males of Otitesella differed in coloration, size and behaviour. Digitata males were aggressors while religiosa males were fighters. Digitata males escaped from the figs whereas religiosa males remained inside the figs, perhaps because only digitata males were attracted to Light. Proportionally more digitata than religiosa males were present in low density populations and females were found to respond differently to the two morphs.
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Interactions between fig wasps and their host figs
- Authors: Nefdt, Rory John Charlton
- Date: 1990
- Subjects: Wasps , Mutualism (Biology)
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5718 , http://hdl.handle.net/10962/d1005404 , Wasps , Mutualism (Biology)
- Description: Fig trees (Ficus spp.) and fig wasps (Hymenoptera: Agaonidae) are partners in an intimate mutualism. The trees provide ovules in which wasp larvae develop while the wasps pollinate the flowers and are therefore indispensible for fig seed production. Agaonid fig wasps oviposit down the styles of fig flowers and it has generally been accepted that they were unable to reach the ovules of "long" styled flowers , which would produce seeds, thus maintaining an evolutionary stable mutualism. African fig species were found to have unimodal style length frequencies, with no separation into long and short styled flowers. In several species the ovipositors of their associated agaonids were long enough to reach the majority of ovules. The number of foundress agaonids entering a fig influenced fig seed set and therefore was an important factor regulating the proportion of flowers producing seeds or pollinators. In the two Ficus species that were studied, entry of more than three agaonid foundresses into one fig resulted in competition for limited oviposition sites and less female - biased offspring sex ratios. It is hypothesised that sequential laying of male eggs followed by female eggs, under variable oviposition site limitation, results in sex ratio adjustment, as predicted by local mate competition theory. Evidence in support of this hypothesis is presented. A number of non - pollinating torymid and pteromalid fig wasps also oviposit into each fig species. The sycophagines and sycoecines oviposit down the styles from inside the fig inflorescences like their agaonid counterparts, while other species insert their ovipositors through the wall of the fig from the outside. Like the agaonids, sycophagines were characterised by being pro - ovigenic, with numerous fully developed eggs at emergence. Sycoecines were able to re - emerge from figs they had oviposited in and lay their eggs in more than one fig. They had short ovipositors, allowing access to a smaller proportion of flowers than agaonids or sycophagines. Externally ovipositing fig wasps were syn-ovigenic, able to develop eggs as adults and invested more energy and time during each oviposition event. Differences in the ovipositor lengths of these species did not segregate their oviposition sites spatially, and therefore does not reduce competition between species. Attack by parasitoids and inquiline fig wasps from the exterior did not constitute a selection pressure against agaonids ovipositing in ovules closer to the periphery of the fig's surface, as predicted by Michaloud's enemy-free-space hypothes is. It cannot therefore explain the preference shown by ovipositing agaonids for shorter styled flowers.
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Interactions between ants, herbivorous insects and bracken (Pteridium aquilinum), a fern with extrafloral nectaries
- Authors: Rashbrook, Vanessa Karin
- Date: 1989
- Subjects: Arthropod pests -- South Africa Ants -- South Africa Ferns -- South Africa
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5592 , http://hdl.handle.net/10962/d1002042
- Description: 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.
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