Thesis (M. Sc.)(Entomology)--University of Pretoria, 2006. / Includes bibliographical references. Available on the Internet via the World Wide Web.

Three methods of communication were examined in the Middle Island tusked weta, Motuweta isolata (Orthoptera: Anostostomatidae); defence behaviour, acoustic and chemosensory signalling. Previous studies had been limited to basic autecology and anecdotal evidence. This study was undertaken to understand the behaviours of this species to assist in conservation efforts. Defensive behaviours were elicited through repeated stimulation while aggressive behaviours were acquired through male-male battles. Femoro-abdominal stridulation was induced within both situations. Defensive stridulation functioned as alarm behaviour and was often accompanied by a visual display. Agonistic stridulation was executed by the eventual winner of combat. Aggressive battles were a progression of behavioural units of increasing risk on injury until an individual was determined the winner. Acoustic analysis was preformed on stridulations observed in aggressive and defensive behaviours. Stridulation was a broad band signal covering a range well above 16 kHz and possibly into the ultrasound range (>20 kHz). Two different forms of stridulation were identified; click train sound and hiss sound. High speed frame-by-frame analysis of stridulation and scanning electron micrographs of the abdomen and medial femur deciphered the mechanism and found the interaction of the cuticle to be unique among weta. Strikingly, micrographs also revealed two morphologies of abdominal projections; truncated ridges and columnar pegs. Both these points were not previously observed. Y-maze and partition tests were utilized for evidence of chemosensory signalling, while gas chromatography-mass spectrometry identified faecal volatiles in M. iisolata and 4 other weta species in a parallel study. Through partition experiments, evidence exists for a species-specific volatile pheromone and a sex-specific chemo-tactile pheromone. Dimethylsulphide, (CH3)2S, was present in the faeces of all 5 weta species, and may be produced by the individual.

Three methods of communication were examined in the Middle Island tusked weta, Motuweta isolata (Orthoptera: Anostostomatidae); defence behaviour, acoustic and chemosensory signalling. Previous studies had been limited to basic autecology and anecdotal evidence. This study was undertaken to understand the behaviours of this species to assist in conservation efforts. Defensive behaviours were elicited through repeated stimulation while aggressive behaviours were acquired through male-male battles. Femoro-abdominal stridulation was induced within both situations. Defensive stridulation functioned as alarm behaviour and was often accompanied by a visual display. Agonistic stridulation was executed by the eventual winner of combat. Aggressive battles were a progression of behavioural units of increasing risk on injury until an individual was determined the winner. Acoustic analysis was preformed on stridulations observed in aggressive and defensive behaviours. Stridulation was a broad band signal covering a range well above 16 kHz and possibly into the ultrasound range (>20 kHz). Two different forms of stridulation were identified; click train sound and hiss sound. High speed frame-by-frame analysis of stridulation and scanning electron micrographs of the abdomen and medial femur deciphered the mechanism and found the interaction of the cuticle to be unique among weta. Strikingly, micrographs also revealed two morphologies of abdominal projections; truncated ridges and columnar pegs. Both these points were not previously observed. Y-maze and partition tests were utilized for evidence of chemosensory signalling, while gas chromatography-mass spectrometry identified faecal volatiles in M. iisolata and 4 other weta species in a parallel study. Through partition experiments, evidence exists for a species-specific volatile pheromone and a sex-specific chemo-tactile pheromone. Dimethylsulphide, (CH3)2S, was present in the faeces of all 5 weta species, and may be produced by the individual.

The eight southern African King Cricket genera, namely Bochus, Borborothis, Henicus, Libanasa, Libanasidus, Nasidius, Onosandridus, and Onosandrus are redescribed from type and museum material and a key to the genera is provided. Additionally, the status of Libanasidus impicta is investigated and species characteristics confirmed. A key to the two Libanasidus species is also provided. No morphological support for these eight southern African anostostomatid genera was attained with cladistic analysis, despite the incorporation of taxonomically important characters. High levels of homoplasy and possible incorrect species placement, resulting in character ambiguity within genera, renders many of the diagnostic characters of this group ineffective for resolving generic relationships. Genetic data based on the large ribosomal subunit (16S) did provide phylogenetic resolution between six of the genera with good bootstrap support. This confirmed speculation by previous authors as to the ancestral nature of Bochus and Borborothis to other genera within the Anostostomatini tribe, as well as the placement of Libanasa within a separate tribe, the Lutosini. The recent merge of the genus Platysiagon with Libanasa is also provisionally supported. Close association was obtained between Libanasidus and Nasidius with Onosandrus being more related to Bochus and Borborothis, supported by the lack of sexual dimorphism in Onosandrus and Borborothis. The phylogenetic position of the genera Henicus and Onosandridus remains unresolved. It is suggested that the designation of the eight anostostomatid genera in southern Africa are valid, but that species placement within these genera need to be revised to resolve character conflict. Focusing on the genetic and morphometric structuring within the famous Parktown Prawn, Libanasidus vittatus from southern Africa consistently suggests two main population assemblages. These correspond to a large North-South ranging population including individuals west of the prominent escarpment, and a smaller population including individuals from the eastern side of the escarpment. COI sequence data recovers two clades representing these two populations with good bootstrap support in likelihood, parsimony, Bayesian and distance analyses. Genetic divergence between the two clades averaged 3.3%, while population parameters estimated using maximum likelihood methods show low migration rates corresponding to less that one female migration per generation. A priori morphometric analyses including PCA&clustering methods show no biologically meaningful variation, suggesting that the two clades recovered represent cryptic sibling species. Inferring a molecular clock of 2% divergence per million years used for sister taxa, signifies isolation of the eastern population at 1.65 mya. A posteriori morphometric analyses confirmed the genetic results, based on 11 size-related measurements. This study provides a starting point for further work on the taxonomy, behaviour and ecology of these fascinating insects. Cytogenetics, multiple genetic loci and geometric morphometrics will provide useful insight into the taxonomic status of the 51 anostostomatid species in southern Africa and is promoted for further studies. / Dissertation (MSc Entomology)--University of Pretoria, 2006. / Zoology and Entomology / unrestricted

Three kinds of processes are known to modify the geographical spatial arrangement of organisms: dispersal, extinction and vicariance. The Southern Hemisphere has an intriguing and complicated geological history that provides an ideal backdrop to study these processes. This thesis focuses on three historical events that illustrate these processes: the proposed marine inundation of New Zealand in the Oligocene, the asteroid impact at the K – Pg boundary, and the continental breakup of Gondwana. It investigates what impact these events had on species diversification by studying the phylogenetic relationships of two groups of taxa – the family Anostostomatidae (insects), and Neoaves (birds). Anostostomatidae were studied in relation to the Oligocene drowning and the break up of Gondwana as they have a wide southern distribution, found on all “Gondwanan” fragments with the exception of Antarctica, and are thought represent an ancient lineage that predates the Gondwanan breakup. Birds, in particular Neoaves, were studied in relation to the asteroid impact at the K – Pg boundary. Although birds are mobile and many circumnavigate the globe between seasons, they are suggested to have originated in the Southern Hemisphere in Gondwanan times, and subsequently undergone range expansion and diversification around the world. In order to address the relationship (if any) between modern biotic diversity and historical geological events, phylogenetic relationships were determined and where possible, molecular clock analysis carried out. Timing information provided by molecular clock analysis is important as it enables distinction between opposing hypotheses such as vicariance and dispersal. In Chapter Two, the phylogenetic relationships within the family Anostostomatidae are investigated. One of the most controversial times in New Zealand’s geological history is during the Oligocene. Some suggest that the lack of fossils and evidence for recent dispersal of numerous taxa support the notion that all modern biota reached the region during the last 25 million years. Anostostomatidae were chosen as they represent a group of insects that are thought to be ancient and there is little published data in the literature. Previous studies focused on the relationships within Hemideina and Deinacrida suggesting that these groups diversified in the early Miocene. The data presented here are from mitochondrial (COI and 12S) and nuclear (18S and 28S) sequences. Molecular dating using a relaxed clock as implemented in BEAST suggest that in fact some lineages were present at or shortly after continental breakup and could have survived throughout this turbulent time. As there were no definitive fossils to use for calibration points, geological events were used as calibration points for the molecular clock. Mutation rates obtained from the different analyses were compared to those published for other insects in an attempt to identify the most likely model. Both maximum likelihood and Bayesian analyses support the presence of three distinct ecological groups in New Zealand; Hemiandrus (ground weta), Anisoura/Motuweta (tusked weta) and Hemideina–Deinacrida (tree–giant weta). With regards to their Australasian relatives (taxa from Australia and New Caledonia) it appears that the family is divided with the most northern New Zealand taxa (tusked weta) more closely related to New Caledonian taxa while all other New Zealand taxa are more closely related to Australian taxa. There does not appear to be any link between the Australian and New Caledonian taxa studied here. Results should be viewed with caution however as an increased mutation rate was observed in the New Caledonian-tusked weta lineage, something future studies will have to address. Chapter Three presents new sequence data and phylogenetic analyses that go towards resolving the apparent basal polytomy of neoavian birds. This chapter includes analyses carried out on previously published data with the addition of nine new mitochondrial genomes. My contribution to this larger project was to perform the phylogenetic analysis and to sequence three of the nine mitochondrial genomes. The genomes I sequenced were the Southern Hemisphere species: dollar bird (Eurystomus orientalis), Owlet nightjar (Aegotheles cristatus cristatus) and great potoo (Nyctibius grandis). The inclusion of these nine new genomes allows assessment and comparison of the six hypothesised groups reported in Cracraft (2001). First an improved conditional down-weighting technique is described reducing noise relative to signal, which is important for resolving deeper divergences. Second, a formula is presented for calculating probabilities of finding predefined groupings in the optimal tree. Maximum likelihood and Bayesian based phylogenetic analyses were carried out and in addition, dating using a relaxed molecular clock was performed in BEAST. Results suggested that the six groups suggested by Cracraft (2001) represent robust lineages. The results suggested that one group, the owls, are more closely related to other raptors, particularly accipitrids (buzzards/eagles) and the osprey rather than the Caprimulgiformes, which could indicate morphological convergent evolution. In addition, a group termed shorebirds appears to be distinct from the large group referred to as ‘Conglomerati’ to which previous publications have suggested they belong. The ‘Conglomerati’ is the least well studied group and may actually comprise of at least three subgroups (as suggested by Cracraft). Within the three suggested groups, Cracraft grouped shorebirds with pigeons and sandgrouse, neither of which (pigeons or sandgrouse) were analysed here. So although the shorebirds are at least close to the ‘Conglomerati’ and may be within that group, their exact position is still not clear. The molecular dating reported here utilised two fossil calibrations (Vegavis and Waimanu), for which there is relatively little dispute as to age or the lineage to which they belong. Calibrations resulting from BEAST analyses suggest that at least 12 distinct lineages were present prior to the K – Pg boundary, a finding supported by previous studies. Robust phylogenies will allow future studies to investigate not only the relationships within Neoaves, but look more closely at the biological and ecological evolution of the group. Chapter Four for the first time investigates whether the phylogenetic relationships within the family Anostostomatidae follow the conventionally accepted order and timing of Gondwanan breakup. Following the initial restults for taxa studied in Australasia (Chapter Two) an attempt to resolve family relationships in a wider spatial (geographic) context was carried out to determine if Australasian taxa are monophyletic when other members of the family are included. Again both maximum likelihood and Bayesian phylogenetic analyses were carried out on both mitochondrial (COI and 12S) and nuclear (18S and 28S) sequences. In this chapter, datasets included samples from across the geographic range of Anostostomatidae (South Africa, Madagascar, South America, Australia, New Caledonia and New Zealand), and two clades were observed, congruent with earlier findings. Sequence divergence within geographic regions was found to be relatively high in the mitochondrial genes (COI and 12S) while low in the nuclear ribosomal RNA genes (18S and 28S) as expected given their relative mutation rates. Under the vicariance paradigm, phylogenetic relationships should follow the order of continental breakup, but this was not found. Further, if dispersal and colonisation were continuous, no geographic substructure is expected, however distinct geographic substructure within clades was consistently observed. This interesting phylogenetic pattern may be a case of convergent evolution or paraphyletic sampling which highlights taxonomic issues of the group. Future studies need to include not only molecular data but information on morphology, ecology and behaviour along with the implementation of biogeographic programs that can test alternative hypotheses (such as dispersal and vicariance) directly. Also, the inclusion of the recently reported fossil from the subfamily Euclydesinae (Martins-Neto 2007) should allow for more accurate date estimates within the family. Taken as a whole the results presented in this thesis suggest that microevolutionary processes are sufficient to explain modern diversity without the need to invoke abiotic events. The three cases investigated here - marine inundation, asteroid impact and continental drift - all appear to have had only a limited effect on the diversity of taxa studied. To reach even stronger conclusions future studies should incorporate different data (for instance nuclear genes, intron position, and genome structure) and use biogeographic software capable of including ecological, morphological and habitat information.