Some parasites may modify the behavior of their hosts. Altered behaviors may: 1) benefit the host in that they defend against the pathogen, 2) benefit the pathogen and represent manipulations of the host response, and 3) benefit neither the host or the pathogen and simply be a product of the host response to infection. In this thesis I examine four host/parasite systems. For each system, I explore host/parasite behavioral interactions, and examine them with regard to selective pressures that may be acting on both the host and the parasite. I test the Hamilton and Zuk hypothese in 26 species of lizards. I find an inverse relationship between a lizard species' brightness and parasite prevalence. My result lend credence to criticisms of the Hamilton and Zuk Hypothesis. If infection does occur, animals may alter their behavior to impair the growth and reproduction of the parasite. To test this prediction, I examine behavioral thermoregulation in two strains of the snail Biomphalaria glabrata, one resistant to, and one susceptible to, the parasite Schistosoma mansoni. The preferred temperature of infected snails drops five weeks after exposure to the parasite. I propose the hypothesis that pathogen-induced host defense responses result in altered host behaviors and enhanced predation. In particular, I examine the effects of the acute phase response (a physiological response whose symptoms include fever, reduced activity and malaise) on antipredatory behavior in bullfrog (Rana catesbeiana) tadpoles. This host response is associated with the preliminary stages of infection with many pathogens yet its behavioral effects have received little attention. I find that the stereotypical effects of the acute phase response can lead to increased predation. I suggest that altered behaviors may afford some parasites a potential pathway to their next host. I examine the behavioral effects of a yeast, Candida spp., a single-host parasite species in its natural host, the red-legged frog (Rana aurora). Infected tadpoles exhibit the same behavioral modifications that are noted in bacteria injected bullfrog tadpoles. These results suggest that some altered behaviors may occur due to a host response to infection and not due to parasitic manipulation. / Graduation date: 1993

From simple beginnings, when only one host was required, numerous parasitic organisms have evolved complex life-cycles involving two or more host species. For example, trematode parasites reproduce in vertebrates, their definitive host, but their current life cycle also typically involves two intermediate hosts that were added during the course of evolution. Vertebrates are often considered to be the ancestral hosts of trematodes although other scenarios exist. While multi-host life cycles are observed in distantly related groups of parasites, their evolution remains largely unexplored. In trematodes, while recent phylogenetic studies have shed light on the sequence along which the different hosts were incorporated in the cycle, conditions that favoured the evolution of such complex life cycles can only be hypothesized. However, one opportunity to understand the force shaping the evolution of complex life cycles is provided by the few trematode species in which the classical three-host cycle facultatively reverts to a shorter cycle (i.e. life cycle abbreviation). In this study, the effects of different environmental factors on the life history strategy of the trematode Coitocaecum parvum were investigated using laboratory and field studies. C. parvum is able to abbreviate its life cycle from three to two hosts by maturing early (i.e. progenesis) and producing eggs inside the second intermediate host; both life history strategies occur simultaneously in C. parvum populations. Environmental factors such as predator densities should strongly influence parasite life history strategies. In fact, this study shows that laboratory reared Coitocaecum parvum adopt preferentially the normal three-host cycle when chemical cues from the definitive host are added to their environment, while the shorter cycle is favoured when these cues are absent. However, in nature, multiple environmental factors are likely to be perceived by parasites. Consequently, C. parvum�s ability to adapt its developmental strategy to definitive host densities may be confounded by the complex combination of various environmental parameters. Within-host competition between parasites sharing a common host is also likely to influence individual life history strategies. Parasites could then use alternative life strategies to adaptively respond to intraspecific and interspecific competition. Indeed, this study found that C. parvum preferentially adopts the abbreviated cycle in the presence of competitors. However, in interspecific competition, C. parvum�s strategy also depends upon the competitor species, possibly influenced by the other species� transmission route. Furthermore, intensity of intraspecific competition proved to constrain C. parvum�s ability to use the abbreviated life cycle. Finally, genetic relatedness between co-infecting C. parvum individuals seems to affect parasite life strategy through kin selection: closely related individuals are more likely to adopt the same developmental strategy, when they share a host, than unrelated ones. C. parvum individuals adopting the abbreviated cycle are enclosed within a cyst in their intermediate host and must produce eggs by self-fertilization, the most severe case of inbreeding. It was hypothesized that their offspring would have reduced fitness due to inbreeding depression, therefore selecting against the shorter cycle. However, this study found no difference in the survival and infection success of offspring produced through the abbreviated and normal cycles. Furthermore, no evidence for a genetic basis of life cycle abbreviation was detected: the same proportion of offspring from both reproductive strategies adopted the shorter life cycle. The work in this thesis provides evidence that although life cycle abbreviation provides Coitocaecum parvum with a viable alternative life strategy, numerous factors promote or restrict the adoption of this strategy. While this life history strategy has no detectable effect on parasite fitness, both environmental parameters and within-host competition affect C. parvum life-history strategies, alternatively selecting for either the shorter or normal life cycle. Overall, the complexity of the parasite environment could maintain both developmental strategies in C. parvum populations and, on a broader scale, could have influenced the evolution of complex life cycles in parasites.

Abstract Experiments in this thesis were conducted to investigate the possibility of bestowing lambs with increased resistance to gastrointestinal parasites through maternal protein and copper supplementation. Reproductive outcomes such as birth weight, haematological parameters and faecal egg counts were used as indices of possible foetal programming. This thesis involved 5 experiments. The first three experiments were done using rats as a preliminary study animal on account of their short generational intervals and high fecundity. The final two experiments involved Merino sheep. The first experiment in Chapter 4 investigated the most optimum larval dose to use in order to elicit a measurable immune response. Weaned offspring were infected with a rat nematode, Nippostrongylus brasiliensis and their response measured by faecal egg counts, parasite recovery from intestines at sacrifice, spleen weights and leucocyte numbers, especially manifested as eosinophilia. There was no significant difference in parasite rejection for rats infected with 1000 larvae/rat. When rats were infected with variable larval doses to determine the optimum dose rate, eosinophilia and spleen weight were significantly increased as dose rate increased from 500 L3 to 2,000 larvae. Based on these results, it was decided to use 1,000 larvae for each rat in Chapter 6. The experiment in Chapter 5 involved feeding diets with 5 graded concentrations of copper (Cu) ranging from deficient (1 ppm diet) to high (16 ppm diet). Rats were fed for 4 weeks before mating after which half of them were sacrificed to determine liver Cu concentrations and haematological parameters. The rest were mated and maintained on their respective Cu diets into the second trimester of pregnancy. Pregnant females were sacrificed on approximately gestational Day 10 to recover foetuses and determine the incidence of foetal defects, foetal Cu status as well as maternal liver copper status. It was determined that most morphological defects occurred for the 1 ppm foetuses and both 2 ppm and 4 ppm had similar incidences of brain enlargements. The 16 ppm copper diet was excessive evidenced by reduced liver iron status and erythrocyte counts to similar levels as for 2 ppm rats although it had no adverse effect on foetal development. Significant differences were found for liver Cu status, erythrocyte counts and spleen weights due to the copper diets. A deficient copper diet containing 1 ppm Cu (LC) and an adequate diet containing 8 ppm Cu (SC) were used for the last rat experiment in Chapter 6 which was funded by the Science and Innovation Award. The LC diets were fed for 4 weeks prior to mating. Rats were then fed LC throughout pregnancy, for the 1st trimester only or for the 1st and 2nd trimesters. Other pregnant females were fed the SC diet throughout pregnancy. Offspring were challenged with 1000 L3 N. brasiliensis and their immune responses measured. Copper deficiency at variable stages of prenatal development caused significant postnatal mortalities but had no effect on response to parasite resistance. However, significant parasite and sex effects were found for parameters such as spleen weight, eosinophilia and weight loss during infection. The foetal brain enlargement caused by the deficient 1 ppm Cu diet was determined to be reversible in vivo upon exposure to a normal 8 ppm Cu diet during gestation. Chapter 7 involved Merino ewes which were fed either a high protein diet (21%) or adequate protein diet (12%) during the first 2 trimesters of pregnancy. Production parameters measured included pregnancy weight gain, fleece yield, protein content in milk as well as birth weight of lambs but none were significantly different. After weaning, the lambs were experimentally infected with 10,000 Haemonchus contortus larvae. Barber’s pole worm is responsible for millions of dollars in production losses in the sheep industry. Responses measured were eosinophilia, faecal egg count, anaemia (PCV) and weight gain/loss during the infection period. No significant differences were found for any parameter tested except for a parasite effect on erythrocyte numbers and PCV. In Chapter 8 Merino ewes were used which were mildly Cu deficient due to grazing on pasture that was copper deficient. Control ewes were supplemented with copper oxide wire particles at mating and mid-pregnancy. The rest of the experiment was the same as for Chapter 7 in terms of Barber’s pole worm larval dose. There were no significant differences in birth weight, weaning weight or ewe fleece weights due to copper status. There were no differences in parasite resistance in the lambs due to maternal Cu status measured by live weights, eosinophil concentrations or faecal egg counts. In conclusion, foetal programming by maternal nutritional supplements for postnatal parasite resistance appears to be impossible. It may be that if a different organ was targeted, such as the spleen, the results would have been different. The thymus appears to be non-programmable during foetal development in rats and sheep. However, it was a worthwhile attempt at conferring resistance to parasites in lambs due to the urgency in combating the global problem of parasite resistance to anthelmintics and the resultant large economic losses that are experienced by the global sheep industry.

Abstract The Benedeniinae, the largest of nine capsalid subfamilies, includes genera with an aseptate, apapillate haptor and a pair of discrete testes. Eight of 13 nominal benedeniine genera, Benedenia Diesing, 1858 (the type genus); Allobenedenia Yamaguti, 1963; Allometabenedeniella Velasquez, 1982; Dioncopseudobenedenia Yamaguti, 1965; Lagenivaginopseudobenedenia Yamaguti, 1966; Oligoncobenedenia Yamaguti, 1965; Pseudallobenedenia Yamaguti, 1966 and Tareenia, Hussey, 1986 are revised using type material of most nominal species with observations for some species from new material. Allobenedenia and Allometabenedeniella are transferred to the Trochopodinae Price, 1936 emend. Sproston, 1946 because all valid species of both genera bear septa on the ventral haptor surface. Allobenedenia ishikawae (Goto, 1894) Yamaguti, 1963 is transferred to Benedenia because its haptor is aseptate. I recognize Menziesia Gibson, 1976, based on the form of the male copulatory organ and associated structures and include five species in it. Tareenia Hussey, 1986 is synonymised with Benedenia because characters used to differentiate the two genera do not indicate discontinuities at the generic level. Benedeniella Johnston, 1929, Calicobenedenia Kritsky and Fennessy, 1999 and Trimusculotrema Whittington and Barton, 1990 are considered to belong in Entobdellinae Bychowsky, 1957, pending further studies on that group. The anatomy of Calicobenedenia is outlined briefly; the other genera are not discussed here in detail. Lachishia n. g. is described, based on the structure of the male copulatory organ and a species is transferred to it from Dioncopseudobenedenia. I describe four new species of benedeniines from teleosts caught at Heron and Green Islands, Australia namely: Benedenia ernsti n. sp. from the gills of Symphorus nematophorus; Benedenia fieldsi n. sp. from the fins of Cephalopholis boenak, C. cyanostigma and C. miniatus; Benedenia haywardi n. sp. from the skin of S. nematophorus and Dioncopsudobenedenia ancoralis n. sp. from the gills of Siganus lineatus. Benedenia akaisaki Iwata, 1990 is synonymised with B. ovata (Goto, 1894) Johnston, 1929, B. kintoki Iwata, 1990 is synonymised with B. elongata (Yamaguti, 1968) Egorova, 1997, B. sargocentron Zhang, Yang and Liu, 2001 is synonymised with B. hawaiiensis Yamaguti, 1968 and Pseudallobenedenia arabica Timofeeva, 1995 is synonymised with P. opakapaka Yamaguti, 1966. Benedenia madai Ishii and Sawada, 1938, B. pagrosomi Ishii and Sawada, 1938 and Allobenedenia pedunculata Raju and Rao, 1980 are considered species inquirendae. I examined an outbreak in aquaculture of the pathogenic benedeniine, Neobenedenia melleni (MacCallum, 1927) Yamaguti, 1963 and report this species for the first time in Australia. I examined the pathogenesis caused by this parasite by histology of host tissue. Possible routes of introduction to the farm in question are investigated and studies are detailed that should be undertaken in Australia to manage future outbreaks of N. melleni. Capsalids usually colonise new hosts by an infective ciliated oncomiracidium. An experiment was conducted to ascertain whether cleaner fish Labroides dimidiatus could transfer adult skin-parasitic monogeneans from one host fish to another. I have shown that transmission of adult monogeneans between two individuals of Hemigymnus melapterus by cleaner fish can occur. In coral reef environments, frequent contact between unrelated hosts involved in cleaning interactions might create previously unsuspected evolutionary pressures. I discuss the implications of this discovery for host-specificity and lateral transmission of monogeneans. General biological studies of monogeneans are necessary to understand the evolution, pathology, epidemiology, phylogeny and taxonomy of these parasites. My study provides a taxonomic system which, when applied to other capsalid subfamilies, should help prevent errors and create a clear system of classification for the entire family.

Abstract The Benedeniinae, the largest of nine capsalid subfamilies, includes genera with an aseptate, apapillate haptor and a pair of discrete testes. Eight of 13 nominal benedeniine genera, Benedenia Diesing, 1858 (the type genus); Allobenedenia Yamaguti, 1963; Allometabenedeniella Velasquez, 1982; Dioncopseudobenedenia Yamaguti, 1965; Lagenivaginopseudobenedenia Yamaguti, 1966; Oligoncobenedenia Yamaguti, 1965; Pseudallobenedenia Yamaguti, 1966 and Tareenia, Hussey, 1986 are revised using type material of most nominal species with observations for some species from new material. Allobenedenia and Allometabenedeniella are transferred to the Trochopodinae Price, 1936 emend. Sproston, 1946 because all valid species of both genera bear septa on the ventral haptor surface. Allobenedenia ishikawae (Goto, 1894) Yamaguti, 1963 is transferred to Benedenia because its haptor is aseptate. I recognize Menziesia Gibson, 1976, based on the form of the male copulatory organ and associated structures and include five species in it. Tareenia Hussey, 1986 is synonymised with Benedenia because characters used to differentiate the two genera do not indicate discontinuities at the generic level. Benedeniella Johnston, 1929, Calicobenedenia Kritsky and Fennessy, 1999 and Trimusculotrema Whittington and Barton, 1990 are considered to belong in Entobdellinae Bychowsky, 1957, pending further studies on that group. The anatomy of Calicobenedenia is outlined briefly; the other genera are not discussed here in detail. Lachishia n. g. is described, based on the structure of the male copulatory organ and a species is transferred to it from Dioncopseudobenedenia. I describe four new species of benedeniines from teleosts caught at Heron and Green Islands, Australia namely: Benedenia ernsti n. sp. from the gills of Symphorus nematophorus; Benedenia fieldsi n. sp. from the fins of Cephalopholis boenak, C. cyanostigma and C. miniatus; Benedenia haywardi n. sp. from the skin of S. nematophorus and Dioncopsudobenedenia ancoralis n. sp. from the gills of Siganus lineatus. Benedenia akaisaki Iwata, 1990 is synonymised with B. ovata (Goto, 1894) Johnston, 1929, B. kintoki Iwata, 1990 is synonymised with B. elongata (Yamaguti, 1968) Egorova, 1997, B. sargocentron Zhang, Yang and Liu, 2001 is synonymised with B. hawaiiensis Yamaguti, 1968 and Pseudallobenedenia arabica Timofeeva, 1995 is synonymised with P. opakapaka Yamaguti, 1966. Benedenia madai Ishii and Sawada, 1938, B. pagrosomi Ishii and Sawada, 1938 and Allobenedenia pedunculata Raju and Rao, 1980 are considered species inquirendae. I examined an outbreak in aquaculture of the pathogenic benedeniine, Neobenedenia melleni (MacCallum, 1927) Yamaguti, 1963 and report this species for the first time in Australia. I examined the pathogenesis caused by this parasite by histology of host tissue. Possible routes of introduction to the farm in question are investigated and studies are detailed that should be undertaken in Australia to manage future outbreaks of N. melleni. Capsalids usually colonise new hosts by an infective ciliated oncomiracidium. An experiment was conducted to ascertain whether cleaner fish Labroides dimidiatus could transfer adult skin-parasitic monogeneans from one host fish to another. I have shown that transmission of adult monogeneans between two individuals of Hemigymnus melapterus by cleaner fish can occur. In coral reef environments, frequent contact between unrelated hosts involved in cleaning interactions might create previously unsuspected evolutionary pressures. I discuss the implications of this discovery for host-specificity and lateral transmission of monogeneans. General biological studies of monogeneans are necessary to understand the evolution, pathology, epidemiology, phylogeny and taxonomy of these parasites. My study provides a taxonomic system which, when applied to other capsalid subfamilies, should help prevent errors and create a clear system of classification for the entire family.

Abstract The Benedeniinae, the largest of nine capsalid subfamilies, includes genera with an aseptate, apapillate haptor and a pair of discrete testes. Eight of 13 nominal benedeniine genera, Benedenia Diesing, 1858 (the type genus); Allobenedenia Yamaguti, 1963; Allometabenedeniella Velasquez, 1982; Dioncopseudobenedenia Yamaguti, 1965; Lagenivaginopseudobenedenia Yamaguti, 1966; Oligoncobenedenia Yamaguti, 1965; Pseudallobenedenia Yamaguti, 1966 and Tareenia, Hussey, 1986 are revised using type material of most nominal species with observations for some species from new material. Allobenedenia and Allometabenedeniella are transferred to the Trochopodinae Price, 1936 emend. Sproston, 1946 because all valid species of both genera bear septa on the ventral haptor surface. Allobenedenia ishikawae (Goto, 1894) Yamaguti, 1963 is transferred to Benedenia because its haptor is aseptate. I recognize Menziesia Gibson, 1976, based on the form of the male copulatory organ and associated structures and include five species in it. Tareenia Hussey, 1986 is synonymised with Benedenia because characters used to differentiate the two genera do not indicate discontinuities at the generic level. Benedeniella Johnston, 1929, Calicobenedenia Kritsky and Fennessy, 1999 and Trimusculotrema Whittington and Barton, 1990 are considered to belong in Entobdellinae Bychowsky, 1957, pending further studies on that group. The anatomy of Calicobenedenia is outlined briefly; the other genera are not discussed here in detail. Lachishia n. g. is described, based on the structure of the male copulatory organ and a species is transferred to it from Dioncopseudobenedenia. I describe four new species of benedeniines from teleosts caught at Heron and Green Islands, Australia namely: Benedenia ernsti n. sp. from the gills of Symphorus nematophorus; Benedenia fieldsi n. sp. from the fins of Cephalopholis boenak, C. cyanostigma and C. miniatus; Benedenia haywardi n. sp. from the skin of S. nematophorus and Dioncopsudobenedenia ancoralis n. sp. from the gills of Siganus lineatus. Benedenia akaisaki Iwata, 1990 is synonymised with B. ovata (Goto, 1894) Johnston, 1929, B. kintoki Iwata, 1990 is synonymised with B. elongata (Yamaguti, 1968) Egorova, 1997, B. sargocentron Zhang, Yang and Liu, 2001 is synonymised with B. hawaiiensis Yamaguti, 1968 and Pseudallobenedenia arabica Timofeeva, 1995 is synonymised with P. opakapaka Yamaguti, 1966. Benedenia madai Ishii and Sawada, 1938, B. pagrosomi Ishii and Sawada, 1938 and Allobenedenia pedunculata Raju and Rao, 1980 are considered species inquirendae. I examined an outbreak in aquaculture of the pathogenic benedeniine, Neobenedenia melleni (MacCallum, 1927) Yamaguti, 1963 and report this species for the first time in Australia. I examined the pathogenesis caused by this parasite by histology of host tissue. Possible routes of introduction to the farm in question are investigated and studies are detailed that should be undertaken in Australia to manage future outbreaks of N. melleni. Capsalids usually colonise new hosts by an infective ciliated oncomiracidium. An experiment was conducted to ascertain whether cleaner fish Labroides dimidiatus could transfer adult skin-parasitic monogeneans from one host fish to another. I have shown that transmission of adult monogeneans between two individuals of Hemigymnus melapterus by cleaner fish can occur. In coral reef environments, frequent contact between unrelated hosts involved in cleaning interactions might create previously unsuspected evolutionary pressures. I discuss the implications of this discovery for host-specificity and lateral transmission of monogeneans. General biological studies of monogeneans are necessary to understand the evolution, pathology, epidemiology, phylogeny and taxonomy of these parasites. My study provides a taxonomic system which, when applied to other capsalid subfamilies, should help prevent errors and create a clear system of classification for the entire family.

Abstract The Benedeniinae, the largest of nine capsalid subfamilies, includes genera with an aseptate, apapillate haptor and a pair of discrete testes. Eight of 13 nominal benedeniine genera, Benedenia Diesing, 1858 (the type genus); Allobenedenia Yamaguti, 1963; Allometabenedeniella Velasquez, 1982; Dioncopseudobenedenia Yamaguti, 1965; Lagenivaginopseudobenedenia Yamaguti, 1966; Oligoncobenedenia Yamaguti, 1965; Pseudallobenedenia Yamaguti, 1966 and Tareenia, Hussey, 1986 are revised using type material of most nominal species with observations for some species from new material. Allobenedenia and Allometabenedeniella are transferred to the Trochopodinae Price, 1936 emend. Sproston, 1946 because all valid species of both genera bear septa on the ventral haptor surface. Allobenedenia ishikawae (Goto, 1894) Yamaguti, 1963 is transferred to Benedenia because its haptor is aseptate. I recognize Menziesia Gibson, 1976, based on the form of the male copulatory organ and associated structures and include five species in it. Tareenia Hussey, 1986 is synonymised with Benedenia because characters used to differentiate the two genera do not indicate discontinuities at the generic level. Benedeniella Johnston, 1929, Calicobenedenia Kritsky and Fennessy, 1999 and Trimusculotrema Whittington and Barton, 1990 are considered to belong in Entobdellinae Bychowsky, 1957, pending further studies on that group. The anatomy of Calicobenedenia is outlined briefly; the other genera are not discussed here in detail. Lachishia n. g. is described, based on the structure of the male copulatory organ and a species is transferred to it from Dioncopseudobenedenia. I describe four new species of benedeniines from teleosts caught at Heron and Green Islands, Australia namely: Benedenia ernsti n. sp. from the gills of Symphorus nematophorus; Benedenia fieldsi n. sp. from the fins of Cephalopholis boenak, C. cyanostigma and C. miniatus; Benedenia haywardi n. sp. from the skin of S. nematophorus and Dioncopsudobenedenia ancoralis n. sp. from the gills of Siganus lineatus. Benedenia akaisaki Iwata, 1990 is synonymised with B. ovata (Goto, 1894) Johnston, 1929, B. kintoki Iwata, 1990 is synonymised with B. elongata (Yamaguti, 1968) Egorova, 1997, B. sargocentron Zhang, Yang and Liu, 2001 is synonymised with B. hawaiiensis Yamaguti, 1968 and Pseudallobenedenia arabica Timofeeva, 1995 is synonymised with P. opakapaka Yamaguti, 1966. Benedenia madai Ishii and Sawada, 1938, B. pagrosomi Ishii and Sawada, 1938 and Allobenedenia pedunculata Raju and Rao, 1980 are considered species inquirendae. I examined an outbreak in aquaculture of the pathogenic benedeniine, Neobenedenia melleni (MacCallum, 1927) Yamaguti, 1963 and report this species for the first time in Australia. I examined the pathogenesis caused by this parasite by histology of host tissue. Possible routes of introduction to the farm in question are investigated and studies are detailed that should be undertaken in Australia to manage future outbreaks of N. melleni. Capsalids usually colonise new hosts by an infective ciliated oncomiracidium. An experiment was conducted to ascertain whether cleaner fish Labroides dimidiatus could transfer adult skin-parasitic monogeneans from one host fish to another. I have shown that transmission of adult monogeneans between two individuals of Hemigymnus melapterus by cleaner fish can occur. In coral reef environments, frequent contact between unrelated hosts involved in cleaning interactions might create previously unsuspected evolutionary pressures. I discuss the implications of this discovery for host-specificity and lateral transmission of monogeneans. General biological studies of monogeneans are necessary to understand the evolution, pathology, epidemiology, phylogeny and taxonomy of these parasites. My study provides a taxonomic system which, when applied to other capsalid subfamilies, should help prevent errors and create a clear system of classification for the entire family.

Thesis (Ph. D.)--University of Adelaide, 1985. / Includes bibliographical references.

Thesis (M. Sc.)--University of Adelaide, 1984. / Includes bibliographical references (leaves 111-137).

Thesis (Ph. D.)--University of Adelaide, 1983. / Includes bibliographical references (leaves 218-251).