Review: Ecology of the free-living stages of major trichostrongylid parasites of sheep

Significant developments over recent decades make it timely to review the ecology of the major gastrointestinal nematode (GIN) species of sheep. These include the relentless development and spread of anthelmintic resistance in all of the major sheep production regions of the world, and the consequent drive towards integrated parasite management (IPM) systems incorporating non-chemotherapeutic strategies such as grazing management. The success of such programs is dependent on a detailed understanding of the environmental influences on the free-living stages of the nematode lifecycle. Major reviews of the subject were conducted prior to 1980, however considerable work has been completed since, including the development of mathematical models describing the epidemiology of GIN infection. Knowledge of the temperature thresholds for free-living development has also improved, while investigations of moisture influences and interactions with temperature have allowed more effective exploitation of environmental effects for IPM. This review re-evaluates our understanding of the factors that determine the success or failure of the free-living phases of the lifecycle in light of these developments. Temperature and moisture are the dominant influences on the free-living stages of Haemonchus contortus, Teladorsagia circumcincta and Trichostrongylus colubriformis, with the effects of pasture conditions playing a significant modulating role. Early in the free-living phase, the developmental success of the three GIN species is limited by susceptibility to cold temperatures. In general, H. contortus is most susceptible, followed by T. colubriformis and then T. circumcincta. The length of the development cycle is dependent largely on temperature, with development rate increasing at warmer temperatures. However, in order for development to proceed to the infective larval stage, addition of moisture is generally required. There has been considerably less work quantifying the effects of moisture on free-living development, although it is clear that H. contortus is most susceptible to desiccation during the pre-infective stages. Once the infective stage is reached, the influences of temperature and moisture on survival are less important, resulting in considerable survival times under conditions lethal to pre-infective stages. However, hot, dry conditions can be lethal for infective larvae of all three species, while extreme cold is also lethal with significant species variatio