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A factorial experiment (3 x 4 x 2 x 3) was conducted in programmable incubators to investigate interaction between the effects of rainfall amount, rainfall distribution and evaporation rate on development of Haemonchus contortus to L3. Sheep faeces containing H. contortus eggs were incubated on sterilised soil under variable temperatures typical of summer in the Northern Tablelands of NSW, Australia. Simulated rainfall was applied in 1 of 3 amounts (12, 24 or 32 mm) and 4 distributions (a single event on the day after deposition, or the same total amount split in 2, 3 or 4 equal events over 2, 3 or 4 days, respectively). Samples were incubated at either a Low or High rate of evaporation (Low: 2.1-3.4 mm/day and High: 3.8-6.1 mm/day), and faeces and soil were destructively sampled at 4, 7 and 14 days post-deposition. Recovery of L3 from the soil (extra-pellet L3) increased over time (up to 0.52% at day 14) and with each increment of rainfall (12 mm: <0.01%; 24 mm: 0.10%; 32 mm: 0.45%) but was reduced under the High evaporation rate (0.01%) compared with the Low evaporation rate (0.31%). All rainfall amounts yielded significantly different recoveries of L3 under Low evaporation rates but there was no difference between the 12 and 24 mm treatments under the High evaporation rate. The distribution of simulated rainfall did not significantly affect recovery of infective larvae. Faecal moisture content was positively associated with L3 recovery, as was the ratio of cumulative precipitation and cumulative evaporation (P/E), particularly when measured in the first 4 days post-deposition. The results show that evaporation rate plays a significant role in regulating the influence of rainfall amount on the success of L3 transmission.

The key influence of moisture availability on development of the free-living stages of Haemonchus contortus is well-documented, although quantitative relationships between moisture variables and development are poorly defined. A factorial experiment (3x2x2) was conducted in programmable incubators to determine the effects of amount and distribution of simulated rainfall on H. contortus development at low evaporation rates (approximately 2 mm/day) under temperatures typical of summers on the Northern Tablelands of NSW, Australia. Sheep faeces containing H. contortus eggs were placed in experimental units containing sterile soil and had one of the three amounts (12, 24 and 32 mm) of simulated rainfall applied, in either a single event on the day after deposition (d 1) or three split events over 6 days (d 1, 3 and 6). Treatments were applied either in week 1 only, or in weeks 1 and 2. Recovery of infective larvae (L3) at 4, 7 and 14 days post-contamination increased with each incremental amount of simulated rainfall over the range of 12-32 mm and was significantly higher under the single "rain" event (2.8%), compared with the three smaller, split events (1.9%). The second application of simulated rainfall in week 2 had only a very small influence on L3 recovery, suggesting that the majority of development to L3 occurred in response to simulated rainfall events in the first 7 days. Both faecal moisture content and the cumulative ratio of precipitation and evaporation (P/E) were strongly and positively correlated with recovery of L3. Recovery of L3 from treatments, which received simulated rainfall in week 1 only was best described by P/E at d 5. Whether the relationships observed in this study hold under field conditions, where macroclimatic conditions such as evaporation rate are substantially more limiting to free-living development, is yet to be determined.

Three studies were undertaken to determine the effects of amount, timing and distribution of simulated rainfall on the developmental success of Haemonchus contortus. Faeces containing H. contortus eggs were deposited onto pasture plots under a rainfall-activated retractable roof which eliminated incident rainfall. In October (spring) 2004 and January (summer) 2005, the effects of amount (6, 12, 18 or 24 mm) and timing (1, 4, 8 or 15 days post-faecal deposition) of a single simulated rainfall event was investigated via manual application of water to plots. More H. contortus pre-infective larvae (L1 and L2) developed under the d 1 simulated rainfall treatment than later treatments. There was no effect of rainfall amount on development in either experiment, and negligible development to infective larvae (L3). In February (summer) 2006, the effects of amount (12, 24 or 32 mm) and distribution (single event or three smaller but equal split events over 32 h) of simulated rainfall events was investigated with water applied via sprinkler. In this experiment L3 were recovered from the herbage in one-third of the plots harvested, however recovery was low (0.08% of eggs deposited) and there were no treatment effects. Recovery of L1 and L2 from faeces increased with simulated rainfall amount at d 4, and more L1 and L2 were recovered from the split distribution treatment at d 4. The results indicate that moisture conditions soon after faecal deposition are key determinants of H. contortus development success, with significant penalties on development when simulated rainfall was applied 7 days or more post-deposition, and when the duration of simulated rainfall was short. High rates of evaporation during both summer experiments resulted in rapid drying of the micro-environment and this appears to have limited development to L3.

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 variation. While the existing body of knowledge is substantial, the interpretation of many studies and comparison between them is complicated by inadequate description of, or variation in, the environmental measurements used. Confounding the effects of environmental variables on development to infective stage is the migration of larvae from the faeces and subsequent survival on pasture. There is a need to build on recent efforts to explore interaction between the effects of temperature and moisture, and also the trend to more closely simulate field conditions in laboratory studies. We propose a logical framework for future ecological investigations to overcome some of these problems, facilitate the development of a more integrated dataset on the subject and improve prediction of free-living development.