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Chapter 1 Literature Review1 1 Chapter 1 Literature review1 1.1 Introduction Australian mosquitoes of the genera Aedes (Kay 1982, Watson and Kay 1998, Boyd and Kay 1999, Watson and Kay 1999b), Culex (Boyd and Kay 2000) and Verrallina (Ryan et al. 2000) have been implicated in the transmission of over 500 arboviruses, including: Ross River, Barmah Forest and dengue viruses. Since 1991, over 77,000 human infections with these viruses have been reported in Australia (Communicable Diseases Network – Australia New Zealand – National Notifiable Diseases Surveillance System), leading to considerable public health concern. As there are no vaccines against these viruses, except for Japanese encephalitis, the community relies on personal protection against mosquito exposure, and on mosquito control to minimise the transmission of disease (Russell 1998a, Russell and Kay 2004). The primary focus of mosquito control in Australia is reducing larval populations and is undertaken by local government authorities in Queensland, but also by State authorities, to varying degrees, in the remaining states. Successful larval control has involved the integration of a number of techniques including: • Public education: the community is urged to remove containers holding water from household gardens and to use personal protection; applicable to Aedes aegypti (Linnaeus) and Aedes notoscriptus (Skuse). 1 Aspects of this review, regarding the use of Bacillus thuringiensis var. israelensis for the control of immature mosquitoes, have been published in the proceedings of the 5th Pacific Rim conference on the biotechnology of Bacillus thuringiensis, pp 57 – 72 (Appendix A). 2 • Physical modification of habitats (runnelling): involves digging shallow channels to link isolated pools with the tidal source, this process increases the movement of low amplitude tides, regularly flushing the saltmarsh (Hulsman et al. 1989); widespread use against Aedes vigilax and some use against Aedes camptorhynchus (Thomson). • Biological control: involves introducing predators (e.g. copepods and fish) into larval habitats (Hurst 2004, Kay and Nam 2005); used primarily against Ae. aegypti, but native fish are applied to constructed wetlands against freshwater species, mainly Culex annulirostris. • Insecticides: include microbials and insect growth regulators, which have been used for the global control of mosquitoes for more than 20 years (Mian and Mulla 1982, Margalit and Dean 1985); broadly used for saltmarsh/mangrove and freshwater species. This thesis will focus on improving the efficacy of insecticide use, while minimising environmental impacts. As such, the following review outlines: the significance of vector- borne disease in Australia, the habitats used by mosquito larvae, microbial insecticide and insect growth regulator use, and possible environmental impacts. 1.2 Mosquito-borne disease in Australia Mosquitoes transmit three subgroups of pathogens: 1) filaria, 2) malaria protozoans, and 3) arboviruses. Human filariasis is no longer a medical concern in Australia; it was eradicated in the 1950s (Boreham and Marks 1986). Malaria was eradicated in 1981; however, up to 1,000 cases are imported annually by infected travellers, occasionally resulting in local transmission (Brookes et al. 1997, Russell 1998a). Arboviruses are of primary concern in Australia. Human disease results from infection with Ross River, Barmah Forest, dengue, Murray Valley encephalitis, Kunjin and Japanese encephalitis viruses (Table 1.1). Ross River and Barmah Forest viruses are closely related Alphaviruses, and can lead to the onset of debilitating polyarthritis (Kay and Aaskov 1989, Mackenzie et al. 1998, Russell and Kay 2004). The severity and duration of symptoms due to Ross River and Barmah Table 1.1: The larval habitats and distribution of Australian mosquitoes and the associated human diseases. Associated Larvalhabitat Species Distributiona arbovirusesb Reference Freshwater Ground-pool Cx. annulirostris All states/territories RRV, BFV, MVEV, Ritchie et al. 1997b, Boyd and Kay 2000, Ryan JEV, KUNV et al. 2000 Ae. procax QLD, NSW, VIC RRV, BFV Ritchie et al. 1997b, Ryan et al. 2000, Ryan and Kay 1999 Cq. linealis QLD, NSW RRV, BFV Jeffery et al. 2002, Kay and Standfast 1987 Container Ae. aegypti Nth QLD DENV Watson and Kay 1999b, Knox et al. 2003 Ae. notoscriptus All states/territories RRV Ritchie et al. 1997b, Watson and Kay 1998 Both Cx. quinquefasciatus All states/territories Nuisance Russell 1998a Saltmarsh/mangrove Ae. vigilax QLD,NSW,NT,WA,SA RRV,BFV,MVEV Kay 1982, Dale et al. 1986, Ritchie et al. 1997b, Boyd and Kay 1999 Ae. camptorhynchus NSW, TAS, SA, VIC, WA RRV, BFV Campbell et al. 1989, Kay and Standfast 1987 Cx. sitiens QLD,NSW,NT,WA RRV Ritchie et al. 1997b Ve. funerea QLD, NSW, NT RRV Ritchie et al. 1997b, Ryan et al. 2000 Notes Literature review 3 a. NT = Northern Territory; QLD = Queensland; WA = Western Australia; NSW = New South Wales; SA = South Australia; VIC = Victoria; TAS = Tasmania. b. RRV = Ross River virus; BFV = Barmah Forest virus; DENV = Dengue virus; KUNV = Kunjin virus; MVEV = Murray Valley encephalitis virus; JEV = Japanese encephalitis virus. 4 Forest virus infection vary greatly among patients and are usually resolved in three months (Mackenzie and Smith 1996, Flexman et al. 1998, Harley et al. 2001). On average 4,800 cases of Ross River virus are reported per year (Miller et al. 2003, Kelly- Hope et al. 2004). Ross River virus has been isolated from wild caught mosquitoes in most regions of Australia. The main vectors are: Ae. camptorhynchus (Campbell et al. 1989), Ae. vigilax (Kay 1982, Ryan et al. 2000), Ae. notoscriptus (Watson and Kay 1998), Cx. annulirostris (Ryan et al 2000), Verrallina funerea (Theobald) (Ryan et al. 2000), Aedes procax (Skuse) (Ritchie et al. 1997b, Ryan et al. 2000), and Coquillettidia linealis (Skuse) (Jeffery et al. 2002). The differing geographical distribution of each vector leads to areas of regional transmission. There are over 700 human infections with Barmah Forest virus reported per year (Miller et al. 2003). The potential vectors of Barmah Forest virus are: Cx. annulirostris (Boyd and Kay 2000), Ae. camptorhynchus (Kay and Standfast 1987), Ae. vigilax (Boyd and Kay 1999), Ae. procax (Ryan and Kay 1999), and Cq. linealis (Kay and Standfast 1987). Infection with dengue can lead to a non-specific febrile illness in most patients, ranging up to a fatal hemorrhagic disease (Gubler 1998b). Aedes aegypti is the major recognised dengue vector on mainland Australia (Watson and Kay 1999b, Knox et al. 2003), and is established in northern Queensland (Sinclair 1992). In addition, members of the Aedes scutellaris group (known dengue vectors), are established on the Torres Strait islands and in Cape York (Rosen et al. 1985, Russell 1998a). Over the past decade there have been almost annual outbreaks of dengue (10 to 3,000 cases per outbreak), each presumably started via infected travellers (Russell 1993, 1998a). In 2005, Aedes albopictus (Skuse) invaded the Torres Strait and threatens to colonise mainland Australia, further increasing the risk of dengue transmission (Russell et al. 2005). Murray Valley encephalitis and Kunjin viruses are closely related flaviviruses, both causing severe fever and encephalitis, and with Murray Valley encephalitis some fatalities (Marshall 1988). The main vector of Murray Valley encephalitis and Kunjin virus is Cx. annulirostris (Kay et al. 1984), although other species such as Culex australicus Dobrotworsky & Drummond (Russell 1998a) and Aedes normanensis (Taylor) (Broom et al. 1989) may have subsidiary roles. Over the past decade, isolated cases of both Murray Valley encephalitis Literature review 5 and Kunjin have been recorded and confined to tropical Western Australia, the Northern Territory and north Queensland (Marshall 1988, Miller et al. 2003, Russell and Kay 2004). Recent research by Hall et al. (2003) has shown that Kunjin is a subtype of West Nile virus. West Nile virus was introduced into the USA in 1999 and to date has caused over 19,700 human infections and over 780 fatalities (Centres for Disease Control and Prevention – Division of Vector-Borne Infectious Diseases). The first recognised outbreak of Japanese encephalitis in Australia occurred in 1995, when three clinical cases (two fatal) were reported at Badu Island in the Torres Strait (Hanna et al. 1996). A further two cases were reported in 1998, one in the Torres Strait and one on the mainland at the Mitchell River (Hanna et al. 1999). Virus isolations have been made from wild-caught Cx. annulirostris and Culex sitiens Wiedemann, indicating these species to be potential vectors in Australia (Ritchie et al. 1997a, Hanna et al. 1999, Johansen et al. 2001, van den Hurk et al. 2003, 2006). Laboratory trials have indicated that Culex gelidus Theobald, Culex quinquefasciatus Say and Ae. notoscriptus may also play roles in localised transmission (van den Hurk et al. 2003). 1.3 Habitats of mosquito larvae In Australia, mosquito control has focused on reducing larval populations, rather than targeting adults which are more dispersed (WHO 1997). There are two broad classes of larval habitats: freshwater (ground-pool and container) and saltmarsh/mangroves. Each habitat type is colonised by the larvae of specific mosquito species, and mosquito control operations must be tailored to suit each habitat. 6 1.3.1 Freshwater 1.3.1.1 Ground-pool In Australia, ground-pool freshwater habitats include permanent and temporary water bodies, e.g. lakes, swamps, streams, wheel ruts
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