Aedes aegypti disregard humidity-related conditions with adequate nutrition.

Weather variations have clear associations with the epidemiology of dengue fever and populations of Aedes aegypti mosquitoes. Data on humidity associations, however, lags with respect to its effect on host-biting, nectar-seeking and survival. This experimental study on Ae. aegypti, sourced from the arid tropics, investigated the effect of low and high relative humidity and diet in relation to host-biting, temporal variations in feeding frequency, and mosquito mortality. In each environmental setting, 10 replicates, containing one male and five female mosquitoes, were challenged with different nutritional sources every six hours over 12 days. Results showed that host-biting did not diminish in low humidity and was six times higher than expected. Sucrose feeding was observed to significantly moderate host-biting and water alone was inadequate for survival. The high host-biting rates help to explain the intensity of dengue epidemics, while the ability of the mosquito to disregard adverse humidity-related conditions helps to explain how dengue epidemics in arid tropical regions can be just as devastating as those in the wet tropics.

Aedes aegypti disregard humidity-related conditions with adequate nutrition

Weather variations have clear associations with the epidemiology of dengue fever and populations of Aedes aegypti mosquitoes. Data on humidity associations, however, lags with respect to its effect on host-biting, nectar-seeking and survival. This experimental study on Ae. aegypti, sourced from the arid tropics, investigated the effect of low and high relative humidity and diet in relation to host-biting, temporal variations in feeding frequency, and mosquito mortality. In each environmental setting, 10 replicates, containing one male and five female mosquitoes, were challenged with different nutritional sources every six hours over 12 days. Results showed that host-biting did not diminish in low humidity and was six times higher than expected. Sucrose feeding was observed to significantly moderate hostbiting and water alone was inadequate for survival. The high host-biting rates help to explain the intensity of dengue epidemics, while the ability of the mosquito to disregard adverse humidity-related conditions helps to explain how dengue epidemics in arid tropical regions can be just as devastating as those in the wet tropics.

Associations of peak shifts in age--prevalence for human malarias with bednet coverage.

Effects of bednet coverage (C) on prevalence of malaria were analysed using data from 1990-92 from 9 Papua New Guinean villages. Effects of coverage varied by age, resulting in a shift in age of peak prevalence from 4.7 (C = 0%) to 11.6 (C = 100%) years for Plasmodium falciparum, from 3.4 to 4.9 years for P. vivax and from 11.0 to 16.8 years for P. malariae. In small areas with no bednets the age distribution of P. falciparum parasitaemia was like that of a holoendemic area. Where coverage was complete the pattern corresponded to mesoendemicity. Thus, protracted use of bednets can result in profound changes in the endemicity of malaria even when coverage is incomplete and without insecticide treatment. Average entomological inoculation rates (EIRs) estimated from indoor landing rates on individuals without bednets were 35, 12 and 10 infectious bites per person per annum for P. falciparum, P. vivax and P. malariae, respectively. Logistic regression analyses indicated that the EIR estimate for P. falciparum was related to prevalence of this species independently of effects of bednet coverage. However, the recent EIR still accounted for much less variation than did the bednets. A similar pattern was seen for P. malariae, while there were no significant relationships between the recent EIR and the parasite positivity for P. vivax. It is concluded that short-term variations in inoculation rate are not important determinants of parasite prevalence in this population.

Area effects of bednet use in a malaria-endemic area in Papua New Guinea.

Relationships between area coverage with insecticide-free bednets and prevalence of Plasmodium falciparum were investigated in 7 community-based surveys over a 33-month period in 1990-93 in 6 villages in the Wosera area of Papua New Guinea. Spatial patterns in circumsporozoite rates for P. falciparum, P. vivax isomorphs K210 and K247, and P. malariae, and the proportions of mosquito blood meals positive for specific human, goat, cat, dog and pig antigens were determined using ELISAs. P. falciparum prevalence in humans was better explained by bednet coverage in the immediate vicinity than by personal protection alone. Circumsporozoite rates for both P. falciparum and P. vivax were also inversely related to coverage with bednets. There was some increase in zoophagy in areas with high coverage, but relatively little effect on the human blood index or on overall mosquito densities. In this setting, protracted use of untreated bednets apparently reduces sporozoite rates, and the associated effects on prevalence are greater than can be accounted for by personal protection. Even at high bednet coverage most anophelines feed on human hosts, so the decreased sporozoite rates are likely to be largely due to reduction of mosquito survival. This finding highlights the importance of local vector ecology for outcomes of bednet programmes and suggests that area effects of untreated bednets should be reassessed in other settings.

Comparison between anopheline mosquitoes (Diptera: Culicidae) caught using different methods in a malaria endemic area of Papua New Guinea.

The mosquito sampling efficiency of CDC (Centers for Disease Control) miniature light traps hung adjacent to mosquito nets, was compared with that of both indoor and outdoor human-bait collections in ten villages in the Wosera area of Papua New Guinea. The most frequently collected anopheline in the matched indoor and light trap samples was Anopheles koliensis Owen, followed by A. punctulatus Dönitz, A. karwari (James), A. farauti Laveran (sensu lato), A. longirostris Brug and A. bancroftii Giles. All species were much less frequent in the light traps than in landing catches. The hypothesis that the numbers of mosquitoes in light traps are proportional to human landing catches was examined using regression models that allowed for sampling error in both entomological measurements. Light traps under-sampled A. punctulatus and A. farauti s.l. at high densities. The models indicated that the ratio of light trap to landing catch females of A. koliensis and A. karwari increased with increasing mosquito density. Light trap catches of A. longirostris were proportional to indoor landing rates but when outdoor landing rates were high this species was under-sampled by light traps. Numbers of A. bancroftii in light traps were found to be proportional to those in outdoor landing catches, but were negatively related to those attempting to bite indoors. Circumsporozoite positivity rates for both Plasmodium falciparum Welch and P. vivax (Grassi & Feletti) in A. punctulatus and A. farauti s.l. were significantly higher in light trap collections than in either indoor or outdoor landing catches, suggesting that light traps may selectively sample older mosquitoes of these species.

Mass treatment with ivermectin for filariasis control in Papua New Guinea: impact on mosquito survival.

Field studies were carried out to determine the impact of mass human treatment with ivermectin on the survival of anthropophagic mosquitoes of the Anopheles punctulatus complex (Diptera: Culicidae), the vectors of lymphatic filariasis and malaria in Papua New Guinea. In a village where mass treatment had been given, using 400 microg/kg ivermectin plus 6 mg/kg diethylcarbamazine citrate (DEC), we performed pre- and post-treatment collections of freshly blood-engorged mosquitoes from the same nine bedrooms. All blood-fed mosquitoes collected less than 4 days after mass treatment died within 9 days, whereas 67% of those collected before treatment survived for >9 days. Comparison (using the log-rank test) of the survival curves for mosquitoes collected (i) before treatment, (ii)<4 days after treatment, and (iii) 28 days after treatment, showed the survival rate of group (ii) to be significantly lower than the other two (chi2=176, df=2, P<0.0001). Pre- and post-treatment all-night landing catches showed no reduction in human biting rates in the experimental village. In another village, where people were mass treated with ivermectin (400 microg/kg) only, the survival rates of freshly blood-engorged An. punctulatus collected from bedroom resting-sites less than 1 day after treatment, were compared to similar collections carried out at the same time in a nearby village where people were not treated with ivermectin. The 48-h survival rate for the ivermectin-treated village was 31% compared to 94% for the other; this difference was highly significant (chi2=32.42, df=1, P<0.0001). Mosquitoes fed 2 months post-treatment with DEC or collected 38 days post-treatment with ivermectin had normal survival rates. We conclude that the duration of the systemic lethal effect of ivermectin on mosquitoes is insufficient to be of epidemiological significance in filariasis control programmes that are based on biannual and annual single-dose treatments, but might reduce vectorial capacity sufficiently to block epidemics of dengue or even malaria.

Effect of diet on biting, oviposition, and survival of Aedes aegypti (Diptera: Culicidae).

The effect of adult diet on host biting, sugar probing and water probing patterns, oviposition behavior, and survival of Aedes aegypti (L.) from Charters Towers, Australia, were tested in the laboratory. The 7 diets were as follows: (1) starvation, (2) water, (3) 10% sugar, (4) blood, (5) blood with water supplement, (6) blood with 10% sugar supplement, and (7) blood with 3% sugar supplement. Biting, probing, oviposition, and survival observations were made every 6 h (0600-0800 hours, 1200-1400 hours, 1800-2000 hours, 2400-0200 hours). Biting frequency on the blood with 3% sugar diet (0.26 feeds per mosquito per 6 h) and the blood with 10% sugar diet (0.23 feeds per mosquito per 6 h) was significantly less than on blood (0.51 feeds per mosquito per 6 h) and 10% sugar (0.40 feeds per mosquito per 6 h) alone. Biting frequency was not significantly different between blood with water (0.47 feeds per mosquito per 6 h) and blood alone (0.51 feeds per mosquito per 6 h). Biting and oviposition occurred throughout the day, peaking between 1800 and 2000 hours and between 2000 and 2400 hours, respectively. Biting frequency with a 3% sugar supplement decreased after the 1st oviposition cycle on day 6 compared with unsupplemented biting. The presence of sugar delayed or inhibited oviposition. Females with access to blood with water bit and oviposited concurrently on days 4-6, 8-9, and 11, indicating a 3-d gonotrophic cycle. Survival on blood alone was not significantly lower than survival on sugar and water supplemented diets. Behavior of the Charters Towers strain proved to be significantly influenced by diet, and biting occurred opportunistically without regard for previously observed crepuscular or diurnal rhythms. The biting frequencies observed were the highest yet recorded for this species, which indicates that the vectorial capacity of the Australian Ae. aegypti may be underestimated severely.

Adaptation of Aedes aegypti (Diptera: Culicidae) oviposition behavior in response to humidity and diet.

Effects of humidity and sugar concentration on the fecundity, temporal oviposition patterns and survival of a tropical strain of Aedes aegypti (L.) were investigated. Fecundity was significantly reduced by low humidity, but was not affected by sugar concentration. Low humidity caused a significant decrease in percentage survival after 19 days as compared to high humidity. Oviposition was inhibited by host availability for eight successive days. When access to a host was no longer provided, oviposition continued for 10 days in three to four distinct cycles without additional bloodmeals. Humidity stress and high sugar concentration caused oviposition to be delayed for one to four days, which is the typical duration of extreme low humidity periods in nature. These responses are hypothesized to protect the eggs of ovipositing females against the environmental hardships of periodic humidity stress and lack of hosts, thus enabling the perpetuation of the vector and the diseases it transmits in hot and dry seasons.

Multiple host-feeding and biting persistence of Aedes aegypti.

The effects of host activity, host defensive behaviour and biting persistence on multiple host-feeding by Aedes aegypti (L.) were investigated, in laboratory conditions, on human volunteers. In four different settings, four or five volunteer hosts displayed inactive, mildly active, mildly defensive and highly defensive behaviour. Into each of the four trials, each with 20 replicates, a single mosquito was released for a period of 10 min. In all settings, the median number of hosts sought per mosquito was constant, regardless of host activity and host defensive behaviour. Host defensive behaviour was thus not seen to exert a selective pressure on mosquitoes to abandon their hosts. No relationship was found between biting persistence and the number of hosts sought, indicating that biting persistence does not affect disease transmission. As one of the female hosts was consistently and significantly less attractive to host-seeking mosquitoes than any other volunteer in all four settings, it seems that some individuals are significantly less at risk from mosquito-borne pathogens than others.

Spatial and temporal variation in abundance of Anopheles (Diptera:Culicidae) in a malaria endemic area in Papua New Guinea.

Abundance of anophelines in 10 villages in the Wosera area of Papua New Guinea was monitored during 1990-1993. Of 85,197 anophelines collected in 1,276 paired indoor and outdoor landing catches, 40.4% were Anopheles koliensis Owen, 36.7% An. punctulatus Donitz, 14.3% An. karwari (James), 4.9% An. farauti s.l. Laveran, 3.1%, An, longirostris Brug, and 0.7% An. bancroftii Giles. Maps of average indoor biting rates were produced using a Bayesian conditional autoregressive model which allowed for heterogeneities in sampling effort over time and space. Differences in spatial distributions among species were observed among and within villages and were related to the distribution of larval habitats and vegetation. Abundance of An. punctulatus and An. koliensis decreased with distance from the main waterway and probably from a sago swamp forest at 6 villages in North Wosera. Abundance of An. punctulatus was associated negatively with those of An. farauti s.l., An. longirostris, and An. bancroftii. The latter 3 species also had relatively low ratios of indoor-to-outdoor biting rates, and earlier biting times than An. punctulatus. Human blood indices of at least 0.79 were observed for all species except An. bancroftii. Abundance of all 6 species was correlated temporally with recent rainfall, but An. koliensis, An. karwari, and An. longirostris showed greater temporal variability than the other species. An punctulatus and An. koliensis tended to occur together in time and space (index of association, I = 0.85). Weaker associations were seen between An. farauti s.l. and An. longirostris (I = 0.44) and An. koliensis and An. karwari (I = 0.34). The most frequently collected species occurred together and were concentrated near the Amugu river; the remaining species tended to occur together but in different parts of the Wosera area. The importance of understanding ecological requirements of the different Anopheles vectors and their association with key household and landscape features are discussed in relation to malaria transmission and control.

Efficacy of carbon dioxide, 1-octen-3-ol, and lactic acid in modified Fay-Prince traps as compared to man-landing catch of Aedes aegypti.

The attractants 1-octen-3-ol and lactic acid significantly decreased catches of Aedes aegypti in Townsville, Australia, by 50% in a controlled laboratory environment and by 100% in the field when compared to carbon dioxide baited bidirectional Fay-Prince trap catches. Evaluation of an omnidirectional alteration on a bidirectional Fay-Prince trap revealed no significant improvement in catch size when compared to both the bidirectional trap and man-landing catch (MLC). Cumulative evening MLC (1730-2000 h) was twice that of the morning MLC (0600-0830 h), which has implications on the precise estimation of the man-biting rate. The MLC sampling method is shown to be a quick, simple, effective and cheap alternative to expensive traps in areas not currently experiencing arbovirus transmission.

Heterogeneities in the transmission of infectious agents: implications for the design of control programs.

From an analysis of the distributions of measures of transmission rates among hosts, we identify an empirical relationship suggesting that, typically, 20% of the host population contributes at least 80% of the net transmission potential, as measured by the basic reproduction number, R0. This is an example of a statistical pattern known as the 20/80 rule. The rule applies to a variety of disease systems, including vector-borne parasites and sexually transmitted pathogens. The rule implies that control programs targeted at the "core" 20% group are potentially highly effective and, conversely, that programs that fail to reach all of this group will be much less effective than expected in reducing levels of infection in the population as a whole.

The Mossie-Buster: a hose-driven insecticide delivery tool for the control of container-breeding mosquitoes.

A novel insecticide delivery tool, the Mossie-Buster, was recently developed to control larval populations from urban breeding sites in Townsville, Australia. This functional and user-friendly control device directly targets the main breeding sites of Aedes aegypti (L.) with a focused delivery of insecticides. The Mossie-Buster comprises a commercially available device and attachments that mix an insecticide solution into the flow of water emitted from a hose. Attached to the device is a trigger for controlled insecticide release. Preliminary laboratory and field trials demonstrated the tool to be effective in eliminating all Ae. aegypti present in various typical breeding containers in different environmental conditions for a minimum of 2 wk in exposed areas to 3 months in an unexposed area.

The gecko: an environmentally friendly biological agent for mosquito control.

Laboratory experiments with Aedes aegypti mosquitoes investigated the effects of light, mosquito density and physiological state on predation rates by the Australian gecko Gehydra dubia and the exotic Asian house gecko Hemidactylus frenatus. For both gecko species a positive correlation was demonstrated between prey density and the predation rate. Using Ae.aegypti males and unfed females as prey in a terrarium (0.054 m3), consumption rates reached 76-108/day for G.dubia and 63-109/day for H.frenatus, with significantly more female mosquitoes than males being eaten in most experiments. Comparing dark with semi-illuminated conditions no consistent contrast of predation rate was demonstrated. Gehyra dubia predation rates on various Australian mosquito species were compared in an experimental room (32 m3) for 24 h exposure with photoperiod L:D 12:12 h Five photophilic species (Aedes vigilax, Anopheles annulipes, Coquillettidia xanthogaster, Culex annulirostris, Cx sitiens) suffered 78-100% predation, compared with only 33-53% predation of four non-photophilic species: Aedes aegypti, Ae.notoscriptus, Ae.vittiger and Cx quinquefasciatus. This demonstrates the potential benefit of domestication for geckoes that learn to hunt at light. When offered a mixture of unfed, freshly blood-fed and gravid females of Ae.aegypti in an illuminated terrarium, both gecko species consumed significantly more unfed than fed or gravid female mosquitoes, presumably because the latter rested whereas the former foraged more actively. H.frenatus consumed significantly more mosquitoes of all categories than did G.dubia per 24 h: unfeds 5.1 +/- 0.1% v. 4.5 +/- 0.5%, blood-feds 4.5 +/- 0.3% v. 4.0 +/- 0.5% and gravids 4.8 +/- 0.4% v. 3.9 +/- 0.5%. Possible relevance of these results to competitive displacement of G.dubia by H.frenatus is discussed.

Sustainability of a successful malaria surveillance and treatment program in a Runggus community in Sabah, east Malaysia.

The district of Kudat has one of the highest and most persistent malaria transmission levels in Sabah, Malaysia, with annual parasite incidence of 102 per 1,000 inhabitants per year. Due to this situation and the failure of DDT spraying to control malaria, a community participation health program (Sukarelawan Penjagaan Kesihatan Primer or SPKP) was developed as an adjunct to current anti-malarial measures during 1987-1991. SPKP is made up of unpaid community workers known as village health volunteers (VHVs). VHVs are selected by a village development and security committees training and supervision a member of the Vector-Borne Diseases Control Program (VBDCP). The beneficiaries of SPKP consisted primarily of Runggus people and other remote, and mobile populations who visit the home of a VHV for diagnosis and treatment. This group of febrile patients and their children who attend a participating school submit finger prick blood and personal details to the VHV. and receive a presumptive treatment for malaria. Thick and thin blood smears are examined by a VBDCP microscopist who then prepare and forward a radical or curative treatment to the VHV so that it can be administered to the microscopically-positive patient free of charge. Between June 1987 to June 1991, VHVs from 32 kampungs (villages) and 22 schools collected 56,245 slides representing 24.7% of total slide collection compared to 74.9% collected by passive case detection (PCD) posts in health centers and district hospital. The average volunteer treated 11.8 (range 10.4-13.4) and 31.4 (range 26-49) patients per month in kampungs and schools respectively. In contrast, non-SPKP posts in a district hospital, health centers and flying doctor service treated an average of 616.3 patients per month (range 134.8-1032.8). The slide positivity rate of blood smears taken by VHVs was 8.43% compared with 7.37% for non-SPKP posts. Average slide collection and slide positivity rates varied considerably from one community to another, despite their close geographic proximity. The monthly number of VHV-diagnosed patients from the school and kampungs communities and the monthly number of true malaria patients in the two groups were significantly correlated. Sustainability of SPKP was linked to an ongoing process of social change which involved co-operative networking between the government health sector and the community. This in turn provided a stimulus for malaria abatement efforts. When Runggus people themselves control and maintain ownership of community-based malaria programs, the function of SPKP as a malaria surveillance system and an antimalarial drug distribution network is vastly improved.

Comparative effects of permethrin-impregnated bednets and DDT house spraying on survival rates and oviposition interval of Anopheles farauti No. 1 (Diptera:Culicidae) in Solomon Islands.

Human-biting, CDC light trap and pig-baited collections were used to monitor changes in the abundance and parity rate of Anopheles farauti No. 1 mosquitoes in three Solomon Islands villages for 30 consecutive nights. Houses in one of the villages were sprayed with DDT, another village was unsprayed but its inhabitants used permethrin-treated bednets and the third village was left completely untreated. Mosquitoes collected each day were dissected to determine follicular maturation and parity. In the second year of vector control, survival rates (determined by time-series analysis of the landing catches and biting parous population) were significantly lower in the village using permethrin-treated bednets than in the other two villages. There was no difference in the estimated survival rates between the untreated and DDT-sprayed villages. The oviposition cycle was extended to 4 days in the permethrin-treated village compared with 3 days in the other villages. The expected infective life was 1.75-fold longer in the untreated village than in the permethrin-treated village. The epidemiological implications of replacing DDT spraying with permethrin for malaria control are discussed.

Observations of human behavior influencing the use of insecticide-impregnated bednets to control malaria in Sabah, Malaysia.

Observational and survey methodologies were used to probe human behavioral factors influencing the use of insecticide-impregnated bednets to control malaria in rural Sabah, Malaysia. One aim was to investigate why a field trial of such nets in an interior area yielded disappointing results. A second aim was to gather baseline data prior to a field trial proposed for a coastal area. Interior villagers reported a significantly higher net usage rate than that observed directly, suggesting that subject self-reports need to be validated in some way. The poor results of the interior field trial appeared related to reluctance to regularly use nets, which were not in wide use previously. Prospects for reducing malaria transmission through bednets appeared better for the coastal area since nearly half of observed villagers were sleeping in them. However, significantly more coastal than interior villagers were observed watching television at night, an activity that may increase malaria risk by keeping villagers awake and out of bednets.

Impact of permethrin-impregnated mosquito nets compared with DDT house-spraying against malaria transmission by Anopheles farauti and An.punctulatus in the Solomon Islands.

In villages of northern Guadalcanal in the Solomon Islands, where the predominant malaria vector is An.farauti No. 1 and An.puctulatus is also involved, malaria transmission rates were compared for three zones: (1) non-intervention: 438 people in seventeen villages; (2) residual DDT house-spraying two cycles per year: 644 people in thirty villages; (3) bednets impregnated with permethrin 0.5 g/m2 twice per year, used by 580 people in sixteen villages. Regular DDT spraying in zones 1 and 3 had been withdrawn 18 months previously. Malariological blood smear surveys of children aged 1-9 years in August 1986 to January 1987 showed a mean baseline malaria parasite rate of 38% (32/84). By February 1988, 18 months after introduction of impregnated bednets, the Plasmodium falciparum infection rate in children was lowest in the zone using impregnated bednets (21% of 29), intermediate in the untreated zone (29% of 34) and highest in the DDT zone (46% of 53), but these differences were not statistically significant. P. vivax infection rates were 9-14%. Using ELISA tests for malaria circumsporozoite antigen in the vectors, overall positivity rates were 0.7% of 49,902 An.farauti and 2.54% of 118 An.punctulatus, comprising 228 P.falciparum and 124 P.vivax infections. In the study zones, vector positivity rates were 0.93% of 31,615 An.farauti in the untreated zone; 0.32% of 16,883 An.farauti in the DDT zone; 0.07% of 1404 An.farauti and 2.54% of 118 An.puctulatus in the impregnated bednet zone. here was no significant correlation between malaria parasite rates in the vectors and the children.(ABSTRACT TRUNCATED AT 250 WORDS)

Population genetic analysis of host seeking and resting behaviors in the malaria vector, Anopheles balabacensis (Diptera: Culicidae).

During the intermonsoon period from mid-September to mid-October 1986, wild-caught Anopheles balabacensis Baisas females were marked and released in a host-choice experiment. Association between capture and recapture of marked mosquitoes from human and bovid hosts and blood meal host identification of recaptured females were determined on a daily basis. Although the mark-recapture and blood meal data indicated behavioral heterogeneity between buffalo and human biters, restriction endonuclease fragment length polymorphism analysis revealed no differences in repeat sequence profiles. Doubly-marked recaptures strongly indicated a "learning" component involved in a separate host preference experiment. In a "habitat loyalty" experiment conducted in January 1987, females of An. balabacensis preferentially returned to the resting sites (indoor surfaces and exit traps) where they were first caught. Of nine isozyme loci found to be polymorphic, the genotypic frequencies of Esterase-3 and Isocitrate dehydrogenase-3 were different in "faithfully" endophilic and exophilic subpopulations. Genetic heterozygosity, as determined by polyacrylamide gel electrophoresis, was greater in exophilic than endophilic population components. These results confirm that genetic and learning components can significantly influence house resting and host seeking behavior and may contribute to local epidemiological patterns of malaria transmission observed in Sabah, Malaysia.

Estimation of survival rate and oviposition interval of Anopheles balabacensis mosquitoes from mark-recapture experiments in Sabah, Malaysia.

An exophilic population of the vector mosquito Anopheles balabacensis Baisas was investigated in two mark-recapture studies (16.ix-13.x.1986 and 6-26.i.1987) at an inland, foothill village in Sabah, Malaysia. Wild female mosquitoes were intercepted as they came to feed on man or buffalo, given a bloodmeal, marked with fluorescent dust and released. The recapture rate was about 12%. A new method of analysis is proposed which uses cross-correlation and a time series model. The estimated survival per oviposition cycle was 0.48-0.54 and the oviposition cycle interval 2-3 days.