Laboratory transmission of La Crosse virus by Ochlerotatus j. japonicus (Diptera: Culicidae)

Ochlerotatus j. japonicus, a recent introduction to the United States, was studied to determine its capability to serve as a vector of La Crosse (LAC) virus. A field-collectedpopulation of Ochlerotatus triseriatus, the primary vector of LAC virus, was similarly tested for comparison. After Oc. j. japonicus ingested virus fromhamsters with viremias of 10(3.6-5.4) plaque-forming units (PFU)/ml of blood, its estimated transmission rates were 35-88%. These rates were slightly lower than, though similar to, those for Oc. triseriatus, 75-100%. Viral titers in Oc. j. japonicus peaked at approximately 10(5.5) PFU/ mosquito about 7 d after ingesting blood meal in which the concentration of LAC virus was PFU/ml of blood; virus had disseminated from the midgut in 100% (8/8) of these specimens. These data, combined with the close association between the habitats of Oc. j. japonicus and Oc. triseriatus and the reported expansion of the range of this newly discovered species in the eastern United States, indicate that Oc. j. japonicus could function as an additional vector of LAC virus.     

Mosquito species richness, composition, and abundance along habitat-climate-elevation gradients in the northern Colorado Front Range

We exploited elevation gradients (1,500-2,400 m) ranging from plains to montane areas along the Poudre River and Big Thompson River in the northern Colorado Front Range to determine how mosquito species richness, composition, and abundance change along natural habitat-climate-elevation gradients. Mosquito collections in 26 sites in 2006 by using CO2-baited CDC light traps yielded a total of 7,136 identifiable mosquitoes of 27 species. Commonly collected species included Aedes vexans (Meigen) (n = 4,722), Culex tarsalis Coquillett (n = 825), Ochlerotatus increpitus (Dyar) (n = 546), Ochlerotatus trivittatus (Coquillett) (n = 303), Aedes cinereus Meigen (n = 280), Ochlerotatus melanimon (Dyar) (n = 146), Ochlerotatus dorsalis (Meigen) (n = 67), Culiseta inornata (Williston) (n = 52), Ochlerotatus pullatus (Coquillett) (n = 38), Ochlerotatus spencerii idahoensis (Theobald) (n = 37), and Culex pipiens L. (n = 29). Species richness was highest in plains habitats at elevations below 1,600 m. Numerous species were found exclusively or predominantly at low elevations below 1,700 m [Anopheles earlei Vargas, Anophelesfreeborni Aitken, Coquilletidia perturbans (Walker), Culex erythrothorax (Dyar), Cx. pipiens, Culex territans Walker, Oc. dorsalis, Ochlerotatus hendersoni (Cockerell), Oc. melanimon, and Oc. trivittatus], whereas others occurred predominantly at high elevations above 2,300 m [Ae. cinereus, Culiseta incidens (Thomson), Culiseta morsitans (Theoblad), Ochlerotatus cataphylla (Dyar), Ochlerotatus intrudens (Dyar), Oc. pullatus, and Ochlerotatus punctor (Kirby)]. Ae. vexans and Cx. tarsalis were abundant in the plains (< 1,600 m; mean June-August temperature > 19.5 degrees C), occurred at low abundances in foothills and low montane areas (1,610-1,730 m; 18.0-19.5 degrees C), and they were collected only sporadically in montane areas above 1,750 m (mean June-August temperature < 17.5 degrees C). These findings suggest that future climate warming may lead to shifts in distribution patterns of West Nile virus vectors (e.g., Cx. tarsalis) toward higher elevations in Colorado.     

Prevalence and distribution of Ochlerotatus japonicus (Diptera: Culicidae) in two counties in southern New York State

The seasonal occurrence and abundance of a newly introduced mosquito in the United States, Ochlerotatus (Finlaya) japonicus (Theobald), are reported for Westchester and Putnam Counties in southern New York State. Adult mosquitoes were sampled at 39 sites distributed throughout the two counties. CDC (Centers forDisease Control and Prevention) light traps (2,107 trap nights) and gravid traps (1,813 trap nights) were used to collect a total of 44,428 mosquitoes from May through October 2000. Oc. japonicus was found at 97.4% of sites sampled and accounted for 9.8% of all specimens collected. Oc. japonicus was recovered from 1.5% (n = 326) of the light trap collections and 18.1% (n = 4,026) of the gravid trap collections. Although gravid traps collected significantly more specimens than light traps, the seasonal activity patterns measured by each trap type were congruent. In all, 30 mosquito species were collected. Unlike other Ochlerotatus or Aedes species, Oc. japonicus was collected throughout the study, indicating a broad seasonal activity period. There was significant regional variation in Oc. japonicus abundance, with higher trap densities occurring in the northern-most trapping sites. This study demonstrates that Oc. Japonicus is established in southern New York State.     

Establishment and abundance of a recently introduced mosquito species Ochlerotatus japonicus (Diptera: Culicidae) in the Southern Appalachians, USA

Ochlerotatus (Finlaya) japonicus japonicus (Theobald; =Aedes japonicus; see Reinert 2000) (Diptera: Culicidae), is an introduced Asian mosquito species first detected in the northeastern United States in 1998. Since its initial discovery, this species has spread to many neighboring states and Canada. It was first identified in North Carolina, South Carolina, and Georgia in 2003. Larval surveys were conducted during 2005 and 2006 in the Southern Appalachians, U.S.A., on a variety of natural and artificial containers to determine extent of Oc.japonicus establishment and larval habitat requirements. Detritus amounts were measured in each container habitat, and co-occurring larval mosquito species were examined as indicators of potential interspecific competition. Data reveal that Oc. japonicus was the most abundant container-inhabiting mosquito species in this survey. It exhibits flexibility in its use of container oviposition habitats, it can persist in a wide range of conditions, and it co-occurs with a wide range of larval mosquito species. Its rapid and successful establishment warrants continued monitoring, because its potential role as a nuisance species or arbovirus vector remains unknown.     

Establishment of Ochlerotatus japonicus (Diptera: Culicidae) in Ontario, Canada

Ochlerotatus (Finlaya) japonicus (Theobold) is newly established in Ontario, Canada. It was first discovered in 2001 during the province-wide West Nile virus mosquito surveillance program implemented by the Ontario Ministry of Health and Long Term Care. Although the numbers of adults trapped in CDC light traps were low in 2001 and 2002, they increased during the 2003 and 2004 surveillance seasons. Oc. japonicus larvae also have been collected in large numbers in the Niagara Peninsula in a variety of natural and artificial containers. The number of health units with records for Oc. japonicus has increased over the 4 yr of surveillance, illustrating the ability of this species to rapidly extend its range. As a potential arboviral bridge vector, its establishment in Ontario requires further study and should be considered a public health concern.     

Experimental transmission of eastern equine encephalitis virus by Ochlerotatus j. japonicus (Diptera: Culicidae)

We evaluated the potential for Ochlerotatus j. japonicus (Theobald), a newly recognized invasive mosquito species in the United States, to transmit eastern equine encephalitis (EEE) virus. Aedes albopictus (Skuse) and Culex pipiens (L.) were similarly tested for comparison. Ochlerotatus j. japonicus and Ae. albopictus became infected and transmitted EEE virus by bite after feeding on young chickens 1 d after they had been inoculated with EEE virus (viremias ranging from 10(7.0-8.7) plaque-forming units [PFU]/ml of blood). No Cx. pipiens (n = 20) had detectable levels of virus 14 d after feeding on an EEE-virus infected chicken with a viremia of 10(8.1) PFU per ml of blood. Depending on the viral titer in the donor chicken, infection rates ranged from 55-100% for Oc. j. japonicus and 93-100% for Ae. albopictus. In these two species, dissemination rates were identical to or nearly identical to infection rates. Depending on the viral titer in the blood meal, estimated transmission rates ranged from 15 to 25% for Oc. j. japonicus and 59-63% for Ae. albopictus. Studies of replication of EEE virus in Oc. j. japonicus showed that there was an "eclipse phase" in the first 4 d after an infectious blood meal, that viral titers peak by day 7 at around 10(5.7) per mosquito, and that virus escaped the mid-gut as soon as 3 d after the infectious blood meal. These data, combined with the opportunistic feeding behavior of Oc. j. japonicus in Asia and the reported expansion of its range in the eastern United States, indicate that it could function as a bridge vector for EEE virus between the enzootic Culiseta melanura (Coquillett)-avian cycle and susceptible mammalian hosts.     

Point source inoculation of mesocyclops (Copepoda: Cyclopidae) gives widespread control of Ochlerotatus and Aedes (Diptera: Culicidae) immatures in service manholes and pits in north Queensland,Australia

This study details the novel application of predacious copepods, genus Mesocyclops, for control of Ochlerotatus tremulus (Theobald) group and Aedes aegypti (L.) mosquito larvae in subterranean habitats in north Queensland, Australia. During June 1997, 50 Mesocyclops sp. 1 were inoculated into one service manhole in South Townsville. Wet season rainfall and flooding in both 1998 and 2000 was responsible for the dispersal of copepods via the underground pipe system to 29 of 35 manholes over an area of 1.33 km2. Significant reductions in Aedes and Ochlerotatus larvae ensued. In these habitats, Mesocyclops and Metacyclops were able to survive dry periods, when substrate moisture content ranged from 13.8 to 79.9%. At the semiarid inland towns of Hughenden and Richmond, cracking clay soil prevents drainage of water from shallow service pits where Oc. tremulus immatures numbered from 292-18,460 per pit. Introduction of Mesocyclops copepods into these sites during May 1999 resulted in 100% control of Oc. tremulus for 18 mo. One uninoculated pit subsequently became positive for Mesocyclops with resultant control of mosquito larvae.     

Seasonal distribution, biology, and human attraction patterns of culicine mosquitoes (Diptera: Culicidae) in a forest near Puerto Almendras, Iquitos, Peru

This study was conducted as part of a field ecology study of arboviral activity in the Amazon Basin, Peru, to determine the taxonomy, frequency, seasonal, and vertical distributions of potential mosquito vectors. In addition, the relative efficiency of human-landing collections and dry ice-baited Centers for Disease Control (CDC)-type light traps was determined for collecting mosquitoes. A total of 70 species of mosquitoes from 14 genera were collected from June 1996 through December 1997 at a forested site near Puerto Almendras, approximately 20 km west-southwest of Iquitos, Peru. Three species [Psorophora (Janthinosoma) albigenu (Peryassu), Ochlerotatus (Ochlerotatus) fulvus (Wiedemann), and Ochlerotatus (Ochlerotatus) serratus (Theobald)] accounted for 70% of all mosquitoes captured in human-landing collections. Overall, biting activity occurred throughout the 24-h cycle but was higher during the daytime, primarily because of large populations of two day-biting species, Ps. albigenu and Oc. serratus. Oc. fulvus was active throughout the 24-h cycle but was more frequently collected during the evening. Oc. fulvus, Ps. albigenu, Culex (Melanoconion) pedroi Sirivanakarn & Belkin, and a mixture of Culex (Melaonoconion) vomerifer Komp, and Culex (Melanoconion) gnomatos Sallum, Huchings & Ferreira, accounted for 73% of the mosquitoes captured during darkness) by human collectors. In general, Ochlerotatus spp. and Psorophora spp. were more commonly captured in human-landing collections, whereas most Culex spp. were more frequently collected in the dry ice-baited CDC-type light traps. In general, mosquito populations were lowest from June through August when river levels were at their lowest. Two large population peaks occurred in November-December and in February-March as a result of "flood water" mosquito populations (e.g., Ps. albigenu). These data provide a better understanding of the taxonomy, population density, and seasonal distribution of potential mosquito vectors within the Amazon Basin region and allow for the development of appropriate vector and disease prevention strategies.     

Infectivity and pathogenicity of a novel baculovirus, CuniNPV from Culex nigripalpus (Diptera: Culicidae) for thirteen species and four genera of mosquitoes

The infectivity and pathogenicity of newly discovered baculovirus, CuniNPV (family Baculoviridae, genus Nucleopolyhedrovirus) originally isolated from the mosquito Culex nigripalpus Theobald, was evaluated in laboratory bioassys against thirteen species and four genera of mosquitoes native to the northeastern U.S. Purified virus at a dosage rate of 1.6 x 10(7) occlusion bodies/ml with 10 mM Mg2+ added was used in exposures with second through fourth instars at temperatures ranging from 17 to 27 degrees C. High infection rates and accompanying mortality were achieved in Cx. pipiens L. (83.0-14.4%), Cx. pipiens f. molestus (80.4% infection), and Cx. salinarius Coquillett (48.0-43.1%). Cx. restuans Theobald was also susceptible but infection rates were lower (21.3-12.5%). The gross pathology associated with infection was identical to that reported in Cx. nigripalpus. Infected larvae were lethargic and were often suspended at the water surface. Development of CuniNPV was observed in the nuclei of the midgut epitheial cells in the gastric caeca and posterior region of the stomach of host larvae. One hundred percent mortality was observed in all larvae that exhibited gross symptoms of infection within 4-d p.i. Cx. territans Walker (subgenus Neoculex Dyar) was the only Culex mosquito that was not susceptible. No infections were obtained with any species of Aedes [Ae. vexans (Meigen)], Culiseta [Culiseta morsitans (Theobald)] or Ochlerotatus [Ochlerotatus canadensis (Theobald), Oc. cantator (Coquillett), Oc. communis (De Geer), Oc. excrucians (Walker), Oc. japonicus (Theobald), Ochlerotatus stimulans (Walker), and Ochlerotatus triseriatus (Coquillett)]. The host range of CuniNPV appears to be restricted to Culex mosquitoes within the subgenus Culex. An inhibitory effect on transmission of CuniNPV was observed when a liver powder/Brewer's yeast mixture was used as a source of food reinforcing the critical role of Mg2+ and sensitivity of the infection process to the presence other divalent cations (Cu2+, Fe2+, and Zn2+) in the larval medium that interfered with the infection process. The high infectivity and pathogenicity of CuniNPV for the principal vectors of West Nile virus in North America make CuniNPV an attractive candidate for future development as a biopesticide.     

Ochlerotatus taeniorhynchus: a probable vector of Dirofilaria immitis in coastal areas of Yucatan, Mexico

Mosquito collections were done on microfilaremic dogs, positive for Dirofilaria immitis (Leidy), for 15 consecutive nights in Celestun, Yucatan, southeastern Mexico, during January 2007. In total, 275 mosquitoes (3 male and 272 female) of five species were collected: Ochlerotatus taeniorhynchus (Wiedemann), Ochlerotatus sollicitans (Walker), Culex quinquefasciatus Say, Culex interrogator (Dyar & Knab), and Aedes aegypti (L.). Oc. taeniorhynchus was the species most frequently collected, and it had the highest rate of feeding success and the highest attack rates. First (L1) and third (L3) instars of Dirofilaria were observed in all mosquito species collected except for Ae. aegypti: 23 of 223 Oc. taeniorhynchus, three of 40 Cx. quinquefasciatus, and one of six Oc. sollicitans and one of one Cx. interrogator were infected with Dirofilaria (10.3% of total examined mosquitoes). This is the first report of Dirofilaria (presumably D. immitis) isolation from wild-caught mosquitoes in Mexico. Results imply that D. immitis can develop from microfilaria to infective L3s in Oc. taeniorhynchus, Cx. quinquefasciatus, and Oc. sollicitans, but L3s (infectious) were only recovered in heads of Oc. taeniorhynchus. Thus, Oc. taeniorhynchus can be considered a potential vector of dirofilariasis in Celestun, because infective L3 Dirofilaria were found in the vicinity of the mouthparts, and this mosquito has a strong association with microfilaremic dogs in this study area.     

Vector competence of Australian mosquitoes (Diptera: Culicidae) for Japanese encephalitis virus

Australian mosquitoes were evaluated for their ability to become infected with and transmit a Torres Strait strain of Japanese encephalitis virus. Mosquitoes, which were obtained from either laboratory colonies and collected using Centers for Disease Control and Prevention light traps baited with CO2 and octenol or reared from larvae, were infected by feeding on a blood/sucrose solution containing 10(4.5 +/- 0.1) porcine stable-equine kidney (PS-EK) tissue culture infectious dose50/mosquito of the TS3306 virus strain. After 14 d, infection and transmission rates of 100% and 81%, respectively, were obtained for a southeast Queensland strain of Culex annulirostris Skuse, and 93% and 61%, respectively, for a far north Queensland strain. After 13 or more days, infection and transmission rates of > 90% and > or = 50%, respectively, were obtained for southeast Queensland strains of Culex sitiens Wiedemann and Culex quinquefasciatus Say, and a far north Queensland strain of Culex gelidus Theobald. Although infection rates were > 55%, only 17% of Ochlerotatus vigilax (Skuse) and no Cx. quinquefasciatus, collected from far north Queensland, transmitted virus. North Queensland strains of Aedes aegypti L., Ochlerotatus kochi (D?nitz), and Verrallina funerea (Theobald) were relatively refractory to infection. Vertical transmission was not detected among 673 F1 progeny of Oc. vigilax. Results of the current vector competence study, coupled with high field isolation rates, host feeding patterns and widespread distribution, confirm the status of Cx. annulirostris as the major vector of Japanese encephalitis virus in northern Australia. The relative roles of other species in potential Japanese encephalitis virus transmission cycles in northern Australia are discussed.     

Habitat preferences and phenology of Ochlerotatus triseriatus and Aedes albopictus (Diptera: Culicidae) in southwestern Virginia

Recently, the number of reported human cases of La Crosse encephalitis, an illness caused by mosquito-borne La Crosse virus (LAC), has increased in southwestern Virginia, resulting in a need for better understanding of the virus cycle and the biology of its vectors in the region. This study examined the spatial and temporal distributions of the primary vector of LAC, Ochlerotatus triseriatus (Say), and a potential secondary vector, Aedes albopictus (Skuse). Ovitrapping surveys were conducted in 1998 and 1999 to determine distributions and oviposition habitat preferences of the two species in southwestern Virginia. Mosquitoes also were collected for virus assay from a tire dump and a human La Crosse encephalitis case site between 1998 and 2000. Oc. triseriatus and Ae. albopictus were collected from all ovitrap sites surveyed, and numbers of Oc. triseriatus eggs generally were higher than those of Ae. albopictus. Numbers of Oc. triseriatus remained high during most of the summer, while Ae. albopictus numbers increased gradually, reaching a peak in late August and declining thereafter. In Wise County, relative Ae. albopictus abundance was highest in sites with traps placed in open residential areas. Lowest numbers of both species were found in densely forested areas. Ovitrapping during consecutive years revealed that Ae. albopictus was well established and overwintering in the area. An oviposition comparison between the yard and adjacent forest at a human La Crosse encephalitis case site in 1999 showed that Ae. albopictus preferentially oviposited in the yard surrounding the home, but Oc. triseriatus showed no preference. LAC isolations from larval and adult collections of Oc. triseriatus females from the same case site indicated the occurrence of transovarial transmission.     

Enhancement or modulation of the vector competence of Ochlerotatus vigilax (Diptera: Culicidae) for ross river virus by temperature

Two different doses of Ross River virus (RR) were fed to Ochlerotatus vigilax (Skuse), the primary coastal vector in Australia; and blood engorged females were held at different temperatures up to 35 d. After ingesting 10(4.3) CCID50/mosquito, mosquitoes reared at 18 and 25 degrees C (and held at the same temperature) had higher body remnant and head and salivary gland titers than those held at 32 degrees C. although infection rates were comparable. At 18, 25, and 32 degrees C, respectively, virus was first detected in the salivary glands on days 3, 2, and 3. Based on a previously demonstrated 98.7% concordance between salivary gland infection and transmission, the extrinsic incubation periods were estimated as 5, 4, and 3 d, respectively, for these three temperatures. When Oc. vigilax reared at 18, 25, or 32 degrees C were fed a lower dosage of 10(3.3) CCID50 RR/mosquito, and assayed after 7 d extrinsic incubation at these (or combinations of these) temperatures, infection rates and titers were similar. However, by 14 d, infection rates and titers of those reared and held at 18 and 32 degrees C were significantly higher and lower, respectively. However, this process was reversible when the moderate 25 degrees C was involved, and intermediate infection rates and titers resulted. These data indicate that for the strains of RR and Oc. vigilax used, rearing temperature is unimportant to vector competence in the field, and that ambient temperature variations will modulate or enhance detectable infection rates only after 7 d extrinsic incubation. Because of the short duration of extrinsic incubation, however, this will do little to influence RR epidemiology, because by this time some Oc. vigilax could be seeking their third blood meal, the latter two being infectious.     

Susceptibility of Ochlerotatus taeniorhynchus (Diptera: Culicidae) to infection with epizootic (subtype IC) and enzootic (subtype ID) Venezuelan equine encephalitis viruses: evidence for epizootic strain adaptation

To test the hypothesis that adaptation to epizootic mosquito vectors mediates emergence of Venezuelan equine encephalitis virus (VEEV) from enzootic progenitors, experimental infection studies were conducted to determine the susceptibility of Ochlerotatus taeniorhynchus (Wiedemann) to epizootic and enzootic strains. Artificial blood meals containing epizootic subtype IC strains isolated during the 1962-1964, 1992-1993, and 1995 Venezuelan/Colombian epizootics and closely related Venezuelan enzootic subtype ID strains were used to compare infectivity and transmission potential. Their greater infectivity and replication suggested that adaptation of epizootic strains to Oc. taeniorhynchus may have enhanced epizootic transmission during the 1962-1964 and 1995 IC coastal epizootics. However, strains from the small 1992-1993 Venezuelan outbreak that did not extend to coastal regions do not seem to infect this species better than closely related subtype ID strains. Adaptation of VEEV to epizootic vectors such as Oc. taeniorhynchus mosquitoes may be a determinant of some but not all VEE emergence events and may influence spread into coastal regions.     

Mosquitoes infected with West Nile virus in the Florida Keys,Monroe County,Florida, USA

More than 30,000 mosquitoes in 22 species or species groups were collected from the Florida Keys, Monroe County, FL, USA, in dry ice-baited light and gravid traps. Dry ice-baited traps collected more mosquitoes than did gravid traps. West Nile virus was detected in pools of Anopheles atropos Dyar & Knab, Deinocerites cancer Theobald, and Ochlerotatus taeniorhynchus (Wiedemann).     

Constitutive differences between natural and artificial container mosquito habitats: vector communities, resources, microorganisms, and habitat parameters

Aquatic containers, including tree holes and vehicle tires, harbor a diverse assemblage of mosquitoes capable of vectoring important diseases. Many studies have examined containers as a mosquito breeding site, although no data exist that have simultaneously compared mosquito communities between tree holes and tires, and few have quantified differences in environmental factors or food resources that may be important for explaining population or community differences. At two times (early and late summer 2009) we sampled two tire and two tree hole sites in south-central Mississippi, and for each container we enumerated mosquito larvae and measured several environmental parameters (canopy cover, water volume, and detritus), and biomass and productivity of fungi and bacteria, and species richness and abundance of protozoans. Tree holes held less water but were more shaded compared with tires; however, after correcting for volume differences, tree holes contained more detritus and were higher in some microorganism measures (protozoan richness, bacterial productivity in the water column). Based on community dissimilarity analysis of mosquitoes, strong differences existed between container types and sampling period; Aedes albopictus (Skuse) and Culex quinquefasciatus (Say) were dominant in tires, whereas Ae. triseriatus (Say) and Orthopodomyia signifera (Coquillett) were dominant in tree holes. This study also reports the use of tires by the invasive mosquito Cx. coronator (Dyar and Knab). Tree holes supported a higher density of larvae but fewer species than tires, though there was variation across time. Our work illustrates that detrital inputs and some microorganisms differ in fundamental ways between tires and tree holes, and because of compositional differences in mosquito communities, these small aquatic habitats cannot be considered to be homogeneous mosquito habitats.     

Autogeny in Ochlerotatus vigilax (Diptera: Culicidae) from southeast Queensland, Australia

Field and laboratory investigations were undertaken to determine the level of expression of autogeny in the mosquito Ochlerotatus vigilax (Skuse) from southeast Queensland, Australia, and whether there was evidence of seasonal variation. At two field sites in southeast Queensland, Wellington Point and Donnybrook, autogeny rates were determined on six occasions between January 2001 and January 2002. The autogeny rate varied between 71 and 100% at Wellington Point and between 63 and 100% at Donnybrook. Autogenous fecundity ranged from 17 to 63 eggs per female at Wellington Point and from 13 to 88 eggs per female at Donnybrook. Positive relationships were found between adult body size (indicated by wing length), autogeny rate, and fecundity. A laboratory study was conducted to investigate the influence of larval nutrition and adult diet (water versus sucrose) on the expression of autogeny. The autogeny rate at a low-diet treatment was between 73 and 90% when sucrose was withheld from females and 100% when sucrose was provided. All high-diet females were autogenous. Autogenous egg development required 80 +/- 6 h from emergence at 27 degrees C. We conclude that autogeny rates are consistently high in Oc. vigilax from the southeast Queensland region.