Calanoid copepods: an overlooked tool in the control of disease vector mosquitoes

Biological control can assist in the management of disease vector mosquitoes. However, we urgently require the identification of novel and effective agents to aid population management strategies. Quantifying interactions strengths between consumers and resources is central to our understanding of trophic stability, and is relevant within the biological control context. Previously, pPredatory biocontrol of disease vector mosquito species has previously focused extensively on cyclopoid copepods, but prey size refuge effects have been identified as a hindrance to their predatory efficacy. Calanoid copepods have yet to be comprehensively examined in the context of mosquito control, despite their high prevalence, diversity and distribution. Here, we apply functional responses (FRs; resource use as a function of resource density) to examine interaction strengthspredation efficiencies of a recently described ephemeral pond specialist species, the freshwater calanoid copepod Lovenula raynerae Suárez-Morales, Wasserman & Dalu 2015, using different size classes of larvae of the disease vector complex Culex pipiens as prey. Lovenula raynerae effectively consumed C. pipiens larvae across their ontogeny. A potentially population destabilising Type II FR was exhibited towards both early and late instar mosquitoes, indicative of a lack of prey refuge across ontogenetic stages. Attack rates were greatest and handling times lowest for early instar larvae compared to late instar larvae. These traits contrast to other copepods, commonly applied in biocontrol, which are only able to handle early instars, and in much smaller numbers. We thus advocate that calanoid copepods can exert particularly marked predatory impact on lower trophic groups, and that their use in disease vector mosquito control strategies should be further explored

Muddy waters: efficacious predation of container-breeding mosquitoes by a newly-described calanoid copepod across differential water clarities

Mosquito-borne diseases induce unrivalled morbidity and mortality in human populations. In recent times, greater urbanisation has facilitated vector population expansion, particularly of those which proliferate in container-style habitats. To combat increased associated disease risk, we urgently require innovative and efficacious control mechanisms to be identified and implemented. Predatory biological control of vectorially-important mosquitoes can be effective. Despite their high prevalence in freshwater ecosystems, predatory calanoid copepods have yet to be examined comprehensively for mosquito control. Moreover, environmental context-dependencies can cause substantial variations in natural enemy impacts on target populations. Accordingly, improved understanding of the effects of context-dependencies upon predatory biocontrol is needed. Here, we use functional responses (FRs) to examine the predatory impact of a newly-described species of calanoid copepod, Lovenula raynerae, upon larval Culex pipiens prey across variations in prey supply and water clarity. Using outdoor field trials, we assess the viability of L. raynerae in reducing mosquito survival in container-style habitats. Lovenula raynerae displayed destabilising Type II FRs towards larval mosquito prey across all water clarities tested, with overall predation rates remaining largely unaffected across all clarity treatments. In the outdoor experiment, L. raynerae applications resulted in substantial reductions in larval C. pipiens populations, with close to total eradication achieved following the experimental period under higher predator densities. These results demonstrate that environmental context such as water clarity may have little effect on vector control by calanoid copepods, which suggests a predatory reliance on hydromechanical signalling. Further, for the first time, we demonstrate the applicability of calanoid copepods to artificial container-style habitats where mosquitoes proliferate. Therefore, our results indicate that further examination into the applicability of this species group to aid vector biocontrol strategies is warranted

Quantifying reproductive state and predator effects on copepod motility in ephemeral ecosystems

Ephemeral wetlands in arid environments are unique ecosystems with atypical trophic structuring, often dominated by invertebrate predation. Copepod behavioural traits and vulnerabilities to predation can vary substantially according to reproductive status. Gravid female copepods may be more vulnerable to predation due to reduced escape speeds or higher visibility to predators. Here, we quantify how reproductive status modulates horizontal motility rates of the predatory ephemeral pond specialist copepod Lovenula raynerae, and the responsiveness of the copepod to predator cues of the notonectid Anisops debilis. Males exhibited significantly higher motility rates than gravid female copepods, however chemical predator cues did not significantly influence activity rates in either sex. The lack of responsiveness to predator cues by specialist copepods in ephemeral wetlands may result from a lack of predation pressure in these systems, or due to time stress to reproduce during short hydroperiods. In turn, this could increase predation risk to copepods from externallyrecruited top predators in ephemeral wetlands, and potentially contribute to the development of skewed sex ratios in favour of females

Assessing multiple predator, diurnal and search area effects on predatory impacts by ephemeral wetland specialist copepods

Predator-prey interaction strengths can be highly context-dependent. In particular, multiple predator effects (MPEs), variations in predator sex and physical habitat characteristics may affect prey consumption rates and thus the persistence of lower trophic groups. Ephemeral wetlands are transient ecosystems and predatory copepods are often numerically dominant. We examine the interaction strengths of a specialist copepod Paradiaptomus lamellatus towards mosquito prey in the presence of conspecifics using a functional response (FR) approach. Further, we examine sex variability in predation rates of P. lamellatus under circadian and surface area variations. Then, we assess the influence of a co-occurring heterospecific predatory copepod, Lovernula raynerae, on total predation rates. We demonstrate MPEs affecting consumption, with negative non-trophic interaction strength and thus antagonism displayed between conspecific predatory units of P. lamellatus. This antagonism was present irrespective of prey density. Furthermore, we show differences between sexes in interaction strengths, with female P. lamellatus significantly more voracious than males, irrespective of time of day and experimental arena surface area. Predation rates by P. lamellatus were significantly lower than the heterospecific calanoid copepod L. raynerae, whilst heterospecific copepod groups exhibited the greatest predatory impact. Our results provide insights into the predation dynamics by specialist copepods, wherein species diversity and sex affect interaction strengths. In turn, this may influence population-level persistence of lower trophic groups under shifting copepod predator composition

Sex demographics alter the effect of habitat structure on predation by a temporary pond specialist

Habitat structure can profoundly influence the strength of interactions between predators and prey. Spatio-temporal habitat structure in temporary wetland ecosystems is particularly variable because of fluctuations in water levels and vegetation colonisation dynamics. Demographic characteristics within animal populations may also alter the influence of habitat structure on biotic interactions, but have remained untested. Here, we investigate the influence of vegetation habitat structure on the consumption of larval mosquito prey by the calanoid copepod Lovenula raynerae, a temporary pond specialist. Increased habitat complexity reduced predation, and gravid female copepods were generally more voracious than male copepods in simplified habitats. However, sexes were more similar as habitat complexity increased. Type II functional responses were exhibited by the copepods irrespective of habitat complexity and sex, owing to consistently high prey acquisition at low prey densities. Attack rates by copepods were relatively unaffected by the complexity gradient, whilst handling times lengthened under more complex environments in gravid female copepods. We demonstrate emergent effects of habitat complexity across species demographics, with predation by males more robust to differences in habitat complexity than females. For ecosystems such as temporary ponds where sex-skewed predator ratios develop, our laboratory findings suggest habitat complexity and sex demographics mediate prey risk

Optical Detection of a Single Nuclear Spin

We propose a method to optically detect the spin state of a 31-P nucleus embedded in a 28-Si matrix. The nuclear-electron hyperfine splitting of the 31-P neutral-donor ground state can be resolved via a direct frequency discrimination measurement of the 31-P bound exciton photoluminescence using single photon detectors. The measurement time is expected to be shorter than the lifetime of the nuclear spin at 4 K and 10 T.Comment: 4 pages, 3 figure