RATIONALE The study of insect migration is problematic due to the small size of insects. Stable isotope analysis can be used to elucidate movement, either by geographic assignment of location of a species, or by simply distinguishing migrant from resident populations. There are few isoscapes of any kind in the UK/Ireland available for interrogation. Thus, I have measured stable isotope ratios (of H, C, N and S) of 299 individuals of the non-migratory Brimstone moth (Opisthograptis luteolata) collected from 93 locations around the UK and Ireland by citizen scientists. METHODS After removing lipids, stable isotope ratios were measured by continuous flow isotope ratio mass spectrometry, using either a conventional elemental analyser (C, N and S) or a high temperature, thermal conversion elemental analyser in reductive mode. RESULTS Maps (isoscapes) were constructed that illustrate the stable isotope spatial distribution of this insect. These are the first isoscapes of H, C, N and S of ...

Oecologia, 1996

Quantifying pathways of energy transfer between plants, pests, and beneficial insects is a necessary step toward maintaining pest stable agroecosystems in the absence of chemical subsidies. A diet switching experiment utilizing a predatory ladybird beetle, Hippodamia variegata (Goeze), evaluated the use of naturally occurring stable C and N isotopes as an economically feasible and safe method for quantifying pathways of energy flow within agroecosystems. Stable isotope values of the ladybird beetle Coleomegilla maculata lengi (Timberlake) collected from an agroecosystem were used to estimate the relative amount of C and N derived from agricultural plants and incorporated by ladybird beetles based on mass balance equations. At the beginning of the diet-switching experiment δ13C and δ15N values of H. variegata (–12.0‰ and 6.3‰, respectively) differed by –0.2‰ and 2.9‰ from the aphids that were provided exclusively as their diet. These data are consistent with previous estimates of trophic level isotope effects. After switching the diet of H. variegata to an alternative food, isotope values of H. variegata gradually shifted toward expected values for individuals fed this diet (–22.9‰ and 8.8‰ for δ13C and δ15N values, respectively). Isotope values of another ladybird beetle, C. maculata, collected from the field indicated that in May, alfalfa and maize (pollen) obtained in the previous year contributed 32% and 68% of the C or N to the diets of these individuals and in August, 52%, 6%, and 42% of the C or N assimilated by these insects was derived from alfalfa, wheat, and maize, respectively. These data are consistent with expectations based on the relative abundance of C. maculata in various crops during the season. The field and laboratory data are a clear indication that isotope values are sensitive to dietary changes on a relatively short time scale (days) and provide a strong basis for the use stable C and N isotope to trace energy flow patterns of these beneficial organisms within agroecosystems.

Entomologia Experimentalis et Applicata, 2007

This is an eclectic review and analysis of contemporary and promising stable isotope methodologies to study the biology and ecology of arthropods. It is augmented with literature from other disciplines, indicative of the potential for knowledge transfer. It is demonstrated that stable isotopes can be used to understand fundamental processes in the biology and ecology of arthropods, which range from nutrition and resource allocation to dispersal, food-web structure, predation, etc. It is concluded that falling costs and reduced complexity of isotope analysis, besides the emergence of new analytical methods, are likely to improve access to isotope technology for arthropod studies still further. Stable isotopes pose no environmental threat and do not change the chemistry or biology of the target organism or system. These therefore represent ideal tracers for field and ecophysiological studies, thereby avoiding reductionist experimentation and encouraging more holistic approaches. Considering (i) the ease with which insects and other arthropods can be marked, (ii) minimal impact of the label on their behaviour, physiology, and ecology, and (iii) environmental safety, we advocate more widespread application of stable isotope technology in arthropod studies and present a variety of potential uses.

Trends in Ecology & Evolution, 2004

Establishing patterns of movement of wild animals is crucial for our understanding of their ecology, life history and behavior, and is a prerequisite for their effective conservation. Advances in the use of stable isotope markers make it possible to track a diversity of animal species in a variety of habitats. This approach is revolutionizing the way in which we make connections between phases of the annual cycle of migratory animals. However, researchers must exercise care in their application of isotopic methods. Here, we review stable isotope patterns in nature and discuss recent tracking applications in a range of taxa. To aid in the interpretation and design of effective and insightful isotope movement studies, we discuss a series of key issues and assumptions. This exciting field will advance rapidly if researchers consider these aspects of study design and interpretation carefully.

Frontiers in Ecology and Evolution

Hydrogen isotope geolocation of insects is based on the assumption that the chitin in the wings of adult migratory insects preserves the hydrogen isotope composition (δ2H) of the larval stages without influence of adult diet. Here, we test this assumption by conducting laboratory feeding experiments for monarch butterflies (Danaus plexippus) including: (1) a starvation treatment where adults were not fed and (2) an enriched treatment where adults were fed a diet isotopically enriched in deuterium (~ +78‰) compared to the larval diet. The δ2H values of adult wings were measured at different time steps along the 24-day experiment. We also investigated intra-wing differences in δ2H values caused by wing pigmentation, absence of wing scales, and presence of major wing veins. We conclude that, although the magnitude of the changes in δ2H values are small (~6‰), wing δ2H values vary based on adult diet and insect age, particularly early after eclosion (i.e., 1–4 days). We found that wing ...

Oecologia, 1999

Wing membranes of laboratory and ®eldreared monarch butter¯ies (Danaus plexippus) were analyzed for their stable-hydrogen (dD) and carbon (d 13 C) isotope ratios to determine whether this technique could be used to identify their natal origins. We hypothesized that the hydrogen isotopic composition of monarch butter¯y wing keratin would re¯ect the hydrogen isotope patterns of rainfall in areas of natal origin where wings were formed. Monarchs were reared in the laboratory on milkweed plants (Asclepias sp.) grown with water of known deuterium content, and, with the assistance of volunteers, on native milkweeds throughout eastern North America. The results show that the stable hydrogen isotopic composition of monarch butter¯ies is highly correlated with the isotopic composition of the milkweed host plants, which in turn corresponds closely with the long-term geographic patterns of deuterium in rainfall. Stable-carbon isotope values in milkweed host plants were similarly correlated with those values in monarch butter¯ies and showed a general pattern of enrichment along a southwest to northeast gradient bisecting the Great Lakes. These ®ndings indicate that natal origins of migratory and wintering monarchs in Mexico can be inferred from the combined dD and d 13 C isotopic signatures in their wings. This relationship es-tablishes that analysis of hydrogen and carbon isotopes can be used to answer questions concerning the biology of migratory monarch butter¯ies and provides a new approach to tracking similar migratory movements of other organisms.

Journal of Insect Physiology, 2010

Determining diet and trophic position of species with stable isotopes requires appropriate trophic enrichment estimates between an animal and its potential foods. These estimates are particularly important for cryptic foragers where there is little comparative dietary information. Nonetheless, many trophic enrichment estimates are based on related taxa, without confirmation of accuracy using laboratory trials. We used stable isotope analysis to investigate diet and to resolve trophic relationships in a large endemic insect, the Auckland tree weta (Hemideina thoracica White). Comparisons of isotopes in plant foods fed to captive wetas with isotope ratios in their frass provided variable results, so frass isotope values had limited usefulness as a proxy indicator of trophic level. Isotopic values varied between different tissues, with trophic depletion of 15N highest in body fat and testes. Tissue fractionation was consistent in captive and wild caught wetas, and isotopic values were not significantly different between the two groups, suggesting that this weta species is primarily herbivorous. Whole-body values in captive wetas demonstrated trophic depletion (Dd) for d15N of about 0.77%and trophic enrichment of 4.28% for d13C. These values differ from commonly estimated trophic enrichments for both insects and herbivores and indicate the importance of laboratory trials to determine trophic enrichment. Isotopic values for femur muscles from a number of local wild weta populations did not vary consistently with body weight or size, suggesting that juveniles eat the same foods as adults. Considerable variation among individuals within and between populations suggests that isotopic values are strongly influenced by food availability and individual foraging traits.

2019

Using stable isotope measurements of inert tissues to determine origins and migratory patterns is well established. However, isotopically determining nutritional origins of lipids, the primary fuel of migration, has not been attempted. I explored isotopic links between diet and stored lipids in captive White-throated Sparrows (Zonotrichia albicollis) and true armyworm moths (Mythimna unipuncta) using δ 13 C and δ 2 H measurements. Isotopic discrimination between body lipids and diet was established as linear calibration functions. Isotopic uptake following a diet switch in moths was used to trace lipid accumulation over time. Isotopic correlations between breath metabolic by-products of fed and fasted sparrows were explored as indicators of lipid use. This study established isotopic (δ 13 C, δ 2 H) linkages between diet and stored lipids for migratory insects and passerines and advocates the use stable isotopes in lipids as a tool to evaluate nutrient origins and allocation strategies in a variety of migratory species.

Animal Migration, 2017

Increasingly, stable isotope measurements are being used to assign individuals to broad geographic origins based on established relationships between animal tissues and tissue-specific isoscapes. In particular, the eastern North American population of the monarch butterfly (Danaus plexippus) has been the subject of several studies using established δ

Animal Migration, 2018

Alternative life history strategies are mechanisms by which organisms are able to maximize fitness across a range of environmental conditions. Fitness is maximized by different strategies depending on context, resulting in trade-offs between life history strategies. Monarch butterflies (Danaus plexippus) employ both migratory and resident life history strategies. Since residents breed throughout the year, but migrants overwinter in reproductive diapause, there are fitness trade-offs between the two strategies. We used stable isotope analysis to evaluate the geographic origins of monarchs in a yearround population in south Florida. Based on stable isotope profiles of hydrogen and carbon (δ2H and δ13C values), we found that 48% (16/33) of monarchs collected in south Florida are migrants that originated from outside the sampling region. Migrants had a larger wing length than residents; thus, switching to a resident strategy could alter their probability of reproductive success. Further...

2008

Stable isotope analyses of tissues have been used to help delineate natal regions and routes of migratory animals. The foundations of such studies are isotopic gradients or differences representing geographic regions and habitat used by the organism that are retained in selected tissues. We sampled peacock butterflies Inachis io on a regional level in southern Sweden to study natural variation and the resolving power of the stable isotope method to delineate individuals from known areas on a smaller scale than has typically been used in previous studies. Hydrogen (dD), carbon (d 13 C) and nitrogen (d 15 N) isotopes were obtained from butterflies at seven different locations in an area of 250 )250 km over three years (2002Á2004). We found sufficient isotopic differences on this regional scale to delineate approximate origins. Of the three isotopes, deuterium showed good discrimination between sites, carbon isotopes showed weaker differentiation, whereas nitrogen isotopes proved unsuitable for small scale studies in this region due to high and unpredictable variation. We found there was enough variation in dD between years to prevent a general application of the technique to resolve sub-regional variation. Substantial part of this variation was probably caused by seasonal changes in dD of precipitation. These differences produce significant variation in dD between years in animals having short and variable tissue development times, and are difficult to estimate in natural situations. We conclude that stable isotopes are potentially powerful predictors for studies of migratory butterflies in Europe. However, without good knowledge about the sampled individuals' previous life-history, a lot of the natural environmental variation in tissue dD cannot be controlled for. In the case of migratory species, this information is difficult to obtain, making the confidence intervals for prediction of natal areas fairly wide and probably only suitable for longer distance migration.

Frontiers in Zoology, 2016

Background: Information on larval diet of many holometabolous insects remains incomplete. Carbon (C) and nitrogen (N) stable isotope analysis in adult wing tissue can provide an efficient tool to infer such trophic relationships. The present study examines whether moth feeding guild affiliations taken from literature are reflected in isotopic signatures. Results: Non-metric multidimensional scaling and permutational analysis of variance indicate that centroids of dietary groups differ significantly. In particular, species whose larvae feed on mosses or aquatic plants deviated from those that consumed vascular land plants. Moth δ 15 N signatures spanned a broader range, and were less dependent on species identity than δ 13 C values. Comparison between moth samples and ostensible food sources revealed heterogeneity in the lichenivorous guild, indicating only Lithosia quadra as an obligate lichen feeder. Among root-feeding Agrotis segetum, some specimens appear to have developed on crop plants in forest-adjacent farm land. Reed-feeding stem-borers may partially rely on intermediary trophic levels such as fungal or bacterial growth. Conclusion: Diagnostic partitioning of moth dietary guilds based on isotopic signatures alone could not be achieved, but hypotheses on trophic relationships based on often vague literature records could be assessed with high resolution. Hence, the approach is well suited for basic categorization of moths where diet is unknown or notoriously difficult to observe (i.e. Microlepidoptera, lichen-feeders).

Measurement of natural stable isotope signatures of insects has proven to be a useful method in determining their natal origin, feeding strategies and mating behavior, and has spawned a new discipline of insect isotope forensics. However the complexity of measuring isotope signatures using elemental analysis-isotope ratio mass-spectrometry (EA-IRMS) has encountered significant technical barriers, which together with costs, have hindered the advancement and wide spread adoption of this potentially useful approach. In the last decade simpler technologies to measure stable isotope ratios in samples based on laser spectroscopy have been developed. We foresee that these instruments will fuel the advancement and adoption of insect isotope forensics into pest management programs because they are easy to use, more affordable, and complement existing marking tools. However prior to the adoption of these new technologies it is necessary to test their fitness for purpose and explore their limitations. In this paper we set out to test the suitability of using combustion module-cavity ring-down spectroscopy (CM-CRDS) for measuring isotope ratios in an entomo-logical context, covering marking of insects, mating behavior and dietary assessment. We carried out a series of experiments to compare the isotope measurements made with the CM-CRDS with the traditional method of EA-IRMS. Initially we compared measurements of carbon isotope values of a variety of common diet components used in insect rearing, which covered a wide range of carbon isotope values. There was good agreement in the isotope values obtained, using the CM-CRDS and the EA-IRMS with similar accuracy and precision. Secondly we compared the 2 methods to measure carbon isotope values of the common cutworm, Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae) samples, and again there was good agreement in the values obtained. Thirdly we tested whether the CM-CRDS could be used to determine the paternity of spermataphores, based on the differential isotope signatures of the paternal and maternal moths, as a result of differences in their rearing diets. The CM-CRDS proved to be fit for the purpose of measuring isotope values in the spermataphores of common cutworm and suggested that the CM-CRDS technology would be suitable for these isotope based mating studies in moths if the spermatophores were of sufficient mass. Finally we explored a number of the issues surrounding CM-CRDS and isotope technologies in general, e.g., logistical considerations, economics of operation, sample size, etc. We conclude that CM-CRDS is a suitable instrument for measuring stable isotope carbon signatures in moths, and most probably other sufficiently large insects and that CM-CRDS could be easily used in both operational and research contexts. Resumen La medición de las firmas de isótopos estables naturales de los insectos ha demostrado ser un método útil para determinar su origen natal, estra-tegias de alimentación y su comportamiento de apareamiento y ha dado lugar a una nueva disciplina de la ciencia forense de isótopos de insectos. Sin embargo, la complejidad de la medición de las firmas isotópicas usando el análisis de espectrometría de masas de ratio elemental de isótopos (AEM-REI) se ha encontrado con obstáculos técnicos importantes, que junto con los costos, han obstaculizado el avance y la adopción generalizada de este enfoque potencialmente útil. En la última decada se han desarrollado tecnologías más simples para medir las proporciones de isótopos estables en muestras basadas en espectroscopia láser. Prevemos que estos instrumentos se alimentan el avance y la adopción del uso de forense de isótopos de insectos en los programas de manejo de plagas, ya que son fáciles de usar, menos costos, y complementan las herramientas para marcar ya existentes. Sin embargo, antes de la adopción de estas nuevas tecnologías es necesario probar su aptitud para el uso y explorar sus limitaciones. En este trabajo nos propusimos evaluar el apropiado del uso de la espectroscopia de combustión de cavidad de módulo de anillo-abajo (ECC-MAA) para medir el ratio de los isótopos en un contexto entomológica, incluyendo el marcar de los insectos, el comportamiento del apareamiento y la evaluación de la dieta. Realizamos una serie de experimentos para comparar las medidas de isótopos realizadas con la ECM-CAA con el método tradicional de AEM-REI. Inicialmente se compararon las medidas de los valores de los isótopos de carbono de una variedad de componentes comunes utilizados en la dieta en la cría de insectos, que cubría una amplia gama de valores de los isótopos de carbono. Hubo buen acuerdo en los valores de isótopos obtenido, utilizando el ECM-CAA y el AEM-REI con exactitud y precisión similar. En segundo lugar se compararon los 2 métodos para medir los valores

PLoS ONE, 2014

Protecting a nation's primary production sector and natural estate is heavily dependent on the ability to determine the risk presented by incursions of exotic insect species. Identifying the geographic origin of such biosecurity breaches can be crucial in determining this risk and directing the appropriate operational responses and eradication campaigns, as well as ascertaining incursion pathways. Reading natural abundance biogeochemical markers using mass spectrometry is a powerful tool for tracing ecological pathways as well as provenance determination of commercial products and items of forensic interest. However, application of these methods to trace insects has been underutilised to date and our understanding in this field is still in a phase of basic development. In addition, biogeochemical markers have never been considered in the atypical situation of a biosecurity incursion, where sample sizes are often small, and of unknown geographic origin and plant host. These constraints effectively confound the interpretation of the one or two isotope geolocation markers systems that are currently used, which are therefore unlikely to achieve the level of provenance resolution required in biosecurity interceptions. Here, a novel approach is taken to evaluate the potential for provenance resolution of insect samples through multiple biogeochemical markers. The international pest, Helicoverpa armigera, has been used as a model species to assess the validity of using naturally occurring d 2 H, 87 Sr/ 86 Sr, 207 Pb/ 206 Pb and 208 Pb/ 206 Pb isotope ratios and trace element concentration signatures from single moth specimens for regional assignment to natal origin. None of the biogeochemical markers selected were individually able to separate moths from the different experimental regions (150-3000 km apart). Conversely, using multivariate analysis, the region of origin was correctly identified for approximately 75% of individual H. armigera samples. The geographic resolution demonstrated with this approach has considerable potential for biosecurity as well as other disciplines including forensics, ecology and pest management.

Florida Entomologist, 2016

In this study we identified a number of moth (Lepidoptera) species that are potential targets for the sterile insect technique (SIT), and we assessed the feasibility of using stable isotope signatures as markers to distinguish mass-reared from wild moth species. Large natural differences in the isotopic signatures of commercially available sugars render them novel markers for mass-reared insects. Sugar beet (Beta vulgaris L.; Caryophyllales: Amaranthaceae), a C 3 plant, has a stable isotopic signature (a measure of the ratio of the stable isotopes 13 C: 12 C) of around −27‰ relative to Vienna Pee Dee Belemnite (VPDB; the international C isotope standard for the stable isotopes, 13 C and 12 C), and sugarcane (Saccharum spp.; Poales: Poaceae), a C 4 plant, has an isotopic signature of around −11‰. Thus by means of such a distinct isotope ratio in the sugar in the diet, mass-reared insects can be easily distinguished from wild insects with a high degree of certainty. It was shown that the method could be extended using a multiple isotope approach, with 15 N or a full suite of C, N, S and O isotopes. Intrinsic isotope marking of mass-reared moths proved to be an accurate means of distinguishing wild from mass-reared populations, based on isotopic differences between the wild host plant species and the diets used in mass-rearing, which where possible, had been manipulated to contain the isotopically divergent sugar type. This intrinsic labeling using stable isotopes could be useful in the assessment of the quality of mass-reared moths, because a stable isotope is a marker that does not affect the insect in any detrimental manner.

Entomologia Experimentalis et Applicata, 2013

Orius majusculus Reuter (Hemiptera: Anthocoridae) is an important component of the pest predatory complex in arable crops in Mediterranean areas. It moves between crops searching for prey, and improving knowledge on its dispersal abilities will help to develop conservation biological control strategies. Stable isotope ratios may be used as a tool for tracking insect movements, as the isotopic composition of insect tissues changes to reflect that of their diet when they undergo dietary shifts on moving between isotopically distinct crops. We carried out laboratory diet switch experiments with a stable isotope approach to infer information on dispersal of O. majusculus individuals among C3 and C4 crops to better understand isotopic field data collections. Switching the aphid food source caused a quick change in d 13 C signatures, regardless of the original and final food source. Changes in the d 13 C ratio of O. majusculus after diet switching fitted with an exponential model that showed similar turnover rates, and thus half-lives, between shifting diets up to 20 days. Subsequently, whereas individuals feeding on C4 aphids did not survive, turnover rate decreased in individuals that switched from C4 to C3 aphids. However, d 13 C traces from the original source remained in the predator until 25 days after switching, and this is enough time to help determine the movement of O. majusculus between crops in the field and to plan the timing of predator sampling and crop practices that may enhance predator ecological services. Orius majusculus that switched to a maize aphid diet showed different turnover rates between sexes, although this did not influence the pattern of switchover.

Die Bodenkultur: Journal of Land Management, Food and Environment

Summary At first glance, there appears little to link nitrogen cycling with entomology other that the use of isotope techniques. Soil management requires a contextual, adaptive, flexible approach that is based on understanding the factors that regulate the soil’s fundamental processes. Using stable isotope techniques for the analysis of the inherent biogeochemical processes can explain the complex soil–plant interactions, the determining factors of the nitrogen cycle, and the impacts of applying external inputs. Using the same stable isotope tools enables an interdisciplinary collaboration between soil science and entomology. Emerging plant pests, for instance, non-native insect species, are a threat to the agroecosystems. The knowledge of the origin and movement of invasive insects by using the intrinsic stable isotope signatures embedded in the environment allows identification of entry pathways and thus the establishment of effective management strategies to reduce the potential ...

Diversity and Distributions, 2005

BMC Ecology

Background The Schreiber’s bat, Miniopterus schreibersii, is adapted to long-distance flight, yet long distance movements have only been recorded sporadically using capture-mark-recapture. In this study, we used the hydrogen isotopic composition of 208 wing and 335 fur specimens from across the species' European range to test the hypothesis that the species migrates over long distances. Results After obtaining the hydrogen isotopic composition (δ2H) of each sample, we performed geographic assignment tests by comparing the δ2H of samples with the δ2H of sampling sites. We found that 95 bats out of 325 showed evidence of long-distance movement, based on the analysis of either fur or wing samples. The eastern European part of the species range (Greece, Bulgaria and Serbia) had the highest numbers of bats that had moved. The assignment tests also helped identify possible migratory routes, such as movement between the Alps and the Balkans. Conclusions This is the first continental-sc...

Movement Ecology

Background Migratory insects are important for the provision of ecosystem services both at the origin and destination sites but – apart from some iconic species – the migration routes of many insect species have not been assessed. Coastlines serve as a funnel where migrating animals including insects accumulate. Migratory behaviour and captures of dragonflies in bird traps suggest autumn migration of dragonflies along coastlines while the origin and regularity of this migration remain unclear. Methods Dragonfly species were caught at the bird observatory Kabli at the Baltic coast in Estonia in 2009, 2010 and 2015. For the 2015 data set, we used a stable hydrogen (H) approach to trace the potential natal origin of the migrant hawker (Aeshna mixta). Results 1079 (2009), 701 (2010) and 88 (2015) A. mixta individuals were caught during the study periods (35, 37 and 11 days in 2009, 2010 and 2015, respectively). The migration period lasted from end of August to end of September. Based on...

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