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It is believed that the high activity of sun with eruptions causing instability of the magnetic field of the Earth cause various health problems in sensitive people, including such events as heart attack. We analysed the daily frequency of patients admitted with heart attack to the Cardiocentre of the Institute of Clinical and Experimental Medicine in relation to daily mean value of the geomagnetic Ap index representing fluctuations in the magnetic field. There were slightly more days with both low Ap and low number of patients than expected by random, and slightly more days with both high Ap and number of patients. However, very high magnetic activity (Ap>30) did not cause high rate of heart attacks.

Hippodamia Chevrolat, 1836 currently comprises 19 species. Four species of Hippodamia are native to the Palearctic region: Hippodamia arctica (Schneider, 1792), H. septemmaculata (DeGeer, 1775), H....

Ladybirds (Coleoptera: Coccinellidae) provide services that are critical to food production, and they fulfill an ecological role as a food source for predators. The richness, abundance, and distribution of ladybirds, however, are compromised by many anthropogenic threats. Meanwhile, a lack of knowledge of the conservation status of most species and the factors driving their population dynamics hinders the development and implementation of conservation strategies for ladybirds. We conducted a review of the literature on the ecology, diversity, and conservation of ladybirds to identify their key ecological threats. Ladybird populations are most affected by climate factors, landscape composition, and biological invasions. We suggest mitigating actions for ladybird conservation and recovery. Short-term actions include citizen science programs and education, protective measures for habitat recovery and threatened species, prevention of the introduction of non-native species, and the maintenance and restoration of natural areas and landscape heterogeneity. Mid-term actions involve the analysis of data from monitoring programs and insect collections to disentangle the effect of different threats to ladybird populations, understand habitat use by taxa on which there is limited knowledge, and quantify temporal trends of abundance, diversity, and biomass along a management-intensity gradient. Long-term actions include the development of a worldwide monitoring program based on standardized sampling to fill data gaps, increase explanatory power, streamline analyses, and facilitate global collaborations.Las catarinas (Coleoptera: Coccinellidae) proporcionan servicios que son críticos para la producción de alimento, y juegan un papel ecológico como fuente de alimento para depredadores. Sin embargo, la riqueza, abundancia y distribución de catarinas están en peligro debido a muchas amenazas antropogénicas. La carencia de conocimiento sobre el estatus de conservación de la mayoría de las especies y los factores que inciden en su dinámica poblacional dificulta el desarrollo e implementación de estrategias de conservación para las catarinas. Realizamos una revisión de la literatura sobre la ecología, diversidad y conservación de catarinas para identificar sus amenazas ecológicas clave. Las poblaciones de catarinas fueron afectadas mayormente por factores climáticos, composición del paisaje e invasiones biológicas. Proponemos acciones de mitigación para la conservación y recuperación de catarinas. Acciones a corto plazo incluyen programas de ciencia y educación ciudadana, medidas de protección para la recuperación de hábitat y de especies amenazadas, prevención de la introducción de especies no nativas y el mantenimiento y restauración de áreas naturales y la heterogeneidad del paisaje. Acciones a mediano plazo implican el análisis de datos obtenidos de programas de monitoreo y colecciones de insectos para desenmarañar el efecto de las diferentes amenazas a las poblaciones de catarinas, comprender el uso del hábitat por taxa de los que se tiene conocimiento limitado y cuantifica las tendencias temporales de la abundancia, diversidad y biomasa a lo largo de un gradiente de intensidad de manejo. Acciones a largo plazo incluyen el desarrollo de un programa de monitoreo a nivel mundial basado en muestreos estandarizados para subsanar la falta de datos, incrementar el poder explicativo, optimizar los análisis y facilitar colaboraciones globales.随着世界范围内人类与野生动物的冲突不断升级, 对野生动物的宽容和接纳等概念变得越来越重要。然而, 当代保护研究表明, 人们对人类与野生动物积极互作的认识有限, 导致可能难以准确描述人类与动物的相遇。如果不解决这些局限性, 就会导致野生动物和景观管理计划的设计和实施存在不足, 以及对本土生态知识的否定。我们利用印度喀拉拉邦瓦亚纳德野生动物保护区森林中的阿迪瓦西族Kattunayakans部落的民族志证据, 来研究印度原住民对于人类与野生动物共存的观点。通过定性的实地研究(包括访谈和在森林样带调查), 我们发现Kattunayakans部落表现出对野生动物的宽容和接纳, 其特点是深入的共存, 包括三个中心思想:野生动物是有理性的交流者;野生动物是神、老师和平等的族群;野生动物是实行 “dharmam”的具有共同起源的亲戚。我们认为, 充分理解以上几点有助于将Kattunayakan部落的观点引入印度的森林管理, 并更广泛地解决人类与野生动物的冲突。.

Wildlife observations submitted by volunteers through citizen science initiatives are increasingly used within research and policy. Ladybirds are popular and charismatic insects, with most species being relatively easy to identify from photographs. Therefore, they are considered an appropriate taxonomic group for engaging people through citizen science initiatives to contribute long-term and large-scale datasets for use in many different contexts. Building on the strengths of a mass participation citizen science survey on ladybirds in the United Kingdom, we have developed a mobile application for ladybird recording and identification across Europe. The main aims of the application are to: (1) compile distribution data for ladybird species throughout Europe, and use this to assess changes in distribution over time; (2) connect and engage people in nature and increase awareness about the diversity and ecological importance of ladybirds. In developing the application we first constructed a database including ladybird species from the United Kingdom, Czech Republic, Slovakia, Italy, Belgium, and Portugal with associated information on relevant morphological features (e.g., size, main color, pronotum pattern) to inform identification. Additionally, the species were assessed on the basis of probability of occurrence within each country which enables users to reduce the number of species to only those with relevance to the location of the recorder. This is amongst the first collaborative citizen science approaches aimed at involving participants across Europe in recording a group of insects. In the near future, we aim to expand the use of the application to all countries in Europe. AOS was supported by FEDER in 85% and by Azorean Public funds by 15% through Operational Program Azores 2020, under the following project AZORESBIOPORTAL -PORBIOTA (ACORES-01-0145FEDER-000072). Further funding for app development was provided by Anglia Ruskin University. info:eu-repo/semantics/publishedVersion

Colour and colour patterns seem to be especially important visual warning signals for predators, which might have innate or learned ability to avoid aposematic prey. To test the importance of larval colour pattern of the aposematic ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), an invasive alien species in Europe, we presented the plasticine models of aposematic larvae to wild and naive birds. We studied the attacks on aposematic larvae of various patterns and colours in nature and in an outdoor aviary. The larvae were cryptic (green), aposematic (resembling those of the H. axyridis larvae), and semi‐aposematic (i.e., black but missing the typical orange patches of H. axyridis larvae). We detected attacks on 71 larvae out of 450 (i.e., 2.6% daily predation). Twenty‐nine attacks were made by birds, 37 by arthropods, and five by gastropods. Wild birds attacked green and black larvae significantly more often than aposematic larvae. Colour did not have an effect on attacks by arthropods. The experiment with naive birds was conducted in an outdoor aviary, where naive great tits, Parus major L., were offered the same artificial larvae as in the first experiment. In total, 57 of 90 exposed larvae were attacked by birds (i.e., 28% daily predation), and green larvae were attacked significantly more than the aposematic larvae (but not more than black larvae). Our results imply that aposematic larvae of H. axyridis are more than 12× less likely to be predated by birds than green larvae in nature. The aposematic pattern represented a more effective signal than the semi‐aposematic signal. The ability to reject aposematic prey seemed to be innate in our birds.

The harlequin ladybird, Harmonia axyridis (Pallas, 1773), is native to Asia but has been introduced to many countries, both intentionally and unintentionally. In the Middle East region, H. axyridis was so far only known from Iran and Turkey. This study reports H. axyridis for the first time from a country with a hot desert climate, Saudi Arabia. The single specimen that was found is most likely the result of unintentional release. Successful spreading of H. axyridis in Saudi Arabia will be limited because of high temperatures during summer and scarcity of prey (aphids). New records from Iran and Turkey suggest fast spreading of H. axyridis in these parts of the Middle East. In addition, we also present new records from Iran and Turkey for H. quadripunctata (Pontoppidan, 1763), the other species in the genus that occurs in the Middle East region. A key and illustrations are provided for both species.

The harlequin ladybird, Harmonia axyridis, is considered to be one of the most invasive insect species worldwide. Its invasion success and extreme speed of range expansion has been partially attributed to weak control of its populations by natural enemies. Previously published data on emergence rates of the hymenopteran parasitoid Dinocampus coccinellae support the enemy release hypothesis: H. axyridis has been consistently less successfully parasitized compared to native ladybird species. In this study, we show that since 2016, i.e., 10 years after its arrival in Central Europe, several populations of H. axyridis in the Czech Republic have a very high prevalence of D. coccinellae parasitism. D. coccinellae emerged from 46% of H. axyridis individuals in the most parasitized population. Moreover, H. axyridis was more parasitized than the native Coccinella septempunctata in seven of nine investigated co-occurring populations. The meta-analytically pooled estimate of D. coccinellae emergence rate from H. axyridis across the Czech populations (this study) is thirteen times higher than the pooled estimate for invasive populations of this beetle elsewhere (historical data up to 2016). We hypothesize that some Central European populations of D. coccinellae have evolved to overcome the immune system of H. axyridis, which was previously thought to be responsible for the high larval mortality of D. coccinellae. As parasitism rates are highly variable in time and space, we encourage future research investigating the determinants of parasitoid prevalence in H. axyridis and other large ladybird species on a continental scale.

1. Few, if any, experimental tests have demonstrated the anti‐predator protection of the developmental stages of ladybirds (Coleoptera: Coccinellidae) against vertebrates, despite the fact that both the visual appearance of ladybirds and the content of defensive compounds fulfil the definition of an aposematic prey. 2. In this study, avian predators of three species were confronted with fourth‐instar larvae and pupae of the harlequin ladybird (Harmonia axyridis) – a large, conspicuous, and toxic ladybird species. 3. The selected predators differed in their individual experience and attitude to ladybirds. Wild‐caught great tits (Parus major) strongly avoided attacking ladybirds in general, whereas wild‐caught tree sparrows (Passer montanus) were willing to include ladybirds in their diet. Domestic chicks (Gallus gallus domesticus) have never been confronted with ladybirds but usually show avoidance of aposematic signals. In this study, great tits and domestic chicks avoided both developmental stages, but in the case of chicks the avoidance had to be learned over the course of repeated encounters. Sparrow avoidance was significantly lower, with more than one‐third of the prey being attacked and eaten. 4. The protection of both developmental stages of ladybirds is similar to adults, despite substantially different visual appearance.

High and low developmental temperatures can induce physiological stress in insects, including oxidative stress that needs to be eliminated by increased antioxidant activity. Alien invasive ladybird Harmonia axyridis (Coleoptera: Coccinellidae) possesses high toxicity and immunity defence abilities, but it is sensitive to temperatures above 35 °C. We quantified the activity of antioxidative enzymes in the adult ladybirds that spent the pupal stage at moderate, low and high temperatures that can cause a physiological stress. Total antioxidant (TAO) and Glutathione S-Transferase (GST) activities of ladybirds H. axyridis were highest in extracts from freshly emerged adults developed at low temperature (17 °C), moderate in beetles from medium temperature (26 °C) and lowest in beetles from high temperature (35 °C). Superoxide dismutase (SOD) activity was highest at the medium temperature. After one day of adult life at the medium temperature, antioxidative activities of individuals originated from the three temperatures became similar: low for TAO and GST and high for SOD. Metabolic needs of the beetle organism during emerging from pupa seem to be stronger than reactions to diverse temperatures.

Harmonia axyridis is an invasive alien ladybird in North America and Europe. Studies show that multiple natural enemies are using Ha. axyridis as a new host. However, thus far, no research has been undertaken to study the effects of simultaneous infections of multiple natural enemies on Ha. axyridis. We hypothesized that high thallus densities of the ectoparasitic fungus Hesperomyces virescens on a ladybird weaken the host’s defenses, thereby making it more susceptible to infection by other natural enemies. We examined mortality of the North American-native Olla v-nigrum and Ha. axyridis co-infected with He. virescens and an entomopathogenic fungus—either Beauveria bassiana or Metarhizium anisopliae. Laboratory assays revealed that He. virescens-infected O. v-nigrum individuals are more susceptible to entomopathogenic fungi, but Ha. axyridis does not suffer the same effects. This is in line with the enemy release hypothesis, which predicts that invasive alien species in new geographic areas experience reduced regulatory effects from natural enemies compared to native species. Considering our results, we can ask how He. virescens affects survival when confronted by other pathogens that previously had little impact on Ha. axyridis.

Based on reviewing the previous relevant literature on Iranian ladybirds as well as identifying and studying coccinellid specimens collected during 2013–2016 from various localities in Lorestan pro...

A species of ladybird beetles: Coccinella leonina transversalis Fabricius, is newly recorded from Iran. Identification keys to Iranian species of the genera Coccinella Linnaeus,, 1758, Pharoscymnus Bedel, 1906, subgenus Scymnus s.str. Kugelann, 1794 and redescription of Pharoscymnus horni (J. Weise, 1901) are given.

Myzus persicae (Sulzer) (Hemiptera: Aphididae) is one of the most common and harmful pest species attacking sweet pepper. Here, we compared the predation rate and performance of Adalia bipunctata (L.), Hippodamia variegata (Goeze), and Scymnus interruptus Goeze (Coleoptera: Coccinellidae) fed on M. persicae infesting sweet pepper. In the laboratory, when offered 150 aphids daily, A. bipunctata was the most voracious species (predation per adult female of 122.7 aphids per 24 h) followed by H. variegata (87.7) and S. interruptus (27.3). When offered 50 aphids, 92% of the S. interruptus larvae developed to adults, 48% of H. variegata, and only 7% of A. bipunctata. In greenhouse experiments, A. bipunctata completely eradicated the aphid population within 2 days following release, whereas H. variegata and S. interruptus initially reduced the aphid population; however, in the long‐term, the infestation was not controlled. Adults of A. bipunctata remained on the plants even in the absence of prey but they laid almost no eggs. Very few adults of H. variegata and S. interruptus were recorded on the plants despite the presence of prey, yet egg laying was observed. Overall, A. bipunctata had an immediate impact on the aphids and is more suitable for curative releases. However, it showed limited potential to establish in the crop. Due to their capacity to complete development with low prey availability, the potential of H. variegata and S. interruptus for preventative releases against aphids in greenhouse crops should be studied further.

Forty-five records of 28 species of ladybird beetles (belonging to 15 genera, 8 tribes and 6 subfamilies of Coccinellidae) as new for particular provinces in Iran are presented. The ten provinces with new records are Alborz, Golestan, Hamadan, Hormozgan, Kohgiluyeh and Boyer-Ahmad, Mazandaran, North Khorasan, Sistan and Baluchistan, Tehran, and Zanjan.

A key to genera of Iranian Coccinellidae, besides species of subfamilies Chilocorinae, Coccinellinae, Epilachinae and Microweiseinae is given.

Understanding the factors restricting the distribution of some insect species to high altitudes is hindered by poor knowledge of temporal changes in their cold hardiness during overwintering. We studied overwintering larvae of five species of Erebia butterflies (Lepidoptera: Nymphalidae: Satyrinae) differing in altitudinal distribution: lowland E. medusa, submountain E. aethiops, subalpine E. pronoe, alpine E. cassioides, and subnivean E. pluto. We subjected them to three treatments, AutumnWarm (13/8°C), imitating conditions prior to overwintering; AutumnCold (5/0°C), imitating late autumn conditions; and WinterCold (5/0°C), differing from AutumnCold by a shorter photoperiod and longer exposure to zero temperatures. Supercooling points (SCP) did not differ between species in the AutumnWarm treatment, despite large differences in the concentrations of cryoprotectants (CrPC; lowest in E. medusa and E. aethiops). Lowland E. medusa was freeze-tolerant, the subalpine, alpine and subnivean species were freeze-avoidant, whereas submountain E. aethiops displayed a mixed strategy. SCPs diverged in the AutumnCold treatment: it increased in the lowland E. medusa (from -16.5 to -10.8°C) and reached the lowest value in E. cassioides (-21.7°C). In WinterCold, SCP increased in subalpine E. pronoe (from -16.1°C in AutumnWarm and -18.7°C in AutumnCold to -12.6°C). E. medusa decreased and E. aethiops increased their CrPCs between autumn and winter; the highest CrPC was recorded in subnivean E. pluto. CrPC did not correlate with SCP across species and treatments. Cryoprotectant profiles corroborated the difference between lowland and freeze-tolerant E. medusa and the three high altitude freeze-avoidant species, with E. aethiops in an intermediate position. Glycerol was surprisingly rare, trehalose was important in all species, and such rare compounds as monopalmitin and monostearin were abundantly present in E. pronoe, E. cassioides and E. pluto.

We report on two lady beetle species, Oenopia conglobata (Linnaeus) and Adalia bipunctata (Linnaeus), sympatric in Chaharmahal and Bakhtiari Province, Iran, closely resembling each other in coloration. This may be an interesting example of Mullerian mimicry. The putative model, O. conglobata, retains its typical pink elytral background with 16 black spots, whereas the locally most common form (forma fasciatopunctata) of the polymorphic mimic, A. bipunctata, exhibits a similar pink background that is otherwise absent in the species. This pink coloration is achieved by a small amount of red pigments deposited over the white color of the lower layer of the elytra.

Intraguild predation (IGP) is a ubiquitous, important and common interaction that occurs in aphidophagous guilds. The effects of extraguild prey (EGP, i.e., aphids) density, predator life stage combinations and duration of the interaction on the level, asymmetry and direction of intraguild predation between lacewing Chrysoperla carnea and ladybird Hippodamia variegata were examined in simple laboratory arena and more complex microcosm environment. Three initial densities of 50, 150 and 400 Aphis fabae third instar nymphs and a control without aphids were provided to six combinations of predator life stages (2nd and 3rd larval instars of lacewing and 3rd and 4th instars and adult females of ladybird). The remaining aphid density and occurrence of IGP were checked after 24, 48 and 72 h. The IGP intensity (IGP level, IL) was similar in the simple arena (reaching 0.6 between larvae in absence of EGP and 0.3 between lacewing larvae and ladybird females) and microcosm environment (0.3 without EGP). In both environments, increasing EGP density lowered IL according to negative exponential relationship. IGP was asymmetric (general average asymmetry was 0.82 in simple arena and 0.93 in microcosm, the difference was not significant) and mostly in favour of larvae of C. carnea, except in the combination of 2nd larvae of C. carnea with the 4th larvae and adults of H. variegata. The direction of IGP, but not other characteristics, partially changed during the duration of the experiment. The incidence of IGP interactions among aphid predators under real conditions and its consequences on aphid biological control are discussed.

The ectoparasitic fungus Hesperomyces virescens was studied on its invasive host, the harlequin ladybird Harmonia axyridis, in the Czech Republic. A primary aim was to examine the relationship between fungal infection and elytral coloration of the ladybird. Furthermore, the role of host sex and mating status of females were analyzed. Beetles (n = 1,102) were sampled during autumn migration, and then sexed, weighed, and screened for infection. Females were dissected for detection of sperm in their spermathecae. Ladybirds were sorted according to color form and absorbance spectrophotometry was used to quantify carotenoid contents in their elytra. In individuals of the nonmelanic succinea form, the degree of melanization was measured using digital photographs and putative age groups were estimated based on background color of elytra. Sexual differences in infection patterns indicated transmission during copulation: males were infected mostly on elytra and venter, and females had infection almost exclusively on elytra. Mated females had higher infection rate than virgins. There was no influence of genetic color form on the fungal infection. Putative age groups (visual sorting to yellow, orange, and red) correlated with fungal infection. Infected individuals had elevated elytral carotenoid levels in comparison to uninfected individuals, which could be explained by host age. Infection-free succinea beetles were extensively melanized because they emerged later in the season at lower temperatures which induced melanization. Overall, we highlight that H. axyridis is a multivoltine species whose age, if not taken into account in ecophysiological studies, might present a considerable confounding factor.

We evaluated the status of native ladybird assemblages in the presence of the non-native ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) in three European countries. To achieve this, we assessed the abundance of ladybirds from common host plants in urban areas of Great Britain, Czech Republic and Slovak Republic in 2014. We determined (i) the effects of season, host plant, location, abundance of H. axyridis and aphids on the ladybird species assemblage, (ii) the relationship between ladybird abundance and aphid density, (iii) the relationship between diversity of native ladybird species and the abundance of H. axyridis. There was a non-linear relationship between abundance of ladybirds and abundance of aphids but we found the abundance of other ladybird species was not correlated with H. axyridis. We highlight the value of large-spatial scale studies for revealing patterns in community assemblages and ultimately informing understanding of ecosystem resilience.

The Asian lady beetle Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is one of the most invasive insects worldwide. It has been introduced into several continents for classical biological control and by accident. The current study is the first report of this species from Iran (new country record). All developmental stages and adults of two morphs, succinea and spectabilis, were collected from fruit trees and ornamental gardens in the city of Rasht in the northern Iranian province Guilan in 2016. We collected 249 adults of the color morph succinea and 31 individuals (11%) of the melanic color morph spectabilis. Only 36% of individuals were males. Larvae and pupae were found from April to September, adults from April to October.

Fungal species identities are often based on morphological features, but current molecular phylogenetic and other approaches almost always lead to the discovery of multiple species in single morpho-species. According to the morphological species concept, the ant-parasitic fungusRickia wasmannii(Ascomycota, Laboulbeniales) is a single species with pan-European distribution and a wide host range. Since its description, it has been reported from ten species ofMyrmica(Hymenoptera, Formicidae), of which two belong to therubra-group and the other eight to the phylogenetically distinctscabrinodis-group. We found evidence forR. wasmanniibeing a single phylogenetic species using sequence data from two loci. Apparently, the original morphological description (dating back to 1899) represents a single phylogenetic species. Furthermore, the biology and host-parasite interactions ofR. wasmanniiare not likely to be affected by genetic divergence among different populations of the fungus, implying comparability among studies conducted on members of different ant populations. We found no differences in total thallus number on workers betweenMyrmicaspecies, but we did observe differences in the pattern of thallus distribution over the body. The locus of infection is the frontal side of the head inMyrmica rubraandM. sabuletiwhereas inM. scabrinodisthe locus of infection differs between worker ants from Hungary (gaster tergites) and the Netherlands (frontal head). Possible explanations for these observations are differences among host species and among populations of the same species in (i) how ant workers come into contact with the fungus, (ii) grooming efficacy, and (iii) cuticle surface characteristics.

International audience; We investigated the mating and fertilization patterns in one Czech local population of the invasive aphidophagous ladybird Harmonia axyridis (Pallas), sampling the same mixed generations before and after overwintering. Our study using eight microsatellite loci provides the first direct evidence that females of H. axyridis are highly polyandrous in the wild and that they not only mate multiple times but also use the sperm from multiple males to fertilize their eggs. Three types of values (minimum number of fathers, estimated number of fathers and effective number of fathers contributing to progeny of one mother) were about two times higher in the spring sample (2.9, 5.2, 3.1), in the peak of their reproduction period, than in the preceding autumn sample (1.6, 2.8, 1.6). The contribution of individual fathers to the number of progeny was highly unbalanced, with a few dominating fathers and many fathers with low number of progeny.

The harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae) has rapidly spread in several continents over the past 30 years and is considered an invasive alien species. The success of H. axyridis as an invader is often attributed to weak control by natural enemies. In this paper, we provide an overview of current knowledge on predators and parasitoids of H. axyridis. The common feature of predators and parasitoids is that they directly kill exploited organisms. Currently available data show that H. axyridis, displaying a variety of chemical, mechanical, and microbiological anti-predator defenses, is usually avoided by predators. However, some birds and invertebrates can eat this ladybird without harmful consequences. The primary defenses of H. axyridis against parasitoids include immune response and physiological and nutritional unsuitability for parasitoid development. These defenses are probably relatively efficient against most ladybird parasitoids, but not against flies of the genus Phalacrotophora. The latter are idiobiont parasitoids and hence can evade the host’s immune response. Indeed, rates of parasitism of H. axyridis by Phalacrotophora in the Palaearctic region (both in the native range in Asia and in Europe) are relatively high. While strong evidence for enemy release on the invasive populations of H. axyridis is lacking, several cases of parasitoid acquisition have been recorded in Europe, North America, and South America. We conclude that enemy release cannot be excluded as a possible mechanism contributing to the spread and increase of H. axyridis in the early stages of invasion, but adaptation of parasitoids may lead to novel associations which might offset previous effects of enemy release. However, further work is required to elucidate the population-level effects of such interactions.