A roadmap for ladybird conservation and recovery.

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.

Effect of Male Body Size on Female Reproduction in Pyrrhocoris apterus (L.) (Heteroptera, Pyrrhocoridae).

Females and males of the abundant heteropteran species Pyrrhocoris apterus (L.) mate with the larger of the available partners. The male benefits from copulation with a large female, which is more productive than a small female. However, the benefit to females from copulation with a large male has not yet been investigated. Overwintered sexually immature adults were transferred from the field to indoors for a long day at 25 °C and subsequently kept as female-male pairs with different partner size ratios. The female lays eggs in several successive clutches. We investigated the influence of male size on the number and weight of eggs in individual clutches and the time elapsed from the transfer to their deposition. These characteristics of reproduction were first regressed on female size, and residuals of this regression were then regressed on male size. The positive effect of increasing male size on the characteristics of female reproduction manifested as a significant value of the latter regressions. The expected positive effect of increasing male body size on female reproductive characteristics was not detected. Several reasons for this deficiency are discussed.

Which Seed Properties Determine the Preferences of Carabid Beetle Seed Predators?

Ground beetles are important invertebrate seed predators in temperate agro-ecosystems. However, there is a lack of information regarding which seed properties are important to carabids when they select seeds for consumption. Therefore, seed properties, such as size, shape, morphological defence, and chemical composition, were measured, and in addition to seed taxonomy and ecology, these data were used to explain carabid preferences. Carabid preferences were assessed using a multi-choice experiment with 28 species of weed seeds presented to 37 species of Carabidae. Multiple regression on distance matrices (MRM) was used to determine the importance of particular sets of seed properties for carabids. The analysis was conducted for the full set of carabids (37 species) as well as for subsets of species belonging to the tribes of Harpalini or Zabrini. For the complete set of species, seed dimensions, seed mass, taxonomy, plant strategy, and seed coat properties significantly explained carabid preferences (proportion of explained variance, R2 = 0.465). The model for Harpalini fit the data comparably well (R2 = 0.477), and seed dimensions, seed mass and seed coat properties were significant. In comparison to that for Harpalini, the model for Zabrini had much lower explanatory power (R2 = 0.248), and the properties that significantly affected the preferences were seed dimensions, seed mass, taxonomy, plant strategy, and seed coat properties. This result suggests that the seed traits that carabids respond to may be specific to taxonomic and likely relate to the degree of specialisation for seeds. This study contributes to understanding the mechanisms that determine the preferences of carabid beetles for seeds.

Behavioural thermoregulation hastens spring mating activity in Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae).

Post-diapause, overwintered adults of the true bug Pyrrhocoris apterus L. (Heteroptera, Pyrrhocoridae) form conspicuous aggregations at warm spots in early spring. Using a combination of laboratory experiments and field observations, we assessed the influence of this behaviour on the seasonal timing of reproduction. In the laboratory, post-diapause pairs mated after an accumulation of 80° days (dd), and females started to lay eggs after an additional 80 dd (considering 10.3 °C as a lower temperature threshold for post-diapause development). In the field, however, females mated as early as late March (based on data from four seasons), which corresponded to the accumulation of <15 dd (based on summing the temperatures recorded at a meteorology station). Such a 'discrepancy' between laboratory and field data is explained by thermoregulation. Aggregated adults bask in sunny spots, thereby increasing their body temperatures to ~25 °C, which is 7-16 °C above the temperature of the surrounding ground. This thermal excess speeds up their post-diapause development and enables early mating and oviposition. As a result, behavioural thermoregulation hastens the seasonal start of reproduction in P. apterus by > 1 month.

Differences in the Phenology of Harmonia axyridis (Coleoptera: Coccinellidae) and Native Coccinellids in Central Europe.

Harmonia axyridis (Pallas), an invasive non-native species in central Europe, can outcompete other aphidophagous species. The distribution and abundance of H. axyridis vary depending on different host plants, and its effects on native coccinellid communities may change accordingly. The distribution and abundance of coccinellids in central Europe (50°N, 14°E) were investigated from 2010 to 2016. Coccinellids were counted at regular intervals on cereals (Avena, Hordeum, and Triticum), herbaceous plants (Matricaria and Urtica) and trees (Acer, Betula, and Tilia). Additionally, the occurrence over time of each species on these plants was assessed and used as an index of persistence. Across all years, the adults and larvae of H. axyridis were the dominant species of coccinellid on trees. However, H. axyridis was less abundant on herbaceous plants and cereals than on trees. Populations of native coccinellids and H. axyridis co-occurred on trees and persisted for the same length of time, while native coccinellids persisted longer than H. axyridis on herbaceous plants and cereals. Compared to 1976-1986, in the 2010s, the abundance of native species decreased on all plants by 50-70%. The presence of H. axyridis could be considered as a factor driving changes in the assemblages of native coccinellids.

Aphids (Homoptera: Aphididae) on Winter Wheat: Predicting Maximum Abundance of Metopolophium dirhodum.

In Central Europe, the most abundant aphid infesting the leaves of small grain cereals is Metopolophium dirhodum (Walker) (Homoptera: Aphididae). Annual variation in its seasonal dynamics was evaluated using a 25-yr series of standardized weekly censuses of winter wheat plots. M. dirhodum made up >50 % of the aphids on the foliage. Date of immigration (8 May-3 July), length of period of population increase (0-9 wk), and date of attaining maximum abundance (28 May-22 July) varied greatly. For the prediction, we regressed maximum numbers/tiller on numbers recorded in the first week after heading. The regression of maximum abundance on nonzero aphid counts revealed a critical number of ≥1.50 aphids/tiller, which if exceeded resulted in a harmful maximum abundance of ≥10 aphids/tiller at the peak. Zero aphid counts resulted in 10% of cases with a harmful maximum abundance. Using this regression for prediction will result in 18% of the recorded cases being false negatives and 9% false positives. Parallel annual variation in the average maximum numbers of M. dirhodum, Sitobion avenae (Fabricius) (Homoptera: Aphididae), and Rhopalosiphum padi (Linné) (Homoptera: Aphididae) indicated the following factors that affected their abundance: temperature in winter and host plant quality. The predictions apply only in areas where M. dirhodum is holocyclic and aphids do not overwinter in wheat stands.

Estimating Prey Consumption in Natural Populations of Harmonia axyridis (Coleoptera: Coccinellidae) Using Production of Feces.

Production of feces (PF) is a useful proxy indicating quantity of ingested food. Although influenced by many uncontrolled factors PF provides insight into food consumption under natural conditions. In Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) variation in PF was investigated in populations of various hostplants at localities of central Bohemia (50N, 14E), throughout the season of 2016. The adults collected from these hostplants were starved for 48 h at standard conditions, and dry mass of feces produced during this period was measured. Despite enormous differences in PF among individuals, significant variation over the season occurred in average PF of both males and females. PF increased with abundance of aphids and was significantly greater in females than males. Gravidity, as manifested through oviposition within 48 h after capture, was associated with increased PF, while hostplant and color morph did not affect variation in PF among individuals. From PF as measured in this study, it can be estimated that at sites hosting abundant aphid populations H. axyridis (as an adult male or female) may consume 19 (male) to 45 (female) aphids per day (assumed body length 1mm). In the absence of aphids, adults may consume one to nine individuals of alternative prey per day (body length 1-2 mm).

Spatial and temporal changes in the abundance and compostion of ladybird (Coleoptera: Coccinellidae) communities.

Because of their services to agriculture most ladybirds (Coleoptera: Coccinellidae) are intensively studied predators of mainly phytophagous pests. The study of the long-term variation in the composition of their communities was stimulated by recent dramatic changes in the abundance of some species. We review and evaluate possible effects of the main causes cited in the literature. Agricultural and habitat changes (particularly urbanization) affect coccinellid abundance, both negatively and positively. In the temperate zone dominant species occur most frequently associated with abundant prey populations on crops, weeds and planted stands of trees resulting from human activity. Invasive non-native species of coccinellids may endanger native species through intraguild predation or competition for resources, but their supposed serious negative effects on native species can differ considerably. Climatic change may influence coccinellid species in several ways, including indirect effects through lower trophic levels and desynchronisation of the phenologies of host plants, prey and coccinellid populations. In the near future we do not expect climate warming to have important effects on ladybird diversity globally, but local changes in the composition of coccinellid communities and abundance of particular species could occur.

Plasticity of the thermal requirements of exotherms and adaptation to environmental conditions.

UNLABELLED: In exothermal organisms, temperature is an important determinant of the rate of ecophysiological processes, which monotonically increase between the minimum (t d min) and maximum (t d max) temperatures typical for each species. In insects, t d min and t d max are correlated and there is a approximately 20°C interval (thermal window W T = t d max - t d min) between them over which insects can develop. We assumed that other exotherms have similar thermal windows because the thermal kinetics of their physiological processes are similar. In this study, we determined the thermal requirements for germination in plants. Seeds of 125 species of Central European wild herbaceous and crop plants were germinated at nine constant temperatures between 5 and 37°C, and the time to germination of 50% of the seeds D and rate of germination R (=1/D) were determined for each temperature and the Lactin model used to determine t d min, t d max, and W T. The average width of the thermal windows for seeds was significantly wider (mean 24°C, 95% CI 22.7-24.2°C), varied more (between 14.5 and 37.5°C) and development occurred at lower temperatures than recorded for insects. The limiting temperatures for germination, t d min and t d max, were not coupled, so the width of the thermal window increased with both a decrease in t d min and/or increase in t d max. Variation in W T was not associated with taxonomic affiliation, adult longevity, or domestication of the different species, but tends to vary with seed size. Plants are poor at regulating their temperature and cannot move to a more suitable location and as a consequence have to cope with wider ranges in temperatures than insects and possibly do this by having wider thermal windows. SYNTHESIS: The study indicated specificity of W T in different exotherm taxa and/or their development stages.

Population dynamics of aphids on cereals: digging in the time-series data to reveal population regulation caused by temperature.

Aphid populations show periodic fluctuations and many causes are attributed to their dynamic. We investigated the regulation by temperature of the aphid populations composed of Metopolophium dirhodum, Sitobion avenae, and Rhopalosiphum padi on winter wheat using a 24 years long time series data. We computed the sum of daily temperatures above 5 °C, the threshold temperature for aphid development, and the sum of daily temperatures within the [0(threshold for wheat development),5] °C interval. Applying Generalised Additive Model framework we tested influences of temperature history expressed via degree days before the start of the aphid immigration on the length of their occurrence. We aimed to estimate the magnitude and direction of this influence, and how far to the past before the start of the aphid season the temperature effect goes and then identify processes responsible for the effect. We fitted four models that differed in the way of correcting for abundance in the previous year and in specification of temperature effects. Abundance in the previous year did not affect the length of period of aphid population growth on wheat. The temperature effect on the period length increased up to 123 days before the start of the current season, i.e. when wheat completed vernalization. Increased sum of daily temperatures above 5 °C and the sum of daily temperatures within the [0,5] °C interval both shortened the length of period of aphid population growth. Stronger effect of the latter suggests that wheat can escape from aphid attacks if during winter temperatures range from 0 to 5 °C. The temperature influence was not homogeneous in time. The strongest effect of past temperature was about 50 to 80 and 90 to 110 days before the beginning of the current aphid season indicating important role of termination of aphid egg dormancy and egg hatching.

The invasion history, distribution and colour pattern forms of the harlequin ladybird beetle Harmonia axyridis (Pall.) (Coleoptera, Coccinellidae) in Slovakia, Central Europe.

The harlequin ladybird beetle Harmonia axyridis (Coleoptera, Coccinellidae) has invaded and established in Slovakia. Following unintentional introduction in 2008, the spread of the alien coccinellid was very fast. By the end of 2009, it was recorded across the whole country, and by the end of 2012 it was widely distributed and common in various habitats, particularly gardens, orchards and urban areas, where it was most frequent on trees. The rate of eastward spread was approximately 200 km year(-1), similar to the overall rate of spread in Europe. Between 2008 and 2012, the coccinellid was recorded in a total of 153 localities, in altitudes ranging from 98 to 1,250 m. Most records of this species were made in lowlands, hilly areas and valleys separating mountain ridges. However, it was only rarely documented in areas above 700 m a.s.l. The non-melanic colour form (f. succinea) was dominant along a longitudinal transect including eight urban areas across Slovakia, with the frequency of melanic forms (f. spectabilis and f. conspicua together) between 6.3 and 19.2% and a median equal to 10.5%. The invasion history and distribution of H. axyridis in Slovakia are discussed with regard to the time sequence of records, rate of spread, altitudinal distribution, anthropogenic dispersal, effective recording, proportion of melanic forms and other relevant aspects associated with the spread of this successful invader.

Geographic variation in seed dormancy among populations of Echinochloa crus-galli.

In 1991-1993, we investigated the incidence of seed dormancy in 25 local populations of barnyard grass, Echinochloa crus-galli (L.) P.Beauv., in the western Czech Republic. The percentage of germination after 4 months afterripening of dry seeds at 25°C varied between 0.0 and 83.6%. Although there were significant annual differences in the percentage of germination at some localities, typical proportions of dormant seeds persisted over 3 years at field sites where the seed bank was not disturbed. One-way ANOVA (using data from 14 cultivated or abandoned fields) revealed that 73.0% of variance in seed dormancy incidence could be attributed to the effect of locality (P<0.001). Incidence of dormancy was not correlated with mother plant stature (dry above-ground biomass, number of tillers, maximal stem height) nor seed mass. There was a significant correlation (r 2=0.403, P<0.005) between dormancy incidence at natural localities in 1991 and in F1 offspring sown at experimental grounds at Praha-Ruzyne in 1992. The results indicate that heredity is important in maintaining local variation in seed dormancy, probably favoured by the self-pollinating reproduction of barnyard grass.