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Oenopia shirkuhensissp. nov. (Coleoptera, Coccinellidae) is described and illustrated. It was found in the mountains around Shirkooh mountain, Yazd province, and in the Kukhbenan Mountains, Kerman province, Iran. It is similar to a common ladybird Adalia bipunctata by the colour pattern on elytra. Congeneric species occurring in Iran, O. conglobata and partly O. oncina are illustrated for comparison, and an identification key is provided.

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.

The genus Coccidula Kugelann includes five species distributed in the Holarctic, with one species in North America and four in Palearctic region. Coccidula belongs to the tribe Coccidulini which historically was treated as a separate subfamily within ladybird beetles, but recent studies confirmed its placement as a tribe within the broadly defined subfamily Coccinellinae. All species are revised and a new synonymy of Lithophilus naviauxi Duverger with C. litophiloides Reitter is proposed. Light and electron microscopy pictures support morphological descriptions. An identification key to all species is also provided.

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.

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.

An updated and corrected checklist of species of ladybird beetles (Coleoptera: Coccinellidae) known in Algeria now contains 75 species belonging in ten tribes. New country records include the European species Oenopiaconglobata and the invasive Asian species Harmoniaaxyridis. Sampling data is provided for 14 species found during a faunistic survey performed mostly in agroecosystems, together with host plant and prey species.

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.

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.

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.

We surveyed ladybirds (Coleoptera: Coccinellidae) in 10 stands of Scots pine (Pinus sylvestris), all monoculture stands 5-100 years old, in western Slovakia, Central Europe, over two successive periods, October 2013 - March 2014 and October 2014 - March 2015. The winter in each period was exceptionally mild. Ladybirds were collected from the lower branches of pine trees using beating trays and were present in 61% of the 1040 samples (one sample containing ladybirds from 20 branches, 1 m long each). In total 3965 individuals of 20 species were recorded. Non-conifer dwelling species associated with broadleaved trees or herbaceous plants prevailed (45% of species), followed by conifer specialists (40%) and generalists (15%). Although 13 species were found at least in one winter month, December, January or February, only four of them, Exochomus quadripustulatus, Coccinella septempunctata, Harmonia axyridis and Hippodamia variegata, were recorded continually during both winters. The number of species, the abundance of all ladybirds and the abundance of dominant species (E. quadripustulatus, C. septempunctata and H. axyridis) decreased from late autumn towards winter and remained lowest during this most adverse time of the year for ladybirds. Overwintering species assemblages of ladybirds changed over time and varied with age of pine stand. Our results suggest that Scots pine in Central Europe supports species rich assemblages of ladybirds from late autumn to early spring and, being widely distributed, it could be suited to winter surveying of ladybirds at large spatial scales to reveal behavioural and ecological responses of species to changing weather or different climates.

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.

Background The tribe Coccinellini is a group of relatively large ladybird beetles that exhibits remarkable morphological and biological diversity. Many species are aphidophagous, feeding as larvae and adults on aphids, but some species also feed on other hemipterous insects (i.e., heteropterans, psyllids, whiteflies), beetle and moth larvae, pollen, fungal spores, and even plant tissue. Several species are biological control agents or widespread invasive species (e.g., Harmonia axyridis (Pallas)). Despite the ecological importance of this tribe, relatively little is known about the phylogenetic relationships within it. The generic concepts within the tribe Coccinellini are unstable and do not reflect a natural classification, being largely based on regional revisions. This impedes the phylogenetic study of important traits of Coccinellidae at a global scale (e.g. the evolution of food preferences and biogeography). Results We present the most comprehensive phylogenetic analysis of Coccinellini to date, based on three nuclear and one mitochondrial gene sequences of 38 taxa, which represent all major Coccinellini lineages. The phylogenetic reconstruction supports the monophyly of Coccinellini and its sister group relationship to Chilocorini. Within Coccinellini, three major clades were recovered that do not correspond to any previously recognised divisions, questioning the traditional differentiation between Halyziini, Discotomini, Tytthaspidini, and Singhikaliini. Ancestral state reconstructions of food preferences and morphological characters support the idea of aphidophagy being the ancestral state in Coccinellini. This indicates a transition from putative obligate scale feeders, as seen in the closely related Chilocorini, to more agile general predators. Conclusions Our results suggest that the classification of Coccinellini has been misled by convergence in morphological traits. The evolutionary history of Coccinellini has been very dynamic in respect to changes in host preferences, involving multiple independent host switches from different insect orders to fungal spores and plants tissues. General predation on ephemeral aphids might have created an opportunity to easily adapt to mixed or specialised diets (e.g. obligate mycophagy, herbivory, predation on various hemipteroids or larvae of leaf beetles (Chrysomelidae)). The generally long-lived adults of Coccinellini can consume pollen and floral nectars, thereby surviving periods of low prey frequency. This capacity might have played a central role in the diversification history of Coccinellini. Electronic supplementary material The online version of this article (doi:10.1186/s12862-017-1002-3) contains supplementary material, which is available to authorized users.

The alien invasive ladybird Harmonia axyridis (Pallas, 1773) (Coleoptera: Coccinellidae) was recorded from Tanzania, Zanzibar for the first time in 2014. This is in total the sixth country of this species in Africa and the second close to the equator. We suggest that repeated introductions may lead to new outbreaks of the species.

The Iranian species of the tribe Hyperaspidini Mulsant, 1846 (Coleoptera: Coccinellidae) are reviewed. The current list includes 12 species, all placed in a single genus Hyperaspis Chevrolat, 1836. Hyperapsis asiatica Lewis, 1896 and H. pumila Mulsant, 1850 are excluded from the Iranian list of Coccinellidae. Diagnoses of the tribe Hyperaspidini and the genus Hyperaspis are given. Images of adult beetles and diagnostic characters of the male genitalia of all species distributed in Iran are shown. A key to identification of the species is presented. Distribution records are provided for each species along with information on host plants and prey species when available.

Harmonia axyridis (Coleoptera: Coccinellidae) has been introduced widely for biological control of agricultural pests. Harmonia axyridis has established in four continents outside of its native range in Asia and it is considered an invasive alien species (IAS). Despite a large body of work on invasion ecology, establishment mechanisms of IAS and their interactions with natural enemies remain open questions. Parasites, defined as multicellular organisms that do not directly kill the host, could potentially play an important role in regulating host populations. This study presents a review of the parasites of H. axyridis, discussing their distributions and effects on host populations across the host’s native and invasive range. These parasites are: Hesperomyces virescens Thaxt. fungi, Coccipolipus hippodamiae (McDaniel and Morrill) mites, and Parasitylenchus bifurcatus Poinar and Steenberg nematodes.

International audience; We investigated factors which may determine realised paternity following multiple mating in a promiscuous ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), namely the mating order and ability of males of different morphs to fertilize females. In experiments using males of two colour morphs, we found that the first male had a reproductive advantage over the second male (sperm precedence P2 = 0.38). Males of the melanic morph spectabilis had a reproductive advantage over males of the semi-melanic morph axyridis (P2 = 0.35). Analysis of paternity using microsatellites applied to young larvae, which avoids possible differential mortality of progeny, confirmed the precedence of the first male and higher paternity rate of colour morphs that are more frequent in the wild. Hatchability of eggs increased from 65% after the first mating to 85% after the second mating delayed by five days. Average duration of mating was 157 min and did not differ between the first and second mating and between the colour morphs of males.

Growing evidence exists that aposematic and toxic prey may be included in a predator’s diet when the predator experiences physiological stress. The tree sparrow Passer montanus is known to have a significant portion of aposematic and toxic ladybirds in its natural diet. Here, we present experiments testing the attack and eating rate of the tree sparrow toward the invasive aposematic harlequin ladybird Harmonia axyridis. We wondered whether the sparrow’s ability to prey on native ladybirds predisposes them to also prey on harlequin ladybirds. We compared the attack and eating rates of tree sparrows of particular age and/or experience classes to test for any changes during ontogeny (hand-reared × young wild-caught ×adult wild-caught) and with differing perceived levels of physiological stress (summer adult × winter adult). Winter adult tree sparrows commonly attacked and ate the offered ladybirds with no evidence of disgust or metabolic difficulties after ingestion. Naïve and wild immature tree sparrows attacked the ladybirds but hesitated to eat them. Adult tree sparrows caught in the summer avoided attacking the ladybirds. These results suggest that tree sparrows are able to cope with chemicals ingested along with the ladybirds. This pre-adaptation enables them to include ladybirds in their diet; though they commonly do this only in times of shortage in insect availability (winter). Young sparrows showed avoidance toward the chemical protection of the ladybirds.