Marina J. Orlova-Bienkowskaja

The harlequin ladybird, Harmonia axyridis: global perspectives on invasion history and ecology

The harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), is native to Asia but has been intentionally introduced to many countries as a biological control agent of pest insects. In numerous countries, however, it has been introduced unintentionally. The dramatic spread of H. axyridis within many countries has been met with considerable trepidation. It is a generalist top predator, able to thrive in many habitats and across wide climatic conditions. It poses a threat to biodiversity, particularly aphidophagous insects, through competition and predation, and in many countries adverse effects have been reported on other species, particularly coccinellids. However, the patterns are not consistent around the world and seem to be affected by many factors including landscape and climate. Research on H. axyridis has provided detailed insights into invasion biology from broad patterns and processes to approaches in surveillance and monitoring. An impressive number of studies on this alien species have provided mechanistic evidence alongside models explaining large-scale patterns and processes. The involvement of citizens in monitoring this species in a number of countries around the world is inspiring and has provided data on scales that would be otherwise unachievable. Harmonia axyridis has successfully been used as a model invasive alien species and has been the inspiration for global collaborations at various scales. There is considerable scope to expand the research and associated collaborations, particularly to increase the breadth of parallel studies conducted in the native and invaded regions. Indeed a qualitative comparison of biological traits across the native and invaded range suggests that there are differences which ultimately could influence the population dynamics of this invader. Here we provide an overview of the invasion history and ecology of H. axyridis globally with consideration of future research perspectives. We reflect broadly on the contributions of such research to our understanding of invasion biology while also informing policy and people.

Expansion of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) to European Russia and adjacent regions

Abstract An invasive alien species, the harlequin ladybird Harmonia axyridis (Pallas, 1773), has quickly expanded its distribution in Eastern Europe. Records of H. axyridis from 31 localities in Lithuania, Latvia, the Ukraine, European Russia, and the Northern Caucasus are summarized and mapped. Within the last few years this species has established in south Latvia, on the Baltic Sea shore (Kaliningrad oblast and Lithuania), in the western and central Ukraine, Crimea, and in the Northern Caucasus. Besides that, individual specimens have been found in 4 more localities in European Russia. The species is recorded from Lipetsk oblast (European Russia), Crimea, and Nikolaev oblast (the Ukraine) for the first time.

Harmonia axyridis (Coleoptera: Coccinellidae) in Asia: a re-examination of the native range and invasion to southeastern Kazakhstan and Kyrgyzstan

Harmonia axyridis (Pallas) is an invasive ladybird spreading in several continents. It is native to East Asia, but its range in Asia has until now been poorly understood. The most complete map of the range of Harmonia axyridis in Asia has been compiled (432 localities). Harmonia axyridis occurs in the south-east of West Siberia, the south of East Siberia, the south of Russian Far East, the east of Kazakhstan, the north of Kyrgyzstan, Mongolia, China, North Korea, South Korea, Japan, and the north of Vietnam. Southeastern Kazakhstan and the north of Kyrgyzstan are not a part of the native range: the large Balkhash Desert is situated between this mountainous region and the nearest part of the native range (Altai mountains). The analysis of dates and places of findings indicates that Harmonia axyridis appeared in southeastern Kazakhstan and Kyrgyzstan after the construction of the Turkestan-Siberian Railway. It is hypothesized that the beetles crossed the desert along this railway.

The life cycle of the emerald ash borer Agrilus planipennis in European Russia and comparisons with its life cycles in Asia and North America

Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), native to Asia, is a destructive invasive pest of ash Fraxinus spp. in U.S.A., Canada and European Russia. It is spreading quickly and will probably soon be detected in other European countries. Information about the life cycle of the pest is needed for detection and survey efforts, development of control options and predicting the potential range in Europe. The life cycle has been studied in North America and Asia, although it has not previously been studied in European Russia. The number of larval instars and the duration of development of A. planipennis in European Russia were determined. Distributions of width of epistome and length of urogomphi indicated four larval instars. The number of excretory ducts and the presence of ledges did not provide a clear differentiation between instars. Development in most of the specimens took 2 years. Generation time is flexible. In warmer regions (Tianjin), most individuals finish development in 1 year, whereas, in colder regions (Moscow, Changchun, Harbin), it takes 2 years. In intermediate climatic regions (Michigan), the ratio of 1 : 2‐year life cycles depends on additional factors. The flexibility of the life cycle allows A. planipennis to establish in regions with different climates.

A revision of the Simocephalus (latirostris) species group (Crustacea: Anomopoda: Daphniidae)

The group of 3 species: Simocephalus latirostris, S. lusaticus and a new species, S. mesorostris, is given the rank of a subgenus. Descriptions and a key for the identification of these species are given. The unknown male of S. latirostris is described. The comparison of the lectotype of S. latitrostis with material from different regions shows that this species is not pantropical as previously supposed. It occurs only in South and Central America. S. mesorostris n. sp., living in Australia, the Malay Archipelago, South-East Asia and Africa, was previously confused with S. latirostris.

World checklist of flea-beetles of the genus Epitrix (Coleoptera: Chrysomelidae: Galerucinae: Alticini)

The world checklist of the genus Epitrix (Coleoptera: Chrysomelidae: Galerucinae: Alticini) is necessary, since many species of this genus are serious pests of potato and other Solanaceae, and since some species have been inadvertedly introduced from one continent to another and established. We have compiled the catalogue of all species described to date. There are 162 species and 11 subspecies in the world. The geographic distribution is indicated for each species.

Are native ranges of the most destructive invasive pests well known? A case study of the native range of the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae)

Knowledge of the native range of invasive pests is vital for understanding their biology, for ecological niche modeling to infer potential invasive distribution, and for searching of natural enemies. Standard descriptions of pest ranges frequently pass from one publication to another without verification. Our goal is to test the reliability of distributional information exemplified by the native range of one of the most destructive and most studied invasive forest insect pests of Asian origin—the emerald ash borer (EAB), Agrilus planipennis Fairmaire. Since the first detections of this notorious insect pest in North America in 2002 and European Russia in 2003, it has killed hundreds of millions of ash trees. Based on the examination of museum specimens and literature sources we compiled the most comprehensive database of records (108 localities) and the most detailed map of the native range of EAB in East Asia to date. There are documented records for 87 mainland localities of EAB in the Russian Far East (Primorskiy, Khabarovskiy Kray), China (Heilongjiang, Jilin, Liaoning, Beijing, Hebei, Tianjin, Shandong and Xinjiang), and South Korea, and 21 localities in Japan. Records from Nei Mongol, Sichuan, Mongolia, and Taiwan are ambiguous since no documented records are available. The example of EAB shows that standard descriptions of pest ranges could include false or ambiguous data. Compilation of the database of documented localities is the only way to obtain reliable information on the range.

Key to Holarctic species of Epitrix flea beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) with review of their distribution, host plants and history of invasions

The genus Epitrix Foudras, 1860a has a worldwide distribution. Some species of Epitrix are major pests of potato, tomato, eggplant, tobacco and other plants in North America and Europe. Some pest species have been inadvertently introduced from North America to Europe, from Europe to North America and from both continents to some islands in Atlantic and Pacific oceans. Therefore, a key for the identification of all Holarctic species is necessary for plant quarantine and protection services. We have compiled the key for distinguishing Epitrix from genera that could be confused with it and a key for all Holarctic species of Epitrix with the figures of spermathecae and aedeagi and the checklist with a review of the geographical distribution, host plants and history of invasions. The following species are included: E. abeillei (Bauduer), E. allardii (Wollaston), E. atropae Foudras, E. brevis Schwarz, E. caucasica (Heikertinger), E. cucumeris (Harris), E. dieckmanni (Mohr), E. ermischi (Mohr), E. fasciata Blatchley, E. flavotestacea Horn, E. fuscula Crotch, E. hirtipennis (Melsheimer), E. humeralis Dury, E. intermedia Foudras, E. krali Döberl, E. lobata Crotch, E. muehlei Döberl, E. priesneri (Heikertinger), E. pubescens (Koch), E. ogloblini (Iablokov-Khnzorian), E. robusta Jacoby, E. setosella (Fairmaire), E. similaris Gentner, E. solani (Blatchley), E. subcrinita (LeConte), E. tuberis Gentner, E. warchalowskii (Mohr) and E. papa Orlova-Bienkowskaja.

Woolly ash aphid Prociphilus fraxinifolii (Riley, 1879) (Hemiptera: Eriosomatidae), a new invasive alien pest of ash trees (Fraxinus) in Russia.

Woolly ash aphid or ash leaf curl aphid Prociphilus fraxinifolii (Riley, 1879) is an alien invasive pest of ash trees native to North America. After its first record in Europe in 2003 in Hungary it has spread to the Ukraine, Serbia, Bulgaria, Great Britain, Spain, Poland and Germany. In 2016 P. fraxinifolii was firstly recorded at the southwestern border of Russia. Now Prociphilus fraxinifolii is firstly recorded in the center of European Russia, namely in Moscow Region, which is more than 700 km far from all other known localities of the species. In September 2017 five groups of ash trees (Fraxinus pennsylvanica) with colonies of Prociphilus fraxinifolii were found in Moscow Region. The example of Prociphilus fraxinifolii shows that alien pest insects can spread in Europe very quickly. Now Moscow region is the only regions of Europe, where the expanding range of Prociphilus fraxinifolii has overlapped with expanding ranges of other invasive alien species established in Europe in the last 20 years and severely damaging ash trees: ash dieback fungus Hymenoscyphus fraxineus Baral et al., 2014 (Ascomycota) and emerald ash borer Agrilus planipennis Fairmaire, 1888 (Coleoptera: Buprestidae).

Co-invasion of the ladybird Harmonia axyridis and its parasites Hesperomyces virescens fungus and Parasitylenchus bifurcatus nematode to the Caucasus

Study of parasites in recently established populations of invasive species can shed lite on sources of invasion and possible indirect interactions of the alien species with native ones. We studied parasites of the global invader Harmonia axyridis (Coleoptera: Coccinellidae) in the Caucasus. In 2012 the first established population of H. axyridis was recorded in the Caucasus in Sochi (south of European Russia, Black sea coast). By 2018 the ladybird has spread to the vast territory: Armenia, Georgia and south Russia: Adygea, Krasnodar territory, Stavropol territory, Dagestan, Kabardino-Balkaria and North Ossetia. Examination of 213 adults collected in Sochi in 2018 have shown that 53% of them are infested with Hesperomyces virescens fungi (Ascomycota: Laboulbeniales) and 8% with Parasitylenchus bifurcatus nematodes (Nematoda: Tylenchida, Allantonematidae). Examined H. axyridis specimens were free of parasitic mite Coccipolipus hippodamiae. An analysis of the phylogenetic relationships of Parasitylenchus bifurcatus based on 18S rDNA confirmed the morphological identification of this species. Hesperomyces virescens and Parasitylenchus bifurcatus are firstly recorded from the Caucasus and Russia, though widespread in Europe. It probably indicates that they appeared as a result of co-invasion with their host. Harmonia axyridis was released in the region for pest control, but laboratory cultures are always free of H. virescens and P. bifurcatus. Therefore, detection of H. virescens and P. bifurcatus indicates that population of H. axyridis in the Caucasus cannot derive exclusively from released specimens. We did not find H. virescens on 400 specimens of 31 other ladybird species collected in the same localities with H. axyridis in the Caucasus. No reliable correlation between infestation by H. virescens and P. bifurcatus has been found. Besides these two parasites an unidentified species of the order Mermithida is recorded. It is the first documented case of H. axyridis infestation by a parasitic nematode of this order in nature.