On the quantitative criteria of subspecies in insects. Case study of Entomoscelis adonidis (Coleoptera: Chrysomelidae) in European Russia and the Caucasus.

Many insect subspecies are described without a statistical assessment of differences. In this work, an attempt is made to statistically process the morphological differences of some recently described subspecies. In this regard, a taxonomic review of Entomoscelis adonidis (Pallas, 1771) (Coleoptera, Chrysomelidae) and related taxa described from European Russia and the Caucasus is presented. Statistical processing of numerous materials (782 specimens) from type localities of the nominal taxa adonidis, spuria Jacobson, 1896, caucasica Kippenberg, 2020, and bashkiriae Kippenberg, 2020, and their environs showed the presence of geographical variability in several characters, which, however, does not reach the level of species or subspecies. For subspecies, the classic "75 %" rule is applied. Therefore, all these specimens are considered to belong to the same species and subspecies. For nomenclature stability and to clarify the taxonomic position of closely related taxa, the neotype of Chrysomela adonidis Pallas is designated from the type locality; the neotype is in accordance with the currently used interpretation of E. adonidis adonidis. Besides that, the lectotype of Entomoscelis adonidis var. spuria Jacobson is designated. The paratypes of E. adonidis caucasica Kippenberg and E. dorsalis bashkiriae Kippenberg were studied. The following new synonyms are proposed: E. adonidis adonidis (Pallas, 1771) = E. adonidis caucasica Kippenberg, 2020 (syn. nov.) = E. dorsalis bashkiriae Kippenberg, 2020 (syn. nov.). The status of E. adonidis var. spuria Jacobson, 1896 as a junior synonym of E. adonidis adonidis is confirmed. The endophallus, particularly, flagellum of E. adonidis was studied for the first time. Several Brassicaceae species, namely Berteroa incana, Barbarea vulgaris, Sisymbrium officinale, Capsella bursa-pastoris, and Thlaspi arvense, as well as Adonis vernalis (Ranunculaceae) are confirmed as host plants of the adults of E. adonidis in cages.

Low Heat Availability Could Limit the Potential Spread of the Emerald Ash Borer to Northern Europe (Prognosis Based on Growing Degree Days per Year).

Emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae) is one of 20 priority quarantine pests of the European Union. It is native to Asia and is established in the USA, Canada, European Russia, and Ukraine. We made the first prognosis of the potential range of A. planipennis in Europe based on heat availability. Mean annual growing degree days base 10 °C (AGDD10) was calculated for each grid square (0.25° × 0.25° latitude x longitude degrees) on the Earth's surface. Minimal AGDD10 recorded in the grid squares currently occupied by A. planipennis was 714° in Asia, 705° in North America, and 711° in European Russia. Agrilus planipennis has never been recorded in localities with AGDD10 below 700°. If the phenotypic plasticity would not allow this species to overcome this threshold, cold regions of Europe would probably not be invaded by A. planipennis. Thus, Fraxinus excelsior could potentially escape from A. planipennis in some regions of Norway, Sweden, Finland, Ireland, and Great Britain.

History of the Biodiversity of Ladybirds (Coccinellidae) at the Black Sea Coast of the Russian Caucasus in the Last 120 Years-Does the Landscape Transformation and Establishment of Harmonia axyridis Have an Impact?

Studies of the history of regional insect fauna are important for understanding the changes in ecosystems. We analyzed the dynamics of ladybird fauna at the main sea resort of Russia over a period of 120 years to determine the following: (1) what species disappeared and what species appeared during landscape transformation; (2) what alien species introduced for pest control have been observed to date; and (3) whether the establishment of the global invader Harmonia axyridis (Coccinellidae) caused the extinction of some ladybird species in the Caucasus. We examined specimens collected by us and 54 other collectors including specimens from old museum collections and detected 62 species, 50 of which were collected in recent years (2011-2020). Landscape transformation and recreational use have caused not a decrease but an increase in ladybird biodiversity. Twenty-nine of 34 species recorded before 1930 have been observed in the region to date. Twenty-three other species have spontaneously spread to the region between 1930 and 2020 because of the creation of suitable anthropogenic habitats or because of unintentional introduction. Rodolia cardinalis, Cryptolaemus montrouzieri, Lindorus lophanthae, and Serangium montazerii were released for pest control, and they occur in gardens and streets but not in natural habitats. Harmonia axyridis, which appeared approximately 10 years ago, is abundant in urban and natural habitats, but there is no evidence that it caused the elimination of any ladybird species.

Invasive Agricultural Pest Drosophila suzukii (Diptera, Drosophilidae) Appeared in the Russian Caucasus.

Spotted-wing drosophila Drosophila suzukii (Matsumura) is one of the most important invasive pests of fruit and wine production worldwide. This species feeds on Prunus spp., Rubus spp., Fragaria spp. (Rosaceae), Vaccinium spp. (Ericaceae), Vitis spp. (Vitaceae), and other soft fruits. It causes significant damage because, unlike most other Drosophila species, it oviposits and feeds on healthy fruits. Drosophila suzukii is a quarantine pest that is included on the European and Mediterranean Plant Protection Organization (EPPO) A2 List. This species is native to East Asia and has been rapidly spreading through Europe since 2008. Herein, we report the first records of D. suzukii in European Russia. In 2017 and 2020, we placed baited traps in different districts of the resort city of Sochi (Black sea Coast of the Caucasus, Krasnodar Territory, Russia). Three specimens of D. suzukii were collected in June 2017, two specimens in September 2017, and 44 specimens in September 2020. Specimens were identified by the typical female ovipositor, spotted wings of the males, and other morphological characters. Krasnodar Territory is one of the main fruit-producing regions of Russia. Therefore, populations of this pest should be monitored and measures to minimize economic damage should be taken.

Predicting the Invasion Potential of the Lily Leaf Beetle, Lilioceris lilii Scopoli (Coleoptera: Chrysomelidae), in North America.

Invasive species are among the leading threats to global ecosystems due to impacts on native flora and fauna through competition and predation. The lily leaf beetle, Lilioceris lilii Scopoli (Coleoptera: Chrysomelidae), is an invasive pest of lilies (Lilium spp.) and other genera of Liliaceae (Liliales). A habitat suitability model was created using Maxent, to help predict if L. lilii will be able to establish in locations were native North American Liliaceae species grow. The model was created using georeferenced occurrence records from the beetle's native, naturalized, and invasive range. Model results indicate that precipitation in the driest quarter and annual average temperatures are most strongly correlated with L. lilii distribution, and suggest that the species will perform poorly in very dry, hot, or cold environments. The model also indicates that the beetle should be able to establish throughout the range of most North American Liliaceae genera, including species of special conservation concern. This model can be used by natural area managers to identify areas of high habitat suitability that overlap with vulnerable North American Liliaceae species, and prioritize L. lilii monitoring and control activities as the beetle continues to expand its range.

Current Distribution and Diagnostic Features of Two Potentially Invasive Asian Buprestid Species: Agrilus mali Matsumura and A. fleischeri Obenberger (Coleoptera: Buprestidae).

Our goal is to analyze the known geographical ranges and diagnostic features of two potentially invasive Asian buprestid species: the quarantine apple tree pest, Agrilus mali Matsumura, and the poplar pest A. fleischeri Obenberger. Based on the examination of museum collections and literature sources, we compiled comprehensive databases of records of the exact collecting sites for both species and generated detailed maps of their ranges. There are 51 documented localities for A. mali in the Russian Far East and East Siberia, Mongolia, China, and the Korean peninsula, and there are 53 documented localities for A. fleischeri in the Far East and Siberia, Kazakhstan, Mongolia, China, and Japan. No evidence of the presence of A. mali in Japan was found. Outbreak sites of A. mali in Xinjiang in the 2000s most likely represent the newly forming invasion areas; their proximity to the wild apple stands in the Kazakh part of the Tien Shan is a direct threat to Kazakhstan and adjacent countries. Sites damaged by A. fleischeri in Liaoning are situated within its native range; the outbreaks were likely triggered by the switch from indigenous to introduced poplars. This situation is similar to the early stages of emerald ash borer (Agrilus planipennis Fairmaire) invasion.

Rigorous Morphological Studies Confirm That the Classical Object of Pest Control Chilocorus kuwanae Is the Same Species as Ch. renipustulatus (Coleoptera: Coccinellidae).

The ladybug Chilocorus kuwanae, which was described in Japan, has been used for biological control of pests for 100 years. Chilocorus kuwanae was recently synonymized with Ch. renipustulatus described in Europe. The synonymy was based on the examination of few specimens. Our aim is to verify this synonymy. We studied all characters previously used to distinguish these taxa: eight metric and nine qualitative characters. Examination of 107 specimens from Japan and Sakhalin and 174 specimens from Europe showed that the ranges of variability in all characters in Asian and European specimens strongly overlap. There are no characters with interspecific hiatuses. Analysis with Amadon's criteria showed that Asian and European specimens also do not represent different subspecies. Conclusions: (1) No differences between the specimens from Asia (Japan and Sakhalin) and Europe were found at specific or subspecific levels. Chilocorus kuwanae is a junior synonym of Ch. renipustulatus. (2) The releases of "Chilocorus kuwanae" in Europe and the Caucasus did not represent classical biological control since the same species was native to these regions. (3) A thorough taxonomical revision with the study of morphological variability should be conducted before the introduction of any species to new regions. (4) Taxonomical conclusions based on morphological studies should be confirmed by statistical methods.

Minimum Winter Temperature as a Limiting Factor of the Potential Spread of Agrilus planipennis, an Alien Pest of Ash Trees, in Europe.

The emerald ash borer, EAB (Agrilus planipennis) is a devastating alien pest of ash trees. It is spreading in European Russia and Ukraine and will appear in other European countries. Our aim was to determine the regions of Europe where the winter temperature drops low enough to prevent A. planipennis establishment. We calculated the minimum daily air temperature from 2003-2019 for each grid square (0.5° × 0.5°) in East Asia, North America and Europe and determined the minimum daily temperature in the grid squares where A. planipennis was recorded. Temperatures of -30 to -33 °C occur in the northern portions of the pest range on all continents. No established population has been recorded in localities where temperatures below -34 °C occur. This temperature is close to the absolute supercooling point of A. planipennis larva (-35.3 °C). It is unlikely that low temperatures could prevent the spread of A. planipennis in northern Western Europe (Sweden, Norway, Finland, etc.), since the temperature in this area did not fall to -34 °C from 2003-2019. However, such temperatures are not rare in eastern European Russia (Kostroma, Vologda, Orenburg regions, etc.), where Fraxinus pennsylvanica and F. excelsior occur. These regions could potentially become refuges for these ash species.

Record of the Emerald Ash Borer (Agrilus planipennis) in Ukraine is Confirmed.

Agrilus planipennis (Coleoptera: Buprestidae) is a devastating invasive pest of ash trees. This wood-boring insect is native to Asia and established in European Russia about 20 years ago. It severely damages Fraxinus pennsylvanica plantations and quickly spreads. In 2019 we first detected A. planipennis in Ukraine. More than 20 larvae were collected from under the bark of F. pennsylvanica trees on 5 September 2019 in the Markivka District of the Luhansk Region. The coordinates of the localities of collection were 49.614991 N, 39.559743 E; 49.614160 N, 39.572402 E; and 49.597043 N, 39.561811 E. The photos of the damaged trees with larval galleries, exit holes and larvae are presented. It indicates that A. planipennis is established in the east of Ukraine. This fact is important for development of quarantine protocols to prevent or at least slow the further spread of this invasive pest in Europe.

Modeling long-distance dispersal of emerald ash borer in European Russia and prognosis of spread of this pest to neighboring countries within next 5 years.

AIM: To develop an approach to model the spatial dynamics of emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae) in European Russia. This tree-killing pest was detected in Moscow 15 years ago and began to spread, posing a threat to ashes all over Europe. The aim was to determine its probable current range and to evaluate the probability of its dispersal to neighboring countries within the next 5 years. LOCATION: Cities and transport hubs of European Russia and neighboring countries. Ash trees in this region occur mainly in urban plantations and along highways. METHODS: Pairwise distances between all locations were used as the main parameter determining the probability of pest spread. For each location, the probability of detection of A. planipennis was calculated using three simulation recurrent models of long-distance dispersal. Parametrization was made by comparison with results of surveys in 2003-2015. Field data on the range of A. planipennis in 2016-2017 were mapped and used for model verification. A prognosis of spread of the pest by 2022 was made. RESULTS: A model based on fat-tailed kernel corresponds to both negative and positive results of surveys. According to the model, the current range is likely to be restricted to Russia, but probability of detection of the pest in the east of Belarus, Ukraine, Estonia, Latvia, and Lithuania by 2022 is 15%-40%. MAIN CONCLUSIONS: The forestry services of neighboring countries probably have about 5 years to prepare for the invasion of this pest, but regular surveys are necessary, since the pest can appear at any time. The case considered shows that the simple approach based on a fat-tailed kernel and just one parameter-pairwise distance between cities-can be used for modeling long-distance dispersal of alien pests of urban plantations.

Coinvasion by the ladybird Harmonia axyridis (Coleoptera: Coccinellidae) and its parasites, Hesperomyces virescens (Ascomycota: Laboulbeniales) and Parasitylenchus bifurcatus (Nematoda: Tylenchida, Allantonematidae), in the Caucasus.

The study of parasites in recently established populations of invasive species can shed light on the sources of invasion and possible indirect interactions between the alien species and native ones. We studied parasites of the global invader Harmonia axyridis (Coleoptera: Coccinellidae) in the Caucasus. In 2012, the first established population of Ha. axyridis was recorded in the Caucasus in Sochi (south of European Russia, Black Sea coast). By 2018, the ladybird had spread to a vast area: Armenia, Georgia and south Russia (Adygea, the Krasnodar territory, the Stavropol territory, Dagestan, Kabardino-Balkaria and North Ossetia). The examination of 213 adults collected in Sochi in 2018 showed that 53% were infested with Hesperomyces virescens fungi (Ascomycota: Laboulbeniales) and that 8% were infested with Parasitylenchus bifurcatus nematodes (Nematoda: Tylenchida, Allantonematidae). The examined Ha. axyridis specimens were free of the parasitic mite Coccipolipus hippodamiae. An analysis of the phylogenetic relationships of P. bifurcatus based on 18S rDNA confirmed the morphological identification of this species. Hesperomyces virescens and P. bifurcatus were first recorded in the Caucasus and Russia, although they are rather widespread in Europe. This likely indicates that they appeared as a result of coinvasion with their host because the populations of Ha. axyridis, He. virescens and P. bifurcatus in the Caucasus are isolated from the main parts of the ranges of these species in Europe. The nearest localities of Ha. axyridis is on another shore of the Black Sea, and the nearest localities of He. virescens and P. bifurcatus are more than 1000 km from the Caucasus. It is impossible to determine whether the first founders of the Caucasian population were infested with the parasites or whether the parasites were introduced by specimens of Ha. axyridis that arrived later from Europe. Harmonia axyridis was released in the region for pest control, but laboratory cultures are always free of He. virescens and P. bifurcatus. Therefore, the detection of He. virescens and P. bifurcatus indicates that the population of Ha. axyridis in the Caucasus could not have derived exclusively from released specimens. We did not find He. virescens on 400 specimens of 29 other ladybird species collected from the same localities as Ha. axyridis in the Caucasus. No reliable correlation between infestation by He. virescens and that by P. bifurcatus has been found. In addition to these two parasites, an unidentified species of the order Mermithida was recorded. This is the first documented case of Ha. axyridis infestation by a parasitic nematode of this order in nature.

Alien leaf beetles (Coleoptera, Chrysomelidae) of European Russia and some general tendencies of leaf beetle invasions.

Invasions of leaf beetles can cause tremendous economic consequences because some of these insects become major pests in invaded territories. We present the first inventory of alien Chrysomelidae of European Russia that appeared in the region in the 20th and 21st centuries (9 species) with analysis of the history of their invasions and detailed maps of distribution. This case study revealed some general tendencies of invasions of leaf beetles: (1) Recently, a dramatic increase in the rate of Chrysomelidae invasions is observed, which reflects the increase in international trade of living plants; (2) Alien leaf beetles can spread quickly, occupying almost all of Europe within several decades; (3) When the range of some leaf beetle species is quickly expanding, or when the species has been recorded established somewhere outside the native range, this species should be regarded as a potential invader worldwide. and (4) Alien leaf beetles usually occur on alien or cultivated plants, but some become naturalized in native communities. The specific information was the following. Two species native to the Mediterranean region, Chrysolina americana (feeds on Rosmarinus and Lavandula) and Leptomona erythrocephala (feeds on Lotus corniculatus) were recorded in European Russia for the first time. A polyphagous pest of floriculture Luperomorpha xanthodera native to China and Korea and a pest of soybeans Medythia nigrobilineata native to east Asia have been in the region since 2016. A pest of tobacco Epitrix hirtipennis native to North America has occurred since 2011. A pest of corn Diabrotica virgifera was intercepted at the border of Russia in 2011 but has not established. Three alien species have been in the region since the 20th century: Zygogramma suturalis introduced from North America for control of Ambrosia, Phyllotreta reitteri native to Afghanistan and Tajikistan and feeding on Lepidium latifolium, and the Colorado potato beetle Leptinotarsa decemlineata.

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.

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.