A literature survey identified 403 primary research publications that investigated the ecological effects of invasive alien insects and/or the mechanisms underlying these effects. The majority of these studies were published in the last 8 years and nearly two-thirds were carried out in North America. These publications concerned 72 invasive insect species, of which two ant species, Solenopsis invicta and Linepithema humile, accounted for 18% and 14% of the studies, respectively. Most publications investigated effects on native biodiversity at population or community level. Genetic effects and, to a lesser extent, effects on ecosystem services and processes were rarely explored. We review the effects caused by different insect invaders according to: their ecosystem roles, i.e. herbivores, predators, parasites, parasitoids and pollinators; the level of biological organisation at which they occur; and the direct and indirect mechanisms underlying these effects. The best documented effects occur in invasive ants, Eurasian forest herbivores invasive in North America, and honeybees. Impacts may occur through simple trophic interactions such as herbivory, predation or parasitism. Alien species may also affect native species and communities through more complex mechanisms such as competition for resources, disease transmission, apparent competition, or pollination disruption, among others. Finally, some invasive insects, particularly forest herbivores and ants, are known to affect ecosystem processes through cascading effects. We identify biases and gaps in our knowledge of ecological effects of invasive insects and suggest further opportunities for research.

Ecological effects of invasive alien insects

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.

/pdf/the-harlequin-ladybird-harmonia-axyridis-global-perspectives-5esz3dhkgy.pdf

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

Omnivory is a frequent feeding strategy in terrestrial arthropods, occurring across a diversity of taxa occupying a wide array of habitats. Because omnivory has important consequences for broad areas of theoretical and applied ecology, it is essential to understand those factors that favor its occurrence. Here we address the limiting role of nitrogen in promoting omnivory, not so much from the historical perspective of herbivores supplementing their nutrient-poor plant diet, but by extending the argument to higher trophic levels where predators feed on each other as well as herbivores. Drawing on the historically documented mismatch in nitrogen stoichiometry between herbivores and their host plants (C:Nplants ≫ C:Nherbivores), and a recently documented, though smaller, difference in nitrogen content between predators and their herbivore prey (C:Nherbivores > C:Npredators), we discuss the existence of a trade-off between nutrient quality and quantity that occurs across trophic levels. The existence of this...

Might nitrogen limitation promote omnivory among carnivorous arthropods

(1985). Analysis of Messy Data, Volume I: Designed Experiments. Technometrics: Vol. 27, No. 4, pp. 440-440.

Analysis of Messy Data, Volume I: Designed Experiments

https://www.zin.ru/animalia/coleoptera/addpages/Andrey_Ukrainsky_Library/References_files/Obrycki09-proof.pdf

Aphidophagy by Coccinellidae: Application of biological control in agroecosystems

Coccinella septempunctata L. (Coleoptera: Coccinellidae), a Palearctic coccinellid, has established and rapidly spread throughout the United States. This quantitative examination of larval interactions between C. septempunctata and Coleomegilla maculata (DeGeer) (Coleoptera: Coccinellidae), a Nearctic coccinellid, was conducted under controlled prey densities. Larvae of both coccinellid species are affected by a limited diet [one pea aphid per day Acyrthosiphon pisum (Harris) (Homoptera: Aphididae)] compared with an excess diet (>20 aphids per day). Larval survival decreased from 86 to 63% in C. maculata and from 84 to 33% in C. septempunctata; mean preimaginal developmental time increased from 20.6 to 26.7 days in C. maculata, and from 18.1 to 32.0 days in C. septempunctata. Additionally, on one aphid per day, mean adult weight was reduced from 12.39 to 9.79 mg in C. maculata, and from 39.57 to 14.44 mg in C. septempunctata. Interspecific interactions, favoring C. septempunctata over C. maculata at a␣low prey density (one aphid per day), take the form of␣reduced survival of C. maculata compared with C.␣septempunctata (14 versus 66%). Reduced survival of␣C. maculata may be the result of competition for aphids or intraguild predation by C. septempunctata on C.␣maculata. No interspecific interactions (measured as effects on larval survival, preimaginal developmental time, and adult weight) were observed between larvae of these two species at a high prey density (>20 aphids per predator per day).

Interactions between an introduced and indigenous coccinellid species at different prey densities.

This experimental field study examined the interactions between larvae of Coccinella septempunctata L., an introduced Palearctic species, and Coleomegilla maculata (Degeer), a Nearctic species. The interactions were examined in 5.8-m3 field cages enclosing potatoes infested with Myzus persicae (Sulzer) (Homoptera: Aphididae). Intra- and interspecific interactions were compared by using 4 measurements: developmental time and survival from 1st instar to adult, adult weight for each coccinellid species, and M. persicae density. No significant differences were found between intra- and interspecific treatments in C. maculata survival: 19.6 and 18.3%, in 1992 and 5.7 and 3.3%, in 1993, respectively; or developmental time: 24.6 and 25.4 d in 1992 and 20.4 and 20.6 d in 1993, respectively. In 1992, no significant differences were observed between the intra- and interspecific treatments in C. septempunctata survival (38.3 and 39.2%) or developmental time (20.0 and 20.5 d). In 1993, no C. septempunctata survived to adults at the relatively lower prey densities. Interspecific interactions between C. septempunctata and C. maculata larvae did not significantly affect M. persicae density, compared with densities in intraspecific cages.

Experimental Assessment of Interactions Between Larval Coleomegilla maculata and Coccinella septempunctata (Coleoptera: Coccinellidae) in Field Cages

Fourteen parasitoid species were recovered from strawberry leafroIler, Ancylis comptana (Froelich), larvae and pupae collected in central Iowa strawberry fields in 1987 and 1989. Parasitism by the two most abundant parasitoid species, Itoplectis conquisitor (Say) (Hymenoptera: Ichneumonidae) and Pediobius facialis (Giraud) (Hymenoptera: Eulophidae), was synchronous with high A. comptana pupal densities. This is the first study in North America in which I. conquisitor and P. facialis were the predominant parasitoids attacking immature A. comptana . In 1987, cumulative percentage parasitism of the first A. comptana generation increased to 7% on 24 June. In 1989, the cumulative percentage parasitism of the first larval generation increased to 5.5% on 23 June. Because strawberries have a relatively high tolerance for A. comptana leaf rolling and feeding activity, naturally occurring parasitoids should be conserved.

Larval and Pupal Parasitism of the Strawberry Leafroller (Lepidoptera: Tortricidae)

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Interactions between an introduced and indigenous coccinellid species at different prey densities.

Experimental Assessment of Interactions Between Larval Coleomegilla maculata and Coccinella septempunctata (Coleoptera: Coccinellidae) in Field Cages

Larval and Pupal Parasitism of the Strawberry Leafroller (Lepidoptera: Tortricidae)