Tag: biocontrol

What is the Asian hornet invasion going to cost Europe?

Since its accidental introduction in 2003 in France, the yellow-legged Asian hornet Vespa velutina nigrithorax is rapidly spreading through Europe. In a new paper, published in the open-access journal Neobiota, French scientists try to estimate the costs of the invasion regarding the potential damage to apiculture and pollination services.

Since its accidental introduction in 2003 in France, the yellow-legged Asian hornet (Vespa velutina nigrithorax) is rapidly spreading through Europe. Both experts and citizen scientists keep on identifying the new invader spreading all over the Old Continent in the last decades.

In a recent study, French scientists led by Prof. Franck Courchamp at the Université Paris-Saclay and the CNRS, tried to evaluate the first estimated control costs for this invasion. Supported by the INVACOST project, their findings are published in the open-access journal Neobiota.

Since its invasion to France in 2004 when it was accidentally introduced from China, the Asian hornet has been spreading rapidly, colonising most of France at an approximate rate of 60-80 km per year, and also invading other European countries: Spain in 2010, Portugal and Belgium in 2011, Italy in 2012, Germany in 2014 and the UK in 2016. In the recent paper, published in the open-access journal Evolutionary Systematics, Dr. Martin Hussemann from CeNaK, University of Hamburg has recorded the northernmost capture of the Asian hornet in Hamburg in September 2019.

These data show that the Asian hornet is spreading all around Europe faster and faster with every year, even in climatically less favourable regions. The rapid invasion of the species is not necessarily caused by human-mediated dispersal, the species can rapidly spread on its own, but nevertheless, it is not uncommon.

Within its native and invasive range, V. velutina nigrithorax actively preys on honeybees, thus, causing harm to apiculture. Due to its active praying on wild insects, the Asian hornet also has a negative impact on ecosystems in general and contributes to the global decline of pollination services and honey production. Furthermore, by nesting in urban areas, the Asian hornet, which is well known for its aggressive behaviour, is a potential threat to human activities.

Currently, the control of the invasion is mainly undertaken by nest destruction and bait trapping, but none of these methods is sufficient enough to achieve complete eradication.

To proceed with the further control of the invasion, there is the need to evaluate economic costs. Those costs are divided into 3 main categories: (1) prevention of the invasion, (2) fighting the invasion and (3) damage caused by the invasion.

The cost of fighting the invasion of the Asian hornet is the cost of nest destruction. To identify those costs, the research team has studied information about the companies providing the services in the nest destruction, extrapolated the cost of nest destruction spatially and modelled the potential distribution of the invasive.

Estimated yearly cost of nest destruction if climatically suitable areas are fully invaded. Grey bars represent countries invasion hasn’t reached yet.
Credit: Prof. Franck Courchamp
License: CC-BY 4.0

As the calculations show, at the moment, the estimated yearly costs for eradication would be €11.9M for France, €9.0M for Italy and €8.6M for the United Kingdom.

“In 2006, only two years after the hornet was first observed in France, three departments were already invaded and the cost of nest destruction was estimated at €408k. Since then, the estimated yearly costs have been increasing by ~€450k each year, as the hornet keeps spreading and invades new departments. Overall, we estimated €23M as the cost of nest destruction between 2006 and 2015. If this temporal trend can be extrapolated for the next few years (i.e. if the hornet keeps spreading at a similar rate), we expect the yearly cost of nest destruction to reach an estimated value of €11.9M (given all suitable areas are invaded) in just 12 years,”
shares Prof. Franck Courchamp.

In Japan and South Korea, where the species has already been observed, the total yearly cost of nest destruction is estimated at €19.5M and €11.9M respectively.

So far, nests eradication is the most effective way to fight the invasion, though, it is not sufficient enough. As a result, so far, only 30-40% of the detected nests are destroyed each year in France. Moreover, rather than the result of a controlled strategy, those destroyed nests are only the ones that have been determined of particular potential harm to human or beekeeping activities. The researchers point out that this is not enough.

Estimated yearly cost of nest destruction in France since the start of the invasion given the yearly invasive range.
Credit: Prof. Franck Courchamp
License: CC-BY 4.0

In conclusion, the scientists call for more active measures and research, related to the invasion of V. velutina nigrithorax. Provided that other countries, including the USA, Australia, Turkey and Argentina appear to be climatically suitable for the species, they are also in danger (e.g., €26.9M for the USA).

The current study presents only the first estimates of the economic costs resulting from the Asian hornet, but definitely more actions need to be taken in order to handle harmful invasive species – one of the greatest threats to biodiversity and ecosystem functioning.

Consensus climate suitability of the yellow-legged hornet predicted from species distribution modelling.
Credit: Prof. Franck Courchamp
License: CC-BY 4.0

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Original source:

Barbet-Massin M, Salles J-M, Courchamp F (2020) The economic cost of control of the invasive yellow-legged Asian hornet. NeoBiota 55: 11-25. https://doi.org/10.3897/neobiota.55.38550

All microgastrinae wasps from around the world finally together in a 1,089-page monograph

With 3,000 known species and thousands more left to describe, the wasps of the subfamily Microgastrinae are the single most important group of parasitoids attacking the larvae of butterflies and moths, many of which are economically important pests. Consequently, these wasps have a significant impact on both the world’s economy and biodiversity.

Due to their affinities, these wasps are widely used in biological control programs to manage agricultural and forestry pests around the globe. Further, they have also been prominently featured in many basic and applied scientific research (e.g. chemical ecology, biodiversity studies, conservation biology, genomics, behavioural ecology). However, the information about Microgastrinae species is scattered across hundreds of papers, some of which are difficult to find. To make matters worse, there has never been an authoritative checklist of the group at a planetary scale.

All currently available information about the group is now brought together in a large monograph of 1,089 pages, published in the open-access, peer-reviewed journal ZooKeys. The publication presents a total of 2,999 valid extant species belonging to 82 genera. On top of that, the monograph features fossil species and genera, unavailable names and the institutions that store the primary types of all listed species.

Moreover, the researchers have included extensive colour illustrations of all genera and many species (thousands of images in 250 image plates); brief characterisation and diagnosis of all genera; detailed species distributions (within biogeographical regions and per individual country); synopsis of what is known on host-parasitoid associations; summary of available DNA barcodes; estimations of the group diversity at world and regional levels; as well as notes on individual species upon request.

“Compiling this annotated checklist was, more than anything, a labour of love,”
says Dr. Jose Fernandez-Triana of the Canadian National Collection of Insects, lead author of the paper.

Monograph paper openly published in *ZooKeys* at
https://doi.org/10.3897/zookeys.920.39128

“For the past six or seven years, we have spent thousands of hours pouring through hundreds of publications, reading original descriptions in old manuscripts, checking type specimens in many collections worldwide, exchanging information with colleagues from all continents. For the past year or so, I basically stopped all other ongoing research projects I was involved with, to focus solely (almost obsessively!) on finishing this manuscript. The work was often tedious and mind-numbing, and many times I had the temptation to delay the completion of the paper for a later time. However, I was lucky that the other co-authors were just as passionate as myself, and we all pushed each other to finish the task when energy ran low.”

Fifteen species of microgastrinae wasps showing the incredible diversity within the subfamily. Note the variety of colours and shapes.
Image by Dr. Jose Fernandez-Triana

“For the past few years, the Microgastrinae wasps have been one of the most intensively studied groups of insects, at least from a taxonomic perspective,” he adds. “Just to give you an idea: between 2014 and 2019 a total of 720 new species of Microgastrinae were described worldwide. That is an average of one new species every three days, sustained over a six-year period and showing no signs of slowing down.”

He also points out that many scientists from many different countries and biogeographical regions have been involved in the description of the new species. The paper recognises them all and their contributions in the Acknowledgements section.

“You could even say that we are witnessing a renaissance in the study of this group of wasps. However, even then, what has been done is only the tip of the iceberg, as we estimated that only 5 to 10% of all Microgastrinae species have been described. That means that we do not have a name, let alone detailed knowledge, for 90-95% of the remaining species out there. Perhaps, there could be up to 50,000 Microgastrinae wasp species worldwide. It is truly humbling when you consider the magnitude of the work that lies ahead.”

Yet, it is not only a matter of counting huge numbers of species. More importantly, many of those species either have already been put in use as biocontrol agents against a wide range of agricultural and forestry pests, or have the potential to be in the future.

For applied scientists, working with hyperdiverse and poorly known groups such as Microgastrinae is even more perplexing. Navigating the maze of old names, synonyms (species described more than one time under different names), homonyms (same names applied to different species), or unavailable names (names that do not conform to the rules of the International Commission of Zoological Nomenclature) is a daunting task. Often, that results in the same species being referred to in several different ways by different authors and academic works. Consequently, many historical references are full of misleading or even plainly wrong information. Meanwhile, it is very difficult to seek out the useful and correct information.

The present annotated checklist could work as a basic reference for anyone working with or interested in the parasitoid wasps of the subfamily Microgastrinae. In the future, the authors hope to produce revised editions, thus continuing to incorporate new information as it is generated, and to also correct possible mistakes.

“We welcome all kinds of criticisms and suggestions. And we hope that biocontrol practitioners will also help us, the taxonomists, to improve future versions of this work. However, for the time being, let me say that it is a tremendous relief to get this first version out!”
concludes Dr. Fernandez-Triana.

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Original source:

Fernandez-Triana J, Shaw MR, Boudreault C, Beaudin M, Broad GR (2020) Annotated and illustrated world checklist of Microgastrinae parasitoid wasps (Hymenoptera, Braconidae). ZooKeys 920: 1-1089. https://doi.org/10.3897/zookeys.920.39128.

Faster than a speeding bullet: Asian hornet invasion spreads to Northern Germany

Known to prey on many insects, including honey bees and other beneficiary species, the Asian hornet, which had recently invaded parts of Europe, presents a serious threat to apiculture and even to ecosystems. In their paper, published in the open-access journal Evolutionary Systematics, German scientists share concerns about this fast invader spreading to the north. In early September 2019, a single specimen was collected alive in Hamburg (Germany), representing the northernmost find of the species so far.

In early September 2019, an Asian hornet (Vespa velutina nigrithorax) was collected alive in Hamburg, Germany, representing the northernmost find of the species so far in Europe and indicating its further spread to the north. The paper by the research group from Hamburg, which also serves to update the occurrence of the dangerous invader, was published in the open-access journal Evolutionary Systematics.

Known to prey on many insects, including honey bees and other beneficiary species, the Asian hornet, which had already invaded parts of Southern and Central Europe, is a potential threat to apiculture and even to ecosystems.

The first specimen was captured in south-western France in 2005 and started to spread quickly. Over the next years, it invaded large parts of France and regions of Spain, Portugal, Belgium, Italy, the Netherlands, Great Britain and south-western parts of Germany. The estimated invasion speed for France has been estimated at around 78 km/year, but in reality, the species spread might be occurring much faster due to anthropogenic factors.

It’s not yet clear if the collected Asian hornet belonged to an already settled population or it’s rather the first record of a new invasion. Nevertheless, considering the fast invasion speed of the species and its relatively high climatic tolerance, it’s quite possible that it had reached Hamburg on natural routes and now reproduces there.

Even though other models suggest that the Hamburg area is not suitable for the species today, the new find might be a sign that the Asian hornet has begun spreading at a speed above that previously known and even in climatically less favourable regions.

“Therefore, the current find needs to be taken seriously, no matter if it is only a single specimen or a member of an established population”, shares the lead researcher Martin Husemann from Centrum für Naturkunde, University of Hamburg.

Invasive species are one of the great challenges in the modern world. Their occurrence can be considered as one of the key important ecological and evolutionary drivers.

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Original source:
Husemann M, Sterr A, Maack S, Abraham R (2020) The northernmost record of the Asian hornet Vespa velutina nigrithorax (Hymenoptera, Vespidae). Evolutionary Systematics 4(1): 1-4.
https://doi.org/10.3897/evolsyst.4.47358

Non-native pest-controlling wasp identified in Canada prior to formal approval

A samurai wasp (Trissolcus japonicus) lays an egg inside a brown marmorated stink bug (Halyomorpha halys) egg. The samurai wasp’s offspring will develop inside the pest’s egg and emerge as an adult wasp. Photo by Warren Wong.

Thought to be Canada’s most promising potential defense against the brown marmorated stink bug – a globally spreading agricultural pest native to Asia – the samurai wasp (another species from Asia and natural parasitoid of the former) has been considered for future release in the country in recent years.

However, prior to any formal decision and regulatory approval, the parasitoid, which is known to be specialized on stink bug eggs, was identified at a heavily infested site in Chilliwack, British Columbia, during a survey of the local enemies of the bug, conducted by a research team led by Dr. Paul Abram of Agriculture and Agri-Food Canada. Their findings are published in the open-access Journal of Hymenoptera Research.

Native to China, Japan, Taiwan and the Korean peninsula, the brown marmorated stink bug (Halyomorpha halys) has already established in areas of the United States and Europe and continues to spread. It is highly damaging to a wide range of vegetable and fruit crops, including peaches, apples, pears, soybeans, cherries, raspberries and pears. Curiously, those infested areas in both the USA and Europe also saw the arrival of the samurai wasp (Trissolcus japonicus) amid assessments whether releasing samurai wasps in the wild should be warranted.

“Classical (importation) biological control of invasive pests, where natural enemies are imported and intentionally introduced from a pest’s area of origin, involves years of research to assess risks and benefits of proposed introductions, followed by regulatory approval,” explain the researchers in their paper.

“However, there is increasing recognition that unintentional introductions of natural enemies are probably common, introducing a high level of uncertainty to the regulatory process for biological control introductions.”

In two consecutive years (2017 and 2018), the team of Dr Abram placed a total of 1,496 egg masses (41,351 eggs) of brown marmorated stink bugs at 16 field sites in coastal and interior British Columbia – already known to host large and well-established breeding populations of the species – in order to monitor and identify the local enemies of the pest. Later on, when the researchers retrieved the eggs and studied their parasitoids, they found three native wasp species, but their parasitism appeared largely unsuccessful.

*Female samurai wasp (Trissolcus japonicus) collected from Chilliwack, British Columbia. Photo by Elijah Talamas.*

According to the scientists, as well as previous studies conducted in both the USA and Europe, native wasps would often lay their eggs in those of the brown marmorated stink bug, but their larvae would rarely complete development. Even when they emerged, they were unlikely to produce their own offspring.

In one of the egg masses, however, the scientists noted that all eggs had been parasitized and, moreover, each produced a viable wasp. Later, the offspring would register a success of >90% in parasitizing brown marmorated stink bug eggs. Following these observations, the team identified these parasitoids as samurai wasps.

While the species is currently being redistributed within some US states on purpose, samurai wasp populations advancing to other localities suggest that much like its host, the parasitoid is also becoming a “global invader”. Therefore, it is quite possible that the samurai wasps in British Columbia have simply crossed a distance of >400 km from nearby Washington State, and the wasp is still at the early stages of its establishment in Canada.

“Nonetheless, the detection of this exotic biological control agent in Canada concurrently with regulatory review of its intentional importation and release is emblematic of the current uncertainty around regulatory control on the movement of biological control agents across borders,” comment the authors of the study.

Field surveys and extensive analyses are currently underway to track the establishment and biological control impact of the samurai wasp in Canada and also reveal how the species ended up in British Columbia.

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Original source:

Abram PK, Talamas EJ, Acheampong S, Mason PG, Gariepy TD (2019) First detection of the samurai wasp, Trissolcus japonicus (Ashmead) (Hymenoptera, Scelionidae), in Canada. Journal of Hymenoptera Research 68: 29-36. https://doi.org/10.3897/jhr.68.32203

Advanced computer technology & software turn species identification interactive

Important group of biocontrol wasps from Central Europe are used to demonstrate the perks and advantages of modern, free-to-use software

Representing a group of successful biocontrol agents for various pest fruit flies, a parasitic wasp genus remains largely overlooked. While its most recent identification key dates back to 1969, many new species have been added since then. As if to make matters worse, this group of visually identical species most likely contains many species yet to be described as new to science.

Having recently studied a species group of these wasps in Central Europe, scientists Fabian Klimmek and Hannes Baur of the Natural History Museum Bern, Switzerland, not only demonstrate the need for a knowledge update, but also showcase the advantages of modern taxonomic software able to analyse large amounts of descriptive and quantitative data.

Published in the open access Biodiversity Data Journal, the team’s taxonomic paper describes a new species – Pteromalus capito – and presents a discussion on the free-to-use Xper3, developed by the Laboratory of Informatics and Systematics of Pierre-and-Marie-Curie University. The software was used to create an openly available updated key for the species group Pteromalus albipennis.

The fully illustrated interactive database covers 27 species in the group and 18 related species, in addition to a complete diagnosis, a large set of body measurements and a total of 585 images, displaying most of the characteristic features for each species.

“Nowadays, advanced computer technology, measurement procedures and equipment allow more sophisticated ways to include quantitative characters, which greatly enhance the delimitation of cryptic species,” explain the scientists.

“Recently developed software for the creation of biological identification keys like Xper3, Lucid or Delta could have the potential to replace traditional paper-based keys.”

To put the statement into context, the authors give an example with one of the studied wasp species, whose identification would take 16 steps if the previously available identification key were used, whereas only 6 steps were needed with the interactive alternative.

One of the reasons tools like Xper3 are so fast and efficient is that the key’s author can list all descriptive characters in a specific order and give them different weight in species delimitation. Thus, whenever an entomologist tries to identify a wasp specimen, the software will first run a check against the descriptors at the top, so that it can exclude non-matching taxons and provide a list of the remaining names. Whenever multiple names remain, a check further down the list is performed, until there is a single one left, which ought to be the one corresponding to the specimen. At any point, the researcher can access the chronology, in order to check for any potential mismatches without interrupting the process.

Being the product of digitally available software, interactive identification keys are not only easy, quick and inexpensive to publish, but they are also simple to edit and build on in a collaborative manner. Experts from all around the world could update the key, as long as the author grants them specific user rights. However, regardless of how many times the database is updated, a permanent URL link will continue to provide access to the latest version at all times.

To future-proof their key and its underlying data, the scientists have deposited all raw data files, R-scripts, photographs, files listing and prepared specimens at the research data Zenodo, created by OpenAIRE and CERN.

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Original source:

Klimmek F, Baur H (2018) An interactive key to Central European species of the Pteromalus albipennis species group and other species of the genus (Hymenoptera: Chalcidoidea: Pteromalidae), with the description of a new species. Biodiversity Data Journal 6: e27722. https://doi.org/10.3897/BDJ.6.e27722

Journal of Hymenoptera Research links Crocodile Dundee, Toblerone, Game of Thrones & Alien

A myriad of species and genera new to science, including economically important wasps drawing immediate attention because of their amusing names and remarkable physical characters, in addition to work set to lay the foundations for future taxonomic and conservation research, together comprise the latest 64th issue of Journal of Hymenoptera Research (JHR).

The species *Qrocodiledundee outbackense*

Two genera (Qrocodiledundee and Tobleronius) named after the action comedy Crocodile Dundee and the chocolate brand Toblerone are only a couple of the 14 new genera from the monograph of the microgastrine wasps of the world’s tropical regions, authored by Dr Jose Fernandez-Triana and Caroline Boudreault of the Canadian National Collection of insects in Ottawa. In their article, the team also describes a total of 29 new species, where five of them carry the names of institutions holding some of the most outstanding wasp collections.

Another curiously named species of microgastrine wasp described in the new JHR issue, is called Eadya daenerys in reference to Daenerys Targaryen, a fictional character known from the best-selling book series A Song of Ice and Fire by George R. R. Martin, and the blockbuster TV show Game of Thrones. Discovered by University of Central Florida‘s Ryan Ridenbaugh, Erin Barbeau and Dr Barbara Sharanowski as a result of a collaboration between biocontrol researchers and taxonomists, the new species might not be in control of three dragons, nor a ruler or protector of whole nations. However, by being a potential biocontrol agent against a particular group of leaf beetle pests, it could spare the lives of many eucalyptus plantations around the world.

Furthermore, a wasp named Dolichogenidea xenomorph, which parasitises other eucalyptus pests, is also named after a character from a sought-after franchise. The scriptwriters of the horror sci-fi movie series Alien are thought to have been thinking of parasitic wasps when they came up with the character Xenomorph, remind authors Erinn Fagan-Jeffries, Dr Steven Cooper and Dr Andrew Austin. Additionally, the team from University of Adelaide and the South Australian Museum point out that the species name translates to ‘strange form’ in Greek, which perfectly suits the characteristic remarkably long ovipositor of the new wasp.

In another paper of the same journal issue, Dr. Jean-Luc Boevé, Royal Belgian Institute of Natural Sciences, Diego Domínguez, Universidad Técnica Particular de Loja, Ecuador, and Dr David Smith, Smithsonian’s National Museum of Natural History, USA, publish an illustrated list of the wasp-related sawflies, which they collected from northern Ecuador a few years ago. They also provide a checklist of the country’s species.

In conclusion, the fifth paper, authored by Serbian scientists Dr Milana Mitrovic Institute for Plant Protection and Environment, and Prof Zeljko

Tomanovic, University of Belgrade, studies ways to extract DNA from dry parasitoid wasps from the natural history archives decades after their preservation. In their work, they make it clear that such projects are of great importance for future taxonomic and conservation research, as well as agriculture.

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The open access Journal of Hymenoptera Research is published bimonthly by the scholarly publisher Pensoft on behalf of the International Society of Hymenopterists.

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Original sources:

Boeve; J, Dominguez D, Smith D (2018) Sawflies from northern Ecuador and a checklist for the country (Hymenoptera: Argidae, Orussidae, Pergidae, Tenthredinidae, Xiphydriidae). Journal of Hymenoptera Research 64: 1-24. https://doi.org/10.3897/jhr.64.24408

Ridenbaugh RD, Barbeau E, Sharanowski BJ (2018) Description of four new species of Eadya (Hymenoptera, Braconidae), parasitoids of the Eucalyptus Tortoise Beetle (Paropsis charybdis) and other Eucalyptus defoliating leaf beetles. Journal of Hymenoptera Research 64: 141-175. https://doi.org/10.3897/jhr.64.24282

Fagan-Jeffries EP, Cooper SJB, Austin AD (2018) Three new species of Dolichogenidea Viereck (Hymenoptera, Braconidae, Microgastrinae) from Australia with exceptionally long ovipositors. Journal of Hymenoptera Research 64: 177-190. https://doi.org/10.3897/jhr.64.25219

Mitrovic M, Tomanovic Z (2018) New internal primers targeting short fragments of the mitochondrial COI region for archival specimens from the subfamily Aphidiinae (Hymenoptera, Braconidae). Journal of Hymenoptera Research 64: 191-210. https://doi.org/10.3897/jhr.64.25399

Parasite vs. Invader: New endoparasitoid wasp can save the Dominican Republic economy

While biocontrol agents come in different shapes, often taking a lot of time for scientists to research, test and produce, natural ones always seem to be the better option. Now that Drs. Taveras and Hansson have discovered a new parasitoid wasp species in the Dominican Republic, they might have not only met the worst natural enemy for a widely spread invasive pest corrupting a large part of the essential pigeon pea crops. They are likely to have found a whole new field for investigation into the potential weapons against the eradicator of up to 76% of the essential crop. Their study is available in the open access Journal of Hymenoptera Research.

The find of this new gregarious endoparasitoid wasp, called Pediobius cajanus, is also the first time representative of this genus has been retrieved from the Asian fly. This invasive pest is estimated to destroy a huge part of the pigeon pea crops, a culture which is an essential food source for tropical America, a large part of the export and even has medicinal value. Therefore, the new species is very important not only in terms of the exploration of the biodiversity in Latin America, but also in economical sense.

Even when treated regularly with insecticides, the Asian fly manages to corrupt as much as 27% of the Dominican pigeon pea crops, a previous study shows. Although a precise figure of the damage cannot be provided due to large variations between localities and the impossibility of tracking the whole span of the pea cultivation areas, there is the case of the town of Rancho Los Vargas, Puerto Plata, where in 2012 the loss of the culture reached 76%.

On the other hand, the new wasp species was found to kill an average of 25% of the Asian fly larvae in the researched areas. In comparison, the previously known enemies of the pest are accountable for only 2%. This is why the scientists are now proposing the new species as a biocontrol agent.

In conclusion, the authors suggest that the parasitoid wasp is likely distributed across a much larger area. They believe that the new species could also be found over the entire island of Hispaniola, on neighbouring islands in the Caribbean and even in the tropical parts of the mainland in the Americas. Its record and distribution both call for a further investigation into the potential implementation of the wasp in controlling the Asian fly.

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Original source:

Taveras R, Hansson C (2015) Pediobius cajanus sp. n. (Hymenoptera, Eulophidae), an important natural enemy of the Asian fly (Melanagromyza obtusa (Malloch)) (Diptera, Agromyzidae) in the Dominican Republic. Journal of Hymenoptera Research 45: 41-54. doi: 10.3897/JHR.45.4964