Euwallacea fornicatus | |
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Adult female of E. fornicatus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Coleoptera |
Family: | Curculionidae |
Genus: | Euwallacea |
Species: | E. fornicatus |
Binomial name | |
Euwallacea fornicatus (Eichhoff, 1868) | |
Species in the complex | |
Tea shot hole borer clade a Contents |
Euwallacea fornicatus, also known as tea shot-hole borer, or polyphagous shot-hole borer (PSHB) is a species complex consisting of multiple cryptic species of ambrosia beetles known as an invasive species in California, Israel, South Africa, and Australia. The species has also been unintentionally introduced into exotic greenhouses in several European countries.
See Wallacea, region of Indonesian islands named after the naturalist Alfred Russel Wallace.
Common names of the species include tea shot-hole borer [1] and polyphagous shot-hole borer (PSHB). [2]
The tea shot hole borer has been known to cause devastating damage to tea ( Camellia sinensis ) in at least ten countries, [1] including India and Sri Lanka where it is a major economic pest. [3]
Since 2007, specimens of E. fornicatus have been documented in Florida on avocado trees. However, it is not considered a health threat because no disease is expressed. [4]
In 2009, specimens matching the description for E. fornicatus were introduced into Israel, where they were documented as vectors of a new fungal plant pathogen in avocado trees. In 2012, similar fungal disease was recorded in avocado trees in California (CA). [5]
The species has also been unintentionally introduced into exotic greenhouses in several European countries. [6]
It was found in South Africa some years before 2022. [7]
Polyphagous shot-hole borer was detected in Perth, Western Australia, in 2021. [2]
Adult females are range between 1.9 and 2.5 mm long. They are bulky, dark brown or black and the frontal edge of the pronotum has a row of saw-like projections. [1] Moreover, specimens have erect setae organized in rows in the elytral declivity with a costa in the posterolateral edge. As many other ambrosia beetles, males are significantly smaller, with non-functional wings.
Larvae are similar to all other larvae in the family Curculionidae; legless, c-shaped and a sclerotized head capsule.
Euwallacea fornicatus is a species complex consisting of multiple cryptic species of ambrosia beetles (Coleoptera: Curculionidae: Scolytinae: Xyleborini), known as an invasive species in California, Israel and South Africa. [8]
Even though several taxa were historically synonymized under the name E. fornicatus due to morphological similarity, some of these taxa differ in terms of economic severity and host preferences. As such, Euwallacea fornicatus is typically considered a species complex, with several clades that occur in separate regions of Southeast Asia and develop in different hosts. [9] [10] Early phylogenetic work using the DNA mitochondrial gene COI suggested that there were three major clades classified as E. fornicatus, supporting the idea of at least three different species with phylogeographic boundaries within the species complex. [11] Later work (in 2018) resolved that there were actually four major lineages, one containing "true" E. fornicatus, another bearing the resurrected name Euwallacea fornicatior , another bearing the resurrected name Euwallacea whitfordiodendrus , and another that had not been named previously, and newly described as Euwallacea kuroshio . [12]
The first two clades, both commonly called the tea shot hole borer, are E. fornicatus and E. fornicatior; these are originally from southern Southeast Asia and introduced into Hawaii and Florida. The third clade is thought to originate from a more northern range in Southeast Asia and to have since been introduced into Los Angeles, California, Israel and South Africa. [13] This clade has been given the common name polyphagous shot hole borer in reference to the very broad host range, and contains E. whitfordiodendrus. It has been severely affecting avocado trees in association with several Fusarium species. The fourth clade, the Kuroshio shot hole borer, is believed to have originated in the Pacific Islands and has since been introduced into San Diego county, California, and contains E. kuroshio. [12] [14] Its impact has impacted many trees in San Diego County, as far south as the Tijuana River Valley Regional Park. [15]
A fundamental problem remains that the morphological variation within the species of each clade is great enough that there is no character or even combination of characters that can be uniquely used for reliable visual identification of the different taxa; however, their DNA sequences differ by from 11-15%, and appear to be far more reliable for identification. [12] Given that past research (until 2018) has generally assumed a single species was being studied, it is hoped that DNA analyses can now be applied to specimens from earlier studies to identify, post facto, which actual species were being examined, so as to better understand the biology of the different lineages. [12]
These beetles are able to reproduce in a number of tree species, including box elder, avocado, castor bean, English oak, coast live oak, silk tree, sweetgum, coral tree, Tītoki, California sycamore, Blue Palo Verde and big leaf maple. [16]
As the rest of the ambrosia beetles, E. fornicatus larvae and adults do not consume the wood itself, but instead feed on a symbiotic fungus ( Fusarium euwallaceae ) carried in a specific structure called mycangium. [8] In E. fornicatus, the mycangium is located in the mandible. The combination of massive numbers of beetles with the symbiotic fungus kills trees, even though the fungus alone is a weak pathogen. [9]
Euwallacea fornicatus breeds in various live hosts and is considered a severe pest of several economically important plants, such as: tea ( Camellia sinensis ), avocado ( Persea americana ), citrus ( Citrus spp.), and cacao ( Theobroma cacao ). [5] The beetle damages the tree by tunnelling deeply, and cultivating fungus which blocks the vascular system of the tree, thus depriving it of the ability to transport water and nutrients to its branches and leaves. [10]
The most recommended management strategies include sanitation of infected hosts and avoiding the spread of infected material. Chemical control can be considered in hosts which are not part of human consumption and some attempts of biological control have been made with little success. Moreover, resistant or tolerant varieties are considered an important aspect of the integrated pest management for this pest. [17]
After its recent introduction to South Africa, a 2022 Stellenbosch University study estimated that the potential economic harm [10] of the pest to be around 18.45 billion international dollars (A$28 billion [10] ), or about 0.66% of the South Africa's GDP. [7]
The pest was first detected in Perth, Australia, in 2021, in two box elder maple trees in East Fremantle. By 2024 the beetle was entrenched in metropolitan parks in Perth such as Hyde Park and Victoria Park, and as of March 2024 [update] scientists are working hard to eradicate it and prevent its movement to the eastern seaboard of the continent. The only way to kill an infestation of the pest is to remove and grind up the affected wood into woodchips, which in most cases requires cutting down entire mature trees. Several large Moreton Bay fig trees had to be cut down, and a 300-year-old paperbark tree was marked for removal, among others. A quarantine zone has been declared, and people are not allowed to take any plant material out of this zone. [10] [18] [19]
Ambrosia beetles are beetles of the weevil subfamilies Scolytinae and Platypodinae, which live in nutritional symbiosis with ambrosia fungi. The beetles excavate tunnels in dead or stressed trees into which they introduce fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases its fungal symbiont. The fungus penetrates the plant's xylem tissue, extracts nutrients from it, and concentrates the nutrients on and near the surface of the beetle gallery. Ambrosia fungi are typically poor wood degraders, and instead utilize less demanding nutrients. Symbiotic fungi produce and detoxify ethanol, which is an attractant for ambrosia beetles and likely prevents growth of antagonistic pathogens and selects for other beneficial symbionts. The majority of ambrosia beetles colonize xylem of recently dead trees, but some colonize stressed trees that are still alive, and a few species attack healthy trees. Species differ in their preference for different parts of trees, different stages of deterioration, and in the shape of their tunnels ("galleries"). However, the majority of ambrosia beetles are not specialized to any taxonomic group of hosts, unlike most phytophagous organisms including the closely related bark beetles. One species of ambrosia beetle, Austroplatypus incompertus exhibits eusociality, one of the few organisms outside of Hymenoptera and Isoptera to do so.
A bark beetle is the common name for the subfamily of beetles Scolytinae. Previously, this was considered a distinct family (Scolytidae), but is now understood to be a specialized clade of the "true weevil" family (Curculionidae). Although the term "bark beetle" refers to the fact that many species feed in the inner bark (phloem) layer of trees, the subfamily also has many species with other lifestyles, including some that bore into wood, feed in fruit and seeds, or tunnel into herbaceous plants. Well-known species are members of the type genus Scolytus, namely the European elm bark beetle S. multistriatus and the large elm bark beetle S. scolytus, which like the American elm bark beetle Hylurgopinus rufipes, transmit Dutch elm disease fungi (Ophiostoma). The mountain pine beetle Dendroctonus ponderosae, southern pine beetle Dendroctonus frontalis, and their near relatives are major pests of conifer forests in North America. A similarly aggressive species in Europe is the spruce ips Ips typographus. A tiny bark beetle, the coffee berry borer, Hypothenemus hampei is a major pest on coffee plantations around the world.
The term mycangium is used in biology for special structures on the body of an animal that are adapted for the transport of symbiotic fungi. This is seen in many xylophagous insects, which apparently derive much of their nutrition from the digestion of various fungi that are growing amidst the wood fibers. In some cases, as in ambrosia beetles, the fungi are the sole food, and the excavations in the wood are simply to make a suitable microenvironment for the fungus to grow. In other cases, wood tissue is the main food, and fungi weaken the defense response from the host plant.
Ambrosia fungi are fungal symbionts of ambrosia beetles including the polyphagous and Kuroshio shot hole borers.
Xyleborini are a tribe of ambrosia beetles, highly specialized weevils of the subfamily Scolytinae. Much of the ambrosia beetle fauna in Eurasia and the Americas consists of Xyleborini species. Some Xyleborini are notorious invasive species.
Xyleborus glabratus, the redbay ambrosia beetle, is a type of ambrosia beetle invasive in the United States. It has been documented as the primary vector of Raffaelea lauricola, the fungus that causes laurel wilt, a disease that can kill several North American tree species in the family Lauraceae, including redbay, sassafras, and avocado.
Xylosandrus compactus is a species of ambrosia beetle. Common names for this beetle include black twig borer, black coffee borer, black coffee twig borer and tea stem borer. The adult beetle is dark brown or black and inconspicuous; it bores into a twig of a host plant and lays its eggs, and the larvae create further tunnels through the plant tissues. These beetles are agricultural pests that damage the shoots of such crops as coffee, tea, cocoa and avocado.
Xyleborus affinis, the sugarcane shot-hole borer, is a species of ambrosia beetle in the family Curculionidae. It is found on all continents with woodlands and is one of the most widespread ambrosia beetles internationally, primarily in areas with humid tropical climates like Florida. Like other ambrosia beetles, Xyleborus affinis is attracted to dead fallen logs in early stages of decay. Due to the presence of X. affinis on moist timber, this species is mistakenly believed to be the cause of tree death. However, there is insufficient evidence to support their presence as a causal factor. Rather, these beetles are believed to target and accelerate the decay process of trees that are already deteriorated and weak.
Xylosandrus germanus, known generally as the alnus ambrosia beetle or black stem borer, is a species of ambrosia beetle in the family Curculionidae. X. germanus poses challenges in woody ornamental species and orchard crops such as apples and pecan. Furthermore, X. germanus is recognized or suspected to act as a vector for plant pathogens to varying extents, potentially leading to the decline of trees. The black stem borer is native to eastern Asia, but is an invasive species in Europe and North America. This species carries and feeds on associated ambrosia fungus, Ambrosiella grosmanniae.
Xyleborinus saxesenii, commonly known as the fruit-tree pinhole borer, is a species of ambrosia beetle in the family Curculionidae. It is native to the Palaearctic region but has been introduced in many locations, including North America. X. saxesenii typically live in freshly dead wood, but it has also been reported to attack live trees. Such attacks on live trees may lead to economic damage.
Euplatypus parallelus, previously known as Platypus parallelus, is a species of ambrosia beetle in the weevil family Curculionidae. The adults and larvae form galleries in various species of tree and logs. It is native to Central and South America but has spread globally, is present in Africa and is well established in tropical Asia.
Cnestus mutilatus, commonly known as the camphor shot borer, camphor shoot borer, or sweetgum ambrosia beetle, is a species of ambrosia beetle in the subfamily Scolytinae of the weevil family Curculionidae. It is native to Asia, but has been established as an invasive species in the United States since 1999.
Xyleborus dispar is a species of bark beetle commonly called the Pear blight beetle, or the European shothole borer. It is an invasive species in North America, and can be a pest in orchards and forests throughout its range.
Euwallacea interjectus is a species of ambrosia beetle in the species complex called Euwallacea fornicatus. It is native to Asia but has been introduced to the Western hemisphere over the last century.
Euwallacea piceus, is a species of weevil native to Oriental Asia but introduced to African and other Westerns Pacific parts of the world. It is a serious pest in tropical and subtropical parts of the Americas.
Debus emarginatus, is a species of weevil widely distributed throughout the Old World tropics.
Euwallacea perbrevis, commonly known as tea shot-hole borer, is a species of weevil native to South and South-East Asia through to Australia, but introduced to Western countries.
Ambrosiella roeperi is the fungal symbiont of the granulate ambrosia beetle, Xylosandrus crassiusculus, facilitating this insect’s capacity to accumulate on and damage a diverse array of woody plants from around the world. It is one of several important nutritional partners derived from order Microascales that sustain and are transported by xylomycetophagous scolytine beetles.
Euwallacea validus is a species of Euwallacea beetle native to Asia. The beetle species was discovered in Long Island, New York in 1975. Like other Euwallacea species beetles, E. validus is known for its mutualistic symbiotic relationship with fungi, acting as a vector for Fusarium oligoseptatum and Raffaelea subfusca, often using Tree of Heaven as a preferred host. Out of the five confirmed species of Euwallacea spp. in the United States, E. validus is the most widespread and longest established, yet much about their second fungal partner, Raffaelea subfusca, is not known.
Euwallacea is a genus of typical bark beetles in the family Curculionidae. They are commonly known as Ambrosia beetles, as all species are symbiotic with Ambrosia fungi. Originally from Asia or Wallacea, they are now found worldwide. Many species are pests, causing damage to, or the death of, tree species valued for their fruit or timber.