Ambrosia beetles are beetles of the weevil subfamilies Scolytinae and Platypodinae (Coleoptera, Curculionidae), 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. [1] 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. [2] The majority of ambrosia beetles colonize xylem (sapwood and/or heartwood) of recently dead trees, but some colonize stressed trees that are still alive, and a few species attack healthy trees. [3] 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.
Until recently ambrosia beetles have been placed in independent families Scolytidae and Platypodidae, however, they are in fact some of the most highly derived weevils, and are now placed in the subfamilies Scolytinae and Platypodinae of Family Curculionidae [4] [5] [6] There are about 3,000 known beetle species employing the ambrosia strategy. [7]
Ambrosia beetles are an ecological guild, but not a phylogenetic clade. The ambrosia habit is an example of convergent evolution, as several groups evolved the same symbiotic relationship independently. [8] The highest diversity of ambrosia beetles is in the tropics. In the Paleotropical region, hundreds of species of Xyleborini and Platypodinae are the main agent initiating dead wood decomposition. In the Neotropics, Platypodinae and Xyleborini are joined by the scolytine tribe Cortylini. Compared to the diversity in the tropics, ambrosia beetle fauna in the temperate zone is rather limited. In the Nearctic region it is dominated by a few species from Cortylini, Xyleborini and Xyloterini. In the Palearctic realm, significant groups are Xyloterini and Xyleborini, joined by Scolytoplatypodini in the Far East.
Beetles and their larvae graze on mycelium exposed on the gallery walls and on bodies called sporodochia, clusters of the fungus' spores. Most ambrosia beetle species don't ingest the wood tissue; instead, the sawdust resulting from the excavation is pushed out of the gallery. Following the larval and pupal stage, adult ambrosia beetles collect masses of fungal spores into their mycangia and leave the gallery to find their own tree.
A few dozen species of ambrosia fungi have been described, currently in the genera Ambrosiella , Meredithiella, and Phialophoropsis (from Microascales), Afroraffaelea and Raffaelea (from Ophiostomatales), Ambrosiozyma (Saccharomycetales), Fusarium and Geosmithia (from Hypocreales), and Flavodon (from Basidiomycota). [3] Many more species remain to be discovered. Little is known about the bionomy or specificity of ambrosia fungi. Ambrosia fungi are thought to be dependent on transport and inoculation provided by their beetle symbionts, as they have not been found in any other habitat. All ambrosia fungi are probably asexual and clonal. [9] Some beetles are known to acquire ("steal") fungal inoculum from fungal gardens of other ambrosia beetle species. [10]
During their evolution, most scolytid and platypodid weevils became progressively more or less dependent on fungi regularly co-habiting dead trees. This evolution had various outcomes in different groups:
The vast majority of ambrosia beetles colonize dead trees, and have minor or no economic effect. A few species are able to colonize living stressed trees (Xylosandrus). [15] A few species are able to attack live and healthy trees, and those can reach epidemic proportions in non-native, invaded regions (Xyleborus glabratus, Euwallacea fornicatus [16] ).
Beetle species that readily colonize lumber, such as sawlogs, green lumber, and stave-bolts, often cause region-specific economic loss from the pinhole and stained-wood defects caused by their brood galleries. In Northern USA and Canada, conifer logs are attractive to Trypodendron lineatum (Oliv.) during the spring swarming flight (Dyer 1967). [17] Previous studies showed that short log sections become attractive more rapidly than corresponding long logs.
The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae.
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.
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.
Austroplatypus incompertus, a type of ambrosia beetle, is endemic to Australia. They are found in the mesic forests, and subtropical and tropical ecosystems along the east coast of Australia. There are many unique characteristics attributable to the A. incompertus, like their gallery excavation in several Eucalyptus species, their obligate eusocial behavior, their relationship with fungi, and their unique sexual dimorphism. These beetles are one of the only insects that display obligate eusocial behavior. Additionally, their sexually dimorphic traits are of interest, since body size is reversed with males having smaller torsos than female a.incompertus beetles.
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.
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.
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.
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.
Sueus niisimai is a species of weevil found in India, Sri Lanka, China, Japan, Taiwan, Korea, Malaysia and Indonesia. It is also found in Australia and Fiji, presumably introduced.
Xyleborinus andrewesi, is a species of weevil widely distributed throughout the Old World tropics and introduced to many New World countries.
Debus emarginatus, is a species of weevil widely distributed throughout the Old World tropics.
Xylosandrus morigerus, is a species of weevil widespread throughout Afrotropical, Australian, Neotropical, Oceania and Oriental regions. It is also introduced to Palearctic regional countries.
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.
Cryphalus mangiferae, the mango bark beetle, is a tiny tropical bark beetle which attacks mango trees. The species belongs to the family Curculionidae, subfamily Scolytinae. It is often cited as a vector of plant pathogenic fungi infecting mango trees and in some countries like Pakistan it is regarded as a serious threat to mango cultivation.
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.