Melittobia Australica
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''Melittobia australica'' is a species of
chalcid wasp Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, me ...
from the family
Eulophidae The Eulophidae are a large family of hymenopteran insects, with over 4,300 described species in some 300 genera. The family includes the genus ''Elasmus'', which used to be treated as a separate family, "Elasmidae", and is now treated as a subf ...
which is a gregarious ecto-
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
of acuealate
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
.


Description

''Melittobia australica'' is a small wasp between 1.1 and 1.4 mm in length but has the typical wasp body plan of a head, thorax, abdomen body structure with the "wasp waist". It is sexually dimorphic with males is normally being larger than females, males are 1.2 to 1.4 mm in length while females are 1.1 to 1.3 mm. Males also have a with a wider head and smaller wings are smaller and their antennal scape is significantly broader. The colour difference is that the males are also a honey brown whereas the females are coloured dark brown. There are at least two morphs of females in ''M. australica'' which differ in the size of the abdomen, the size of the eyes and the extent of wing development. The "crawler" morph have a normal abdomen, small eyes and underdeveloped wings. The "flier" and possibly the "jumper" morphs have larger eyes and wings and a smaller abdomen. The "crawlers" remain within the host nest for the whole of their life cycle while the other morphs disperse as adults.


Distribution

''Melittobia australica'' was first described by the American entomologist
Alexandre Arsène Girault Alexandre Arsène Girault ( ; 9 January 1884 – 2 May 1941) was an American entomologist specializing in the study of chalcid wasps. An eccentric and controversial figure, Girault was also a prolific and dedicated entomologist. He published mor ...
in 1912 from Australia but it has since been recorded from North America, Central America, northern South America, the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
, Africa, eastern Asia, Europe (
Sicily (man) it, Siciliana (woman) , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = Ethnicity , demographics1_footnotes = , demographi ...
) and New Zealand. Some authorities believe that it is indigenous to Australia but that human commercial activity has allowed it to spread and become cosmopolitan.


Habitat

In Australia ''Melittobia australica'' was original reared from the nest of the
sphecid The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be ...
wasp '' Pison spinolae'' by Girault and rainforest was the original habitat.


Biology

''Melittobia australica'' is a parasitoid and its primary hosts are solitary bees and wasps. Its life cycle starts with the female finding the nest of a suitable host where the progeny are in the prepupal stage. The female feeds on the prepupa, she punctures it with the
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
and feeds on the exuded body fluids, using the proteins in the ingested fluid for ovogenesis. She lays 10–50 eggs per day under the pupal covering of the host. Fertilised eggs develop into female wasps and unfertilised eggs into males, a behaviour known as
arrhenotoky Arrhenotoky (from Greek -τόκος ''-tókos'' "birth of -" + ἄρρην ''árrhēn'' "male person"), also known as arrhenotokous parthenogenesis, is a form of parthenogenesis in which unfertilized eggs develop into males. In most cases, parthen ...
. The eggs are normally laid on a single host within the nest and many females may lay eggs on the same host which can be completely covered in larval ''M. australica'' of different ages which emerge from the eggs a few days after oviposition. The larvae feed on the tissue of the host and their development of the fames into the differing morphs is determined by the density of larvae feeding on the host. Where there is a low density of larvae the females will mainly be "crawlers" whereas intermediate densities will result in "jumpers" or and high densities of larvae will cause mostly "fliers" to develop. Where the host larvae are small the female may lay her eggs on more than one host. Male ''M. australica'' produce a very powerful
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
to attract females. As they have been laid on the same host at the same time the males mature at the same time as their sisters and often mate with them. If males encounter other males they will fight, often resulting in the death of the losing male. It has also been observed that the first male to emerge will often decapitate any other males either before the merge or immediately after emergence and I another observation the first male to emerge stopped the development of the other males by touching them with his antennae. There are up to 50 females per male and it is not known why the males fight when there are so many females. Once the male finds a receptive female they start a courtship ritual which involves tactile, chemical and auditory signals including the release of pheromone from the abdomen, the raising and lowering of the male's legs and the male rubbing the female with his antennae which also have a large gland which may produce pheromones too. The males hold the female's head in their mandibles, just below the
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
and maintain antennal contact throughout the ritual which usually lasts 15 minutes. The ritual is followed by mating which fertilises the females eggs. The female then finds a new host, in the same nest if she is a "crawler" or in a new nest if she is a "jumper" or a "flier" and begins to lay eggs on the new host. Each female lays a mean of 10.9 eggs on a host and these take 2–3 days to hatch, the larvae take 20–30 days to develop into adults, depending on conditions, and begin the life cycle again. Under laboratory conditions females live up to 9 days as adults and the total livespan is between 31 and 37 days. The female will normally mate only once while the males are
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
. If no males are present or an inseminated female has used up all of the sperm from mating she may lay a few unfertilised eggs to produce males, often mating with her own son. ''M. australica'' is a gregarious ectoparasitoid and when the female oviposits on a host she releases a pheromone which attracts other females to that site. All of the progeny resulting form a mating are laid on the same host and they develop together, consuming the host's tissues and eventually killing it. The newly mature females move to the original oviposition site and release a pheromone which induces nearby females to co-operate in forming a group, called a "chewing circle", which chews through the cuticle of the host allowing the adult wasps to escape. The adults, however are solitary and after mating the males die and the females move to a new host. Females seem to detect the cells of the hosts by chemical means and once a nest is detected the female remains in its vicinity but they do not appear to be attracted to the cells, rather they are "chemically arrested". Female ''M. australica'' have been observed excavating access holes through the mud walls of the nests of '' Pison'' sp. with their mandibles, just one hole per cell was excavated and several females were observed working on each hole. Only a single female at a time worked on the digging, each taking turns. Adult females are responsive to light,
phototaxis Phototaxis is a kind of taxis, or locomotory movement, that occurs when a whole organism moves towards or away from a stimulus of light. This is advantageous for phototrophic organisms as they can orient themselves most efficiently to receive li ...
, with the crawlers being negatively phototaxic and the fliers positively phototaxic, i.e. attracted to light, so that they leave the host nest after mating. The females can delay feeding and egg laying for several weeks until the host reaches an appropriate developmental stage, they will even wait in empty cells until the host uses them. ''M. australica'' is totally dependent on the host wasps for food, the females feed on the host before oviposition and the larvae consume the host. Adult males probably do not feed. The primary hosts are solitary wasps and bees but it will also parasitise
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
s of other insect orders where these are present in the host's nest.
Bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
s have also been found to be hosts of this wasp. Where populations of ''M. australica'' are high it can have a noticeable effect on the populations of host species and many of the host species have evolved defences against ''M. australica'' including nest location, chemical defences, physical barriers and defensive behaviours. Where ''M. autralica'' is common there can be a high mortality of its host species, which are often pollinators, and so ''M. australica'' parasitism can lead to an inhibition in the reproductive and dispersal capabilities of many plants. There is very little information about predators and parasites of ''M. australica'', although an egg hyperparasitoid, '' Anagrus putnamii'', has been recorded it has not been confirmed that it was using ''M. australica'' as its host. In addition, there have been observations of parasitic
mites Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
which feed on larvae of the
sphecid The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be ...
wasp ''
Sceliphron ''Sceliphron'', also known as black mud daubers or black mud-dauber wasps, is a genus of Hymenoptera of the Sphecidae family of wasps. They are solitary mud daubers and build nests made of mud. Nests are frequently constructed in shaded niches, o ...
'' spp. competing with ''M. australica'' larvae and even feeding upon them.


Hosts

The first recorded host of ''Melittoboa australica'' was ''Pison spinolae''; however, the primary hosts are leaf cutting bees ''
Megachile The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pe ...
'' spp, mason wasps and mud dauber wasps but it is wide-ranging in its choice of hosts. There are no indigenous
social wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s in Australia but in New Zealand and North America it has been recorded in the nests of ''
Vespula ''Vespula'' is a small genus of social wasps, widely distributed in the Northern Hemisphere. Along with members of their sister genus ''Dolichovespula'', they are collectively known by the common name yellowjackets (or yellow jackets) in North ...
'' species, as well as nests of ''
Polistes Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella ...
'', ''
Bombus A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
'' and
honeybees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmo ...
''
Apis mellifera The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
''. The bombyliid fly '' Anthrax angularis'' and the
ichneumonid The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family (biology), family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 2 ...
'' Stenarella victoriae'' are parasites in the nests of ''Sceliphron'' spp. and have been recorded as being viable hosts for ''M. australica''. In addition, ''M. australica'' were found on dipteran puparia found in the nests of ''Sceliphron'' which are thought to be parasites on the spiders stocked in empty cells by the wasps.


References

{{Taxonbar, from=Q13833896 Eulophidae Insects described in 1912 Agricultural pest insects Hymenoptera of New Zealand Hymenoptera of Europe