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, m ...
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Eulophidae
The Eulophidae is 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 subfa ...
. It is one of the largest subfamilies of the Eulophidae containing over 100 genera and nearly 3,000 species. The species of the family Tetrastichinae are found in almost any type of terrestrial habitat and have a worldwide distribution, except Antarctica. They show a varied biology and hosts for Tetrastichinae wasps have been identified from over 100 different insect families, across 10 different orders and they have also been recorded as being
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 ...
s on
nematode
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
s, mites and spiders' eggs. Some species are even
phytophagous
A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet (nutrition), diet. These more broadly also encompass an ...
, while others are
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 ...
s and yet others are
gall
Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or war ...
formers.
Biology
The Tetrastichinae contains species which are mainly parasitic, although some species are known to be phytophages. Phytophagy has so far been shown to be secondary, especially in species in which the larvae are inquilines with gall forming insects and primary phytophagy has not been demonstrated. Endoparasitism is more frequent than ectoparasitiusm and taxa of either parasitic type may be gregarious or solitary. In the gregarious forms a single host may support very few parasitoids or it may host many of them, for example in a single
chrysalis
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages the ...
parasitised by '' Aprostocetus xanthopus'' over 2000 larvae were found. Sometimes hyperparasitism is shown, with the species in the genus '' Eutetrastichus'' (now synonymised with '' Baryscapus'') in particular being hyperparasites, and hyperparasitism may be obligate but it can also occur opportunistically in response to circumstance.
Some species are highly host specific while other species appear to be able to vary their hosts, this may be due to otherwise cryptic taxa which have not yet been identifies or described. The most common hosts are insects but
arachnid
Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s are also hosts for some taxa. Some of the groups within the Tetrastchinae all share similar hosts, such as the genus '' Tamarixia'' which parasitise ''
Psylloidea
Psylloidea is a Taxonomic rank, superfamily of Hemiptera, true bugs, including the jumping plant louse, jumping plant lice and others which have recently been classified as distinct families. Though the group first appeared during the Early Jura ...
'' while the majority of the species in ''
Aprostocetus
''Aprostocetus'' is a genus of hymenopteran insects of the Family (biology), family Eulophidae. The genus was erected by John O. Westwood in 1833. This very large group (about 800 described species) of parasitoid wasps has a global distribution.
...
'' solely parasitise
Cecidomyiidae
Cecidomyiidae is a family of diptera, flies known as gall midges or gall gnats. As the name implies, the larvae of most gall midges feed within plant tissue, creating abnormal plant growths called galls. Cecidomyiidae are very fragile small in ...
. In other genera it is the ecological or behavioural traits of the host which are important, the '' Minotetrastichus'' species parasitise leaf miners whether these are
Coleoptera
Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
,
Hymenoptera
Hymenoptera is a large order of insects, comprising the 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 parasitic.
Females typi ...
or
Lepidoptera
Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
; while other groups target galls irrespective of the nature of the gall former.
In many species reproduction is solely through
thelytoky
Thelytoky (from the Ancient Greek, Greek θῆλυς ''thēlys'' "female" and τόκος ''tókos'' "birth") is a type of parthenogenesis and is the absence of mating and subsequent production of all female diploid offspring as for example in aph ...
, i.e. female eggs are produced by
parthenogenesis
Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
, while in others varying proportions of males are produced from unfertilised eggs while females are produced from fertilised eggs. This can differ within species, for example there are no records of males in North American specimens of '' Tetrastichus asparagi'' but are recorded in small numbers in European samples. Another reason for the scarcity of males is that in some taxa (e.g. '' Melittobia'') they have been observed to stay within the host pupa on emergence and aggressively attack and kill each other until only a few survive. The surviving males then have to complete a complicated courtship ritual before they are able to copulate with the females.
Taxonomy
These small wasps are rather uniform in morphology and the difficulty of preserving specimens makes them difficult to study and classify and traditionally most species were placed in a single large genus ''
Tetrastichus
''Tetrastichus'' is a genus of hymenopteran insects of the family Eulophidae.
'' Tetrastichus planipennisi'' is a parasitoid of the emerald ash borer, a wood boring insect native to Asia which is an invasive species in North America. ''T. pla ...
'' until a study in 1987 by M.W.R. de V. Graham split this large genus into a number of smaller, more natural groupings, fifteen of which were named as new
genera
Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
. All subsequent taxonomic work on this subfamily has been based on Graham's revision.
Aprostocetus
''Aprostocetus'' is a genus of hymenopteran insects of the Family (biology), family Eulophidae. The genus was erected by John O. Westwood in 1833. This very large group (about 800 described species) of parasitoid wasps has a global distribution.
...
Chouioia
''Chouioia'' is a genus of endoparasitic wasp of the family Eulophidae. '' Chouioia cunea'' is considered an important parasite of the fall webworm
The fall webworm (''Hyphantria cunea'') is a moth in the family Erebidae known principally for ...
Tetrastichus
''Tetrastichus'' is a genus of hymenopteran insects of the family Eulophidae.
'' Tetrastichus planipennisi'' is a parasitoid of the emerald ash borer, a wood boring insect native to Asia which is an invasive species in North America. ''T. pla ...
Thymus
The thymus (: thymuses or thymi) is a specialized primary lymphoid organ of the immune system. Within the thymus, T cells mature. T cells are critical to the adaptive immune system, where the body adapts to specific foreign invaders. The thymus ...