Agathidinae
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Agathidinae is a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of braconid
parasitoid wasp Parasitoid wasps are a large group of hymenopteran Superfamily (zoology), superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, ...
s. Some species have been used in biological control programs.


Description

Agathidines are among the larger braconids, and diurnal members of this subfamily are often brightly patterned. They belong to the noncyclostome group. Several genera, such as ''Agathis'', '' Cremnops'' and '' Disophrys'', are characterized by elongate genae. Agithidines can be recognized by the following combinations of features: forewing veins M + CU not tubular in basal third or more, forewing vein RS complete to the wing margin (except '' Mesocoelus'' and '' Plesiocoelus),'' occipital carina absent, and second submarginal cell of forewing usually present (90% of species.)


Distribution

They are found worldwide, but are more diverse in the tropics. However, some genera such as ''Agathis'' and '' Earinus'' are more speciose in temperate regions.


Biology

Most Agathidines are solitary koinobiont endoparasitoids of concealed
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 ...
larvae. However, the tribe Disophrini attacks free living Lepidoptera larvae. The tribes Agathidini and Earinini attack the first and second
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
larvae of their hosts, contrasting with the Disophrini, which attack later instars, as well as the Cremnoptini, which parasitize every larval stage. Most species are diurnal, but many Disophrini are
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, with pale coloration and enlarged ocelli. Most species attack semi-concealed hosts, like leaf-rollers. Agathidines have three larval
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s. They likely seek out and kill competitors as first instars. As final instars, they emerge from their host caterpillar, feed externally, then spin their cocoons.
Temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
species overwinter in their hosts during their first instar. Some species, such as '' Earinus elator'', seek out hosts very early in the year, "about the time of bud-burst of their hosts' host plants." In relation to this, they have evolved to complete their metamorphosis before winter so they can more easily seek out hosts early. To prevent their freezing from their
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
-filled Meconium onward, they secrete a goo rich in
glycerol Glycerol () is a simple triol compound. It is a colorless, odorless, sweet-tasting, viscous liquid. The glycerol backbone is found in lipids known as glycerides. It is also widely used as a sweetener in the food industry and as a humectant in pha ...
and
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s over the waste. This reduces the freezing point of the water in the waste, preventing ice formation. Most tropical species are brightly colored and are involved in
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
. Studies of the species '' Alabagrus texanus'' have found that males that arrive early to a site where females recently emerged from were often successful, but the very young and very old males were always unsuccessful. Males also had
short-term memory Short-term memory (or "primary" or "active memory") is the capacity for holding a small amount of information in an active, readily available state for a short interval. For example, short-term memory holds a phone number that has just been recit ...
regarding sites of female emergence. Agathidines have been noted to be more abundant in ecosystems where fluids are harder to obtain. Many have concealed nectar extraction adaptations (CNEAs) that make them more adapted to gaining nectar from flowers, "perhaps particularly from
Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...
." These CNEAs include elongate malar regions, galea, glossa, and maxillary palps that form a tube. These elongate mouthparts have independently evolved in the subfamily six times. '' Agathis clavatus'' has been found to practice the mating system of males searching for females at feeding sites (SFF). This strategy is hardly found in Ichneumonoidea.


Taxonomy and phylogeny

The first attempt at a tribal arrangement for Agathidinae consisted of 2 tribes, Agathidini and Microdini. Sharkey revised the tribes in 1992 and considered 5 tribes, adding Cremnoptini, Disophrini, and Earini. The Microdini, then treated as Eumicrodini, was synonymized under Agathidini by Simbolotti and van Achterberg in 1999 to render 4 tribes. Phylogenetic analysis by Sharkey and Chapman in 2017 revealed that 4 tribes were insufficient and instead divided the subfamily into 7 tribes. They further found that the genera ''Asperagathis'', ''Bassus'', and ''Zosteragathis'' were not monophyletic. One problematic species, the Nearctic '' Bassus annulipes'', was found to represent a complex of species. Further, ''Bassus annulipes'' was rendered under the tribe Lytopylini while the Old World species fell under Agathidini.


Relationship with humans

Agathidines have been used for biological pest control. Notable examples include '' Agathis pumila'' for the western larch case-bearer, '' Alabagrus stigma'' for the sugarcane borer, and '' Bassus unicoloratus'' and Agathis gibbosa for the potato tuberworm. Other species play important roles in their native ecosystems to suppress forestry pests, such as Bassus tumidulus which controls the population of Gypsonoma aceriana.


Genera

The 63
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 ...
placed here include:


Tribe Agathidini Haliday, 1833

* '' Aerophilus'' Szépligeti, 1902 * '' Agathacrista'' Sharkey, 2013 * '' Agathigma'' Sharkey, 2017 * '' Agathis'' Latreille, 1804 * '' Alabagrus'' Enderlein, 1920 * '' Aphelagathis'' Sharkey, 2015 * '' Asperagathis'' Sharkey, 2017 * '' Bassus'' Fabricius, 1804 (
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
) * '' Braunsia'' Kriechbaumer, 1894 * '' Camptothlipsis'' Enderlein, 1920 * '' Chimaeragathis'' Sharkey, 2017 * '' Cymagathis'' Sharkey, 2017 * '' Gyragathis'' van Achterberg & Long, 2010 * '' Ischnagathis'' Cameron, 1909 * '' Leuroagathis'' Sharkey, 2017 * '' Liragathis'' Sharkey, 2017 * '' Neothlipsis'' Sharkey, Parys & Clutts, 2011 * '' Pharpa'' Sharkey, 1986 * '' Plesiocoelus'' van Achterberg, 1990 * '' Pneumagathis'' Sharkey, 2015 * '' Scabagathis'' Sharkey, 2017 * '' Trachagathis'' Viereck, 1913 * '' Trochantagathis'' Sharkey, 2017 * '' Xanthagathis'' Sharkey, 2017 * '' Zamicrodus'' Viereck, 1912 * '' Zosteragathis'' Sharkey, 2017


Tribe Agathirsini Sharkey, 2017

* '' Agathirsia'' Westwood, 1882 * '' Crassomicrodus'' Ashmead, 1900 * '' Gelastagathis'' Sharkey, 2015 * '' Marjoriella'' Sharkey, 1983


Tribe Cremnoptini Sharkey, 1992

* '' Cremnops'' Foerster, 1862 * '' Cremnoptoides'' van Achterberg & Chen, 2004 * '' Hyrtanommatium'' Enderlein, 1920 * '' Labagathis'' Enderlein, 1920 * '' Megalagathis'' Schulz, 1906 * '' Mesoagathis'' Cameron, 1905 * '' Zacremnops'' Sharkey & Wharton, 1985


Tribe Disophrini Sharkey, 1992

* '' Coccygidium'' Saussure, 1892 * '' Coronagathis'' van Achterberg & Long, 2010 * '' Disophrys'' Förster, 1863 * '' Euagathis'' Szépligeti, 1900 * '' Gyrochus'' Enderlein, 1920 * '' Hemichoma'' Enderlein, 1920 * '' Hypsostypos'' Baltazar, 1963 * '' Liopisa'' Enderlein, 1920 * '' Macroagathis'' Szépligeti, 1908 * '' Monophrys'' van Achterberg, 1988 * '' Oreba'' Cameron, 1900 * '' Pelmagathis'' Enderlein, 1920 * '' Protroticus'' van Achterberg, 1988 * '' Troticus'' Brullé, 1846 * '' Zelodia'' van Achterberg, 2010 * '' Zelomorpha'' Ashmead, 1900


Tribe Earinini Sharkey, 1992

* '' Chilearinus'' Sharkey, 2022 * '' Earinus'' Wesmael, 1837


Tribe Lytopylini Sharkey, 2017

* '' Amputoearinus'' Sharkey, 2006 * '' Austroearinus'' Sharkey, 2006 * '' Lytopylus'' Förster, 1862 * '' Sesioctonus'' Viereck, 1912 * '' Smithagathis'' Sharkey, 2017


Tribe Mesocoelini Achterberg, 1990

* '' Aneurobracon'' Brues, 1930 (formerly in Orgilinae) * '' Mesocoelus'' Schulz, 1911 (formerly in Orgilinae) * '' Therophilus'' Wesmael, 1837 * Undescribed genus (former '' Bassus annulipes'' group)


Gallery

Alabagrus texanus ♀ (40447935543).jpg, Agathidini - ''Alabagrus texanus'' Agathirsia (26449415).jpg, Agathirsini - ''Agathirsia'' sp. Biroia.jpg, Cremnoptini - ''Megalagathis'' sp. Amputostypos.jpg, Disophrini - ''Coccygidium '' sp. Earinus elator (Braconidae) - (imago), Arnhem, the Netherlands.jpg, Earini - ''Earinus elator'' Lytopylus.jpg , Lytopylini - ''Lytopylus'' sp. Therophilus.jpg , Mesocoelini - ''Therophilus'' sp.


References


External links


Photos on BugGuide

DNA barcodes at BOLD systems
{{Taxonbar, from=Q1949213 Braconidae Apocrita subfamilies