Megarhyssa Praecellens
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''Megarhyssa'', also known as giant ichneumonid wasps, giant ichneumons, or stump stabbers, is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of large
ichneumon wasp The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25, ...
s, with some species known for having the longest
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 typica ...
s of any
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s. They are idiobiont ectoparasitoids of the larvae of wood-boring
horntail Horntails or wood wasps are any of the 150 non-social species of the hymenopteran family Siricidae, a type of xylophagous, wood-eating sawfly. The common name "horntail" derives from the stout, spine-like structure at the end of the adult's abdom ...
wasps. The ovipositor can be mistaken for a large stinger. This is a genus of
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the clade Holometabola. Immatur ...
insects Insects (from Latin ') are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed ...
within subfamily Rhyssinae that includes 37 species and belongs to
Ichneumonidae The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25 ...
, the family of
wasps 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 ...
with the highest biodiversity in the world.


Geographical range and habitat

''Megarhyssa'' species occur all over the world. These are the only four ''Megarhyssa'' species known to inhabit the paleartic region inhabiting decidious forests. They are widespread across the United States, and Canada. The species '' M. macrurus'', '' M. atrata'', and '' M. greenei'' are known to be
sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
in the northeastern United States.  ''M. macrurus'' is known to inhabit further southern regions as well, reaching Mexico. '' M. nortori'' has been introduced to South Africa, Australia, and New Zealand as a biological control agent.


Description and identification

''Megarhyssa'' male adults reach body lengths going from 2.3 to 3.8 cm (0.9 to 1.6 inches), while female adults can measure from 3.5 to 7.5 cm (1.5 to 3 inches). Both sexes can be distinguished because females have an extremely slender, and long organ to lay eggs called 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 typica ...
. This organ is much longer than the body itself, its length can range from 5 to 10 cm (2 to 4 inches). ''Megarhyssa'' adults show variation in coloration, including dark brown, bluish black, reddish brown and/or bright yellow.


Species of the United States and Canada

The species '' M. atrata'' (Fabricius) shows a bright yellow head, and an almost completely black body. The wings are black as well. It can be easily distinguished from the species '' M. macrurus'', '' M. greenei'', and '' M. nortori'', which show yellow and brown striped color patterns that resemble much, reason for which these other species require identification by specialists.


Ecology

Family
Ichneumonidae The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25 ...
is composed of
parasitoid wasps Parasitoid wasps are a large group of hymenopteran 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, sooner or later causi ...
. The adults do not often feed. When they do so, they feed on water droplets found on leaves that often contain
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
. The
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
feed on flesh from hosts the adults locate. In the Neartic, Paleartic, and Australian regions, some ''Megarhyssa'' species such as ''M. emarginatoria'', ''M. jezoensis'', and '' M. nortori'' are known to parasitize
Siricidae Horntails or wood wasps are any of the 150 non-social species of the hymenopteran family Siricidae, a type of wood-eating sawfly. The common name "horntail" derives from the stout, spine-like structure at the end of the adult's abdomen which is ...
. The species '' M. atrata'', '' M. macrurus'', and '' M. greenei'' show diurnal activity and have undergone
allopatric speciation Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
and parasitize the
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
of one species of woodwasp: ''
Tremex columba ''Tremex columba'', also known as the pigeon tremex or pigeon horntail, is a species of horntail that is native to eastern and western North America. Appearance and behavior The females are larger than the males, with females growing to 25-3 ...
''. These three species are capable of coexisting with each other because their
ovipositors The ovipositor is a tube-like organ (anatomy), organ used by some animals, especially insects, for the laying of Egg (biology), eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of t ...
have different lengths. As a result, they find their
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
at different depths within trees, or logs. The distance the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
is within the bark is equal to the length of the ovipositor of each species. The ovipositor of '' M. atrata'' is known to puncture up to 14 cm inside the wood, and such length makes this
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 ...
the largest species in
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 ...
order.


Reproduction


Reproductive cycle

The reproductive cycle of ''Megarhyssa'' begins with the female locating a
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
of ''
Tremex columba ''Tremex columba'', also known as the pigeon tremex or pigeon horntail, is a species of horntail that is native to eastern and western North America. Appearance and behavior The females are larger than the males, with females growing to 25-3 ...
'' inside the bark of a tree. She lays an egg close by or on the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
, the ''Megarhyssa'' larva devours it, it pupates under the bark, and emerges the following year as an adult. Males often emerge first. The species '' M. atrata'', '' M. macrurus'', and '' M. greenei'' share their territory, and habitat, showing the same behavior: males emerge, and remain close by the trees where more
wasps 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 ...
from these species emerge. ''Megarhyssa'' males often try to squeeze inside the holes of the bark of trees even before females emerge because preemergence
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. '' Copulation'' is the union of the sex organs of two sexually repr ...
is easier to carry out than postemergence
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. '' Copulation'' is the union of the sex organs of two sexually repr ...
. In females the
genital A sex organ, also known as a reproductive organ, is a part of an organism that is involved in sexual reproduction. Sex organs constitute the primary sex characteristics of an organism. Sex organs are responsible for producing and transporting ...
opening is oriented anteriorly; in males, posteriorly. In preemergence mating, the male inserts his
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
inside the hole and inside the genital opening of the female. In postemergence mating, the male must be on the abdomen of the female, and bend his
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
all around it to reach her
genitalia A sex organ, also known as a reproductive organ, is a part of an organism that is involved in sexual reproduction. Sex organs constitute the primary sex characteristics of an organism. Sex organs are responsible for producing and transporting ...
properly. Afterward, females look for a spot in the same area where the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
they need to feed their
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
is found. Once they locate a
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
, females must pierce the bark of trees using 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 typica ...
to reach the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
. The exact movements of the ovipositor remain unclear, although it is known females carry out a series of movements with it: she puts 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 typica ...
as vertically as possible relative to the wood. The intersegmentary segments must be fully unfolded and coupled to the rotation of the last abdominal segment. Then, the stylus can start penetrating the wood. The
wasps 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 ...
always follow a very straight line without deviating to reach the chamber where the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
lies.


Mate choice and competition

''Megarhyssa'' males have been known to detect other individuals to emerge before the emergence happened. Males of the three species aggregate around the hole to find out the
sex Sex is the biological trait that determines whether a sexually reproducing organism produces male or female gametes. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inheri ...
of the individual. Males do so hearing how the females bite, and chew through the wood to find their way out. If the individual is a male, many males often show little interest, and fly away. In case it is a female, males try to
mate Mate may refer to: Science * Mate, one of a pair of animals involved in: ** Mate choice, intersexual selection *** Mate choice in humans ** Mating * Multi-antimicrobial extrusion protein, or MATE, an efflux transporter family of proteins Pers ...
with her as soon as possible. The
sensory organs A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as s ...
that allow the
wasps 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 ...
to detect their mates, and hosts is close to the antennae.


Evolution of flight

The movement that flight makes possible for
parasitoids 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 ...
is crucial to reproduce, so it is related directly with fitness. In '' M. nortori'', it is known females perform longer single flights and overall longer flights than males. It is thought this is because females are the individuals that reach new habitats, redistribute the progeny, and locate hosts. These activities require longer-range flights. Males of ''M. nortori'' spend most of their time in aggregations around sites from which females are about to emerge. Even if males are disturbed and fly away from the site, they return and regroup around the same point of the tree. They have been known to be flying around a specific tree and patrolling it over time. Such features lead to the evolution of shorter-range flights.


Species

Species within the genus: *'' Megarhyssa arisana'' *'' Megarhyssa atomistica'' *'' Megarhyssa atrata'' *'' Megarhyssa aurantia'' *'' Megarhyssa babaulti'' *'' Megarhyssa belulliflava'' *'' Megarhyssa bicolor'' *'' Megarhyssa bonbonsana'' *'' Megarhyssa cultrimacularis'' *'' Megarhyssa fulvipennis'' *'' Megarhyssa gloriosa'' *''
Megarhyssa greenei ''Megarhyssa greenei'', also known as Greene's giant ichneumonid wasp, is a species of large Ichneumonidae, ichneumon wasp. It is known from the United States and Canada. Description and identification It is very similar in appearance to ''Meg ...
'' *'' Megarhyssa hainanensis'' *'' Megarhyssa indica'' *'' Megarhyssa insulana'' *'' Megarhyssa jezoensis'' *'' Megarhyssa laniaria'' *'' Megarhyssa lenticula'' *'' Megarhyssa longitubula'' *'' Megarhyssa macrurus'' *'' Megarhyssa middenensis'' *'' Megarhyssa mirabilis'' *'' Megarhyssa nortoni'' *'' Megarhyssa obtusa'' *'' Megarhyssa perlata'' *'' Megarhyssa praecellens'' *'' Megarhyssa recava'' *'' Megarhyssa rixator'' *'' Megarhyssa rotundamacula'' *'' Megarhyssa strimacula'' *'' Megarhyssa superba'' *'' Megarhyssa taiwana'' *'' Megarhyssa vagatoria'' *''
Megarhyssa verae ''Megarhyssa'', also known as giant ichneumonid wasps, giant ichneumons, or stump stabbers, is a genus of large ichneumon wasps, with some species known for having the longest ovipositors of any insects. They are idiobiont ectoparasitoids of the ...
'' *'' Megarhyssa weixiensis'' *''
Megarhyssa wugongensis ''Megarhyssa'', also known as giant ichneumonid wasps, giant ichneumons, or stump stabbers, is a genus of large Ichneumonidae, ichneumon wasps, with some species known for having the longest ovipositors of any insects. They are idiobiont ectopara ...
''


References


External links

* *
Bugguide.net




* ttp://naturedocumentaries.org/3843/giant-ichneumon-megarhyssa-macrurus-ovipositing/ Video: Giant Ichneumon Wasp Ovipositing {{Authority control Insects used as insect pest control agents Biological pest control wasps Ichneumonidae genera Ichneumonidae Taxa named by William Harris Ashmead