The Stephanidae, sometimes called crown wasps,
are a
family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
parasitoid wasp
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 causin ...
s. They are the only living members of the superfamily
Stephanoidea. Stephanidae has at least 345 living species in 11
genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
.
The family is considered
cosmopolitan
Cosmopolitan may refer to:
Food and drink
* Cosmopolitan (cocktail), also known as a "Cosmo"
History
* Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953
Hotels and resorts
* Cosmopoli ...
in distribution, with the highest species concentrations in
subtropical
The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
and moderate climate zones.
Stephanidae also contain four extinct genera described from both
compression fossil
A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason fo ...
s and
inclusions in
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
.
Biology
Stephanids are noted for their ocellar corona, a semicircular to circular set of projections around the middle
ocellus
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 ...
, forming a "crown" on the head.
Only stephanids and the similarly old Hymenoptera family
Orussidae
The Orussidae or the parasitic wood wasps represent a small family of sawflies (" Symphyta"). Currently, about 93 extant and four fossil species are known. They take a key position in phylogenetic analyses of Hymenoptera, because they form the s ...
have ocellar coronae, and it is uncertain if they developed the structure separately or if a common ancestor of both developed it and it was then lost in all but the two families. Weakly developed grooves starting at the base of the antennae and extending past the eyes to the back of the head capsule are present. This feature is seen more developed in hymenopteran families in which the adults emerge from pupal chambers in wood.
All genera of Stephanidae have a
pronotum
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
that is modified to some extent. They bear highly modified hind legs, with a swollen hind
femur
The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
that has large teeth on the underside, and the tibiae have a tip end that widens distinctly. The largest species, reaching up to in length, are found in the genus ''
Megischus
''Megischus'' is a genus of crown wasp. It was circumscription (taxonomy), circumscribed by Gaspard Auguste Brullé in 1846. The genus has a cosmopolitan distribution, and over eighty species are recognized.
It is the type genus of the tribe (bio ...
''. Stephanids are noted as
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 of
xylophagous
Xylophagy is a term used in ecology to describe the habits of an herbivorous animal whose diet consists primarily (often solely) of wood. The word derives from Greek ''ξυλοφάγος'' (''xulophagos'') "eating wood", from ''ξύλον'' (') ...
beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
e, with a majority of the stephanids hosts coming from the families
Cerambycidae
The longhorn beetles (Cerambycidae), also known as long-horned or longicorns, are a large family of beetles, with over 35,000 species described. Most species are characterized by extremely long antennae, which are often as long as or longer than ...
and
Buprestidae
Buprestidae is a family of beetles known as jewel beetles or metallic wood-boring beetles because of their glossy iridescent colors. Larvae of this family are known as flatheaded borers. The family is among the largest of the beetles, with some ...
, though some
Curculionidae
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.
T ...
and occasional hymenopteran hosts are taken. One species, ''
Schlettererius cinctipes'', is a known parasitoid of
horntail
Horntail or wood wasp 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 u ...
wasps and has been introduced to
Tasmania
)
, nickname =
, image_map = Tasmania in Australia.svg
, map_caption = Location of Tasmania in AustraliaCoordinates:
, subdivision_type = Country
, subdi ...
as a
biological pest control
Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
agent.
Members of the genus ''
Foenatopus'' are parasitoids of ''
Agrilus sexsignatus'',
wood-boring beetle larvae found infesting
eucalyptus
''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
in the
Philippines
The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no),
* bik, Republika kan Filipinas
* ceb, Republika sa Pilipinas
* cbk, República de Filipinas
* hil, Republ ...
. The rate of parasitism for an ''A. sexsignatus'' population was recorded to vary from only 2% up to 50% of the population.
Taxonomy and fossil record
The family is noted to be the most
basal group of hymenopterans in the suborder
Apocrita
Apocrita is a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" ( petiole) formed ...
. They are the only living group left over from the early diversification of Apocrita. In general, the family is considered rare, with close to 95% of the species known to have been described from single specimens. Until the early 1800s, members of Stephanidae were grouped into the parasitic wasp superfamily
Ichneumonoidea
The superfamily Ichneumonoidea contains one extinct and three extant families, including the two largest families within Hymenoptera: Ichneumonidae and Braconidae. The group is thought to contain as many as 100,000 species, many of which have not ...
based on the superficial resemblance between some members of the two groups.
William Elford Leach
William Elford Leach Royal Society, FRS (2 February 1791 – 25 August 1836) was an English zoologist and marine biologist.
Life and work
Elford Leach was born at Hoe Gate, Plymouth, the son of an attorney. At the age of twelve he began a me ...
suggested a new family grouping for the stephanids in the 1815 edition of ''
Edinburgh Encyclopædia
The ''Edinburgh Encyclopædia'' is an encyclopaedia in 18 volumes, printed and published by William Blackwood and edited by David Brewster between 1808 and 1830. In competition with the Edinburgh-published ''Encyclopædia Britannica'', the ''Edin ...
''.
The name Stephanidae was first published by
Alexander Henry Haliday
Alexander Henry Haliday (1806–1870, also known as Enrico Alessandro Haliday, Alexis Heinrich Haliday, or simply Haliday) was an Irish entomologist. He is primarily known for his work on Hymenoptera, Diptera, and Thysanoptera, but worked on ...
in his 1839 ''Hymenoptera Britannica''. About 110 years later, the stephanids were placed into a separate superfamily, Stephanoidea, by P.L.G. Benoit, along with the proposed family "Stenophasmidae". The latter group was moved out of Stephanoidea in 1969 by
Alexandr Pavlovich Rasnitsyn
Alexandr Pavlovich Rasnitsyn (Russian: Александр Павлович Расницын) is a Russian entomologist, expert in palaeoentomology, and Honored Scientist of the Russian Federation (2001).
His scientific interests are centered on ...
, who transferred the "Stenophasmidae" to the family
Braconidae
The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
and
synonymized the two families.
Fossil specimens related to the family are uncommon, and most are dated to the
Tertiary
Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago.
The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
. The oldest confirmed members of the family are ''
Kronostephanus zigrasi'', ''
Lagenostephanus lii'', and ''
Phoriostephanus exilis'' all known from the
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
(
Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
)
Burmese amber
Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
.
Other early member of the family is the
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus ''
Archaeostephanus'', which is known from a single species ''
Archaeostephanus corae'' found in the
late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
New Jersey amber and first described in 2004. The first species to be described from the fossil record was ''
Protostephanus ashmeadi'', which was first published in 1906 by paleoentomologist
Theodore Dru Alison Cockerell
Theodore Dru Alison Cockerell (1866–1948) was an American zoology, zoologist, born at Norwood, England, and brother of Sydney Cockerell. He was educated at the Middlesex Hospital Medical School, and then studied botany in the field in Colorad ...
. The specimen is also the youngest fossil found, dating from the Late Eocene
Florissant Formation
The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
.
All other extinct species in the family are known from fossils preserved in
Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
.
Based on the fossil record of the family, Li ''et al.'' (2017) infer that the family originated during the
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987.
In European lithostratigraphy, the name ...
or
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
.
Taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of the family as outlined by
Michael S. Engel and Jaime Ortega-Blanco in 2011:
Subfamilies and Tribes
*Subfamily
Schlettereriinae Orfila
**Tribe †
Phoriostephanini Engel & Huang
***Genus
†''
Phoriostephanus''
Engel & Huang Burmese amber
Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Late Cretaceous (
Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
)
**Tribe Schlettereriini
Orfila
***Genus
†''
Archaeostephanus''
Engel & Grimaldi New Jersey amber
New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of th ...
, Late Cretaceous (
Turonian
The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
)
***Genus †''
Kronostephanus''
Engel & Grimaldi Burmese amber
Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Late Cretaceous (
Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
)
***Genus ''
Schlettererius''
Ashmead
*Subfamily †
Electrostephaninae Engel
**Genus †''
Electrostephanus''
Brues Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
,
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
***Subgenus †''
Electrostephanodes''
Engel & Ortega-Blanco
***Subgenus †''
Electrostephanus''
Brues
*Subfamily
Stephaninae Leach
**Tribe
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
***Genus †''
Protostephanus
''Protostephanus'' is an extinct genus of crown wasp in the Hymenoptera family Stephanidae known from an Eocene fossil found in the United States of America. The genus contains a single described species, ''Protostephanus ashmeadi'' placed in th ...
''
Cockerell Baltic amber, Eocene
***Genus †''
Denaeostephanus''
Engel & Grimaldi Baltic amber, Eocene
***Genus †''
Lagenostephanus''
Li et al. Burmese amber, Myanmar, Late Cretaceous (Cenomanian)
**Tribe Stephanini
Leach
***Genus ''
Stephanus (insect)
''Stephanus'' is a genus of parasitoid wasps in the family Stephanidae. Records of species are from Europe and Asia.
Species
The following list may be incomplete:
* ''Stephanus anijimensis'' Watanabe & van Achterberg, 2014
* ''Stephanus antinor ...
''
Jurine
**Tribe Megischini
Engel & Grimaldi
***Genus ''
Hemistephanus''
Enderlein
***Genus ''
Megischus
''Megischus'' is a genus of crown wasp. It was circumscription (taxonomy), circumscribed by Gaspard Auguste Brullé in 1846. The genus has a cosmopolitan distribution, and over eighty species are recognized.
It is the type genus of the tribe (bio ...
''
Brullé
***Genus ''
Pseudomegischus''
Achterberg
****Subgenus ''
Pseudomegischus''
Achterberg
****Subgenus ''
Callomegischus''
Achterberg
**Tribe Foenatopodini
Enderlein
***Subtribe Madegafoenina
Engel & Grimaldi
****Genus ''
Madegafoenus''
Benoit
****Genus ''
Afromegischus''
Achterberg
***Subtribe Foenatopodina
Enderlein
****Genus ''
Parastephanellus''
Enderlein
****Genus ''
Comnatopus''
Achterberg
****Genus ''
Profoenatopus''
Achterberg
****Genus ''
Foenatopus''
Smith
References
{{Taxonbar, from=Q140717
Apocrita families