Xyelidae
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The Xyelidae are a comparatively species-poor family of sawflies comprising about 80
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
species in five genera worldwide and is the only family in the superfamily Xyeloidea.Taeger, A., Liston, A.D., Prous, M., Groll, E.K., Gehroldt, T. & Blank S.M. 2018: ECatSym – Electronic World Catalog of Symphyta (Insecta, Hymenoptera). Program version 5.0 (19 Dec 2018), data version 40 (23 Sep 2018). Senckenberg Deutsches Entomologisches Institut (SDEI), Müncheberg.
/ref> The fossil record of the family is extensive, comprising more than 120 species and including the oldest fossil
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 ...
species dating back to the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, between 245 and 208 million years ago. Xyelidae are to be regarded as
living fossils A living fossil is an extant taxon that cosmetically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of origin of the extant clade. Living fossi ...
since they represent one of the oldest lineages of insects and include still extant forms. The extant species occur in the Northern Hemisphere, especially in
boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
regions of the
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical region ...
, though there are a few
Oriental The Orient is a term for the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of ''Occident'', the Western World. In English, it is largely a metonym for, and coterminous with, the ...
species. Two genera and about 15 species occur in Europe.Taeger, A., Blank, S.M. & Liston, A. D. 2006: European Sawflies (Hymenoptera: Symphyta) – A Species Checklist for the Countries. Pp. 399-504. In: Blank, S.M., Schmidt, S. & Taeger, A. (eds): Recent Sawfly Research: Synthesis and Prospects. Goecke & Evers, Keltern.Blank, S.M. 2002: Biosystematics of the extant Xyelidae with particular emphasis on the Old World taxa (Insecta: Hymenoptera). Dissertation, Freie Universität Berlin. Considering additional fossil records from Australia, South Africa and Argentina, the extant species display a relict distribution. The species in the subfamily Xyelinae are associated with
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s (esp. '' Pinus'' and ''
Abies Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family Pinaceae. They are found on mountains throughout much of North and Central America, Europe, Asia, and North Africa. The genus is most closely related to ...
''), where the
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 feed on
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
or within buds. The larvae of the subfamily Macroxyelinae feed on the leaves of
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
trees (various Juglandaceae and ''
Ulmus Elms are deciduous and semi-deciduous trees comprising the flowering plant genus ''Ulmus'' in the plant family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical-montane regions of Nor ...
''). The family is characterized by the appendages of the head, which are remarkable in that the antennae and
palpi Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ...
are nearly leg-like in structure, with a long basal segment followed by a series of tiny segments, as in the tibia-tarsus.


Description


Imago (Adult)

Xyelidae are small Hymenoptera. Most species are 3 to 5 mm long, but species of '' Macroxyela'' and ''
Megaxyela ''Megaxyela'' is a genus of sawflies in the family Xyelidae. There are about 13 described species in ''Megaxyela'', found in the eastern Nearctic and in the southeastern part of East Asia. Two fossil species have been discovered, in Colorado and S ...
'' of East Asia and North America are larger, measuring 10 to 15 mm. The imagines display a number of ancestral characters of Hymenoptera, which may be absent in more derived lineages of Hymenoptera. Those include the absence of a wasp waist (thorax and abdomen abut without constriction), presence of cenchri on the
metathorax The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
to fix the wings at rest, presence of an antennal grooming apparatus on
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
and first
tarsomere The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip, plur ...
of the fore leg, and presence of a molar tooth on the mandible. Most intriguing is the morphology of the
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
which bears a long and thick third article followed by a number of shorter and more slender
antennomeres Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for Sensory system, sensing in arthropods. Antennae are connected to the first one or two Segmentation (biology), segments of the arthropod head. They vary widel ...
. This so-called synantennomere 3 is the product from the
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the st ...
fusion of several antennal articles, and it is unique among the extant Hymenoptera species. In '' Pleroneura'', '' Xyelecia'' and most species of ''
Xyela ''Xyela'' is a genus of sawflies, belonging to the family Xyelidae. The genus was described in 1819 by Dalman. The genus has cosmopolitan distribution. List of species Subgenus ''Xyela'' * ''Xyela alberta'' (Curran, 1923) * ''Xyela altenhofe ...
'' the maxillary palps are strongly enlarged and bear specialized
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
on the distal articles. The wing venation is the most complete among Hymenoptera: Only in Xyelidae the radial sector ''Rs'' furcates into the veins ''Rs1'' and ''Rs2'', while in other Hymenoptera ''Rs1'' is absent. The females bear a more or less long
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 ...
, which in some species of ''Xyela'' may be as long as the body. Morphology of the ovipositor and the ovipositor sheath are important characters for identification to species level. The penis valves of the males are densely setulous, which is a rare character state among the basal lineages of Hymenoptera. Females and males mate with the bodies directing in opposite direction. In Xyelinae the genital capsule of the males are revolved for 180° after disclosure from the pupal skin ( strophandry). Macroxyelinae are orthandrous after emergence. They mate in the same position as Xyelinae, but the male genital capsule is rotated yet in course of mating (facultative strophandry).


Larva

Like in many other sawflies, the
larvae 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 ...
of Xyelidae are superficially similar to caterpillars of
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
(" eruciform" type of larvae). Larvae of species feeding inside plants are whitish, those of free-feeding species whitish green or yellow. Larvae of ''Megaxyela'' bear a conspicuous pattern of black spots (see plate 21 figure 3 inTaeger, A.; Blank, S.M.; Liston, A.D. 2010: World Catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1-1064.) or they resemble bird droppings.Shinohara, A., Hara, H., Kramp, K., Blank, S. M. & Kameda, Y. 2017: Bird droppings on chestnut leaves or sawfly larvae: DNA barcodes verify the occurrence of the archaic ''Megaxyela togashii'' (Hymenoptera, Xyelidae) in Hokkaido, Japan. Zootaxa 4221(2): 220-232. The roundish head capsule bears a larval eye (
stemma Stemma (plural stemmata) may refer to: * In stemmatics, an approach to textual criticism, a stemma or stemma codicum is a diagram showing the relationships of the various versions of a text to earlier versions or manuscripts * Tree-like diagrams ...
) on each side, which is reduced in mining species, and short antennae comprising five articles. The
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
bears short legs comprising three articles. Unlike the larvae of all other Hymenoptera and Lepidoptera which lack
prolegs A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few other types of ins ...
on at least the first abdominal segment, larvae of Xyelidae have prolegs on all abdominal segments. In free feeding Xyelidae (''Macroxyela'', ''Megaxyela'') the abdominal prolegs are conspicuous and consist of two articles, while in the mining species (''Pleroneura'', ''Xyela'') they are reduced to inconspicuous transverse bulges.


Pupa

Unique among the Hymenoptera, the
pupa A pupa ( la, pupa, "doll"; plural: ''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 ...
of Xyelidae are of the decticous and exarate types, in which the antennae, legs and mandibles are free and mobile. This stage represents the already developed (
pharate A pupa ( la, pupa, "doll"; plural: ''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 ...
) imago which is still enclosed by the pupal skin. At this stage, the wings are not expanded, and in females, the ovipositor and its sheath curve dorsally above the tip of the abdomen. The decticous pupae are capable of biting open the cocoon, digging to the surface of the ground, moving around on the surface, and drinking (figs 22-23 in). Burdick, D.J. 1961: A taxonomic and biological study of the genus ''Xyela'' Dalman in North America. University of California Publications in Entomology 17(3): 285-355.This pupal form is an ancestral character of endopterygote insects. In other orders, it occurs in including
Neuropterida The Neuropterida are a clade, sometimes placed at superorder level, of holometabolous insects with over 5,700 described species, containing the orders Neuroptera (lacewings, antlions), Megaloptera (alderflies, dobsonflies), and Raphidioptera (sna ...
, Mecoptera, Trichoptera and in the basal lineages of
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
. The Hymenoptera excluding Xyelidae are characterized by immobile adecticous pupae.


Ecology

The larvae of all Xyelidae are phytophagous and associated with trees. Larvae of the comparatively species-rich ''Xyela'' live inside the growing staminate cones of
pines A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
and feed on the sporophylls and the pollen. The North American '' Xyela gallicaulis'' is exceptional in causing galls on fresh shoots of some pine species, inside which the larva feeds. Larvae of ''Pleroneura'' feed inside young shoots of
fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...
s. Only the Japanese '' Pleroneura piceae'' is associated with
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
. Larvae of Macroxyelinae are free feeders of deciduous tree species. The two North American species of ''Macroxyela'' feed on
elm Elms are deciduous and semi-deciduous trees comprising the flowering plant genus ''Ulmus'' in the plant family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical-montane regions of North ...
s, the East Asian and North American species of ''Megaxyela'' on Juglandaceae like walnuts,
hickory Hickory is a common name for trees composing the genus ''Carya'', which includes around 18 species. Five or six species are native to China, Indochina, and India (Assam), as many as twelve are native to the United States, four are found in Mexi ...
and
wingnuts ''WingNuts: Temporal Navigator'' is a 2001 top-down multi-directional shooter for Macintosh made by Freeverse Software. It was inspired by arcade games ''Time Pilot'' and ''1942.'' In ''WingNuts'', the player navigates an aircraft around a two-di ...
.Shinohara, A. 1992: The Sawfly Genus ''Megaxyela'' (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 60(4): 783-796. For '' Xyelecia nearctica'' an endophagous life style and an association with firs is supposed. Many species of Xyelidae are host specific, with only a single larval host plant known.
Monophagy Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
is the prevailing life style in most species of ''Xyela''. For this genus, the date of oviposition is closely correlated with the development of the staminate cones of the host pines, which might prevent a host shift in many cases. For some North American species of ''Xyela'' the association with each several species of pines (
oligophagy Oligophagy refers to the eating of only a few specific foods, and to monophagy when restricted to a single food source. The term is usually associated with insect dietary behaviour.morphologically similar species of ''Xyela'' properly. Through molecular access, true oligophagy could be demonstrated recently for '' Xyela bakeri'' (associated at least with '' Pinus contorta'', ''
Pinus ponderosa ''Pinus ponderosa'', commonly known as the ponderosa pine, bull pine, blackjack pine, western yellow-pine, or filipinus pine is a very large pine tree species of variable habitat native to mountainous regions of western North America. It is the ...
'' and ''
Pinus sabiniana ''Pinus sabiniana'' (sometimes spelled ''P. sabineana''), with vernacular names including towani pine, foothill pine, gray pine, bull pine, and digger pine, is a pine endemic to California in the United States. Some sources discourage using the n ...
'') and '' Xyela brunneiceps'' (associated with ''
Pinus flexilis ''Pinus flexilis'', the limber pine, is a species of pine tree-the family Pinaceae that occurs in the mountains of the Western United States, Mexico, and Canada. It is also called Rocky Mountain white pine. A limber pine in Eagle Cap Wilderness ...
'' and ''
Pinus strobiformis ''Pinus strobiformis'', commonly known as southwestern white pine, Mexican white pine or Chihuahua white pine, is a medium-sized white pine tree whose native habitat is in southwestern United States and Mexico. It is typically a high-elevation p ...
''). Lack of reliable, reproducible host data could cause inaccurate reports of monophagy or oligophagy in many taxa of Xyelidae, but at least some of the species included in ''Megaxyela'' and ''Pleroneura'' are truly oligophagous. After completing feeding, Xyelidae larvae dig into the ground and form an earthen cell, where they usually spin a cocoon and develop into a pupa. Absence of a cocoon has been observed in ''Megaxyela togashii'' from Japan. The imago emerges in the following spring to mate. Subsequently, the female deposits eggs with the ovipositor. In most species of ''Xyela'', the tip of valvula 3 of the ovipositor sheath is equipped with specialized sensory structures called sensilla trichodea and sensilla campaniformia, which are involved in the oviposition process. In ''Pleroneura'', unlike practically all other Hymenoptera, the hard and conical ovipositor sheath is used in addition to the ovipositor proper to penetrate the resinous buds of firs. '' Megaxyela gigantea'' and most other species of ''Megaxyela'' has strikingly long hind legs which are used to fold a newly growing leaf to provide shelter for their eggs which are fixed with glutinous material between on the upper sides of the folded leaf.Saito, K., 1941: in dendro-entomologischer Beitrag.(In Japanese.) Bull. agric. For. Coll. Suigen, (6): 1-235. Many species of Xyelidae facultatively
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
for several years. In '' Xyela alpigena'' and '' Xyela obscura'' a diapause of at least two years is obligate. This strategy might have evolved to ensure well developed cones are available for oviposition: the host plants ''
Pinus cembra ''Pinus cembra'', also known as Swiss pine, Swiss stone pine or Arolla pine or Austrian stone pine or just stone pine, is a species of a pine tree in the subgenus ''Strobus''. Description The Swiss pine is a member of the white pine group, ...
'' and '' Pinus mugo'' produce cones very irregularly in the subalpine zone of the
European Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Sw ...
.Pschorn-Walcher, H. & Altenhofer, E. 2000: Langjährige Larvenaufsammlungen und Zuchten von Pflanzenwespen (Hymenoptera, Symphyta) in Mitteleuropa. Linzer biologische Beiträge 32(1): 273-327 Adults of ''Xyela'' are pollen feeders of a variety of plants, which bear flowers with easily accessible pollen (e.g., '' Betula'', ''
Cercocarpus ''Cercocarpus'', commonly known as mountain mahogany, is a small genus of at least nine species of nitrogen-fixing flowering plants in the rose family, Rosaceae. They are native to the western United States and northern Mexico, where they grow i ...
'', '' Ostrya'', '' Pinus'', ''
Purshia ''Purshia'' (bitterbrush or cliff-rose) is a small genus of 5–8 species of flowering plants in the family Rosaceae which are native to western North America. Description ''Purshia'' species form deciduous or evergreen shrubs, typically reach ...
'', ''
Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...
'', '' Salix).'' The enlarged maxillary palps of most ''Xyela'' (and supposedly also of ''Pleroneura'' and ''Xyelecia'') serve for the extraction of pollen from the flowers. Although they feed upon often economically important tree species, Xyelidae are usually of only small significance as pest organisms. ''Pleroneura piceae'' damages the growth of Sakhalin spruce, since the larvae destroy the young shoots. Larvae of ''Megaxyela major'' (and supposedly also of ''Megaxyela langstoni'') feed on leaves of
pecan The pecan (''Carya illinoinensis'') is a species of hickory native to the southern United States and northern Mexico in the region of the Mississippi River. The tree is cultivated for its seed in the southern United States, primarily in Georgia, ...
and are regarded as a pest of pecan plantations in the Southeastern USA.


Systematics and taxonomy

Xyelidae represent the most basal lineage of Hymenoptera and very likely the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of all other extant Hymenoptera. This assertion is supported by
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses of both morphological characters and DNA sequences. The great age of the family is supported by numerous fossil records. All Hymenoptera recorded from the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
are classified as Xyelidae, while representatives of other hymenopterous families have been found no earlier than the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. During the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
and 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 Xyelidae obviously were much more species-rich and more widely distributed than today. Thus, the comparatively few extant species can be regarded a relict group.


Extant taxa

Th
Electronic World Catalog of Symphyta
ref name=ECatSym /> provides a complete account of the valid extant genera and species, their synonyms, the concerning references to original descriptions and distribution data on the level of countries and provinces. * Xyelinae **'' Pleroneura'' Konow, 1897: at least eight species in EurasiaShinohara, A. 1995: The Sawfly Genus ''Pleroneura'' (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 63(4): 825-840Blank, S.M. 2002: The Western Palaearctic Xyelidae (Hymenoptera). Pp. 197-233. In: Viitasaari, M. (ed.): Sawflies (Hymenoptera, Symphyta) I. A review of the suborder, the Western Palaearctic taxa of Xyeloidea and Pamphilioidea. Tremex, Helsinki.
read online
Shinohara, A. 2016: The sawfly genus ''Pleroneura'' (Hymenoptera, Xyelidae) of Japan: ''P. itoi'' n. sp. and a key to species. Zootaxa 4121(4): 495-500. and five in North AmericaSmith, D.R., Ohmart, C.P. & Dahlsten, D.L. 1977: The fir shoot-boring sawflies of the genus ''Pleroneura'' in North America (Hymenoptera: Xyelidae). Annals of the Entomological Society of America 70: 761-767. **''
Xyela ''Xyela'' is a genus of sawflies, belonging to the family Xyelidae. The genus was described in 1819 by Dalman. The genus has cosmopolitan distribution. List of species Subgenus ''Xyela'' * ''Xyela alberta'' (Curran, 1923) * ''Xyela altenhofe ...
'' Brébisson, 1819 (synonyms: ''Pinicola'' Dalman, 1818 reoccupied by the genus-group name of birds ''Pinicola'' Vieillot, 1808">Pinicola.html" ;"title="reoccupied by the genus-group name of birds ''Pinicola">reoccupied by the genus-group name of birds ''Pinicola'' Vieillot, 1808 ''Xyelatana'' Benson, 1938) ***''Xyela'' ('' Pinicolites'') Meunier, 1920: ''Xyela lata'' D.R. Smith, 1990 in western North America ***''Xyela'' (''
Xyela ''Xyela'' is a genus of sawflies, belonging to the family Xyelidae. The genus was described in 1819 by Dalman. The genus has cosmopolitan distribution. List of species Subgenus ''Xyela'' * ''Xyela alberta'' (Curran, 1923) * ''Xyela altenhofe ...
'') Brébisson, 1819: 29 species in EurasiaBlank, S.M., Kramp, K. & Shinohara, A. 2017: ''Xyela fusca'' spec. nov. from Japan elucidates East Asian–North American relationships of ''Xyela'' (Hymenoptera, Xyelidae). Zootaxa 4303(1): 103–121. and 20 in North America,Blank, S.M. & Kramp, K. 2017: ''Xyela davidsmithi'' (Hymenoptera, Xyelidae), a New Pine Catkin Sawfly with an Unusual Host Association from the Sierra Nevada. Proceedings of the Entomological Society of Washington 119: 703- 717. but many additional species have not yet been described. * Macroxyelinae **'' Macroxyela'' W.F. Kirby, 1882: '' Macroxyela aenea'' (Norton, 1872) und '' Macroxyela ferruginea'' (Say, 1824) in eastern and central North AmericaSmith, D. R. & Schiff, N. M. 1998: The genera ''Macroxyela'' Kirby and ''Megaxyela'' Ashmead (Hymenoptera: Xyelidae) in North America. - Proceedings of the entomological Society of Washington 100(4): 636-657. **''
Megaxyela ''Megaxyela'' is a genus of sawflies in the family Xyelidae. There are about 13 described species in ''Megaxyela'', found in the eastern Nearctic and in the southeastern part of East Asia. Two fossil species have been discovered, in Colorado and S ...
'' Ashmead, 1898: six species in eastern Eurasia,Blank, S.M., Kramp, K., Smith, D.R., Sundukov, Y.N., Wei, M. & Shinohara, A. 2017: Big and beautiful: the ''Megaxyela'' species (Hymenoptera, Xyelidae) of East Asia and North America. European Journal of Taxonomy 348: 1–46. seven species in North America. **'' Xyelecia'' Ross, 1932: two species, '' Xyelecia japonica'' Togashi, 1972 in JapanShinohara, A. 1998: Collection records of two rare xyelid sawflies (Hymenoptera, Symphyta) in Japan. Japanese Journal of Systematic Entomology 4: 389-390. and '' Xyelecia nearctica'' in Western North AmericaSmith, D. R. 1964: Description of the Male of ''Xyelecia nearctica'', with Comments on the Genus (Hymenoptera: Xyelidae). Pan-Pacific Entomologist 40(1): 54-56.Smith, D.R. 1967: A review of the larvae of Xyelidae, with notes on the family classification (Hymenoptera). Annals of the Entomological Society of America 60: 376-384. The European species can be identified with help of ''„The Western Palaearctic Xyelidae“'' of Blank (2002), all Eurasian species of ''Xyela'' with Blank et al. (2013).Blank, S.M., Shinohara, A. & Altenhofer, E. 2013: The Eurasian species of ''Xyela'' (Hymenoptera, Xyelidae): taxonomy, host plants and distribution. Zootaxa 3629: 1-10
PDF (Abstract)
The North American Macroxyelinae were revised by Smith & Schiff (1998), the North American species of ''Xyela'' by Burdick (1961), the North American species of ''Pleroneura'' by Smith et al. (1977), the East Asian species of ''Megaxyela'' by Shinohara (1992), the East Asian species of ''Pleroneura'' by Shinohara (1995), and the species of ''Megaxyela'' of the World by Blank et al. (2017).


Extinct taxa

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Electronic World Catalog of Symphyta
ref name=ECatSym /> provides a complete account of the valid fossil genera and species, their synonyms and the concerning references to original descriptions. Genus without classification into a subfamily of Xyelidae **†'' Potrerilloxyela'' Lara, Rasnitsyn & Zavattieri, 2014, 1 fossil species † Archexyelinae **†'' Archexyela'' Riek, 1955, 2 fossil species **†'' Asioxyela'' Rasnitsyn, 1964, 4 fossil species **†'' Dinoxyela'' Rasnitsyn, 1969, 1 fossil species **†'' Euryxyela'' Rasnitsyn, 1964, 1 fossil species **†'' Ferganoxyela'' Rasnitsyn, 1969, 2 fossil species **†'' Leioxyela'' Rasnitsyn, 1969, 5 fossil species **†'' Lithoxyela'' Rasnitsyn, 1969, 1 fossil species **†'' Madygenius'' Rasnitsyn, 1969, 2 fossil species **†'' Moltenia'' Schlüter, 2000, 1 fossil species **†'' Oryctoxyela'' Rasnitsyn, 1969, 2 fossil species **†'' Triassoxyela'' Rasnitsyn, 1964, 3 fossil species **†'' Xaxexis'' Pagliano & Scaramozzino, 1990 ''Euryxyela'' Hong, 1984 1 fossil species **†'' Xiphoxyela'' Rasnitsyn, 1969, 2 fossil species **†'' Xyelinus'' Rasnitsyn, 1964, 2 fossil species Macroxyelinae *Genera without classification into a tribe of Macroxyelinae **†'' Bolboxyela'' Rasnitsyn, 1990, 1 fossil species **†'' Brachyoxyela'' Gao, Zhao & Ren, 2011, 2 fossil species *† Angaridyelini **†'' Angaridyela'' Rasnitsyn, 1966, 8 fossil species **†'' Baissoxyela'' Rasnitsyn, 1969, 1 fossil species **†'' Ceratoxyela'' J. Zhang & X. Zhang, 2000, 1 fossil species **†'' Lethoxyela'' J. Zhang & X. Zhang, 2000, 2 fossil species **†'' Liaoxyela'' J. Zhang & X. Zhang, 2000, 1 fossil species **†'' Nigrimonticola'' Rasnitsyn, 1966, 1 fossil species **†'' Ophthalmoxyela'' Rasnitsyn, 1966, 1 fossil species *† Ceroxyelini **†'' Ceroxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Isoxyela'' J. Zhang & X. Zhang, 2000, 1 fossil species **†'' Sinoxyela'' J. Zhang & X. Zhang, 2000, 1 fossil species *† Gigantoxyelini **†'' Abrotoxyela'' Gao, Ren & Shih, 2009, 2 fossil species **†'' Chaetoxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Chionoxyela'' Rasnitsyn, 1993, 1 fossil species **†'' Gigantoxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Heteroxyela'' J. Zhang & X. Zhang, 2000, 1 fossil species **†'' Platyxyela'' Wang, Shih & Ren, 2012, 1 fossil species **†'' Shartexyela'' Rasnitsyn, 2008, 1 fossil species * Macroxyelini **†'' Anthoxyela'' Rasnitsyn, 1977, 4 fossil species **''
Megaxyela ''Megaxyela'' is a genus of sawflies in the family Xyelidae. There are about 13 described species in ''Megaxyela'', found in the eastern Nearctic and in the southeastern part of East Asia. Two fossil species have been discovered, in Colorado and S ...
'' Ashmead, 1898, 2 fossil species in addition to extant species * Xyeleciini **†'' Microxyelecia'' Rasnitsyn, 1969, 1 fossil species **†'' Uroxyela'' Rasnitsyn, 1966, 1 fossil species **'' Xyelecia'' Ross, 1932, 1 fossil species in addition to extant species **†'' Xyelites'' Rasnitsyn, 1966, 2 fossil species † Madygellinae **†'' Chubakka'' Kopylov, 2014, 1 fossil species **†'' Madygella'' Rasnitsyn, 1969, 5 fossil species **†'' Samarkandykia'' Kopylov, 2014, 2 fossil species Xyelinae *† Liadoxyelini **†'' Aequixyela'' Wang, Rasnitsyn & Ren, 2014, 1 fossil species **†'' Anomoxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Cathayxyela'' Wang, Rasnitsyn & Ren, 2014, 1 fossil species **†'' Kirghizoxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Liadoxyela'' Martynov, 1937, 3 fossil species **†'' Lydoxyela'' Rasnitsyn, 1966, 1 fossil species **†'' Orthoxyela'' Rasnitsyn, 1983, 1 fossil species * Xyelini **†'' Enneoxyela'' Rasnitsyn, 1966, 4 fossil species **†'' Eoxyela'' Rasnitsyn, 1965, 5 fossil species **†'' Spathoxyela'' Rasnitsyn, 1969, 2 fossil species **''
Xyela ''Xyela'' is a genus of sawflies, belonging to the family Xyelidae. The genus was described in 1819 by Dalman. The genus has cosmopolitan distribution. List of species Subgenus ''Xyela'' * ''Xyela alberta'' (Curran, 1923) * ''Xyela altenhofe ...
'' Dalman, 1819, 7 fossil species in addition to extant species, including ''Xyela'' ('' Pinicolites'') Meunier, 1920 **†'' Xyelisca'' Rasnitsyn, 1969, 1 fossil species **†'' Yanoxyela'' Ren, Lu, Guo & Ji, 1995, 1 fossil species


References

{{Taxonbar, from=Q147974 Sawfly families Sawflies