The Xyelidae are a comparatively species-poor family of

sawflies Sawflies are wasp-like insects that are in the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plant ...

, comprising about 80

extant Extant or Least-concern species, least concern 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 * Exta ...

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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

of the family is extensive, comprising more than 120 species and including the oldest fossil

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 ...

species dating back to the

Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...

, between 245 and 208 million years ago. Xyelidae are the most basal contemporary lineage of Hymenoptera and have many ancestral morphological features. The extant species occur in the Northern Hemisphere, especially in boreal 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 reg ...

, though there are a few

Oriental The Orient is a term referring to the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of the term ''Occident'', which refers to the Western world. In English, it is largely a meto ...

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 A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...

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 sin ...

s (esp. ''

Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...

'' and ''

Abies Firs are evergreen coniferous trees belonging to the genus ''Abies'' () in the family Pinaceae. There are approximately 48–65 extant species, found on mountains throughout much of North and Central America, Eurasia, and North Africa. The genu ...

''), 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 ...

e feed on

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

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 Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...

trees (various

Juglandaceae The Juglandaceae are a plant family known as the walnut family. They are trees, or sometimes shrubs, in the order Fagales. Members of this family are native to the Americas, Eurasia, and Southeast Asia. The nine or ten genera in the family have ...

and ''

Ulmus Elms are deciduous and semi-deciduous trees comprising the genus ''Ulmus'' in the family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical- montane regions of North America and Eurasia, ...

''). The family is characterized by the appendages of the head, which are remarkable in that the antennae and palpi 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 East Asia is a geocultural region of Asia. It include ...

'' 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 Wasp waist is a women's fashion silhouette, produced by a style of corset and girdle, that has experienced various periods of popularity in the 19th and 20th centuries. Its primary feature is the abrupt transition from a natural-width rib cag ...

(thorax and abdomen abut without constriction), presence of cenchri on the metathorax to fix the wings at rest, presence of an antennal grooming apparatus on

tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...

and first tarsomere of the fore leg, and presence of a

molar tooth The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone toot ...

on the mandible. Most intriguing is the morphology of the antenna which bears a long and thick third article followed by a number of shorter and more slender antennomeres. 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 stu ...

fusion of several antennal articles, and it is unique among the extant Hymenoptera species. In '' Pleroneura'', '' Xyelecia'' and most species of '' Xyela'' the maxillary palps are strongly enlarged and bear specialized

seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...

e 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 typica ...

, 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

e of Xyelidae are superficially similar to

caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...

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 ...

(" 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) on each side, which is reduced in mining species, and short antennae comprising five articles. The

thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...

bears short legs comprising three articles. Unlike the larvae of all other Hymenoptera and Lepidoptera which lack

proleg 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 (biology), order Lepidoptera, though they can also be found on larvae of insects such as symphyta, sawflies. In ...

s 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 (; : 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 th ...

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 (; : 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 ...

) 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 holometabolan insects. In other orders, it occurs in including

Neuropterida The Neuropterida are a clade of holometabolous insects, sometimes placed at the superorder level. The clade contains the orders Neuroptera (lacewings, antlions), Megaloptera (alderflies, dobsonflies), and Raphidioptera (snakeflies), and includes ...

Mecoptera Mecoptera (from the Greek language, Greek: ''mecos'' = "long", ''ptera'' = "wings") is an Order (biology), order of insects in the superorder Holometabola with about six hundred species in nine Family (biology), families worldwide. Mecopterans a ...

Trichoptera The caddisflies (order Trichoptera) are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the basis ...

and in the basal lineages of

. 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

pine 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...

s 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 are evergreen coniferous trees belonging to the genus ''Abies'' () in the family Pinaceae. There are approximately 48–65 extant species, found on mountains throughout much of North and Central America, Eurasia, and North Africa. The genu ...

s. Only the Japanese '' Pleroneura piceae'' is associated with

spruce A spruce is a tree of the genus ''Picea'' ( ), a genus of about 40 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal ecosystem, boreal (taiga) regions of the Northern hemisphere. ''Picea'' ...

. Larvae of Macroxyelinae are free feeders of deciduous tree species. The two North American species of ''Macroxyela'' feed on elms, the East Asian and North American species of ''Megaxyela'' on

walnuts A walnut is the edible seed of any tree of the genus ''Juglans'' (family Juglandaceae), particularly the Persian or English walnut, '' Juglans regia''. They are accessory fruit because the outer covering of the fruit is technically an invo ...

hickory Hickory is a common name for trees composing the genus ''Carya'', which includes 19 species accepted by ''Plants of the World Online''. Seven species are native to southeast Asia in China, Indochina, and northeastern India (Assam), and twelve ...

and wingnuts.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 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 contorta'', with the common names lodgepole pine and shore pine, and also known as twisted pine, and contorta pine, is a common tree in western North America. It is common near the ocean shore and in dry montane forests to the subalpine ...

'', '' Pinus ponderosa'' and '' Pinus sabiniana'') and '' Xyela brunneiceps'' (associated with '' Pinus flexilis'' and '' Pinus strobiformis''). 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'' and '' Pinus mugo'' produce cones very irregularly in the

subalpine Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...

zone of the European Alps.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 A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' contains 3 ...

'', '' Cercocarpus'', ''

Ostrya ''Ostrya'' is a genus of eight to 10 small deciduous trees belonging to the birch family Betulaceae. Common names include hop-hornbeam and hophornbeam. It may also be called ironwood, a name shared with a number of other plants. The genus is na ...

'', ''

'', '' Purshia'', ''

Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...

'', ''

Salix Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...

).'' 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 primarily in the U.S. states of 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 ...

of all other extant Hymenoptera. This assertion is supported by

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

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

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 143.1 Mya. ...

. During the

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

and the

Tertiary Tertiary (from Latin, meaning 'third' or 'of the third degree/order..') may refer to: * Tertiary period, an obsolete geologic period spanning from 66 to 2.6 million years ago * Tertiary (chemistry), a term describing bonding patterns in organic ch ...

, the Xyelidae obviously were much more species-rich and more widely distributed than today. Thus, the comparatively few extant species can be regarded a

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'' 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'') 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. **''

'' 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

Th
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 **''

'' 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'' 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

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