Sawflies are the insects of 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 plants where they lay their eggs. The name is associated
especially with the Tenthredinoidea, by far the largest superfamily,
with about 7,000 known species; in the entire suborder, there are
8,000 described species in more than 800 genera. The suborder Symphyta
is paraphyletic, consisting of several basal groups within the order
The primary distinction between sawflies and their relatives the
Apocrita – the ants, bees, and wasps – is that the adults lack a
"wasp waist", and instead have a broad connection between the abdomen
and the thorax. Some sawflies are Batesian mimics of wasps and bees,
and the ovipositor can be mistaken for a stinger.
Sawflies vary in
length, most measuring 2.5 millimetres (0.1 in) to 20 millimetres
(0.8 in); the largest known sawfly measured 55 millimetres
(2.2 in). The larvae are caterpillar-like, but can be
distinguished by the number of prolegs and the absence of crochets in
sawfly larvae. The great majority of sawflies are plant-eating, though
the members of the superfamily
Orussoidea are parasitic. The adults
feed on pollen and nectar. Predators include birds, insects and small
animals. The larvae of some species have anti-predator adaptations
such as regurgitating irritating liquid and clustering together for
safety in numbers.
Sawflies are hosts to many parasitoids, most of
which are Hymenoptera, the rest being Diptera.
Adult sawflies are short-lived, with a life expectancy of 7–9 days,
though the larval stage can last from months to years, depending on
the species. Parthenogenetic females, which do not need to mate to
produce fertilised eggs, are common in the suborder, though many
species have males.
Sawflies go through a complete metamorphosis with
four distinct life stages – egg, larva, pupa and adult. The female
uses her ovipositor to drill into plant material (or, in the case of
Orussoidea, other insects) and then lays eggs in groups called rafts
or pods. After hatching, larvae feed on plants, often in groups. As
they approach adulthood, the larvae seek a protected spot to pupate,
typically in bark or the soil. Large populations of species such as
the pine sawfly can cause substantial damage to economic forestry,
while others such as the iris sawfly are important pests in
horticulture. Outbreaks of sawfly larvae can defoliate trees and may
cause dieback, stunting or death.
Sawflies can be controlled through
the use of insecticides, natural predators and parasites, or
Sawflies first appeared 250 million years ago in the Triassic. The
oldest superfamily, the Xyeloidea, has existed into the present.
Sawflies are distributed globally, though they are more diverse in the
5 Behaviour and ecology
5.2 Life cycle and reproduction
6 Relationship with humans
8 Further reading
9 External links
Sawfly laying eggs in a plant, using the serrated saw-like ovipositor
for which the group is named
The suborder name "Symphyta" derives from the Greek word symphyton,
meaning "grown together", referring to the group's distinctive lack of
a wasp waist between prostomium and peristomium. Its common name,
"sawfly", derives from the saw-like ovipositor that is used for
egg-laying, in which a female makes a slit in either a stem or plant
leaf to deposit the eggs. The first known use of this name was in
Sawflies are also known as "wood-wasps".
Carl Gerstäcker established the suborder Symphyta
In his original description of
Hymenoptera in 1863, German zoologist
Carl Gerstäcker divided them into three groups,
Hymenoptera apocrita and
Hymenoptera phytophaga. But four years
later in 1867, he described just two groups, H. apocrita syn. genuina
and H. symphyta syn. phytophaga. Consequently, the name Symphyta is
given to Gerstäcker as the zoological authority. In his description,
Gerstäcker distinguished the two groups by the transfer of the first
abdominal segment to the thorax in the Apocrita, compared to the
Symphyta. Consequently, there are only eight dorsal half segments in
the Apocrita, against nine in the Symphyta. The larvae are
distinguished in a similar way.
The Symphyta have therefore traditionally been considered, alongside
the Apocrita, to form one of two suborders of Hymenoptera.
Symphyta are the more primitive group, with comparatively complete
venation, larvae that are largely phytophagous, and without a
"wasp-waist", a symplesiomorphic feature. Together, the Symphyta make
up less than 10% of hymenopteran species. While the terms sawfly
and Symphyta have been used synonymously, the Symphyta have also been
divided into three groups, true sawflies (phyllophaga), woodwasps or
xylophaga (Siricidae), and Orussidae. The three groupings have been
distinguished by the true sawflies' ventral serrated or saw-like
ovipositor for sawing holes in vegetation to deposit eggs, while the
woodwasp ovipositor penetrates wood and the
Orussidae behave as
external parasitoids of wood-boring beetles. The woodwasps themselves
are a paraphyletic ancestral grade. Despite these limitations, the
terms have utility and are common in the literature.
While most hymenopteran superfamilies are monophyletic, as is
Hymenoptera, the Symphyta has long been seen to be
Cladistic methods and molecular phylogenetics
are improving the understanding of relationships between the
superfamilies, resulting in revisions at the level of superfamily and
family. The Symphyta are the most primitive (basal) taxa within
Hymenoptera (some going back 250 million years), and one of the
taxa within the Symphyta gave rise to the monophyletic suborder
Apocrita (wasps, bees, and ants). In cladistic analyses the
Orussoidea are consistently the sister group to the Apocrita.
The oldest unambiguous sawfly fossils date back to the Middle or Late
Triassic. These fossils, from the family Xyelidae, are the oldest of
all Hymenoptera. One fossil, Archexyela ipswichensis from
Queensland is between 205.6 and 221.5 million years of age, making it
among the oldest of all sawfly fossils. More Xyelid fossils have
been discovered from the Middle
Jurassic and the Cretaceous, but the
family was less diverse then than during the
Mesozoic and Tertiary.
The subfamily Xyelinae were plentiful during these time periods, in
Tertiary faunas were dominated by the tribe Xyelini; these are
indicative of a humid and warm climate.
The cladogram is based on Schulmeister 2003.
Symphyta within Hymenoptera
Cephoidea (stem sawflies)
Siricoidea (horntails or wood wasps)
Xiphydrioidea (wood wasps)
Orussoidea (parasitic wood wasps)
wasp waist 200mya
APOCRITA (ants, bees, wasps)
Symphyta (red bar) are paraphyletic as
Apocrita are excluded.
There are approximately 8,000 species of sawfly in more than 800
genera, although new species continue to be discovered.
However, earlier studies indicated that 10,000 species grouped into
about 1,000 genera were known. Early phylogenies such as that of
Alexandr Rasnitsyn, based on morphology and behaviour, identified nine
clades which did not reflect the historical superfamilies. Such
classifications were replaced by those using molecular methods,
starting with Dowton and Austin (1994). As of 2013, the Symphyta
are treated as nine superfamilies (one extinct) and 25 families. Most
sawflies belong to the
Tenthredinoidea superfamily, with about 7,000
Tenthredinoidea has six families, of which
Tenthredinidae is by far the largest with some 5,500 species.
Extinct taxa are indicated by a dagger (†).
Superfamilies and families
Anaxyeloidea Martynov, 1925
Anaxyelidae Martynov, 1925 (1 species) & †12 genera
Cephoidea Newman, 1834 (1 & †1family)
Cephidae Newman, 1834 (21 genera, 160 spp. & †3 genera
†Superfamily Karatavitoidea Rasnitsyn, 1963 (1 family)
Orussoidea Newman, 1834 (1 & †1 family)
Orussidae Newman, 1834 (16 genera, 82 spp.) & †3 genera
Pamphilioidea Cameron, 1890 (2 & †1 families) (syn.
Megalodontesidae Konow, 1897 (1 genera, 42 spp.) & †1
Pamphiliidae Cameron, 1890 (10 genera, 291 spp.) & †3
Siricoidea Billberg, 1820 (2 & †5 families)
Siricidae Billberg, 1820 (11 genera, 111 spp.) & †9
Tenthredinoidea Latreille, 1803 (6 & †2 families)
Argidae Konow, 1890 (58 genera, 897 spp.) and †1 genus
Blasticotomidae Thomson, 1871 (2 genera, 12 spp.) & †1
Cimbicidae W. Kirby, 1837 (16 genera, 182 spp.) & †6
Diprionidae Rohwer, 1910 (11 genera, 136 spp.) & †2
Pergidae Rohwer, 1911 (60 genera, 442 spp.)
Tenthredinidae Latreille, 1803 (400 genera, 5,500 spp.) &
Xiphydrioidea Leach, 1819
Xiphydriidae Leach, 1819 (28 genera, 146 spp.)
Xyeloidea Newman, 1834
Xyelidae Newman, 1834 (5 genera, 63 spp.) & †47genera
Giant horntail, Urocerus gigas, a Batesian mimic of a hornet,
ovipositing. It does not sting.
Many species of sawfly have retained their ancestral attributes
throughout time, specifically their plant-eating habits, wing veins
and the unmodified abdomen, where the first two segments appear like
the succeeding segments. The absence of the narrow wasp waist
distinguishes sawflies from other members of hymenoptera, although
some are Batesian mimics with coloration similar to wasps and bees,
and the ovipositor can be mistaken for a stinger. Most sawflies
are stubby and soft-bodied, and fly weakly.
Sawflies vary in
length: Urocerus gigas, which can be mistaken as a wasp due to its
black-and-yellow striped body, can grow up to 20 millimetres
(0.8 in) in length, but among the largest sawflies ever
discovered was Hoplitolyda duolunica from the Mesozoic, with a body
length of 55 millimetres (2.2 in) and a wingspan of 92
millimetres (3.6 in). The smaller species only reach
lengths of 2.5 millimetres (0.1 in).
European hornet is a wasp-waisted Apocritan with a sting, not a
Heads of sawflies vary in size, shape and sturdiness, as well as the
positions of the eyes and antennae. They are characterised in four
head types: open head, maxapontal head, closed head and genapontal
head. The open head is simplistic, whereas all the other heads are
derived. The head is also hypognathous, meaning that the lower
mouthparts are directed downwards. When in use, the mouthparts may be
directed forwards, but this is only caused when the sawfly swings its
entire head forward in a pendulum motion. Unlike most primitive
insects, the sutures (rigid joints between two or more hard elements
on an organism) and sclerites (hardened body parts) are obsolescent or
absent. The clypeus (a sclerite that makes up an insects "face") is
not divided into a pre- and postclypeus, but rather separated from the
front. The antennal sclerites are fused with the surrounding head
capsule, but these are sometimes separated by a suture. The number of
segments in the antennae vary from six in the Accorduleceridae to 30
or more in the Pamphiliidae. The compound eyes are large with a
number of facets, and there are three ocelli between the dorsal
portions of the compound eyes. The tentorium comprises the whole
inner skeleton of the head.
Three segments make up the thorax: the mesothorax, metathorax and
prothorax, as well as the exoskeletal plates that connect with these
segments. The legs have spurs on their fourth segments, the
Sawflies have two pairs of translucent wings. The fore and
hind wings are locked together with hooks. Parallel development in
sawfly wings is most frequent in the anal veins. In all sawflies, 2A
& 3A tend to fuse with the first anal vein. This occurs in several
families including Argidae,
Diprionidae and Cimbicidae.
Aposematically coloured caterpillar-like larva of Abia sericea
The larvae of sawflies are easily mistaken for lepidopteran larvae
(caterpillars). However, several morphological differences can
distinguish the two: while both larvae share three pairs of thoracic
legs and an apical pair of abdominal prolegs, lepidopteran
caterpillars have four pairs of prolegs on abdominal segments 3-6
while sawfly larvae have five pairs of prolegs located on abdominal
segments 2-6; crochets are present on lepidopteran larvae, whereas on
sawfly larvae they are not; the prolegs of both larvae gradually
disappear by the time they burrow into the ground, therefore making it
difficult to distinguish the two; and sawfly larvae only have a single
pair of minute eyes, whereas lepidopteran larvae have four to six eyes
on each side of the head.
Sawfly larvae behave like
lepidopteran larvae, walking about and eating foliage. Some groups
have larvae that are eyeless and almost legless; these larvae make
tunnels in plant tissues including wood. Many species of sawfly
larvae are strikingly coloured, exhibiting colour combinations such as
black and white while others are black and yellow. This is a warning
colouration because some larvae can secrete irritating fluids from
glands located on their undersides.
Sawflies are widely distributed throughout the world. The largest
family, the Tenthredinidae, with some 5,000 species, are found on all
continents except Antarctica, though they are most abundant and
diverse in the temperate regions of the northern hemisphere; they are
absent from New Zealand and there are few of them in Australia. The
next largest family, the Argidae, with some 800 species, is also
worldwide, but is commonest in the neotropics, especially in Africa,
where they feed on woody and herbaceous angiosperms. Of the other
Megalodontidae are Palearctic; the
Xyelidae, Pamphilidae, Diprionidae, Cimbicidae, and
Holarctic, while the
Siricidae are mainly
Holarctic with some tropical
species. The parasitic
Orussidae are found worldwide, mostly in
tropical and subtropical regions. The wood-boring
worldwide, but most species live in the subtropical parts of Asia.
Behaviour and ecology
Rose stem sawfly (Hartigia trimaculata) larva in a rose stem
Sawflies are mostly herbivores, feeding on plants that have a high
concentration of chemical defences. These insects are either resistant
to the chemical substances, or they avoid areas of the plant that have
high concentrations of chemicals. The larvae primarily feed in
groups; they are folivores, eating plants and fruits on native trees
and shrubs, though some are parasitic. However, this is not
always the case; Monterey pine sawfly (Itycorsia) larvae are solitary
web-spinners that feed on Monterey pine trees inside a silken web.
The adults feed on pollen and nectar.
Anti-predator adaptation: spitfire sawfly larvae grouped together for
safety in numbers
Sawflies are eaten by a wide variety of predators. While many birds
find the larvae distasteful, some such as the currawong (Strepera) and
stonechats (Saxicola) eat both adults and larvae. The larvae
are an important food source for the chicks of several birds,
Sawfly and moth larvae form one third of the
diet of nestling corn buntings (Emberiza calandra), with sawfly larvae
being eaten more frequently on cool days.
Black grouse (Tetrao
tetrix) chicks show a strong preference for sawfly larvae.
Sawfly larvae formed 43% of the diet of chestnut-backed chickadees
(Poecile rufescens). Small carnivorous mammals such as the masked
shrew (Sorex cinereus), the northern short-tailed shrew (Blarina
brevicauda) and the deer mouse (Peromyscus maniculatus) predate
heavily on sawfly cocoons. Insects such as ants and certain
species of predatory wasps (Vespula vulgaris) eat adult sawflies and
the larvae, as do lizards and frogs. Pardalotes, honeyeaters
and fantails (Rhipidura) occasionally consume laid eggs, and several
species of beetle larvae prey on the pupae.
The larvae have several anti-predator adaptations. While adults are
unable to sting, the larvae of species such as the spitfire sawfly
regurgitate a distasteful irritating liquid, which makes predators
such as ants avoid the larvae. In some species, the larvae
cluster together, reducing their chances of being killed, and in some
cases form together with their heads pointing outwards or tap their
abdomens up and down. Some adults bear black and yellow
markings that mimic wasps.
Parasitoidal chalcid wasps (Dahlbominus fuscipennis (sv)) emerging
from a sawfly cocoon
Sawflies are hosts to many parasitoids, most of which are parasitic
Hymenoptera; more than 40 species are known to attack them. However,
information regarding these species is minimal, and fewer than 10 of
these species actually cause a significant impact on sawfly
populations. Many of these species attack their hosts in the grass
or in other parasitoids.[clarification needed] Well known and
important parasitoids include Braconidae,
Eulophidae and Ichneumonidae
wasps. Braconid wasps attack sawflies in many regions throughout the
world, in which they are ectoparasitoids, meaning that the larvae live
and feed outside of the hosts body; braconids have more of an impact
on sawfly populations in the New World than they do in the Old World,
possibly due to no known ichneumonid parasitoids living in North
America.[clarification needed] Some braconid wasps that attack
sawflies include Bracon cephi, B. lisogaster, B. terabeila and
Heteropilus cephi. Female braconids locate sawfly larvae
through the vibrations they produce when feeding, followed by
inserting the ovipostior and paralysing the larva before laying eggs
inside the host. These eggs hatch inside the larva within a few days,
where they feed on the host. The entire host's body may be consumed by
the braconid larvae, except for the head capsule and epidermis. The
larvae complete their development within two or three weeks.
Ten species of wasps in the family
Ichneumonidae attack sawfly
populations, although these species are usually rare. The most
important parasitoids in this family are species in the genus
Collyria. Unlike Braconid wasps, the larvae are endoparasitoids,
meaning that the larvae live and feed inside the hosts body. One
well known Ichneumonid is Collyria coxator, which is a dominant
parasitoid of C. pygmaeus. Recorded parasitism rates in Europe are
between 20 – 76%, and as many as eight eggs can be found in a single
larva, but only one Collyria individual will emerge from its host. The
larva may remain inside of their host until spring, where it emerges
Several species in the family
Eulophidae attack sawflies, although
their impact is low. Two species in the genus
Pediobius have been
studied; the two species are internal larval parasitoids and have only
been found in the northern hemisphere. Parasitism of sawflies by
Eulophids in grass exceeds 50%, but only 5% in wheat. It is unknown as
to why the attack rate in wheat is low. Furthermore, some fungal
and bacterial diseases are known to infect eggs and pupa in warm wet
Outbreaks of certain sawfly species, such as Diprion polytomum, have
led scientists to investigate and possibly collect their natural
enemies to control them. Parasites of D. polytomum have been
extensively investigated, showing that 31 species of hymenopterous and
dipterous parasites attack it. These parasites have been used in
successful biological control against pest sawflies, including Cephus
cinctus throughout the 1930s and 1950s and C. pygmaeus in the 1930s
Life cycle and reproduction
Adult male newly emerged from its cocoon
Like all other hymenopteran insects, sawflies go through a complete
metamorphosis with four distinct life stages – egg, larva, pupa and
adult. Many species are parthenogenetic, meaning that females do
not need to reproduce to fertilise their eggs. Unfertilized eggs
develop as male, fertilized eggs develop into females, see
Arrhenotoky.The lifespan of an individual sawfly is two months to two
years, though the adult life stage is often very short (approximately
7 – 9 days), only long enough for the females to lay their
eggs. The female uses its ovipositor to drill into plant
material to lay her eggs (though the family
Orussoidea lay their eggs
in other insects). Plant-eating sawflies most commonly are associated
with leafy material but some specialize on wood, and the ovipositors
of these species (such as the family Siricidae) are specially adapted
for the task of drilling through bark. Once the incision has been
made, the female will lay as many as 30 to 90 eggs. Females avoid the
shade when laying their eggs because the larvae develop much slower
and may not even survive, and they may not also survive if they are
laid on immature and glaucous leaves. Hence, female sawflies search
for young adult leaves to lay their eggs on.
These eggs hatch in two to eight weeks, but such duration varies by
species and also by temperature. Until the eggs have hatched, some
species such as the small brown sawfly will remain with them and
protects the eggs by buzzing loudly and beating her wings to deter
predators. There are six larval stages that sawflies go through,
lasting 2 – 4 months, but this also depends on the species. When
fully grown, the larvae emerge from the trees en masse and burrow
themselves into the soil to pupate. During their time outside, the
larvae may link up to form a large colony if many other individuals
are present. They gather in large groups during the day which gives
them protection from potential enemies, and during the night they
disperse to feed. The emergence of adults takes awhile, with some
emerging anywhere between a couple months to 2 years. Some will reach
the ground to form pupal chambers, but others may spin a cocoon
attached to a leaf. Larvae that feed on wood will pupate in the
tunnels they have constructed. In one species, the jumping-disc sawfly
(Phyllotoma aceris) forms a cocoon which can act like a parachute. The
larvae live in sycamore trees and do not damage the upper or lower
cuticles of leaves that they feed on. When fully developed, they cut
small perforations in the upper cuticle to form a circle. After this,
they weave a silk hammocks within the circle; this silk hammock never
touches the lower cuticle. Once inside, the upper-cuticle's disc
separates and descends towards the surface with the larvae attaching
themselves to the hammock. Once they reach the round, the larvae work
their way into a sheltered area by jerking their discs along.
The majority of sawfly species produce a single generation per year,
but others may only have one generation every two years. Most sawflies
are also female, making males rare.
Relationship with humans
The pine sawfly
Diprion pini is a serious pest of forestry.
Caterpillar-like larvae of
Iris sawfly on yellow flag, showing damage
to host plant
Sawflies are major economic pests of forestry. For example, species in
the Diprionidae, such as the pine sawflies,
Diprion pini and
Neodiprion sertifer, cause serious damage to pines in regions such as
Scandinavia. D. pini larvae defoliated 500,000 hectares (1,200,000
acres) in the largest outbreak in Finland, between 1998 and 2001. Up
to 75% of the trees may die after such outbreaks, as D. pini can
remove all the leaves late in the growing season, leaving the trees
too weak to survive the winter. Little damage to trees only occurs
when the tree is large or when there is minimal presence of larvae.
Eucalyptus trees can regenerate quickly from damage inflicted by the
larvae; however, they can be substantially damaged from outbreaks,
especially if they are young. The trees can be defoliated completely
and may cause "dieback", stunting or even death.
Sawflies are serious pests in horticulture. Different species prefer
different host plants, often being specific to a family or genus of
hosts. For example,
Iris sawfly larvae, emerging in summer, can
quickly defoliate species of Iris including the yellow flag and other
freshwater species. Similarly the rose sawflies,
Arge pagana and
A. ochropus, defoliate rose bushes.
The giant woodwasp or horntail, Urocerus gigas, has a long ovipositor,
which with its black and yellow colouration make it a good mimic of a
hornet. Despite the alarming appearance, the insect cannot sting.
The eggs are laid in the wood of conifers such as Douglas fir, pine,
spruce, and larch. The larvae eat tunnels in the wood, causing
Alternative measures to control sawflies can be taken. Mechanical
methods include removing larvae from trees and killing them by
squishing or dropping them into boiling water or kerosene, although
this is not practical in plantations. Predators can also be used to
eliminate larvae, as well as parasites which have been previously used
in control programs. Small trees can be sprayed with a number
of chemicals, including maldison, dimethoate and carbaryl, if removing
larvae from trees is not effective enough.
^ a b Gerstäcker, C.E.A. (1867). "Ueber die Gattung Oxybelus Latr.
und die bei Berlin vorkommenden Arten derselben". Zeitschrift für die
Gesammten Naturwissenschaften (in German). 30 (7): 1–144.
^ a b Aguiar, A.P.; Deans, A.R.; Engel, M.S.; Forshage, M.; Huber,
J.T.; Jennings, J.T.; Johnson, N.F.; Lelej, A.S.; Longino, J.T.;
Lohrmann, V.; Mikó, I.; Ohl, M.; Rasmussen, C.; Taeger, A.; Yu,
D.S.K. (2013). "Order
Hymenoptera In: Zhang, Z.-Q. (ed.) Animal
biodiversity: an outline of higher-level classification and survey of
taxonomic richness". Zootaxa. 3703 (1): 51–62.
^ "Symphyta". Merriam-Webster. Retrieved 2 December 2016.
^ a b c Australian Museum (20 October 2009). "
Sawflies". Retrieved 11 August 2015.
^ "Sawfly". Merriam-Webster Online Dictionary. Retrieved 29 November
^ Gordh, G.; Headrick, D.H. (2011). A Dictionary of Entomology (2nd
ed.). Wallingford: CABI. p. 1344.
^ Carus, J.V.; Gerstäcker, C.E.A. (1863). Handbuch der zoologie: bd.
Arthropoden, bearb. von A. Gerstäcker. Raderthiere, würmer,
echinodermen, coelenteraten und protozoen, bearb. von J. Victor Carus.
1863 (in German). Leipzig, Germany: Engelmann. p. 189.
doi:10.5962/bhl.title.1399. OCLC 2962429.
^ Dallas, W.S. (1867). Insecta In: Günther, A.C.L.G The Zoological
Record: Insecta, Volumes 3-4. London, UK: John van Voorst.
p. 307. OCLC 6344527.
^ a b Goulet & Huber 1993, p. 5.
^ a b c Sharkey, M.J. (2007). "Phylogeny and classification of
Hymenoptera" (PDF). Zootaxa. 1668: 521–548.
^ a b c Mao, M.; Gibson, T.; Dowton, M. (2015). "Higher-level
phylogeny of the
Hymenoptera inferred from mitochondrial genomes".
Molecular Phylogenetics and Evolution. 84: 34–43.
doi:10.1016/j.ympev.2014.12.009. PMID 25542648.
^ Sharkey, M.J.; Carpenter, J.M.; Vilhelmsen, L.; Heraty, J.;
Liljeblad, J.; Dowling, A.P.G.; Schulmeister, S.; Murray, D.; Deans,
A.R.; Ronquist, F.; Krogmann, L.; Wheeler, W.C. (2012). "Phylogenetic
relationships among superfamilies of Hymenoptera". Cladistics. 28 (1):
^ a b Song, S.-N.; Tang, P.; Wei, S.-J.; Chen, X.-X. (2016).
"Comparative and phylogenetic analysis of the mitochondrial genomes in
basal hymenopterans". Scientific Reports. 6: 20972.
PMC 4754708 . PMID 26879745.
^ Hennig, W. (1969). Die Stammesgeschichte der Insekten. Frankfurt:
Waldemar Kramer. pp. 291, 359. ASIN B0000BRK5P.
^ Hermann, Henry R. (1979). Social Insects. 1. Oxford: Elsevier
Science. p. 85. ISBN 978-0-323-14979-2.
^ Engel, M.S. (2005). "A new sawfly from the
Triassic of Queensland
(Hymenoptera: Xyelidae)". Memoirs of the Queensland Museum. 51 (2):
^ Wang, M.; Rasinitsyn, A.P.; Ren, Dong (2014). "Two new fossil
sawflies (Hymenoptera, Xyelidae, Xyelinae) from the Middle
China". Acta Geologica Sinica. 88 (4): 1027–1033.
^ Wang, M.; Gao, T.; Shih, C.; Rasinitsyn, A.P.; Ren, D. (2016). "The
diversity and phylogeny of
Mesozoic Symphyta (Hymenoptera) from
Northeastern China". Acta Geologica Sinica. 90 (1): 376–377.
^ Rasnitsyn, A.P. (2006). "Ontology of evolution and methodology of
taxonomy". Paleontological Journal. 40 (S6): S679–S737.
^ Schulmeister, S. (2003). "Simultaneous analysis of basal Hymenoptera
(Insecta), introducing robust-choice sensitivity analysis". Biological
Journal of the Linnean Society. 79: 245–275.
^ Schulmeister, S. "'Symphyta'". Retrieved 28 November 2016.
^ a b Capinera 2008, pp. 3250–3252.
^ Taeger, A.; Blank, S.M.; Liston, A. (2010). "World catalog of
symphyta (Hymenoptera)". Zootaxa. 2580: 1–1064.
^ Skvarla, M.J.; Smith, D.R.; Fisher, D.M.; Dowling, A.P.G. (2016).
"Terrestrial arthropods of Steel Creek, Buffalo National River,
Sawflies (Insecta: Hymenoptera: "Symphyta")".
Biodiversity Data Journal. 4: e8830. doi:10.3897/BDJ.4.e8830.
PMC 4867044 . PMID 27222635.
^ Taeger, A.; Blank, S.M. (1996). "Kommentare zur Taxonomie der
Symphyta (Hymenoptera): Vorarbeiten zu einem Katalog der
Pflanzenwespen, Teil 1". Beiträge zur Entomologie (in German). 46
^ Rasnitsyn, A.P. (1988). "An outline of evolution of hymenopterous
insects (order Vespida)". Oriental Insects. 22: 115–145.
^ Dowton, M.; Austin, A.D. (1994). "Molecular phylogeny of the insect
order Hymenoptera: apocritan relationships". Proceedings of the
National Academy of Sciences of the United States of America. 91 (21):
9911–9915. Bibcode:1994PNAS...91.9911D. doi:10.1073/pnas.91.21.9911.
PMC 44927 . PMID 7937916.
^ Goulet & Huber 1993, p. 104.
^ Goulet & Huber 1993, pp. 5–6.
^ a b c d e f g h Burton, M.; Burton, R. (2002). International
Wildlife Encyclopedia. 16 (3rd ed.). Tarrytown, New York: Marshall
Cavendish. pp. 2240–2241. ISBN 978-0-7614-7282-7.
^ a b Goulet & Huber 1993, p. 6.
^ Gao, T.; Shih, C.; Rasnitsyn, A.P.; Ren, D.; Laudet, V. (2013).
"Hoplitolyda duolunica gen. et sp. nov. (Insecta, Hymenoptera,
Praesiricidae), the hitherto largest sawfly from the
China". PLoS ONE. 8 (5): e62420. Bibcode:2013PLoSO...862420G.
doi:10.1371/journal.pone.0062420. PMC 3643952 .
^ Benson, R.B. (1952). Handbooks for the Identification of British
Hymenoptera 2 Symphyta Section (b) (PDF). London: Royal
Entomological Society of London. p. 51. OCLC 429798429.
Archived from the original (PDF) on 24 September 2015.
^ Ross 1937, p. 11.
^ Ross 1937, p. 9.
^ a b Ross 1937, p. 10.
^ Ross 1937, p. 21.
^ Ross 1937, p. 13.
^ Ross 1937, pp. 22–29.
^ Ross 1937, p. 27.
^ Adams, C.; Early, M.; Brook, J.; Bamford, K. (2014). Principles of
Horticulture: Level 2. New York, New York: Routledge. p. 18.
^ Ross 1937, p. 29.
^ Australian Museum (9 July 2009). "Metamorphosis: a remarkable
change". Retrieved 8 December 2016.
^ Looney, C.; Smith, D.R; Collman, S.J.; Langor, D.W.; Peterson, M.A.
Sawflies (Hymenoptera, Symphyta) newly recorded from
Washington State". Journal of
Hymenoptera Research. 49: 129–159.
^ Rosenthal, G.A.; Berenbaum, M.R. (1991). Herbivores: Their
Interactions with Secondary Plant Metabolites (2nd ed.). Oxford:
Elsevier Science. p. 190. ISBN 978-0-323-13940-3.
^ a b "
Sawflies (Tenthredinoidae)". BBC. 2014. Retrieved 11 August
^ Bandeili, B.; Müller, C. (2009). "Folivory versus
florivory—adaptiveness of flower feeding". Naturwissenschaften. 97
(1): 79–88. Bibcode:2010NW.....97...79B.
doi:10.1007/s00114-009-0615-9. PMID 19826770.
^ a b Kleintjes, P.K.; Dahlsten, D.L. (1994). "Foraging behaviour and
nestling diet of Chestnut-Backed chickadees in monterey pine" (PDF).
The Condor. 96 (3): 647–653. doi:10.2307/1369468.
^ Cummins, S.; O'Halloran, J. (2002). "An assessment of the diet of
nestling Stonechats using compositional analysis: Coleoptera
Hymenoptera (sawflies, ichneumon flies, bees, wasps and
ants), terrestrial larvae (moth, sawfly and beetle) and Arachnida
(spiders and harvestmen) accounted for 81% of
diet". Bird Study. 49 (2): 139–145.
^ a b c d e f g h i j k Phillips, C. (1992). "Spitfires –
Defoliating Sawflies" (PDF). Department of Primary Industries and
Resources. Government of South Australia. Retrieved 11 August
^ Campbell, L.H.; Avery, M.I.; Donald, P.; Evans, A.D.; Green, R.E.;
Wilson, J.D. (1997). A Review of the Indirect Effects of Pesticides on
Birds (PDF) (Report). Peterborough, UK: Joint Nature Conservation
Committee, Report. no 227. p. 27. ISSN 0963-8091.
^ Brickle, N.W.; Harper, D.G.C. (1999). "Diet of nestling Corn
Buntings Miliaria calandra in southern England examined by
compositional analysis of faeces". Bird Study. 46 (3): 319–329.
^ Starling-Westerberg, A. (2001). "The habitat use and diet of Black
Grouse Tetrao tetrix in the Pennine hills of northern England". Bird
Study. 48 (1): 76–89. doi:10.1080/00063650109461205.
^ Cayford, J.T. (1990). "Distribution and habitat preferences of Black
Grouse in commercial forests in Wales: conservation and management
implications". Proceedings of the International Union Game of
Biologists Congress. 19: 435–447.
^ Holling, C.S. (1959). "The components of predation as revealed by a
study of small-mammal predation of the European
Pine Sawfly" (PDF).
The Canadian Entomologist. 91 (5): 293–320.
^ Müller, Caroline; Brakefield, P.M. (2003). "Analysis of a chemical
defense in sawfly larvae: easy bleeding targets predatory wasps in
late summer". Journal of Chemical Ecology. 29 (12): 2683–2694.
doi:10.1023/B:JOEC.0000008012.73092.01. ISSN 1573-1561.
^ Petre, C.-A.; Detrain, C.; Boevé, J.-L. (2007). "Anti-predator
defence mechanisms in sawfly larvae of Arge (Hymenoptera, Argidae)".
Insect Physiology. 53 (7): 668–675.
doi:10.1016/j.jinsphys.2007.04.007. PMID 17540402.
^ Phillips, Charlma (December 1992). "Spitfires - Defoliating
Sawflies". PIRSA. Archived from the original on 6 November 2009.
^ Hairston, N.G. (1989). Ecological Experiments: Purpose, Design and
Execution. Cambridge, Massachusetts: Cambridge University Press.
p. 96. ISBN 978-0-521-34692-4.
^ a b c d e Capinera 2008, p. 1827.
^ Alberta Agriculture (1988). Guide to Crop Protection in Alberta. 2.
Alberta: University of Alberta. p. 73.
^ Nelson, W.A.; Farstad, C.W. (2012). "Biology of Bracon cephi (Gahan)
(Hymenoptera: Braconidae), an important native parasite of the wheat
stem sawfly, Cephus cinctus Nort. (Hymenoptera: Cephidae), in Western
Canada". The Canadian Entomologist. 85 (3): 103–107.
^ Capinera 2008, p. 1827–1828.
^ a b Capinera 2008, p. 1828.
^ Morris, K.R.S.; Cameron, E.; Jepson, W.F. (1937). "The insect
parasites of the spruce sawfly (Diprion polytomum, Htg.) in Europe".
Bulletin of Entomological Research. 28 (03): 341–393.
^ Hartman, J.R.; Pirone, T.P.; Sall, M.A. (2000). Pirone's Tree
Maintenance (7th ed.). New York, New York: Oxford University Press.
p. 235. ISBN 978-0-19-802817-8.
^ Müller, C.; Barker, A.; Boevé, J.-L.; De Jong, P.W.; De Vos, H.;
Brakefield, P.M. (2004). "Phylogeography of two parthenogenetic sawfly
species (Hymenoptera: Tenthredinidae): relationship of population
genetic differentiation to host plant distribution". Biological
Journal of the Linnean Society. 83 (2): 219–227.
^ Krokene, Paal (6 December 2014). "The common pine sawfly – a
troublesome relative". Science Nordic. Retrieved 28 November
^ "Iris sawfly". Royal Horticultural Society. Retrieved 28 November
^ "Large rose sawfly". Royal Horticultural Society. Retrieved 28
^ "Great Wood Wasps". UK Safari. Retrieved 28 November 2016.
^ "Giant Woodwasp". Massachusetts Introduced Pests Outreach Project.
Retrieved 28 November 2016.
Capinera, J.L. (2008). Encyclopedia of Entomology (2nd ed.).
Dordrecht, Netherlands: Springer Science & Business Media.
Goulet, H.; Huber, J.T. (1993).
Hymenoptera of the World: An
Identification guide to families (PDF). Ottawa, Ontario: Agriculture
Canada. ISBN 978-0-660-14933-2.
Ross, H.H. (1937). A Generic Classification of the Nearctic Sawflies
(Hymenoptera, Symphyta). Urbana, Illinois: University of Illinois.
Blank, S.M.; Schmidt, S.; Taeger, A. (2006). Recent
Synthesis and Prospects. Keltern, Germany: Goecke und Evers.
Smith, D.R. (1969). Nearctic
Sawflies I. Blennocampinae: Adults and
Larvae (Hymenoptera: Tenthredinidae) (Technical Bulletin 1397).
Washington, D.C.: US Department of Agriculture.
Smith, D.R. (1969). Nearctic
Sawflies II. Selandriinae: Adults and
Larvae (Hymenoptera: Tenthredinidae) (Technical Bulletin 1398).
Washington, D.C.: US Department of Agriculture.
Smith, D.R. (1971). Nearctic
Sawflies III. Heterarthrinae: Adults and
Larvae (Hymenoptera: Tenthredinidae) (Technical Bulletin 1420).
Washington, D.C.: US Department of Agriculture.
Smith, D.R. (1979). Nearctic
Sawflies IV. Allantinae: Adults and
Larvae (Hymenoptera: Tenthredinidae) (Technical Bulletin 1595).
Washington, D.C.: US Department of Agriculture.
Wagner, M.R.; Raffa, K.F. (1993).
Sawfly Life History Adaptations to
Woody Plants. San Diego, California: Academic Press.
Wikispecies has information related to Symphyta
Wikimedia Commons has media related to Symphyta.
Symphyta: Encyclopædia Britannica
Sawflies: a close relative of wasps at CSIRO
Symphyta" - Sawflies, Horntails, and Wood Wasps at BugGuide
Hymenoptera – Chrysis.net
ECatSym - Electronic World Catalog of Symphyta (Insecta, Hymenoptera)
– Digital Entomological Information
Checklist of British and Irish
Hymenoptera - Sawflies, ‘Symphyta’
Biodiversity Data Journal 2: e1168
Archaeognatha (jumping bristletails)
Thysanura (Zygentoma) (silverfish, firebrats)
Odonata (dragonflies, damselflies)
Phasmatodea (stick and leaf insects)
Notoptera (ice-crawlers, gladiators)
Orthoptera (crickets, wetas, grasshoppers, locusts)
Zoraptera (angel insects)
Blattodea (cockroaches, termites)
Psocodea (barklice, lice)
Hemiptera (cicadas, aphids, true bugs)
Hymenoptera (sawflies, wasps, ants, bees)
Strepsiptera (twisted-winged parasites)
Megaloptera (alderflies, dobsonflies, fishflies)
Neuroptera (net-winged insects: lacewings, mantidflies, antlions)
Mecoptera (scorpionflies) + Siphonaptera (fleas)
Diptera (gnats, mosquitoes, flies)
Lepidoptera (moths, butterflies)
Four most speciose orders are marked in bold
Italic are paraphyletic groups
Based on Sasaki et al. (2013)
Extinct incertae sedis families and genera are marked in italic
Extant Hymenopteran families
Pamphiliidae (web-spinning sawflies)
Argidae (argid sawflies)
Blasticotomidae (fern sawflies)
Cimbicidae (cimbicid sawflies)
Diprionidae (conifer sawflies)
Pergidae (pergid sawflies)
Tenthredinidae (common sawflies)
Cephidae (stem sawflies)
Anaxyelidae (cedar wood wasps)
Xiphydriidae (wood wasps)
Orussidae (parasitic wood wasps)
Ichneumonidae (ichneumon wasps)
Cynipidae (gall wasps)
Agaonidae (fig wasps)
Evaniidae (ensign wasps)
Chrysididae (cuckoo wasps)
Rhopalosomatidae (rhopalosomatid wasps)
Vespidae (paper wasps, potter wasps, pollen wasps, yellowjackets,
Tiphiidae (tiphiid wasps)
Thynnidae (flower wasps)
Sierolomorphidae (sierolomorphid wasps)
Mutillidae (velvet ants)
Pompilidae (spider wasps)
Sapygidae (sapygid, or club-horned wasps)
Scoliidae (scoliid wasps)
Ampulicidae (cockroach wasps)
Crabronidae (sand wasps, bee wolves)
Sphecidae (thread-waisted wasps)
Andrenidae (mason bees)
Apidae (honey bees, bumblebees, cuckoo bees, carpenter bees, orchid
bees, stingless bees)
Colletidae (plasterer bees)
Halictidae (sweat bees)
Megachilidae (mason bees, leafcutter bees)
Italic are paraphyletic groups
Based on Malm and Nyman (2015) and Peters et al (2017)
Fauna Europaea: 11278