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
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
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
* 1 Etymology
* 2 Phylogeny
* 2.1 Taxonomy
* 3 Description
* 4 Distribution
* 5 Behaviour and ecology
* 5.1 Parasites
* 5.2 Life cycle and reproduction
* 6 Relationship with humans
* 7 References
* 7.1 Bibliography
* 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
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 paraphyletic.
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 the
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
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
Hymenoptera , 250mya
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.)
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.) ">
Syntexis libocedrii )
(Janus femoratus )
(Orussus coronatus )
Sirex noctilio )
Tenthredo notha )
(Xiphydria camelus )
(Xyela julii )
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 tibiae.
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 ">
Aposematically coloured caterpillar -like larva of
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
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
Palearctic ; the
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, including
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.
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
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. Well known and important parasitoids include
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. 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 and
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
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
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
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 economic damage.
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.
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