Caddisfly

The caddisflies (order Trichoptera) are a group of insects

with aquatic larvae and terrestrial adults. There are Caddisflies
approximately 14,500 described species, most of which can be Temporal range:
divided into the suborders Integripalpia and Annulipalpia on
the basis of the adult mouthparts. Integripalpian larvae
construct a portable casing to protect themselves as they move
around looking for food, while annulipalpian larvae make
themselves a fixed retreat in which they remain, waiting for
food to come to them. The affinities of the small third suborder
Spicipalpia are unclear, and molecular analysis suggests it may
not be monophyletic. Also called sedge-flies or rail-flies, the Scientific classification
adults are small moth-like insects with two pairs of hairy
membranous wings. They are closely related to the Lepidoptera Domain: Eukaryota
(moths and butterflies) which have scales on their wings; the Kingdom: Animalia
two orders together form the superorder Amphiesmenoptera. Phylum: Arthropoda
The aquatic larvae are found in a wide variety of habitats such Class: Insecta
as streams, rivers, lakes, ponds, spring seeps and temporary (unranked): Amphiesmenoptera
waters (vernal pools), and even the ocean.[1][2][3] The larvae of
Order: Trichoptera
many species use silk to make protective cases, which are often
strengthened with gravel, sand, twigs, bitten-off pieces of Kirby, 1813
plants, or other debris. The larvae exhibit various feeding Superfamilies
strategies, with different species being predators, leaf
shredders, algal grazers, or collectors of particles from the Annulipalpia
water column and benthos. Most adults have short lives during
which they do not feed. Hydropsychoidea
Philopotamoidea
In fly fishing, artificial flies called dry flies are tied to imitate
adults, while larvae and pupae are imitated with artificial flies Integripalpia
called wet flies or nymphs. It is also possible to use them as
Leptoceroidea
bait, though this is not as common as artificial flies and is
known as bait fishing. Common and widespread genera such as Limnephiloidea
Helicopsyche and Hydropsyche are important in the sport, Phryganeoidea
where caddisflies are known as "sedges". Caddisflies are useful Sericostomatoidea
as bioindicators, as they are sensitive to water pollution and are
large enough to be assessed in the field. In art, the French artist Tasimioidea
Spicipalpia
Hubert Duprat has created works by providing caddis larvae Hydroptiloidea
with small grains of gold and precious stones for them to build
Rhyacophiloidea
into decorative cases.

Etymology
The name of the order "Trichoptera" derives from the Greek: θρίξ (thrix, "hair"), genitive trichos +
πτερόν (pteron, "wing"), and refers to the fact that the wings of these insects are bristly. The origin of
the word "caddis" is unclear, but it dates back to at least as far as Izaak Walton's 1653 book The
Compleat Angler, where "cod-worms or caddis" were mentioned as being used as bait. The term
cadyss was being used in the fifteenth century for silk or cotton cloth, and "cadice-men" were
itinerant vendors of such materials, but a connection between these words and the insects has not
been established.[4]

Evolution and phylogeny

Fossil history
Fossil caddisflies have been found in rocks dating back to the Triassic.[5]
The largest numbers of fossilised remains are those of larval cases, which
are made of durable materials that preserve well. Body fossils of
caddisflies are extremely rare, the oldest being from the Early and Middle
Triassic, some 230 million years ago, and wings are another source of
fossils.[6] The evolution of the group to one with fully aquatic larvae
seems to have taken place sometime during the Triassic.[7] The finding of
fossils resembling caddisfly larval cases in marine deposits in Brazil may
push back the origins of the order to the Early Permian period.[6]
Eocene fossil in Baltic
Evolution amber, Lithuania (44mya)

Nearly all adult caddisflies are terrestrial, but their larvae and pupae are
aquatic. They share this characteristic with several distantly-related groups, namely the dragonflies,
mayflies, stoneflies, alderflies and lacewings.[7] The ancestors of all these groups were terrestrial, with
open tracheal systems, convergently evolving different types of gills for their aquatic larvae as they
took to the water to avoid predation.[7] Caddisflies was the only group of these insects to use silk as
part of their lifestyle, which has been a contributing factor to their success and why they are the most
species-rich order of aquatic insects.[8]

About 14,500 species of caddisfly in 45 families have been recognised worldwide,[9] but many more
species remain to be described. Most can be divided into the suborders Integripalpia and Annulipalpia
on the basis of the adult mouthparts. The characteristics of adults depend on the palps, wing venation
and genitalia of both sexes. The latter two characters have undergone such extensive differentiation
among the different superfamilies that the differences between the suborders is not clear-cut.[10] The
larvae of Annulipalpians are campodeiform (free-living, well sclerotized, long legged predators with
dorso-ventrally flattened bodies and protruding mouthparts). The larvae of Integripalpians are
polypod (poorly sclerotized detritivores, with abdominal prolegs in addition to thoracic legs, living
permanently in tight-fitting cases).[10] The affinities of the third suborder, Spicipalpia, are unclear;
the larvae are free-living with no cases, instead creating net-like traps from silk.[4]

Phylogeny
The cladogram of external relationships, based on molecular analysis, shows the order as a clade,
sister to the Lepidoptera, and more distantly related to the Diptera (true flies) and Mecoptera
(scorpionflies).[11]

⁠ Hymenoptera (sawflies, wasps)

⁠

⁠ Coleoptera (beetles)
⁠
⁠Coleopterida
Strepsiptera (twisted-wing parasites)
⁠ ⁠
⁠

⁠ Raphidioptera (snakeflies)
⁠Neuropteroidea ⁠
⁠
Megaloptera (alderflies and allies)
⁠
⁠Neuropterida
⁠
⁠
⁠
⁠ Neuroptera (Lacewings and allies)
⁠
⁠
Holometabola
⁠
⁠Aparaglossata Lepidoptera (butterflies, moths)
⁠ ⁠
⁠
⁠Amphiesmenoptera
⁠ Trichoptera (caddisflies)
⁠
⁠

⁠Panorpida ⁠Antliophora
⁠ ⁠ Diptera
⁠

⁠ ⁠
 Mecoptera (scorpionflies)
⁠ ⁠
⁠
⁠ Siphonaptera (fleas)
⁠

The cladogram of relationships within the order is based on a 2002 molecular phylogeny using
ribosomal RNA, a nuclear elongation factor gene, and mitochondrial cytochrome oxidase. The
Annulipalpia and Integripalpia are clades, but the relationships within the Spicipalpia are unclear.[12]

⁠ Annulipalpia (fixed-retreat makers)

⁠
Trichoptera ⁠ Integripalpia (portable-case makers)
⁠ ⁠ ⁠
⁠ ⁠ "Spicipalpia" (paraphyletic?)
⁠

Distribution
Caddisflies are found worldwide, with the greater diversity being in warmer regions. They are
associated with bodies of freshwater, the larvae being found in lakes, ponds, rivers, streams and other
water bodies.[13] The land caddis, Enoicyla pusilla (family: Limnephilidae), lives in the damp litter of
the woodland floor. In the United Kingdom it is found in and around the county of Worcestershire in
oakwoods.[14]

Ecology
Caddisfly larvae can be found in all feeding guilds in freshwater
habitats. Most early stage larvae and some late stage ones are
collector-gatherers, picking up fragments of organic matter from
the benthos. Other species are collector-filterers, sieving organic
particles from the water using silken nets, or hairs on their legs.
Some species are scrapers, feeding on the film of algae and other
periphyton that grows on underwater objects in sunlight. Others
0:26
are shredder-herbivores, chewing fragments off living plant
material while others are shredder-detritivores, gnawing at rotting Larva in its underwater habitat
wood or chewing dead leaves that have been pre-processed by
bacteria and fungi; most of the nutrients of the latter group come
from consumption of the bacteria and fungi. The predatory species either actively hunt their prey,
typically other insects, tiny crustaceans and worms, or lie in wait for unwary invertebrates to come too
close. A few species feed opportunistically on dead animals or fish, and some Leptoceridae larvae feed
on freshwater sponges.[15]
One such opportunistic species is Gumaga nigricula (family: Sericostomatidae) which has been
observed scavenging fish carcasses and even bits of deer flesh.[16] This particular family of caddisflies
is typically classified among the shredders, suggesting caution when classifying macroinvertebrates
into strict ecological functional groups, as some may shift their diets opportunistically.[16]

Like mayflies, stoneflies and dragonflies, but to a somewhat lesser extent, caddisflies are an indicator
of good water quality; they die out of streams with polluted waters.[17] They are an important part of
the food web, both larvae and adults being eaten by many fish. The newly hatched adult is particularly
vulnerable as it struggles to the surface after emerging from the submerged pupa, and as it dries its
wings. The fish find these new adults easy pickings, and fishing flies resembling them can be
successful for anglers at the right time of year.[18]

The adult stage of a caddisfly may only survive for a few weeks; many species do not feed as adults and
die soon after breeding, but some species are known to feed on nectar.[19] The winged insects are
nocturnal and provide food for night-flying birds, bats, small mammals, amphibians and arthropods.
The larval stage lasts much longer, often for one or more years, and has a bigger impact on the
environment.[20] They form an important part of the diet of fish such as the trout. The fish acquire
them by two means, either plucking them off vegetation or the stream-bed as the larvae move about,
or during the daily behavioural drift; this drift happens during the night for many species of aquatic
larvae, or around midday for some cased caddisfly species, and may result from population pressures
or be a dispersal device. The larvae may drift in great numbers either close to the bottom, in mid-
water or just below the surface. The fish swallow them whole, case and all.[21]

Underwater structures

Cases
Caddisflies are best known for the portable cases created by their larvae. About thirty families of
caddisfly, members of the suborder Integripalpia, adopt this stratagem. These larvae eat detritus,
largely decaying vegetable material, and the dead leaf fragments on which they feed tend to
accumulate in hollows, in slow-moving sections of streams and behind stones and tree roots. The
cases provide protection to the larvae as they make their way between these resources.[22]

The case is a tubular structure made of silk, secreted from salivary glands near the mouth of the larva,
and is started soon after the egg hatches. Various reinforcements may be incorporated into its
structure, the nature of the materials and design depending on the larva's genetic makeup; this means
that caddisfly larvae can be recognised by their cases down to family, and even genus level. The
materials used include grains of sand, larger fragments of rock, bark, sticks, leaves, seeds and mollusc
shells. These are neatly arranged and stuck onto the outer surface of the silken tube. As the larva
grows, more material is added at the front, and the larva can turn round in the tube and trim the rear
end so that it does not drag along the substrate.[22]

Caddisfly cases are open at both ends, the larvae drawing oxygenated water through the posterior end,
over their gills, and pumping it out of the wider, anterior end. The larvae move around inside the
tubes and this helps maintain the water current; the lower the oxygen content of the water, the more
active the larvae need to be. This mechanism enable caddisfly larvae to live in waters too low in
oxygen content to support stonefly and mayfly larvae.[19]

Larva with portable Larva emerging from Larval case of Case of Limnephilus
case of rock case made of plant Limnephilidae made flavicornis made of
fragments material of bitten-off plant snail shells
pieces

Fixed retreats
In contrast to larvae that have portable cases, members of the Annulipalpia have a completely
different feeding strategy. They make fixed retreats in which they remain stationary, waiting for food
to come to them. Members of the Psychomyiidae, Ecnomidae and Xiphocentronidae families
construct simple tubes of sand and other particles held together by silk and anchored to the bottom,
and feed on the accumulations of silt formed when suspended material is deposited. The tube can be
lengthened when the growing larva needs to feed in new areas.[23] More complex tubes, short and
flattened, are built by Polycentropodidae larvae in hollows in rocks or other submerged objects,
sometimes with strands of silk suspended across the nearby surface. These larvae are carnivorous,
resembling spiders in their feeding habits and rushing out of their retreat to attack any unwary small
prey crawling across the surface.[23]

Silk domes
Larvae of members of the family Glossosomatidae in the suborder Spicipalpia create dome-shaped
enclosures of silk which enables them to graze on the periphyton, the biological film that grows on
stones and other objects, while carrying their enclosure around like turtles.[24] In the family
Philopotamidae, the nets are sac-like, with intricate structure and tiny mesh. The larvae have
specialised mouthparts to scrape off the microflora that get trapped in the net as water flows
through.[25]

Nets
The larvae of other species of caddisfly make nets rather than
cases. These are silken webs stretching between aquatic vegetation
and over stones. These net-making larvae usually live in running
water, different species occupying different habitats with varying
water speeds. There is a constant drift of invertebrates washed
downstream by the current, and these animals, and bits of debris,
accumulate in the nets which serve both as food traps and as
retreats.[26] Net made by a larva of the suborder
Spicipalpia

Development and morphology

Caddisfly larvae are aquatic, with six pairs of tracheal gills on the underside of the abdomen. The eggs
are laid above water on emergent twigs or vegetation or on the water surface although females of some
species enter water to choose sites. Although most species lay eggs, a few in the genus Triplectides are
ovoviviparous. Some species lay eggs on land and although most are associated with freshwater, a few
like Symphitoneuria are found in coastal saline water. Philanisus plebeius females lay their eggs into
the coelomic cavity of intertidal starfish.[27] The larvae are long and roughly cylindrical, very similar
to those of lepidoptera but lacking prolegs.[27] In case-bearing species, the heads are heavily
sclerotised while the abdomen is soft; the antennae are short and the mouthparts adapted for biting.
Each of the usually ten abdominal segments bears a pair of legs with a single tarsal joint. In case-
bearing species, the first segment bears three papillae, one above and two at the sides, which anchor
the larva centrally in the tube. The posterior segment bears a pair of hooks for grappling.[19] There are
five to seven larval instars, followed by an aquatic pupa which has functional mandibles (to cut
through the case), gills, and swimming legs.[9]

The pupal cocoon is spun from silk, but like the larval case, often has other materials attached. When
pupating, species that build portable cases attach them to some underwater object, seal the front and
back apertures against predators while still allowing water to flow through, and pupate within it. Once
fully developed, most pupal caddisflies cut through their cases with a special pair of mandibles, swim
up to the water surface, moult using the exuviae as a floating platform, and emerge as fully formed
adults. They can often fly immediately after breaking from their pupal cuticle. Emergence is mainly
univoltine (once per year) with all the adults of a species emerging at the same time. Development is
within a year in warm places, but takes over a year in high latitudes and at high elevation in mountain
lakes and streams.[9]

The adult caddisfly is a medium-sized insect with membranous, hairy wings, which are held in a tent-
wise fashion when the insect is at rest. The antennae are fairly long and threadlike, the mouthparts
are reduced in size and the legs have five tarsi (lower leg joints).[19] Adults are nocturnal and are
attracted to light. Some species are strong fliers and can disperse to new localities,[26] but many fly
only weakly.[19] Adults are usually short-lived, most being non-feeders and equipped only to breed.
Once mated, the female caddisfly lays eggs in a gelatinous mass, attaching them above or below the
water surface depending on species. The eggs hatch in a few weeks.[28]
 Egg mass on leaf Pupa with oar-like Pupae Parachiona picicornis
away from water swimming legs adult emerging from
aquatic pupa

Adult

Relationship with humans

In angling
Adult caddisflies are called sedges by anglers. Individual species emerge
en masse at different times, and are used one after the other, often for
only a few days each year, as models for artificial fishing flies for fly
fishing in trout streams.[14] A mass emergence is known as a hatch.[29]
Each type has its own angling name, so for example Mystacides is the
dancer; Sericostoma the caperer; Leptocerus the silverhorn; Phryganea
the murragh or great red sedge; Brachycentrus subnubilis the grannom;
Lepidostoma the silver sedge;[14] Oecetis the longhorn sedge; "Silver Sedge" fishing fly
Cheumatopsyche the little sister sedge; Helicopsyche the speckled Peter, mimicking Lepidostoma
an important fishing fly in North America; and Hydropsyche the specked caddisfly, from Trout fly-
sedge, perhaps the most important caddisfly genus for anglers with over fishing in America
50 species of net-makers.[29]

As bioindicators
Caddisflies are useful as bioindicators (of good water quality), since they are sensitive to water
pollution, and are large enough to be assessed conveniently in the field.[30] Some species indicate
undisturbed habitat, and some indicate degraded habitat.[31] Although caddisflies may be found in
waterbodies of varying qualities, species-rich caddisfly assemblages are
generally thought to indicate clean water bodies, such as lakes, ponds,
and marshes. Together with stoneflies and mayflies, caddisflies feature
importantly in bioassessment surveys of streams and other water
bodies.[32]

In art
While caddisflies in the wild construct their cases out of twigs, sand,
aquatic plants, and rocks, the French artist Hubert Duprat makes art by
providing wild caddisflies with precious stones and other materials. He
collected caddisfly larvae from the wild and put them in climate-
controlled tanks. He removes the larvae from their original cases and
adds precious and semi-precious items such as grains of gold into the
tank. The larvae then build new cases out of precious items, creating a
unique form of artwork. The resulting works are sold across the world.[33]
"Limnephilus elegans the
Elegant Grannom", from
British Entomology by John
As food
Curtis, c. 1840
In Japan the larvae of Stenopsyche marmorata are eaten as a delicacy
called Zazamushi.[34]

Taxonomy
There are roughly 16,266 extant species in 618 genera and 51 families worldwide.[35]

Suborder Annulipalpia
Superfamily Hydropsychoidea
Family Hydropsychidae
Superfamily Psychomyioidea
Family Dipseudopsidae
Family Ecnomidae
Family †Electralbertidae
Family Polycentropodidae
Family Psychomyiidae
Family Xiphocentronidae
Superfamily Philopotamoidea
Family Philopotamidae
Family Stenopsychidae
Suborder Integripalpia
Superfamily Leptoceroidea
Family Atriplectididae
 Family Calamoceratidae
Family Molannidae
Family Leptoceridae
Family Limnocentropodidae
Family Odontoceridae
Family Philorheithridae
Superfamily Limnephiloidea
Family Apataniidae
Family Brachycentridae
Family Goeridae
Family Limnephilidae
Family Lepidostomatidae
Family Oeconesidae
Family Pisuliidae
Family Rossianidae
Family †Taymyrelectronidae
Family Uenoidae
Superfamily †Necrotaulioidea
Family †Necrotauliidae
Superfamily Phyrganeoidea
Family †Baissoferidae
Family †Dysoneuridae
Family †Kalophryganeidae
Family Phryganeidae
Family Phryganopsychidae
Family Plectrotarsidae
Superfamily Sericostomatoidea
Family Anomalopsychidae
Family Antipodoeciidae
Family Barbarochthonidae
Family Beraeidae
Family Calocidae
Family Chathamiidae
Family Conoesucidae
Family Helicophidae
Family Helicopsychidae
Family Hydrosalpingidae
Family Kokiriidae
Family Petrothrincidae
Family Sericostomatidae
Superfamily Tasimioidea
Family Tasimiidae
 Superfamily †Vitimotaulioidea
Family †Vitimotauliidae
Family †Cladochoristidae
Family †Microptysmatidae
Family †Prosepididontidae
Family †Protomeropidae
Family †Uraloptysmatidae
Suborder Spicipalpia
Superfamily Hydroptiloidea
Family Glossosomatidae
Family Hydroptilidae
Family Ptilocolepidae
Superfamily Rhyacophiloidea
Family Hydrobiosidae
Family Rhyacophilidae

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1).

Further reading
A useful reference to the larvae of the British Trichoptera is "Caddis Larvae" Norman E. Hickin (1967)
Hutchinson & Co. Ltd. London.

Caddisfly (http://troutster.com/caddis-fly-adult-larva-and-pupa-life-cycle/), Troutster.com

External links
The Nature Explorers (https://web.archive.org/web/20120322123316/http://www.thenatureexplorer
s.com/fossil_creek.html) - Videos of some very interesting caddisfly species in Arizona.
Caddisflies and Fly Fishing (http://www.troutnut.com/hatch/12/Insect-Trichoptera-Caddisflies) -
Photos, limited species life history descriptions.
Adult caddisfly and caddisfly larva (casemaker) (http://www.cirrusimage.com/Trichoptera_caddisfl
y.htm) Archived (https://web.archive.org/web/20180729230546/http://www.cirrusimage.com/Tricho
ptera_caddisfly.htm) 2018-07-29 at the Wayback Machine — diagnostic photographs and
information
Kendall Bioresearch Services Trichoptera page (http://www.kendall-bioresearch.co.uk/trich.htm)
 Tree of Life Trichoptera page (http://tolweb.org/tree?group=Trichoptera&contgroup=Endopterygot
a)
Trichoptera world checklist (https://web.archive.org/web/20050207114739/http://entweb.clemson.e
du/database/trichopt/)
Animaldiversity.ummz.umich.edu (http://animaldiversity.ummz.umich.edu/site/accounts/classificati
on/Trichoptera.html#Trichoptera)
Artwork with caddis fly larvae and precious metals! (http://www.cabinetmagazine.org/issues/25/du
prat.php) "An unusual artistic collaboration between the French artist Hubert Duprat and a group
of caddis fly larvae."
Caddisfly larvae (https://web.archive.org/web/20130726014637/http://cabinetoffreshwatercuriositie
s.com/2011/03/01/caddis-larvae-trichoptera/), The BioFresh Cabinet of Freshwater Curiosities.

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