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Sisyridae, commonly known as spongeflies or spongillaflies, are a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of winged insects in the order
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
. There are approximately 60 living
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
described, and several extinct species identified from the fossil record.


Description

In general appearance, the adults resemble some brown lacewings (
Hemerobiidae Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long (som ...
). The forewings of spongillaflies have a span of 4–10 millimetres. The greyish or brownish wings have few cross veins except in the
costa Costa may refer to: Biology * Rib (Latin: ''costa''), in vertebrate anatomy * Costa (botany), the central strand of a plant leaf or thallus * Costa (coral), a stony rib, part of the skeleton of a coral * Costa (entomology), the leading edge of th ...
l field, and most of these are not forked. The subcostal (Sc) and radial (R1) veins are fused near the wingtip. The
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e of spongillaflies look rather bizarre. Similar to those of some osmylids (
Osmylidae Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids, also called lance lacewings, stream lacewings or giant lacewings, are found all over the world. There are around 225 extant species. Descr ...
) at first glance, they have spindly legs on a bulky
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
, long
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
e, and flexible, threadlike mouthparts. However, the second and third
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s carry seven pairs of jointed, movable
tracheal The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
s beneath their plump
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
. These gills are possessed by no other extant insect family, and readily distinguish them from osmylid larvae.


Ecology and life cycle

Adult spongillaflies are
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
or
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
. They are
omnivores An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutri ...
, sometimes hunting small
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
, but mainly
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
on such animals' carcasses, as well as on
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
and honeydew. The females deposit their eggs singly or as small
clutch A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...
es on plants that droop over
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s or slow-moving
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of wate ...
s. A protective web is spun to cover the eggs. When the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e hatch, they drop down into the water where they develop until
pupation A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
. They use their mouthparts to parasitize
Spongillidae Spongilidae is a family of sponges that live in freshwater lakes and rivers. The following genera are recognized in the family: * ''Anheteromeyenia'' Schröder, 1927 * ''Corvoheteromeyenia'' Ezcurra de Drago, 1979 * ''Corvospongilla'' Annandale, ...
freshwater
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s (e.g. of the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
Spongilla Overview ''Spongilla'' is a genus of freshwater sponges with over 200 different species. Spongilla was first publicly recognized in 1696 by Leonard Plukenet and can be found in lakes, ponds and slow streams.''Spongilla'' have a leuconoid body f ...
'', hence the name "spongillaflies") and
Phylactolaemata Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore, and like nearly all bryoz ...
freshwater
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns by stinging into the host animals' body and sucking out
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
contents. The antennae are stouter than they look and are used to aid in locomotion. Development to pupation takes between several weeks and one year. Spongillafly larvae leave the water and go to hidden places nearby to
pupate A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
, choosing locations like beneath rocks or behind tree bark. They spin a cocoon for pupation, but in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
climates they overwinter in the cocoon as larvae, pupating only the following spring.


Systematics and taxonomy

Spongillaflies were formerly placed in the
Osmyloidea Osmyloidea is a euneuropteran superfamily in the lacewing order Neuroptera sister to the superfamilies Dilaroidea, Mantispoidea, and the clade Neoneuroptera. The superfamily includes three living families and two extinct families described ...
, as their closest relatives were held to be the osmylids (
Osmylidae Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids, also called lance lacewings, stream lacewings or giant lacewings, are found all over the world. There are around 225 extant species. Descr ...
) and the
Nevrorthidae The Nevrorthidae, (often incorrectly spelled "Neurorthidae"), are a small family of winged insects of the order Neuroptera. Extant species may be described as living fossils. There are 19 extant species in four genera, with a geographically di ...
. This was due to the similar
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
s of the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e of spongillaflies and osmylids. But this is apparently due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
; actually, the spongillaflies seem to be closer to the brown lacewings (
Hemerobiidae Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long (som ...
) than to the osmylids, but even more closely related to the dusty wings ( Coniopterygidae). And even though they are not often placed in the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Coniopterygoidea yet, they most likely form a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
with the Coniopterygidae and thus it would seem that the Coniopterygoidea, rather than being an unnecessarily
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, is expanded to signify that the spongillaflies and the dusty wings are each other's closest relatives. A few
fossil 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 ...
genera are known, mainly from the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
like ''"Sisyra" amissa'' which may or may not be the first record of the living genus. The genus '' Cratosisyrops'', from the Aptian
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous ( Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly duri ...
, was suggested to be a spongillafly member, but later authors Perkovsky and Makarkin in 2015 noted the fossil lacks details to confidently place it in the family. The oldest members of the family are species in the subfamily Paradoxosisyrinae from
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
and '' Prosisyrina'' from
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
Taimyr amber Taymyr or Taimyr may refer to: Places *Taymyr Peninsula, a peninsula in Siberia *Taymyr Gulf *Taymyra, a river in the Taymyr Peninsula *Lake Taymyr *Taymyr Island, an island in the Kara Sea *Taymyr Autonomous Okrug, a former federal subject of Rus ...
. The family is currently divided into two subfamilies with the four extant genera and two of the extinct genera placed into Sisyrinae, while five extinct genera are placed into the subfamily Paradoxosisyrinae. Sisyridae *†'' Stictosisyra'' Yang et al. 2018
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Late Cretaceous (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
) *†Paradoxosisyrinae Burmese amber, Myanmar, Late Cretaceous (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
) **†'' Buratina'' **†'' Khobotun'' **†'' Paradoxosisyra'' **†'' Protosiphoniella'' **†'' Sidorchukatia'' *Sisyrinae **'' Climacia'' **†'' Paleosisyra''
Oise amber Oise ( ; ; pcd, Oése) is a department in the north of France. It is named after the river Oise. Inhabitants of the department are called ''Oisiens'' () or ''Isariens'', after the Latin name for the river, Isara. It had a population of 829,419 ...
, France,
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
(
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
)
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
, Eocene
Bembridge Marls The Bouldnor Formation is a geological formation in the Hampshire Basin of southern England. It is the youngest formation of the Solent Group and was deposited during the uppermost Eocene and lower Oligocene. Stratotype and occurrence The Bo ...
, United Kingdom, Eocene (
Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage of t ...
) **†'' Prosisyrina''
Taimyr amber Taymyr or Taimyr may refer to: Places *Taymyr Peninsula, a peninsula in Siberia *Taymyr Gulf *Taymyra, a river in the Taymyr Peninsula *Lake Taymyr *Taymyr Island, an island in the Kara Sea *Taymyr Autonomous Okrug, a former federal subject of Rus ...
, Russia, Late Cretaceous (
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
) **'' Sisyborina'' **'' Sisyra'' **'' Sisyrina''


References

:''This article draws heavily on the corresponding article in the German-language Wikipedia.'' {{Taxonbar, from=Q1497564 Neuroptera Coniopterygoidea Neuroptera families