Sisyridae, commonly known as spongeflies or spongillaflies, are a
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
winged insects in the
order Neuroptera
The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
. There are approximately 60 living
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
described, and several extinct species identified from the fossil record.
Description

In general appearance, the adults resemble some brown lacewings (
Hemerobiidae). The forewings of spongillaflies have a span of 4–10 millimetres. The greyish or brownish wings have few
cross veins except in the
costal field, and most of these are not forked. The subcostal (Sc) and radial (R1) veins are fused near the wingtip.
The
larva
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e of spongillaflies look rather bizarre. Similar to those of some osmylids (
Osmylidae) at first glance, they have spindly legs on a bulky
thorax
The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen.
In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
, long
antennae, 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, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s carry seven pairs of jointed, movable
tracheal gill
A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...
s beneath their plump
abdomen
The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. 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 (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
or
nocturnal
Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite.
Nocturnal creatur ...
. They are
omnivores, sometimes hunting small
invertebrates
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
, but mainly
scavenging on such animals' carcasses, as well as on
pollen
Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
and
honeydew.
The females deposit their eggs singly or as small
clutch
A clutch is a mechanical device that allows an output shaft to be disconnected from a rotating input shaft. The clutch's input shaft is typically attached to a motor, while the clutch's output shaft is connected to the mechanism that does th ...
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. The term excludes seawater and brackish water, but it does include non-salty mi ...
lake
A lake is often a naturally occurring, relatively large and fixed body of water on or near the Earth's surface. It is localized in a basin or interconnected basins surrounded by dry land. Lakes lie completely on land and are separate from ...
s or slow-moving
river
A river is a natural stream of fresh water that flows on land or inside Subterranean river, caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of ...
s. A protective web is spun to cover the eggs. When the
larva
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e hatch, they drop down into the water where they develop until
pupation
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
. They use their mouthparts to
parasitize Spongillidae freshwater
sponge
Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a basal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and a ...
s (e.g. of the
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''
Spongilla'', hence the name "spongillaflies") and
Phylactolaemata freshwater
bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns by stinging into the host animals' body and sucking out
cell 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, 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 (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
climates they overwinter in the cocoon as larvae, pupating only the following spring.
Systematics and taxonomy
Spongillaflies were formerly placed in the
Osmyloidea, as their closest relatives were held to be the osmylids (
Osmylidae) and the
Nevrorthidae. 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 p ...
s of the
larva
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e 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 comm ...
; actually, the spongillaflies seem to be closer to the brown lacewings (
Hemerobiidae) than to the osmylids, but even more closely related to the dustywings (
Coniopterygidae
The dustywings, Coniopterygidae, are a family (biology), family of Pterygota (winged insects) of the net-winged insect order (biology), order (Neuroptera). About 460 living species are known.Engel & Grimaldi (2007) These tiny insects can usually ...
). And even though they are not often placed in the
superfamily Coniopterygoidea yet, they most likely form a
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 "unisp ...
taxon
In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...
, is expanded to signify that the spongillaflies and the dustywings 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 preserve ...
genera are known, mainly from the
Eocene
The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
like ''"Sisyra" amissa'' which may or may not be the first record of the living genus.
The genus ''
Cratosisyrops'', from the Aptian
Crato Formation, 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 International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
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. Th ...
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 m ...
Taimyr amber.
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. Th ...
, Myanmar, Late Cretaceous (
Cenomanian
The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
)
*†Paradoxosisyrinae Burmese amber, Myanmar, Late Cretaceous (
Cenomanian
The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
)
**†''
Buratina''
**†''
Khobotun''
**†''
Paradoxosisyra''
**†''
Protosiphoniella''
**†''
Sidorchukatia''
*Sisyrinae
**''
Climacia''
**†''
Paleosisyra''
Oise amber
Oise amber () is a type of amber found near the Oise river near Creil in northern France. Oise amber is around 53 million years old, dating to the Early Eocene (Ypresian). Oise amber is softer than Baltic amber, although Oise amber is older ...
, France,
Eocene
The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(
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
Baltic amber or succinite is amber from the Baltic region, home of its largest known deposits. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that this forested region provided the re ...
, Eocene
Bembridge Marls, 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 ...
)
**†''
Prosisyrina''
Taimyr amber, 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 m ...
)
**''
Sisyborina''
**''
Sisyra''
**''
Sisyrina''
References
:''This article draws heavily on the
corresponding article in the German-language Wikipedia.''
{{Taxonbar, from=Q1497564
Neuroptera
Coniopterygoidea
Neuroptera families