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The Capniidae, the small winter stoneflies, are a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s in the
stonefly Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mos ...
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
(Plecoptera). It constitutes one of the largest stonefly families, containing some 300
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 appropriat ...
distributed throughout the
holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
. Their closest relatives are the rolled-winged stoneflies (
Leuctridae The Leuctridae are a family of stoneflies. They are known commonly as rolled-winged stoneflies
). Many species are
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to small ranges, perhaps due to the family's tendency to evolve tolerance for cold (isolating populations in mountain valleys) and winglessness (inhibiting dispersal). Indeed, some wingless Capniidae – e.g. the
Lake Tahoe Lake Tahoe (; was, Dáʔaw, meaning "the lake") is a freshwater lake in the Sierra Nevada of the United States. Lying at , it straddles the state line between California and Nevada, west of Carson City. Lake Tahoe is the largest alpine lake i ...
benthic stonefly ('' "Capnia" lacustra'', ''Capnia'' is not monophyletic and this species is suspected to belong elsewhere) or '' Baikaloperla'' spp. – spend their entire lifecycles under water and do not disperse from their native lakes at all.


Description and ecology

Adult Capniidae, as their common name implies, are typically small Plecoptera; while most are less than 1 cm long with some measuring just 4 mm as adults, a few are as large as at adulthood. The adults emerge from the water in winter and are often found walking around on the snow. Characteristic are the
wings A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expre ...
with at most one cubital crossvein, and the paraprocts (anal lobes), the inner lobes of which form a tube closed on the underside by the outer lobes.
Nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
s of small winter stoneflies typically have very elongated and slender bodies, similar to those of
Leuctridae The Leuctridae are a family of stoneflies. They are known commonly as rolled-winged stoneflies
. However, the groove along the
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 to ...
, from segment 1 to 9, is generally very pronounced. The nymphs dwell in the
hyporheic zone The hyporheic zone is the region of sediment and porous space beneath and alongside a stream bed, where there is mixing of shallow groundwater and surface water. The flow dynamics and behavior in this zone (termed hyporheic flow or underflow) is re ...
, the interface between stream water and
groundwater Groundwater is the water present beneath Earth's surface in rock and Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit ...
. Only immediately before
moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
into the adult form will the nymphs move out of the substrate and appear on the
stream bed A stream bed or streambed is the bottom of a stream or river ( bathymetry) or the physical confine of the normal water flow ( channel). The lateral confines or channel margins are known as the stream banks or river banks, during all but fl ...
. Thus, although they may be plentiful in clean rivers and streams, they are seldom encountered in standard samples of
benthos Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.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 "unispe ...
as to
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
, with the Capniinae being inserted at that rank. However, this is rather pointless; in any case, the
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...
,
systematics Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic t ...
, and
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of the small winter stoneflies are highly confused. There appear to be two very basal genera and presumably two larger
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 ter ...
s which conceivably could be considered subfamilies, but the phylogeny of the Capniidae is by no means robustly resolved, with about one-third of the named genera of uncertain position. Hence, any subdivision beyond the generic level is premature. The
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal ...
''Capnia'' has generally been considered
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
. Three new or resurrected genera have recently been created, partially or wholly, from some of its species: '' Arsapnia'', '' Sierracapnia'', and '' Zwicknia''. In addition, seven ''Capnia'' species were previously included the genus '' Bolshecapnia'' when it was elevated from subgenus to genus, and three of those species were placed in the new genera '' Eurekapnia'' and '' Sasquacapnia'' in 2019.


Genera

These genera belong to the family Capniidae: * '' Allocapnia'' Claassen, 1928 * '' Apteroperla'' Matsumura, 1931 * '' Arsapnia'' Banks, 1897 * '' Baikaloperla'' Zapekina-Dulkeit & Zhiltzova, 1973 * '' Bolshecapnia'' Ricker, 1965 * '' Capnia'' Pictet, 1841 * '' Capniella'' Klapálek, 1920 * '' Capnioneura'' Ris, 1905 * '' Capnopsis'' Morton, 1896 * '' Capnura'' Banks, 1900 * '' Eocapnia'' Kawai, 1955 * '' Eucapnopsis'' Okamoto, 1922 * '' Eurekapnia'' Stark & Broome, 2019 * '' Isocapnia'' Banks, 1938 * '' Mesocapnia'' Raušer, 1968 * '' Nemocapnia'' Banks, 1938 * '' Paracapnia'' Hanson, 1946 * '' Sasquacapnia'' Baumann & Broome, 2019 * '' Sierracapnia'' Bottorff & Baumann, 2015 * '' Sinocapnia'' Murányi, Li & Yang, 2015 * '' Takagripopteryx'' Okamoto, 1922 * '' Utacapnia'' Gaufin, 1970 * '' Zwicknia'' Murányi, 2014


Extinct genera

* †'' Dobbertiniopteryx'' Ansorge, 1993 oooo Green Series, Germany, Early Jurassic (
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toar ...
),
Daohugou The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerates ...
, China, Middle/Late Jurassic * †'' Rovnocapnia'' Sinitshenkova 2009
Rovno amber Rivne amber, occasionally called Ukrainian amber, is amber found in the Rivne Oblast and surrounding regions of Ukraine and Belarus. The amber is dated between Late Eocene and Early Miocene, and suggested to be contemporaneous to Baltic amber. ...
, Ukraine,
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...


References

{{Taxonbar, from=Q998398 Plecoptera families