Hydroptilini

Holzenthal, Ralph W., Blahnik, Roger J., Prather, Aysha L. & Kjer, Karl M., 2007, Order Trichoptera Kirby, 1813 (Insecta), Caddisflies *, Zootaxa 1668, pp. 639-698 : 670-686

Tribe Hydroptilini Stephens

Acanthotrichia Wells, 1982 ....................................... Au Acritoptila Wells, 1982 ........................................... Au Agraylea Curtis, 1834 ........................................... Hol Allotrichia McLachlan, 1880 ...................................... Pa Austratrichia Wells, 1982 ......................................... Au Cyclopsiella Kjaerandsen, 1997 ................................... Afr Dhatrichia Mosely, 1948 ........................................ Afr Hellyethira Neboiss, 1977 ..................................Au, Pa, Or Hydroptila Dalman, 1819 ............................... Cosmopolitan Hydroptilina Martynov, 1934 ...................................... Pa Jabitrichia Wells, 1990 ................................... Afr, Au, Or Microptila Ris, 1897 ...................................... Afr, Or, Pa Missitrichia Wells, 1991 .......................................... Au Mulgravia Wells, 1982 ........................................... Au Oxyethira Eaton, 1873 .................................. Cosmopolitan Paroxyethira Mosely, 1924 ....................................... Au Paucicalcaria Mathis and Bowles, 1989 ....................Na (Arkansas) Tangatrichia Wells and Andersen, 1995 ............................ Afr Tricholeiochiton Kloet and Hincks, 1944 .................. Au, Neo, Or, Pa Ugandatrichia Mosely, 1939 ..............................Afr, Au, Neo Vietrichia Olah, 1989 ................................... Or ( Vietnam) Wlitrichia Kjaerandsen, 1997 ..................................... Afr Xuthotrichia Mosely, 1934 ........................................ Au

Tribe Leucotrichiini Flint, 1970

Abtrichia Mosely, 1939 ......................................... Neo Acostatrichia Mosely, 1939 ...................................... Neo Alisotrichia Flint, 1964 ................................ Neo (+sw USA) Anchitrichia Flint, 1970 ......................................... Neo Ascotrichia Flint, 1983 .......................................... Neo Betrichia Mosely, 1939 ......................................... Neo Celaenotrichia Mosely, 1934 ..................................... Neo Cerasmatrichia Flint, Harris , and Botosaneanu, 1994.................. Neo Ceratotrichia Flint, 1992 ........................................ Neo Costatrichia Mosely, 1937 ....................................... Neo Eutonella Müller, 1921 .......................................... Neo Leucotrichia Mosely, 1934 ....................................Neo, Na Mejicanotrichia Harris and Holzenthal, 1997 ................ Neo ( Mexico ) Peltopsyche Müller, 1879 ........................................ Neo Scelobotrichia Harris and Bueno-Soria, 1993 ........................ Neo Zumatrichia Mosely, 1937 ............................. Neo (+sw USA)

Tribe Neotrichiini Ross, 1956

Kumanskiella Harris and OS Flint, 1992............................ Neo Mayatrichia Mosely, 1937 ....................................Neo, Na Neotrichia Morton, 1905 .....................................Neo, Na Taraxitrichia Flint and Harris, 1991 ................................ Neo

Tribe Ochrotrichiini Marshall, 1979

Metrichia Ross, 1938 ................................. Neo (+sw USA) Ochrotrichia Mosely, 1934 ...................................Neo, Na Rhyacopsyche Mueller, 1879 ..................................... Neo

......c ontinued Tribe Orthotrichiini Nielsen, 1948

Ithytrichia Eaton, 1873 ....................................... Hol, Or Nothotrichia Flint, 1967 ...................................... Neo, Na Orthotrichia Eaton, 1873 ................................ Cosmopolitan

Tribe Stactobiini Botosaneanu, 1956

Bredinia Flint, 1968 ............................................ Neo Byrsopteryx Flint, 1981 ......................................... Neo Catoxyethira Ulmer, 1912 ..................................... Au, Or Chrysotrichia Schmid, 1958 .................................... Au, Or Flintiella Angrisano, 1995 ....................................... Neo Niuginitrichia Wells, 1990 ........................................ Au Orinocotrichia Harris, Flint , and Holzenthal, 2002.................... Neo Parastactobia Schmid, 1958 ...................................... Or Plethus Hagen, 1887 ............................................. Or Scelotrichia Ulmer, 1951 ................................... Old World Stactobia McLachlan, 1880 ................................. Old World Stactobiella Martynov, 1924 .................................. Hol, Or Tizatetrichia Harris, Flint , and Holzenthal, 2002...................... Neo

incertae sedis

Caledonotrichia Sykora 1967 ........................Au ( New Caledonia) Dibusa Ross, 1939 .............................................. Na Dicaminus Mueller, 1879 ........................................ Neo Macrostactobia Schmid, 1958 ..................................... Or Maydenoptila Neboiss, 1977 ...................................... Au Orphninotrichia Mosely, 1934 ..................................... Au

Mathis and Bowles (Arkansas, USA), and Vietrichia Olah ( Vietnam) . Agraylea Curtis occurs across the Holarctic and has about 20 species. The other Hydroptilini genera are more widespread across several biogeographical regions and include 2 large cosmopolitan genera, Hydroptila Dalman with about 400 described species and Oxyethira with about 200. Orthotrichiini is a small tribe of 3 genera, of which Orthotrichia is cosmopolitan and contains over 150 species. Stactobiini is a heterogeneous assemblage of genera endemic to a single region or more broadly distributed across several regions; Stactobia is the most species rich with about 150 species found only in the Old World. Nielsen (1948) studied the biology of Hydroptilidae . Hydroptilid larvae are highly diverse in form, habitat, and feeding behavior. Although most construct cases of silk or sand, some construct flat, fixed shelters, while others remain free-living until pupation. Many genera remain unknown in the larval stage. The family is the terra incognita of Trichoptera ( Flint 1992b) .

Rhyacophilidae : Rhyacophilidae is a relatively large family, originally established by Stephens (1836). At one time the family included also Glossosomatidae and Hydrobiosidae and other taxa, but its definition has progressively become more restricted. Evolutionary relationships of the family were discussed by Ross (1956) and the family was the subject of a large revision by Schmid (1970). The family is predominantly north temperate and is found in North America, Europe, and Asia, but also extends into India and the tropical areas of southeastern Asia. Currently most of the diversity is included in a single genus, Rhyacophila Pictet , the largest genus in Trichoptera , with over 700 species and additional ones regularly being described. In addition to the landmark works of Ross and Schmid on Rhyacophila, Prather and Morse (2001) studied the phylogeny of the R. invaria group from eastern North America and Mey (1999b) investigated the biogeography of Southeast Asian members of the genus. Other genera include Himalopsyche Banks (ca. 50 species, predominantly in the eastern Palearctic and Oriental regions, but with 1 species from western North America), Philocrena Lepneva (1 species from Georgia, western Palearctic), and Fansipangana Mey (a single species recently described from Vietnam). The family is 1 of 2 (the other being Hydrobiosidae ) that includes species that are free-living and predaceous as larvae, constructing a domed pupal chamber of rocks at maturity. As the etymology of the family name indicates, the larvae frequent cool, fast flowing rivers and streams. Larvae in the genus Himalopsyche , and some in the genus Rhyacophila , possess abdominal and thoracic gills, quite different from those in Integripalpia or Hydropsychidae .

INTEGRIPALPIA

PLENITENTORIA

Apataniidae : This is a northern and montane group found in North America, Europe, and Asia. The family names dates to Wallengren (1886), but for most of its history it was included as a subfamily of Limnephilidae . Wiggins (1996) treated the group as a distinct family and subsequent workers have accepted this designation. There are nearly 200 species in 18 genera, divided into 2 subfamilies. The Apataniinae contains the largest genus, Apatania Kolenati (nearly 100 species, Holarctic), as well as Apataniana Mosely (Palearctic, Oriental) , Apatidelia Mosely ( China) , 4 monotypic genera: Talgara Mey ( Kazakhstan) , Radema Martynov ( Russia) , Thamastes Hagen (Siberia) , Proradema Mey (Siberia) , and 5 small genera endemic to Lake Baikal: Baicalina Martynov (5 species), Protobaicalina Ivanov (4 species), Protoradema Ivanov (2 species), Baicalinella Martynov (monotypic), and Baicaloides Martynov (monotypic). The subfamily Moropsychinae contains the genera Moropsyche Banks (30 species, East Palearctic and Oriental), and Notania Mosely (5 species, Oriental). Four genera, Allomyia Banks (Nearctic and eastern Palearctic, 23 species), Manophylax Wiggins (Nearctic and eastern Palearctic, 6 species), Moselyana Denning (Oregon, monotypic), and Pedomoecus Ross (Pacific northwest of North America, monotypic), form a monophyletic group ( Gall 1994) separate from either subfamily. Apataniid larvae construct cases of small rock pieces, although Manophylax larvae also add plant pieces to the upper surface ( Wiggins 2004). Corbet (1966) documented parthenogenesis in some species of Apatania . Larvae occur in cool running waters, but at high elevations and extreme northern latitudes, some species of Apatania are found in lakes. Most larvae graze periphyton from rocks with scraper mandibles. Some species also occur in hygropetric habitats, some of which are dry for much of the year. The larvae of Moselyana are found in spring seepages, and are detritivores with toothed mandibles.

Brachycentridae : This is a Northern Hemisphere family found in both the Old and New Worlds. Ulmer (1903) originally established this group as a subfamily of Sericostomatidae . It now contains 6 genera and a little over 100 species. Three of these genera are monotypic: Adicrophleps Flint (Nearctic) , Amiocentrus Ross (Nearctic) , and Dolichocentrus Martynov (southeastern Siberia). Eobrachycentrus Wiggins ( Japan and western North America) contains only half a dozen species. Brachycentrus Curtis (ca. 30 species) and Micrasema McLachlan (ca. 75 species) are both widespread across the Holarctic and Oriental regions. Larvae construct cases from plant or rock materials, and some species use silk alone for part of the case. Several genera build 4-sided cases. The family is ecologically diverse. They inhabit running waters, but may be found in slow-flowing marshy channels. Some genera feed on aquatic moss; others are filter-feeders. Some North American species of Brachycentrus can be found in thermal streams with temperatures as high as 34°C that smell strongly of hydrogen sulfide ( Wiggins 2004).

Goeridae : This is a widely distributed family, found on all continents except South America and Australia. Ulmer (1903) originally described this group as a subfamily of Sericostomatidae . Flint (1960) and other North American workers considered it a subfamily of Limnephilidae , but other authors either always considered it a separate family ( Schmid 1980) or elevated the group to its place as a separate family ( Wiggins 1996). The Goeridae are divided into 3 subfamilies. Goerinae Ulmer contains most of the genera, each with 1 or only a few species: Archithremma Martynov (central eastern Siberia), Gastrocentrella Ulmer (Sumatra) , Silonella Fischer ( France, Spain), Gastrocentrides Ulmer ( Burma, Indonesia), Goeracea Denning and Goerita Ross (North America), and Lithax McLachlan (widespread across the western Palearctic). Silo Curtis is the second largest genus with over a dozen western Palearctic species. The largest genus Goera Stephens (ca. 130 species) is found in all biogeographic regions except the Neotropical, but with scant representation in the Afrotropics (1 species in southern Africa) and Australasia (2 species from the southwest Pacific). Larcasinae Navás contains 1 genus, Larcasia Navás (6 species, Palearctic and Oriental); and Lepaniinae Wiggins contains only 1 species endemic to northwestern North America, Lepania cascada Ross. Parker (1998) reviewed the genus Goerita , established its monophyly, and discussed the phylogenetic relationships among its 3 species. Larvae of Goeridae construct cases entirely of rock fragments; some genera incorporate larger rock fragments laterally. Most larvae live in cool running waters and are grazers on periphyton. Lepania larvae are detritivores in spring seepages ( Wiggins 1973b). Archithremma ulachensis Martynov is unusual in having a terrestrial pupa ( Levanidova & Vshivkova 1984).

Kokiriidae : McFarlane (1964) erected the plectrotarsid subfamily Kokiriinae when he described Kokiria miharo from New Zealand. Subsequently, Ross (1967) raised it to family status and included the Chilean species Rhynchopsyche fusca Schmid (originally described in Brachycentridae ) in the new family. Neboiss (1974) described Tanjistomella verna , the first record of the family in Australia; in that work he also referred the New Caledonian genus Mecynostomella Kimmins (originally placed in Sericostomatidae ) to Kokiriidae . Neboiss later described 2 more Australasian genera, Taskiria and Taskiropsyche . Flint et al. (1999) considered Rhynchopsyche fusca a junior synonym of Pangullia faziana Navás (originally described in Limnephilidae ). Johanson (2003b) recently revised Mecynostomella and nearly doubled the described species diversity of Kokiriidae , so that it now consists of 15 species described from New Zealand, New Caledonia, Chile, and Australia. The larvae are predatory, and live in sandy deposits of small streams and lakes. Larval cases are constructed from sand, and are dorsoventrally depressed and flanged around the edge. This family has been considered by various authors to be closely allied with either limnephiloid or leptoceroid families, in the latter case possibly because of the similarity of the larval cases to those of molannids and some Ceraclea (Leptoceridae) . However, the characters proposed by Frania and Wiggins (1997) to support a close relationship with Molannidae have not held up to re-examination ( Prather 2002), nor are they corroborated in recent molecular studies (Holzenthal et al. 2007, Kjer et al. 2001, 2002).

Lepidostomatidae : This family is widely distributed throughout the Northern Hemisphere, and extends southward to Panama, New Guinea, and the Afrotropical region. It was originally described by Ulmer (1903) as a subfamily of Sericostomatidae . It is divided into 2 subfamilies. The nominotypical subfamily contains 3 genera and most of the species: Hummeliella Forsslund is a monotypic genus from China; Lepidostoma Rambur contains most of the diversity in the family (ca. 380 species; Afrotropical, Australasian, Palearctic, and Nearctic); and Paraphlegopteryx Ulmer (ca. 20 species) is widespread in the East Palearctic and Oriental regions. The subfamily Theliopsychinae Weaver, 1993 contains 4 genera: Crunoecia McLachlan and Martynomyia Fischer are West Palearctic genera with only a handful of species each; Theliopsyche Banks is a Nearctic genus with half a dozen species; and Zephyropsyche Weaver is a small genus (4 species) from South and Southeast Asia. Larval cases are generally square in cross section and constructed of quadrate leaf or bark pieces. Some species build cylindrical cases of sand grains as early instars and switch to 4-sided cases as they mature; a few retain the sand grain cases throughout larval development. Larvae are generally inhabitants of cool streams and springs, but they may also occur along the shorelines of lakes. They are primarily detritivores. Weaver (1988) provided a synopsis of the North American species and a review of the world species ( Weaver 2002), where he synonymized several genera, formerly separated by secondary sexual characters of the male, with Lepidostoma . Myers and Sperling (2002) looked at the relationships of the subgenera of Lepidostoma , based on mitochondrial DNA sequence data.

Limnephilidae : This is the largest family in the Plenitentoria, with approximately 900 described species. At higher latitudes and elevations, it is the dominant group in much of the Northern Hemisphere. The family was first established by Kolenati (1848) and includes species described by Linnaeus in Systema Naturae, 10th ed. (Table 1). Schmid (1955) resolved the family into its current classification ( Table 4), with refinements by Wiggins and colleagues ( Vineyard & Wiggins 1988, Wiggins 1973a, Wiggins et al. 1985). The family is divided into 4 subfamilies, Dicosmoecinae Schmid, Drusinae Banks, Limnephilinae Kolenati, and Pseudostenophylacinae Schmid. The Dicosmoecinae , with fewer than 100 described species, are considered the most primitive of the limnephilid subfamilies, and include the only Southern Hemisphere taxa in the family; of its 19 genera, 7 are endemic to South America and 1, Archaeophylax Kimmins , is endemic to Australia ( Wiggins 2002). The Drusinae are restricted to the Palearctic region. Of the 8 genera in this subfamily, only Drusus Stephens contains more than half a dozen species; many of these are micro-endemics. Recent molecular studies have questioned the generic classification of Drusinae (Pauls et al. 2007). The nominotypical subfamily contains over 60 genera, divided into 4 tribes. Chaetopterygini Hagen, with 10 genera, are a Palearctic group with about 60 species. Chilostigmini Schmid are a group of 11 small genera, with approximately 40 Old and New World species. The tribe Limnephilini Kolenati (21 genera, ca. 300 species) includes most of the lentic genera of the Limnephilidae ; it also includes Limnephilus Leach , the most species-diverse genus, with nearly 200 described species widely distributed across the Holarctic region and as far south as Central America; 2 anomalous genera, Sphagnophylax Wiggins and Winchester , and Thermophylax Nimmo have been tentatively assigned to the Limnephilini , but this remains in some dispute ( Morse 2006). The Stenophylacini Schmid (ca. 200 species) is primarily Old World in distribution, although 4 of its 23 genera are endemic to North America; 1 genus Mesophylax McLachlan , is found in Ethiopia and Arabia ( Malicky 1998, 1999). Pseudostenophylacinae is a small subfamily of 5 genera and about 100 species ( Schmid 1990), with predominantly Oriental and Asian Palearctic distribution; the largest genus Pseudostenophylax Martynov (80 species, primarily Oriental) is represented in North America by 3 species.

This is arguably the most ecologically diverse caddisfly family, as larvae occupy the full range of habitats. Limnephilid larvae are found in lakes, streams, and marshes. Some species of Ironoquia live in temporary pools and streams. Desmona larvae have been observed leaving the water at night to feed on shoreline plants ( Erman 1981, Wiggins & Wisseman 1990), and a North American species of Philocasca Ross has an entirely terrestrial larva. Limnephilid larvae use both plant and mineral materials in their cases; the general trend in the family is that larvae in cool running waters use rock material, while those in warmer lentic habitats use plant material ( Wiggins 1996).

Oeconesidae : This is a small family of 6 genera and fewer than 20 described species. Tillyard (1921) described the family originally as a tribe of Sericostomatinae. The monotypic genus Tascuna Neboiss is found in Tasmania. The other genera, Oeconesus McLachlan (5 species), Pseudoeconesus McLachlan (9 species), Zelandopsyche Tillyard (2 species) and the monotypic genera Zepsyche McFarlane and Tarapsyche McFarlane , are endemic to New Zealand. Larval cases are of plant and rock materials ( Cowley 1978). Larvae feed on plant debris in small forested streams ( Cowley 1978, Winterbourn & Davis 1976).

Subfamily Dicosmoecinae Schmid, 1955

Allocosmoecus Banks, 1943 ....................................... Na

Amphicosmoecus Schmid, 1955 .................................... Na

Anomalocosmoecus Schmid, 1957 ................................. Neo

Antarctoecia Ulmer, 1907 ....................................... Neo

Archaeophylax Kimmins, 1953 ....................................Au

Austrocosmoecus Schmid, 1955 ................................... Neo

Cryptochia Ross, 1950 ........................................... Na

Dicosmoecus McLachlan, 1875 ................................... Hol

Ecclisocosmoecus Schmid, 1964 .................................. Hol

Ecclisomyia Banks, 1907 ........................................ Hol

Eocosmoecus Wiggins and Richardson, 1989 ......................... Na

Evanophanes Banks, 1940 ........................................ Or

Ironoquia Banks, 1916 .......................................... Hol

Metacosmoecus Schmid, 1955 .................................... Neo

Monocosmoecus Ulmer, 1906 .................................... Neo

Nothopsyche Banks, 1906 ...................................... Pa, Or

Onocosmoecus Banks, 1943 ...................................... Hol

Platycosmoecus Schmid, 1964 .................................... Neo

Verger Navas, 1918 ............................................ Neo

Subfamily Drusinae Banks, 1916

Anomalopterygella Fischer, 1966 ................................... Pa

Cryptothrix McLachlan, 1867 ..................................... Pa

Drusus Stephens, 1833 ........................................... Pa

Ecclisopteryx Kolenati, 1848 ...................................... Pa

Hadimina Sipahiler, 2002......................................... Pa

Leptodrusus Schmid, 1955 ........................................ Pa

Metanoea McLachlan, 1880 ....................................... Pa

Monocentra Rambur, 1842 ........................................ Pa

Subfamily Limnephilinae Kolenati, 1848

Tribe Chaetopterygini Hagen, 1858

Annitella Klapalek, 1907 ...................................... Pa Badukiella Mey, 1979 ........................................ Pa Chaetopterna Martynov, 1913 ................................. Pa Chaetopteroides Kumanski, 1987 ............................... Pa Chaetopterygopsis Stein, 1874 ................................. Pa Chaetopteryx Stephens, 1829 .................................. Pa Kelgena Mey, 1979 .......................................... Pa Pseudopsilopteryx Schmid, 1952 ............................... Pa Psilopteryx Stein, 1874 ....................................... Pa Rizeiella Sipahiler, 1986 ...................................... Pa

Tribe Chilostigmini Schmid, 1955

Brachypsyche Schmid, 1952 ................................... Pa Chilostigma McLachlan, 1876 ................................ Hol Chilostigmodes Martynov, 1914 ............................... Hol Desmona Denning, 1954 ...................................... Na Frenesia Betten and Mosely, 1940 .............................. Na Glyphopsyche Banks, 1904 ................................... Na Grensia Ross, 1944 ........................................ Hol

......c ontinued Tribe Chilostigmini Schmid, 1955

Homophylax Banks, 1900 ..................................... Na Phanocelia Banks, 1943 ...................................... Na Pielus Navás, 1935 .......................................... Or Psychoglypha Ross, 1944 ..................................... Na

Tribe Limnephilini Kolenati, 1848

Anabolia Stephens, 1837 ................................. Hol, Or Arctopora Thomson, 1891 ................................... Hol Asynarchus McLachlan, 1880 .............................. Hol, Or Clistoronia Banks, 1916 .......................... Na, Neo ( Mexico ) Crenophylax Ruiter and Nishimoto 2007 ......................... Na Glyphotaelius Stephens, 1833 ................................. Pa Grammotaulius Kolenati, 1848 ............................. Hol, Or Halesochila Banks, 1907 ..................................... Na Hesperophylax Banks, 1916 ................................... Na Lenarchus Martynov, 1914 .................................. Hol Lepnevaina Wiggins, 1987 .................................... Pa Leptophylax Banks, 1900 ..................................... Na Limnephilus Leach, 1815 ............................. Hol, Neo, Or Nemotaulius Banks, 1906 ................................. Hol, Or Philarctus McLachlan, 1880 .............................. Hol, Or Platycentropus Ulmer, 1905 ................................... Na Psychoronia Banks, 1916 ..................................... Na Rhadicoleptus Wallengren, 1891 ............................... Pa Rivulophilus Nishimoto, Nozaki , and Ruiter, 2001.................. Pa Sphagnophylax Wiggins and Winchester, 1984 .................... Na Thermophylax Nimmo, 1995 .................................. Pa

Tribe Stenophylacini Schmid, 1955

Acrophylax Brauer, 1867 ..................................... Pa Allogamus Schmid, 1955 ..................................... Pa Anisogamodes Martynov, 1924 ................................ Pa Anisogamus R McLachlan, 1874 ............................... Pa Chionophylax Schmid, 1951 .................................. Pa Chyranda Ross, 1944 ........................................ Na Clostoeca Banks, 1943 ....................................... Na Consorophylax Schmid, 1955 ................................. Pa Enoicyla Rambur, 1842 ...................................... Pa Enoicylopsis Navás, 1917 ..................................... Pa Halesus Stephens, 1836 ..................................... Hol Hydatophylax Wallengren, 1891 .............................. Hol Isogamus Schmid, 1955 ...................................... Pa Leptotaulius Schmid, 1955 .................................... Pa Melampophylax Schmid, 1955 ................................. Pa Mesophylax McLachlan, 1882 .............................. Pa, Or Parachiona Thomson, 1891 ................................... Pa Philocasca Ross, 1941 ....................................... Na Platyphylax McLachlan, 1871 ................................. Pa Potamophylax Wallengren, 1891 ............................... Pa Psilopterna Martynov, 1915 ................................ Pa, Or Pycnopsyche Banks, 1905 .................................... Na

......c ontinued Tribe Stenophylacini Schmid, 1955

Stenophylax Kolenati, 1848 ................................ Pa, Or

Subfamily Pseudostenophylacinae Schmid, 1955

Aplatyphylax Kimmins, 1950 ..................................... Or Astenophylina Mosely, 1936 ...................................... Or Astratodina Mosely, 1936 ..................................... Pa, Or Phylostenax Mosely, 1935 ..................................... Pa, Or Pseudostenophylax Martynov, 1909 ............................ Hol, Or Phryganeidae : Linnaeus’s original taxon is now a relatively small family confined to the more northern latitudes. Leach (1815) circumscribed the Linnaean genus Phryganea to the species P. grandis , and placed the genus Phryganea in the tribe Phryganides with Limnephilus . Burmeister (1839) was the first to use the name Phryganeidae , as a subfamily of Phryganeodea; Burmeister’s Phryganeidae included species currently placed in Sericostomatidae and Limnephilidae . By the late 19th century, most workers recognized a unit similar to the modern concept of Phryganeidae . Wiggins (1998) published a landmark treatise on the Phryganeidae , the only full-length book devoted to an entire family of caddisflies, which serves as the definitive reference. The family currently contains some 80 extant species in 15 genera. One genus, the monotypic Yphria Milne , from the Sierra Nevada of California and southern Oregon, is assigned to its own subfamily, Yphriinae . All other genera are in Phryganeinae . The nominotypical genus Phryganea , as currently recognized, contains only a handful of species from Asia, Europe, and North America. The largest genera (neither with more than 20 described species) are Agrypnia Curtis , found across the northern latitudes of Europe, Asia, and North America, and Eubasilissa Martynov , which is entirely Asian. Species in the latter genus include the largest extant caddisflies. Most genera contain only a handful of species: Banksiola Martynov (Nearctic) , Hagenella Martynov (Holarctic) , Neurocyta Navás (mountains of northern India and bordering countries), Semblis Fabricius (Palearctic) , Oligotricha Rambur (Palearctic, with 1 species extending into Alaska and the Yukon). Four genera are monotypic: Agrypnetes McLachlan (Palearctic) , Beothukus Wiggins (Nearctic) ; Fabria Milne (Nearctic) ; and Trichostegia Kolenati (northern and central Europe). Except in Yphria , which incorporates rock fragments into its case, phryganeid larvae construct cases of plant material, cut to size and fastened together in rings or a continuous spiral. Maybe because of their conspicuous size, adults of many phryganeid species have developed chemical and/or mechanical defense systems; many species produce an odiferous fluid from the anal opening when handled, and - uniquely within the Trichoptera - at least some species of Eubasilissa have urticating setae on the thorax and wings. Larvae are generally found among aquatic plants in ponds and marshes; some occur in slow flowing waters, a few are found in temporary pools and deep lake waters. Predation and herbivory are common larval feeding strategies in this family.

Phryganopsychidae : This family contains a single genus, Phryganopsyche Wiggins , with only a few Asian species found from the Himalayas to Japan and the Russian Far East. These species were originally placed in Phryganeidae (as Phryganopsis , a name preoccupied in the Lepidoptera ). The larvae, previously unknown, turned out to be very different from phryganeid larvae, and Wiggins (1959) erected a new family to accommodate these anomalous species. Wiggins and Gall (1993) concluded that the family was a “phylogenetic relict,” which could not be allied with certainty to any of the families of Plenitentoria. Wiggins (2004) placed it in its own superfamily. In the most recent molecular analysis (Holzenthal et al. 2007), Phryganopsychidae formed a clade with Kokiriidae and Pisuliidae . The larval case, constructed of plant debris, is very different from that of other case-makers: it is much longer than the larva and is not rigid, and in some ways bears more resemblance to an annulipalpian tube than an integripalpian case. Before pupation, the larva constructs a rigid case of woody materials ( Wiggins 2004). Larvae are detritivores, and live in marginal pools of streams and springs.

Pisuliidae : This small family is found in tropical Africa and Madagascar. Ross (1967) was the first to consider this group a family. It contains 2 genera, Pisulia Marlier (6 species) and Silvatares Navás (10 species); the latter was originally described in Calamoceratidae . Stoltze (1989) revised the family in its entirety. Larvae construct cases of leaf and wood pieces. Pisulia larvae occupy hygropetric habitats; Silvatares larvae are found in small streams. Larvae are detritivorous shredders.

Plectrotarsidae : This is a very small Australasian family of 3 genera and 5 species. The family was erected by Mosely (1953). At one time it included Kokiria , now in its own family. The family now consists of 2 monotypic genera, Liapota Neboiss and Nanoplectrus Neboiss , and Plectrotarsus Kolenati (3 species). The phylogenetic position of the family is equivocal ( Gall & Wiggins 1999). In the most recent molecular hypothesis (Holzenthal et al. 2007), it emerged as sister to all other Plenitentoria, but not allied with the Phryganeidae or Phryganopsychidae as indicated in other studies ( Frania & Wiggins 1997, Ivanov & Sukatcheva 2002, Wiggins 2004). Larval cases are constructed of plant pieces; the larvae are shredding detritivores, found in shallow flowing waters with abundant vegetation.

Rossianidae : This family contains only 2 species Goereilla baumanni Denning and Rossiana montana Denning , described originally in Goeridae and Limnephilidae , respectively. Gall (1997) erected the family to accommodate these 2 species after his phylogenetic analysis recovered them as a sister clade to Limnephilidae + Apataniidae , Uenoidae , and Goeridae ( Gall 1994) . Both species are known only from western North America; neither is commonly collected. Goereilla baumanni larvae are found in the organic muck of spring seeps. Rossiana montana larvae occur in stream gravel deposits under moss or in hygropetric habitats. Larvae of both species use rock fragments to construct stout, slightly curved cases.

Uenoidae : This family is found in North America, eastern Asia, and southern Europe. It was originally described by Iwata (1927) as a subfamily of Sericostomatidae . The family was revised by Wiggins et al. (1985); several taxonomic and phylogenetic works were published subsequently ( Vineyard & Wiggins 1987, 1988, Vineyard et al. 2005, Wiggins & Erman 1987, Wiggins & Wisseman 1992). The 7 genera are divided into 2 subfamilies. Thremmatinae Martynov contains the genera Neophylax McLachlan (ca. 40 species, Holarctic), Oligophlebodes Ulmer (7 species, western North America), and Thremma McLachlan (7 species, Mediterranean region). The nominotypical subfamily includes Uenoa Iwata (11 species) and 3 genera endemic to western North America: Farula Milne (11 species), Neothremma Dodds and Hisaw (7 species), and Sericostriata Wiggins, Weaver , and Unzicker (monotypic). Larvae of Thremmatinae construct stout cases of coarse rock particles (e.g., Neophylax ) or fine-grained, flattened, cases resembling the freshwater limpet Ancylus (e.g., Thremma ). Larvae of Uenoinae are more slender and construct cases of fine sand or silk ( Wiggins et al. 1985). The larval diet in Uenoidae is diatoms and fine organic particles scraped from rock surfaces. Generally, larvae are found in cool, fast-flowing headwaters; however, in the genus Neophylax , some species occur in downstream warmer waters.

INTEGRIPALPIA

BREVITENTORIA

“Leptoceroidea”

Atriplectididae : The family shows a disjunct Gondwanan distribution with species in Australia, including Tasmania, the Neotropics (northern Andes, southeast Brazil), and the Seychelles. It was erected by Neboiss (1978) to accommodate the Australasian species Atriplectides dubius Mosely , originally described in the Leptoceridae subfamily Triplectidinae and later transferred to the Odontoceridae ( Mosely & Kimmins 1953) . Upon discovery of the very unusual larval stage, Neboiss established the family and also transferred the monotypic Seychelles genus Hughscottiella auricalla Ulmer , originally described in Odontoceridae , to Atriplectididae . Neboiss (1999) described a 2nd Australasian species Atriplectides ikmaleus and Holzenthal (1997) described a new genus and species from Peru, Neoatriplectides froehlichi , such that the family now contains 4 species. The larval stages of all 3 genera are known (Holzenthal 1997, Marlier 1978, Neboiss 1978) and are unique within the Trichoptera in that the head, pro- and mesonota are narrow, elongate, and retractile. Larvae feed as scavengers by cutting a small opening in the body of dead arthropods, thus allowing them to insert their head and anterior thorax to feed on the internal tissues ( Malicky 1997). They are found in sandy bottom sediments of small streams and lakes.

Calamoceratidae : The family has long been recognized within the Trichoptera , being first established by Ulmer (1906). The nominotypical genus was included in a “section” of Leptoceridae by McLachlan and a few other early workers. The 8 genera are well defined and together comprise about 175 species from around the world. Anisocentropus McLachlan and Phylloicus Müller are the largest genera in the family with over 60 species each. The former is widespread in the Paleotropics of Africa, Asia, and Australia, with 1 outlying species in eastern North America. Phylloicus is endemic to the Neotropics, with several species extending their range into the southwestern USA. Banyallarga Navás is another Neotropical endemic of less than 20 species. The Neotropical fauna was recently revised in its entirety by Prather (2003, 2004). The other speciesrich genus in the family is Ganonema McLachlan , with about 20 species in the Oriental and eastern Palearctic regions. Smaller genera, with no more than 2 or 3 species each, include: Ascalaphomerus Walker ( China) , Calamoceras Brauer (Europe) , Georgium Fischer ( Japan, Thailand), and Heteroplectron McLachlan (eastern and western North America, Japan). Larvae of the family are well known for their flattened cases made of large pieces of excised leaves that completely camouflage the larva from above. Others build tubular cases of sand grains or hollow a twig to use as a case. The larvae inhabit the slower, depositional areas of small streams and rivers where they feed as shredders of leaf litter and other plant detritus. Larvae of a Brazilian species inhabit the “tanks” of water trapped by the leaf axils of bromeliads. Adults of many species have very brightly colored and patterned wings imparted by thickened hairs or scales. Many are more active during the day than most Trichoptera , as they engage in diurnal mating behavior.

Leptoceridae : The long-horned caddisflies comprise 1 of the 3 largest families in the order with about 1,800 described species. The family is about equal in diversity to the Hydropsychidae and only surpassed by the microcaddisflies, Hydroptilidae , in total known species richness. In all of these families, many more new species assuredly await discovery and description, especially from tropical regions around the world. The family was first established by Leach (1815) and includes several species described by Linnaeus in Systema Naturae, 10th ed. (Table 1). Nineteenth century workers also included species now in Odontoceridae , Molannidae , Calamoceratidae , and Beraeidae in Leptoceridae , but by the early 20th century modern family concepts were for the most part established. Forty-seven genera are known at present in the family, but new genera are still being described, for example Fernandoschmidia Holzenthal and Andersen (2007) , and recent generic synonymies have also occurred, for example Ylodes is a junior synonym of Triaenodes ( Holzenthal & Andersen 2004) . The family is divided into 2 subfamilies, the nominotypical subfamily Leptocerinae Leach of cosmopolitan distribution and Triplectidinae Ulmer, having a primarily Southern Hemisphere distribution in the Neotropics and Australasia, with a putative triplectidine larva in southern Africa (de Moor 1997). Two genera in particular are widespread and diverse on all continents, Oecetis McLachlan with about 400 described species and Triaenodes McLachlan with about 230 known species. Setodes Rambur , with about 220 species in very diverse in the Old World, especially India and Southeast Asia, but in the New World it is represented by only a handful of eastern North American species and 1 on Cuba. Other large genera, with about 100-150 species each, include Adicella McLachlan (Palearctic, Afrotropical, Oriental) , Athripsodes Billberg (Afrotropical, Palearctic) , Ceraclea Stephens (Nearctic, Afrotropical, Palearctic) , and Leptocerus Leach (Palearctic, Afrotropical, Oriental, with a single species in the New World). Nectopsyche Müller , with about 60 species, is a characteristic component of the Neotropical fauna; species in the genus also occur well into North America. The Neotropics is home to a number of endemic genera, including Achoropsyche Holzenthal , Amazonatolica Holzenthal & Oliveira Pes , Amphoropsyche Holzenthal , Atanatolica Mosely , Brachysetodes Schmid , Grumichella Müller , and Neoathripsodes Holzenthal. The Australasian region holds the largest diversity of endemic genera, including Condocerus Neboiss , Gracilipsodes Sykora , Lectrides Mosely , Leptorussa Mosely , Notoperata Neboiss , Russobex St. Clair , Symphitoneuria Ulmer , Symphitoneurina Schmid , Triplectidina Mosely , Triplexa Mosely , Triplexina Mosely , and Westriplectides Neboiss; all but Leptorussa are members of the Triplectidinae . Three additional triplectidine genera show a trans-Antarctic pattern between Australasia and the Neotropics, Hudsonema Mosely , Notalina Mosely , and Triplectides Kolenati , the latter the largest genus in the subfamily with about 65 species, some of which occur in India, Southeast Asia, and Japan. The Afrotropics has a rich fauna of Leptocerinae , especially within Athripsodes , Ceraclea , Leptocerus , Oecetis , and Triaenodes , but it also has several endemic genera, including Axiocerina Ross, Leptocho Barnard , Leptocerina Mosely , Blyzophilus Andersen & Kjaerandsen, Ptochoectis Ulmer , Hemileptocerus Ulmer , and Sericodes Schmid. The Oriental region has a staggering diversity of Setodes and Oecetis species, especially in India, and also harbors a few endemic genera as well, including Fernandoschmidia Holzenthal & Andersen , Leptoceriella Schmid , and Poecilopsyche Schmid. Erotesis McLachlan with 2 European and 1 Japanese species is the only genus endemic to the Palearctic region. Remaining genera within the family, all members of Leptocerinae , occur across 2 or 3 biogeographical regions and include Homilia McLachlan (Europe, Afrotropical), Mystacides Berthold (Holarctic, Oriental) , Parasetodes McLachlan (Palearctic, Afrotropical, Oriental) , Tagalopsyche Banks (Afrotropical, Oriental) , and Trichosetodes Ulmer (Palearctic, Afrotropical, Oriental) . The placement of Nietnerella Kimmins , with a single species from Sri Lanka, within the family is equivocal. Phylogenetic studies within the family, in addition to those reviewed by Morse (1997), include Calor et al. (2006), Manuel et al. (2005), Morse and Yang (2002), Stuart and Currie (2002), and Yang and Morse (2000). Larvae of the family construct a wide diversity of cases, perhaps the most diverse in the order. Cases are fundamentally tubular, but can be made entirely of silk secretions, of plant pieces arranged spirally or laid transversely, or of large leaf fragments to form a flattened case. Others make simple tubular cases of sand grains, strongly or only slightly tapered towards the posterior ends; sometimes there are larger stones placed laterally. Mystacides larvae incorporate long conifer needles or leaf stems that trail off the end of the case. Some genera make irregular cases of plant fragments, while others hollow a twig or use the abandoned cases of other caddisflies as their own. Some Ceraclea build flat, limpet-like cases of sand grains, while those that feed on freshwater sponge incorporate sponge pieces and spicules in their cases. The larvae of Leptecho helicotheca Scott from South Africa build snail-shaped cases remarkably similar to those of Helicopsyche . Larvae are found everywhere, from high mountain torrents, through all orders of streams, to large lowland rivers. In northern latitudes, they are common in lakes and in the tropics they occur in oxbow lakes and other standing waters, even temporary ones; some are semi-terrestrial and inhabit the sides of waterfalls where they are wet by the splash. Larvae feed as leaf detritus shredders, periphyton scrapers, and predators, even on freshwater sponge. Adults are often very abundant and come to lights by the 1000s. Their long, slender antennae and generally narrow forewings are distinguishing features. There are quite a few genera that have brightly colored and iridescent hairs and scales on the wings, making them among the most beautiful of caddisflies.

Limnocentropodidae : The family contains a single genus, Limnocentropus Ulmer , and 15 species occurring in India and Nepal, China, Southeast Asia (including Borneo), and Japan. The genus was created for the Japanese species L. insolitus Ulmer , and originally included in the Phryganeidae ( Ulmer 1907a) . The genus Kitagamia Iwata, and its coordinate family Kitagamiidae Tsuda, is a synonym of Limnocentropus . The family Limnocentropidae was established by Tsuda (1942) as a replacement name for Kitagamiidae, later emended to Limnocentropodidae by Kimmins (1950). Larvae live in torrential waters and attach their cases to rocks by a strong, silken peduncle, about as long as or longer than the case. The case itself is made of small rocks with silken denticles incorporated in some species. Larvae are large, robust, and predaceous. The case is positioned to extend in the current so that the larva can collect drifting insects with its strong, stout, outstretched spiny legs ( Wiggins 1969). Unlike the vast majority of Trichoptera , adults have well developed, sclerotized mandibles.

Molannidae : The family contains only 2 genera, Molanna Curtis with about 2 dozen species and Molannodes McLachlan with about 1 dozen species. Indomolannodes Wiggins is a junior synonym of Molannodes according to Malicky (2000). Members of the family were at one time included as a “section” or tribe of Leptoceridae by McLachlan and others, but were established as a family by Wallengren (1891). The family occurs across the Holarctic region as well as India, Sri Lanka, and Southeast Asia, including parts of the Indonesian archipelago. Larvae construct heavy, depressed cases of sand grains with large lateral and anterior expansions that serve to hide the larva from above and facilitate protection from predators as well as from overturning by waves ( Otto 2000) or sinking in soft sediments. They live on the sandy bottoms of lakes, often at considerable depths, and the sandy depositional areas of streams and springs. The larval food consists of algae, including diatoms, leaf litter detritus, and aquatic invertebrates. Adults of some species tend to roll the wings around the body and hold themselves at an angle while at rest, thus resembling a small twig or tiny branch.

Odontoceridae : In earlier days, the family was considered a “section” of Leptoceridae (e.g., Walker) or as a subfamily of Leptoceridae (e.g., Ulmer), but later workers considered Odontocerinae Wallengren (1891) to be a distinct family. The family contains about 115 extant species in 14 genera, scattered about the Old and New Worlds. Four genera are endemic to North America, 3 in the West ( Namamyia Banks , Nerophilus Banks , and Parthina Denning ) and 1 in the East ( Pseudogoera Carpenter ). Of the 2 additional genera found in North America, Marilia Müller reaches its greatest diversity in the Neotropics (ca. 45 species), with additional species in China, Southeast Asia, and Australia, and Psilotreta Banks occurs in eastern North America, India, Nepal, Southeast Asia, China, Korea, and Japan. In addition to many species of Marilia , the Neotropics harbors 2 endemic monotypic genera, both from southeastern Brazil, Barypenthus Burmeister and Anastomoneura Huamantinco and Nessimian. The Australasian fauna is poorly represented by only a couple of species of Marilia and 2 species of the endemic genus Barynema Banks. Southeast Asia is home to 2 endemic genera, Inthanopsyche Malicky and Lannapsyche Malicky , in addition to those mentioned above. Only the nominotypical genus, Odontocerum Leach , occurs in Europe. The family is not known from Africa, but a single monotypic genus with equivocal placement in the family ( Neboiss 1978) was described from the Seychelles, Leptodermatopteryx Ulmer. Larvae live in springs and small to medium-sized streams and rivers, some are associated with waterfalls. They seem to prefer slow flowing areas or depositional zones, where they may burrow in the sandy substrate. Cases are made of sand grains or larger mineral fragments and are very resistant to crushing due to reinforcing silken mortar applied by the larva between sand grains. Larvae are omnivorous, feeding on organic detritus, vascular plants, moss, and algae as well as aquatic arthropods.

Philorheithridae : This is another small family of about 25 species that shows a trans-Antarctic distribution, with genera endemic to Australia, Tasmania, New Zealand, or southern Chile and adjacent Argentina. There are 8 genera, with about 2-6 species each, distributed as follows: southern Chile, Argentina ( Mystacopsyche Schmid , Psilopsyche Ulmer ), southeast Australia, Tasmania ( Austrheithrus Mosely , Aphilorheithrus Mosely , Kosrheithrus Mosely, Ramirheithrus Neboiss , Tasmanthrus Mosely ), and New Zealand ( Philorheithrus Hare ). The 2 earliest named genera, Psilopsyche Ulmer and Philorheithrus Hare , were originally described in the Odontoceridae and Sericostomatidae , respectively. Mosely (1936) established the family, with Philorheithrus as its type genus. The semi-raptorial morphology of the fore- and midlegs of the larvae reveal their predatory behavior. Larvae construct stout, tubular cases of sand grains and live on or in sandy bottom sediments in small to medium-sized rivers. In the males of most species, there is a pair of “pilifers:” digitate, semimembranous structures, emerging dorsad of the maxillary palps and held in front of the face. Some males have pectinate antennae. In general, these are rather large caddisflies, with wing lengths of about 1-1.5 cm.

Tasimiidae : This small family contains only 4 genera, 2 in southern Chile and 2 in southeast Australia and Tasmania, for a total of 9 species. The Chilean fauna is composed of 2 species, each in a monotypic genus, Charadropsyche penicillata Flint and Trichovespula macrocera Schmid. Tasimia Mosely contains 5 species from southeastern Australia and Tasmania, while Tasiagma Neboiss has 1 species in Australia and Tasmania and 1 on Lord Howe Island, lying between Australia and New Zealand. The family was established by Riek (1968) for the genus Tasimia , originally described in the Sericostomatidae ; Trichovespula was first included in Lepidostomatidae , and later transferred to Tasimiidae by Flint (1969). Larvae all seem to build narrow to broad, flattened, tubular cases of sand grains with anterior and lateral flanges of larger mineral fragments ( Flint 1967, 1999). They live in small, shallow streams where they cling to the faces of rocks. They probably feed on periphyton and organic sediments growing or adhering to the rock surfaces.

INTEGRIPALPIA

BREVITENTORIA

Sericostomatoidea

Anomalopsychidae : The family was established by Flint (1981) for 2 Chilean species formerly included in the Sericostomatidae : Contulma cranifer Flint and Anomalopsyche minuta Schmid. It is the only caddisfly family fully endemic to the Neotropics and now contains 26 species ( Holzenthal & Flint 1995, Holzenthal & Robertson 2006), distributed in the mountainous regions from Costa Rica south to Chile and in the highlands of southeastern Brazil. The larvae of both genera have been described ( Flint 1981, Holzenthal & Flint 1995) and inhabit seeps, spring-runs, and small to medium-sized streams in forested areas as well as those above the tree line in the northern Andes. Many species frequent the splash zone of waterfalls and cascades, where they are often found in aquatic moss. The larvae have scraping mandibles and more than likely feed on periphyton. They build cylindrical, slightly curved cases of sand grains. As a whole, members of the family are rare and infrequently collected; adults fly to lights, but are as easily collected with an aerial net during the day.

Antipodoeciidae : The family is endemic to eastern Australia and still contains a single species, Antipodoecia turneri Mosely. The species was originally described in the Sericostomatidae , but Ross (1967) established a new family, Antipodoeciidae , to accommodate it in his attempt to rectify the then polyphyletic composition of the Sericostomatidae , sensu lato. It is a small insect with a forewing length of only 3.5 mm; nothing of substance has been published on its biology. The larvae build slightly curved and tapered, cylindrical cases of sand grains. It is the only caddisfly family not yet included in a molecular phylogenetic analysis of family relationships within the order.

Barbarochthonidae : This endemic South African family was established by Scott (1985, 1993) to accommodate a single species, Barbarochthon brunneum Barnard , first described in the Sericostomatidae ( Barnard 1934) . The small dark brown adults have a conspicuous cream colored pronotum and are common in the western and southern Cape region at mid- to high elevations. Larvae occur in both fast-flowing torrents as well as pools, and are typically associated with marginal vegetation, including clumps of submerged Scirpus ; they feed as leaf detritivores. The long, slender, curved, tapered case is made entirely of darkened silk except for some transverse rows of small sand grains towards the posterior end.

Beraeidae : This small family, established by Wallengren (1891), is comprised of 7 genera and about 50 species. It reaches its greatest diversity in the western Palearctic region, where 5 genera occur ( Beraea Stephens , Beraeamyia Mosely , Ernodes Wallengren , Beraeodes Eaton , and Beraeodina Mosely , the latter 2 monotypic). The genus Beraea also occurs in eastern North America, where 3 species are known. A single genus, Notoernodes , with 2 species, occurs in Tanzania. Elsewhere the family is found only in Japan, where it is represented by the genus, Nippoberaea Botosaneanu, Nozaki , & Kagaya, containing a single species, N. gracilis (Nozaki & Kagaya) . Most beraeid larvae build slightly to strongly tapered cases of small sand grains and live in springs, seeps, and small streams, usually among aquatic mosses, leaf litter deposits, roots of emergent plants, and other dense, marginal vegetation or in the marginal organic sediments ( Hamilton 1985). Gut contents of Beraea gorteba Ross in southeastern North America included small pieces of vascular plants, fungal mycelia, and other organic material, but no animal parts ( Hamilton 1985), foodstuffs probably typical of most species in the family.

Calocidae : This is another family, endemic to Australia and New Zealand, established by Ross (1967) to accommodate genera originally placed in Sericostomatidae , Beraeidae , or Odontoceridae . As with other new families established in this paper, Ross gave no family diagnosis or indication of included genera. In the same paper, he placed Pycnocentrella eruensis Mosely , originally described in Beraeidae , in his newly created family Pycnocentrellidae, later synonymized with Calocidae by Neboiss (1977). The New Zealand genus Alloecentrella Wise , first described in Beraeidae and at times included in the Helicophidae was formally transferred from Calocidae to Helicophidae by Henderson and Ward (2007); their phylogenetic analysis revealed its close affinity to other helicophid genera. In its present composition, the family now contains 6 genera endemic to Australia ( Caenota Mosely , Caloca Mosely , Calocoides Neboiss , Pliocaloca Neboiss , Tamasia Mosely ) and 1 endemic to New Zealand ( Pycnocentrella ), with a total of 19 species. One Tasmanian species is terrestrial and lives under moss and leaf litter in wet sclerophyll forest, but others occur in small, woodland streams among plant roots and accumulations of detritus ( Jackson 1998, Neboiss 1979). They construct slightly curved and tapered cylindrical cases of small rock fragments or somewhat flattened cases of 2 dorsal and 2 ventral rows of leaf material ( Jackson 1998). Adults are small to rather large (forewing lengths 5-14 mm) and have dark forewings patterned with irregular white spots. In males of some genera, the antennal scape has expandable lobes and the head bears long, expandable filaments ( Neboiss 1986).

Chathamiidae : The family was first described as a subfamily within Rhyacophilidae by Tillyard (1925) to accommodate the species Chathamia brevipennis Tillyard , a species endemic to the Chatham Islands, a group of 10 small islands 800 km east of New Zealand. It was later moved to the Philanisidae by Wise (1965), a family erected by Mosely (in Mosely & Kimmins 1953) to accommodate Philanisus plebeius Walker , first described in Hydropsychidae and later included in Sericostomatidae by Ulmer (1907b). Riek (1976) synonymized Philanisidae with Chathamiidae and provided a comprehensive review of the family, including a discussion of phylogeny and the description of 2 new species. Ward (1995) described a 3rd species of Philanisus , such that the family contains 5 species, distributed as follows: Chathamia brevipennis Tillyard (Chatham Islands), C. integripennis Riek ( New Zealand), Philanisus plebeius Walker ( New Zealand, southeast Australia), P. fasciatus Riek (Kermadec Islands, ca. 1000 km NNE of New Zealand), and P. mataua Ward ( New Zealand). Ulmer (in Mosely & Kimmins 1953) described the larva of P. plebeius , long known to inhabit marine intertidal rock pools ( Hudson 1904), but it was not until the late 1970s when the remarkable biology and life-history of the species was fully revealed ( Anderson & Lawson-Kerr 1977, Winterbourn & Anderson 1980). The larvae and pupae of the species, and assumed all members of the family, are among the very few fully marine insects. The females oviposit through the papular pores of intertidal starfish where the eggs undergo embryonic development. After hatching, the 1st instar larvae leave the starfish through the same pores and construct cases of calcareous algae. Larvae feed on non-calcareous Rhodophyceae. Adult females have long oviscapts, probably facilitating the insertion of eggs in the starfish. Adults of the Chatham Island species are brachypterous, but those of other species are fully winged.

Conoesucidae : Ross (1967) established the subfamily Conoesucinae within the Sericostomatidae for Australasian genera with “atrophied” scutal warts, but he did not name the included genera or offer any other diagnosis. Later, Neboiss (1977) elevated the subfamily to family status, provided a detailed diagnosis, and included 6 Australasian genera in the family, all formerly included in the Sericostomatidae . Additional sericostomatid genera have been transferred to Conoesucidae such that the family now contains a dozen genera and ca. 40 species, endemic to either southeastern Australia and Tasmania ( Coenoria Mosely , Conoesucus Mosely , Costora Mosely , Hampa Mosely , Lingora Mosely , Matasia Mosely ) or New Zealand ( Beraeoptera Mosely , Confluens Wise , Olinga McLachlan , Periwinkla McFarlane , Pycnocentria McLachlan , Pycnocentrodes Tillyard ). The larvae live in small, cool, fast-flowing streams where they feed on leaf litter detritus, algae, and moss. Their cases are made of sand, small rocks, plant parts, or silk; cases are tubular and only slightly curved. Adult males have shortened, membranous maxillary palps that are held out in front of the face. Ward (1995) reported that adults of a New Zealand species of Pycnocentria were common and active during hot summer days on streamside sedges, herbs, and grasses.

Helicophidae : The helicophids are a small family of caddisflies found in Australia, New Zealand, and now New Caledonia as well as austral South America (southern Chile and adjacent Argentina), one of several caddisfly families showing this trans-Antarctic biogeographical distribution pattern. The family was created by Mosely (in Mosely and Kimmins 1953) to accommodate a new genus and 2 new species, Helicopha astia Mosely and H. hortena Mosely , both from New South Wales. Additional genera have been transferred to the family from Beraeidae , Calocidae , and Sericostomatidae and described within the family itself, the most recent of these being Briama Johanson and Ward from New Caledonia and Heloccabus Neboiss from eastern Australia, the later placed provisionally in the family ( Johanson & Ward 2001, Neboiss 2002). In addition, the endemic New Zealand genus Alloecentrella was just transferred to the family from Calocidae by Henderson & Ward (2007). Thus, the current accounting of the 11 genera in the family, for a total of about 35 species, is as follows: Alloecella Banks ( Australia) , Alloecentrella Wise ( New Zealand), Alloecentrellodes Flint ( Chile) , Austrocentrus Schmid ( Argentina, Chile), Briama Johanson and Ward ( New Caledonia), Eosericostoma Schmid ( Argentina, Chile), Helicopha Mosely ( Australia, New Caledonia), Heloccabus Neboiss ( Australia) , Microthremma Schmid ( Chile) , Pseudosericostoma Schmid ( Chile) , and Zelolessica McFarlane ( New Zealand). Henderson (2007), Johanson (2003a), and Neboiss (2002) studied phylogenetic relationships among some members of the family; interestingly Henderson & Ward’s (2007) cladogram did not group Heloccabus with other helicophids included in their data matrix. The current placement of many genera in the family is equivocal and a revision of the entire family and others in the Sericostomatoidea is needed ( Flint 1992a). Helicophid larvae build tubular cases of sand grains, plant material, including almost entirely of pieces of moss, or entirely of silk; some Chilean species construct broad, flat cases of small mineral fragments. They live in clear, fast flowing, forested streams and spring runs, often associated with aquatic moss. Adults of many species are small, rare, and infrequently collected, although those of the Chilean genus Eosericostoma are common and widespread (Flint 1992).

Helicopsychidae : The snail-case caddisflies of the family Helicopsychidae were first recognized as the subfamily Helicopsychinae of Sericostomatidae by Ulmer (1906) and were retained there by a number of European workers well into the 1950s, most notably Ulmer himself ( Ulmer 1955). Ross (1944) and other American workers considered the group a distinct family, reflecting its current status. As presently constituted, the family contains only 2 genera, the cosmopolitan Helicopsyche von Siebold with about 250 species, and the New Zealand endemic genus Rakiura McFarlane , with a single species, R. vernale McFarlane. Several previously recognized genera, including Cochliopsyche Müller (Neotropical) , Petrotrichia Ulmer (Afrotropical, including Madagascar and the Seychelles, but absent from southern Africa), and Saetotrichia Brauer ( Australia, New Zealand, New Caledonia), were relegated as subgenera of Helicopsyche by Johanson (1998). In the same paper, Johanson described 2 additional subgenera of Helicopsyche: Feropsyche (Nearctic, Neotropical) and Galeopsyche ( Korea, Vietnam). The nominotypical subgenus occurs in the Palearctic and Oriental regions. As a whole the family is poorly represented in the Northern Hemisphere, but reaches its greatest diversity in the tropics of the Old and New Worlds (Johanson 1997); the Neotropics alone hosts about 100 species. Larvae of the genus are the familiar and remarkable snail-case builders. These helical, sand grain cases are so similar to snails that early workers described these insects as molluscs. Lea (1834) went so far as to say of Valvata arenifera (= Helicopsyche borealis ), “It has the singular property of strengthening its whirls by the agglutination of particles of sand, and by which it is entirely covered.” While all helical, there is great diversity in the height of cases, the number and openness of the whorls, the size of mineral material, and the amount of silk incorporated. All helicopsychid larvae appear to feed as scrapers on periphyton and other organic matter on the exposed surfaces of rocks. They are found in slow flowing lowland streams as well as springs, small fast-flowing streams, and the wave-washed shores of lakes in temperate regions; they also occur in the hyporheic zone ( Williams et al. 1983) and in thermal springs (Resh et al. 1984). The biology of the North American species, H. borealis (Hagen) is well known ( Vaughn 1985a, b, 1987). Since Morse’s (1997) review of phylogenetic studies within the Trichoptera, Johanson has undertaken significant analyses of evolutionary relationships within Helicopsyche ( Johanson 1998, 2001, 2002, Johanson & Willassen 1997).

Hydrosalpingidae : The family contains only a single species Hydrosalpinx sericea described by Barnard (1934). He placed the species only within the “Aequipalpia” in the “neighbourhood of Molannidae- Beraeidae ,” but not within a specific family. Fischer (1970) had the genus listed under Helicopsychidae , but noted Scott’s (1967) opinion that it may belong to the Beraeidae . It was not until 1985 that a new family, Hydrosalpingidae , was established for the genus ( Scott 1985), which was more fully diagnosed and described some years later ( Scott 1993). The species is endemic to the western and southwestern Cape Province of South Africa. Once common in cool acidic mountain streams, the species is now considered rare, possibly due to predation by introduced trout. Larvae feed on algae and detritus. Their tubular cases characteristically have a slightly flared anterior end and are made entirely of golden-brown silk. The discarded cases are often occupied by species of Athripsodes (Leptoceridae) . Adults are medium-sized, densely hairy, golden brown caddisflies. Males have unusually long and slender maxillary and labial palps.

Petrothrincidae : The family was established by Scott (1985) for 2 species from South Africa, Petrothrincus circularis Barnard and P. triangularis (Hagen) (originally described from the case only and included in Molanna ); Scott (1993) later described a 3rd South African species, P. demoori and expanded the diagnosis and description of the family. All of the South African species are endemic to the Cape Province. In his original description of the genus, Barnard (1934) could not place Petrothrincus within any known family and referred it only to “Aequipalpia.” Fischer (1964, 1972) and others continued to catalog the genus within the Molannidae . Weaver (1997) recorded the family from Madagascar for the first time and described 3 additional species in a new genus, Gyrocarisa Weaver , to which 2 other species were added later by Malm and Johanson (2005). Recently, Gyrocarisa was synonymized with Petrothrincus by Johanson and Olah (2006), who described 5 additional Malagasy species. As of now, the family contains a single genus, Petrothrincus , containing 14 species from South Africa and Madagascar. The larvae live in small, cool streams in the mountains and foothills in fast flowing riffles as well as pools. They construct broad, depressed, limpet-like cases similar to those seen in European Thremma , some North American Ceraclea , and in Chilean Eosericostoma ; at least among the South African species the case can be circular or triangular. Larvae feed as scrapers on periphyton and the organic sediment that collects on the surfaces of rocks. Weaver (1997) noted several modifications of the female abdomen and wings, including long hairs on the hind wings, for holding the large eggmass above the abdomen and between the folded wings, possibly in a protective posture.

Sericostomatidae : The family was established by Stephens (1836) as Sericostomidae (later emended by McLachlan 1874 to Sericostomatidae ). Over the years, the family has been used as a “dumping ground” for genera unable to be placed with confidence in other families. Fischer (1970) listed 26 genera in Sericostomatidae and stated, “Several of these genera may belong to the Lepidostomatidae , a few others probably to the Beraeidae . For some of the genera from the Australian region one or more subfamilies will have to be created.” In fact, all of the Australian genera once included in Sericostomatidae have been moved to other families, most newly created for them (e.g., Antipodoeciidae , Conoesucidae , Tasimiidae ), such that the family no longer occurs in the Australasian region. In other regions, other families were established for genera originally described in Sericostomatidae (e.g., Anomalopsychidae from the Neotropics, Barbarochthonidae from South Africa). As presently constituted the family contains 19 genera and only 100 or so species. The distribution of these genera is cosmopolitan, except for Australia, New Zealand, and their biogeographically associated islands, but the genera are for the most part restricted within their regions. In Africa, the family occurs only in South Africa where 5 endemic genera occur ( Aclosma Morse , Aselas Barnard , Cheimacheramus Barnard , Petroplax Barnard , and Rhoizema Barnard , the later also recently described from Madagascar). In the Neotropics, the genera are endemic to southern Chile and adjacent Argentina ( Chiloecia Navás , Myotrichia Schmid , Notidobiella Schmid , and Parasericostoma Schmid ) and to southern and southeastern Brazil and adjacent Argentina ( Grumicha Müller ). In North America, 2 genera are endemic to the eastern half of the continent ( Agarodes Banks and Fattigia Ross ) and 1 genus, Gumaga Tsuda occurs in the western portion of the region. Gumaga is also found in the Oriental region where a genus endemic to India also occurs ( Asahaya Schmid ). Five genera occur in the western Palearctic region from northern and southern Europe, northern Africa, east to the Caucasus, Iran, and the Arabian peninsula ( Cerasma McLachlan , Notidobia Stephens , Oecismus McLachlan , Schizopelex McLachlan , and Sericostoma Latreille ). In addition to the 19 genera formally assigned to the family, several additional anomalous genera are known within the superfamily Sericostomatoidea that have not been assigned to a family. For completeness of coverage, these genera are: Ceylanopsyche Fischer from Sri Lanka, Karomana Schmid from India, Mpuga Schmid from India, Ngoya Schmid from India, and Seselpsyche Malicky from the Seychelles. Schmid (1993) and Malicky (1993) discuss the status of these enigmatic genera. The larvae of Sericostomatidae build tubular, strongly to slightly curved and tapered cases of sand grains or of silk alone. In Brazil, the long, slender silken cases of Grumicha were used as adornments by the Tupí-Guarani Indians. Sericostomatid larvae inhabit streams and lakes, the latter especially in temperate regions; they often burrow in sandy deposits. The primary food source is leaf litter detritus. Males of many species have modified antennal scapes with scent scales or scent glands, eversible glands on the face, or mask-like maxillary palps, or a combination of these.