Skip to main content
NCBI home page
ZooKeys logoLink to ZooKeys
. 2025 Dec 10;1263:47–68. doi: 10.3897/zookeys.1263.148069

Functional traits of ancestral caddisfly (Trichoptera) larvae and pupae

Xinyu Ge 1, John C Morse 2,✉
PMCID: PMC12712640  PMID: 41426527

Abstract

Recent phylogenomic studies have concluded that the ancestor of order Trichoptera and suborder Integripalpia probably had a larva that was “free living,” without a portable case or fixed retreat. Phylogenies inferred from those investigations regarding hypotheses for other probable functional traits of larvae and pupae of the Trichoptera ancestor and its immediate descendants were considered, especially with reference to the extant amphiesmenopteran sister lineage Lepidoptera. To test our hypotheses an Ancestral Character State Reconstruction by Parsimony Analysis was performed to explore functional traits for five habitat and behavioral traits. Like the larva of Micropterigidae, the basal lineage of Lepidoptera, the ancestral caddisfly larva was not only “free living” but also was a shredding herbivore of bryophytes. Like that larva, it may have been often submerged, perhaps as a semi-aquatic sprawler in madicolous or hygropetric habitats, but it could also have been a clinger in lotic-erosional habitats. Also, the characteristics of the pupal cocoon are not clear; it may have been closed and permeable like that of Micropterigidae, or it was closed and semipermeable like that of Hydroptilidae, or it was open in a long-dome shelter like that of the Annulipalpia ancestor.

Key words: Bryophytes, case, cocoon, pupa, retreat, silk

Introduction

Ecologically, larvae of caddisflies (the insect order Trichoptera) are more diverse than those of almost any other order of freshwater macroinvertebrates and are similar in ecological diversity to larvae of freshwater flies, or Diptera (Merritt et al. 2019). Larvae of the megadiverse moths, the trichopteran sister order Lepidoptera according to Misof et al. (2014), are mostly terrestrial herbivores. Therefore, a long-standing question has been how caddisfly larvae made the very dramatic evolutionary transition from a purely terrestrial habitat to an aquatic habitat. What did the ancestral caddisfly larvae look like? What did they eat? In what habitat did they live and what behaviors allowed them to live there successfully? In what type of enclosure did they pupate? With the less-well-developed phylogenetic evidence known at the time of their studies, Milne and Milne (1939) thought ancestral Trichoptera larvae inhabited ponds and slow streams and Ross (1956) concluded “that the primitive caddisflies were cool-adapted forms living in running streams.” Either of these ideas would have required multiple morphological, physiological, and behavioral adaptations to have evolved simultaneously, a seemingly improbable scenario. With regard to pupae, Wiggins (2004) inferred that in the ground plan of Trichoptera “pupation [occurred] at [the] end of [the] final larval instar in [a] domed enclosure, open to moving water” and beneath this dome they constructed a “closed, semipermeable cocoon free from [the] wall of [the] dome.”

What were the functional traits of ancestral caddisfly larvae and pupa and what were the structural and functional evolutionary characteristics that enabled their successful invasion of resource-rich freshwater habitats? Recent revisions of moth phylogeny provided by Mitter et al. (2017) and Kawahara et al. (2019) placed Micropterigidae and Agathaphagidae as the earliest successive lineages of Lepidoptera. Recent revisions of caddisfly phylogeny also were provided by Thomas et al. (2020), Frandsen et al. (2024), and Ge et al. (2024a), placing Hydropsychoidea as the basal lineage for all other Annulipalpia and Hydroptiloidea as the basal lineage for all other Integripalpia. These recent authors also determined the time of origin for those orders as Late Carboniferous (~300 MA) for Lepidoptera (Kawahara et al. 2019) and Permian (~275 MA by Thomas et al. 2020; ~295 MA by Frandsen et al. 2024; 281.16–302.52 MA by Ge et al. 2024a) for Trichoptera. Through the investigations of many colleagues summarized in publications such as those by Graf et al. (2008) and Merritt et al. (2019), we also have a much improved understanding of the functional traits of the basal lineages of Lepidoptera and Trichoptera and their included families.

Based on those resources, we presented our hypothesis at the 18th International Symposium on Trichoptera (Quito, Ecuador, 1–5 July 2024) that the larva of the ancestral caddisfly was not only free-living (Wiggins 2004; Frandsen et al. 2024; Ge et al. 2024a), but also a shredding herbivore feeding on bryophytes in wet and sometimes submerged or hygropetric environments. Reviewers for the manuscript submitted for the Proceedings of the symposium encouraged us to conduct a parsimony analysis to test this hypothesis, an analysis which we agreed to undertake.

Currently, the superorder Amphiesmenoptera Kiriakoff, 1948, consists of four orders and ~211 extant and fossil (†) families (Morse 2025; van Nieukerken et al. 2011):

† Order Tarachoptera Mey, Wichard, Müller & Wang, 2017a;

† Family Tarachocelidae Mey, Wichard, Ross & Ross, 2017b;

† Order Protomeropina Tillyard, 1926;

† Family Cladochoristidae Riek, 1953;

† Family Kalophryganeidae Haupt, 1956;

† Family Karaungiridae Novokshonov & Sukatsheva, 1997;

† Family Microptysmatidae Martynova, 1958;

† Family Protomeropidae Tillyard, 1926;

† Family Terminoptysmatidae Melnitsky & Ivanov, 2020 (in Kopylov et al. 2020);

† Family Uraloptysmatidae Ivanov, 1992;

Order Trichoptera ~51 extant fam., 14 fossil fam.;

Order Lepidoptera ~134 extant fam., 4 fossil fam.

Among these four orders, Mey et al. (2017b) and Mey and Wichard (2023) considered Lepidoptera and Trichoptera (and their respective stem groups and sister groups) as sister lineages. For Tarachoptera and Protomeropina, DNA sequences are unavailable for phylogenetic analysis, and nothing is known about the structures, functions, and habitats of their immature stages.

Order Lepidoptera

Among the families of Lepidoptera, Mitter et al. (2017) considered that Micropterigidae is probably the basal lineage but said that “the question is not fully settled.” If correct, however, then they concluded that the gymnosperm-seed-feeding family Agathiphagidae is the next successive family in the order and the sister family of all other Lepidoptera, the clade Angiospermavora. The phylogenomic study of Kawahara et al. (2019) supported these relationships.

Larvae of micropterigids feed on foliose liverworts, mosses, or detritus and live in or on the ground (Mitter et al. 2017) without constructed cases or shelters (i.e., are “free living”). Davis and Landry (2012) provided further details for two of the three species of the North American micropterigid genus Epimartyria: E. auricrinella Walsingham, 1898, and E. pardella (Walsingham 1880). Epimartyria auricrinella larvae feed on liverworts (Bazzania trilobata) in shaded, wet locations (swampy woods, boggy ditches, creek margins), “which probably differed little from those of ancestral ‘Amphiesmenoptera’ ” (Kristensen 1997), and that can be flooded periodically or seasonally. They are peripneustic with spiracles on abdominal segments I–VIII and a plastron supporting a thin layer of air on each side around the spiracles and over the posterior pronotum, permitting survival when submerged for at least short periods. Eggs of E. pardella are laid on the ventral surfaces of liverwort (Conocephalum conicum) and larvae also spend days on the undersides of liverwort thalli; “pupation occurs within a thin-walled, tightly woven brown cocoon close to the ground and attached to vegetation with strands of coarse silk” (Davis and Landry 2012); no other pupal shelter was mentioned. For basal lineages of Lepidoptera, silk production is confined to construction of cocoons by last-instar larvae (Kristensen 1997).

Order Trichoptera

In multigene and phylogenomic studies, Thomas et al. (2020), Frandsen et al. (2024), and Ge et al. (2024a) all concluded that Trichoptera and suborders Annulipalpia and Integripalpia are each monophyletic. Furthermore, they concluded that the relationships within Annulipalpia were (Hydropsychoidea (Philopotamoidea + Psychomyioidea). Within Integripalpia, those three studies also concluded that the basal lineage is a clade of Hydroptiloidea sister families Ptilocolepidae and Hydroptilidae (or Hydroptilidae only, Ptilocolepidae having not been studied by Ge et al. 2024a). Frandsen et al. (2024) and Ge et al. (2024a) also inferred that the remaining families of Integripalpia are a clade with relationships (Phryganides (Glossosomatidae + Rhyacophiloidea)), for which subterorder Phryganides Latreille, 1805, consists of all families with tube-case-making larvae.

In modern Annulipalpia, campodeiform larvae usually construct silken retreats in all instars to provide physical protection from predators, usually attached to stable substrate surfaces in lotic waterways. These retreats may be augmented with specialized filter nets to one side of the upstream end (Hydropsychoidea and Stenopsychidae) or much of the inner surface of the retreat itself may filter food particles (Philopotamidae and some Psychomyioidea). Some Psychomyioidea larvae use their retreats as snares resembling spider webs for capturing drifting prey, as lairs from which to pounce on unsuspecting prey or to graze on periphyton, as reinforcement for subterranean tunnels with filter nets (Wiggins 2004), or as substrate for culturing algae (Ings et al. 2010, 2017).

In modern Integripalpia, Hydroptiloidea (Hydroptilidae + Ptilocolepidae) larvae undergo hypermetamorphosis, with the first four instars having a typical campodeiform shape (Nielsen 1948) and then, in the fifth and final instar, becoming hypergastric, developing a much-enlarged abdomen. Notably, the early-instar campodeiform larvae are “free-living,” with only the last instar constructing a sagitally compressed or transversely depressed case of silk or of silk and plant or mineral substrate sealed laterally on two longitudinal seams; this case is usually portable but sometimes fixed to substrate (Wiggins 2004). In particular, much like at least some lepidopteran micropterigid larvae, Ptilocolepidae larvae shred living mosses and liverworts in hygropetric or madicolous, damp streamside habitats, typically above the water line. Furthermore, the final instar, with its much-enlarged abdomen, constructs a portable, transversely depressed case of pieces of liverwort (Thienemann 1904; Ito and Higler 1993; González et al. 2000; Wiggins 2004; Ito et al. 2014). The final instar of a hydroptiloid larva pupates in a silken cocoon that is either semipermeable (Hydroptilidae) or permeable (Ptilocolepidae) (Wiggins 2004).

In all larval instars, Glossosomatidae larvae construct so-called “tortoise” cases of mineral pieces from which they graze periphyton off the upper surfaces of rocks in rapidly flowing water. Each glossosomatid case consists of a dome-like dorsal covering attached laterally to a flat, transverse, ventral strap or “plastron” that leaves identical, interchangeable openings for the head and legs at one end and the tip of the abdomen at the other; prior to pupation, the fifth (last) larval instar removes the ventral strap, seals the remaining dome to the rock, and spins a semipermeable cocoon beneath it (Wiggins 2004). Campodeiform Rhyacophiloidea (Hydrobiosidae + Rhyacophilidae) larvae build no cases until just prior to pupation but roam freely over the substrate; they feed generally as predators, but sometimes scrape periphyton, collect fine particulate organic matter, or shred living plant tissue (Cummins et al. 2019). The dome-like shelter built by the mature larva resembles that of Glossosomatidae, under which the rhyacophiloid larva also spins a semipermeable cocoon before pupating (Wiggins 2004).

Tube-case making Phryganides larvae typically construct their cases in all instars. The earliest lineages of infraorder Plenitentoria Weaver, 1984, do so with angiosperm pieces and those of infraorder Brevitentoria Weaver, 1984, make cases with mineral materials. When preparing for pupation, they usually attach their larval cases to stable substrate and close the ends with tough silken mesh sieves to deter predators while allowing flow of oxygenated water (Wiggins 2004).

With apparent radiation of so much structural and functional diversity in freshwater ecosystems, we were curious to learn how the ancestral caddisfly larva invaded its aqueous habitat and what evolutionary adaptations allowed it to succeed in that environment.

Methods

For each of the families for which phylogenetic relationships were inferred by Ge et al. (2024a) and with the addition of Ptilocolepidae (from Frandsen et al. 2024), and the basal lineages of Lepidoptera (Mitter et al. 2017; Kawahara et al. 2019), we determined from the most recent available studies the basal lineages of Lepidoptera and families of Trichoptera and the traits of those lineages (Tables 1–3). From among these, we selected five trait categories of the larvae, including the Case-Retreat, Food (= Feeding or Trophic Relationships), Habitat, Cocoon Construction, and Habit using terminology mainly from Cummins et al. (2019). For each trait category, the known trait of the ancestral lineage of each family was encoded for analysis as in Tables 1–3.

Table 1.

Ancestral traits for case-retreat and food-feeding-trophic relationships of larvae of basal taxa of Lepidoptera (Kawahara et al. 2019) and the Trichoptera families for which phylogenetic relationships were inferred by Ge et al. (2024a), with Ptilocolepidae from Frandsen et al. (2024), showing color codes for inferred ancestral nodes in Fig. 1.

Taxon Case-Retreat1 (Trait 1) Codes for Trait 1 Food-Feeding-Trophic Relationships2 (Trait 2) Codes for Trait 2 Reference(s) for Inferred Basal Lineage(s) of Terminal Taxa Reference(s) for Traits of Basal Lineage(s)
Angiospermavora None (free living, fl). 0 (white) Shredding herbivores (angiosperms) (shh). 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Stehr 1987 (Angiospermavora families).
Heterobathmiidae None (free living, fl). 0 (white) Shredding herbivores (Nothofagus, angiosperms) (shh). 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Kristensen and Nielsen 1983.
Agathiphagidae None (free living, fl). 0 (white) Shredding herbivores (Kauri seeds, gymnosperms) (shh). 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Dumbleton 1952.
Micropterigidae None (free living, fl). 0 (white) Shredding herbivores (bryophytes) (shh). 5 (black) Mitter et al. 2017, Kawahara et al. 2019. Davis and Landry 2012.
Ptilocolepidae None (free living early instars) →portable rigid purse case (last instar; fl, pc). 0 (white) Shredding herbivores (bryophytes) (shh). 5 (black) Palaeagapetus and Ptilocolepus (the only genera). Ito et al. 2014; Waringer and Graf 2002.
Hydroptilidae None (free living early instars) →portable rigid purse case (last instar; fl, pc). 0 (white) Grazers (gra). 1 (brown) Stactobia (Marshall 1979). Graf et al. 2008; Waringer and Graf 2011.
Glossosomatidae Portable rigid tortoise case (ps). 4 (black) Grazers (gra). 1 (brown) Anagapetus and Glossosoma (Robertson and Holzenthal 2013). Wiggins 1996; Cummins et al. 2019.
Hydrobiosidae None (free living, fl). 0 (white) Predators (pre). 2 (green) Apsilochorema (Ward et al. 2004). Holzenthal et al. 2007.
Rhyacophilidae None (free living, fl). 0 (white) Predators (pre). 2 (green) Fansipangana (larva unknown) and Rhyacophila (McLaughlin et al. 2019). Graf et al. 2008.
Phryganopsychidae Limp plant tube case (pl). 1 (brown) Shredding detritivores (shd). 3 (yellow) Phryganopsyche (only genus). Wiggins 2004.
Pisuliidae Rigid plant tube case (plant, pl). 1 (brown) Shredding detritivores (shd). 3 (yellow) Pisulia and Silvatares (Stoltz 1989). Stoltz 1989.
Phryganeidae Rigid mineral and plant tube case (mn and pl). 1 (brown) Predators (pre). 2 (green) Yphria (Wiggins 1998). Wiggins 1998.
Brachycentridae Rigid plant tube case (pl). 1 (brown) Shredding herbivores (shh). 0 (white) Eobrachycentrus (Flint 1984). Wiggins 1965, 1996.
Lepidostomatidae Rigid tube case (mineral or plant, mn or pl). 1 (brown) Shredding detritivores (shd). 3 (yellow) Basal lineages of Lepidostomatinae and Theliopsychinae have not been inferred. Wiggins 2004; Cummins et al. 2019.
Uenoidae Rigid silk tube case (mineral, mn; si). 2 (green) Grazers and gatherers (gra, gat). 1 (brown) Sericostriata (Wiggins et al. 1985). Wiggins et al. 1985.
Goeridae Rigid mineral tube case (mn). 2 (green) Grazers (gra). 1 (brown) Basal lineages of Goerinae, Larcasinae, and Lepaniinae have not been inferred. Traits are almost universal in the family. Wiggins 1996, 2004.
Apataniidae Rigid mineral tube case (mn). 2 (green) Grazers (shd). 1 (brown) Apataniana (Ge et al. 2024b). Waringer and Malicky 2016.
Limnephilidae Rigid mineral tube case (mn; plant, pl). 2 (green) Shredding detritivores; gatherers (shd, gat). ? (stripe) Ecclisomyia, Philocasca (Vshivkova et al. 2007). Wiggins 2004; Cummins et al. 2019.
Limnocentropodidae Rigid mineral tube case (mn). 2 (green) Predators (pre). 2 (green) Limnocentropus (Wiggins 2004). Wiggins 2004.
Odontoceridae Rigid mineral tube case (mn). 2 (green) Predators (pre). 2 (green) Two subfamilies: monotypical Pseudogoerinae and polytypical Odontocerinae, but basal lineages of Odontocerinae have not been inferred. (Wallace and Ross 1971). Wallace and Ross 1971 (Pseudogoera); Wiggins 1996; Holzenthal et al. 2007 (Odontocerinae).
Helicopsychidae Rigid mineral tube case (mn). 2 (green) Grazers (gra). 1 (brown) Rakiura (Johanson 1998; Johanson et al. 2017). Traits of Rakiura not described, apparently as for Helicopsyche (Johanson 1998).
Sericostomatidae Rigid mineral tube case (mn). 2 (green) Shredding herbivores, (shh). 0 (white) [Larvae of basal Cheimacheramus, Rhoizema, Petroplax, and Notidobiella unknown] Gumaga (Johanson et al. 2017). Resh et al. 1997.
Calamoceratidae Rigid plant tube case (pl). 1 (brown) Shredding detritivores (shd). 3 (yellow) Two subfamilies: Monotypical Anisocentropinae and polytypical Calamoceratinae, but basal lineages of Calamoceratinae have not been inferred (Prather 2002). Wallace and Sherberger 1970 (Anisocentropus).
Molannidae Rigid mineral tube case (mn). 2 (green) Collecting gatherers, predators (gat, pre). ? (stripe) Molanna and Mollanodes (the only genera). Graf et al. 2008; Cummins et al. 2019.
Leptoceridae Rigid mineral or plant or silk tube case (mn or pl or si). ? (stripe) Collecting gatherers, shredding herbivores, grazers, predators (gat, shh, gra, pre). ? (stripe) Grumichella and Leptorussa (Malm and Johanson 2011); Russobex (St Clair 1988). Case-retreat and trophic relations of Grumichella, Leptorussa, and Russobex unknown.
Hydropsychidae Fixed silk+substrate retreat with filter net (rt, fn). 3 (yellow) Collecting filterers (pff). 4 (red) Arctopsychinae: Arctopsyche, Parapsyche (Geraci et al. 2005). Cummins et al. 2019; Graf et al. 2008.
Stenopsychidae Fixed silk+substrate retreat with filter net (rt, fn). 3 (yellow) Collecting filterers (pff). 4 (red) Pseudostenopsyche and Stenopsychodes larvae unknown; Stenopsyche is the only other genus in the family. Tanida 2002.
Philopotamidae Fixed silk retreat used as filter net (rt, fn). 3 (yellow) Collecting filterers (pff). 4 (red) Larvae of basal subfamily Rossodinae (Rossodes) are unknown (Gibon 2013; Blahnik 2005); larvae of both remaining subfamilies ecologically similar. Graf et al. 2008; Cummins et al. 2019.
Dipseudopsidae Fixed silk+substrate retreat in buried tube (tm, fn). 3 (yellow) Collecting filterers (pff). 4 (red) Subfamilies Dipseudopsinae and Hyalopsychinae (the only subfamilies) (Ross and Gibbs 1973). Ulmer 1957; Marlier 1962; Gibbs 1968 (Dipseudopsinae: Dipseudopsis, Protodipseudopsis); Wallace et al. 1976 (Hyalopsychinae: Phylocentropus).
Pseudoneureclipsidae Short fixed silk retreat in sand tube on ventral surface of rock (tm, rt). 3 (yellow) Collecting filterers (pff) possibly using vortices (“eddying currents”--Gibbs 1973). 4 (red) Pseudoneureclipsis and Antillopsyche (the only genera). Gibbs 1973; Tachet et al. 2001 (Pseudoneureclipsis).
Xiphocentronidae Long, fixed meandering, tubular silk+substrate retreat on substrate surface often extending above water line (tm, rt). 3 (yellow) Collecting gatherers (gat). 1 (brown) Proxiphocentroninae: Proxiphocentron, larva unknown (Schmid 1982); Xiphocentroninae: Drepanocentron (Vilarino et al. 2022). Genco et al. 2020 (Drepanocentron).
Psychomyiidae Long, fixed, meandering, tubular silk+substrate retreat on substrate surface (tm, rt). 3 (yellow) Grazers (gra). 1 (brown) (Psychomyiinae: Metalype), (Tinodinae: Lype) (Li and Morse 1997). Edington and Hildrew 1995 (Metalype, Lype).
Ecnomidae Long, fixed, meandering, tubular silk+substrate retreat on substrate surface (tm, rt). 3 (yellow) Predators (pre). 2 (green) Daternomina, Parecnomina (Johanson and Espeland 2010). Cartwright 1997 (Daternomina); Gibbs 1973 (Parecnomina).
Polycentropodidae Fixed silk retreat with filtering function (rt, fn). 3 (yellow) Collecting filterers, predators (pff, pre). 2 (green) Neureclipsis (Johanson et al. 2012). Edington and Hildrew 1995; Inaba et al. 2014; Wiggins 1996 (Neureclipsis).
Open in a new tab

1fl = free-living; fn = filternet with retreat; mn = tubular mineral case; pc = purse case; pl = tubular plant case; ps = plastron-strap “tortoise” case; rt = silk retreat; si = tubular silk case; tm = tubular retreat with mineral particles. 2gat = gatherers-scrapers of sedimented fine particulate organic matter (FPOM = organic particles <10μ3); gra = grazers-scrapers of endolithic and epilithic algal tissues, biofilm, some FPOM, some tissues of living plants; pff = passive filter feeders of suspended FPOM and coarse particulate organic matter (CPOM = mainly fallen, entrained, and microbially conditioned leaves and other dead organic tissue >10μ3) and meiofauna and microfauna from moving water using nets or leg hairs and specialized mouthparts; pie = piercers of filamentous algae; pre = predators; shd = shredding detritivores of CPOM; shh = shredding herbivores of living plant tissue.

Table 3.

Ancestral traits for habits of larvae of basal taxa of Lepidoptera (Kawahara et al. 2019) and the Trichoptera families for which phylogenetic relationships were inferred by Ge et al. (2024a), with Ptilocolepidae from Frandsen et al. (2024), showing color codes for inferred ancestral nodes in Fig. 3.

Taxon Habit (Trait 5) Codes for Trait 5 Reference(s) for Inferred Basal Lineage(s) of Terminal Taxa Reference(s) for Traits of Basal Lineage(s)
Angiospermavora Terrestrial sprawlers 4 (yellow) Mitter et al. 2017, Kawahara et al. 2019. Stehr 1987 (Angiospermavora families).
Heterobathmiidae Burrower (miner in Nothfagus leaves) 1 (brown) Mitter et al. 2017, Kawahara et al. 2019. Kristensen and Nielsen 1983.
Agathiphagidae Burrower (in Kauri seeds) 1 (brown) Mitter et al. 2017, Kawahara et al. 2019. Dumbleton 1952.
Micropterigidae Semi-aquatic sprawlers 3 (black) Mitter et al. 2017, Kawahara et al. 2019. Davis and Landry 2012.
Ptilocolepidae Semi-aquatic sprawlers 3 (black) Palaeagapetus and Ptilocolepus (the only genera). Ito et al. 2014; Waringer and Graf 2002 .
Hydroptilidae Submerged sprawlers 2 (green) Stactobia (Marshall 1979). Graf et al. 2008; Waringer and Graf 2011.
Glossosomatidae Clingers 0 (white) Anagapetus and Glossosoma (Robertson and Holzenthal 2013). Wiggins 1996; Cummins et al. 2019.
Hydrobiosidae Clingers 0 (white) Apsilochorema (Ward et al. 2004). Holzenthal et al. 2007.
Rhyacophilidae Clingers 0 (white) Fansipangana (larva unknown) and Rhyacophila (McLaughlin et al. 2019). Graf et al. 2008.
Phryganopsychidae Submerged sprawlers 2 (green) Phryganopsyche (only genus). Wiggins 2004.
Phryganeidae Clingers, sprawlers 0 (white) Yphria (Wiggins 1998). Wiggins 1998.
Brachycentridae Clingers 0 (white) Eobrachycentrus (Flint 1984). Wiggins 1965, 1996.
Lepidostomatidae Climbers, sprawlers, clingers 0 (white) [Basal lineages of Lepidostomatinae and Theliopsychinae have not been inferred.] Wiggins 2004; Cummins et al. 2019.
Uenoidae Clingers 0 (white) Sericostriata (Wiggins et al. 1985) Wiggins et al. 1985.
Goeridae Clingers 0 (white) [Basal lineages of Goerinae, Larcasinae, and Lepaniinae have not been inferred. Traits are almost universal in the family.] Wiggins 1996, 2004.
Apataniidae Clingers 0 (white) Apataniana (Ge et al. 2024b). Waringer and Malicky 2016.
Limnephilidae Clingers 0 (white) Ecclisomyia, Philocasca (Vshivkova et al. 2007). Wiggins 2004; Cummins et al. 2019.
Limnocentropodidae Clingers 0 (white) Limnocentropus (Wiggins 2004) Wiggins 2004
Odontoceridae Burrowers 1 (brown) Two subfamilies: monotypical Pseudogoerinae and polytypical Odontocerinae, but basal lineages of Odontocerinae have not been inferred. (Wallace and Ross 1971). Wallace and Ross 1971 (Pseudogoera); Wiggins 1996; Holzenthal et al. 2007 (Odontocerinae).
Helicopsychidae Clingers 0 (white) Rakiura (Johanson 1998; Johanson et al. 2017). Traits of Rakiura not described, apparently as for Helicopsyche (Johanson 1998).
Sericostomatidae Sprawlers 2 (green) Larvae of basal Cheimacheramus, Rhoizema, Petroplax, and Notidobiella unknown; Gumaga (Johanson et al. 2017). Resh et al. 1997.
Calamoceratidae Sprawlers 2 (green) Two subfamilies: Monotypical Anisocentropinae and polytypical Calamoceratinae, but basal lineages of Calamoceratinae have not been inferred (Prather 2002). Wallace and Sherberger 1970 (Anisocentropus).
Molannidae Sprawlers 2 (green) Molanna and Mollanodes (the only genera). Graf et al. 2008; Cummins et al. 2019.
Leptoceridae Climbers, sprawlers, clingers, swimmers 0 (white) Grumichella and Leptorussa (Malm and Johanson 2011); Russobex (StClair 1988). Habits of Grumichella, Leptorussa, and Russobex unknown.
Hydropsychidae Clingers 0 (white) Arctopsychinae: Arctopsyche, Parapsyche (Geraci et al. 2005). Cummins et al. 2019; Graf et al. 2008.
Stenopsychidae Clingers 0 (white) Pseudostenopsyche and Stenopsychodes larvae unknown; Stenopsyche is the only other genus in the family. Tanida 2002.
Philopotamidae Clingers 0 (white) Larvae of basal subfamily Rossodinae (Rossodes) are unknown--Gibon 2013; Blahnik 2005; larvae of both remaining subfamilies ecologically similar. Graf et al. 2008; Cummins et al. 2019.
Dipseudopsidae Burrowers (miners in sand) 1 (brown) Subfamilies Dipseudopsinae and Hyalopsychinae (the only subfamilies) (Ross and Gibbs 1973). Ulmer 1957; Marlier 1962; Gibbs 1968 (Dipseudopsinae: Dipseudopsis, Protodipseudopsis); Wallace et al. 1976 (Hyalopsychinae: Phylocentropus).
Pseudoneureclipsidae Clingers 0 (white) Pseudoneureclipsis and Antillopsyche (the only genera). Gibbs 1973; Tachet et al. 2001 (Pseudoneureclipsis).
Xiphocentronidae Clingers 0 (white) Proxiphocentroninae: Proxiphocentron, larva unknown (Schmid 1982); Xiphocentroninae: Drepanocentron (Vilarino et al. 2022). Genco et al. 2020 (Drepanocentron).
Psychomyiidae Clingers 0 (white) Psychomyiinae: Metalype; Tinodinae: Lype (Li and Morse 1997). Edington and Hildrew 1995 (Metalype, Lype).
Ecnomidae Clingers 0 (white) Daternomina, Parecnomina (Johanson and Espeland 2010) Cartwright 1997 (Daternomina); Gibbs 1973 (Parecnomina)
Polycentropodidae Clingers 0 (white) Neureclipsis (Johanson et al. 2012) Edington and Hildrew 1995; Inaba et al. 2014; Wiggins 1996 (Neureclipsis)
Open in a new tab

The Case/Retreat category (Table 1) was segregated into five groups of traits for the families: free-living (code 0, without case or retreat), portable plant-tube case (1), portable mineral-tube case (2), fixed silk retreat (3), or portable tortoise case (4). These traits are color-coded white, brown, green, yellow, and black, respectively, in Fig. 1A. Because early instars of Ptilocolepidae and Hydroptilidae live freely and because of the biogenetic law, we included those families in the free-living group.

Figure 1.

Figure 1.

Open in a new tab

Evolution of case/retreat traits and food traits for Trichoptera and basal Lepidoptera families on phylogeny from Ge et al. (2024a), with modifications from Frandsen et al. (2024) and Kawahara et al. (2019). A. Evolution of case-retreat traits (Character 1), with code 0 (white) = free-living or without case or retreat, 1 (brown) = portable plant-tube case, 2 (green) = portable mineral-tube case, 3 (yellow) = fixed silk retreat, 4 (black) = portable tortoise case, and 5 (stripe) = unknown trait; B. Evolution of food-feeding-tropic relationships traits (Character 2), with code 0 (white) = shredding herbivores-vascular plants, 1 (brown) = grazers and gatherers, 2 (green) = predators, 3 (yellow) = shredding detritivores, 4 (red) = collecting filterers, 5 (black) = shredding herbivores-bryophytes, and 6 (stripe) = unknown trait.

The Food category (Table 1) was segregated into six groups of traits for the families: shredders-herbivores-vascular plants (code 0), grazers and gatherers (1), predators (2), shredders-detritivores (3), collecters-filterers (4), and shredders-herbivores-bryophytes (5). These traits are color-coded white, brown, green, yellow, red, black, respectively, in Fig. 1B. The food of the ancestral lineages of Limnephilidae is unknown (stripe).

The Habitat category (Table 2) was segregated into four groups of traits for the families: madicolous/hygropetric (code 0), lotic-erosional (1), lotic-depositional (2), and terrestrial (3). These traits are color-coded white, green, black, and brown, respectively, in Fig. 2A.

Table 2.

Ancestral traits for habitat and cocoon of larvae and pupae of basal taxa of Lepidoptera (Kawahara et al. 2019) and the Trichoptera families for which phylogenetic relationships were inferred by Ge et al. (2024a), with Ptilocolepidae from Frandsen et al. (2024), showing color codes for inferred ancestral nodes in Fig. 2.

Taxon Habitat (Trait 3) Codes for Trait 3 Cocoon1 (Trait 4) Codes for Trait 4 Reference(s) for Inferred Basal Lineage(s) of Terminal Taxa Reference(s) for Traits of Basal Lineage(s)
Angiospermavora Terrestrial. 3 (brown) New, closed, permeable, free cocoon. 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Stehr 1987 (Angiospermavora families).
Heterobathmiidae Terrestrial. 3 (brown) New, closed, permeable, free cocoon. 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Kristensen and Nielsen 1983.
Agathiphagidae Terrestrial. 3 (brown) No cocoon. 4 (stripe) Mitter et al. 2017, Kawahara et al. 2019. Dumbleton 1952.
Micropterigidae Wet soil-hygropetric. 0 (white) New, closed, permeable, free cocoon. 0 (white) Mitter et al. 2017, Kawahara et al. 2019. Davis and Landry 2012.
Ptilocolepidae Madicolous-hygropetric. 0 (white) Closed, permeable cocoon in larval purse case. 0 (white) Palaeagapetus and Ptilocolepus (the only genera). Ito et al. 2014; Waringer and Graf 2002.
Hydroptilidae Madicolous-hygropetric, lotic, lentic. 0 (white) Closed, semipermeable cocoon in larval purse case. 3 (black) Stactobia (Marshall 1979). Graf et al. 2008; Waringer and Graf 2011.
Glossosomatidae Lotic-erosional. 1 (green) Closed, semipermeable cocoon in larval dome case. 3 (black) Anagapetus and Glossosoma (Robertson and Holzenthal 2013). Wiggins 1996; Cummins et al. 2019.
Hydrobiosidae Lotic-erosional. 1 (green) Closed, semipermeable cocoon in new dome case. 3 (black) Apsilochorema (Ward et al. 2004). Holzenthal et al. 2007.
Rhyacophilidae Lotic-erosional. 1 (green) Closed, semipermeable cocoon in new dome case. 3 (black) Fansipangana (larva unknown) and Rhyacophila (McLaughlin et al. 2019). Graf et al. 2008.
Phryganopsychidae Lotic-depositional. 2 (black) Closed, permeable cocoon in new tube case. 0 (white) Phryganopsyche (only genus). Wiggins 2004.
Pisuliidae Lotic-depositiional (Silvatares), hygropetric (Pisulia). 2 (black) Open, in larval silk-lined tube case. 1 (brown) Pisulia and Silvatares (Stoltz 1989). Stoltz 1989.
Phryganeidae Lotic-depositional. 2 (black) Open, in new silk-lined tube case. 1 (brown) Yphria (Wiggins 1998). Wiggins 1998.
Brachycentridae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Eobrachycentrus (Flint 1984). Wiggins 1965,1996.
Lepidostomatidae Lotic-erosional or depositional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) [Basal lineages of Lepidostomatinae and Theliopsychinae have not been inferred.] Wiggins 2004; Cummins et al. 2019.
Uenoidae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Sericostriata (Wiggins et al. 1985). Wiggins et al. 1985.
Goeridae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) [Basal lineages of Goerinae, Larcasinae, and Lepaniinae have not been inferred. Traits in bold are almost universal in the family.] Wiggins 1996, 2004
Apataniidae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Apataniana (Ge et al. 2024b). Waringer and Malicky 2016.
Limnephilidae Lotic-erosional (small mountain streams), lentic. 1 (green) Open, in new silk-lined tube case. 1 (brown) Ecclisomyia, Philocasca (Vshivkova et al. 2007). Wiggins 2004; Cummins et al. 2019.
Limnocentropodidae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Limnocentropus (Wiggins 2004). Wiggins 2004.
Odontoceridae Lotic-depositional. 2 (black) Open, in larval silk-lined tube case. 1 (brown) Two subfamilies: monotypical Pseudogoerinae and polytypical Odontocerinae, but basal lineages of Odontocerinae have not been inferred. (Wallace and Ross 1971). Wallace and Ross 1971 (Pseudogoera); Wiggins 1996; Holzenthal et al. 2007 (Odontocerinae).
Helicopsychidae Lotic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Rakiura (Johanson 1998; Johanson et al. 2017). Traits of Rakiura not described, apparently as for Helicopsyche (Johanson 1998).
Sericostomatidae Lotic-depositional. 2 (black) Open, in larval silk-lined tube case. 1 (brown) Larvae of basal Cheimacheramus, Rhoizema, Petroplax, and Notidobiella unknown; Gumaga (Johanson et al. 2017). Resh et al. 1997.
Calamoceratidae Lotic-depositional. 2 (black) Open, in larval silk-lined tube case. 1 (brown) Two subfamilies: monotypical Anisocentropinae and polytypical Calamoceratinae, but basal lineages of Calamoceratinae have not been inferred (Prather 2002). Wallace and Sherberger 1970 (Anisocentropus).
Molannidae Lentic-erosional. 1 (green) Open, in larval silk-lined tube case. 1 (brown) Molanna and Mollanodes (the only genera). Graf et al. 2008; Cummins et al. 2019.
Leptoceridae Lotic-erosional (fast, cool, mountain streams). 1 (green) Open, in larval silk-lined tube case. 1 (brown) Grumichella and Leptorussa (Malm and Johanson 2011); Russobex (StClair 1988). Habitat of Leptorussa unknown; habitat of Grumichella and Russobex mentioned by Calor et al. (2016) and St Clair (1988).
Hydropsychidae Lotic-erosional. 1 (green) Open cocoon in new, long-dome shelter. 2 (green) Arctopsychinae: Arctopsyche, Parapsyche (Geraci et al. 2005). Cummins et al. 2019; Graf et al. 2008.
Stenopsychidae Lotic-erosional. 1 (green) Open cocoon in new, long-dome shelter. 2 (green) Pseudostenopsyche and Stenopsychodes larvae unknown; Stenopsyche is the only other genus in the family. Tanida 2002.
Philopotamidae Lotic-erosional (Cummins et al. 2019), lotic-depositional (Graf et al. 2008). 1 (green) Open or closed cocoon in new, long-dome shelter. 2 (green) Larvae of basal subfamily Rossodinae (Rossodes) are unknown--Gibon 2013; Blahnik 2005; larvae of both remaining subfamilies ecologically similar. Wiggins 2004 (pupal cocoon); Graf et al. 2008 and Cummins et al. 2019 (larval habitat).
Dipseudopsidae Lotic-depositional. 2 (black) Open cocoon in buried larval sand tube. 2 (green) Subfamilies Dipseudopsinae and Hyalopsychinae (the only subfamilies) (Ross and Gibbs 1973). Ulmer 1957; Marlier 1962; Gibbs 1968 (Dipseudopsinae: Dipseudopsis, Protodipseudopsis); Wallace et al. 1976 (Hyalopsychinae: Phylocentropus).
Pseudoneureclipsidae Lotic-depositional. 2 (black) Open cocoon in new or larval sand tube on substrate surface. 2 (green) Pseudoneureclipsis and Antillopsyche (the only genera). Gibbs 1973; Tachet et al. 2001 (Pseudoneureclipsis).
Xiphocentronidae Lotic-depositional, including above water surface (hygropetric?). 2 (black) Open cocoon in larval sand tube, possibly hygropetric--Sturm 1960. 2 (green) Proxiphocentroninae: Proxiphocentron, larva unknown (Schmid 1982); Xiphocentroninae: Drepanocentron (Vilarino et al. 2022). Genco et al. 2020 (Drepanocentron).
Psychomyiidae Lotic. 1 (green) Open cocoon in new, long-dome shelter. 2 (green) (Psychomyiinae: Metalype), (Tinodinae: Lype) (Li and Morse 1997) Edington and Hildrew 1995 (Metalype, Lype).
Ecnomidae Lotic-depositional, lentic. 2 (black) Open or closed cocoon in new, long-dome shelter. 2 (green) Daternomina, Parecnomina (Johanson and Espeland 2010). Cartwright 1997 (Daternomina); Gibbs 1973 (Parecnomina).
Polycentropodidae Lotic-depositional. 2 (black) Open cocoon in new, long-dome shelter. 2 (green) Neureclipsis (Johanson et al. 2012). Edington and Hildrew 1995; Inaba et al. 2014; Wiggins 1996 (Neureclipsis).
Open in a new tab

1 Data mainly from Wiggins 2004.

Figure 2.

Figure 2.

Open in a new tab

Evolution of habitat traits and cocoon traits for Trichoptera and basal Lepidoptera families on phylogeny from Ge et al. (2024a), with modifications from Frandsen et al. (2024) and Kawahara et al. (2019). A. Evolution of habitat traits (Character 3), with code 0 (white) = madicolous/ hygropetric, 1 (green) = lotic-erosional, 2 (black) = lotic-depositional, 3 (brown) = terrestrial; B. Evolution of cocoon traits (Character 4), with code 0 (white) = closed permeable cocoon, 1 (brown) = open in silk-lined tube case, 2 (green) = open in long-dome shelter or sand tube, 3 (black) = closed semipermeable cocoon, 4 (stripe) = unknown trait.

The Cocoon category (Table 2) was segregated into four groups of known traits for the families: closed permeable cocoon (code 0); open in silk-lined tube (1); open in long-dome shelter or sand-tube (2); closed semipermeable cocoon (3), sensu Wiggins 2004). These traits are color-coded white, green, black, and red, respectively, in Fig. 2B. The cocoon trait for Agathiphagidae is unknown (stripe).

The Habit category (Table 3) was segregated into five groups of traits for the families: clingers (code 0), burrowers (1), submerged sprawlers (2), semi-aquatic sprawlers (3), and terrestrial sprawlers (4). These traits are color-coded white, brown, green, black, and yellow, respectively, in Fig. 3.

Figure 3.

Figure 3.

Open in a new tab

Evolution of habit traits (Character 5) for Trichoptera and basal Lepidoptera families on phylogeny from Ge et al. (2024a), with modifications from Frandsen et al. (2024) and Kawahara et al. (2019). Code 0 (white) = clingers, 1 (brown) = burrowers, 2 (green) = submerged sprawlers, 3 (black) = clingers, 4 (yellow) = terrestrial sprawlers or unknown trait.

An Ancestral Character State Reconstruction (ACSR) was conducted for each category of traits. Mesquite v.3.7.0 (http://mesquiteproject.org) was used to perform the ACSR Parsimony Analysis on the encoded family nodes of the phylogeny.

Results

The ACSR results are shown at the hypothetical ancestral nodes of Figs 1–3 with the same color notations as for the taxa in Tables 1–3. Regarding larval case or retreat, our analysis supports the conclusion of Frandsen et al. (2024) and Ge et al. (2024a) that the free-living condition, without any type of shelter (e.g., similar to the life-style of early instars of Hydroptilidae and Ptilocolepidae), was probably the ancestral state for Trichoptera (Fig. 1A). Our analysis also supports our hypothesis that caddisfly ancestral larvae were shredding herbivores of bryophytes (Fig. 1B). The ancestral habitat is inconclusive, likely either madicolous/hygropetric or lotic-erosional (Fig. 2A). The type of pupal cocoon was equivocal, as well, either a closed and permeable cocoon or a closed and semipermeable cocoon, or an open silk-lined chamber in a long-dome shelter or sand tube (Fig. 2B). The ancestral larval habit was likely that of a clinger or a semi-aquatic sprawler (Fig. 3).

Discussion

Trichoptera ancestor and its descendants

Based on their phylogenomic analyses, both Frandsen et al. (2024) and Ge et al. (2024a) agreed that the ancestral larva of Integripalpia in all instars and probably of Trichoptera in at least early instars was “free-living,” i.e., without a portable case or fixed retreat. Now, by comparing the functional traits of the larva of the hypothesized basal lepidopteran lineage (Micropterigidae) and those of modern caddisfly lineages as summarized above, we can infer some additional functional traits of larvae of their Amphiesmenopteran ancestor and the Trichoptera ancestor and of the evolutionary route taken by the Trichoptera ancestor to invade freshwater habitats.

Our analysis concludes that larvae of the free-living caddisfly ancestor probably fed on bryophytes, as those of Micropterigidae and Ptilocolepidae do today. Bryophytes (liverworts, hornworts, mosses) require moist environments for reproduction and are often found living in or immediately beside flowing water and subject to immersion, attached by rhizomes to rocks and other stable substrate on stream edges (Glime 2023). Clinging to or sprawling on semiaquatic bryophytes was probably the common lifestyle of the caddisfly larval ancestor and may have been that of the amphiesmenopteran ancestor as well. The analysis indicated that lotic-erosional or terrestrial habitats are also possibilities. Whatever its habitat may have been, this ancestral caddisfly larva lost its spiracles and either lost or never had a plastron, respiring instead directly through its integument from the cold, oxygen-rich water, apparently making the terrestrial habitat option unlikely. This evolutionary development may also have been possible because of their small size and consequent high surface-to-volume ratio, allowing effective respiration in a freshwater medium. Thus, this ancestral caddisfly larva evolved from a semiterrestrial environment (as in Micropterigidae and Ptilocolepidae) into a permanently submerged lifestyle (as in most Hydroptiloidea and other Trichoptera), radiating to access the many food and habitat resources available under water in the Permian Period.

Precocious production of silk by the active larva, well before time for pupation, apparently evolved for various purposes in the ancestral Trichoptera larva, whether for retreat building (as in modern Annulipalpia) or larval case construction (as in most modern Integripalpia). The primitive caddisfly larva also constructed, apparently for the first time in Amphiesmenoptera, a dome-like pupation shelter of silk and substrate materials, an evolutionary development or synapomorphy of Trichoptera that is observed in basal lineages of both Annulipalpia and Integripalpia. The larva then spun a closed permeable or semipermeable cocoon inside this shelter, fusing the cocoon to its inner walls (Hydroptilidae) or leaving it mostly free (other basal lineages of modern Integripalpia), or the larva spun an open or closed cocoon as in modern Annulipalpia. In Integripalpia, larvae of the tube-case-making Phryganides pupated in their tube cases after lining them with silk and constructing tough silken mesh at either end, then making undulating ventilation movements to propel the unidirectional flow of water through the tube (Wiggins 2004).

Integripalpia ancestor and its descendants

Considering the recently published phylogenetic hypotheses (Thomas et al. 2020; Frandsen et al. 2024; Ge et al. 2024a), semipermeable cocoons may have evolved independently in Hydroptilidae and the Glossosomatidae + Rhyacophiloidea lineage; alternatively, semipermeable cocoons may have first appeared in the Integripalian ancestor and been abandoned in the Ptilocolepidae lineage and Phryganides. A further development in the Integripalpia lineage was the precocious construction of the pupal shelter in early instars (immediately on molting into the hypogastric last instar of Hydroptiloidea and in all instars of Glossosomatidae). Wiggins (2004) suggested that the hydroptiloid purse case and glossosomatid tortoise case are homologous because they are both based on two sheets of case materials fastened together at lateral seams; if these case-construction behaviors are homologous, the precocious construction of a pupal shelter apparently was abandoned in Rhyacophiloidea to facilitate rapid movement for capturing prey, delaying shelter construction until immediately before pupation; if they are not homologous, as Frandsen et al. (2024) concluded, the larval ancestor of Glossosomatidae+Rhyacophiloidea apparently was free-living, with the Glossosomatidae ancestor evolving tortoise case construction behavior independently.

The portable tubular cases of Phryganides larvae do not appear to be homologous with the portable cases of Hydroptiloidea and/or Glossosomatidae (Wiggins 2004). Nevertheless, this tubular type of case may also, like those of hydroptiloids and glossosomatids, have evolved as a precocious pupation shelter, initially for larval protection. For pupation, the tubular case was retained and modified by the production of silken sieve plates at the vulnerable ends, or the larval case was abandoned and a new protective tube shelter was constructed as in Phryganopsyche (Phryganopsychidae), Yphria (Phryganeidae), and Ecclisomyia (Limnephilidae) (Wiggins 2004; Givens 2018). Besides its value as camouflage and physical protection, a selective advantage of the tube was for improved respiration by larval and pupal undulations inside the tube, continuously directing a fresh supply of oxygenated water over the body. This adaptation allowed various lineages of Phryganides to invade quiet-water habitats with lower concentrations of dissolved oxygen (Mackay and Wiggins 1979). Most case-making larvae of basal Integripalpia included minerals of various sizes or filamentous algae into their larval/pupal cases. The incorporation of angiosperm leaf pieces in some later lineages, especially in earlier lineages of infraorder Plenitentoria, became possible after the evolution of that major plant clade (Morse et al. 2019).

Annulipalpia ancestor and its descendants

Like the larvae of Rhyacophiloidea and early instar larvae of Hydroptiloidea, the larvae of most Annulipalpia have a campodeiform shape. They also evolved strong anal prolegs and tarsal claws for maneuvering efficiently on the substrate. Hydropsychidae are presently thought to be the basal lineage of Annulipalpia (Frandsen et al. 2024; Ge et al. 2024a) and species of their subfamily Arctopsychinae are considered the descendants of the basal lineage of Hydropsychidae (Geraci et al. 2005). Modern species of Arctopsychinae characteristically live in some of the fastest flowing mountain-stream habitats. The functional changes that allowed a free-living, wet-habitat ancestral caddisfly larva to invade lotic-erosional ecotones characteristically occupied by modern arctopsychines are unclear and must have been dramatic, a scenario inviting future investigation.

Conclusions

An appreciation of the importance of functional traits for understanding freshwater communities and their necessary biotic and abiotic characteristics has increased in recent years (Graf et al. 2008; Merritt et al. 2019). Inference of ancestral functional traits helps put modern traits in their historical context and provides a foundation for predicting and confirming yet-unknown traits still being investigated for the modern fauna (Morse and White 1979).

Acknowledgements

We are grateful for helpful advice from Dr Paul Frandsen, Brigham Young University, and two anonymous reviewers.

Citation

Ge X, Morse JC (2025) Functional traits of ancestral caddisfly (Trichoptera) larvae and pupae. In: Ríos-Touma B, Frandsen PB, Holzenthal RW, Houghton DC, Rázuri-Gonzales E, Pauls SU (Eds) Proceedings of the 18th International Symposium on Trichoptera. ZooKeys 1263: 47–68. https://doi.org/10.3897/zookeys.1263.148069

Funding Statement

The authors have no funding to report.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

We ensure that all research conducted will respect participant confidentiality, adhere to ethical guidelines, and contribute to the academic community with honesty and integrity.

Use of AI

No tools based on artificial intelligence technologies were used in the conduct of this research or in the preparation or editing of the resulting manuscript.

Funding

This research was funded by the National Natural Science Foundation of China (32400357) and the USA National Science Foundation (DEB-2325925).

Author contributions

Conceptualization G, M; Data curation G, M; Formal analysis G, M; Methodology G, M; Resources G, M; Validation G, M; Visualization G; Writing—original draft M; Writing—review & editing G, M.

Author ORCIDs

Xinyu Ge https://orcid.org/0000-0001-7375-504X

John C. Morse https://orcid.org/0000-0003-3187-4045

Data availability

All of the data that support the findings of this study are available in the main text.

References

  1. Blahnik RJ. (2005) Alterosa, a new caddisfly genus from Brazil (Trichoptera: Philopotamidae). Zootaxa 991(1): 1–60. 10.11646/zootaxa.991.1.1 [DOI] [Google Scholar]
  2. Calor AR, Holzenthal RW, Froehlich CG. (2016) Phylogeny and revision of the Neotropical genus Grumichella Müller (Trichoptera: Leptoceridae), including nine new species and a key. Zoological Journal of the Linnean Society 176(1): 137–169. 10.1111/zoj.12310 [DOI] [Google Scholar]
  3. Cartwright DI. (1997) Preliminary guide to the identification of late instar larvae of Australian Ecnomidae, Philopotamidae and Tasmiidae (Insecta: Trichoptera). Co-operative Research Center for Freshwater Ecology, Albury, Australia. Identification Guide 10: 1–33. [Google Scholar]
  4. Cummins KW, Wallace JR, Merritt RW, Wiggins GB, Morse JC, Holzenthal RW, Robertson DR, Rasmussen AK, Currie DC, Berg MB. (2019) Table 19A: Summary of ecological and distributional data for Trichoptera (caddisflies). In: Merritt RW, Cummins KW, Berg MB. (Eds) An Introduction to the Aquatic Insects of North America (5th edn.). KendallHunt Publishing Company, Dubuque, Iowa, 726–764.
  5. Davis DR, Landry J-F. (2012) A review of the North American genus Epimartyria (Lepidoptera, Micropterigidae) with a discussion of the larval plastron. ZooKeys 183: 37–83. 10.3897/zookeys.183.2556 [DOI] [PMC free article] [PubMed] [Google Scholar]
  6. Dumbleton LJ. (1952) A new genus of seed-infesting micropterygid moths. Pacific Science 6: 17–29. [Google Scholar]
  7. Edington JM, Hildrew AG. (1995) A Revised Key to the Caseless Caddis Larvae of the British Isles, with Notes on Their Ecology. Scientific Publication – Freshwater Biological Association 53: 1–134. [Google Scholar]
  8. Flint Jr OS. (1984) The genus Brachycentrus in North America, with a proposed phylogeny of the genera of Brachycentridae (Trichoptera). Smithsonian Contributions to Zoology 398(398): 1–58. 10.5479/si.00810282.398 [DOI] [Google Scholar]
  9. Frandsen PB, Holzenthal R, Espeland M, Breinholt JW, Thomas Thorpe JA, Simon S, Kawahara AY, Plotkin D, Hotaling S, Li Y-y, Nelson CR, Niehuis O, Mayer C, Podsiadlowski L, Donath A, Misof B, Mariarty E, Lemmon A, Morse JC, Liu S, Pauls SU, Zhou X. (2024) Phylogenomics recovers multiple origins of portable case-making in caddisflies (Insecta: Trichoptera), nature’s underwater architects. Proceedings. Biological Sciences 291(2026): e20240514. 10.1098/rspb.2024.0514 [DOI] [PMC free article] [PubMed]
  10. Ge X-y, Peng L, Morse JC, Wang J-y, Zang H, Yang L, Sun C, Wang B. (2024a) Phylogenomics resolves a 100-year-old debate regarding the evolutionary history of caddisflies (Insecta: Trichoptera). Molecular Phylogenetics and Evolution 201(108196): 1006–1020. 10.1016/j.ympev.2024.108196 [DOI] [PubMed] [Google Scholar]
  11. Ge X-y, Wang J-y, Zang H, Chai L, Liu W, Zhang J, Yan C, Wang B. (2024b) Mitogenomics provide new phylogenetic insights of the family Apataniidae (Trichoptera: Integripalpia). Insects 15(12): e973. 10.3390/insects15120973 [DOI] [PMC free article] [PubMed]
  12. Genco MS, Morse JC, Caterino MS, Murray-Stoker KM, Pham TH. (2020) Larvae and adults of Vietnamese species of Drepanocentron and Hydromanicus (Trichoptera: Xiphocentronidae, Hydropsychidae). In: Laudee P, Morse JC. (Eds) Proceedings of the 16th International Symposium on Trichoptera.Zoosymposia 18: 72–92. 10.11646/zoosymposia.18.1.11 [DOI]
  13. Geraci CJ, Kjer KM, Morse JC, Blahnik RJ. (2005) Phylogenetic relationships of Hydropsychidae subfamilies based on morphology and DNA sequence data. In: Tanida K, Rossiter A. (Eds) Proceedings of the 11th International Symposium on Trichoptera, Osaka & Shiga.Tokai University Press, Hadano-shi, Kanagawa 257-0003, Japan, 131–136.
  14. Gibbs DG. (1968) The larva, dwelling-tube and feeding of a species of Protodipseudopsis (Trichoptera: Dipseudopsidae). Proceedings of the Royal Entomological Society of London. Series A, General Entomology 43(4–6): 73–79. 10.1111/j.1365-3032.1968.tb01030.x [DOI] [Google Scholar]
  15. Gibbs DG. (1973) The Trichoptera of Ghana. Deutsche Entomologische Zeitschrift 20 (IV–V): 363–424. 10.1002/mmnd.4810200410 [DOI]
  16. Gibon F-M. (2013) Une sous-famille caractéristique des forêts humides primaires malgaches: Les Rossodinae (Trichoptera, Philopotamidae). Zoosystema 35(2): 151–174. 10.5252/z2013n2a2 [DOI] [Google Scholar]
  17. Givens DR. (2018) The Nearctic Ecclisomyia species (Trichoptera: Limnephilidae). Zootaxa 4413(2): 201–259. 10.11646/zootaxa.4413.2.1 [DOI] [PubMed] [Google Scholar]
  18. Glime JM. (2023) Bryophyte Ecology (Vol. 4). Habitats and Roles. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 22 March 2023. http://digitalcommons.mtu.edu/bryophyte-ecology/ [accessed 02 June 2025]
  19. González MA, Vieira-Lanero R, Cobo F. (2000) The immature stages of Ptilocolepus extensus McLachlan, 1884 (Trichoptera: Hydroptilidae: Ptilocolepidae) with notes on biology. Aquatic Insects 22(1): 27–38. 10.1076/0165-0424(200001)22:1;1-Z;FT027 [DOI] [Google Scholar]
  20. Graf W, Murphy J, Dahl J, Zamora-Muñoz C, López Rodríguez MJ. (2008) Trichoptera. In: Schmidt-Kloiber A, Hering D. (Eds), Distribution and ecological preferences of European freshwater organisms (Vol. 1). Pensoft, Sofia, 388 pp. [Google Scholar]
  21. Haupt H. (1956) Beitrag zur Kenntnis der eozänen Arthopodenfauna des Geiseltales. Nova Acta Leopoldina 18(128): 1–89. [Google Scholar]
  22. Holzenthal RW, Blahnik RJ, Kjer KM, Prather AL. (2007) An update on the phylogeny of caddisflies (Trichoptera). In: Bueno-Soria J, Barba-Álvarez R, Armitage B. (Eds) Proceedings of the XIIth International Symposium on Trichoptera, June 18–22, 2006.The Caddis Press, Columbus, Ohio, 143–153.
  23. Inaba S, Nozaki T, Kobayashi S, Tanida K. (2014) Discovery of immature stages of Neureclipsis mandjurica (Martynov, 1907) (Trichoptera, Polycentropodidae) from Japan. Biogeography 16: 63–70. [Google Scholar]
  24. Ings NL, Hildrew AG, Grey J. (2010) Gardening by the psychomyiid caddisfly Tinodes waeneri: Evidence from stable isotopes. Oecologia 163(1): 127–139. 10.1007/s00442-009-1558-8 [DOI] [PubMed] [Google Scholar]
  25. Ings NL, Grey J, King L, McGowan S, Hildrew AG. (2017) Modification of littoral algal assemblages by gardening caddisfly larvae. Freshwater Biology 62(3): 507–518. 10.1111/fwb.12881 [DOI] [Google Scholar]
  26. Ito T, Higler LWG. (1993) Biological notes and description of little-known stages of Ptilocolepus granulatus (Pictet) (Trichoptera: Hydroptilidae). In: Otto C. (Ed.) Proceedings of the 7th International Symposium on Trichoptera, Umeå, Sweden, 3–8 August 1992.Backhuys Publishers, Leiden, The Netherlands, 177–181.
  27. Ito T, Wisseman RW, Morse JC, Colbo MH, Weaver JS III. (2014) The genus Palaeagapetus Ulmer (Trichoptera, Hydroptilidae, Ptilocolepinae) in North America. Zootaxa 3794(2): 201–221. 10.11646/zootaxa.3794.2.1 [DOI] [PubMed] [Google Scholar]
  28. Ivanov VD. (1992) A new family of caddisflies (Insecta, Trichoptera) from the Permian of the middle Urals. Палеонтологический журнал 4: 31–35. [Google Scholar]
  29. Johanson KA. (1998) Phylogenetic and biogeographic analysis of the family Helicopsychidae (Insecta: Trichoptera). Entomologica Scandinavica (Supplement 53): 1–172.
  30. Johanson KA, Espeland M. (2010) Phylogeny of the Ecnomidae (Insecta: Trichoptera). Cladistics 26(1): 36–48. 10.1111/j.1096-0031.2009.00276.x [DOI] [PubMed] [Google Scholar]
  31. Johanson KA, Malm T, Espeland M, Weingartner E. (2012) Phylogeny of the Polycentropodidae (Insecta: Trichoptera) based on protein-coding genes reveal non-monophyletic genera. Molecular Phylogenetics and Evolution 65(1): 126–135. 10.1016/j.ympev.2012.05.029 [DOI] [PubMed] [Google Scholar]
  32. Johanson KA, Malm T, Espeland M. (2017) Molecular phylogeny of Sericostomatoidea (Trichoptera) with the establishment of three new families. Systematic Entomology 42(1): 240–266. 10.1111/syen.12209 [DOI] [Google Scholar]
  33. Kawahara AY, Plotkin D, Espeland M, Meusemann K, Toussaint EFA, Donath A, Gimnich F, Frandsen PB, Zwick A, dos Reis M, Barber JR, Peters RS, Liu S, Zhou X, Mayer C, Podsiadlowski L, Storer C, Yack JE, Misof B, Breinholt JW. (2019) Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the United States of America 116(45): 22657–22663. 10.1073/pnas.1907847116 [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Kiriakoff SG. (1948) A classification of the Lepidoptera and related groups with some remarks on taxonomy. Biologisch Jaarboek 15: 118–143. [Google Scholar]
  35. Kopylov DS, Rasnitsyn AP, Aristov DS, Bashkuev AS, Bazhenova NV, Dmitriev VY, Gorochov AV, Ignatov MS, Ivanov VD, Khramov AV, Legalov AA, Lukashevich ED, Mamontov YS, Melnitsky SI, Ogłaza B, Ponomarenko AG, Prokin AA, Ryzhkova OV, Shmakov AS, Sinitshenkova ND, Solodovnikov AY, Strelnikova OD, Sukacheva ID, Uliakhin AV, Vasilenko DV, Wegierek P, Yan EV, Zmarzły M. (2020) The Khasurty fossil insect Lagerstätte. Paleontological Journal 54(11): 1221–1394. 10.1134/S0031030120110027 [DOI] [Google Scholar]
  36. Kristensen NP. (1997) Early evolution of the Lepidoptera + Trichoptera lineage: Phylogeny and the ecological scenario. In: Grandcolas P. (Ed.) The Origin of Biodiversity in Insects: Phylogenetic Tests of Evolutionary Scenarios.Mémoires du Museum National d’Histoire Naturelle 173: 253–271.
  37. Kristensen NP, Nielsen ES. (1983) The Heterobathmia life history elucidated: Immature stages contradict assignment to suborder Zeugloptera (Insecta, Lepidoptera). Journal of Zoological Systematics and Evolutionary Research 21(2): 101–124. 10.1111/j.1439-0469.1983.tb00280.x [DOI] [Google Scholar]
  38. Latreille PA. (1805) Histoire Naturelle, Générale et Particulière, des Crustacés et des Insects (Vol. 13). F. Dufart, Paris, 432 pp. [Google Scholar]
  39. Li Y-j, Morse JC. (1997) Phylogeny and classification of Psychomyiidae (Trichoptera) genera. In: Holzenthal RW, Flint Jr OS. (Eds) Proceedings of the 8th International Symposium on Trichoptera.Ohio Biological Survey, Columbus, Ohio, 271–276.
  40. Mackay RJ, Wiggins GB. (1979) Ecological diversity in Trichoptera. Annual Review of Entomology 24(1): 185–208. 10.1146/annurev.en.24.010179.001153 [DOI] [Google Scholar]
  41. Malm T, Johanson KA. (2011) A new classification of the long-horned caddisflies (Trichoptera: Leptoceridae) based on molecular data. Evolutionary Biology 11: 1–10. 10.1186/1471-2148-11-10 [DOI] [PMC free article] [PubMed] [Google Scholar]
  42. Marlier G. (1962) Genera des trichopteres de l’Afrique. Annales de Musée Royal de l’Afrique Centrale. Tervuren 109: 1–263. [Google Scholar]
  43. Marshall JE. (1979) A review of the genera of the Hydroptilidae. Bulletin of the British Museum (Natural History). Entomology Series 39(3): 135–239. [Google Scholar]
  44. Martynova OM. (1958) Novye nasekomye iz permskikh i mezozoiskikh otlozhenii SSSR. Materialy k Osnovam Paleontologii 2: 69–94. [In Russian] [Google Scholar]
  45. McLaughlin JE, Frandsen PB, Mey W, Pauls SU. (2019) A preliminary phylogeny of Rhyacophilidae with reference to Fansipangana and the monophyly of Rhyacophila. In: Mendez PK, Spagna JC, Holzenthal RW, Houghton DC. (Eds) Proceedings of the 15th International Symposium on Trichoptera, New Brunswick, New Jersey, 4–7 June 2015.Zoosymposia 14: 189–192. 10.11646/zoosymposia.14.1.20 [DOI]
  46. Merritt RW, Cummins KW, Berg MB [Eds] (2019) An Introduction to the Aquatic Insects of North America (5th Edn. ) KendallHunt Publishing Company, Dubuque, Iowa, 1480 pp. [Google Scholar]
  47. Mey W, Wichard W. (2023) Tarachoptera: The extinct and enigmatic cousins of Trichoptera and Lepidoptera, with descriptions of two new species. Contributions to Entomology 73(2): 137–146. 10.3897/contrib.entomol.73.e110233 [DOI] [Google Scholar]
  48. Mey W, Wichard W, Müller P, Wang B. (2017a) The blueprint of the Amphiesmenoptera—Tarachoptera, a new order of insects from Burmese amber (Insecta, Amphiesmenoptera). Fossil Record 20: 129e145. 10.5194/fr-20-129-2017 [DOI]
  49. Mey W, Wichard W, Ross E, Ross A. (2017b) On the systematic position of a highly derived amphiesmenopteran insect from Burmese amber (Insecta, Amphiesmenoptera). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 107(2–3): 249–254. 10.1017/S1755691017000330 [DOI] [Google Scholar]
  50. Milne MJ, Milne LJ. (1939) Evolutionary trends in caddis worm case construction. Annals of the Entomological Society of America 32(3): 533–542. 10.1093/aesa/32.3.533 [DOI] [Google Scholar]
  51. Misof B, Liu S, Meusemann K, Peters RS, Donath A, Mayer C, Frandsen PB, Ware J, Flouri T, Beutel RG, Niehuis O, Petersen M, Izquierdo-Carrasco F, Wappler T, Rust J, Aberer AJ, Aspöck U, Aspöck H, Bartel D, Blanke A, Berger S, Böhm A, Buckley TR, Calcott B, Chen J, Friedrich F, Fukui M, Fujita M, Greve C, Grobe P, Gu S, Huang Y, Jermiin LS, Kawahara AY, Krogmann L, Kubiak M, Lanfear R, Letsch H, Li Y-y, Li Z-y, Li J, Lu H, Machida R, Mashimo Y, Kapli P, McKenna DD, Meng G, Nakagaki Y, Navarrete-Heredia JL, Ott M, Ou Y, Pass G, Podsiadlowski L, Pohl H, von Reumont BM, Schütte K, Sekiya K, Shimizu S, Slipinski A, Stamatakis A, Song W, Su X, Szucsich NU, Tan M, Tan X, Tang M, Tang J, Timelthaler G, Tomizuka S, Trautwein M, Tong X, Uchifune T, Walzl MG, Wiegmann BM, Wilbrandt J, Wipfler B, Wong TKF, Wu Q, Wu G, Xie Y, Yang S, Yang Q, Yeates DK, Yoshizawa K, Zhang Q, Zhang R, Zhang W, Zhang Y, Zhao J, Zhou C, Zhou L, Ziesmann T, Zou S, Li Y, Xu X, Zhang Y, Yang H, Wang J, Wang J, Kjer KM, Zhou X. (2014) Phylogenomics resolves the timing and pattern of insect evolution. Science 346(6210): 763–767. 10.1126/science.1257570 [DOI] [PubMed] [Google Scholar]
  52. Mitter C, Davis DR, Cummings MP. (2017) Phylogeny and evolution of Lepidoptera. Annual Review of Entomology 62(1): 265–283. 10.1146/annurev-ento-031616-035125 [DOI] [PubMed] [Google Scholar]
  53. Morse JC. (2025) Trichoptera World Checklist. https://trichopt.app.clemson.edu/ [accessed 03 June 2025]
  54. Morse JC, White Jr DF. (1979) A technique for analysis of historical biogeography and other characters in comparative biology. Systematic Zoology 28(3): 356–365. 10.2307/2412588 [DOI] [Google Scholar]
  55. Morse JC, Frandsen PB, Graf W, Thomas JA. (2019) Diversity and ecosystem services of Trichoptera. In: Morse JC, Adler PH (Eds) Diversity and ecosystem services of aquatic insects. Insects 10: e125. 10.3390/insects10050125 [DOI] [PMC free article] [PubMed]
  56. Nielsen A. (1948) Postembryonic development and biology of the Hydroptilidae: A contribution to the phylogeny of the caddis flies and to the question of the origin of the case-building instinct. Biologiske Skrifter 5(1): 1–200. [pls I–III.] [Google Scholar]
  57. Novokshonov VG, Sukatsheva I. (1997) On the knowledge of the Paleozoic Promeropidae—presumably the first caddisflies. In: Novokshonov VG. (Ed.) Problems of Origin, Systematics and Ecology of Caddisflies of Russia and adjacent Territories.Proceedings of the 5th All-Russian Trichopterological Symposium. Kyadrat, Voronezh, 21–24.
  58. Prather AL. (2002) Phylogenetic analysis of Leptoceroidea and Calamoceratidae, and revisions of the Neotropical genera Banyallarga and Phylloicus (Insecta: Trichoptera). PhD thesis, University of Minnesota, Saint Paul, xix + 485 pp.
  59. Resh VH, Wood JR, Bergey EA, Feminella JW, Jackson JK, McElravy EP. (1997) Biology of Gumaga nigricula (McL.) in a northern California stream. In: Holzenthal RW, Flint Jr OS. (Eds) Proceedings of the 8th International Symposium on Trichoptera: Minneapolis and Lake Itasca, Minnesota, U.S.A., 9–15 August 1995. Ohio Biological Survey, Columbus, Ohio, 401–410.
  60. Riek EF. (1953) Fossil Mecopteroid insects from the Upper Permian of New South Wales. Records of the Australian Museum 23(2): 55–87. 10.3853/j.0067-1975.23.1953.621 [DOI] [Google Scholar]
  61. Robertson DR, Holzenthal RW. (2013) Revision and phylogeny of the caddisfly subfamily Protoptilinae (Trichoptera: Glossosomatidae) inferred from adult morphology and mitochondrial DNA. Zootaxa 3723(1): 1–99. 10.11646/zootaxa.3723.1.1 [DOI] [PubMed] [Google Scholar]
  62. Ross HH. (1956) Evolution and Classification of the Mountain Caddisflies. University of Illinois Press, Urbana, 213 pp. [Google Scholar]
  63. Ross HH, Gibbs DG. (1973) The subfamily relationships of the Dipseudopsinae (Trichoptera, Polycentropodidae). Journal of the Georgia Entomological Society 8(4): 312–316. [Google Scholar]
  64. Schmid F. (1982) La famille des xiphocentronides (Trichoptera: Annulipalpia). Memoirs of the Entomological Society of Canada 121(S121): 1–127. 10.4039/entm114121fv [DOI]
  65. St Clair R. (1988) The adult and immatures of Russobex gen. nov., a new monotypic genus from Victoria (Trichoptera: Leptoceridae). Proceedings of the Royal Society of Victoria 100: 47–52. [Google Scholar]
  66. Stehr FW. (1987) 26, Order Lepidoptera. In: Stehr FW. (Ed.) Immature Insects.Kendall/Hunt Publishing Company, Dubuque, Iowa, 288–596.
  67. Stoltz M. (1989) The Afrotropical caddisfly family Pisuliidae, Systematics, zoogeography, and biology (Trichoptera: Pisuliidae). Steenstrupia (Copenhagen) 15(1): 1–49. [Google Scholar]
  68. Sturm H. (1960) Die terrestrischen Puppengehäuse von Xiphocentron sturmi Ross (Xiphocentronidae, Trichoptera). Zoologische Jarbücher Systematik 87(4/5): 387–394.
  69. Tachet H, Morse JC, Berly A. (2001) The larva and pupa of Pseudoneureclipsis lusitanicus Malicky, 1980 (Trichoptera: Hydropsychoidea): Description, ecological data and taxonomical considerations. Aquatic Insects 23(2): 93–106. 10.1076/aqin.23.2.93.4917 [DOI] [Google Scholar]
  70. Tanida K. (2002) Stenopsyche (Trichoptera: Stenopsychidae): Ecology and biology of a prominent Asian caddis genus. In: Mey W. (Ed.) Proceedings of the 10th International Symposium on Trichoptera.Nova Supplementa Entomologica, Keltern 15: 595–606.
  71. Thienemann A. (1904) Ptilocolepus granulatus Pt., eine Übergangsform von den Rhyacophiliden zu den Hydroptiliden. Allgemein Zeitschrift für Entomologie 9(21–22): 418–441. [Google Scholar]
  72. Thomas JA, Frandsen PB, Prendini E, Zhou X, Holzenthal RW. (2020) A multigene phylogeny and timeline for Trichoptera (Insecta). Systematic Entomology 45(3): 670–686. 10.1111/syen.12422 [DOI] [Google Scholar]
  73. Tillyard RJ. (1926) Upper Permian insects of New South Wales. Part II: The orders Mecoptera, Paramecoptera and Neuroptera. Proceedings of the Linnean Society of New South Wales 51(3): 265–282. [pls xv, xvi.] [Google Scholar]
  74. Ulmer G. (1957) Köcherfliegen (Trichopteren) von den Sunda-Inseln, Teil III. Archiv für Hydrobiologie Supplement 23(2/4): 109–470.
  75. van Nieukerken EJ, Kaila L, Kitching IJ, Kristensen NP, Lees DC, Minet J, Mitter C, Mutanen M, Regier JC, Simonsen TJ, Wahlberg N, Yen S-h, Zahiri R, Adamski D, Baixeras J, Bartsch D, Bengtsson BÅ, Brown JW, Bucheli SR, Davis DR, Prins JD, Prins WD, Epstein ME, Gentili-Poole P, Gielis C, Hättenschwiler P, Hausmann A, Holloway JD, Kallies A, Karsholt O, Kawahara AY, Koster SJC, Kozlov MV, Lafontaine JD, Lamas G, Landry J-F, Lee S, Nuss M, Park K-T, Penz C, Rota J, Schintlmeister A, Schmidt BC, Sohn J-C, Solis MA, Tarmann GM, Warren AD, Weller S, Yakovlev RV, Zolotuhin VV, Zwick A. (2011) Order Lepidoptera Linnaeus, 1758. Zootaxa 3148: 212–221. 10.11646/zootaxa.3148.1.41 [accessed 03 June 2025] [DOI] [Google Scholar]
  76. Vilarino A, Dias ES, Bispo PC. (2022) Phylogeny indicates polyphyly in Cnodocentron (Trichoptera: Xiphocentronidae): Biogeography and revision of New World species (Caenocentron). Zoological Journal of the Linnean Society 194(4): 1341–1373. 10.1093/zoolinnean/zlab077 [DOI] [Google Scholar]
  77. Vshivkova TS, Morse JC, Ruiter D. (2007) Phylogeny of Limnephilidae and composition of the genus Limnephilus (Limnephilidae: Limnephilinae, Limnephilini). In: Bueno-Soria J, Barba-Alvarez R, Armitage B. (Eds) Proceedings of the XIIth International Symposium on Trichoptera, June 18–22, 2006.The Caddis Press, Columbus, Ohio, 309–319.
  78. Wallace JB, Ross HH. (1971) Pseudogoerinae: A new subfamily of Odontoceridae (Trichoptera). Annals of the Entomological Society of America 64(4): 890–894. 10.1093/aesa/64.4.890 [DOI] [Google Scholar]
  79. Wallace JB, Sherberger FF. (1970) The immature stages of Anisocentropus pyraloides (Trichoptera: Calamoceratidae). Journal of the Georgia Entomological Society 5(4): 217–224. [Google Scholar]
  80. Wallace JB, Woodall WR, Staats AA. (1976) The larval dwelling-tube, capture net and food of Phylocentropus placidus (Trichoptera: Polycentropodidae). Annals of the Entomological Society of America 69(1): 149–154. 10.1093/aesa/69.1.149 [DOI] [Google Scholar]
  81. Walsingham TdG. (1880) On some new and little known species of Tineidae. Proceedings of the Zoological Society of London, 77–93. [pls XI–XII.]
  82. Walsingham TdG. (1898) Descriptions of a new micropterigid genus and species and a new eriocraniad species from N. America. Entomologist’s Record and Journal of Variation 10: 161–163. [pl. II.] [Google Scholar]
  83. Ward JB, Leschen RAB, Smith BJ, Dean JD. (2004) Phylogeny of the caddisfly (Trichoptera) family Hydrobiosidae using larval and adult morphology, with the description of a new genus and species from Fiordland, New Zealand. Records of the Canterbury Museum 18: 23–43. [Google Scholar]
  84. Waringer J, Graf W. (2002) Ecology, morphology and distribution of Ptilocolepus granulatus (Pictet 1834) (Insecta: Trichoptera) in Austria. Lauterbornia 43: 121–129. [Google Scholar]
  85. Waringer J, Graf W. (2011) Atlas of Central European Trichoptera larvae. Eric Mauch Verlag, Dinkelscherben, vi + 468 pp.
  86. Waringer J, Malicky H. (2016) The larvae of the European species of genus Apataniana Mosely, 1936 (Trichoptera: Apataniidae): Descriptions, key and ecology. ZooKeys 586: 121–134. 10.3897/zookeys.586.7758 [DOI] [PMC free article] [PubMed] [Google Scholar]
  87. Weaver JS III. (1984) The evolution and classification of Trichoptera, part I: The groundplan of Trichoptera. In: Morse JC. (Ed.) Proceedings of the 4th International Symposium on Trichoptera, Clemson, South Carolina, 11–16 July 1983. W.Junk Publishers, The Hague, Series Entomologica 30: 413–419.
  88. Wiggins GB. (1965) Additions and revisions to the genera of North American caddisflies of the family Brachycentridae with special reference to the larval stages (Trichoptera). Canadian Entomologist 97(10): 1089–1106. 10.4039/Ent971089-10 [DOI] [Google Scholar]
  89. Wiggins GB. (1996) Larvae of the North American Caddisfly Genera (Trichoptera). Second Edition. University of Toronto Press, Toronto, 457 pp. 10.3138/9781442623606 [DOI] [Google Scholar]
  90. Wiggins GB. (1998) The Caddisfly Family Phryganeidae. University of Toronto Press, Toronto, 320 pp. [Google Scholar]
  91. Wiggins GB. (2004) Caddisflies: The Underwater Architects. University of Toronto Press, Toronto, 292 pp. 10.3138/9781442623590 [DOI] [Google Scholar]
  92. Wiggins GB, Weaver JS III, Unzicker JD. (1985) Revision of the caddisfly family Uenoidae (Trichoptera). Canadian Entomologist 117(6): 763–800. 10.4039/Ent117763-6 [DOI] [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

All of the data that support the findings of this study are available in the main text.

Articles from ZooKeys are provided here courtesy of Pensoft Publishers

RESOURCES