Open Access Open Access  Restricted Access Subscription or Fee Access


Revision and phylogeny of the caddisfly subfamily Protoptilinae (Trichoptera: Glossosomatidae) inferred from adult morphology and mitochondrial DNA



Protoptilinae Ross, 1956, is the most diverse subfamily belonging to the saddle- or tortoise-case-making caddisfly family Glossosomatidae Wallengren, 1891. The subfamily has a disjunct distribution: 5 genera are known from the East Palaearctic and Oriental regions; the remaining 13 are restricted to the Nearctic and Neotropical regions. Monophyly of Protoptilinae and each of 17 genera was tested using 80 taxa, 99 morphological characters, and mitochondrial DNA (COI). Additionally, homologies of morphological characters were assessed across genera and a standardized terminology for those structures was established. Mitochondrial DNA data were unavailable for 55 of the 80 taxa included in this study. To test the effects of the missing molecular data, 5 different datasets were analyzed using both parsimony and Bayesian methods. There was incongruence between the COI and morphological data, but results suggest the inclusion of COI data in a combined analysis, although incomplete, improved the overall phylogenetic signal. Bayesian and parsimony analyses of all 5 datasets strongly supported the monophyly of Protoptilinae. Monophyly of the following genera was also supported: Canoptila Mosely, 1939; Culoptila Mosely, 1954; Itauara Müller, 1888; Mastigoptila Flint, 1967; Mortoniella Ulmer, 1906; Protoptila Banks, 1904; and Tolhuaca Schmid, 1964. Several taxonomic changes were necessary for classification to reflect phylogeny accurately. Accordingly, Matrioptila Ross, 1938; Poeciloptila Schmid, 1991; Temburongpsyche Malicky, 1992; and Nepaloptila Kimmins, 1964, are designated new junior synonyms of Padunia Martynov, 1910. Additionally, the endemic Caribbean genera Campsiophora Flint, 1964, and Cubanoptila Sykora, 1973, are designated new junior synonyms of Cariboptila Flint, 1964. Diagnoses and a key to the subfamilies of Glossosomatidae and world genera of Protoptilinae incorporating these taxonomic changes are provided.


taxonomy, identification key, male genitalia, female genitalia, morphology, COI, missing data, combined analyses

Full Text:

PDF/A (6.27 MB)


Alfaro, M.E., Zoller, S. & Lutzoni, F. (2003) Bayes or bootstrap? A simulation study comparing the performance of Bayesian Markov Chain Monte Carlo sampling and bootstrapping in assessing phylogenetic confidence. Molecular Biology and Evolution, 20, 255–266.

Angrisano, E.B. (1993) Contribution al conocimiento del genero Antoptila Mosely (Trichoptera: Glossosomatidae). Revista de la Sociedad Entomológica Argentina, 52, 57–62.

Angrisano, E.B. (1997) Los Trichoptera del Uruguay. III. Familias Philopotamidae, Hydrobiosidae y Glossosomatidae. Revista de la Sociedad Entomológica Argentina, 56, 55–58.

Ball, S.L., Hebert, P.D.N., Burian, S.K. & Webb, J.M. (2005) Biological identifications of mayflies (Ephemeroptera) using DNA barcodes. Journal of the North American Benthological Society, 24, 508–524.

Banks, N. (1904) Two new species of Hydroptilidae. Entomological News, 15, 116.

Blahnik, R.J. & Holzenthal, R.W. (2004) Collection and curation of Trichoptera, with an emphasis on pinned material. Nectopsyche, Neotropical Trichoptera Newsletter, 1, 8–20.

Blahnik, R.J. & Holzenthal, R.W. (2006) Revision of the genus Culoptila (Trichoptera: Glossosomatidae). Zootaxa, 1233, 1–52.

Blahnik, R.J. & Holzenthal, R.W. (2008) Revision of the Mexican and Central American species of Mortoniella (Trichoptera: Glossosomatidae: Protoptilinae). Zootaxa, 1711, 1–72.

Blahnik, R.J. & Holzenthal, R.W. (2011) Revision of the austral South American species of Mortoniella (Trichoptera: Glossosomatidae: Protoptilinae). Zootaxa, 2851, 1–75.

Blahnik, R.J., Holzenthal, R.W. & Prather, A.L. (2007) The lactic acid method for clearing Trichoptera genitalia. In: Bueno-Soria, J., Barba-Alvarez, R. & Armitage, B.J. (Eds.) Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press, Columbus, Ohio, pp. 9–14.

Bond, J.E. & Hedin, M. (2006) A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference. Molecular Phylogenetics and Evolution, 41, 70–85.

Botosaneanu, L. (1977) Trichoptères (imagos) de Cuba, capturés par moi-meme en 1973 (Insecta, Trichoptera). Fragmenta Entomologica, 13, 231–284.

Botosaneanu, L. (1994) A study of the larvae of caddisflies (Trichoptera) from Cuba. Tropical Zoology, 7, 451–475.

Botosaneanu, L. (1996) Caddis flies (Trichoptera) from the Dominican Republic (West Indies). II. All families except Hydroptilidae; with general observations for Hispaniola. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Entomologie, 66, 5–26.

Botosaneanu, L. (1998) On several Hydropsyche from Iran - with description of a new species (Trichoptera: Hydropsychidae). Entomologische Zeitschrift, 108, 496–500.

Botosaneanu, L. & Alkins-Koo, M. The caddis flies (Insecta: Trichoptera) of Trinidad and Tobago, West Indies. Bulletin de l’Insitut Royal des Sciences Naturelles de Belgique, Entomologie, 63, 5–45.

Botosaneanu, L. & Sykora, J. (1973) Sur quelques Trichoptéres (Insecta: Trichoptera) de Cuba. In: Résultats des Expéditions Biospéologiques cubano-roumaines Cuba. Editura Academiei Republicii Socialiste România., Bucharest, pp. 379–407.

Brandley, M.C., Schmitz, A. & Reeder, T.W. (2005) Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards. Systematic Biology, 54, 373–390.

Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795–803.

Bryant, H.N. (1995) Why autapomorphies should be removed: A Reply to Yeates. Cladistics, 11, 381–384.

Buckley, T.R. (2002) Model misspecification and probalistic tests of topology: Evidence from empirical data sets. Systematic Biology, 51, 509–523.

Buckley, T.R., Arensburger, P., Simon, C. & Chambers, G.K. (2002) Combined data, Bayesian phylogenetics, and the origins of the New Zealand cicada genera. Systematic Biology, 51, 4–18.

Bueno-Soria, J. (1983) Five new species of caddisflies (Trichoptera) from Mexico. Proceedings of the Entomological Society of Washington, 85, 450–455.

Bueno-Soria, J. (1984) Three new species of the genus Protoptila from Mexico and Costa Rica (Trichoptera: Glossosomatidae). Proceedings of the Biological Society of Washington, 97, 392–394.

Bueno-Soria, J. & Santiago-Fragoso, S. (1979) Una nueva especie del género Protoptila Banks (Trichoptera: Glossosomatidae) de Veracruz, México. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoología, 50, 477–480.

Bueno-Soria, J. & Santiago-Fragoso, S. (1996) Studies in aquatic insects X: Descriptions of five new species of genus Culoptila Mosely (Trichoptera: Glossosomatidae) from México. Proceedings of the Biological Society of Washington, 109, 446–452.

Cassis, G. & Schuh, R.T. (2009) Systematic methods, fossils, and relationships within Heteroptera (Insecta). Cladistics, 26, 262–280.

Castlebury, L.A., Rossman, A.Y., Jaklitsch, W.J. & Vasilyeva, L.N. (2002) A preliminary overview of the Diaporthales based on large subunit nuclear ribosomal DNA sequences. Mycologia, 94, 1017–1031.

Colwell, R.K. (2003) Biota 2: The Biodiversity Database Manager. Sunderland, Massachusetts.

DeBry, R.W. (2001) Improving interpretation of the decay index for DNA sequence data. Systematic Biology, 50, 742–752.

deWaard, J.R., Ivanova, N.V., Haijbabaei, M. & Hebert, P.D.N. (2008) Assembling DNA barcodes: Analytical protocols. In: Martin, C.C. (Ed.) Environmental Genomics, Methods in Molecular Biology. Humana Press, pp. 275–283.

Dos Santos, C.M. & Falaschi, R.L. (2007) Missing data in phylogenetic analysis: Comments on support measures. Darwiniana, 45, 25–26.

Durbin, R., Eddy, S., Krogh, A. & Mitchison, G. (1998) Biological Sequence Analysis: Probalistic Models of Proteins and Nucleic Acids. Cambridge University Press, Cambridge, U.K., 368 pp.

Efron, B., Halloran, E. & Holmes, S. (1996) Bootstrap confidence levels for phylogenetic trees. Proceedings of the National Academy of Sciences of the United States of America, 93, 7085–7090.

Egge, J.D. & Simons, A.M. (2009) Molecules, morphology, missing data and the phylogenetic position of a recently extinct madom catfish (Actinopterygii: Ictaluridae). Zoological Journal of the Linnean Society, 155, 60–75.

Erixon, P., Svennblad, B., Britton, T. & Oxelman, B. (2003) Reliability of Bayesian posterior probabilities and bootstrap frequencies in phylogenetics. Systematic Biology, 52, 665–673.

Felsenstein, J. (1978) Cases in which parsimony and compatibility methods will be positively misleading. Systematic Zoology, 27, 401–410.

Felsenstein, J. (1985) Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39, 783–791.

Felsenstein, J. (2004) Inferring Phylogenies. Second Edition. Sinauer Associates, Sunderland, Massachusetts, 664 pp.

Fischer, F.C.J. (1966) Leptoceridae Pars 2. Trichopterorum Catalogus VII. Nederlandsche Entomologische Vereeniging, Amsterdam, iii + 163 pp.

Fischer, F.C.J. (1971) Supplement to Vols. I and II, Trichopterorum Catalogus XII. Nederlandsche Entomologische Vereeniging, Amsterdam, vii + 311 pp.

Fitch, W.M. (1971) Toward defining the course of evolution: Minimum change for a specific tree topology. Systematic Zoology, 20, 406–416.

Fitch, W.M. & Bruschi, M. (1987) The evolution of prokaryotic ferredoxins – with a general method correcting for unobserved substitutions in less branched lineages. Molecular Biology and Evolution, 4, 381–394.

Flint, O.S. Jr. (1962) The immature stages of Matrioptila jeanae (Ross) (Trichoptera: Glossosomatidae). Journal of the New York Entomological Society, 70, 64–67.

Flint, O.S. Jr. (1963) Studies of Neotropical caddis flies, I: Rhyacophilidae and Glossosomatidae (Trichoptera). Proceedings of the United States National Museum, 114, 453–478.

Flint, O.S. Jr. (1964) The caddisflies (Trichoptera) of Puerto Rico. University of Puerto Rica, Agricultural Experiment Station, Technical Paper, 40, 1–80.

Flint, O.S. Jr. (1967a) Studies of Neotropical caddis flies, IV: New species from Mexico and Central America. Proceedings of the United States National Museum, 123, 1–24.

Flint, O.S. Jr. (1967b) Trichoptera collected by Prof. J. Illies in the Chilean subregion. Beiträge zur Neotropischen Fauna, 5, 45–68.

Flint, O.S. Jr. (1968) The caddisflies of Jamaica. Bulletin of the Institute of Jamaica, Science Series, 19, 1–68.

Flint, O.S. Jr. (1971) Studies of Neotropical caddisflies, XII: Rhyacophilidae, Glossosomatidae, Philopotamidae, and Psychomyiidae from the Amazon Basin (Trichoptera). Amazoniana, 3, 1–67.

Flint, O.S. Jr. (1974) Studies of Neotropical caddisflies, XVIII: New species of Rhyacophilidae and Glossosomatidae (Trichoptera). Smithsonian Contributions to Zoology, 169, 1–30.

Flint, O.S. Jr. (1981) Studies of Neotropical caddisflies, XXVIII: The Trichoptera of the Río Limón Basin, Venezuela. Smithsonian Contributions to Zoology, 330, 1–61.

Flint, O.S. Jr. (1983) Studies of Neotropical caddisflies, XXXIII: New species from austral South America (Trichoptera). Smithsonian Contributions to Zoology, 377, 1–100.

Flint, O.S. Jr. (1991) Studies of Neotropical caddisflies, XLV: The taxonomy, phenology, and faunistics of the Trichoptera of Antioquia, Colombia. Smithsonian Contributions to Zoology, 520, 1–113.

Flint, O.S. Jr. (1992) New species of caddisflies from Puerto Rico (Trichoptera). Proceedings of the Entomological Society of Washington, 94, 379–389.

Flint, O.S. Jr., Holzenthal, R.W. & Harris, S.C. (1999) Nomenclatural and systematic changes in the Neotropical caddisflies. Insecta Mundi, 13, 73–84.

Flint, O.S. Jr. & Sykora, J.L. (2004) Caddisflies of Hispaniola, with special reference to the Dominican Republic (Insecta: Trichoptera). Annals of Carnegie Museum, 73, 1–60.

Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxydase subunit from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.

Frania, H.E. & Wiggins, G.B. (1997) Analysis of morphological and behavioural evidence for the phylogeny and higher classification of Trichoptera (Insecta). Life Sciences Contributions, Royal Ontario Museum, 160, 1–67.

Goloboff, P.A. (1999) Analyzing large data sets in reasonable times: Solutions for composite optima. Cladistics, 15, 415–428.

Goloboff, P.A. (2002) Chapter 4. Techniques for analyzing large data sets. In: DeSalle, R. (Ed.) Methods and Tools in Biosciences and Medicine: Techniques in Molecular Systematics and evolution. Birkhauser Verlag Basel, Switzerland.

Goloboff, P.A. & Farris, J.S. (2001) Methods for quick consensus estimation. Cladistics, 17, 526–534.

Haijbabaei, M., deWaard, J.R., Ivanova, N.V., Ratnasingham, S., Dooh, R.T., Kirk, S.L., Mackie, P.M. & Hebert, P.D.N. (2005) Critical factors for assembling a high volume of DNA barcodes. Philisophical Transactions of the Royal Society B - Biological Sciences, 360, 1959–1967.

Hawkins, J.A., Hughes, C.E. & Scotland, R.W. (1997) Primary homology assessment, characters and character states. Cladistics, 13, 275–283.

Heath, T.A., Hedtke, S.M. & Hillis, D.M. (2008) Taxon sampling and the accuracy of phylogenetic analyses. Journal of Systematics and Evolution, 46, 239–257.

Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, Series B, 270, 313–321.

Hebert, P.D.N., Penton, E.H., Burns, J.M., Janzen, D.H. & Hallwachs, W. (2004) Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101, 14812–14817.

Hedtke, S.M., Townsend, T.M. & Hillis, D.M. (2006) Resolution of phylogenetic conflict in large data sets by increased taxon sampling. Systematic Biology, 55, 522–529.

Holder, M. & Lewis, P.O. (2003) Phylogeny estimation: Traditional and Bayesian approaches. Nature Reviews, 4, 275–284.

Holzenthal, R.W. (2004) Three new species of Chilean caddisflies (Insecta: Trichoptera). Proceedings of the Entomological Society of Washington, 106, 110–117.

Holzenthal, R.W. & Andersen, T. (2004) The caddisfly genus Triaenodes in the Neotropics (Trichoptera: Leptoceridae). Zootaxa, 511, 1–80.

Holzenthal, R.W. & Blahnik, R.J. (2006) The caddisfly genus Protoptila in Costa Rica (Trichoptera: Glossosomatidae). Zootaxa, 1197, 1–37.

Holzenthal, R.W., Blahnik, R.J., Kjer, K.M., Prather, A. (2007) An update on the phylogeny of caddisflies (Trichoptera). In: Bueno-Soria, J., Barba-Alvarez, R., Armitage, B. (Eds.), Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press, Columbus, Ohio, pp. 143–153.

Holzenthal, R.W., Blahnik, R.J., Prather, A.L. & Kjer, K.M. (2007) Order Trichoptera Kirby, 1813 (Insecta), caddisflies. Zootaxa, 1668, 639–698.

Houghton, D.C. & Stewart, K.W. (1998a) Immature life history descriptions and distribution of Culoptila cantha (Ross) (Trichoptera: Glossosomatidae). Proceedings of the Entomological Society of Washington, 100, 511–520.

Houghton, D.C. & Stewart, K.W. (1998b) Life history and case-building behavior of Culoptila cantha (Trichoptera: Glossosomatidae) in the Brazos River, Texas. Annals of the Entomological Society of America, 91, 59–70.

Huelsenbeck, J.P. (1991) When are fossils better than extant taxa in phylogenetic analysis? Systematic Zoology, 40, 458–469.

Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.

Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314.

Ivanova, N.V., deWaard, J.R. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002.

Kagaya, T. & Nozaki, T. (1998) Notes on Japanese Padunia Martynov (Trichoptera: Glossosomatidae), with the description of a new species. Aquatic Insects, 20, 97–107.

Kearney, M. (2002) Fragmentary taxa, missing data, and ambiguity: Mistaken assumptions and conclusions. Systematic Biology, 51, 369–381.

Kearney, M. & Clark, J.M. (2003) Problems due to missing data in phylogenetic analyses including fossils: A critical review. Journal of Vertebrate Paleontology, 23, 263–274.[0263:pdtmdi];2

Kimmins, D.E. (1964) On the Trichoptera of Nepal. Bulletin of the British Museum (Natural History) Entomology, 15, 33–55.

Kitching, I.J., Forey, P.L., Humphries, C.J. & Williams, D.J. (1998) Cladistics, 2nd edition. The Theory and Practice or Parsimony Analysis. Oxford University Press, Oxford, 228 pp.

Kjer, K.M., Blahnik, R.J. & Holzenthal, R.W. (2001) Phylogeny of Trichoptera (Caddisflies): Characterization of signal and noise within multiple datasets. Systematic Biology, 50, 781–816.

Kolaczkowski, B. & Thornton, J.W. (2009) Long-branch attraction bias and inconsistency in Bayesian phylogenetics. PLoS ONE, 4:e7891.

Kumanski, K.P. (1987) On caddisflies (Trichoptera) of Cuba. Acta Zoologica Bulgarica, 34, 3–35.

Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez, R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics, 23, 2947–2948.

Larson, A. (1994) The comparison of morphological and molecular data in phylogenetic systematics. In: Schierwater, B., Streit, B., Wagner, G.P. & DeSalle, R. (Eds.) Molecular Ecology and Evolution: Approaches and Applications. Birkhauser, Basel, Switzerland.

Leach, A.D. & Reeder, T.W. (2002) Molecular systematics of the eastern fence lizard (Sceloporus undulatus): A comparison of parsimony, likelihood, and Bayesian approaches. Systematic Biology, 51, 44–68.

Lee, D.C. & Bryant, H.N. (1999) A reconsideration of the coding of inapplicable characters: Assumptions and problems. Cladistics, 15, 373–378.

Lemmon, A.R., Brown, J.M., Stanger-Hall, K. & Lemmon, E.M. (2009) The effect of ambiguous data on phylogenetic estimates obtained by Maximum Likelihood and Bayesian inference. Systematic Biology, 58, 130–145.

Lewis, P.O. (2001a) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology, 50, 913–925.

Lewis, P.O. (2001b) Phylogenetic systematics turns over a new leaf. TRENDS in Ecology & Evolution, 16, 30–37.

Maddison, D.R. & Maddison, W.P. (2000) MacClade 4: Analysis of phylogeny and character evolution, Version 4.0. Sinauer Associates, Sunderland, Massachusetts.

Maddison, W.P. (1993) Missing data versus missing characters in phylogenetic analysis. Systematic Biology, 42, 576–581.

Malicky, H. (1995) Weitere neue Kocherfliegen (Trichoptera) aus Asien. Braueria, 22, 11–26.

Malicky, H. (2013) Synonyms and possible synonyms of Asiatic Trichoptera. Braueria, 40, 41–54.

Malicky, H. & Chantaramongkol, P. (1992) Neue Köcherfliegen (Trichoptera) aus Thailand und angrenzenden Ländern. (Arbeiten über thailändische Köcherfliegen Nr. 11.) Braueria, 19, 13–23.

Malicky, H. & Chantaramongkol, P. (2009) Beiträge Kenntnis asiatischer Trichopteren. Braueria, 36, 11–58.

Malicky, H., Chantaramongkol, P., Changthong, N., Nuntakwang, A. & Thamsenanupap, P. (2006) Beschreibungen einiger Köcherfliegen aus Nord-Thailand (Trichoptera) (Arbeit Nr. 41 über thailändische Köcherfliegen.) Braueria, 33, 39–42.

Malicky, H. & Silamon, S. (2012) Eine neue Gattung und vier neue Arten von Trichopteren aus Thailand. Braueria, 39, 22–24.

Marlier, G. (1955) Un trichoptère pélagique nouveau du lac Tanganika. Revue de Zoologie et de Botanique Africaines, 52, 150–155.

Marshall, J.E. (1979) A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the British Museum (Natural History) Entomology, 39, 135–239.

Martynov, A.B. (1910) Les Trichoptères de la Sibèrie et des régions adjacentes. II. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de Saint Pétersbourg, 15, 351–429.

Martynov, A.B. (1913) Къ познанію фауны Trichoptera Кавказа I. (Қъ отчету о командировкъ). [Contribution to the knowledge of the Trichopterous fauna of the Caucasus.] (In Russian). Raboti iz zoologicheskago Kabineta Varshavskago de Varsovie [Travaux du Cabinet zoologique de l’Universit de Varsovie], 1912, 1–111, pls. I–IX.

Martynov, A.B. (1929) On a collection of Trichoptera from the River Bija and from the vicinities of the Lake Teletzkoje. Konowia, Vienna, 8, 293–311, 216 figs.

Martynov, A.V. (1934) Tableaux analytiques de la faune de l'URSS, publiés par l'institut zoologisque de l'Académie des sciences. Trichoptera Annulipalpia. I. Travaux de I'nstitute Zoologique de l'Académie des Science de l'URSS, 343 pp.

Morse, J.C. (1988) A new species of Ceraclea (Athripsodina) (Trichoptera: Leptoceridae) from Lake Tanganyika. Revue de Zoologie Africaine, 102, 43–45.

Morse, J.C. (2013) Trichoptera World Checklist. Available from (accessed 18 February 2013).

Morse, J.C. & Holzenthal, R.W. (2008) Chapter 18, Caddisfly genera. In: Merritt, R.W., Cummins, K.W. & Berg, M.A. (Eds.) An Introduction to the Aquatic Insects of North America, 4th edition. Kendall/Hunt, Dubuque, pp. 481–552.

Morse, J.C. & Yang, L. (1993) Higher classification of the Chinese Glossosomatidae (Trichoptera). In: Otto, C. (Ed.) Proceedings of the 7th International Symposium on Trichoptera. Backhuys Publishers, Leiden, The Netherlands, pp. 139–148.

Morse, J.C. & Yang, L. (2004) The world subgenera of Glossosoma Curtis (Trichoptera: Glossosomatidae), with a revision of the Chinese species of Glossosoma subgenera Synafophora Martynov and Protoglossa Ross. Proceedings of the Entomological Society of Washington, 106, 52–73.

Mosely, M.E. (1934) New exotic Hydroptilidae. Transactions of the Royal Entomological Society of London, 82, 137–163.

Mosely, M.E. (1937) Mexican Hydroptilidae (Trichoptera). Transactions of the Royal Entomological Society of London, 86, 151–189.

Mosely, M.E. (1939) The Brazilian Hydroptilidae (Trichoptera). Novitates Zoologicae, 41, 217–239.

Mosely, M.E. (1954) The Protoptila group of the Glossosomatinae (Trichoptera: Rhyacophilidae). Bulletin of the British Museum (Natural History) Entomology, 3, 317–346.

Müller, F. (1888) Larven von Mücken und Haarflüglern mit zweierlei abwechselnd thätigen Athemwerkzeugen. Entomologische Nachrichten, 14, 273–277.

Müller, F. (1921) Briefe un noch nicht veröffentliche Abhandlungen aus dem nachlass 1854–1897. In: Möller, A. (Ed.) Fritz Müller: Werke, Briefe und Leben, 2. G. Fischer, Jena, pp. 383–642.

Müller, J. & Reisz, R.R. (2006) The phylogeny of early eureptiles: Comparing parsimony and Bayesian approaches in the investigation of a basal fossil clade. Systematic Biology, 55, 503–511.

Nichols, S.W. (1989) The Torre-Bueno Glossary of Entomology by J.R. de la Torre-Bueno, Revised Edition. New York Entomological Society, New York, 840 pp.

Nielsen, A. (1957) A comparative study of the genital segments and their appendages in male Trichoptera. Biologiske Skrifter, 8, 1–159.

Nielsen, A. (1980) A comparative study of the genital segments and the genital chamber in female Trichoptera. Kongelige Danske Videnskabernes Selskab Biologiske Skrifter, 23, 1–200.

Nishimoto, H. & Nozaki, T. (2007) New species of Padunia Martynov (Trichoptera: Glossosomatidae) from Japan. In: Bueno-Soria, J., Barba-Álvarez, R. & Armitage, B.J. (Eds.) Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press, Columbus, Ohio, pp. 229–242.

Nylander, J.A.A., Ronquist, F., Huelsenbeck, J.P. & Joséluis, N.-A. (2004) Bayesian phylogenetic analysis of combined data. Systematic Biology, 53, 47–67.

Nyman, T., Zinovjev, A.G., Vikberg, V. & Farrell, B.D. (2006) Molecular phylogeny of the sawfly subfamily Nematinae (Hymenoptera: Tenthredinidae). Systematic Entomology, 31, 569–583.

Patterson, C. (1982) Morphological characters and homology. In: Friday, J.J.A. & Friday, A.E. (Eds.) Problems in phylogenetic reconstruction. Academic Press, London, pp. 21–74.

Pauls, S., Blahnik, R.J., Zhou, X., Wardwell, C.T., & Holzenthal, R.W. (2010) DNA barcode data confirm new species and reveal cryptic diversity in Chilean Smicridea (Smicridea) (Trichoptera: Hydropsychidae). Journal of the North American Benthological Society, 29, 1058–1074.

Pleijel, F. (1995) On character coding for phylogeny reconstruction. Cladistics, 11, 309–315.

Posada, D. & Crandall, K.A. (1998) MODELTEST: Testing the model of DNA substitution. Bioinformatics, 14, 817–818.

Prendini, L. (2001) Species or supraspecific taxa as terminals in cladistic analysis? Groundplans versus exemplars revisited. Systematic Biology, 50, 290–300.

Rambaut, A. (2007) FigTree v1.3. Available from (Accessed 15 August 2009)

Rambaut, A. & Drummond, A.J. (2007) Tracer v1.4, Available from (accessed 5 August 2009).

Remane, A. (1952) Die Grundlagen des Naturlichen Systems der Vergleichenden Anatomie und der Phylogenetik. Akademische Verlagsgesellschaft, Leipzig, Germany

Robertson, D.R. & Holzenthal, R.W. (2005) The Neotropical caddisfly genus Tolhuaca (Trichoptera: Glossosomatidae). Zootaxa, 1063, 53–68.

Robertson, D.R. & Holzenthal, R.W. (2006) The Neotropical caddisfly genus Canoptila (Trichoptera: Glossosomatidae). Zootaxa, 1272, 45–59.

Robertson, D.R. & Holzenthal, R.W. (2008) Two new species and a new record of Protoptila from Bolivia (Trichoptera: Glossosomatidae: Protoptilinae). Annals of the Entomological Society of America, 101, 465–473.[465:tnsaan];2

Robertson, D.R. & Holzenthal, R.W. (2011) Revision of the Neotropical caddisfly genus Itauara Müller, 1888 (Trichoptera, Glossosomatidae). ZooKeys, 114, 41–100.

Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

Ronquist, F., Huelsenbeck, J.P. & van der Mark, P. (2005) MrBayes 3.1 Manual. Florida State University. Available from (Accessed 5 October 2009)

Ross, H.H. (1938) Descriptions of Nearctic caddis flies (Trichoptera) with special reference to the Illinois species. Bulletin of the Illinois Natural History Survey, 21, 101–183.

Ross, H.H. (1941) Descriptions and records of North American Trichoptera. Transactions of the American Entomological Society, 67, 35–126.

Ross, H.H. (1944) The caddisflies or Trichoptera of Illinois. Bulletin of the Illinois Natural History Survey, 23, 1–326.

Ross, H.H. (1956) Evolution and Classification of the Mountain Caddisflies. University of Illinois Press, Urbana, 213 pp.

Rueda Martin, P.A. & Gibon, F.M. (2008) New species of Glossosomatidae from Bolivia with new records from Bolivia and northwestern Argentina (Trichoptera: Glossosomatidae). Annales de Limnologie, 44, 215–225.

Santos, A.P.M. & Nessimian, J.L. (2009) A new species of Protoptila Banks (Trichoptera: Glossosomatidae: Protoptilinae) from Brazil. Acta Amazonica, 39, 723–726.

Schmid, F. (1958) Contribution a l'etude des Trichoptères Neotropicaux III. Mitteilungen aus dem Zoologischen Museum in Berlin, 34, 183–217.

Schmid, F. (1959) Contribution a l'etude des trichoptères Neotropicaux IV. Acta Zoologica Lilloana, 17, 477–483.

Schmid, F. (1964) Contribution a l'etude des trichopteres neotropicaux V. Tijdschrift voor Entomologie, 107, 307–339.

Schmid, F. (1989) Les hydrobiosides (Trichoptera, Annulipalpia). Bulletin de l'Institute Royal des Sciences Naturelles de Belgique, Entomologie, 59, 1–154.

Schmid, F. (1990 [1991]) Quelques nouveaux trichoptères indiens (Trichoptera). Naturaliste Canadien, 117, 239–251.

Schmid, F. (1998) The insects and arachnids of Canada, Part 7. Genera of the Trichoptera of Canada and adjoining or adjacent United States. NRC Research Press, Ottawa, 319 pp.

Schuh, R.T. & Brower, A.V.Z. (2009) Biological Systematics: Principles and Applications, 2nd edition. Cornell University Press, Ithaca, 311 pp.

Simmons, M.P., Pickett, K.M. & Miya, M. (2004) How meaningful are Bayesian support values? Molecular Biology and Evolution, 21, 188–199.

Simon, C., Buckley, T.R., Frati, F., Stewart, J.B. & Beckenbach, A. (2006) Incorporating molecular evolution into phylogeneitc analysis, and a new compilation of conserved polymerase chain reaction primers for animal mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics, 37, 545–579.

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 652–682.

Stein, J.P.E.F. (1874) Beitrag zur Kenntniss der Phryganeiden des Altvaters und einiger anderer. Stettiner Entomologische Zeitung, 35, 244–253.

Stocks, I.C. (2010) Comparative and functional morphology of wing coupling structures in Trichoptera: Annulipalpia. Journal of Morphology, 271, 152–168.

Strong, E.E. & Lipscomb, D. (1999) Character coding and inapplicable data. Cladistics, 15, 363–371.

Swofford, D.L. (2003) PAUP* Phylogenetic Analysis Using Parsimony (* and Other Methods), Version 4. Sinauer Associates, Sunderland, Massachusetts.

Swofford, D.L., Olsen, G., J., Waddell, P.J. & Hillis, D.M. (1996) Phylogenetic Inference. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.) Molecular Systematics, 2nd edition. Sinauer Associates, Sunderland, Massachusetts, pp. 407–514.

Sykora, J.L. (1999) Genus Mortoniella and its distribution in South America (Trichoptera, Glossosomatidae, Protoptilinae). In: Malicky, H. & Chantaramongkol, P. (Eds.) Proceedings of the 9th International Symposium on Trichoptera. Faculty of Science, Chiang Mai University, Chiang Mai, Thailand, pp. 377–387.

Tian, L. & Li, Y. (1986) Four new species of the caddisflies on Hengduan Mountain, China. Journal of Nanjing Agricultural University, 2, 50–54.

Tsuda, M. (1942) Zur Kenntnis den Koreanischen Trichopteren. Memoirs of the College of Science, Kyoto Imperial University, Series B, 17, 227–237.

Ulmer, G. (1906) Neuer beitrag zur kenntnis aussereuropäischer Trichopteren. Notes from the Leyden Museum, 28, 1–116.

Valverde, A.d.C. & Miserendino, M.L. (1998) Aportos al conocimiento do los estados preimaginales de Mastigoptila longicornuta (Trichoptera: Glossosomatidae). Revista de la Sociedad Entomológica Argentina, 57, 49–55.

Walker, F. (1852) Cataloque of the Specimens of Neuropterous Insects in the Collection of the British Museum, Part I: Phryganides-Perlides. British Museum, London, 192 pp.

Wallengren, H.D.J. (1891) Skandinaviens Neuroptera. Andra afdelningen. Svenska Akademien Handlingar, 24, 1–173.

Ward, J.B. & Pollard, S.D. (2002) Mating plugs and exaggerated genitalia: Possible consequences of sexual selection on two species of Trichoptera from New Caledonia. In: Mey, W. (Ed.), Nova Supplementa Entomologica (Proceedings of the 10th International Symposium on Trichoptera), 15, 249–252.

Wichard, W. (1989) Köcherfliegen des Dominikanischen Bernsteins. VII. Fossile Arten der Gattung Cubanoptila Sykora, 1973. Mitteilungen Münchener Entomologischen Gesellschaft, 79, 91–100.

Wichard, W. (1995) Köcherfliegen des Dominikanischen Bernsteins VIII. Cubanoptila grimaldii n. sp., eine weitere fossile Art der Glossosomatidae (Insecta, Trichoptera). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg, 78, 159–165.

Wichard, W. (2007) Overview and descriptions of caddisflies (Insecta, Trichoptera) in Dominican amber (Miocene). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie), 366, 1–51.

Wichard, W., Kraemer, M.M.S. & Luer, C. (2006) First caddisfly species from Mexican amber (Insecta: Trichoptera). Zootaxa, 1378, 37–48.

Wiens, J.J. (1998) Does adding characters with missing data increase or decrease phylogenetic accuracy? Systematic Biology, 47, 625–640.

Wiens, J.J. (2003a) Incomplete taxa, incomplete characters, and phylogenetic accuracy: Is there a missing data problem? Journal of Vertebrate Paleontology, 23, 297–310.

Wiens, J.J. (2003b) Missing data, incomplete data, and phylogenetic accuracy. Systematic Biology, 52, 528–538.

Wiens, J.J. (2005) Can incomplete taxa rescue phylogenetic analyses from long-branch attraction? Systematic Biology, 54, 731–742.

Wiens, J.J. (2006) Missing data and the design of phylogenetic analyses. Journal of Biomedical Informatics, 39, 34–42.

Wiens, J.J. (2009) Paleontology, genomics, and combined-data phylogenetics: Can molecular data improve phylogeny estimation for fossil taxa? Systematic Biology, 58, 87–99.

Wiens, J.J. & Reeder, T.W. (1995) Combining data sets with different numbers of taxa for phylogenetic analysis. Systematic Biology, 44, 548–558.

Wiggins, G.B. (1996) Larvae of the North American Caddisfly Genera (Trichoptera), 2nd Edition. University of Toronto Press, Toronto, 457 pp.

Wiggins, G.B. (2004) Caddisflies: The Underwater Architects. University of Toronto Press, Toronto, 292 pp.

Wilcox, T.P., Zwickl, T.A., Heath, T.A. & Hillis, D.M. (2002) Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25, 361–371.

Wiley, E.O. (1981) Phylogenetics: The Theory and Practice of Phylogenetic Systematics. John Wiley & Sons, Inc., New York, 439 pp.

Wilkinson, M. (1992) Ordered versus unordered characters. Cladistics, 8, 375–385.

Wilkinson, M. (1995) Coping with abundant missing entries in phylogenetic inference using parsimony. Systematic Biology, 44, 501–514.

Wilkinson, M. (2003) Missing entries and multiple trees: Instability, relationships, and support in parsimony analysis. Journal of Vertebrate Paleontology, 23, 311–323.[0311:meamti];2

Yang, Z. & Rannala, B. (2012) Molecular phylogenetics: principals and practice. Nature Reviews Genetics, 13, 303–314.

Yeates, D. (1992) Why remove autapomorphies? Cladistics, 8, 387–389.

Zander, R.H. (2004) Minimal values for reliability of bootstrap and jackknife proportions, decay index, and Bayesian posterior probability. PhyloInformatics, 2, 1–13.

Zhou, X., Kjer, K.M. & Morse, J.C. (2007) Associating larvae and adults of Chinese Hydropsychidae caddisflies (Insecta: Trichoptera) using DNA sequences. Journal of the North American Benthological Society, 26, 719–742.


  • There are currently no refbacks.

ISSN 1175-5326 (Print Edition) & ISSN 1175-5334 (Online Edition)
Published by Magnolia Press, Auckland, New Zealand
A founding journal of Biotaxa