Systematic studies of the caddisfly subfamily Protoptilinae (Trichoptera: Glossosomatidae)

Abstract

Protoptilinae Ross 1956, is the most diverse subfamily [268 species (including 5 fossil spp.) and 17 genera] belonging to the saddle- or tortoise-case making family Glossosomatidae. The subfamily has a disjunct distribution: 4 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 the 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 (COI) data was unavailable for 55 of the 80 taxa included in this study. To test the effects of including a large set of highly incomplete taxa, 5 different datasets were analyzed using both parsimony and Bayesian methods: TOTAL COMBO (80 taxa, morphology and COI); TOTAL MORPH (80 taxa, morphology); SUBSET COMBO (25 taxa, morphology and COI); SUBSET MORPH (25 taxa, morphology); and SUBSET COI (25 taxa, COI). 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, Culoptila, Itauara, Mastigoptila, Mortoniella, Protoptila, and Tolhuaca. Monophyly for the genera Campsiophora, Cariboptila, Cubanoptila, Nepaloptila, Padunia, and Poeciloptila was not supported. Several taxonomic changes were necessary for classification to reflect phylogeny. Accordingly, I proposed that Matrioptila (Ross 1938), Poeciloptila Schmid 1991, Temburongpsyche Malicky 1992, and Nepaloptila Kimmins 1964 all become new junior synonyms of Padunia Martynov 1910. Additionally, I proposed to that the endemic Caribbean genera Cariboptila Flint 1964 and Cubanoptila Sykora 1973 become new junior synonyms of Campsiophora Flint 1964. A key to the world genera of Protoptilinae incorporating these taxonomic changes was provided. Using a fossil-calibrated relaxed molecular clock analysis (based on mitochondrial DNA and ribosomal RNA), independent geologic evidence, climatology, and Bayesian dispersal-vicariance analysis (DIVA), I constructed a biogeographic scenario for Protoptilinae. Divergence time estimates indicated that Protoptilinae arose at the Paleocene-Eocene Boundary, well after the initial Mesozoic break-up of Gondwanaland. Most major protoptiline lineages diverged during the Eocene and most modern genera arose during the Oligocene. DIVA analyses suggested Tolhuaca to be a relict of a more widespread protoptiline ancestor whose distribution encompassed Central and South America. The timing of many lineage divergences were correlated to geologic events that would have facilitated faunal movement between land-masses. Island corridors between South and Central America allowed migration northward. The GAARlandia landspan (Greater Antilles + Aves Ridge) provided a route for protoptiline range expansion from northern South America to the Greater Antilles and its subsequent re-submergence led to the divergence of Campsiophora. The presence of Padunia in Asia resulted from migration across Beringia. Pleistocene glaciation may have caused the extinction of northern populations, resulting in Padunia’s present disjunct distribution. Nearly a third of all speciation events appear to be the result of dispersal, range expansion, or subsequent re-colonization. Species-level revisions of Canoptila, Itauara, and Tolhuaca were also presented, including generic diagnoses, illustrations, and descriptions of males. Females of Canoptila and Tolhuaca were described for the first time. Additionally, the first known record of Protoptila in Bolivia was reported and a review of the occurrence of scales and androconia in Trichoptera was provided. A total of 28 species were treated, 22 vii distribution. Nearly a third of all speciation events appear to be the result of dispersal, range expansion, or subsequent re-colonization. Species-level revisions of Canoptila, Itauara, and Tolhuaca were also presented, including generic diagnoses, illustrations, and descriptions of males. Females of Canoptila and Tolhuaca were described for the first time. Additionally, the first known record of Protoptila in Bolivia was reported and a review of the occurrence of scales and androconia in Trichoptera was provided. A total of 28 species were treated, 22 described as new: Canoptila williami (Brazil), new species, Itauara alexanderi, new species (Brazil), I. bidentata, new species (Guyana), I. blahniki, new species (Brazil), I. charlotta, new species (Brazil), I. emilia, new species (Brazil), I. flinti, new species described as new: Canoptila williami (Brazil), new species, Itauara alexanderi, new species (Brazil), I. bidentata, new species (Guyana), I. blahniki, new species (Brazil), I. charlotta, new species (Brazil), I. emilia, new species (Brazil), I. flinti, new speciesdescribed as new: Canoptila williami (Brazil), new species, Itauara alexanderi, new species (Brazil), I. bidentata, new species (Guyana), I. blahniki, new species (Brazil), I. charlotta, new species (Brazil), I. emilia, new species (Brazil), I. flinti, new species (Brazil), I. guianensis, new species (Guyana), I. jamesii, new species (Brazil), I. julia, new species (Brazil), I. lucinda, new species (Brazil), I. ovis, new species (Guyana, Venezuela), I. peruensis, new species (Peru), I. rodmani, new species (Brazil), I. simplex, new species (Brazil), I. spiralis, new species (Guyana), I. stella, new species (Brazil), I. tusci, new species (Brazil), I. unidentata, new species (Guyana), Protoptila diablita, new species (Bolivia), P. julieta, new species (Bolivia), and Tolhuaca brasiliensis, new species (Brazil).