Systematics of the family Polycentropodidae (Inseecta:Trichoptera: Psychomyioidea) and taxonomic revisions of New World Polyplectropus Ulmer.

Abstract

The monophyly and phylogenetic relationships of subfamilies and genera traditionally classified in Polycentropodidae Ulmer, 1903, one of the most diverse families in the suborder Annulipalpia, with more than 700 species in 3 subfamilies, were tested. Particular emphasis was placed on testing the monophyly of the cosmopolitan genus Polyplectropus. Larval information is unknown for 46% of the taxa included in this study. To understand the effects of including characters with large sets of missing data, three alternative datasets [TOTAL (all available data for all taxa)= 49 ingroup taxa, 122 characters (including highly incomplete characters); LPA (larval, pupal, adult) = 20 ingroup taxa, 122 characters; ADULT (only adult characters) = 49 ingroup taxa, 86 adult characters] were analyzed under parsimony and Bayesian methods. The five outgroup taxa, representing all four extant families in the Psychomyioidea and the single family in the Hydropsychoidea, remained constant in all datasets. The TOTAL and ADULT datasets included all 20 currently recognized polycentropodid genera placed in 3 subfamilies, and the LPA and TOTAL datasets included characters interpreted from structures of the larvae, pupae, and adults. Results rejected the monophyly of Polycentropodidae, as currently defined; however, the monophyly of the three largest cosmopolitan genera, Polycentropus, Polyplectropus, and Nyctiophylax, could not be rejected nor confirmed. The monophyly of the following taxa was strongly supported in all analyses: Cernotina, Cyrnellus, Kambaitipsyche, Neureclipsis, Paranyctiophylax, New World Polyplectropus sensu stricto, Placocentropus, Neotropical Nyctiophylax, and in the outgroup, Psychomyia + Xiphocentron; while monophyly was strongly supported in some, but not all analyses for the following taxa: Cyrnus, Antillopsyche, Pseudoneureclipsis, Polycentropus sensu stricto, Pseudoneureclipsinae, New Zealand Polyplectropus, Polycentropodinae, Cyrnodes scotti + Pahamunaya jihmita. The implementation of two different analytical methods revealed some areas of conflict which would not have been detected under a single method of analysis. Contradictory results among the datasets were primarily due to either inclusion or exclusion of key sets of characters (i.e., immature characters); and second, missing data negatively affected

phylogenetic reconstruction when proportions of characters with missing data were high and characters without missing data were unable to provide adequate phylogenetic signal due to high variation in rates of evolution among characters. Therefore, a combination of few overall characters that have high variation in rates of change, plus an abundance of missing data may be problematic and may lead to poorly resolved trees, thus decreased accuracy. This study also emphasized the importance in phylogenetic reconstruction of including data from all available sources. Several taxonomic changes were necessary in order for classification to properly reflect phylogeny. Three new genera, all from the Neotropical region, will be described in future publications. The redefinition of Paranyctiophylax as a valid genus in Polycentropodinae was confirmed. Additionally, the recommendation was made that North American Polycentropus species previously belonging in Plectrocnemia or Holocentropus be recognized as such (either Plectrocnemia or Holocentropus depending on original designation) and not as belonging in Polycentropus. Furthermore, species described in Polycentropus post-1944 in North America are transferred to either Holocentropus or Plectrocnemia to reflect previously hypothesized sister relationships. The following new or reinstated combinations were proposed: Plectrocnemia albipuncta Banks, 1930 combinatio revivisco; Plectrocnemia aureola Banks, 1930 comb. rev.; Plectrocnemia cinerea (Hagen), 1861 comb. rev.; Plectrocnemia clinei Milne, 1936 comb. rev.; Plectrocnemia crassicornis (Walker), 1852 comb. rev.; Plectrocnemia jenula (Denning), in Denning & Sykora, 1966 combinatio nova; Plectrocnemia icula (Ross), 1941 comb. nov.; Plectrocnemia nascotia (Ross), 1941 comb. nov.; Plectrocnemia remota (Banks), 1911 comb. rev.; Plectrocnemia sabulosa (Leonard & Leonard), 1949 comb. nov.; Plectrocnemia smithae (Denning), 1949 comb. nov.; Plectrocnemia vigilatrix Navás, 1933 comb. rev.; Plectrocnemia weedi (Blickle & Morse), 1955 comb. nov.; Holocentropus chellus (Denning), 1964 comb. nov.; Holocentropus flavus Banks, 1908 comb. rev.; Holocentropus glacialis Ross, 1938 comb. rev.; Holocentropus grellus Milne, 1936 comb. rev.; Holocentropus interruptus Banks, 1914 comb. rev.; Holocentropus melanae Ross, 1938 comb. rev.; Holocentropus milaca (Etnier), 1968 comb. nov.; Holocentropus picicornis (Stephens), 1836 comb. rev. Additional taxonomic changes proposed based on current findings were: 1) the elevation

of Pseudoneureclipsinae to family status: Pseudoneureclipsidae Ulmer status novus; and 2) the resurrection of Placocentropus Schmid, nomen revivisco, to include the following species: Placocentropus aspinosus (Schmid), 1964 comb. nov.; Placocentropus chilensis (Yamamoto), 1966 comb. nov.; Placocentropus obtusus Schmid, 1955 comb. rev.; Placocentropus quadriappendiculatus (Schmid), 1964 comb. nov.; Placocentropus quadrispinosus (Schmid), 1964 comb. nov.; Placocentropus tuberculatus (Flint), 1983 comb. nov.; Placocentropus valdiviensis (Flint), 1983 comb. nov. A phylogeny of New World Polyplectropus species was inferred. Characters were interpreted from structures of the male and female genitalia as well as the fore- and hind wings. Parsimony and Bayesian phylogenetic analyses of 89 ingroup taxa (97% of the known New World diversity in the genus), two outgroup taxa, and 59 morphological characters were performed. Results of the parsimony and Bayesian analyses were similar, although the Bayesian tree was less resolved. Monophyly of the panamensis and charlesi Groups, as currently defined, was rejected. A total of 10 lineages, with varying amounts of support, were recognized. These groups are the alienus Group (2 species), annulicornis Group (11 species, 8 new), bredini Group (19, 7), charlesi Group (3), fuscatus Group (3, 2), guyanae Group (2, 2), manuensis Group (3, 3), narifer Group (5, 3), santiago Group (25, 6), and thilus Group (15, 7). Four species remain unassigned to any species-group: P. beccus, P. beutelspacheri, P. kanukarum, and P. nayaritensis. The distribution of the genus is mostly restricted to the Mexican and Brazilian subregions of the Neotropics. Most of the species and species-groups are regional endemics. The taxonomy of New World species of Polyplectropus Ulmer, 1905 was revised to include detailed male and female diagnoses, descriptions, illustrations, distribution records, and keys to males of all species and species-groups. A key to genera of New World Polycentropodidae, including a redescription of Polyplectropus, was provided. The homology of the male genitalia of species in the genus was discussed, as well as reassessment and diagnoses of 10 species groups, 6 newly established. A total of 92 species were treated, 39 described as new: Polyplectropus adamsae, sp. nov. (Peru), P. alatespinus, sp. nov. (Brazil), P. amazonicus, sp. nov. (Brazil), P. andinensis, sp. nov. (Argentina, Bolivia), P. blahniki, sp. nov. (Venezuela), P. bolivianus, sp. nov. (Bolivia), P. brasilensis, sp. nov. (Brazil), P. vii brborichorum, sp. nov. (Ecuador), P. cressae, sp. nov. (Venezuela), P. colombianus, sp. nov. (Colombia), P. corniculatus, sp. nov. (Peru), P. cuzcoensis, sp. nov. (Peru), P. ecuadoriensis, sp. nov. (Ecuador), P. flintorum, sp. nov. (Venezuela), P. gaesum, sp. nov. (Brazil), P. guyanae, sp. nov. (Guyana, Venezuela), P. holzenthali, sp. nov. (Brazil), P. hystricosus, sp. nov. (Brazil), P. insularis, sp. nov. (Panama), P. julitoi, sp. nov. (Brazil), P. kanukarum, sp. nov. (Guyana), P. maculatus, sp. nov. (Venezuela), P. manuensis, sp. nov. (Peru), P. matatlanticus, sp. nov. (Brazil), P. minensium, sp. nov. (Brazil), P. novafriburgensis, sp. nov. (Brazil), P. peruvianus, sp. nov. (Peru), P. petrae, sp. nov. (Brazil), P. pratherae, sp. nov. (Brazil), P. puyoensis, sp. nov. (Ecuador), P. robertsonae, sp. nov. (Bolivia), P. rodmani, sp. nov. (Brazil), P. rondoniensis, sp. nov. (Brazil), P. tragularius, sp. nov. (Brazil), P. tripunctatum, sp. nov. (Peru), P. venezolanus, sp. nov. (Venezuela), P. woldai, sp. nov. (Panama), P. zamoranoensis, sp. nov. (Honduras), and P. zuliae, sp. nov. (Venezuela). Polyplectropus buchwaldi is designated as a nomen dubium.