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Issue Information Systematic Entomology (2017), 42, 240–266 DOI: 10.1111/syen.12209 Molecular phylogeny of Sericostomatoidea (Trichoptera) with the establishment of three new families KJELL ARNE JOHANSON1, TOBIAS MALM1 andMARIANNE ESPELAND2 1Department of Zoology, Swedish Museum of Natural History, Stockholm, Sweden and 2Arthropoda Department, Zoological Research Museum Alexander Koenig, Bonn, Germany Abstract. We inferred the phylogenetic relationships among 58 genera of Sericostom- atoidea, representing all previously accepted families as well as genera that were not placed in established families. The analyses were based on five fragments of the protein coding genes carbamoylphosphate synthetase (CPSase of CAD), isocitrate dehydroge- nase (IDH), Elongation factor 1a (EF-1a), RNA polymerase II (POL II) and cytochrome oxidase I (COI). The data set was analysed using Bayesian methods with a mixed model, raxml, and parsimony. The various methods generated slightly different results regarding relationships among families, but the shared results comprise support for: (i) a monophyletic Sericostomatoidea; (ii) a paraphyletic Parasericostoma due to inclusion of Myotrichia murina, leading to synonymization of Myotrichia with Parasericostoma; (iii) a polyphyletic Sericostomatidae, which is divided into two families, Sericostom- atidae sensu stricto and Parasericostomatidae fam.n.; (iv) a polyphyletic Helicophidae which is divided into Helicophidae sensu stricto and Heloccabucidae fam.n.; (v) hypoth- esized phylogenetic placement of the former incerta sedis genera Ngoya, Seselpsyche and Karomana; (vi) a paraphyletic Costora (Conoesucidae) that should be divided into several genera after more careful examination of morphological data; (vii) reinstatement of Gyrocarisa as a valid genus within Petrothrincidae. A third family, Ceylanopsychi- dae fam.n., is established based on morphological characters alone. A hypothesis of the relationship among 14 of the 15 families in the superfamily is presented. A key to the families is presented based on adults (males). Taxonomic history, diagnosis, habitat preference and distribution data for all sericostomatoid families are presented. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid: zoobank.org:pub:CF6A6B9F-6A72-4265-BD09-3A710DFCD7B1. Introduction just above 600 described extant species in 12 established families (Morse, 2016). The number of species is unevenly The first described members of the superfamily Sericostoma- distributed among the 68 recognized genera, and the largest toidea, Beraeodes minutus (Linnaeus) and Notidobia ciliaris genus, Helicopsyche von Siebold, comprises 273 species (45% (Linnaeus), were described by Linnaeus (1761), both originally of the species in the superfamily), and 50% of the genera in the classified together with all other Trichoptera species in the genus superfamily have one or two species only, indicating variable Phryganea (Fig. 1). Presently, the Sericostomatoidea contain evolutionary success in diversification or a tendency to use Helicopsyche as a kind of waste basket. The superfamily has a cosmopolitan distribution, but individual families have restricted Correspondence: Kjell Arne Johanson, Department of Zoology, distributions. Five genera – Ceylanopsyche Fischer, Karomana Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Schmid, Mpuga Schmid, Ngoya Schmid and Seselpsyche Sweden. E-mail: [email protected] 240 © 2016 The Authors. Systematic Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society. This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. Phylogeny of Sericostomatoidea 241 Fig. 1. Picture of species in Sericostomatoidea. (A) Sericostoma personatum (Spence) male, Norway, representing the type genus Sericostoma Latreille of the family Sericostomatidae. (B) Beraea pullata (Curtis) female, Sweden, representing the type genus Beraea Stephens of the family Beraeidae. Malicky – are classified as sericostomatoids but with uncertain among families based on analysis of morphological data using family affinities (Morse, 2016). Several attempts have been parsimony was that by Weaver (1983), who also was the first to made to outline the relationship among the sericostomatoid group eight of the families of Ross (1967) ‘leptocerid branch’ families and genera, starting with Ulmer more than 100 years into the superfamily Sericostomatoidea. He listed two synapo- ago (Ulmer, 1912), who hypothesized that the three families, morphies for the superfamily: loss of the abdominal tergite IX Sericostomatidae, Beraeidae and Helicopsychidae, presently in the larvae, and loss of the mid-leg preapical spurs in adults. classified as sericostomatoids, were not believed to be most He stated (pp. 51 and 244) that the Anomalopsychidae forms the closely related to each other (Fig. 2A), but no arguments were sister group to the rest of the families in the superfamily based given for that interpretation. Ross (1956) offered a first furcated on the presence of ocelli in the adults, which were lost in the phylogenetic hypothesis of Trichoptera families, included the sister group comprising the other seven families (Fig. 2E). The same three sericostomatoid taxa as Ulmer, and retained them as family Antipodoeciidae was included in the ‘leptocerid branch’ polyphyletic (Fig. 2B). In a similar analysis a decade later, Ross as a distinct family by Ross (1967) but was synonymized with (1967) included seven sericostomatoid families and grouped Beraeidae by Ross (1978), and not included in the analysis by them in the ‘leptocerid branch’ (Fig. 2C). The Sericostomatidae Weaver (1983) as a distinct family. As a first effort to classify was revised by Ross (1978), who reduced the number of fami- the Hydrosalpingidae, Petrothrincidae and Barbarochthonidae lies in Sericostomatoidea to five, but the superfamily remained in relation to the nine other families in the superfamily, deMoor polyphyletic (Fig. 2D). The first hypothesis of the relationship (1993) performed a parsimony analysis of 59 morphological © 2016 The Authors. Systematic Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society. 42, 240–266 242 K. A. Johanson et al. Fig. 2. Hypotheses of evolutionary relationships among families in Sericostomatoidea, after: (A) Ulmer (1912); (B) Ross (1956); (C) Ross (1967); (D) Ross (1978); (E) Weaver (1983); (F) Johanson (1998) derived from deMoor (1993); (G) Frania & Wiggins (1997); (H) Kjer et al. (2001); (I) Neboiss (2002); (J) Holzenthal et al. (2007a); (K) Malm et al. (2013). In Fig. 2A–D taxon names in bold belong to the Sericostomatoidea; taxa in plain are classified outside the Sericostomatidea. © 2016 The Authors. Systematic Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society. 42, 240–266 Phylogeny of Sericostomatoidea 243 characters, and included all 12 families. In his two most par- Material and methods simonious trees, three monophyletic groups were detected, Petrothrincidae + Barbarochthonidae + Hydrosalpingidae, Taxon sampling Calocidae + Chathamiidae, and Conoesucidae + Helicophidae, but a reanalysis of the data matrix (Johanson, 1998) forced most The data comprise representatives from all sericostomatoid of the branches into collapsed polytomies (Fig. 2F). Frania families and the majority of genera recognized in the group. & Wiggins (1997) performed a comprehensive analysis of When possible, two or more species from each genus were the relationship among Trichoptera families, and included six included and for some genera two individuals of the same sericostomatoids. They found (Fig. 2G) reproduced Sericos- species were represented. We extracted and sequenced repre- tomatoidea to be monophyletic and the family Sericostomatidae sentatives of 58 of the 70 Sericostomatoidea genera described was the sister group to remaining five families, of which the at present (Table 1). Twelve genera (Notoernodes Andersen previously assumed most basal taxon, Anomalopsychidae, & Kjærandsen, Alloecentrellodes Flint, Microthremma Schmid, formed the sister group to Helicopsychidae. Molecular data Pseudosericostoma Schmid, Aclosma Morse, Asahaya Schmid, Aselas Barnard, Mpuga, Ceylanopsyche, Latarima Shackleton, was first analysed in the context of Trichoptera evolution Webb, Lawler & Suter and Cerasma McLachlan, Chiloecia by Kjer et al. (2001), who applied a combination of mor- Navás) were excluded from our analyses due to difficulties phological data and DNA sequences from both coding and of obtaining high-quality DNA. Outgroup taxa were selected noncoding genes. Their support for monophyly of Sericos- across the suborder Integripalpia based on the evolutionary tomatoidea was obtained as very low, and almost absent for hypothesis presented by Malm et al. (2013). The outgroup the internal relationships among the eight families included comprises the genera Phryganopsyche Wiggins (Phryganopsy- of the analysis, in practice resulting in collapse of branches chidae), Phryganea Linnaeus (Phryganeidae), Taskiropsyche within the superfamily (Fig. 2H). Attempting to correctly Neboiss and Kokiria McFarlane (Kokiriidae), Anisocentropus place the genus Heloccabus Neboiss in the sericostomatoid McLachlan and Phylloicus Müller (Calamoceratidae), Molanna tree,
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