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

Phylogeny and revision of the Neotropical genus Grumichella Müller (Trichoptera: Leptoceridae), including nine new species and a key

The systematics of the Neotropical caddisfly genus Grumichella Müller (Leptoceridae: Grumichellinae) are reviewed. Diagnoses, descriptions and illustrations are provided for four previously described species, G. aequiunguis Flint, 1983, G. flaveola (Ulmer, 1911), G. pulchella (Banks, 1910) and G. rostrata Thienemann, 1905, and nine new species: G. blahniki sp. nov. (Peru), G. boraceia sp. nov. (Brazil), G. cressae sp. nov. (Venezuela), G. jureia sp. nov. (Brazil), G. leccii sp. nov. (Brazil), G. muelleri sp. nov. (Brazil), G. paprockii sp. nov. (Brazil), G. parati sp. nov. (Brazil) and G. trujilloi sp. nov. (Venezuela). The monophyly of the genus is corroborated (16 synapomorphies) and the phylogenetic relationships of its included species, based on analysis of 66 adult, larval, and pupal characters, are inferred as (G. aequiunguis ((G. boraceiae (G. leccii, G. parati)) (G. rostrata ((G. flaveola, G. pulchella) (G. muelleri, G. paprockii)) (G. jureia (G. trujilloi (G. cressae, G. blahniki)))))). A taxonomic key to the males of the species is presented

Taxonomy and systematics: contributions to benthology and J-NABS

Systematics, or taxonomy, is the study of the diversity of life on Earth. Its goals are to discover and describe new biological diversity and to understand its evolutionary and biogeographic origins and relationships. Here we review the contributions to the field of systematics and taxonomy published over the last 25 y in J-NABS and its predecessor Freshwater Invertebrate Biology (FIB). We examined a total of 64 studies that we considered to be largely taxonomic in nature. We classified these studies into 2 major categories: morphological (e.g., descriptive taxonomy, taxonomic revisions) and molecular (e.g., deoxyribonucleic acid [DNA] barcoding, population genetics). We examined studies in 5-y increments for J-NABS. We also studied the period 1982 to 1985, during which FIB was published. On average, 12 taxonomic studies were published within each 5-y period. Molecular studies first appeared in 1986 and have slowly increased, reaching their greatest number within the last 5 y. Studies also were classified by their individual attributes. Morphological studies were, by far, the most common, but studies also included molecular data, biological information, distributional data, keys, and biogeographical analyses. Most studies included .1 of these attributes. Overall, the role of J-NABS in the development of benthic taxonomy has been minimal in terms of number of publications, but as part of the nexus of taxomonic literature, all contributions have been important to the discipline. We discuss these contributions and their impact on the following subject areas: taxonomy and revisionary systematics, phylogenetic and molecular systematics, taxonomic resources, taxonomic resolution, conservation and taxonomy, professional training, taxonomic certification, and graduate education. We also give an overview of new developments in the taxonomists’ toolbox. These developments include DNA barcoding, online taxonomic resources, digital identification keys, cybertaxonomy, and modern museum collections and resources

The Trichoptera barcode initiative: a strategy for generating a species-level Tree of Life

DNA barcoding was intended as a means to provide species-level identifications through associating DNA sequences from unknown specimens to those from curated reference specimens. Although barcodes were not designed for phylogenetics, they can be beneficial to the completion of the Tree of Life. The barcode database for Trichoptera is relatively comprehensive, with data from every family, approximately two-thirds of the genera, and one-third of the described species. Most Trichoptera, as with most of life’s species, have never been subjected to any formal phylogenetic analysis. Here, we present a phylogeny with over 16 000 unique haplotypes as a working hypothesis that can be updated as our estimates improve. We suggest a strategy of implementing constrained tree searches, which allow larger datasets to dictate the backbone phylogeny, while the barcode data fill out the tips of the tree. We also discuss how this phylogeny could be used to focus taxonomic attention on ambiguous species boundaries and hidden biodiversity. We suggest that systematists continue to differentiate between ‘Barcode Index Numbers’ (BINs) and ‘species’ that have been formally described. Each has utility, but they are not synonyms. We highlight examples of integrative taxonomy, using both barcodes and morphology for species description. This article is part of the themed issue ‘From DNA barcodes to biomes’

A new Ecuadorian species of the rare Neotropical caddisfly genus Amphoropsyche Holzenthal (Trichoptera, Leptoceridae)

Volume: 640Start Page: 59End Page: 6

Contulma boliviensis Holzenthal & Robertson, 2006, new species

Contulma boliviensis, new species Figs. 1 A–E Contulma boliviensis is a member of the Cranifer Group of Holzenthal & Flint (1995) based on the possession of an elongate dorsolateral process posteriorly on segment IX of the male genitalia. It stands isolated within the group as no other species has such a deeply excavated, C-shaped segment IX, but it shares with C. ecuadorensis Holzenthal and Flint and C. cranifer elongate, semimembranous sub-phallic processes. The phallus of this new species, with its flap-like, downturned, lateral flanges, is also unique. Length of forewing: 4.6 mm. Forewing (in alcohol) completely denuded, forewing membrane brown, with 4 light spots in membrane along costal margin, another near nygma; head, thorax, and appendages brown. Posterior foretibial spur very small, much shorter than anterior spur. Male genitalia. Segment IX in lateral view narrow, C-shaped, with anterior margin strongly rounded, produced anteriorly; posterior margin highly excavated medially, greatly produced ventrally, to form prominent, rounded, setose lobe; posterodorsally with pair of sclerotized, elongate, sinuate, dorsolateral processes, apices acute, turned medially; dorsally segment IX narrow mesally with short, shelflike extension; sternum IX in ventral view with small, posteromesal, shelf-like projection. Inferior appendage in lateral view short, digitate, sparsely setose apically. Processes of subphallic membranes present, digitate, setose, longer than inferior appendages and lateral to inferior appendages. Segment X elongate, membranous, with lightly sclerotized lateral flanges. Phallus relatively simple; phallobase sclerotized, tubular, elongate, straight; in dorsal view not broad posteriorly, laterally with flap-like, downturned flanges, subapicoventrally with paired membranous lobes; apex of phallus with membranous and semimembranous lobes as in Figs. 1 D, E; phallotremal sclerite not evident. Female. Unknown. Holotype male: BOLIVIA: Santa Cruz: Parque Nacional Amboró, 17 ° 50 ’ 15 ”S, 64 ° 23 ’ 29 ”W, 2030 m, flight intercept trap, AMB 21 FIT002, 17–20.x. 2001, Spector, Ledezma (UMSP 000086401) (UASC). Etymology. Named for the type country, in recognition of the first species in the genus Contulma to be described from Bolivia.Published as part of Holzenthal, Ralph W. & Robertson, Desiree R., 2006, Four new species of Contulma from South America (Trichoptera: Anomalopsychidae), pp. 49-59 in Zootaxa 1355 on pages 50-52, DOI: 10.5281/zenodo.17458

Contulma tripui Holzenthal & Robertson, 2006, new species

Contulma tripui, new species Figs. 4 A–E Contulma tripui is most similar to C. fluminensis based on the similarly complex morphology of the phallic apparatus, but it differs primarily in having the anterodorsal margin of segment IX prominently produced anteriorly and the lobe of the posterolateral margin of IX situated much more nearly medially than in C. fluminensis. Length of forewing: 4.5 mm. Forewing color dark brown, immaculate; head, thorax, and appendages brown. Posterior foretibial spur about 1 / 2 length of anterior spur. Male genitalia. Segment IX in lateral view broad, with anterodorsal margin prominently produced anteriorly, heavily setose ventrally; posterolateral margin produced medially to form broad, prominent, subtriangular, heavily setose lobe; posterodorsally with pair of lightly sclerotized, elongate, curved, ventrally directed, dorsomesal processes with rugose apices; dorsally segment IX highly reduced; sternum IX in ventral view with posteromesal, sclerotized, spatulate projection, surface longitudinally striate, apically cleft, anteromesally bearing about 4 flat, fused, toothlike setae. Inferior appendage in lateral view short, subtriangular, dorsal margin straight, bearing about 4 or 5 short setae, ventral margin heel-like; inferior appendages apparently fused to base of sternum IX projection, together forming highly complex structure as in Figs. 4 A, C, and inset. Processes of subphallic membranes absent (or not visible). Segment X highly reduced, entirely membranous. Phallus complex; phallobase sclerotized, tubular, curved; in dorsal view broad posteriorly, ventromedially with posterior membranous lobe, apicoventrally with lightly sclerotized lobe; apex of phallus with complex membranous lobes and sclerites as in Figs. 4 D, E; dorsally with paired, very large, highly convoluted, membranes; apicolaterally with paired, highly convoluted, membranous lobes, apically with large, broad, spatulate sclerites, bearing prominent keel-like longitudinal striae on their outer surfaces; phallotremal sclerite present, small, subspherical. Female. Unknown. Holotype male: BRAZIL: Minas Gerais: Estação Ecológica do Tripuí, Córrego Tripuí, 20 ° 23 ’ 22 ”S, 43 ° 32 ’ 32 ”W, el. 1070 m, 20.ix. 1998, Paprocki, Braga (UMSP 000046938) (MZUSP). Etymology. Named for the small stream in the ecological station of the same name, known for harboring an endemic species of onychophoran, Peripatus acacioi. Tripuí is the indigenous Tupi word for fast or quick water.Published as part of Holzenthal, Ralph W. & Robertson, Desiree R., 2006, Four new species of Contulma from South America (Trichoptera: Anomalopsychidae), pp. 49-59 in Zootaxa 1355 on pages 56-57, DOI: 10.5281/zenodo.17458

Tolhuaca brasiliensis Robertson & Holzenthal, 2005, new species

Tolhuaca brasiliensis, new species Figs. 4, 7, 8 This new species lacks the sclerotized conical endothecal spines found in T. cupulifera. Tergum X is quadrate in lateral view, whereas in T. cupulifera it is triangular. Tergum X is also shallowly excavate apicomesally, while it is deeply excavate in T. cupulifera. Tolhuaca brasiliensis is about half the size of T. cupulifera and also has narrower wings. The species is known only from southeastern Brazil. Adult. Length of forewing: male 2.7–2.8 mm (n= 2), female 2.7 mm (n= 1). Body, wings, and appendages nearly uniformly fuscous, tibia and tarsi yellowish brown. Forewing (Fig. 4 A) narrow, margins nearly parallel; with erect setae along Cu 2; Sc reaching anterior margin; fork I emerging beyond cord; fork II emerging at cord; forks III and IV shorter than their stems; A 3 gradually intersecting A 2; crossveins r, d, r­m, and mcu faintly visible, forming nearly straight line. Hind wing (Fig. 4 B) relatively narrow, tappering slightly past anastomosis, subacute apically; Sc and R 1 fused; fork II subequal to its stem; fork III shorter than its stem; crossvein r­m faintly visible, r and m­cu absent. Male genitalia (Fig. 7). Sternum VI (Fig. 7 C) with somewhat thickened, basally curved, mesal process, projecting downward. Abdominal segment IX (Fig. 7 A) well developed dorsally and laterally, but extremely reduced ventrally, forming very thin sclerotized strap; anterior margin broadly rounded; tergum IX, in dorsal view (Fig. 7 B), with posteromesal margin rounded; membranous connection between segments IX and X distinct. Tergum X (Figs. 7 A, 7 B) covered with fine microtrichia; in lateral view (Fig. 7 A), parallel sided, with broad lateral flange, setose apically and laterally, with rounded apicoventral processes, apex subtruncate; in dorsal view (Fig. 7 B), with lateral margins nearly straight and subparallel; apex slightly excavate medially. Phallobase (Figs. 7 A, 7 D) large and tubular, projecting apicodorsally, with basal extension and apparent suture and constriction medially; lightly sclerotized, but rugose ventrally and dorsally, with small, stout setae laterally and ventrally; endotheca (Figs. 7 A, 7 D) entirely membranous, greatly enlarged and convoluted when evaginated, with 3 convoluted tubular sclerites of varying lengths. Female genitalia (Fig. 8). Sternum V with oblique, slightly raised, sclerotized mesal linear ridge. Sternum VI process associated with slightly oblique apodeme. Segment VII normally developed. Abdominal segment VIII synscleritous, anterior margin membranous and receding ventrally, merging with intersegmental membrane, posterodorsal and posteroventral margins distinct and lightly sclerotized. Tergum IX lightly sclerotized. Tergum X, in lateral view, elongate, slightly bulbous; in dorsal and ventral views, bulbous, bearing cerci. Holotype male: BRAZIL: São Paulo: Parque Estadual de Campos do Jordão, 1 st order trib. to Rio Galharada, 22 ° 41 ' 40 "S, 045° 27 ' 47 "W, 1530 m, 14–16.ix. 2002, Blahnik, Prather, Huamantinco (UMSP 000087908) (MZUSP). Paratypes: BRAZIL: same data as holotype — 1 female (UMSP); same except Rio Galharada, 13–15.ix. 2002, Blahnik, Prather, Melo, Huamantinco — 1 male (UMSP). Distribution. Brazil. This species is known only from the type locality in the Serra da Mantiqueira. Etymology. Named for the type locality, in recognition of first species of the genus Tolhuaca to be discovered in Brazil.Published as part of Robertson, Desiree R. & Holzenthal, Ralph W., 2005, The Neotropical caddisfly genus To l h u a c a (Trichoptera: Glossosomatidae), pp. 53-68 in Zootaxa 1063 on pages 62-65, DOI: 10.5281/zenodo.17019

Tolhuaca

Position of Tolhuaca within the subfamily Protoptilinae Schmid (1964) originally placed Tolhuaca in the family Sericostomatidae, commenting on the similarity of the bifid tergum X of the male genitalia to that of Brachycentridae, at that time a subfamily within Sericostomatidae. Flint (1967) later transferred Tolhuaca to Protoptilinae upon discovery that the wing figures in the original description were mislabelled with those of Austrocentrus griseus Schmid 1964 (Helicophidae). Since that time, there has been nothing published regarding the placement of Tolhuaca in relation to the other protoptiline genera. When Schmid (1964) first described the genus he wrote, “Dans l’état actuel de nos connaissances, il est impossible d’assigner une position phylétique au genre Tolhuaca, dont la nervulation assez complète, surtout aux ailes antérieures, contraste avec l’extrême simplification et spécialisation des génitalia…” [“In the current state of our knowledge, it is impossible to assign a phyletic position to the genus Tolhuaca, whose rather complete venation, especially in the forewings, contrasts with the extreme simplification and specialization of the genitalia…” Translation from Schmid (1964)]. The male genitalia (Figs. 5 A, 7 A) have completely lost both the inferior and preanal appendages, and sternum IX has been reduced to nothing more than a thin, ventral strap. The phallic apparatus (Figs. 5 A, 5 D, 5 F, 7 A, 7 D) consists of a greatly enlarged, but simple tubular phallobase, and an eversible membranous endotheca. These apparently derived conditions of the male genitalia contrast with other features that imply a basal placement of Tolhuaca within Protoptilinae. Published works regarding protoptiline phylogeny are few and far between. Morse and Yang (1993) discussed some genera from the East Palaearctic and Oriental regions, and also provided a very useful table comparing wing venation among 15 recognized genera. Ross (1956) provided an early discussion of possible phylogenetic relationships among protoptiline genera and considered the genus Matrioptila Ross 1956 to be most primitive based on characteristics of the male genitalia and wing venation. In Matrioptila, segment IX is complete and ringlike, tergum X consists of a pair of simple lobes, and inferior appendages are present. Forewing veins Cu 1 and Cu 2 are often fused in Protoptilinae, but in Matrioptila, they are separate and distinct along their entire lengths and in the hind wing, Cu 1 is branched. These genitalic and venational characters are primitive according to Ross (1956). Kimmins (1964) described a new genus, Nepaloptila, from Nepal and placed it in Protoptilinae. The genus shares some characteristics of the male genitalia with Matrioptila, but Kimmins (1964) considered Nepaloptila to be more primitive, based on its retention of apical fork V in the forewing, which is usually absent in other Protoptilinae. The venation of Tolhuaca (Figs. 3, 4) is quite similar to that of Nepaloptila (Kimmins 1964), differing only slightly. In Tolhuaca, Sc is distinct from R 1 in the forewing, whereas in Nepaloptila, Sc and R 1 are fused near the wing margin. The 2 genera also differ in the position of crossveins in the forewing: those of Tolhuaca form a relatively straight transverse cord along the anastomosis whereas in Nepaloptila, they do not. In the hind wing, A 2 is present in Tolhuaca, but absent in Nepaloptila. Although Tolhuaca would seem to have more specialized male genitalia than Nepaloptila and Matrioptila, other characters suggest Tolhuaca is the more primitive genus. In Nepaloptila and Matrioptila, the foretibial spur has been lost, whereas in Tolhuaca, although reduced considerably, the spur is retained (Fig. 2). The genus Tolhuaca also has two small setal warts on the mesoscutellum (Fig. 1), which Ross (1956) considered to be primitive; these warts are absent in all other known protoptilines. The female genitalia (Figs. 6, 8) consist of an elongate oviscapt and 2 pairs of long, rod­like apodemes on segments VIII and IX. The presence of these apodemes is pleisiomorphic within Amphiesmenoptera (Kristensen 1984). The retention of the foretibial spur, presence of mesoscutellar setal warts, and the structure of the female genitalia suggest that Tolhuaca deserves a basal placement within the subfamily Protoptilinae.Published as part of Robertson, Desiree R. & Holzenthal, Ralph W., 2005, The Neotropical caddisfly genus To l h u a c a (Trichoptera: Glossosomatidae), pp. 53-68 in Zootaxa 1063 on pages 55-56, DOI: 10.5281/zenodo.17019