Systematics of the Neotropical caddisfly genus Notidobiella Schmid (Trichoptera, Sericostomatidae), with the description of 3 new species

Abstract Three new species of Notidobiella Schmid (Insecta: Trichoptera) are described from South America: Notidobiella amazoniana sp. n. (Brazil), Notidobiella brasiliana sp. n. (Brazil), and Notidobiella ecuadorensis sp. n. (Ecuador). In addition, the 3 previously described species in the genus, Notidobiella chacayana Schmid, Notidobiella inermis Flint, and Notidobiella parallelipipeda Schmid, all endemic to southern Chile, are redescribed and illustrated, including the females of each species for the first time, and a key to males of the species in the genus is provided. The occurrence of Notidobiella in Brazil and Ecuador represents a significant extension of the range of the genus beyond southern Chile where it previously was thought to be endemic. The biogeography of Sericostomatidae and other austral South American Trichoptera is reviewed. The presence of the family in South America may not be part of a “transantarctic” exchange, but instead may represent an earlier occurence in the region. The distribution of Notidobiella in tropical South America likely represents recent dispersal from southern South America to the north.


Introduction
Th e caddisfl y family Sericostomatidae occurs in all biogeographic regions, except the Australasian, but its species diversity is very unevenly distributed across these regions (Morse 2010). Nineteen genera and 100 species have been described world wide, with half of the species occurring in the western Palearctic (50 species in 5 genera: Cerasma McLachlan, Notidobia Stephens, Oecismus McLachlan, Schizopelex McLachlan, Sericostoma Latreille) (Holzenthal et al. 2007b). South Africa harbors 12 species in 5 endemic genera (Aclosma Morse, Aselas Barnard, Cheimacheramus Barnard, Petroplax Barnard, Rhoizema Barnard), but no species are found in tropical Africa. Th e genus Agarodes Banks contains 12 species confi ned largely to the southeastern United States, where a second monotypic genus, Fattigia Ross, also occurs. Th ere is a single species known from India, Asahaya asambaddha Schmid, and the 6 species in the genus Gumaga Tsuda occur in Mexico (Baja California) and the western United States (3 species) and in southern and eastern Asia (3 species). In the Neotropics, the family is represented in the Chilean subregion by 4 genera, Chiloecia Navás (1 species, nomen dubium), Myotrichia Schmid (1), Notidobiella Schmid (3), and Parasericostoma Schmid (10) (Flint et al. 1999b). Until now, only a single sericostomatid species, Grumicha grumicha (Vallot), was known from the Brazilian subregion (Flint et al. 1999a).
In this paper, we describe 3 new species of Notidobiella, 1 from the Amazon basin, Brazil, 1 from southeastern Brazil, and 1 from Ecuador, thus extending the range of this genus well beyond its Chilean representation. In addition, we provide illustrations and diagnoses of males and females (the latter for the fi rst time) of the 3 previously described species of Notidobiella, N. chacayana Schmid, N. inermis Flint, and N. parallelipipeda Schmid, and a key to males of species in the genus. Th e Neotropical species of Sericostomatidae, including those in the genus Notidobiella, appear to be members of a southern Gondwana fauna (de Moor and Ivanov 2008).

Material and methods
Techniques and procedures used in the preparation and examination of specimens are those outlined by Blahnik and Holzenthal (2004) and Blahnik et al. (2007). Terminology for genitalia and wing venation follows that presented by Holzenthal et al. (2007b) and morphological structures are labeled in Figs 3-5. Th e species are presented in alphabetical order, except for the type species of the genus, N. parallelipipeda, which is presented fi rst, after the generic diagnosis. Material examined and types are deposited in the collections of the University of Minnesota Insect Collection, St. Paul, Minnesota, USA (UMSP), the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP), the Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil (INPA), and the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (NMNH). UMSP barcode accession label numbers for holotypes are included in the list of material examined, but not for paratypes.
Diagnosis: (modifi ed from Schmid 1955; characters pertain to the male of the type species, except where noted): Head short, but broad, with large, projecting eyes, with conspicuous setae between ommatidea (Fig. 1); in most species interocular distance equal to or slightly less than diameter of eye, when viewed frontally (eyes of females smaller, interocular distance about 2× diameter of eye). According to Schmid (1955) bordering the eyes medially is an elongate, convex cephalic tubercle, but this structure was not evident in the material of the type species examined by us. Head dorsally with prominent, elongate occipital and retroccipital warts; ocellar, antennal, frontal, anterolateral, and hypomedial setal warts absent (terminology of Ivanov 1990) (although females with small anterolateral setal wart). Antennal scape much shorter than head and about as wide as long, with oval setal warts on both its dorsal and ventral surfaces; scapes almost touch medially. Maxillary palp very short, composed of enlarged, heavily setose basal article held against the face and sclerotized on its lateral side only, medial side membranous, with small, oval, sclerotized, setose apical article ( Fig. 2) Flint (1967) described the immature stages of N. chacayana. Larvae construct slightly tapered and curved cases of small mineral fragments embedded in silk and occur on the bottoms of small streams. Th e 3 previously known species, N. chacayana Schmid, 1957, N. inermis Flint, 1983, and N. parallelipipeda Schmid, 1955, are endemic to southern Chile. Description. Of the species in the genus with broad, spatulate inferior appendages (all species except N. amazoniana), the type species is the most distinctive based on the parallel-sided inferior appendages with their prominent, mesally directed, mesal processes.
Material  Description. Th is is the smallest species in the genus. Its wings are short and broad with venation typical for the genus except forewing crossveins r and s are absent, leaving the discoidal cell open (Fig. 7). Its genitalia are the most atypical in the genus in that the inferior appendages are not broadly spatulate, but sinuous in shape and uniform in width. Schmid. 4 Male genitalia A segments IX, X, inferior appendages, lateral B segments IX, X, dorsal C segment IX, inferior appendages ventral D inferior appendage, dorsal E phallus, lateral F phallus, ventral G sternum VII posteromesal process, ventral. 5. Notidobiella parallelipipeda Schmid. Female genitalia, segments IX, X, dorsal.

Figures 4-5. Notidobiella parallelipipeda
Tergum IX bears a short triangular, posteromesal process, rather than short, paired processes, as found in the other species. Otherwise, the genitalia are typical for the genus.
Etymology: Named for the state and region where the specimens were collected, which represents a signifi cant northern extension of the range of the genus. Description. Th is new species is most similar to N. chacayana in the overall shape and structure of the inferior appendages. Both species possess an elongate mesal process on the ventromesal margin of the inferior appendage. In N. ecuadorensis sp. n., the ventromesal process is also present, but is shorter and broader in ventral view; in the other 2 Chilean species, N. inermis and N. parallelipipeda, the ventromesal processes are either very reduced (N. inermis) or long (N. parallelipipeda), but not nearly as long as in N. brasiliana sp. n. Setting N. brasiliana sp. n., apart from all of its congeners is the pair of elongate posteromesal processes on sternum IX; in all other species these processes are much shorter and broader. Furthermore, forewing crossveins r and s are absent, leaving the discoidal cell open (Fig. 10).

Notidobiella brasiliana
Adult. Forewing length 7.0 mm male (n=1); 7.9-8.2 mm female (n=4). Color medium to dark brown, palps and legs light brown; forewings dark brown with scattered golden hairs, pale golden spot on anal margin at about midlength. Sternum VII of male with broad, fi ngernail-like, posteromesal process.
Female genitalia (Fig. 12). Tergum VIII quadrate; pleural membranes extensive, highly folded; sternum VIII broad, anterior margin with apodemal ridge, extending dorsolaterally; posterolateral corners rounded, heavily setose, especially posteriorly. Tergum IX with heavily setose, posterolateral lobes, rounded to subtriangular in dorsal view; with lateral, microsetose, elevated ridge; sternum IX highly membranous, the membranes with parallel pleats or folds; dorsally tergum  Description. Th e combination of broadly spatulate inferior appendage, thumb-like mesal process on the ventromesal margin of the inferior appendage, and short posteromesal processes on sternum IX separate this species from its congeners. Th e wing venation (Fig. 14) is similar to that of the type species.
Male genitalia (Fig. 13). Segment IX with anterior margin broadly produced midlaterally; tergum IX narrow, elevated, mound-like; sternum IX with pair of short, triangular, posteromesal processes, bearing very long apical setae. Tergum X simple, triangular in lateral view, with slight dorsomesal excavation, setose apically. Preanal appendage short, ovate, setose. Inferior appendage prominent, heavily setose, very broadly spatulate, narrow basally, with short, thumb-like mesal process on ventromesal margin. Phallic apparatus simple, tubular, relatively straight from base to apex; endophalic membranes prominent, with paired apical membranous lobes; elongate, lightly sclerotized band internally (perhaps the phallotremal sclerite).  Etymology: Named for Ecuador, the country of the holotype, which represents a signifi cant northern extension of the range of the genus. of the inferior appendages, as illustrated, and in the possession of a narrow posteromesal process on sternum VII. Adult. Forewing length 6.0-6.5 mm male (n=2); 8.0 mm female (n=2). Color brown, palps and legs stramineous; forewings brown, with scattered golden setae. Sternum VII of male with narrow, fi ngernail-like, posteromesal process.

Biogeographic considerations
As defi ned most recently by de Moor and Ivanov (2008), the Trichoptera fauna of southern Chile and adjacent patagonian Argentina exhibits a strong biogeographical affi nity to Australia, New Zealand, and other southern Pacifi c islands (e.g., New Caledonia). Th is "Temperate Gondwana" (de Moor and Ivanov 2008) or "transantarctic" pattern (e.g., Brundin 1966) also includes the temperate, southernmost part of Africa and Madagascar. In the Neotropical Trichoptera, the affi nity to the southern African fauna is very weak, perhaps exhibited only at the family level within Sericostomatoidea (de Moor and Ivanov 2008) and potentially among genera within Sericostomatidae (although relationships among genera within this family are yet to be inferred).
On the other hand, the biogeographical affi nity between southern South America and Australasia is strong. Within South America, nearly all species in Chile and adjacent Argentina are endemic, prompting Flint (1976) to divide the Neotropics into 2 distinct subregions, the Chilean (southern Chile/Argentina) and the Brazilian (the rest of the Neotropics as defi ned by Wallace 1876); these regions are equivalent to the "Patagonian" and "Neotropical" (sensu stricto) Trichoptera regions of de Moor and Ivanov (2008). Four Trichoptera families are representative of a temperate Gondwanan pattern: Helicophidae, Kokiriidae, Philorheithridae, and Tasimiidae, each family with genera endemic to Australia/New Zealand/New Caledonia, southern South America, or Madagascar (no genera are shared) (Neboiss 1986, Flint et al. 1999b, Holzenthal et al. 2007b, Weaver et al. 2008 (Table 1). Sukatcheva and Jarzembowski (2001) questionably placed a fossil (fragment of a forewing) from the early Cretaceous of southern England in the Helicophidae.
Th e presence of Helicophidae and Hydrobiosidae in Eocene Baltic amber (Botosaneanu and Wichard 1983) and of fossil Plectrotarsidae (extant taxa endemic to Australasia) and a putative helicophid from late Cretaceous deposits in England (Sukatsheva and Jarzembowski 2001) suggests, as hypothesized by de Moor and Ivanov (2008), that certain southern temperate Trichoptera may be relicts of a more widespread fauna which included now extinct (but still extant in Sericostomatidae) north temperate elements. Two species of Triplectides in Baltic amber (Ulmer 1912) suggests the same scenario for this southern Gondwanan genus. In addition, the putative triplectidine larva from South Africa (de Moor 1997) and the recent discovery of Philorheithridae in Madagascar (Weaver et al. 2008) suggest at least a Gondwanan origin for these taxa (category 3 of Amorin et al. 2009) and, by inference, other endemic austral Trichoptera.
Evidence suggests that the contemporary distribution of the Patagonian and Australasian temperate Gondwanan Trichoptera fauna refl ects a past dispersal corridor between Australia and southern South America via Antarctica (Sanmartín and Ronquist 2004). Th is "transantarctic exchange" pattern also has been demonstrated for other insects, including aquatic taxa (Cranston and Edward 1999, Amorin et al. 2009, Daugeron et al. 2009). However, it may be that the current distribution of other southern Gondwana caddisfl ies, such as Sericostomatidae, refl ects an older dispersal prior to the breakup of southern Gondwana, now represented by relict Southern Hemisphere distributions. Th e southeast Brazilian sericostomatid Grumicha grumicha might represent evidence to support the relict hypothesis.
Th e now widespread occurrence of Notidobiella in temperate southern Chile and tropical South America (Ecuador, southeast Brazil, Amazonian Brazil) suggests a more recent dispersal of the genus to northern tropical South America from Patagonia and its subsequent diversifi cation. Th e data from insects analyzed by Sanmartín and Ronquist (2004, table 4) found a signifi cantly higher frequency of dispersal from southern South America to northern South America than from the other direction. Th is may be true for other caddisfl ies with both Patagonian and Neotropical (sensu de Moor and Ivanov) distributions, including Antarctoecia (Huamantinco and Nessimian 2003), Atopsyche and Cailoma (Hydrobiosidae) (Ross and King 1952, Flint 1974, Schmid 1989 [although Atopsyche is absent from Patagonia, its putative sister genus is Patagonian]), Contulma (Anomalopsychidae) (Holzenthal and Flint 1995), Smicridea (Smicridea) (Hydropsychidae) (Flint 1989), Tolhuaca (Glossosomatidae) (Robertson and Holzenthal 2005), and Triplectides (Leptoceridae).
As confi rmed by Crisci et al. (1991), Sanmartín and Ronquist (2004), and Amorin et al. (2009) the historical biogeography of southern South America is complex. Th e distribution of the austral South American caddisfl ies support this conclusion, with a fauna pertaining strongly to a transantarctic pattern, but with perhaps older Gondwana elements, as exhibited by Sericostomatidae. Th e current weight of evidence described above, however, supports a more recent dispersal of this southern fauna to northern South America.
Other than in a few studies, phylogenetic hypotheses are lacking for most of the taxa reviewed above. Phylogenies of Southern Hemisphere caddisfl y taxa inferred from molecular data are even fewer (e.g., Johanson and Keijsner 2008) and only one ) has used events-based models (e.g., Sanmartín at al. 2001, Ronquist andSanmartín 2004) or divergence time estimates (but see Amorin et al. 2009 for a critique of molecular dating) to address historical biogeography. Given the current ease with which molecular sequence data can be obtained and with the availability of newer analytical methods (e.g., Ronquist 1997, Sanderson 2002, Zaldivar-Riverón et al. 2008, there is a wealth of hypotheses that can be tested regarding the historical biogeography of the austral caddisfl y fauna once phylogenetic information is available (Santos and Amorin 2007).