Grumichella, MULLER, 1879

Calor, Adolfo R., Holzenthal, Ralph W. & Froehlich, Claudio G., 2016, Phylogeny and revision of the Neotropical genus Grumichella Müller (Trichoptera: Leptoceridae), including nine new species and a key, Zoological Journal of the Linnean Society 176 (1), pp. 137-169 : 139-140

GRUMICHELLA MÜLLER, 1879

Grumichella Müller, 1879: 407 (Type species: Grumichella rostrata Thienemann, 1905 , first includ- ed species). – Holzenthal, 1988a:88 [male, female, larva, pupa, revision, phylogeny, distribution]. – Flint, Holzenthal & Harris, 1999: 128 [catalog]. – Morse, 1981: 259 [classification, phylogeny]. – Morse & Holzenthal, 1987: 140 [classification, phylogeny]. – Calor & Holzenthal, 2008: 255 [phylogeny]. – Malm & Johanson, 2011: 3, 7 [classification, phylogeny].

Leptocellodes Ulmer, 1911: 21 [Type species: Leptocellodes flaveola Ulmer, 1911 , original designation (the designation of G. flaveola Ulmer, 1911 as type species by Ulmer (1955) is not valid (ICZN 1985, Art. 69(a) sensu Holzenthal, 1988a)]; – Ulmer, 1955: 499 [to synonymy].

Grumichella Müller, 1879 (= Leptocellodes Ulmer, 1955 ) comprises 13 species, including the nine new species described here: G. aequiunguis Flint, 1983 ; G. blahniki sp. nov.; G. boraceia sp. nov.; G. cressae sp. nov.; G. flaveola (Ulmer) Holzenthal, 1988a , G. jureia sp. nov.; G. leccii sp. nov.; G. muelleri sp. nov.; G. paprockii sp. nov.; G. parati sp. nov.; G. pulchella (Banks) Holzenthal, 1988a ; G. rostrata Thienemann, 1905 ; G. trujilloi sp. nov.

Müller (1879a, b, 1880a, b, 1888, 1921) illustrated the larvae of an unknown species and discussed its biology. Pupae and cases of two species, G. rostrata and G. aequiunguis , were described by Thienemann (1905). Roback (1966) described and illustrated the larvae of G. flaveola from Peru, but included it under ‘unknown family 2’. Ulmer (1955) also described immature forms of the genus. Morse (1981) erected the tribe Grumichellini to accommodate Grumichella and Atanatolica in his family classification. Holzenthal (1988a) reviewed the genus, redescribed the four valid species, and described the larvae and pupae, including those of eight probable unknown species of larvae (Holzenthal’s ‘species A–H’). He also corroborated the monophyly of Grumichella based on 20 synapomorphies and proposed the phylogenetic relationships among two species groups: G. pulchella and G. rostrata groups. The first species group, composed of G. flaveola and G. pulchella , is characterized by nine spots on the forewings. The G. rostrata group, comprising G. aequiunguis and G. rostrata , bears only two spots on forewings. Holzenthal & Pes (2004), in their description of Amazonatolica , another Grumichellini genus, discussed other characters and the phylogenetic position of Grumichella . Calor & Holzenthal (2008) described another related genus, Osflintia , and proposed a phylogeny of Grumichellini , placing Osflintia as sistergenus of all other grumichelline genera.

Biological remarks: Müller (1879a, b) provided the first records of the habitat of larvae, including their occurrence in small waterfalls and rock surfaces in fast flowing waters of small mountain streams. These observations were corroborated by others authors (e.g. Holzenthal, 1988a; Flint et al., 1999). In our fieldwork, the larvae (and pupae) were often collected from aggregations of more than 50 individuals positioned side by side or in a small clump (e.g. Parque Estadual de Campos do Jordão, São Paulo State, Brazil). Müller (1879a, b) also observed the use of the posterior silken projection on the larval cases to help to hold the larva in the current. The ability of larvae to maintain position in fast flow is aided by the stout legs with modified tarsal claws, as emphasized by Holzenthal (1988a) and by other authors observing larvae of other grumichelline genera (St. Clair, 1994; Ward, 2001). Müller (1879a, b) ‘speculated’ that the water current prevented the pupae from crawling out of the case, and that the anterior attachment pedicle of the case needed to be severed by the emerging pupae. The loose case then drifted to slack current, where the adult could emerge ( Holzenthal, 1988a). In laboratory rearings and rarely in natural habitats (e.g. G. boraceia sp. nov.), a behaviour of flotation and drift was observed in the fifth instar larvae, with air bubbles inside the anterior portions of cases (our pers. observ.).

The use of abandoned larval cases by other genera of caddisflies is not uncommon, especially in Triplectides ( Holzenthal, 1988b; Flint et al., 1999; Crisci-Bispo, Bispo & Froehlich, 2004; Calor & Froehlich, 2008), and includes observations by A.R.C. and by P.A. Rueda- Martín (pers. commun.) in Brazil and Argentina, respectively, of Marilia (Odontoceridae) occupying Grumichella cases.