Genus Pamphilius Latreille, 1803

Pamphilius Latreille, 1803: 303 .

See Taeger et al. (2010) for more synonymy.

Pamphilius is a large and rather heterogeneous genus, characterized as follows: anterior part of malar space without setiferous area (female) or pit (male) (Fig. 2: 3aa); supraocular area in male without narrow, dull, densely pubescent patch; facial crests in male various in shape, nearly flattened, rounded or carinate (e.g., Figs 55e, 63e, 65e); ventral inner surface of fore tibia without disconnected row of scale-like setae; tarsal claws without acute basal lobe (e.g., Figs 56g, 74i, 89g). It is distinguishable from the related genera by the features given in the key (see Shinohara 2002b for more details), but as discussed above, the monophyly of this genus is uncertain based on both morphological and molecular data.

Based on morphology, Shinohara (2002b) divided the world species of this genus into 12 species groups, of which four major groups ( P. vafer, P. sylvaticus, P. alternans and P. histrio groups) contained about 80% of all the species then known. Each of the remaining eight species groups contained only one to four species. The P. ochreipes group, represented by two Nearctic species, is here treated as a part of the P. vafer group. In the Russian Far East and Korea, the P. inanitus, P. latifrons and P. sulphureipes groups have been recorded in addition to the four major species groups.

For the molecular analyses of the present study, representatives of the seven species groups mentioned above, except for the P. latifrons group, and additionally the P. basilaris group, were used. The Pamphilius basilaris group contains two species associated with Juglandaceae from Japan and China (Shinohara 2003b; Shinohara et al. 2012). It has not been recorded from the Russian Far East and Korea, but the species group is likely to be found particularly in Korea.

The monophyly of each species group and the relationship among the species groups were discussed by Shinohara (2002b) based on morphology and will be examined further below in the light of the results of molecular analyses. Generally speaking, the monophyly of each species group was at least partly supported also by molecular data (see discussion under respective species group below) but the relationship among the species groups is still poorly clarified, as the hypotheses based on three different data sets (morphology, Fig. 14; COI sequences, Fig. 15; NaK sequences, Fig. 16) differ drastically (see also discussion under Pamphiliini above).

Pamphilius is distributed in the Holarctic region and contains 123 valid species and subspecies (Taeger et al. 2010; Shinohara & Wei 2012, 2016; present work), of which 43 are recorded for the Russian Far East and Korea.

The larvae of Pamphilius species feed on the host leaves singly or gregariously in simple leaf-rolls or in webs. Known host plants include Rosaceae ( Cerasus, Padus, Sorbus, Rosa, Rubus, Filipendula, Fragaria, Aruncus, Spiraea, etc.), Betulaceae ( Alnus, Betula, Corylus, Carpinus, etc.), Salicaceae ( Populus, Salix), Spindaceae ( Acer), Caprifoliaceae ( Lonicera, Macrodiervilla), Adoxaceae (Viburnum), Fagaceae (Quercus), Cornaceae (Cornus), Juglandaceae ( Juglans, Platycarya) and Saxifragaceae (Astilbe) (Shinohara 2002b, 2003b; Shinohara & Hara 2011; Shinohara & Kojima 2011; Shinohara et al. 2012, 2016a, b, 2019; Shinohara & Wei 2016). In some cases, distinct association between a species group and host plant family exists as noted under each species group below.