Neduba inversa Cole, Weissman, & Lightfoot, sp. n.

Fig. 19 (distribution), Fig. 27 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 3C (live habitus), Plate 5G (male calling song), Plate 8D (male ventral sclerite), Plate 10G (male titillators), Plate 12E (female subgenital plate).

Common name. Kings Canyon Shieldback.

History of recognition. Likely confused with N. sierranus .

Type material. HOLOTYPE MALE:, USA, CA, Fresno Co., Bretz Mill Campground, Sierra National Forest, 24 mi. NE Trimmer on Big Creek Rd., 37.03775N, 119.24040W, 871 m, 13-VIII-2015, JA Cole, JCT15-15 [karyotype], 150820_01 [recording], 206 [teeth], 3.3 [mm file count], tegmen in gel capsule and genitalia in vial below insect deposited in CAS, Entomology type #19710 . PARATYPES (n = 19): Fresno Co., 1♁, same data as holotype, CAS; 2♁, same data as holotype, LACM; 3♁, 1♀, Bretz Mill Campground, Sierra National Forest, 24 mi. NE Trimmer on Big Creek Rd., 37.0375N, 119.2388W, 1006 m, 29-30-VII-2012, JA Cole, LACM ; 3♁, Princess Campground, Sequoia National Forest, 36.80456N, 118.94154W, 1797 m, 25-27-VII-2017, JA Cole, CAS ; 7♁, 1♀, same data except LACM; 1♁, same data except JAC .

Measurements. (mm, ♁n = 17, ♀ n = 2) Hind femur ♁20.22–24.12, ♀ 23.15–23.40, pronotum total length ♁8.17–10.54, ♀ 8.48–9.42, prozona length ♁3.57–5.11, ♀ 4.06–4.98, metazona dorsal length ♁4.29–5.95, ♀ 3.50– 5.36, pronotum constriction width ♁2.22–2.95, ♀ 2.43–2.94, metazona dorsal width ♁5.68–6.63, ♀ 6.25–6.42, head width ♁4.84–5.47, ♀ 5.24–5.59, ovipositor length ♀ 15.81–16.78.

Distribution. West slope of the Sierra Nevada, between the San Joaquin and Kings River watersheds in the vicinity of Kings Canyon National Park.

Habitat. Understory of mixed conifer forests.

Seasonal occurrence. Available records are midsummer from July (1-VII-1935, EE Honeycutt, CAS) through mid-August (13-VIII-2015, JA Cole, LACM) .

Stridulatory file. (n = 7) length 3.0–4.0 mm, 206–266 teeth, tooth density 66.3 ± 6.9 (55.5–78.3) teeth/mm.

Song. (n = 24) Qualitatively like Sierranus Group taxa, high frequency (PTF 15.9 ± 1.2 kHz) with a fluttering quality caused by multiple OPT (PTN 9.2 ± 3.4). PTR is significantly slower (1.3 ± 0.3 s- 1) than N. sierranus and faster than N. radocantans (ANCOVA, P = 3.96 × 10 -9) but is indistinguishable from that of N. arborea .

Karyotype. (n = 5) 2n♁= 21 (2m + 16t + XtXtYm). JCT15-15, paratopotype.

Recognition. Males have a high stridulatory file tooth density (64–68 teeth/mm) like N. sierranus and N. radocantans . A weakly constricted pronotum separates N. sierranus from this species. N. radocantans is morphologically separable only by its slightly higher stridulatory file tooth density (68–75 teeth/mm). This species belongs to the Sequoia Group lineage but has a Sierranus Group song with multiple OPT. The high PTN of N. radocantans will separate that otherwise cryptic species from N. inversa . Males of the neighboring species to the north, N. sierranus, have songs with a faster PTR than those of N. inversa . Songs from all population of the Sequoia Group to the south have only one OPT. Karyotypes also separate N. inversa from the geographically proximal species mentioned above. The distribution of this species lies between the San Joaquin and Kings River drainages in the vicinity of Kings Canyon National Park.

Etymology. l. inversa “to change, to pervert, to turn upside down,” in reference to the mosaic of characters possessed by this species, with the song of one clade but the genetics of another.

Notes. At the type locality the species was common, but the quiet male songs were drowned out by the incessant loud calls of the shield-back katydid Cyrtophyllicus chlorum Hebard. The distribution of N. inversa lies where the Sierranus and Sequoia Groups meet in the central Sierra Nevada (Figs. 8, 19). This species combines characters from the two lineages and gene flow between neighboring lineages has occurred during its evolutionary history (Fig. 4). The stridulatory file and the calling song with a fluttering sound, caused by numerous OPT, is similar to N. sierranus in the Sierranus Group, which is distributed to the north in the Yosemite Valley region. DNA places this species with Sequoia Group species to the south, however, and this species shares a karyotype with N. prorocantans . The remaining Sequoia Group species have simple songs with one OPT between any pair of MPT. Song alleles may have introgressed across species boundaries in this contact zone region (e.g. Cole 2016).

Material examined. Type series only, see Type material above.