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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., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 64-66, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800

Neduba castanea (Scudder, 1899) Fig. 19 (distribution), Fig. 21 (male and female habitus, calling song, drumming, male and female tremulation karyotype), Plate 2 B–C (live habitus), Plate 5A (male calling song), Plate 7I (male ventral sclerite), Plate 10B (male titillators), Plate 11K (female subgenital plate). Common name. Chestnut Shieldback. History of recognition. Scudder (1899) described Tropizaspis castanea from Los Angeles County, California. Caudell (1907) transferred this species to Aglaothorax based on habitus, in particular the pronotal shape, but the species was moved to Neduba once the male genitalia were examined (Rentz 1988; Rentz & Birchim 1968). Our fieldwork shows that N. castanea is the only Neduba species found in mainland Los Angeles County (i.e. exclusive of the Channel Islands), where it is limited to the fringes of the Mojave Desert. Type material. The holotype male is housed at ANSP. Images of the type are available at OSFO (Cigliano et al. 2020). PARATYPE (n = 1): USA, CA , Los Angeles Co., 1♀, Los Angeles Co., VII, Coquillett, USNM; TOPOTYPES EXAMINED (n = 26): USA, CA, Los Angeles Co., 2♁, County road N4 near Llano, 34.505828N, 117.817841W, 1280 m, 6-VIII-1988, DB Weissman & DC Lightfoot, CAS; 1♁, County Road N6, 2 mi. NW of Devil’s Punchbowl, 34.4267N, 117.8697W, 1250 m, 6-VIII-1988, DB Weissman & DC Lightfoot, CAS; 1♁, Devil’s Punchbowl County Park, 34.4138N, 117.8587W, 1445 m, 17-VIII-1982, DB Weissman, CAS; 1♁, Devil’s Punchbowl County Park, 34.4138N, 117.8587W, 1445 m, 30-VIII-1975, PH Sullivan, CAS; 1♁, Devil’s Punchbowl Road 1 mi. NW of Devil’s Punchbowl, 34.42382N, 117.86816W, 1341 m, 20-VI-2018, JA Cole, JAC; 1♁, Juniper Hills, Devil’s Punchbowl Road (County Road N6), 2 miles east of Longview Road, 34.4267N, 117.8697W, 1316 m, 2005, JA Cole, LACM; 3♁, 1♀ nymph, same data except 13-VI-2003, JA Cole, JN Hogue, LACM; 1♁, same data except JAC; 2♁, same data except 14-VI-2015, JA Cole, GE Bell, T Farwell, LACM; 1♁, same data except JAC; 2♁, same data except 14-VI-2017, JA Cole, LACM; 3♁, 1♀, same data except 18-VI-2008, JA Cole, LACM; 1♀ nymph, same data except 20-VI-2003, JA Cole, JN Hogue, LACM; 1♁, same data except 7-VII-2004, JA Cole, LACM; 1♁, 1♀, Largo Vista Rd. (county Road N4), 1 mi. S of Fort Tejon Rd., 34.4530N, 117.7649W, 1277 m, 22-VI-2008, JA Cole, LACM; 1♀, same data except 22-VI-2008, JA Cole, JAC. Measurements. (mm, ♁n = 14, ♀ n = 6) Hind femur ♁16.99–21.20, ♀ 18.50–23.87, pronotum total length ♁9.90–13.20, ♀ 10.08–13.20, prozona length ♁3.84–5.69, ♀ 4.83–6.92, metazona dorsal length ♁5.60–8.67 ♀ 4.72– 6.76, pronotum constriction width ♁2.70–3.68, ♀ 2.95–3.85, metazona dorsal width ♁7.24–8.49 ♀ 6.75–8.25, head width ♁4.94– 5.85, ♀ 5.05–6.81, ovipositor length ♀ 14.92–20.25. Habitat. Pinyon-juniper (Juniperus spp.- Pinus monophylla Torr & Frém.), Joshua tree (Yucca brevifolia Engelm.) woodlands, Ceanothus scrub, and yellow pine forest. Specimens were taken from pinyon pine, juniper, mountain mahogany (Cercocarpus sp.), and Joshua tree. Distribution. Dry north slope of Transverse Ranges of California including the San Gabriel, San Bernardino, Liebre, and San Emigdio Mountains. Seasonal occurrence. Adult specimens have been taken from mid-June in foothill regions (13-VI-2003, JA Cole & JN Hogue, LACM) into September at high elevation (12-IX-2015, DA Gray, CAS). Females were last instar nymphs at the earliest date of occurrence listed above. Stridulatory file. (n = 5), length 3.7–4.6 mm, 79–99 teeth, tooth density 21.6 ± 1.1 (19.8–22.7) teeth/mm. Song. (n = 24) A continuous series of long MPTL (373.1 ± 51.5 ms) with widely spaced toothstrikes delivered at PTR of 2.6 ± 0.7 s- 1. The sound resembles a finger running along the teeth of a comb. The PTF of 13.7 ± 2.1 kHz extends comfortably into the audible range. There is a significant high frequency component, however, with 18.7 kHz PTF measured in one high frequency laboratory recording. Males begin calling at sunset. The population from McGill Campground on Mount Pinos, Ventura County, California has a significantly slower PTR than that of the type locality (2-sample t -test, P = 0.002). Males may also produce audible substrate vibrations via drumming. Drums precede bouts of stridulation (Fig. 21). Karyotype. (n = 5) 2n♁ = 22 (4m + 16t + XmYt), shared only with N. macneilli. T88-69, S88-72, topotype. Recognition. Morphological and color pattern differences thought to separate N. castanea from N. macneilli (per Rentz & Birchim 1968) are unreliable. The development of the styli on the subgenital plate, from absent to slight swellings to articulated, may vary within a population and even between the left and right sides of a single specimen. Both species exhibit the full range of variation in color patterns. Female N. castanea have round, highly convex subgenital plates, whereas those of N. macneilli are subtriangular with a bluntly pointed apex. Geographically N. castanea replaces N. macneilli, which is a Sierra Nevada species, in the Transverse Ranges of southern California. Notes. We treat N. castanea and N. macneilli as closely related sister species based on reproductive isolation via allopatry. Both inhabit the same life zone in different geographic regions of California. N. castanea and N. macneilli are sympatric with Aglaothorax ovata, and all three species share a similar robust habitus suggesting convergent evolution of body form in their habitat. Enlarged pronotal size may be adaptive for signal transmission in open habitats such as pinyon-juniper and Joshua tree woodlands, where trees and bushes are widely spaced, compared with the dense forest understories inhabited by other Neduba species Groups. Drumming was observed during courtship in the field at the Largo Vista Road locality (see Type material above), during which a male alternated drumming and stridulating. The signaling did not result in copulation in this observed instance (JAC pers. obs.). Material examined. (n = 26) All USA, CA, Kern Co., 3♀, McGill Campground, Los Padres National Forest, 34.82539N, 119.09791W, 12-IX-2015, DA Gray, LACM; 11♁, 1♀, McGill Campground, Los Padres National Forest, 34.81505N, 119.10014W, 2271 m, 8-9-VIII-2017, JA Cole, LACM; 1♁, 1♀, same data except JAC; Los Angeles Co., in addition to type material (above), 5♁, Sierra Highway, 0.6 miles west of Crown Valley Road, Acton, 34.4936N, 118.1834W, 929 m, 17-VI-2003, JA Cole, JN Hogue, LACM; 1♁, same data except 18-19-VI-2008, JA Cole, JAC sound record; 1♁, Spunky Canyon, 2 mi. SW of Green Valley, 34.6023N, 118.3863W, 1047 m, 16-VI- 2012, JA Cole, LACM; 2♁, same data except 26-VII-2001, JA Cole, LACM., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 49-52, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Scudder, S. H. (1899) Two genera of North American Decticinae. Proceedings of the American Academy of Arts and Sciences, 35, 81 - 93. https: // doi. org / 10.2307 / 25129903","Caudell, A. N. (1907) The Decticinae (a group of Orthoptera) of North America. Proceedings of the United States National Museum, 32, 285 - 410. https: // doi. org / 10.5479 / si. 00963801.32 - 1530.285","Rentz, D. C. (1988) The shield-backed katydids of southern Africa: their taxonomy, ecology and relationships to the faunas of Australia and South America (Orthoptera: Tettigoniidae: Tettigoniinae). Invertebrate Taxonomy, 2, 223 - 335. https: // doi. org / 10.1071 / IT 9880223","Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0."]}

Neduba extincta Rentz, 1977 Fig. 19 (distribution). Common name. Extinct Shieldback. History of recognition. Described from a single male museum specimen deposited at CAS (Rentz 1977). Type material. The holotype male is the only specimen known. Images of the holotype are available at OSFO (Cigliano et al. 2020). Measurements. See Rentz (1977). Distribution. Antioch Sand Dunes, Contra Costa County, California, on the western edge of the Central Valley. Habitat. Historically known from the sandy banks of the San Joaquin River, elevation 9 m. The 55 acre Antioch Dunes National Wildlife Refuge is the only National Wildlife Refuge in the country established to protect endangered plants and insects. Seasonal occurrence. The only known specimen was collected 1-VII-1937 (ES Ross, CAS). Stridulatory file. (n = 1) length 3.2 mm, 167 teeth, tooth density 52.2 teeth/mm, Song. Unknown. Karyotype. Unknown. Recognition. Description indicates large body size and absence of styli on subgenital plate. The stridulatory file tooth density places this species at the upper end of variation for the Sequoia Group. The tooth density of the single known specimen is less than the closest geographic relative, N. inversa (density 64–68 teeth/mm). The single specimen is the only Sequoia Group individual collected in the Central Valley west of the Sierra Nevada. Notes. This species is one of four extinct North American Orthoptera species (Hoekstra 1998) and that status has not changed; David Rentz and DBW have searched for this species at the type locality on several occasions over the last few decades, visiting during summer months when Neduba are active and using a bat detector. On no occasion were individuals found. The lack of molecular, bioacoustical, and cytogenetic characters make this species difficult to place in context of this revision, but the stridulatory file tooth density is consistent with the Sequoia Group. Minimally destructive molecular work may be undertaken in the future to place N. extincta into phylogenetic context. Among the extant Sequoia Group species, N. inversa is distributed near the San Joaquin River watershed and is therefore a possible relative, and this lineage could have colonized the western edge of the Central Valley across riparian corridors. The description of N. arborea in this work reports the only other Sierranus or Sequoia Group members known west of the Sierra Nevada. Many Neduba populations were no doubt extirpated as the eastern slope of the Coast Ranges became more arid and as humans modified the Central Valley for agriculture. In the case of the Antioch dunes, sand mining and commercial development drove extinction of this species. Thorough collecting efforts are needed in the eastern slopes of the Coast Ranges to search for possible unknown populations., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 63-64, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Rentz, D. C. (1977) A new and apparently extinct katydid from Antioch Sand Dunes (Orthoptera: Tettigoniidae). Entomological News, 88, 241 - 245.","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0.","Hoekstra, J. M. (1998) Conserving Orthoptera in the wild: lessons from Trimerotropis infantilis (Oedipodinae). Journal of Insect Conservation, 2, 179 - 185. https: // doi. org / 10.1023 / A: 1009612317241"]}

Neduba lucubrata Cole, Weissman, & Lightfoot, sp. n. Fig. 19 (distribution), Fig. 23 (male and female habitus, calling song, drumming, male and female terminalia, karyotype), Plate 2E (live habitus), Plate 5C (male calling song), Plate 7 G–H (male ventral sclerites), Plate 10D (male titillators), Plate 12A (female subgenital plate). Common name. Midnight Shieldback. History of recognition. None. Type material. HOLOTYPE MALE, USA, CA, Monterey Co., Miller’s Lodge, 0.6 miles west jct. Arroyo Seco Road and G16, 36.25466N, 121.43208W, 192 m, 22-VII-2015, JA Cole, DB Weissman, JAC000002197 [specimen barcode], DNA161 [tissue], SING0503 [DNA extraction], JCT15-08 [karyotope], genitalia in vial under specimen, deposited in CAS, Entomology type #19712. PARATYPES (n = 32): All USA, CA , Monterey Co., 4♁, 1♀, same data as holotype, LACM; 2♁, 1♀, Arroyo Seco Rd., 0.6 mi. W of intersection with G16, 36.235139N, 121.473392W, 274 m, 29-VII-1983, DB Weissman, CAS; 3♁, Bottcher’s Gap, Los Padres National Forest, 19 miles north of Big Sur off SR1 on Palo Colorado Road, 36.355N, 121.8138W, 652 m, 20-21-VIII-2012, JA Cole, LACM; 1♁, same data except 7-8-IX-2002, JA Cole, LACM; 7♁, Nacimiento-Ferguson Rd., at bridge of Nacimiento River, 8.4 miles east of SR1, 36.0135N, 121.4216W, 587 m, 19-20-VIII-2012, JA Cole, LACM; San Benito Co., 9♁, Short Fence Trailhead, Coalinga Road, Laguna Mountain Recreation Area, BLM, 36.36403N, 120.8784W, 670 m, 10-11-VIII-2017, JA Cole, LACM; 2♁, same data except 10-11-VIII-2017, JA Cole, JAC; 2♁, Upper Sweetwater Campground, Coalinga Road, Laguna Mountain Recreation Area, BLM, 36.36067N, 120.85256W, 848 m, 10-11-VIII-2017, JA Cole, LACM. Measurements. (mm, ♁n = 24, ♀ n = 2) Hind femur ♁16.35–20.74, ♀ 21.61–22.90, pronotum total length ♁7.04–9.03, ♀ 8.95–9.17, prozona length ♁2.65–4.78, ♀ 4.02–4.18, metazona dorsal length ♁3.85–5.10, ♀ 4.77– 5.15, pronotum constriction width ♁1.85–2.66, ♀ 2.70–3.35, metazona dorsal width ♁5.36–6.25, ♀ 5.76–6.35, head width ♁3.90–4.70, ♀ 5.11–5.46, ovipositor length ♀ 16.50–16.60. Distribution. Santa Lucia and Diablo Ranges in the South Coast Range, California. Habitat. Mixed woodland and chaparral. Taken from twigs in tangles, poison oak, and California sage (Artemisia californica Less.). Males call approximately 1 m above ground level in thick tangles. At dusk two males and one female emerged for nocturnal activity from a pack rat nest, suggesting that this structure served as a daytime shelter for an aggregation of individuals. These individuals retreated into the nest when disturbed. Seasonal occurrence. Adults from late July (22-VII-2015, JA Cole and DB Weissman, LACM) through early September (8-IX-2002, JA Cole, LACM). Stridulatory file. (n = 7) length 3.2–3.8 mm, 121–186 teeth, tooth density 45.9 ± 4.2 (38.4–50.0) teeth/mm. Song. (n = 31) Brief bouts separated by long intervals between bouts. PTR is 4.0 ± 0.5 s- 1. Males may add one PT to each successive bout, for example three successive bouts may consist of 3 PT, 4 PT, then 5 PT, and then the cycle recommences at 3 PT. PTF is 14.9 ± 0.8 kHz. Males may accompany stridulation with audible drumming, which is generated by the abdomen striking the substrate (Weissman 2001; JAC pers. obs.). Drumming coincides with partial PT at the beginning of a bout (Fig. 23). Drumming does not occur frequently and may not occur in all populations. Drumming was observed at the Monterey County localities of Arroyo Seco (one of two males, DBW, JAC, pers. obs.) and Bottcher’s Gap (JAC, pers. obs.), but not at any San Benito County localities. Karyotype. (n = 8) Unique. 2n♁ = 24 (2m + 20t + XtYt). T83-37, S83-107, paratopotype. Recognition. With a single apical protibial spine, dark apical tegminal spots and prosternal spines, this species may be confused only with N. propsti, a larger species that does not occur on the California mainland. The female subgenital plate length and width are subequal, in contrast with the elongated subgenital plate of N. propsti. The song, which consists of short stridulation bouts accompanied by drumming, is unique. Etymology. l. lucubrata burning the midnight oil, descriptive of male acoustic activity continuing throughout the night. Notes. This species shares morphological and genitalic characteristics with N. propsti as well as with the species of the Sierranus and Castanea Groups. Santa Lucia Range and Diablo Range populations are genetically distinct (Figs. 3–5) but are separable neither morphologically nor bioacoustically. The chirping song structure combined with abdominal drumming makes N. lucubrata the most acoustically distinct species of Neduba. Selection for mate recognition may have driven the evolution of distinctive song phrasing in N. lucubrata, as within its distribution are found two sympatric nedubines: N. carinata and a small Aglaothorax species. Females may require silent periods of an appropriate intermediate length between bouts to recognize a conspecific male signal (Cole 2016): intermediate length gaps between PT bouts will contrast with the continuous PT production of sympatric N. carinata as well and the long periods of silence between pulse production in Aglaothorax. Material examined. Type series only. See Type material above. Sierranus Group The Sierranus Group is composed of 3 species (sierranus, arborea, and radocantans). Like the Sequoia Group, all Sierranus Group species have a single spine on the posterior margin of the forefemur, a pair of prosternal spines, and the entire male tegmen ivory or white. The male subgenital plate has lateral carinae that converge to the apex, which is devoid of styli, petal-like and often reflexed. The stridulatory file tooth density is the highest of all Neduba species groups (59–77 teeth/mm), and this character alone separates it from all Sequoia Group species except N. inversa. The Sierranus Group is distributed in the central and northern Sierra Nevada while the Sequoia Group occupies the southern portions of that mountain range (Figs. 8, 19). Within this Group are morphologically cryptic species defined by song and/or karyotype. Body part measurements and stridulatory files offer the only means to identify males that lack song data. Females may also be identified by body part measurements and sometimes the shape of the subgenital plate. Species are parapatric in the Sierra Nevada (Fig. 19) and thus geography will serve to narrow species possibilities. Molecular data show hybridization (Fig. 4) between species with adjacent ranges., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 55-57, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Weissman, D. B. (2001) Communication and reproductive behaviour in North American Jerusalem crickets (Stenopelmatus) (Orthoptera: Stenopelmatidae). In: Field, L. H. (Ed.), The Biology of Wetas, King Crickets and Their Allies. CAB International, New York, pp. 351 - 373. https: // doi. org / 10.1079 / 9780851994086.0351"]}

Neduba duplocantans Cole, Weissman, & Lightfoot, sp. n. Fig. 19 (distribution), Fig. 30 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 3H (live habitus), Plate 5J (male calling song), Plate 12H (female subgenital plate). Common name. Doubletime Shieldback. History of recognition. None. Type material. HOLOTYPE MALE: USA, CA, Tulare Co., South Fork Road, 9.75 miles southeast of Three Rivers, 36.35334N, 118.78451W, elev. 970 m, 25-VI-2017, JA Cole, W Chatfield-Taylor, 170625_03 [recording], JCT17-1 [karyotype], 173 [teeth], 3.4 [mm], tegmen in gelcap below specimen, deposited in CAS, Entomology type #19709. PARATYPES (n = 24): Fresno Co., 1♀, Big Creek Road, 5 mi. S of Bretz Mill Campground, 36.97546N, 119.21088W, 810 m, 13-VIII-2015, JA Cole, LACM; Tulare Co., 3 ♁, 8♀, same data as holotype, LACM; 2♁, same data as holotype, CAS; 1♀, same data as holotype, JAC; 1♁, 1♀, South Fork Campground, Sequoia National Park, 36.35209N, 118.76511W, 1112 m, 12-13-VIII-2015, JA Cole, CAS; 1♁, 6♀, same data except LACM. Measurements. (mm, ♁n = 7, ♀ n = 17) Hind femur ♁19.46–21.95, ♀ 21.44–22.69, pronotum total length ♁8.20–9.28, ♀ 8.10–8.95, prozona length ♁3.30–4.71, ♀ 3.48–5.28, metazona dorsal length ♁4.42–5.44, ♀ 3.09– 4.75, pronotum constriction width ♁2.00–2.35, ♀ 2.03–2.95, metazona dorsal width ♁5.60–6.05, ♀ 5.20–6.41, head width ♁4.60–5.12, ♀ 4.85–5.85, ovipositor length ♀ 13.09–17.60. Distribution. At present, known only from the vicinity of the Kaweah River in the Sequoia National Forest. Habitat. Found in tangles and among grass along a stream flowing out of a side canyon. Also collected on bushes along the road paralleling the Kaweah River. Seasonal occurrence. Males were active and females were a mixture of last instar nymphs, teneral adults, and mature adults in late June (25-VI-2017, JA Cole & W Chatfield-Taylor, LACM). By August this species was rare (12-VIII-2015, JA Cole, LACM). This species may be active earlier in the season than sympatric N. sequoia. Stridulatory file. (n = 7) length 2.7–3.4 mm, 126–173 teeth, tooth density 51.0 ± 4.9 (45.8–60.7) teeth/mm. Song. (n = 7) Unique. The PTR of 5.1 ± 0.3 s- 1 is much faster than any other Sequoia or Sierranus Group species, and twice as fast as sympatric N. sequoia. The MPTL is brief at 72.3 9.1 ms. The short MPTL contributes to a low PTdc of 35.8 ± 4.9%. PTF of 17.0 ± 1.0 kHz is higher than other Sequoia Group species and is difficult to hear except at short range. Karyotype. (n = 3) 2n♁= 21 (18t + XtXtYm), shared with N. sierranus, JCT 15-12, paratopotype. Recognition. Male N. duplocantans are separated from sympatric N. sequoia by stridulatory file tooth density (47–52 teeth/mm vs. 55–59 teeth/mm, respectively). To the north, N. inversa has a similarly constricted pronotum but that species also has a higher tooth density (64–68 teeth/mm). Male N. prorocantans to the south have similar stridulatory files but have a weakly constricted pronotum. The fast song PTR is unique, and readily separates this species by ear from syntopic N. sequoia as well as from neighboring N. prorocantans to the south, as well as N. inversa to the north due to the multiple OPT in that species. Adult activity apparently commences earlier in the season than N. sequoia in areas where they co-occur, but the two species may be found together in midsummer. Etymology. l. duplo “twice as much, double” + cantans “singing,” referring to the rate of PT production that is twice as fast as a sympatric congener. Notes. This species was discovered by song while collecting a series of N. sequoia, and its distinctiveness was appreciated further when cytogenetic analysis revealed a distinct karyotype. The discovery of two morphologically cryptic Sequoia Group species (N. duplocantans and N. sequoia) that are both sympatric and syntopic is of considerable evolutionary interest. The species pair has diverged in several aspects: calling song, karyotype, and phenology. The calling song PTR differed by a factor of two. Considering the remarkably different songs and distinct karyotypes, stasipatric and secondary contact hypotheses should be tested in the Sequoia Group. With N. duplocantans adult activity commencing before N. sequoia have matured, allochronic isolation may also be evolving between the two species. Two females among the paratype series were identified through rDNA sequence (Figs. 4-5). Introgression of mtDNA among the Sequoia Group species (Fig. 4) is expected if young, geographically proximal species have a history of hybridization as observed in Aglaothorax (Cole 2016). Material examined. (n = 2) Tulare Co., in addition to Type material (above), 1♀, Hammond, Kaweah Powerhouse 3 Forebay, 36.48618N, 118.83553W, 853 m, 17-VIII-2010, W Chatfield-Taylor, LACM; 1♀, Ash Mountain, Kaweah Power Station 3, 36.48606N, 118.83586W, 7-VI-1986, DJ Burdick, CAS.

Neduba macneilli Rentz & Birchim, 1968 Fig. 19 (distribution), Fig. 22 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 2D (live habitus), Plate 5B (male calling song), Plate 7 J–K (male ventral sclerites), Plate 10C (male titillators), Plate 11L (female subgenital plate). Common name. MacNeill’s Shieldback. History of recognition. N. macneilli was described from the eastern Sierra Nevada mountains of California by Rentz & Birchim (1968). The type locality is 1 mile west of Tom’s Place, Mono County, California. Tinkham (1944) referred to this species under the name carinata. Type material. The male holotype is housed at ANSP. Images of the holotype are available at OSFO (Cigliano et al. 2020). PARATYPES (n = 2): USA, CA , Mono Co., 2♁, Rock Creek, 1 mi. W Tom’s Place, 37.5586N, 118.7025W, 2143 m, 10-IX-1966, DC & KA Rentz, CAS; TOPOTYPES (n = 16): 4♁, 1♀, Tom’s Place, 37.5586N, 118.7025W, 2143 m, 28-VIII-1986, DB & BI Weissman, DCF Rentz, CAS; 8♁, 1♀, Tom’s Place, 1 mi. W, 37.5586N, 118.7025W, 2143 m, 15-16-VII-2012, JA Cole, LACM; 2♁, same data except JAC. Measurements. (mm, ♁n = 19, ♀ n = 4) Hind femur ♁16.35–21.50, ♀ 22.55–28.10, pronotum total length ♁9.89– 11.85, ♀ 10.15–12.13, prozona length ♁4.59–6.30, ♀ 4.61–6.68, metazona dorsal length ♁4.90–6.20, ♀ 4.78–5.54, pronotum constriction width ♁2.58–3.85, ♀ 3.15–3.42, metazona dorsal width ♁6.65–7.81, ♀ 6.96–7.55, head width ♁4.13–5.83, ♀ 5.93–6.25, ovipositor length ♀ 17.14–19.35. Distribution. Eastern slope of the Sierra Nevada Mountains of California. Habitat. Yellow pine forest and pinyon-juniper woodlands. Specimens were taken from Purshia tridentata Pursh (DC) and Pinus monophylla. Seasonal occurrence. Adult specimens have been taken from mid-July (10-VII-2003, JA Cole & JF Eguizabal, LACM) through October (22-X-1939, ER Tinkham, CAS). Stridulatory file. (n = 19) length 3.4–4.4 mm, 71–99 teeth, tooth density 23.1 ± 1.8 (19.8–26.4) teeth/mm. Song. (n = 43) The song of this species was published in Morris et al. (1975). The calling song of N. macneilli is not distinguishable from that of N. castanea (see discussion under the former species). In addition to calling song, 2 of 5 recorded males of N. macneilli (identified as N. castanea in Weissman (2001)) from 1.3 m E Walker Pass, Kern Co. (S88-68), produced an audible drumming by visibly tapping the substrate in the laboratory simultaneously with both hind legs. Of many males recorded from various populations, these were the only males of N. macneilli ever heard drumming. A courting topotypic male produced short PT accompanied by obvious body movements (tremulations) that could have caused substrate vibration. This male was displaying to a female nymph (JAC, pers. obs., 15-VII-2012). Karyotype. (n = 14) 2n♁ = 22 (4m + 16t + XmYt), identical to that of N. castanea, and corrects the result of Ueshima and Rentz (1979). T86-83, S86-98, topotype. Recognition. Males are indistinguishable from N. castanea. The female subgenital plate is subtriangular, in contrast to the rounded plate of N. castanea. This species inhabits the southeastern Sierra Nevada, whereas N. castanea inhabits the Transverse Ranges. Notes. Color pattern features that were used to separate N. macneilli and N. castanea (Rentz & Birchim 1968) are variable and are correlated with habitat rather than with lineage. The darker, mottled coloration of N. macneilli matches the background in higher elevation pinyon-juniper woodlands, while the lighter coloration typical of N. castanea is more cryptic in Joshua tree woodlands of the lower, arid foothill slopes. Both N. castanea and N. macneilli may have greenish or bluish abdominal pleura when alive (Plate 2C), colors that are invariably lost in museum specimens even if gutted and stuffed. Only freeze-drying retains such colors. Material examined. (n = 110). All USA, CA, Inyo Co, 1♁, 4 Jeffrey Campground, Inyo National Forest, 37.24759N, 118.56942W, 2479 m, 10-11-IX-2016, JA Cole, J Bailey, JAC sound record; 1♁, 8.1 mi. W of Big Pine on road to Sage Flat Camp, 37.164839N, 118.43694W, 2195 m, 28-VIII-1986, DB & BI Weissman, DCF Rentz, CAS; 11♁, 1♀, Big Pine Canyon, 37.164931N, 118.289546W, 16-VIII-1938, ER Tinkham, CAS; 1♁, same data except J Davis, CAS; 2♁, same data except 22-VIII-1948, ER Tinkham, CAS; 9♁, same data except 24-X-1939, ER Tinkham, CAS; 5♁, Glacier Lodge Rd. SR168, 1.38 mi. NE of Aspendell, 37.25047N, 118.58169W, 2497 m, 10-IX-2016, JA Cole, J Bailey, LACM; 1♀, Glacier Lodge, 11 mi. W of Big Pine, 37.164762N, 118.48971W, 4- VIII-1931, ER Tinkham, CAS; 5♁, 5♀, Lone Pine Canyon, 36.606044N, 118.062865W, 3-VIII-1931, ER Tinkham, CAS; 2♁, Lone Pine, 9 mi. W, 36.605934N, 118.225457W, 7-VIII-1961, JS Buckett, BMED; 2♁, Mouth Big Pine Canyon, 37.164931N, 118.289546W, 22-X-1939, ER Tinkham, CAS; 1♁, Sage Flat Camp, 8 mi. W of Big Pine, 37.082856N, 118.392482W, 2243 m, 19-VIII-1982, DB Weissman, CAS; 1♁, Sage Flat, 6 mi. SW of Big Pine on Glacier Lodge Rd., 37.1279N, 118.4037W, 2204 m, 13-VII-2003, JA Cole, JF Eguizabal, LACM; 1♁, same data except JAC; 1♁, same data except 4-VIII-2004, JA Cole, LACM; 1♀, Saline Valley, Grapevine Canyon Road Station 3, 36.58917N, 117.58472W, 1958 m, 18-VIII-1959, B. Banta, CAS; Kern Co., 6♁, 2♀, 1.3 mi. E of Walker Pass on SR178, 35.66245N, 118.003532W, 1460 m, 5-VIII-1988, DB Weissman & DC Lightfoot, CAS; 1♁, 1♀, Freeman Canyon, 1.5 miles southeast of Walker Pass on SR178, 35.6509N, 118.0055W, 1470 m, 10-VII-2003, JA Cole, JF Eguizabal, LACM; 2♁, 2♀, same data except 14-VII-2005, JA Cole, LACM; 4♁, SR178, 4.1 mi. W of Walker Pass, 35.662431N, 118.099936W, 1280 m, 5-VIII-1988, DB Weissman & DC Lightfoot, CAS; 2♁, 1♀, Walker Pass, 35.662453N, 118.02674W, 1600 m, 18-VIII-1982, DB Weissman, CAS; 1♁, Walker Pass, 35.67669N, 118.04383W, 1416 m, 21-22-VII-2015, JA Cole, DB Weissman, JAC sound record; 1♁, Walker Pass, 35.662453N, 118.02674W, 1537 m, 21-IX-1967, RE Love, CAS; 4♁, 4♀, Walker Pass, 35.662453N, 118.02674W, 1537 m, 22- VIII-1938, ER Tinkham, CAS; 2♁, same data except 29-IX-1960, JR Helfer, CAS; 3♁, Walker Pass, 35.662453N, 118.02674W, 1524 m, 7-IX-1966, DC & KA Rentz, CAS; 1♁, Walker Pass Campground, BLM, 15 mi. E of Onyx off SR178, 35.689934N, 117.952753W, 1537 m, 13-14-VII-2012, JA Cole, JAC sound record; 2♁, Walker Pass Recreation Area, BLM, 15 miles east of Onyx off SR178, 35.6646N, 118.037W, 1537 m, 13-14-VII-2005, JA Cole, LACM; 1♁, same data except 23-24-VI-2008, JA Cole, JAC sound record; Mono Co., in addition to type material (above), 2♁, 13 mi. N of Lee Vining, 38.145913N, 119.121816W, 2164 m, 24-VIII-1957, ER Tinkham, CAS; 1♁, French Camp, Inyo National Forest, 0.25 mi. S of Tom’s Place on Rock Creek Rd., 37.5505N, 118.6836W, 2249 m, 16-VII-2012, JA Cole, LACM; 11♁, Mono Lake, 38.007604N, 119.014763W, 1958 m, 11-VIII-1938, ER Tinkham, CAS; 7♁, Mouth of Tioga Pass, nr. Mono Lake, 38.007604N, 119.014763W, 1-VIII-1931, ER Tinkham, CAS; Tulare Co., 2♁, Kennedy Meadow, 26 miles northwest of Pearsonville via Nine Mile Canyon Road and Kennedy Meadows Road, 36.0518N, 118.1288W, 1883 m, 11-13-VII-2003, JA Cole, JF Eguizabal, LACM. Lucubrata Group The Lucubrata Group contains one early branching species (Figs. 3–5) that inhabits the South Coast Ranges of California (Figs. 8, 19). This group is defined by having one spine on the posterior margin of the fore femora, a pair of well-developed prosternal spines, and in having the apices of the tegmina darkened. This suite of characters is shared only with the Propsti Group. The lateral carinae of the male subgenital plate taper regularly to a narrow apex and styli are rudimentary. Male subgenital plates of the Carinata, Propsti, and Castanea Groups have subparallel lateral carinae and well-developed articulate styli. The song of the Lucubrata Group is delivered in bouts (Plate 5C), as opposed to the continuous PT of the Propsti Group (Plate 4J). The karyotype is unique., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 52-54, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Tinkham, E. R. (1944) Biological, taxonomic and faunistic studies on the shield-back katydids of the North American deserts. American Midland Naturalist, 31, 257 - 328. https: // doi. org / 10.2307 / 2421073","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0.","Morris, G. K., Aiken, R. B. & Kerr, G. E. (1975) Calling songs of Neduba macneilli and N. sierranus (Orthoptera: Tettigoniidae: Decticinae). Journal of the New York Entomological Society, 83, 229 - 234.","Weissman, D. B. (2001) Communication and reproductive behaviour in North American Jerusalem crickets (Stenopelmatus) (Orthoptera: Stenopelmatidae). In: Field, L. H. (Ed.), The Biology of Wetas, King Crickets and Their Allies. CAB International, New York, pp. 351 - 373. https: // doi. org / 10.1079 / 9780851994086.0351","Ueshima, N. & Rentz, D. C. (1979) Chromosome systems in the North American Decticinae with reference to Robertsonian changes. Cytologia, 44, 693 - 714. https: // doi. org / 10.1508 / cytologia. 44.693"]}

Neduba ambagiosa Cole, Weissman, & Lightfoot sp. n. Fig. 9 (distribution), Fig. 12 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 1 B– C (live habitus), Plate 4C (male calling song), Plate 6I (male ventral sclerite), Plate 6 G–H (male ventral sclerites, ambagiosa x carinata hybrids), Plate 9D (male titillators), Plate 11C (female subgenital plate). Common name. Ambiguous Shieldback. History of recognition. Individuals of this species were encompassed in the type series of N. convexa (Caudell 1907). Type material. HOLOTYPE MALE: USA, CA, Lake Co., Middle Creek, 2.7 mi. S of Middle Creek Campground on Elk Mountain Road, 39.21569N, 122.93381W, 430 m, 6-VIII-2014, JA Cole, DB Weissman, JAC000001944 [specimen barcode], S14-64 [stop], JCR140918_00 [recording], 110 T [teeth] 3.4 [mm stridulatory file length], genitalia extracted and cleared in vial and excised tegmen in gelcap below specimen, deposited at CAS, Entomology type #19706. PARATYPES (n = 27): USA, CA, Lake Co., 6♁, 3♀, Middle Creek Campground, 7 miles northwest of Upper Lake on Elk Mountain Road, 39.2537N, 122.9502W, 610 m, 3-6-VII-2003, JA Cole, LACM; 1♁, 1♀, same data except JAC; 5♁, 4♀, Middle Creek, 2.7 miles south of Middle Creek Campground on Elk Mountain Road, 39.21569N, 122.93381W, 430 m, 6-VIII-2014, JA Cole, DB Weissman, LACM; Mendocino Co., 4♁, 3♀, Tranquility, 1 mile south of Caspar, 39.349009N, 123.815849W, 7-VII-1982, JR Helfer, CAS. Measurements. (mm, ♁n = 16, ♀ n = 9) Hind femur ♁17.59–22.4, ♀ 19.95–22.36, pronotum total length ♁7.82– 11.14, ♀ 6.90–8.84, prozona length ♁3.26–4.91, ♀ 3.75–5.12, metazona dorsal length ♁4.35–6.23, ♀ 2.93–3.87, pronotum constriction width ♁1.55–2.93, ♀ 2.30–3.15, metazona dorsal width ♁5.43–7.00, ♀ 4.20–6.15, head width ♁3.75–4.80, ♀ 4.42–5.10, ovipositor length ♀ 13.43–18.81. Distribution. North Coast Ranges of California. The type locality is in the vicinity of Clear Lake. Habitat. Understory of chaparral and mixed conifer forests, riparian corridors, and canyon bottoms. On leaf litter, poison oak, and on tree trunks. Seasonal occurrence. mid-June (16-VI-1978, DB Weissman, CAS) through November (15-XI-1953 JR Helfer, CAS). Nymphs from December through July. Stridulatory file. (n = 13) length 2.6–3.9 mm, 94–139 teeth, tooth density 34.2 ± 2.6 (30.8–40.9) teeth/mm. Song. (n = 11) PTR 2.9 ± 0.3 s- 1, identical to topotype N. convexa. PTF 13.0 ± 0.9 kHz. Males are nocturnal singers. A genetically identified hybrid population (Fig. 4; see also Notes under N. carinata) from Solano County, California has a significantly faster pulse rate of 3.1 ± 0.2 s- 1 (ANCOVA, P = 1.08×10 -5). Karyotype. (n = 9) 2n♁ = 26 (2m + 22t + XtYt). T14-24, S14-64, paratopotype. Recognition. A low stridulatory file tooth density (30–40) will separate this species from N. carinata and N. diabolica distributed to the south. The ventral sclerite apex is usually conical to pyramidal, rarely flat, with a short to medium-length lateral process. The length of the lateral process is shorter than most N. convexa, a species of the Sierra Nevada, northern Central Valley and Cascade Range, but is longer than that of N. oblongata in the Diablo Range. The lateral process is shorter than the shaft, separating this taxon morphologically from N. longiplutea distributed farther north in the North Coast Ranges. Curved apical margins and a distinct central groove on the triangular subgenital plate will separate females of this species from N. carinata, N. diabolica, and N. oblongata which are pentagonal with straight apical margins. The subgenital plate of N. longiplutea to the north is longer than wide with a bifurcate apex in contrast to the subequal length and width and entire, acute apex of this species. Calling song will separate this species from N. carinata, N. oblongata, and N. diabolica as all are species with similar length MPT and OPT and fast PTR. Etymology. l. ambagiosa circuitous, indirect, roundabout, referring to the distribution being the southwestern extent of a ring of Convexa Clade taxa around the Central Valley (Fig. 8). Notes. This species has exchanged genes with populations of N. carinata (see also Notes section under N. carinata species account). In addition to DNA evidence (Fig. 4), males from a population along Putah Creek in Solano County have a high stridulatory file tooth density as in N. carinata but sing with N. convexa song, albeit with a faster PTR that may be influenced by N. carinata alleles. The geography of this hybrid population suggests that other populations from the northern and eastern Bay Area may also have hybrid characteristics. Stridulatory files from north Bay Area populations that we have examined are within the range for this species. Populations in inland locations such as Solano County may experience gene flow that is otherwise prevented across the Coast Ranges by San Francisco Bay.A precopulatory pair was observed (JAC pers. obs.) at the type locality (Fig. 2G, Plate 1B). After phonotaxis the female oriented directly behind the male and palpated the male with her antennae. Material examined. DETERMINED (n = 51): All USA, CA , Alameda Co., 1♁, Berkeley, Strawberry Canyon, 37.871593N, 122.272747W, 4-VII-1959, RW Thorp, CAS; Humboldt Co., 1♁, Scotia, 40.482357N, 124.100881W, 18-VII-1958, JR Helfer, CAS; Lake Co., in addition to type material (above), 2♁, 1♀, Anderson Springs, 38.775N, 122.69194W, 29-VII-1961, JS Buckett, BMED; 1♀, Anderson Springs, 38.775N, 122.69194W, 7-VIII-1955, JS Buckett, BMED; 1♁, Blue Lakes, 39.17111N, 123.01111W, 414 m, 30-VII-1949, HA Hurt, BMED; 1♁, SR20, mile marker 114, 39.01101N, 122.4908W, 423 m, 5-VIII-2014, JA Cole, DB Weissman, JAC sound record; Marin Co., 1♁, 2.9 km Samuel P Taylor State Park, 38.02583N, 122.72667W, 61 m, 27-IX-1981, VF Lee, CAS; 1♁, Lagunitas, 38.011312N, 122.702208W, 5-VII-2009, EC VanDyke, CAS; 1♁, 1♀, Lake Lagunitas, 37.946869N, 122.594742W, 24-VI-1958, DC Rentz, CAS; 1♁, McClures Beach, 38.187421N, 122.965276W, 15-VII-1960, JS Buckett, BMED; 1♁, same data except CAS; 1♁, same data except 2-X-1960, JS Buckett, BMED; 1♁, same data except 6-VIII-1964, JS Buckett, BMED; 2♁, Tomales Bay State Park, beach area, 38.166667N, 122.916667W, 16-VI-1978, DB Weissman, CAS; Mendocino Co., 5♁, Mendocino, 39.307674N, 123.799459W, 37 m, 1960, ER Tinkham, CAS; 1♀, same data except 10-XI-1959, JR Helfer, CAS; 1♁, same data except, 10-XI-1961, JR Helfer, CAS; 1♁, same data except 15-X-1959, JR Helfer, CAS; 1♀, same data except 15-XI-1953, JR Helfer, CAS; 1♁, 1♀, same data except 20-X-1962, JR Helfer, CAS; 1♀, same data except 22-X-1961, JR Helfer, CAS; 1♀, same data except, 3-X-1958, JR Helfer, CAS; 1♁, same data except 3-XI-1959, JR Helfer, CAS; 1♁, same data except 3-XI-1961, JR Helfer, CAS; 1♁, same data except 30-X-1959, JR Helfer, CAS; 1♀, same data except 5-XII-1958, JR Helfer, CAS; 3♁, Ukiah, 39.15017N, 123.20778W, 196 m, 2014, JN Hogue, CSUN; 1♁, 3♀, Mendocino Co., 20-VII-1923, ER Leach, CAS; Napa Co., 1♁, end of Atlas Peak Road 10 mi. airline N of Napa, 38.450743N, 122.262753W, 6-VII-1973, JA Sutro, CAS; 1♀, north side Howell Mountain, 2 miles NNE Angwin, 38.602527N, 122.435787W, 396 m, 18-X-1974, HB Leech, CAS; San Francisco Co., 5♁, Lands End, 37.787707N, 122.505528W, 17-X-1909, FX Williams, CAS; Solano Co., 1♀ nymph, Green Valley, 38.227515N, 122.151579W, 34 m, 16-VI-1953, AA Grigarick, BMED; Sonoma Co., 1♁, Bodega Bay, 38.33325N, 123.048057W, 20-VII-1974, LD & MD Anderson, UCR; QUESTIONABLE PLACEMENT (n = 20): Alameda Co., 1♀, Oakland Hills, 37.804372N, 122.270803W, 6-IX-1948, CD MacNeill, CAS; Marin Co., 1♁, McClures Beach, Pt. Reyes Peninsula, 38.187421N, 122.965276W, 2-X-1960, JS Buckett, CAS; 1♀, Mill Valley, 37.89076N, 122.523586W, 2-VIII-1925, EP VanDuzee, CAS; 1♀ nymph, same data except 24-VI-1923, EP VanDuzee, CAS; 1♀ nymph, same data except 25-VI-1925, EP VanDuzee, CAS; 1♀, same data except 26-VII-1926, EP VanDuzee, CAS; 1♀, same data except 27-VI-1925, EP VanDuzee, CAS; 1♀, same data except 5-VII-1925, EP VanDuzee, CAS; 1♀, same data except 7-XI-1925, EP VanDuzee, CAS; 2♁, Mount Tamalpais, NE slope, 37.927425N, 122.591924W, 700 m, R Hunt, BMED; Mendocino Co., 1♀, Navarro, 39.151843N, 123.541956W, 18-XII-1973, DO Clark, CSCA; 1♁, VanDamme State Park, 39.27639N, 123.77333W, 13-X-1979, T Tyler, CSCA; Napa Co., 1♀, Hwy 121, 6 mi. NW of Napa, 38.403064N, 122.4184W, no collector, CSCA; 1♀ nymph, Lake Berryessa, 38.58903N, 122.229496W, 20-V-1961, CD Macneill, CAS; Solano Co., 1♁, Vallejo, 38.104086N, 122.256637W, 18 m, 6-VIII-1956, D Haug, CSCA; Sonoma Co., 1♁ nymph, Cazadero, 38.533246N, 123.08528W, 36 m, 5-VII-1962, R Bartges, BMED; 1♀, Jenner, 38.449636N, 123.115559W, 17-X- 1937, no collector, BMED; 1♀ nymph, Kenwood, 38.4138N, 122.546094W, 126 m, 14-V-1960, TH Gantenbein, BMED; 1♀ nymph, Stillwater Cove, 38.545833N, 123.298611W, 23-V-1954, EI Schlinger, BMED; HYBRIDS WITH N. CARINATA: Napa Co., 1♀, Monticello Dam, 38.513394N, 122.104351W, 8-X-1969, F Andrews, CSCA; Solano Co., 3♁, Lake Solano County Park, 5 mi. SW of Winters off SR128, 38.49576N, 122.03399W, 51 m, 30- VII-2016, JA Cole, LACM; 12♁, 3♀, same data except 31-VII-1-VIII-2013, JA Cole, LACM; 7♁, same data except 31-VII-2014, JA Cole, DB Weissman, LACM; 2♁, 1♀, same data except 31-VII-1-VIII-2013, JA Cole, JAC; 1♀, same data except 25-VII-2019, JA Cole, JAC, Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 30-32, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Caudell, A. N. (1907) The Decticinae (a group of Orthoptera) of North America. Proceedings of the United States National Museum, 32, 285 - 410. https: // doi. org / 10.5479 / si. 00963801.32 - 1530.285"]}

Neduba steindachneri (Herman, 1874) Fig. 9 (distribution), Fig. 18 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 1E (live habitus), Plate 4I (male calling song), Plate 7D (male ventral sclerite), Plate 9I (male titillators), Plate 11I (female subgenital plate). Common name. Steindachner’s Shieldback. History of recognition. Originally described in Tropizaspis from Fox Island, Puget Sound, Washington. Caudell (1907) synonymized this species under N. carinata. This species was subsequently removed from synonymy on the basis of the wide distance between type localities, but without examination of specimens (Rentz & Birchim 1968). NEW COMBINATION: N. picturata (Scudder 1899) is transferred under the synonymy of this species, rather than N. diabolica, given the type was collected during the Northwestern Boundary Survey (Caudell 1907). This survey, from 1857-1861, was led by B.R. Kennerly, along the 409-mile-long Canadian-US border between the Rocky Mountains and Point Roberts on coastal Washington. Type material. The syntype series of males and females is in NMW Vienna. Lectotype male here chosen as the adult whose images appear, along with collection labels, on the OSFO (Cigliano et al. 2020). Label data of this specimen is as follows: Small square label: “Stein-/dachner /1869[typed]/Califor [handwritten over label] Wide rectangular label [handwritten]: Type n. sp. steindachneri/Fox Island Pouget Sound/1874 (http:// orthoptera.speciesfile.org/Common/basic/ShowImage.aspx?TaxonNameID=1141747&ImageID=208936) TOPOTYPES EXAMINED (n = 4): WA, Pierce Co., 4♁, Puget Sound, Fox Island, 47.235925N, 122.626948W, 16-VIII-1986, DB Weissman, CAS Measurements. (mm, ♁n = 20, ♀ n = 1) Hind femur ♁16.72–21.00, ♀ 19.80, pronotum total length ♁7.61–9.55, ♀ 7.56, prozona length ♁3.34–4.41, ♀ 1.94, metazona dorsal length ♁4.15–5.83, ♀ 5.62, pronotum constriction width ♁1.95–2.55, ♀ 2.60, metazona dorsal width ♁5.65–6.90, ♀ 5.50, head width ♁3.71– 4.45, ♀ 4.69, ovipositor length ♀ 14.18. Distribution. Central Oregon Cascade and Coast Ranges north to coastal British Columbia. Habitat. Understory of fir and fir-oak forests. On grasses, low branches, Ribes, and leaf litter. Seasonal occurrence. Summer through fall, from July (6-VII-1926, CL Hubbs, CAS) through September (19- IX-2015, JA Cole, LACM). Stridulatory file. (n = 11) length 3.5–4.0 mm, 108–127 teeth, tooth density 31.3 ± 2.3 (27–35.3) teeth/mm. Song. (n = 26) Male songs are of the zwee-zwee type with PTR 3.7 ± 0.4 s- 1, faster than all other Convexa Clade species except some males of N. radicata. Males sometimes add partial PT when initiating song bouts. MPTL is 250.4 ± 47.6 ms. The OPT is poorly developed or absent. PTF is 13.1 ± 1.7 kHz. Males are nocturnal singers. Karyotype. (n = 5) 2n♁ = 26 (2m + 22t + XtXt) T86-74, S86-95, topotype. Recognition. The male ventral sclerite is mushroom-shaped with a short, broad shaft and a wide, evenly curved apex with a short, blunt, recurved anterolateral projection. Tubercles are large, sparse, and confined to the apex. The titillators of N. steindachneri begin curvature after the midpoint of the shaft and are not as strongly curved dorsally (>90 o) as seen in most other Carinata Group species (Plate 9). Stridulatory file tooth density (27–35 teeth/mm) is lower than that of N. convexa (2-sample t -test, P = 5.48 ×10 -3) and also trends lower than N. cascadia (33–39 teeth/ mm). The female subgenital plate is pentagonal, flat, subequal in length and width, and with curved apical margins and a strongly pointed apex, which separates this species from N. cascadia females which have straight apical margins and from N. convexa and N. longiplutea females which have plates that are longer than wide. The song is unique in having reduced or absent OPT. The PTR is faster than all other convexa clade species. This species has the most northern range of any Neduba (Fig. 9). Notes. N. steindachneri is locally common in the forest understory of the western slopes of Oregon and Washington Cascade Range and primarily the eastern slopes of the Oregon and Washington coastal ranges. A phylogeographic break separates populations north and south of the Willamette River (Figs. 3–4) that suggests historical isolation that indicates historical isolation. Hybridzation with N. cascadia is possible and should be sought from populations in central Oregon, where the ranges of these two species overlap. Material examined. (n = 28) USA, OR, Benton Co., 8♁, McDonald-Dunn Forest, Corvallis, 44.60382N, 123.33253W, 144 m, 18-IX-2015, JA Cole, LACM; 1♁, same data except JAC; 5♁, same data except 19-VII-1982, DC Lightfoot, CAS; Columbia Co., 1♀, Wilson Creek, 46.07701N, 123.02315W, EH Nast, CAS; Hood River Co., 1♁, 1♀, Hood R., 45.70556N, 121.52028W, 122 m, 17-VII-1931, RH Beamer, CAS; Marion Co., 1♁, Sublimity, Silver Creek Falls, 45.000267N, 122.840916W, 167 m, 19-VIII-1945, W Blehm, CAS; Multnomah Co., 3♁, Dabney State Park, 45.51766N, 122.35429W, 22 m, 19-IX-2015, JA Cole, JAC; 5♁, Oxbow Regional Park Day Use Area, SE Gordon Creek Rd., 1.2 mi. SE jct. Hurlburt Rd., 45.49821N, 122.27871W, 59 m, 19-IX-2015, JA Cole, JAC; 1♁ same data except JAC; 1♁, SE Gordon Creek Rd., 2.7 mi. S jct. Hurlburt Rd., 45.4784N, 122.28107W, 257 m, 19-IX-2015, JA Cole, JAC sound record; Tillamook Co., 1♁, Tillamook, 45.51667N, 123.71667W, 30- VII-1926, CL Hubbs, CAS; WA, Kitsap Co., 1♁, Holly, 47.557593N, 122.978488W, 3 m, 6-VII-1926, CL Hubbs, CAS; 5♁, Margarete Ranch, Port Orchard, 47.47137N, 122.65009W, 1525 m, 13-14-VIII-2013, JA Cole, LACM; 2♁, same data except JAC; Mason Co., 1♁, 1♀, Lake Cushman, 47.487964N, 123.266471W, 244 m, 27-VIII-1919, P Putnam, CAS, Pierce Co., see Type material (above). QUESTIONABLE PLACEMENT: OR, Douglas Co., 1♁ nymph, Drain, 43.658731N, 123.318699W, 9-VI-2014, EC VanDyke, CAS. Material from databases not examined by us (n = 7): CANADA, BC, Departure Bay, 49.20, 123.98W, 1925, GJ Spencer, Symbiota Collections of Arthropods Network (SCAN, https://scan-bugs.org/portal); UBC Campus 49.26N, 123.25W, 3-X-1946, D Evans, SCAN; Gailano Island, 48.93N, 123.45W, 4-IX-1971, J Scudder, SCAN; same data except 1971, J Scudder, SCAN; same data except 11-X-1976, GGE Scudder, SCAN; same data except 1976, GGE Scudder, SCAN; Vancouver Island, 49.66667N, 125.83333W, 9-VIII-2007, J Miskelly, SINA. Propsti Group The Propsti Group contains a single species, the Santa Catalina Island endemic (Fig. 8) N. propsti. The Group is defined by having a single spine on the fore tibia located on the posterior margin, a pair of prosternal spines, darkened male tegminal apices, and a female subgenital plate that is over half again as long as wide. Only N. lucubrata from mainland California shares this suite of characters, but this species has a smaller body size, a male subgenital plate that narrows from base to apex along the lateral carinae, and a song that consists of a bout (chirp) structure (Plate 5C) whereas the song of N. propsti is continuous (Plate 4J). The karyotype is unique. Propsti Group The Propsti Group contains a single species, the Santa Catalina Island endemic (Fig. 8) N. propsti. The Group is defined by having a single spine on the fore tibia located on the posterior margin, a pair of prosternal spines, darkened male tegminal apices, and a female subgenital plate that is over half again as long as wide. Only N. lucubrata from mainland California shares this suite of characters, but this species has a smaller body size, a male subgenital plate that narrows from base to apex along the lateral carinae, and a song that consists of a bout (chirp) structure (Plate 5C) whereas the song of N. propsti is continuous (Plate 4J). The karyotype is unique., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 44-46, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Herman, O. (1874) Die Decticiden der Brunner von Wattenwyl'schen Sammlung. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 24, 191 - 210.","Caudell, A. N. (1907) The Decticinae (a group of Orthoptera) of North America. Proceedings of the United States National Museum, 32, 285 - 410. https: // doi. org / 10.5479 / si. 00963801.32 - 1530.285","Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Scudder, S. H. (1899) Two genera of North American Decticinae. Proceedings of the American Academy of Arts and Sciences, 35, 81 - 93. https: // doi. org / 10.2307 / 25129903","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0."]}

Neduba propsti Rentz & Weissman, 1981 Fig. 19 (distribution), Fig. 20 (male and female habitus, calling song, drumming, male and female tremulation karyotype), Plate 2A (live habitus), Plate 4J (male calling song), Plate 7F (male ventral sclerite), Plate 10A (male titillators), Plate 11J (female subgenital plate). Common name. Santa Catalina Island Shieldback. History of recognition. Described as a Santa Catalina Island endemic (Rentz & Weissman 1981). Listed under Aglaothorax in OSFO for unspecified reasons (Cigliano et al. 2020). Type material. The holotype male is in the CAS collection. Images of the holotype are available at OSFO (Cigliano et al. 2020). Measurements. (mm, ♁n = 9, ♀ n = 5) Hind femur ♁20.86–23.47, ♀ 24.10–25.89, pronotum total length ♁8.30– 9.45, ♀ 9.15–9.89, prozona length ♁4.15–5.55, ♀ 5.15–5.93, metazona dorsal length ♁3.90–4.90, ♀ 3.88– 4.15, pronotum constriction width ♁2.44–3.13, ♀ 2.98–3.22, metazona dorsal width ♁6.37–7.01, ♀ 6.27–7.22, head width ♁4.25–4.95, ♀ 5.18–5.54, ovipositor length ♀ 18.04–19.14. Distribution. Restricted to Santa Catalina Island, Los Angeles County, California, USA. Probably found throughout the island in suitable habitat. Habitat. Dense chaparral vegetation, often on steep hillsides and in canyon bottoms. Individuals prefer to sit on the inner twigs of tangles. Also, in gardens on non-native vegetation. One individual taken from ornamental eucalyptus (JAC, pers. obs.). Seasonal occurrence. Adult records from mid-June (14-VI-1985, S Bennett, CAS) through July (28-VII-1981 DB Weissman, CAS). Stridulatory file. (n = 5) length 3.2–3.9 mm, 94–111 teeth, tooth density 30.4 ± 2.1 (28.2–33.0) teeth/mm. Song. (n = 10) Continuous 200 ms MPTL at a brisk PTR of 4.4 ± 0.4 s- 1. PT consist of the least amount of pulses (toothstrikes) of any species (~20). PTF approaches the ultrasonic at 18.5 ± 3.5 kHz. A captive male drummed at irregular intervals while stridulating (Fig. 20); the drums were audible and induced considerable substrate vibration in the cage. Karyotype. (n = 4) Unique. 2n♁ = 24 (4m + 18t + XmYt). T85-12, S85-70, topotype. Recognition. Shares the following morphological characters with N. lucubrata: a single apical spine on the fore tibiae, prosternal spines, and tegmina darkened apically. The stridulatory file has a lower tooth density (28–33 teeth/ mm) than any other species except those of the Castanea Group. Male genitalia of N. propsti are similar to those of N. castanea and N. macneilli, but the arms of the titillators of N. propsti have a shaft that is straight and not swollen at a distance of 1/6 from the base as in the latter two species (Plate 10). The song is unique in having short MPTL produced continually at a rapid PTR. Females have the longest subgenital plate of any species, approximately 1.5 times longer than wide. This is the only nedubine on Santa Catalina Island (Figs. 8, 19) with the most southerly distribution of any Neduba. Notes. N. propsti is an early branching lineage (Figs. 3–5) that has apparently been isolated on Santa Catalina Island for a long time. The island has never been connected to mainland California (Legg et al. 2004) being the product of tectonic uplift. Males may be wary and cease calling at the slightest disturbance, as much as a single leaf falling, and jump with little provocation (JAC pers. obs.). This is one of a handful of Neduba species that drum (Weissman 2001; see also N. castanea, N. macneilli, and N. lucubrata below). Drumming was observed in captivity by a single male without a female present (JAC pers. obs) and not in the field; the context of drumming in the mating system is not known in this species. Material examined. (n = 14) All USA, CA, Los Angeles Co.: 3♁, Hermit Gulch Campground, Avalon Canyon, Santa Catalina Island, 33.38265N, 118.33951W, 91 m, 9-10-VII-2013, JA Cole, LACM; 1♁ same data except JAC; 2♀, Santa Catalina Island, 33.383361N, 118.417576W, 1-VII-1983, DB Weissman, CAS; 1♀, same data except 20-VII-1982, S Bennett, CAS; 1♀, same data except 28-VII-1981, S Bennett, LACM; 1♁, same data except 30-VI-1973, DB Weissman, CAS; 4♁, Santa Catalina Island, Toyon Bay, 33.383N, 118.416W, 14-VI-1985, S Bennett, CAS; 1♀, same data except 5-VII-1986, S Bennett, CAS. Castanea Group The Castanea Group is readily recognizable on account of the robust habitus, the short hind femora, by having only one spine on the anterior margin of the fore tibiae, and by the lack of prosternal spines. The lateral carinae of the male subgenital plate converge apically, although not as dramatically as in the Sierranus and Sequoia Groups, and the styli vary from articulate to rudimentary to absent. The posterior margins of the abdominal tergites have only slight crenulations. Superficially the species of this group resemble sympatric Aglaothorax ovata. Karyotypes also separate the two species in this group from all other Neduba. Castanea Group species occupy the dry slopes of central and southern California mountain ranges (Fig. 8).

Neduba convexa Caudell, 1907 Fig. 9 (distribution), Fig. 16 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 4G (male calling song), Plate 7 A–C (male ventral sclerites), Plate 9H (male titillators), Plate 11G (female subgenital plate). Common name. Convex Shieldback. History of recognition. This taxon was described as a variety of N. carinata (Caudell 1907). The type series included specimens from widely separated localities in California: Mount Shasta and Napa County. We resolved the taxonomy in relation to topotypes from the Mount Shasta type locality designated by Rentz & Birchim (1968). The North Coast Range type material from Napa County belongs to N. ambagiosa. (see species account above). Type material. The type specimen is a lectotype male from Mt. Shasta, Siskiyou Co., in the USNM. Images of the type are available at OSFO (Cigliano et al. 2020). TOPTYPES EXAMINED (n = 19): USA, CA, Siskiyou Co., 7♁, McBride Camp, 4 mi. E of Mount Shasta City on Rd A10, 41.409294N, 122.117727W, 1433 m, 2-IX-1983, DB Weissman, CAS; 1♁, same data except 29-VI-1992, DB Weissman, CAS; 7♁, McBride Springs Campground, 4 mi. NE of Mount Shasta City, 41.3529N, 122.28328W, 1487 m, 1-2-VIII-2013, JA Cole, LACM; 3♁, same data except JAC; 1♁, same data except 22-25-VIII-2019, JA Cole, JAC. Measurements. (mm, ♁n = 27, ♀ n = 5) Hind femur ♁18.38–20.60, ♀ 21.42–24.22, pronotum total length ♁8.50–10.36, ♀ 7.78–9.50, prozona length ♁2.95–4.45, ♀ 3.39–4.60, metazona dorsal length ♁5.03–6.55, ♀ 4.39– 5.17, pronotum constriction width ♁2.35–3.06, ♀ 2.57–3.04, metazona dorsal width ♁5.38–6.95, ♀ 5.55–6.05, head width ♁4.06– 4.87, ♀ 5.03–5.80, ovipositor length ♀ 15.80–21.19. Distribution. Widely distributed in the northern Central Valley and Cascade Range of Northern California, extending east along the junction of the Sierra Nevada and Cascade Range to the Diamond Mountains. Habitat. Understory vegetation, tangles, and leaf litter of chaparral and mixed conifer forests. Also, in remaining northern Central Valley riparian forests. Seasonal occurrence. Specimens suggest a long summer and fall adult activity from July (1-VII-2000, JA Cole, LACM) through November (10-XI-1948, HP Chandler, CAS). Nymphs have been collected from early April through late August. Adult activity will commence earlier in the lower elevations of the Central Valley. Stridulatory file. (n = 8) length 3.1–3.9 mm, 106–145 teeth, tooth density 35.3 ± 2.8 (29.7–38.2) teeth/mm. Song. (n = 14) The male song is of the “zwee-zwee type with PTR 2.6 ± 0.3 s- 1 at 25 oC, MPTL 308.1 ± 51.1 ms, and PTF 14.0 ± 0.8 kHz. OPT are well developed. A distinct amplitude increase commences at the middle of each MPT. Males are nocturnal and may sing in aggregations. Karyotype. (n = 15) 2n♁ = 26 (2m + 22t + XtYt). T83-48, S83-119, topotype. Recognition. N. convexa can be confused with N. longiplutea distributed to the west, but the latter has a male ventral sclerite lateral process that is longer than the shaft, unlike the reverse condition in the former. To the south and north of N. convexa are N. radicata and N. cascadia, respectively, both of which have small lateral process on the male ventral sclerite. N. ambagiosa is also similar but males usually have a more conical ventral sclerite apex and that is a Coast Range species. Females may often be separated from N. longiplutea by the lack of a bifurcate subgenital plate apex, but this is variable in N. convexa. The N. radicata subgenital plate is wider than long with a deep notch, and that of N. cascadia does not have curved apical margins. The song of N. radicata to the south has a faster PTR, whereas the song of N. cascadia to the north has a slower PTR. Notes. This species has a wide range and can be locally abundant. N. convexa was probably widespread in the Central Valley riparian forests before modification of the region for agriculture. Local variation is evident. Males from the populations along the Feather River Drainage and Laufman Campground in the Diamond Mountains tend to have flatter ventral sclerite apices (Plate 7C) and are genetically distinct (Figs. 3–5), but these populations are not reliably separable using any character. Further study may conclude that cryptic species are present along this east-west transect across the northern rim of the Central Valley and the Sierra Nevada. The physiology of this species is apparently adapted for activity at cold temperatures. On Mount Shasta, males sang between 10–15 oC. When brought into the laboratory at standard temperature, individuals became hyperactive and did not survive long (JAC pers. obs.). Material examined. (n=88) All USA, CA, Butte Co., 1♀ nymph, Chico, 39.728494N, 121.837478W, 59 m, 7-V-1968, T Kone, B Wilkey, W Wiard, CSCA; 6♁, Sacramento River State Park, Indian Fishery Day Use Area, 39.70522N, 121.93905W, 42 m, 18-VII-2015, JA Cole, DB Weissman, LACM; Lassen Co., 3♁, 2♀, Dyer Mountain Rdg., 40.239055N, 121.03246W, 2279 m, 27-VIII-1968, FL Blanc, CSCA; 1♁, Laufman Campground, Plumas National Forest, 3.5 mi. S of Milford on Milford Grade Rd., 40.135N, 120.3483W, 1554 m, 13- 14-VIII-2002, JA Cole, LACM; 1♁, same data except JA Cole, JAC; 2♁, same data except 18-VII-2012, JA Cole, LACM; 5♁, same data except 8-VIII-2014, JA Cole, DB Weissman, LACM; Plumas Co., 1♁, Almanor, 40.217386N, 121.17413W, 1377 m, 16-VII-1940, no collector, CAS; 2♁, Hallsted Campground, 0.25 mi. W of Twain on SR 70, 40.0174N, 121.0745W, 864 m, 1-4-VII-2000, JA Cole, LACM; 3♁, same data except 13-16-VII- 2004, JA Cole, LACM; 1♁, 2♀, same data except JAC; 1♁, same data except 15-17-VIII-1997, JA Cole, LACM; 4♀, same data except 7-10-IX-2001, JA Cole, LACM; 1♁, same data except 15-17-VIII-1998, JA Cole, LACM; 1♁, Johnsville, 39.76073N, 120.695498W, 1573 m, 16-VII-1973, RA Belmont, BMED; 1♁, same data except 9-VIII-1961, JS Buckett, BMED; 1♁, same data except 25-VIII-1961, JS Buckett, BMED; 1♁, same data except 9-X-1974, H Pini, BMED; 1♀, Meadow Valley, 39.929612N, 121.060791W, 1153 m, 30-X-1954, B Forbs, BMED; 2♁ adults, 1♁ nymph, same data except 5-VII-1924, EC Van Dyke, CAS; 1♀, Quincy, 39.936836N, 120.947176W, 27-IX-1967, J Badaj, CSCA; Shasta Co., 1♁, Bridge Bay Rd., yacht area, 40.75611N, 122.32167W, 4-VIII-1980, no collector, CAS; 1♁, Burney Fall, 41.010716N, 121.652765W, 853 m, 10-XI-1948, HP Chandler, CAS; 1♁, Crystal Lake, 40.934999N, 121.556861W, 1585 m, 2-IX-1953, HP Chandler, CAS; 1♀ nymph, Hat Creek, 40.829684N, 121.508994W, 1003 m, 10-VII-1952, GF Pronin, CAS; 2♁, same data except 11-VIII-1964, RE Pinger, CSCA; 1♀, same data except 19-VIII-1951, no collector, CAS; 1♀ nymph, same data except 20-VIII-1951, GF Pronin, CAS; 1♁, 4♀, same data except 22-VII-1965, S Seminoff, CSCA; 1♀ nymph, same data except 23-VI-1951, GF Pronin, CAS; 1♁, same data except 25-VIII-1951, GF Pronin, CAS; 1♀ nymph, Iron Mountain Mine, 40.675427N, 122.52807W, 305 m, 30-VII-1947, HP Chandler, CAS; 1♁, Lamoine, 40.977927N, 122.430847W, 377 m, 2-IX- 1970, RE Whipp, CSCA; 1♀, Whiskeytown, 40.63876N, 122.559737W, 390 m, 10-X-1976, TR Haig, CSCA; Siskiyou Co., in addition to type material (above), 1♀, Castle Lake, 41.227294N, 122.383254W, 1554 m, IX-1953, HP Chandler, CAS; 1♀, Dunsmuir, 41.208209N, 122.271953W, 698 m, 7-VII-1970, V Pierce, CSCA; 1♀, Specimen Gulch, 41.13986N, 123.124485W, 25-VII-1967, AD & GJ Keuter, CAS; Tehama Co., 1♁ nymph, 7 mi. NE Red Bluff, 40.25019N, 122.141936W, 4-IV-1961, T Gallion, CSCA; 11♁, 1♀, Potato Patch Campground, Lassen National Forest, 21.5 mi. SW of Chester off SR36 and SR32, 40.1894N, 121.5315W, 1128 m, 19-20-VII-2012, JA Cole, LACM; 4♁, same data except JAC; ♁3, ♀ 2, same data except 7-VIII-2014, JA Cole, DB Weissman, CAS., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on page 40, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Caudell, A. N. (1907) The Decticinae (a group of Orthoptera) of North America. Proceedings of the United States National Museum, 32, 285 - 410. https: // doi. org / 10.5479 / si. 00963801.32 - 1530.285","Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0."]}

Neduba diabolica (Scudder, 1899) Fig. 9 (distribution), Fig. 13 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 4D (male calling song), Plate 6F (male ventral sclerite), Plate 9E (male titillators), Plate 11D (female subgenital plate). Common name. Mount Diablo Shieldback. History of recognition. Originally described in Tropizaspis from a female taken at “Monte Diablo, California ” (Scudder 1899). Rentz & Birchim (1968) assigned the majority of South Coast Range Neduba populations to this species and synonymized variety picturata (Scudder 1899) under this name. We reassign all other South Coast Range populations to N. carinata. The type of picturata was collected during the Northwestern Boundary Survey (Caudell 1907), which makes picturata a synonym of N. steindachneri (see discussion under that species) and not N. diabolica. Type material. The female holotype is housed in ANSP. Type not examined by us but images available at OSFO (Cigliano et al. 2020). TOPOTYPES EXAMINED (n = 23): USA, CA, Contra Costa Co., 1♀, Mount Diablo State Park, 37.862703N, 121.93107W, 1021 m, 1-IX-2002, DB Weissman, CAS; 3♁, Mount Diablo State Park, 37.862703N, 121.93107W, 670-1160 m, 30-VI-1990, DB & DD Weissman, CAS; 3♁, Mount Diablo State Park, Fire Interpretive Trail, 37.8807N, 121.9172W, 1131 m, 20-21-VII-2005, JA Cole, JF Eguizabal, LACM; 1♁, Mount Diablo State Park, Juniper Camp, 37.8785N, 121.9339W, 886 m, 27-28-VI-2008, JA Cole, W Chatfield-Taylor, W Ericson, JAC; 1♁, 1♀, same data except LACM; 1♁, Mount Diablo State Park, Juniper Camp, 37.862703N, 121.93107W, 886 m, 28-VIII-1982, DB Weissman, CAS; 1♁, 1♀, Mount Diablo State Park, Lookout Point, 37.862703N, 121.93107W, 7-VI-1967, DC & KA Rentz, CAS; 2♁, Mount Diablo State Park, near North Gate Entrance, 37.862703N, 121.93107W, 229 m, 13-VII-1982, DB Weissman, CAS; 1♁, 2♀, Mount Diablo State Park, summit, 37.862703N, 121.93107W, 1021 m, 11-VIII-1990, DB Weissman & DW Weissman, CAS; 1♁, same data except 13-VII-1982, DB Weissman, CAS; 2♁ nymphs, 1♀ nymph, Russelman Park, Mount Diablo East Slope, 19- IV-1931, EP VanDuzee, CAS; 1♁ nymph, same data except 24-IV-1932, EC VanDyke, CAS. Measurements. (mm, ♁n = 8, ♀ n = 3) Hind femur ♁20.09–22.49, ♀ 21.80–23.09, pronotum total length ♁9.50– 11.02, ♀ 9.40–9.98, prozona length ♁4.38–4.92, ♀ 4.81–5.34, metazona dorsal length ♁4.86–6.10, ♀ 4.59–4.75, pronotum constriction width ♁2.55–3.21, ♀ 2.55–3.10, metazona dorsal width ♁6.30–6.86, ♀ 5.89–6.05, head width ♁4.60–5.95, ♀ 4.95–5.39, ovipositor length ♀ 16.47–18.04. Distribution. Known only from Mount Diablo at the north end of the Diablo Range, California. Not known from Mt. Hamilton, also in the Diablo Range, some 67 km southeast of Mt. Diablo where N. oblongata occurs. The Vallecitos Valley is a possible biogeographic break between these two species (See N. oblongata species account above). Habitat. Oak woodland and chaparral habitats. Males call from the interior of large bushes, especially in canyon bottoms. Seasonal occurrence. Adults have been taken from late June (27-VI-2008, JA Cole, W Chatfield-Taylor, W Ericson, LACM) to September (1-IX-2002, DB Weissman, CAS). Nymphs were collected 7-VI-1967 (Rentz & Birchim 1968). Stridulatory file. (n = 4) length 3.0– 3.5 mm, 132–145 teeth, tooth density 43.2 ± 4.5 (37.7–48.3) teeth/mm. Song. (n = 12) The song of N. diabolica consists of bouts of uniform rate “lisping” as in N. carinata. PTR 10.8 ± 0.7 s- 1 is identical to N. carinata. PTF 13.3 ± 2.3 kHz; a high frequency lab recording measured PTF at 19.2 kHz. PTdc 80.4 ± 8.0% is significantly higher than that found in N. carinata songs (ANOVA, P = 2.36 ×10 -3). PTdc is a temperature-invariant song character (linear regression, P = 0.188), and thus can be compared among recordings that lack temperature control. Males are nocturnal singers. Bout length is variable, but the bouts of N. diabolica males tend to be shorter and more even in length than those of N. carinata males. Karyotype. (n = 4) 2n♁ = 26 (2m + 22t + XtYt) T90-12, S90-61, topotype. This corrects the information of Ueshima and Rentz (1979). Recognition. The high stridulatory file tooth density (37–48) is shared only with N. carinata in the Carinata Group. The ventral sclerite is narrow with a high convex to pyramidal apex and a poorly developed anterolateral process. In contrast, ventral sclerites of N. oblongata have a low convex apex and a minute lateral process. Except for N. radicata, all other Convexa Clade taxa have long anterolateral processes. The female subgenital plate is pentagonal and flat as in N. carinata and N. oblongata, but those two species lack a distinct medial groove, which is present in this species. Songs of N. diabolica males are qualitatively similar to those of N. carinata but have a higher duty cycle. The distribution is restricted to the vicinity of Mount Diablo, California. Notes. This species exhibits a mosaic of characters. DNA places N. diabolica with the Convexa Clade, the song type and stridulatory file are like those of N. carinata, and the genitalia resemble those of N. radicata. Mount Diablo has remained above sea level since the early Miocene (Bartow 1991) and lies at the junction of the distributions of the southern Carinata and northern Convexa Clades. The mixture of characters found in N. diabolica suggests past introgression. By maintaining its species status, we recognize that N. diabolica is a phylogenetically distinct, philopatric lineage that inhabits an ancient region of high endemism and provided it remains protected (Mount Diablo is a California State Park), this lineage may persist into the future with its mixture of characters. The name picturata Scudder was originally described to differentiate insects with a mottled color pattern as opposed to uniform or striped coloration. These color pleomorphisms are shared by all species in the Carinata Group. Material examined. See Type material above., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 33-35, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Scudder, S. H. (1899) Two genera of North American Decticinae. Proceedings of the American Academy of Arts and Sciences, 35, 81 - 93. https: // doi. org / 10.2307 / 25129903","Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Caudell, A. N. (1907) The Decticinae (a group of Orthoptera) of North America. Proceedings of the United States National Museum, 32, 285 - 410. https: // doi. org / 10.5479 / si. 00963801.32 - 1530.285","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0.","Ueshima, N. & Rentz, D. C. (1979) Chromosome systems in the North American Decticinae with reference to Robertsonian changes. Cytologia, 44, 693 - 714. https: // doi. org / 10.1508 / cytologia. 44.693","Bartow, J. A. (1991) The Cenozoic Evolution of the San Joaquin Valley, California. United States Geological Survey, United States Government Printing Office, Washington, 40 pp. https: // doi. org / 10.3133 / pp 1501"]}

Neduba oblongata Cole, Weissman, & Lightfoot sp. n. Fig. 9. (distribution), Fig. 11 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 4B (male calling song), Plate 6E (ventral sclerite), Plate 11B (female subgenital plate). Common name. Mount Hamilton Shieldback. History of recognition. None. Type material. HOLOTYPE MALE: USA, CA, Santa Clara Co., Mount Hamilton, 2.6 road mi. W observatory, 37.341883N, 121.643002W, 1036 m, 12-VI-1982, DB Weissman, S 82-23 [stop], R82-47 [recording], T82-2 [karyotype], 127 [teeth] 3.5 [mm stridulatory file length], deposited at CAS, Entomology type #19713. PARATYPES (n=10): USA, CA, Santa Clara Co., 1♁, Mount Hamilton, 0.3 mi. W observatory, 37.341883N, 121.643002W, 1260 m, 6-VII-1997, DB Weissman, CAS; 3♁, Mount Hamilton, 15 mi. W observatory, 37.341883N, 121.643002W, 488 m, 12-VI-1982, DB Weissman, CAS; 3♁, 1♀, Mount Hamilton, 2.6 mi. W observatory, 37.341883N, 121.643002W, 1036 m, 12-VI-1982, DB Weissman, CAS; 2♁, Mount Hamilton, 6.5 mi. W observatory, 37.32864N, 121.65729W, 686 m, 8-VIII-2015, DB & DW Weissman, LACM Measurements. (mm, ♁n = 8, ♀ n = 1) Hind femur ♁18.44–20.61, ♀ 22.01, pronotum total length ♁8.65–11.01, ♀ 9.66, prozona length ♁3.65–5.21, ♀ 4.95, metazona dorsal length ♁4.80–6.27, ♀ 4.71, pronotum constriction width ♁2.50–3.14, ♀ 3.55, metazona dorsal width ♁6.00–7.39, ♀ 6.43, head width ♁4.26–4.85, ♀ 5.18, ovipositor length ♀ 18.58. Distribution. Known only from Mount Hamilton in the Diablo Range, Santa Clara County, California. Habitat. Oak woodland and chaparral. Seasonal occurrence. Available records indicate adult activity extends throughout the summer from mid-June (12-VI-1982, DB Weissman, CAS) through mid-August (8-VIII-2015, DB & DW Weissman, CAS). Stridulatory file. (n = 4) length 2.8–3.5 mm, 114–134 teeth, tooth density 39.1 ± 2.3 (36.3–41.3) teeth/mm. Song. (n = 9) Unique. PTR is bimodal and switches between a slow rate 7.45 ± 0.59 s- 1 and a fast rate of 11.2 ± 1.28 s- 1. Karyotype. (n = 4) 2n♁ = 26 (2m + 22t + XtYt), S82-23, T82-2, paratopotype. Recognition. Stridulatory file, male genitalia, song, and geography. A low stridulatory file tooth density separates this species (36–41 teeth/mm) from N. carinata, which has a significantly higher density (38–55 teeth/mm; two-sample t -test, P = 0.007). The ventral sclerite is robust with a straight shaft, low convex apex, and a minute lateral process. Other Carinata Group species have a longer lateral process, and the shaft of Convexa Clade species is curved. Santa Lucia Mountains N. carinata males may have a minute lateral process but the whole sclerite is not as robust as in N. oblongata. The ventral sclerite of N. diabolica, which inhabits the same mountain range, has a more conical apex, a longer lateral process, and a curved shaft. The calling song is unique among Carinata Group species in having a bimodal PTR. The female subgenital plate is pentagonal and flat, identical to N. carinata, both of which are separated from N. diabolica by the lack of a medial groove. This species is geographically restricted to the Mount Hamilton vicinity in the Diablo Range of California. Etymology. l. oblongata oblong, referring to the fusiform habitus and enlarged, oval pronotum. Notes. The two highest peaks in the Diablo Range, Mount Hamilton (1284 m) and Mount Diablo (1173 m) are mountain habitat islands separated by a mere 67 km, yet each harbors a distinct Carinata Group species (see N. diabolica below). This contrasts with a single (albeit variable) species, N. carinata, distributed across 200 km of the South Coast Ranges. The Santa Clara Valley is thus implicated as an isolating barrier between the Diablo Range and the Coast Ranges, while the Vallecitos Valley is a potential biogeographic break between populations in the north and south of the Diablo Range. The bimodal PTR in the male calling song is reminiscent of the songs of Sierranus Group species (see below). Given the evolutionary distance between the Carinata Group and the Sierranus Group (Figs. 3–5), the mostly likely explanation for shared bimodal PTR is convergent evolution. A preexisting receiver response may exist in Neduba females (e.g. Basolo 1996; Ryan & Rand 1999) that selects for male songs with an elaborate pattern or additional OPT. Material examined. Type series only. See Type material above., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 28-30, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Basolo, A. L. (1996) The phylogenetic distribution of a female preference. Systematic Biology, 45, 290 - 307. https: // doi. org / 10.1093 / sysbio / 45.3.290","Ryan, M. J. & Rand, A. S. (1999) Phylogenetic influence on mating call preference in female tungara frogs, Physalaemus pustulosus. Animal Behaviour, 57, 945 - 956. https: // doi. org / 10.1006 / anbe. 1998.1057"]}

Neduba prorocantans Cole, Weissman, & Lightfoot, sp. n. Fig. 19 (distribution), Fig. 28 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 3 D���E (live habitus), Plate 5H (male calling song), Plate 8E (male ventral sclerite), Plate 10H (male titillators), Plate 12F (female subgenital plate). Common name. Incessant Shieldback History of recognition. Records from Dougherty Creek, Kern River Canyon, CA (Rentz & Birchim 1968) were confused with N. sierranus. Type material. HOLOTYPE MALE: USA, CA, Kern Co., Tehachapi, Water Canyon Rd., 0.15 mi. S of intersection with Highline Rd., 35.307739N, 118.021738W, elev. 1463 m, 28-VIII-1983, DB Weissman, S 83-115, R83- 295 [recording], T83-46 [karyotype], 150 [teeth], 3.3 [mm file count], tegmen in gelcap and genitalia in vial below specimen, deposited at CAS, Entomology type #19681. PARATYPES (n = 30): USA, CA, Kern Co., 3���, 1♀, same data as holotype; 6���, Hobo Campground overflow area, 3 miles west of Bodfish on Kern River Canyon Road, 35.5752N, 118.5305W, 700 m, 23-VI-2003, JA Cole, LACM; 1���, horse trough spot, Tejon Ranch Conservancy, 34.97927N, 118.69159W, 1223 m, 3-VI-2017, L Pavliscak, LACM; 1���, Kern River Rd. 1.1 mi. from jct. Caliente-Bodfish Rd., 35.59429N, 118.5141W, 1470 m, 21- VII-2015, JA Cole, DB Weissman, LACM; 1���, Kernville, 37 Sierra Way, 35.7561N, 118.4203W, 828 m, 26-28-VII- 2002, JA Cole, JAC; 2���, same data except LACM; 2���, same data except 27-VII-2004, JA Cole, LACM; 1���, same data except 28-VII-2004, JA Cole, JAC; 1♀, Lopez Flat, Tejon Ranch Conservancy, 34.94264N, 118.63381W, 816 m, 3-VI-2017, JA Cole, K Halsey, LACM; 3���, 1♀, Paradise Valley, Tehachapi Mountains, 34.91664N, 118.66759W, 7-VIII-1931, ER Tinkham, CAS; 1���, Tehachapi Mountain Park, 35.06861N, 118.4825W, 1470 m, 20-VII-2015, JA Cole, DB Weissman, LACM; 2���, Tehachapi Mountain Park, 35.06861N, 118.4825W, 1707 m, 28-VIII-1983, DB Weissman, CAS; 1♀, Tejon Ranch Conservancy, Indian Schoolhouse, Tejon Creek, 35.04512N, 118.67052W, 647 m, 1-IV-2018, K Moore, LACM; 4���, Water Canyon Rd., 1.4 mi. below entrance to Tehachapi Mountain Park, 35.08258N, 118.49486W, 1571 m, 20-VII-2015, JA Cole, DB Weissman, LACM. Measurements. (mm, ���n = 14, ♀ n = 2) Hind femur ���19.60���22.59, ♀ 22.55���23.50, pronotum total length ���8.25���10.15, ♀ 9.25���9.65, prozona length ���3.83���5.09, ♀ 4.54���5.65, metazona dorsal length ���4.07���5.35 ♀ 4.00��� 4.71, pronotum constriction width ���2.44���3.50, ♀ 3.24���3.29, metazona dorsal width ���4.99���6.80, ♀ 5.60���5.89, head width ���4.70���5.57, ♀ 5.47���5.60, ovipositor length ♀ 8.75���9.41. Distribution. West slope of the southern Sierra Nevada south of the Tule River watershed, south to the north slope of the Tehachapi Mountains. Habitat. Forest understory, often in thick tangles of vegetation. Taken from poison oak, willow (Salix spp.), and thorny brambles in riparian areas. Nymphs were collected from under bark of fallen tree limbs and from leaf litter under a valley oak tree (Quercus lobata N��e). Museum specimens (ER Tinkham, CAS) also indicate association with oaks. Seasonal occurrence. Adults collected in the field from late June (23-VI-2003, JA Cole, LACM) through August (28-VIII-1983, DB Weissman, CAS). Last instar nymph collected in early June (3-VI-2017, LA Pavliscak, LACM) matured 19-VI-2017. Stridulatory file. (n = 5) length 3.3���3.8 mm, 150���183 teeth, tooth density 49.1 �� 2.7 (45.5���52.4) teeth/mm. Song. (n = 12) A continuous train of alternating MPT and OPT delivered at PTR 2.4 �� 0.3 s- 1. The PTdc is 62.1 �� 9.7%, which is lower than that of N. sequoia (ANCOVA, P = 3.75��10 -4). The PTF is high at 15.6 �� 1.2 kHz. Karyotype. (n = 7) 2n��� = 21 (2m + 16t + XtXtYm). T83-44, S83-115, type locality. Recognition. This is the most morphologically distinctive species of the Sierranus and Sequoia Groups. Males combine a weakly constricted pronotum and a low stridulatory file tooth density (below 53 teeth/mm), lower than all other Sierranus and Sequoia Group species except N. duplocantans (47���52 teeth/mm) which has a more strongly constricted pronotum. Females have the shortest ovipositor of any Sierranus or Sequoia Group species, 10 mm or less in length. The song has a lower PTdc than that of the related N. sequoia. The karyotype is shared only by N. inversa. This species ranges the farthest south of any Sequoia or Sierranus Group species and occurs in the Tehachapi Mountains in addition to the Sierra Nevada. Etymology. l. proro ���to prolong, keep going��� + cantans ���singing,��� describing the incessant and repetitive nature of the male calling song. Notes. This species may be common in years of adequate rainfall and scarce in dry years, during which populations are localized around water sources. During the summer of 2001, a wet year for California, katydids were abundant in the Kernville area and males were seen walking and singing on bare soil on hillsides some distance from riparian or forested areas (JAC pers. obs). Material examined. Type series only, see Type material above., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warcha��owska-��liwa, El��bieta, Marya��ska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 66-68, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851"]}

Neduba radicata Cole, Weissman, & Lightfoot, sp. n. Fig. 9 (distribution), Fig. 15 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 1D (live habitus), Plate 4F (male calling song), Plate 6K (male ventral sclerite), Plate 9G (male titillators), Plate 11F (female subgenital plate). Common name. Lake Tahoe Shieldback. History of recognition. Confused with N. convexa, specimen from CA: El Dorado Co., Ice House Road, 22-X- 1965, D.C. Rentz, 1 male (Rentz & Birchim 1968). Type material. HOLOTYPE MALE: USA, CA, El Dorado Co., Stanford University Sierra Camp, Fallen Leaf Lake, 38.901076N, 120.061626W, 1940 m, 9-VII-1988, DB Weissman, CAS, S88-60 [stop], R88-75 A [recording], T88-8 [karyotype], genitalia extracted and cleared in vial and excised tegmen in gelcap below specimen, deposited in CAS, Entomology type #19679. PARATYPES (n = 36): USA, CA, El Dorado Co., 8♁, 1♀, China Flat Campground, El Dorado National Forest, 2 mi. SE of Kyburz off US50, 38.7535N, 120.2671W, 1470 m, 20-21-VII-2012, JA Cole, LACM; 1♁, same data except JAC; 4♁, same data except 12-VIII-2002, JA Cole, LACM; 1♁, same data JAC; 2♁, same data except 19-VII- 2015, JA Cole, DB Weissman, LACM; 1♁, Emerald Bay State Park, Lake Tahoe, 38.964888N, 120.090884W, 1890 m, 8-VII-2016, DI Weissman, CAS; 1♀ nymph, same data as holotype except 10-VII-1935, FE Blaisdell, CAS; 1♀ nymph, same data as holotype except 20-VII-1936, FE Blaisdell, CAS; 1♀, same data as holotype except 3-VIII- 1989, BI Weissman, CAS; 2♁, 2♀, same data as holotype except 9-VII-1988, DB Weissman, CAS; 1♁ nymph, same data as holotype except VII-1915, LS Rosenbaum, CAS; 1♀ nymph, same data as holotype except VII-1931, OH Swezey, CAS; 6♁, 1♀, Loon Lake, 38.997163N, 120.3095W, 1950 m, VIII-1991, BI Weissman, CAS; Placer Co., 1♁, Finning Mill Rd. at Forest Hill Rd., 39.0586N, 120.7696W, 1097 m, 20-VII-2012, JA Cole, JAC; 1♁, Squaw Valley, foot trail by Squaw Creek, 39.21169N, 120.198611W, 1920 m, 1-VIII-1988, VF Lee, CAS. Measurements. (mm, ♁n = 20, ♀ n = 4) Hind femur ♁18.00–20.91, ♀ 19.05–22.36, pronotum total length ♁8.70–10.16, ♀ 7.35–8.46, prozona length ♁3.38–4.36, ♀ 3.80–4.62, metazona dorsal length ♁4.82–6.47, ♀ 3.32– 4.66, pronotum constriction width ♁2.10–2.65, ♀ 2.55–2.87, metazona dorsal width ♁6.35–7.55, ♀ 5.28–5.62, head width ♁4.20–4.75, ♀ 4.50–5.22, ovipositor length ♀ 12.78–15.54. Distribution. High elevations in the central and northern Sierra Nevada of California. Habitat. Mixed conifer woodland. Taken from understory tangles and leaf litter in coniferous forest, under logs, in a small shrub in an open field, and from riverbanks. One male paratype sang from 2.4 m above ground in thicket of branches. He dropped to the ground when approached (JAC pers. obs.). Seasonal occurrence. Midsummer through fall, from July (1-VII-1950, HL McKenzie, CSCA) through November (17-XI-1967, HR Ingham, CSCA). Nymphs occur from early June through August, thus overlapping broadly with adult activity. Stridulatory file. (n = 6) length 3.3–3.9 mm, 115–135 teeth, tooth density 35.4 ± 3.4 (30.8–40.9) teeth/mm. Song. (n = 19) PTR is significantly faster than all other Convexa Clade lineages (ANCOVA, P = 1.08×10 -5), owing to significantly shorter MPTL of 202.5 ± 53.6 ms (ANCOVA, P = 4.27×10 -11). MPT often have a characteristic amplitude modulation pattern, with a gradual increase in amplitude followed by an abrupt increase at the middle of the PT (Plate 4F). Males may sing in the late afternoon as well as at night. Karyotype. (n = 7) 2n♁ = 26 (2m + 22t + XtYt), T88-8, S88-60, paratype. Recognition. The male ventral sclerite has straight, thick shaft, the apex rounded with the highest point lateral to the central axis, and the short lateral process directed 45 o anterior to plane of shaft. This genital morphology may only be confused with N. diabolica and some N. carinata, both of which have higher stridulatory file tooth densities and lisping songs and are distributed in the South Coast Ranges. The female subgenital plate is distinct from all other Carinata Group species: wider than long with a strongly bifurcate apex. The song PTR is faster and MPTL shorter than all other Convexa Clade taxa. This species is the only Neduba found in high elevation yellow pine forest in the central and northern Sierra Nevada Mountains of California. Etymology. l. radicata having roots, having found a home. Notes. At China Flat Campground, El Dorado County, California, two males were observed emerging from leaf litter to sing in understory tangles.Acoustical activity commenced before dusk at this locality during overcast conditions after a rainstorm. At the southern extent of the range along Finning Mill Road, males were singing before sunset at 2019 h. This species is sympatric with N. radocantans (Sierranus Group) in El Dorado County, California. Specimens examined. (n = 21) All USA, CA, El Dorado Co., in addition to type material (above), 1♁, Kelsey, 38.798791N, 120.820768W, 24-VII-1939, J Labadie, CSCA; 1♀, Pollock Pines, 38.761292N, 120.586594W, 1207 m, 1-VII-1950, HL McKenzie, CSCA; 1♀ nymph, Snowline Camp, 38.746292N, 120.624373W, 21-VI-1948, CD MacNeill, CAS; 1♁, Strawberry, 38.796852N, 120.145187W, 1768 m, 1-X-1954, ME Gardner, BMED; 1♀, same data except WJ Wall, BMED; 1♀ nymph, same data except 28-VII-1950, ME Gardner, BMED; 1♀ nymph, same data except 3-VIII-1952, WJ Wall, BMED; 1♁ nymph, Strawberry Valley, 38.796582N, 120.145187W, 12-VIII- 1912, EC VanDyke, CAS; 1♁ nymph, same data except 5-VIII-1912, EC VanDyke, CAS; 1♁, US50 3.7 mi. W Kyburz, 38.76427N, 120.35897W, 1112 m, 19-VII-2015, JA Cole, DB Weissman, JAC sound record; Nevada Co., 1♁, junction SR20&I-80, 4-X-1972, AM Shapiro, BMED; Placer Co., Cisco, 39.301569N, 120.546874W, VI-1910, C VanGeldern, CAS; 3♁, Emigrant Gap, 39.300456N, 120.668268W, 1567 m, 17-XI-1967, HR Ingham, CSCA; 1♀, Michigan Bluff, 39.042956N, 120.741319W, 1073 m, 11-V-1962, G Buxton, CSCA; Sierra Co., 1♁, 10 mi. S Downieville, 39.41439N, 120.826891W, 28-VII-1962, WJ Turner, CSCA; 1♁, Goodyears Creek, 39.540911N, 120.88824W, 10-VII-1925, EB Nast, CAS; 1♁, St. Charles Hill, 39.56795N, 120.911061W, 7-VII-1925, EB Nast, CAS; Yuba Co., 1♁ nymph, Challenge, 39.487389N, 121.223572W, 1-VII-1963, J Vercamp, CAS; 1♁, same data except 20-VIII-1964, R Whitely, CAS; 1♀, same data except VIII-1963, E Ball Jr, CAS., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 37-38, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851"]}

Neduba carinata Walker, 1869 Figure 9 (distribution), Fig. 10 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 1A (live habitus), Plate 4A (male calling song), Plate 6 A–D (ventral sclerites), Plate 9 A–C (male titillators), Plate 11A (female subgenital plate). Common name. Carinate Shieldback. History of recognition. As the type species of Neduba, establishing the identity of N. carinata is of central taxonomic importance. In a situation resembling that of Gryllus assimilis (Rehn & Hebard 1915), N. carinata was at times used as a “trash can” taxon (Rentz & Birchim 1968), into which multiple species were binned (e.g. Tinkham 1944). The broad type locality of “ California ” was narrowed to Fremont Peak, San Benito County, California, USA (Rentz & Birchim 1968). We accepted this type locality designation and determined taxonomic and geographic relationships relative to this locality. The Fremont Peak type locality is in the Gabilan Range of the South Coast Ranges of California. In over 40 years of fieldwork by orthopterists at Fremont Peak (e.g. Helfer 1987; Rentz & Birchim 1968; Strohecker et al. 1968), sympatric Neduba species have not been found at this locality; however, the correct assignment of the name N. carinata requires that all other possibilities are eliminated. This is a valid concern considering that an additional Neduba species occurs in the South Coast Ranges (see N. lucubrata). Images of the N. carinata holotype show a subgenital plate with parallel lateral carinae and long, articulate styli (Cigliano et al. 2020), a condition that is found in all topotypic males that we have examined. In contrast, the subgenital plate of N. lucubrata has lateral carinae that converge to the apex and terminate in rudimentary knob-like styli. Thus, we conclude that the name N. carinata indeed belongs to our topotypic series. Variety picturata (Scudder 1899) was synonymized under N. diabolica (Rentz & Birchim 1968). Type material. F. Walker’s male holotype of N. carinata resides in the British Museum. Type not examined by us but images available at OSFO (Cigliano et al. 2020). TOPOTYPES EXAMINED (n = 25): USA, CA, San Benito Co., 2♁, Fremont Peak State Park, peak area, 36.760793N, 121.502442W, 457 m, 25-VI-1982, DB Weissman, CAS; 6♁, 1♀, same data except 762 m, 25-VI-1982, DB Weissman, CAS; 14♁, 1♀, Fremont Peak State Park, Valley View Campground, 11 mi. S of San Juan Bautista on County Road G1, 36.7597N, 121.505W, 829 m, 18–19- VII-2005, JA Cole, JF Eguizabal, LACM; 1♁, same data except JAC. Measurements. (mm, ♁n = 34, ♀ n = 7) Hind femur ♁18.33–28.80, ♀ 21.55–26.21, pronotum total length ♁8.23–10.72, ♀ 8.35–10.05, prozona length ♁3.43–4.88, ♀ 4.02–5.20, metazona dorsal length ♁4.45–6.45, ♀ 3.67– 4.85, pronotum constriction width ♁2.00–2.78, ♀ 2.35–3.00, metazona dorsal width ♁5.11–7.25, ♀ 5.18–6.45, head width ♁3.89–4.84, ♀ 4.60–5.32, ovipositor length ♀ 16.00–18.27. Distribution. South Coast Ranges of California, from the southern San Francisco Bay Area south along the Santa Lucia Range to Morro Bay. Habitat. Understory vegetation, tangles, and leaf litter in mixed woodland, riparian woodland, and coastal redwood (Sequoia sempervirens (Lamb. ex D. Don) Endl.) forest. Males call near the forest floor from plants such as poison oak (Toxicodendron diversilobum (Torr. & A. Gray)), introduced Mediterranean grasses, and peavines (Lathyrus sp.). Seasonal occurrence. Adults from late June (23-VI-1957, TJ Cohn, CAS) through December (18-XII-2016, DW Weissman, CAS). Nymphs collected in May and alongside adults in late June (CAS). Stridulatory file. (n = 24) length 2.4–3.4 mm, 115–159 teeth, tooth density 44.7 ± 3.9 (38.5–55.3) teeth/mm. Song. (n = 52) Frequent sustained bouts of low frequency lisping. Males call incessantly at night and sometimes during the afternoon. PTR 10.9 ± 2.1 s- 1. The MPT and OPT are subequal in length. Males spend a lot of time singing with PTdc 73.1 ± 9.7% (calculated by combining both MPT and OPT). PTF 11.4 ± 1.7 kHz is among the lowest carrier frequencies in the genus. Karyotype. (n = 22) 2n♁ = 26 (2m + 22t + XtYt). T82-6, S82-26, topotype. Recognition. This species is separated morphologically from all other species in the Carinata Group except N. diabolica by the high stridulatory file tooth density (38–55 teeth/mm). The male ventral sclerite throughout most of the range has the shaft broad and straight (narrow, curved shaft in Convexa Clade), the apex broad and convex to subconical covered with tubercles of large size, and a short lateral process (often long and shelf-like in the Convexa Clade). In N. carinata populations found near the Convexa Clade in the South Bay Area, the ventral sclerite of the former species resembles the latter and is an unreliable diagnostic feature; songs remain diagnostic. Acoustically, the low frequency, rasping or lisping bouts of song with a fast uniform PTR (~11 s-1) separate this species from all other Neduba except N. diabolica. Females have a pentagonal subgenital plate without sinuous distal margins and no medial groove, characters shared only with N. oblongata. Notes. This is a common species in California Coast Range woodlands throughout summer and fall. The proximity of the range to the heavily populated San Francisco Bay Area and the low frequency calling song that many people can hear makes this one of the most commonly encountered species. Considerable variation exists in the male internal genitalia, which we attribute to historical episodes of population isolation and contact as the California coastline and Coast Ranges changed repeatedly and dramatically over the Cenozoic Era. The Santa Lucia Range and portions of the Santa Cruz Mountains have changed between islands, peninsulas, and broad mainland connections (Bartow 1991). A distinct ventral sclerite shape is found in the Santa Lucia Range (Fig. 10, Plate 6B), which may reflect separation from the Coast Ranges north of Monterey Bay by a marine embayment that lasted from 18 to 2 Ma (Hall 2002). Santa Cruz Mountains N. carinata have ventral sclerites with long lateral processes (Fig. 10, Plate 6D) that resemble those of several Convexa Clade taxa (Plates 6–7). The Carinata and Convexa Clades meet geographically in the San Francisco Bay Area, where mtDNA introgression (Fig. 4) suggests gene flow as the explanation for shared ventral sclerite morphology despite distinct calling song differences. Material examined (n = 116). All USA, CA, Monterey Co., 1♁, 1.5 mi. N of Carmel, 36.58N, 121.92W, 213 m, 23-VI-1957, TJ Cohn, CAS; 1♁, 1♀, 11.2 mi. N Pfeiffer Big Sur State Park off Hwy 1, 36.411288N, 121.782732W, 5-VII-1986, DB & BI Weissman, CAS; 1♀, Asilomar, 1.3 mi. W of Pacific Grove, 36.617735N, 121.94011W, 10-VII-1957, TJ Cohn, CAS; 1♀ nymph, Big Sur River Trail, 36.280942N, 121.860028W, 21–23- V-1971, FS Bartholomew, CAS; 6♁, 1♀, Big Sur, SR1, 36.3547N, 121.8136W, 685 m, 20-VIII-2012, JA Cole, LACM; 2♁, Bottcher’s Gap, Los Padres National Forest, 19 miles north of Big Sur off SR1 on Palo Colorado Road, 36.355N, 121.8138W, 652 m, 20-VIII-2012, JA Cole, LACM; 2♁, same data except 7-IX-2002, JA Cole, LACM; 1♁, 1♀, same data except JAC; 1♁, Nacimiento-Ferguson Rd., at bridge of Nacimiento R., 8.4 mi. E of SR1, 36.0135N, 121.4216W, 587 m, 19–20-VIII-2012, JA Cole, JAC sound record; 2♁, 1♀, Pacific Grove, 36.617738N, 121.916622W, 4-VII-1925, HH Keifer, CAS; 1♁, Palo Colorado Rd., 3 mi. E of SR1, 36.3864N, 121.8687W, 327 m, 20-VIII-2012, JA Cole, JAC sound record; 1♁ nymph, Pebble Beach, 36.56635N, 121.946622W, 18-V-1969, RP Allen, CSCA; 1♀ nymph, same data except 27-V-1921, EC VanDyke, CAS; 1♁, Pfeiffer Big Sur State Park, 36.248852N, 121.782732W, 60 m, 23-IX-1995, DB Weissman, CAS; 3♁, same data except 27-IX-1985, DB Weissman, CAS; 4♁, 3♀, same data except 5-VII-1986, DB & BI Weissman, CAS; San Benito Co., see Type material above; San Francisco Co., 1♁, San Francisco, 37.77493N, 122.419416W, 24-X-1909, FX Williams, CAS; San Mateo Co., 2♁, Jasper Ridge, 37.407995N, 122.22691W, 12-VII-1982, DB Weissman, CAS; 1♁, Kings Mount, 37.44083N, 122.32333W, 14-X-1946, ES Ross, CAS; 1♁, 1♀, same data except 17-VIII-1946, ES Ross, CAS; 1♁, Portola State Park, 37.2527N, 122.21854W, 8-VIII-1962, RP Allen, CSCA; Santa Clara Co., 1♀, Los Gatos, 12-XII-1965, J Harville, CAS; 2♁, Los Gatos, 15431 Francis Oaks Way, 37.22661N, 121.97468W, 22-VIII-2013, DB Weissman, CAS; 1♁, same data except VI-1986, DB Weissman, CAS; 7♁, 2♀, Los Gatos, Francis Oaks Way, 37.226611N, 121.97468W, 120 m, 6-VII-1987, DB Weissman, CAS; 2♁, Uvas Canyon County Park, 15 miles west of Morgan Hill on Croy Road, 37.0855N, 121.7954W, 349 m, 19-20-VII-2005, JA Cole, JF Eguizabal, JAC; 6♁, 1♀, same data except LACM; 1♁, 2♀, same data except 2-3-VII-2003, JA Cole, LACM; 2♁, 2♀, same data except 22-VIII-2014, DB & DW Weissman, CAS; 1♁, same data except 6-IX-2002, JA Cole, JAC; 2♁, same data except LACM; 1♁, Mount Umunhum, 37.15255N, 121.89897W, 1008 m, 5-VII-2018, DW & DB Weissman, CAS; Santa Cruz Co., 1♀, Loma Prieta, 37.110781N, 121.844676W, 2-VIII-1959, CAS; 1♀, Santa Cruz, 36.974117N, 122.030796W, 25-XI-1955, RL Lewis, CSCA; 1♁, Soquel, 36.98801N, 121.95663W, 18-XII-2016, DW Weissman, CAS; 4♁, same data except 9-IX-2014, DW Weissman, CAS; 1♁, Summit above Saratoga, 37.263832N, 122.023015W, 26-VIII-1968, J Smith, BMED; 6♁, 1♀, U.C. Santa Cruz Campus, 36.974117N, 122.030796W, 25- VI-1982, SI Weissman, CAS., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 26-28, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800, {"references":["Walker, F. (1869) 2 Catalogue of the Specimens of the Dermaptera Saltatoria in the Collection of the British Museum. Printed for the Trustees of the British Museum, London, 216 pp.","Rehn, J. A. G. & Hebard, M. (1915) The genus Gryllus (Orthoptera) as found in America. Proceedings of the Academy of Natural Sciences of Philadelphia, 67, 293 - 322.","Rentz, D. C. & Birchim, J. D. (1968) Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society, 3, 1 - 173. https: // doi. org / 10.5962 / bhl. title. 150851","Tinkham, E. R. (1944) Biological, taxonomic and faunistic studies on the shield-back katydids of the North American deserts. American Midland Naturalist, 31, 257 - 328. https: // doi. org / 10.2307 / 2421073","Helfer, J. R. (1987) How to Know the Grasshoppers, Crickets, Cockroaches and Their Allies. Dover Publications, Inc., New York, ix + 364 pp.","Strohecker, H. F., Middlekauf, W. W. & Rentz, D. C. (1968) The grasshoppers of California. Bulletin of the California Insect Survey, 10, 1 - 177.","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2020) Orthoptera Species File Online v. 5.0 / 5.0.","Scudder, S. H. (1899) Two genera of North American Decticinae. Proceedings of the American Academy of Arts and Sciences, 35, 81 - 93. https: // doi. org / 10.2307 / 25129903","Bartow, J. A. (1991) The Cenozoic Evolution of the San Joaquin Valley, California. United States Geological Survey, United States Government Printing Office, Washington, 40 pp. https: // doi. org / 10.3133 / pp 1501","Hall, C. A. (2002) Nearshore marine paleoclimatic regions, increasing zoogeographic provinciality, molluscan extinctions, and paleoshorelines, California: late Oligocene (27 Ma) to late Pliocene (2.5 Ma). Geological Society of America, Special Papers, 357, 1 - 489. https: // doi. org / 10.1130 / SPE 357"]}

Neduba sequoia Cole, Weissman, and Lightfoot sp. n. Fig. 19 (distribution), Fig. 29 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 3 F–G (live habitus), Plate 5I (male calling song), Plate 8 F–H (male ventral sclerite), Plate 10I (male titillators), Plate 12G (female subgenital plate). Common name. Big Trees Shieldback History of recognition. Likely confused with N. sierranus. Type material. HOLOTYPE MALE: USA, CA, Tulare Co., Coy Flat Campground, Sequoia National Forest, 0.5 miles south of Camp Nelson off SR190, 36.1269N, 118.6183W, 1524 m., 30-31-VII-2012, JA Cole, DNA67 [tissue], SING357 [DNA extraction], JCR120914_00 [recording], genitalia in vial below specimen, deposited in CAS, Entomology type #19714. PARATYPES (n = 29): Tulare Co., 9♁, 1♀, same data as holotype, LACM; 1♁, same data as holotype JAC; 2♁, same data as holotype except, 20-VIII-2006, DB Weissman, DC Lightfoot, CAS; 11♁, 3♀, South Fork Campground, Sequoia National Park, 36.35029N, 118.76511W, 1112 m, 12-13-VIII-2015, JA Cole, LACM; 1m, same data except JAC; 1♁, Hwy 190 7 mi. E Springville, 36.156806N, 118.724278W, 707 m, 5-V-2011, DB Weissman, DBW. Measurements. (mm, ♁n = 23, ♀ n = 5) Hind femur ♁17.48–24.26, ♀ 22.03–24.14, pronotum total length ♁8.36–10.66, ♀ 9.50–10.05, prozona length ♁3.34–5.40, ♀ 4.78–5.96, metazona dorsal length ♁4.08–6.05, ♀ 3.79– 4.72, pronotum constriction width ♁2.15–3.15, ♀ 2.57–2.96, metazona dorsal width ♁5.40–6.97, ♀ 5.85–6.65, head width ♁4.45–5.72, ♀ 5.10–5.90, ovipositor length ♀ 14.68–17.42. Distribution. Western slope of the southern Sierra Nevada Mountains between the Kaweah River and Tule River watersheds, in the vicinity of Sequoia National Park. Habitat. Understory of mixed conifer forests, especially in riparian habitats and mesic areas. At the South Fork localities, many adults were feeding on green leaves of mountain mahogany (Cercocarpus). Singing males at this locality were observed and recorded in young dead cedars at the forest edge. Seasonal occurrence. Adults from late July (27-VII-1986, DJ Burdick, CAS) through August (20-VIII-2006, DB Weissman, CAS). Nymphs from May through mid-June. Stridulatory file. (n = 18) length 2.7–3.7 mm, 152–203 teeth, tooth density 57.5 ± 3.4 (47.5–61.6) teeth/mm. Song. (n = 19) A continuous series of alternating MPT and OPT as in other Sequoia Group species (except N. inversa). The PTR of 3.0 ± 0.5 s- 1 is significantly faster than N. prorocantans, which is distributed to the south, and is half the rate of syntopic N. duplocantans. PTdc of 75.1 ± 8.7% is significantly higher than all other Sequoia Group species (ANCOVA, P = 3.75×10 -4). PTF is 14.3 ± 1.0 kHz. Karyotype. (n = 4) Unique. 2n♁ = 22 (2m + 18t + XtYt). T06-12, S06-80, paratopotype. Recognition. This is a plastic species that is difficult to recognize. A higher stridulatory file tooth density (55– 59 teeth/mm) as well as larger body size will separate this species from the lower density (47–52 teeth/mm) of both N. prorocantans distributed to the south and N. duplocantans, with which it is sympatric. To the north, N. inversa has a greater stridulatory file tooth density (64–68 teeth/mm). The song PTR is faster than that of N. prorocantans but half that of N. duplocantans, whereas N. inversa sings with numerous OPT between MPT as in Sierranus Group taxa. The karyotype is unique. N. sequoia inhabits an area between the Kaweah River and Tule River watersheds, a range that it shares only with N. duplocantans. Etymology. Named after Sequoia National Park and Sequoia National Forest of the southern Sierra Nevada of California. Notes. This species is common at all localities where collected. Taxidermy of South Fork specimens showed the gut contents to be full of Cercocarpus. Collections were also made at oatmeal trails. Material examined. DETERMINED (n = 2): Tulare Co., in addition to type material (above), 1♀, Ash Mountain, Kaweah Power Station 3, 36.48606N, 118.83586W, 27-VII-1996, DJ Burdick, CAS; 1♀ nymph, Ash Mountain, Kaweah Power Station 3, 36.48606N, 118.83586W, 3-VI-1984, DJ Burdick, CAS. QUESTIONABLE PLACEMENT (n = 8): Tulare Co., 2♀ nymphs, Sequoia National Park, Potwisha Campground, 36.517446N, 118.799821W, 13-VI-1923, EC VanDyke, CAS; 3♀ nymphs, same data except 13-VI-1929, EC VanDyke, CAS; 1♀ nymph, same data except 27-V-1928, EC VanDyke, CAS; 1♁, 1♀ nymphs, same data except 8-V-1931, EC VanDyke, CAS., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 68-70, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800

Neduba cascadia Cole, Weissman, & Lightfoot, sp. n. Fig. 9 (distribution), Fig. 17 (male and female habitus, calling song, male and female terminalia, karyotype), Plate 1 F–H (live habitus), Plate 4H (male calling song), Plate 7E (male ventral sclerite), Plate 11H (female subgenital plate). Common name. Cascade Shieldback. History of recognition. None. Type material. HOLOTYPE MALE: USA, OR, Jackson Co., Wildcat Campground, Hyatt Lake Complex, Cascade-Siskiyou National Monument, 42.18283N, 122.44775W, 1531 m, 27-29-VII-2016, JA Cole, JAC000002023 [specimen barcode], DNA218 [genomic], SING0616 [DNA extraction], JCR160727_02 [recording], 125 [teeth], 3.7 [mm stridulatory file length], tegmen in gelcap below specimen, deposited in CAS, Entomology type #19708. PARATYPES (n = 12): 5♁, same data as holotype, LACM; 4♁, same data as holotype, CAS; 1♁, same data as holotype, JAC; 1♀, same data as holotype except 28-VIII-2019, DB Weissman & DC Lightfoot, CAS; 1♀, Woodruff Meadow, 30 mi. SW Crater L., 42.885125N, 122.509206W, 6-VIII-1960, JR Helfer, CAS. Measurements. (mm, ♁n = 7, ♀ n = 1) Hind femur ♁18.05–18.95 ♀ 19.91, pronotum total length ♁9.20–9.81, ♀ 8.85, prozona length ♁3.01–4.65, ♀ 4.55, metazona dorsal length ♁5.10–6.80, ♀ 4.30, pronotum constriction width ♁2.07–2.30, ♀ 2.55, metazona dorsal width ♁6.50–7.25, ♀ 6.49, head width ♁4.10–4.68, ♀ 5.02, ovipositor length ♀ 16.00. Distribution. Southern Cascade and Siskiyou mountain ranges of southern Oregon. Habitat. Understory and edges of coniferous forest. Seasonal occurrence. Scant records are from mid-July through late August. Adult activity probably lasts from midsummer through fall until first frosts. Stridulatory file. (n = 4) length 3.4–3.7 mm, 125–133 teeth, tooth density 36.7 ± 2.1 (33.8–38.8) teeth/mm. Song. (n = 6) Typical Convexa Clade “zwee-zwee” song type but with a significantly slower PTR of 1.8± 0.1 s- 1 (ANCOVA P = 1.08×10 -5) and longer MPTL of 498.2 ± 65.5 ms (ANCOVA P = 4.27×10 -11) than all other Convexa Clade taxa. PTF is 13.9 ± 2.2 kHz. Karyotype. (n=2) 2n♁ = 26 (2m + 22t + Xt +Yt) T19-16, S19-90, paratopotype. Recognition. The male ventral sclerite of N. cascadia has a blunt apex with scattered large tubercles and a short lateral process that blends into the shaft by an obtuse angle. This contrasts with N. convexa and N. longiplutea, both distributed to the south, which have the ventral sclerite apex low if not flat and a long lateral process with numerous fine tubercles that are arranged in rows. N. steindachneri to the north has a mushroom-shaped ventral sclerite with few large tubercles and a blunt, recurved lateral process. The female subgenital plate is unique among the Convexa Clade in having straight apical margins. The slow PTR separates the song of N. cascadia from all other Carinata Group species; N. steindachneri to the north has the fastest PTR in this species Group. Etymology. cascadia, reflecting the southern Cascade Range distribution. Notes. The distribution of this species lies between that of N. steindachneri to the north and N. convexa to the south. Further studies are needed to determine the limits of the ranges of northern Carinata Group taxa. Museum specimens from intermediate localities are nymphs and are only tentatively identified as this species. The calling songs and genitalia of all three species differ, so sympatry may be expected if these differences confer reproductive isolation. That calling song extremes of PTR occur in two species with adjacent ranges suggests prezygotic reproductive isolation mediated by calling song. Material examined. In addition to type material (above), QUESTIONABLE PLACEMENT (n = 3): USA, OR, Jackson Co., 2♀ nymphs, Little Applegate River, 42.198722N, 123.045356W, 701 m, 6-VIII-1950, B Malkin, CAS; 1♀ nymph, Union Creek, 42.906905N, 122.445598W, 7-31-VIII-1950, B Malkin, CAS., Published as part of Cole, Jeffrey A., Weissman, David B., Lightfoot, David C., Ueshima, Norihiro, Warchałowska-Śliwa, Elżbieta, Maryańska-Nadachowska, Anna & Chatfield-Taylor, Will, 2021, A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini), pp. 1-92 in Zootaxa 4910 (1) on pages 42-44, DOI: 10.11646/zootaxa.4910.1.1, http://zenodo.org/record/4448800

Okanagana boweni Chatfield-Taylor and Cole, new species Table 1 (measurements), Table 2 (song statistics), Table 3 (molecular vouchers), Fig. 1 (habitus), Fig. 2 (male timbal), Fig. 3 (male terminalia), Fig. 4 (female terminalia), Fig. 5 (calling song), Fig. 6 (geographic distribution), Fig. 7 (phylogenetic hypothesis). Zoobank ID: 9C190D25-DE35-406D-AC4C-803AF864F25C Type material. HOLOTYPE: 1 male, USA, NV, Washoe Co., Pyramid Lake, Tamarack Beach, 39.91333 oN, 119.54083 oW, 1134 m, 10-VII-2019, JA Cole, J Bailey, W Chatfield-Taylor, JF Eguizabal leg., ex tamarisk, rabbitbrush, recording WCT3737, barcode JAC000000230, deposited at LACM, LACM ENT 456101. PARATYPES: 1 female, same data as holotype, LACM; 1 male, same data as holotype, CAS; 1 male, 1 female, same data as holotype, DNA extractions SING0873, SING0874, LACM; 3 males, same data as holotype, SEMK; 1 male, USA, CA, Inyo Co., Lone Pine Campground, 7 miles west of Lone Pine on Whitney Portal Road, 36.5977 oN, 118.1838 oW, 1785 m, 9-VII-2008, JA Cole leg., ex Artemisia tridentata, DNA extraction SING0246, LACM; 1 male, same data as previous, CAS; 2 males, same data, SEMK; 1 male, CA, Inyo Co., Lone Pine 15 mi. S, 36.3981 oN, 118.0256 oW, 1129 m, 18-VI-1969, PM Jump leg., LACM; 1 male, CA, Inyo Co., Lone Pine Cp. Gr., 36.5971 oN, 118.1769 oW, 1744 m, 28-VI-1970, DS Verity leg., LACM; 1 male, CA, Inyo Co., Tinemaha Rd. 2 mi. E US 395 nr. Independence, 36.8563 oN, 118.2145 oW, 1165 m, 28-V-1978, N Lee leg., LACM; 5 males, CA, Lassen Co., Doyle 4.5 mi. S, 39.9792 oN, 120.0622 oW, 1219 m, 15-VI-1969, RR Snelling & RA Snelling leg., LACM (Supp. Table 1). Description. Head width narrower than front of pronotum. Deep median groove in yellow triangle at base of epicranium continues between lateral ocelli to form the epicranial suture. Front strongly produced with a median sulcus. Frons hirsute with deep frontocylpeal suture. Supra-antennal plates orange (Fig. 1). Lateral margins of pronotum distinctly subparallel with broad posterior margin entire, giving a quadrate aspect. Humeral angles broadly rounded, anterior angles acute. Pronotum completely bordered with yellow. Rugose ridges lateral to pronotal midline marked with variable amounts of brown or yellow. Anterior half of pronotum with strongly sulcate broad longitudinal yellow line. Mesonotum black, with broad yellow posterior margin and cruciform elevation partly (holotype) to completely (paratypes) yellow. Four yellow spots form a semicircle with yellow marks at base of each wing. Size and shape of mesonotal markings variable (extensive in holotype). Metanotum bordered by yellow that is interrupted by black anterior to timbals (Figs. 1, 2). Forewings with yellow costa turning black and slightly fuscous past the node. Venation variegated with yellow to marginal cells (holotype) or mostly black (some paratypes). Posterior margin of membrane gently curved, wing tip rounded (feature visible only on spread, intact specimens). Basal cell opaque, varying from black (holotype) to yellow (some paratypes). Hind wings lightly fuscous at base. Wing membranes orange. Meso- and metasternum hairy, marked with black and yellow. Profemora striped black and yellow, other legs variably marked (Fig. 1). Tergum sparsely hairy, shining black, with posterior margin of last two tergites variably bordered with yellow (Fig. 1). Male timbals with 2 major and 2 minor ribs (Fig. 2). Sternites yellow with extensive variable black markings edging posterior margins; in holotype, yellow markings are narrow at the center and broaden laterally, giving the impression of a central longitudinal black stripe bordered by two yellow stripes on the abdominal venter. The 7th abdominal sternite hourglass-shaped, medially constricted with rounded apical margins (Fig. 1). Uncus in dorsal view diamond-shaped, formed from straight lateral margins that widen for one half to two thirds of the length and then converge towards the apex (Fig. 3 shows typical condition). In lateral view, upper surface with distinct curvature, not parallel with respect to lower surface. Hook forms abruptly after short and deep posteroventral excavation. Aedeagus elongated, attached at an angle perpendicular to the plane of the uncus with posterior edge distinctly incurved to varying degrees. Male valve roughly triangular with posterior edge slightly curved. Female with apical prongs on 7th abdominal sternite broad and rounded with a wide, faint secondary notch (Fig. 4). Diagnosis. Okanagana boweni is a large-bodied species most similar to allopatric O. simulata in body size and habitus but to sympatric O. utahensis in color pattern (Fig. 1). Diagnosis of museum specimens may be accomplished by a suite of characters, some of which are visible on spread specimens with pulled genitalia and others on unprepared specimens. In unprepared specimens a character combination is necessary for diagnosis of the three similar species. A quadrate pronotum with parallel lateral margins separates O. boweni and O. simulata from O. utahensis, which has an anteriorly constricted pronotum, and a broad longitudinal yellow sulcus on the anterior half of the pronotum separates O. boweni and O. utahensis from O. simulata, in which the sulcus is barely indicated and faintly marked (Fig. 1). Thus, O. boweni possesses both a quadrate pronotum with a broad yellow longitudinal sulcus. Less reliable diagnostic characters include the semicircle of mesonotal markings, which tend to be more extensive in O. boweni and O. utahensis compared with those of O. simulata, the head width narrower than the anterior margin of the pronotum in O. boweni but subequal in O. simulata, and the posterior pronotal margin appearing complete in O. boweni but incomplete in O. simulata due to a darkened median constriction in the latter (Fig. 1). On prepared specimens the uncus and aedeagus are valuable characters for diagnosing the two species (Fig. 3). Uncus shape should be used in conjunction with other characters due to intraspecific variation that is found generally throughout Okanagana (WCT pers. obs.). In dorsal view, the uncus of O. boweni has parallel lateral margins that converge towards the apex with a distinct angle that begins at one half to two thirds of its length, compared with the gradually and smoothly curved lateral margins of O. simulata and O. utahensis. When exposed in lateral view, the aedeagus is a singular character that diagnoses O. boweni from both O. simulata and O. utahensis. The posterior edge of the aedeagus is long and incurved in O. boweni, forms an acute angle in O. simulata, and is uniformly straight and short in O. utahensis with its posterior margin situated perpendicular to the plane of the uncus (Fig. 3). Female O. boweni may be diagnosed by the lack of the deep secondary notch on the 7th abdominal sternite found in O. simulata and to a lesser extent in O. utahensis females (Fig. 4). For O. utahensis without clearly indicated secondary notches, diagnosis may require examination of pronotal shape or the condition of the longitudinal sulcus (Fig. 1). O. boweni is sympatric with O. utahensis but can be diagnosed from the latter in the field by the larger body size (Table 1) and quadrate aspect of the pronotum of O. boweni, in contrast with the smaller body size and trapezoidal aspect lent by the anterior convergence of the lateral pronotal margins in O. utahensis (Fig. 1). The morphology of the male (Fig. 3) and female (Fig. 4) terminalia are also diagnostic. Bioacoustics. The male calling song of O. boweni is a continuous train of syllables with a single dominant frequency that is similar to the calls of both O. simulata and O. utahensis (Table 2; Fig. 5; see Chatfield-Taylor & Cole 2019). Amplitude modulation at a rate of 7.82 �� 0.47 s- 1 (n = 4) may elaborate the song structure (Fig. 5), and when this occurs a wavering quality is imparted to the song as it oscillates in loudness. Amplitude modulations often began or became more pronounced when males were disturbed or approached, behavior that was also observed in O. simulata and O. utahensis (JAC, pers. obs.). Neither syllable rate (ANCOVA, P = 0.111) nor peak frequency (P = 0.653) varied with temperature. Songs differed significantly between species (MANOVA, P = 4.74x10 -4) but not between O. boweni populations (P = 0.543). Of the three species, O. utahensis was most divergent in song, but songs of O. boweni and O. simulata also differed significantly (P = 5.2x10 -3), differences that were again not explained by population (P = 9.08x10 -2). Per the methodology of Chatfield-Taylor and Cole (2019), the calls of O. boweni and O. simulata were separated by 0.09 standard deviations (SD) in parameter space while separation between O. boweni and O. utahensis ranged from 0.85 to 2 SD. Distribution. The distribution extends from the southern extent of the Owens Valley in eastern California north to Pyramid Lake, Nevada (Fig. 6). This distribution coincides with a geologically defined region of major transform faulting (Carlson et al. 2013; Wesnousky 2005) that has been termed the Walker Lane (Locke et al. 1940). The Walker Lane extends from the Garlock Fault at the southeastern corner of the Sierra Nevada and Tehachapi Range, which forms the northern border of the Mojave Desert, northeast to the Pyramid Lake Block in Nevada, and northwest to the Northern California Shear Zone that borders the Modoc Plateau in northeastern California (Carlson et al. 2013; Smith 1962; Stewart 1988). Etymology. The species is named after Charles Bowen, a naturalist from Belize who kindled the senior author���s interest in the biological sciences. Seasonal occurrence. Available records indicate activity from late spring (28 May, N. Lee, LACM) through midsummer (10 July, holotype, LACM). Habitat and notes. Habitat at the type locality is primarily alkali flats dominated by sages in the Tridentate group (Artemisia spp.) and tamarisk (Tamarix sp.; J. Bailey, pers. comm.). At the type locality males called from tamarisk and other available vegetation in their open habitat as they engaged in sing-fly behavior, while females were found exclusively on Artemisia tridentata. Besides O. boweni, we observed three other hooked uncus species in association with Artemisia tridentata during our 2019 field season: O. simulata, O. utahensis, and O. vanduzeei Distant. Hooked uncus taxa have been associated previously with Artemisia (Davis 1919; Kondratieff et al. 2002; Sanborn et al. 2002). Molecular phylogenetics. Alignment lengths were 510 bp for 16S, 198 bp for ARD1 UTR, 414 bp for ARD1 CDS, 423 bp for EF1�� exons, 305 bp for EF1�� introns, and 1013 indel characters. PartitionFinder returned two partitions: 1) the HKY+I model applied to a partition consisting of the 3rd codon positions of coding regions and all noncoding regions, and 2) the F81 model applied to all first and second codon positions. The standard deviation of split frequencies was 3.63x10 -3, which indicated significant convergence of Markov chains. A utahensis - vanduzeei clade resolved sister to a boweni - simulata clade (Fig. 7) with strong support (posterior probability 98%). Whereas the former pair were reciprocally monophyletic, O. simulata rendered O. boweni paraphyletic. A relatively long branch separated O. simulata from O. boweni., Published as part of Chatfield-Taylor, Will & Cole, Jeffrey A., 2020, A new species of Okanagana from the Walker Lane region of Nevada and California (Hemiptera: Auchenorrhyncha: Cicadidae), pp. 515-530 in Zootaxa 4868 (4) on pages 517-525, DOI: 10.11646/zootaxa.4868.4.3, http://zenodo.org/record/4418072, {"references":["Davis, W. T. (1919) Cicadas of the genera Okanagana, Tibicinoides and Okanagodes, with descriptions of several new species. Journal of the New York Entomological Society, 27, 179 - 222.","Davis, W. T. (1921) Records of cicadas from North America with descriptions of new species. Journal of the New York Entomological Society, 29, 1 - 16. https: // doi. org / 10.5962 / bhl. title. 9382","Chatfield-Taylor, W. & Cole, J. A. (2019) Noisy neighbours among the selfish herd: a critical song distance mediates mate recognition within cicada emergences (Hemiptera: Cicadidae). Biological Journal of the Linnean Society, 128, 854 - 864. https: // doi. org / 10.1093 / biolinnean / blz 132","Carlson, C. W., Pluhar, C. J., Glen, J. M. G. & Farner, M. J. (2013) Kinematics of the west-central Walker Lane: Spatially and temporally variable rotations evident in the Late Miocene Stanislaus Group. Geosphere, 9, 1530 - 1551. https: // doi. org / 10.1130 / GES 00955.1","Wesnousky, S. G. (2005) The San Andreas and Walker Lane fault systems, western North America: transpression, transtension, cumulative slip and the structural evolution of a major transform plate boundary. Journal of Structural Geology, 27, 1505 - 1512. https: // doi. org / 10.1016 / j. jsg. 2005.01.015","Locke, A., Billingsley, P. & Mayo, E. (1940) Sierra Nevada tectonic patterns. Geological Society of America Bulletin, 51, 513 - 540. https: // doi. org / 10.1130 / GSAB- 51 - 513","Smith, G. I. (1962) Large lateral displacement on Garlock Fault, California, as measured from offset dike swarm. Bulletin of the American Association of Petroleum Geologists, 46, 85 - 104. https: // doi. org / 10.1306 / BC 74375 F- 16 BE- 11 D 7 - 8645000102 C 1865 D","Stewart, J. (1988) Tectonics of the Walker Lane Belt, western Great Basin Mesozoic and Cenozoic deformation in a zone of shear. In: Ernst, W. (Ed.), Metamorphism and Crustal Evolution of the Western US, (Rubey Volume). Prentice Hall, Englewood Cliffs, New Jersy, pp. 685 - 713.","Kondratieff, B. C., Ellingson, A. R. & Leatherman, D. A. (2002) Insects of Western North America 2. The Cicadas of Colorado (Homoptera: Cicadidae, Tibicinidae). Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, iii + 63 + xx pp.","Sanborn, A. F., Breitbarth, J. H., Heath, J. E. & Heath, M. S. (2002) Temperature responses and habitat sharing in two sympatric species of Okanagana (Homoptera: Cicadoidea). Western North American Naturalist, 62, 437 - 450."]}

Okanagana boweni sp. n. is described from the western margin of the Great Basin of North America. The new species is diagnosed from allopatric O. simulata Davis and sympatric O. utahensis Davis using morphological, bioacoustical, and molecular characters. The distribution of this new species coincides with the Walker Lane region that lies along the border of California and Nevada, USA. Based on geography, bioacoustics, morphology, and molecular phylogenetics, we hypothesize that O. boweni sp. n. is the allopatric sister species of O. simulata.