Neduba ambagiosa Cole, Weissman, & Lightfoot, 2021

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), Zootaxa 4910 (1), pp. 1-92 : 30-32

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

Figures

PLATE 1. Live habitus of Carinata Group. A. N. carinata male, B. N. ambagiosa paratypes in phonotaxis, C. N. ambagiosa male (photo credit Jim Hogue), D. N. radicata paratype male, E. N. steindachneri male, F. N. cascadia paratopotype male showing mottled color form, G. N. cascadia paratopotype male singing (note elevated pronotal shield to give tegmina room to move, and perhaps, also create a parabola effect), H. N. cascadia paratyopotype male showing striped color form.

FIGURE 2. Neduba typical male calling song structure exemplified by N. convexa, recording JCR130808_01, 23.3oC (A-C), and song structure with multiple minor pulse trains exemplified by N. sierranus recording JCR120805_00, 21.7oC (D-E). A. 5 s oscillogram window showing 9 full wingstroke cycles (and a partial 10th major pulse train); B. 1.5 s oscillogram window showing individual pulses within major pulse trains; C. spectrogram of B with dashed line at limit of human hearing at 20 kHz; D. 5 s oscillogram window showing 6 full wingstroke cycles (and a partial series of minor pulse trains); E. 1.5 s oscillogram window of D showing individual pulses within major pulse trains; F. Male N. sierranus singing, Madera Co., CA. Note elevated pronotum exposing tegmina beneath; G. Phonotaxis in N. ambagiosa sp. n., Lake Co., CA.

PLATE 4. Male calling songs of the Carinata and Propsti Groups.. A. N. carinata, B. N. oblongata, C. N. ambagiosa, D. N. diabolica, E. N. longiplutea, F. N. radicata, G. N. convexa, H. N. cascadia, I. N. steindachneri, J. N. propsti. Left column shows 4 s oscillogram windows, right column are 1 s windows.

FIGURE 4. Comparison of A. rDNA, and B. mtDNA trees. Nodes with <85% posterior probability are collapsed. Species Groups are color coded.

PLATE 6. Variation in the ventral sclerites of the titillators of the male genitalia: Carinata Group.. A–D. N. carinata, E. N. oblongata, F. N. diabolica, G–H. N. ambagiosa × carinata hybrids, I. N. ambagiosa, J. N. longiplutea, K. N. radicata.

FIGURE 8. Distribution of Neduba species groups generated from records in this work.

FIGURE 9. Distribution of Carinata Group species showing records in this work.

PLATE 9. Titllator arms of Carinata Group species.

PLATE 11. Female subgenital plates of Carinata (A–I), Propsti (J), and Castanea (K–L) Group species.

FIGURE 12. N. ambagiosa male and female habitus, calling song, male and female terminalia, and karyotype.