Beameria ansercollis Sanborn and M. Heath, 2011

Sanborn, Allen F., Heath, Maxine S., Phillips, Polly K. & Heath, James E., 2011, A new species of the genus Beameria (Hemiptera: Cicadidae) from North America, Journal of Natural History 45 (25 - 26), pp. 1589-1605 : 1593-1602

Beameria ansercollis Sanborn and M. Heath sp. nov.

Type series

Holotype male. “ UTAH: San Juan Co. / Gooseneck St. Reserve / 26 June 1982 coll. / J.E. and M.S. Heath ”, male ( INHS).

Paratypes. Eight males same data as holotype ( MSHC) ; eight males “ UTAH: San Juan Co. // 1 mi N of Mexican / Hat on US 163 / 22 June 1984? ( MSHC); two males “ UTAH: San Juan Co. / 1 / 4 mi. N. Mexican Hat / 3 July 1986 / Al Sanborn coll. // recorded” ( MSHC) ; eight males “ UTAH: San Juan Co. / 1 / 4 mi. N. Mexican Hat / 3 July 1986 / Al Sanborn coll.” ( MSHC) ; one male “ UTAH: San Juan Co. / 1 / 4 mi. N. Mexican Hat / 3 July 1986 / J.E. Heath coll.” ( MSHC) ; one female “ UTAH: San Juan Co. / 1 / 4 mi. N. Mexican Hat / 3 July 1986 / M.S. Heath coll.” ( INHS) ; six males “ UTAH: San Juan Co. / Junction Utah 261 and / U.S. 163; 3 July 1986 / Al Sanborn coll.” ( MSHC) ; six males and one female “ UTAH: San Juan Co. / Gooseneck St. Reserve / 3 July 1986 / Al Sanborn coll.” ( MSHC) ; one male “ UTAH: San Juan Co. / Goosenecks of the / San Juan. 18 June / 1990. M.S., J.E., and C.M. / Heath coll. Ex. Grass // recorded specimen” ( MSHC) ; five males “ UTAH: San Juan Co. / Goosenecks of the / San Juan. 18 June / 1990. M.S., J.E., and C.M. / Heath coll. Ex. Grass // recorded specimen” ( MSHC) ; two males “ UTAH: San Juan Co. / Goosenecks of the / San Juan River. 2 July / 1989. Al Sanborn coll. // recorded” ( MSHC) ; one male “ UTAH: San Juan Co. / Goosenecks State Park / 30 June 1997 / James Diewald coll.” ( MSHC) ; one male “ UTAH: San Juan Co. / Goosenecks State Park / 30 June 1997 / Barbara Diewald coll.” ( MSHC) ; two males “ UTAH: San Juan Co. / Goosenecks State Park / 21 June 2007 / Kathy Hill and Dave / Marshall coll.” ( MSHC) ; one male “ UTAH: San Juan Co. / Goosenecks State Park / 21 June 2007 / Dave Marshall and / Kathy Hill coll.” ( MSHC) ; seven males and one female “ UTAH San Juan Co. / 9 mi. W Montezuma Cr. / 8 July 1970 / J.L. Petty ” ( BYUC 5 males and one female, AFSC 2 males) ; one male “ USA: UTAH: San Juan Co. / Goosenecks of the San Juan. / 3.6 mi SW of Rt 261 on Rt 316 / 37 ◦ 10.504N 109 ◦ 55.635W 5052 ft / 21 Jun 2007 D. Marshall, K. Hill, / M. and J. Heath US. UT.GOS // HILL / MARSHALL VOUCHER / pinned specimen, legs in EtOH / 07. US. UT.GOS.01 / Beameria venosa ? / Specimen recorded” ( AFSC) ; one male “ USA: UTAH: San Juan Co. / Goosenecks of the San Juan. / 3.6 mi SW of Rt 261 on Rt 316 / 37 ◦ 10.504N 109 ◦ 55.635W 5052 ft / 21 Jun 2007 D. Marshall, K. Hill, / M. and J. Heath US. UT.GOS // HILL / MARSHALL VOUCHER / pinned specimen, legs in EtOH / 07. US. UT.GOS.02 / Beameria venosa ? / Specimen recorded” ( AFSC) ; one male “ USA: UTAH: San Juan Co. / Goosenecks of the San Juan. / 3.6 mi SW of Rt 261 on Rt 316 / 37 ◦ 10.504N 109 ◦ 55.635W 5052 ft / 21 Jun 2007 D. Marshall, K. Hill, / M. and J. Heath US. UT.GOS” ( AFSC) ; nine males and two females “ UTAH: San Juan Co. / 1 mi. E of US 163 on Co. 216A / 14 Jul 1991 / A. Sanborn, P. Phillips coll.” ( AFSC) ; one male and one female “ UTAH: San Juan Co. / 3.5 mi. W of UT 261 on UT 316 / 14 Jul 1991 / A. Sanborn, P. Phillips coll.” ( AFSC) ; one male and one female “ UTAH: San Juan Co. / 9.5 mi. E of Goosenecks State Park / on UT 261 37 ◦ 11.73’N 109 ◦ 52.29W / 28 Jun 1998 / A. Sanborn and P. Phillips coll.” ( AFSC) ; four males and one female “ UTAH: San Juan Co. / 9.5 mi. E of Goosenecks State Park / on UT 261 37 ◦ 11.73’N 109 ◦ 52.29W / 28 Jun 1998 / A. Sanborn coll.” ( AFSC) ; one male “ UTAH: San Juan Co. / 19 mi. SE of Hite Crossing, Colorado River / on UT 95 37 ◦ 41.44’N 110 ◦ 13.56’W / 28 Jun 1998 / A. Sanborn coll.” ( AFSC) ; one male “ UTAH: San Juan Co. / 17.3 mi. SE of Hite Crossing, Colorado River / on UT 95 37 ◦ 43.52’N 110 ◦ 14.94’W / 28 Jun 1998 / P. Phillips coll.” ( AFSC) .

Etymology

The species name is a composition of anser (Latin for goose) and collis (Latin for neck). The name is in reference to the type location overlooking the entrenched meander forming the Goosenecks of the San Juan River.

Measurements (mm)

N = 25 males or 7 females, mean (range). Length of body: male 13.11 (12.0–14.5), female 14.0 (12.8–14.8); length of tegmina: male 16.08 (14.2–17.6), female 16.69 (15.9– 17.2); width of tegmina: male 5.43 (4.9–7.0), female 5.71 (5.3–6.0); length of head: male 2.24 (2.1–2.4), female 2.31 (2.1–2.6); width of head including eyes: male 4.37 (4.1–4.6), female 4.52 (4.35–4.7); width of pronotum including suprahumeral plates: male 5.04 (4.6–5.2), female 5.10 (4.8–5.4); width of mesonotum: male 4.36 (3.9–4.6), female 4.52 (4.35–4.7).

Comparative morphometric data are summarized in Table 1. Mean body mass for the three species of Beameria differs significantly ( ANOVA, F = 178.32, d.f. = 2, 427, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 13.274, P <0.001), B. ansercollis sp. nov. and B. wheeleri (q = 25.188, P <0.001) and B. venosa and B. wheeleri (q = 10.600, P <0.001). However, the B. venosa population did not pass the Kolmogorov and Smirnov test for Gaussian distributions ( KS = 0.1161, P = 0.0006) so a nonparametric test was performed. The Kruskal– Wallis test is also highly significant ( KW = 171.37, P <0.0001). The Dunn’s multiple comparisons test shows significant differences between B. ansercollis sp. nov. and B. venosa (P <0.001), B. ansercollis sp. nov. and B. wheeleri (P <0.001) and B. venosa and B. wheeleri (P <0.001). Body length is significantly different in the three species of Beameria ( ANOVA, F = 43.471, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 5.639, P <0.001), B. ansercollis sp. nov. and B. wheeleri (q = 9.446, P <0.001) and B. venosa and B. wheeleri (q = 12.977, P <0.001). Wing length is significantly different in the three species of Beameria ( ANOVA, F = 40.492, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 8.255, P <0.001), B. ansercollis sp. nov. and B. wheeleri (q = 6.486, P <0.001) and B. venosa and B. wheeleri (q = 12.680, P <0.001). Wing width differs significantly in the three species of Beameria ( ANOVA, F = 44.009, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 5.234, P <0.01), B. ansercollis sp. nov. and B. wheeleri (q = 9.836, P <0.001) and B. venosa and B. wheeleri (q = 12.963, P <0.001). Head length is significantly different in the three species of Beameria ( ANOVA, F = 18.361, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there is no difference between B. ansercollis sp. nov. and B. venosa (q = 2.093, P> 0.05) but there are differences between B. ansercollis sp. nov. and B. wheeleri (q = 7.182, P <0.001) and B. venosa and B. wheeleri (q = 7.979, P <0.001). Head width differs significantly in the three species of Beameria ( ANOVA, F = 17.366, d.f. = 2, 46, P <0.0001). A Tukey– Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 4.643, P <0.01), B. ansercollis sp. nov. and B. wheeleri (q = 5.042, P <0.01) and B. venosa and B. wheeleri (q = 8.331, P <0.001). Pronotum width differs significantly in the three species of Beameria ( ANOVA, F = 46.347, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 12.596, P <0.001) and B. venosa and B. wheeleri (q = 11.529, P <0.001) but not B. ansercollis sp. nov. and B. wheeleri (q = 0.8065, P> 0.05). Mesonotum width differs significantly in the three species of Beameria ( ANOVA, F = 60.466, d.f. = 2, 46, P <0.0001). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. venosa (q = 12.682, P <0.001), B. ansercollis sp. nov. and B. wheeleri (q = 4.402, P <0.01) and B. venosa and B. wheeleri (q = 14.696, P <0.001).

Diagnosis

Beameria ansercollis sp. nov. is intermediate in size between B. venosa and B. wheeleri . The male operculum of B. ansercollis sp. nov. reaches to the middle of sternite II while it does not reach sternite II in B. venosa and reaches the anterior margin of sternite II in B. wheeleri . The lateral uncus lobes of B. ansercollis sp. nov. do not curve away from the median uncus lobes laterally so they are less lobate than those of B. venosa , nor do they have a lateral notch as found in B. wheeleri . Female sternite VII is not deeply notched nor is the lateral margin expanded posteriorly as is found in the other Beameria species. The medial angle of the female operculum extends posterior to the meracanthus in B. ansercollis sp. nov. but does not expand posterior to the meracanthus in the other species of Beameria . In addition, any of the morphological measurements provided above can be used to differentiate the species from its cogeners, with the exception of head length differentiating B. ansercollis sp. nov. from B. venosa , or pronotum width differentiating B. ansercollis sp. nov. from B. wheeleri .

Description

Colouration. Ground colour testaceous marked with tawny ( Figure 1A ).

Head ( Figure 1B ). Not as wide as mesonotum. Testaceous except tawny marks surrounding ocelli, spot on either side of midline of frontoclypeal suture, light mark on medial supra-antennal plate, two small spots between lateral ocellus and posterior eye and lateral vertex, between frontal margin and medial angle of eye, which extends along medial ventral margin of eye and gena terminating at anterior fore coxa. Long silvery pile posterior to and posteriomedial to eye. Scape and proximal pedicel testaceous, remaining antenna tawny. Gena testaceous except for tawny longitudinal mark. Lorum tawny with testaceous lateral and anterior margins. Ventral head covered with long silvery pile. Postclypeus with testaceous spot along frontoclypeal suture and medial elongate spot on apex. Transverse grooves tawny, transverse ridges testaceous. Short silvery pile in lateral transverse grooves and transversly across posterior postclypeus. Anteclypeus tawny except testaceous anterior border and midline. Rostrum testaceous laterally, tawny medially, tip piceous reaching to middle of hind coxae.

Thorax ( Figure 1B ). Pronotum testaceous with tawny mark along lateral ambient fissure, connecting to spot on pronotal collar anterior angle. A pair of tawny marks on either side of the anterior pronotum in most paratypes. Fine silvery pile medial to paramedian fissure on anterior pronotum and lateral pronotal collar. Mesonotum with tawny irregular spots in lateral sigilla and submedian sigilla. Tawny spots in scutal depression and across anterior portion of anterior arm of cruciform elevation. Long silvery pile in posterior wing groove and depressions of cruciform elevation. Ventral surface with fine, white pile and light waxy pubescence. Ventral plates testaceous except tawny basisternum 2, medial meron 2, medial basisternum 2 and spots on posterior trochantins 2 and 3.

Legs. Testaceous striped with tawny. Coxae with small tawny spot anteriorly and proximally, middle and hind coxae with small distal spot medially and laterally. Coxae with fine, white pile. Trochanters testaceous. Femora with tawny stripes, distally with small medial and lateral spot. Primary spine slightly angled with tawny tip, secondary spine upright, apical spine small. Tibial spurs tawny. Tarsi testaceous with tawny spot on distal mesotarsus and distal third of pretarsus tawny. Claws dark testaceous proximally, tips tawny.

Tegmina and wings ( Figure 1A ). Tegmina hyaline with eight apical cells, infuscation on radial, radiomedial and medial cross-veins. Venation ochraceous proximally, darkening past node to black distally. Anal vein 2 + 3 black. Small linear tawny mark at wing base. Wings hyaline with six apical cells. Venation ochraceous.

Operculum. Male operculum testaceous reaching to anterior sternite II ( Figure 1D ). Medial margins well separated, posterior border smoothly rounded. Female operculum ochraceous covered with white pile and white pruinosity ( Figure 1E ). Posterior margin smoothly curved, extending posteriorly to middle of sternite II and medially to lateral meracanthus. Meracanthus ochraceous with tawny spot at base. Operculum covered with fine, white pile, thicker anteriorly and laterally. Meracanthus testaceous with white pile at base.

Abdomen ( Figure 1A ). Dorsal abdomen tawny except testaceous tergite 1, medial tergite 2, posterior medial tergites 2–6 and posterior tergites 7 and 8. Timbal cover testaceous with tawny spot medially and tawny mark on posterior lateral surface, incomplete covering less than one quarter of timbal ( Figure 1C ). Timbal with three ribs. Fine silvery pile dorsally, thicker lateral to midline on tergites 2–4 and on anterior of lateral tergite 3. Sparse, long pile on tergite 8. White, waxy pubescence across anterior tergite 2 posterior to tergite 1 and lateral tergite 8. Sternite II testaceous, sternites III–VII tawny anteriorly and testaceous posteriorly, sternite VIII testaceous. Epipleurites same coloration as sternites. Ventral abdomen covered with fine, silvery pile.

Male genitalia ( Figures 1F,G ). Pygofer tawny anteriorly, testaceous posteriorly. Dorsal beak absent. Distal shoulder with fuscous recurved point. Pygofer basal lobes small, rounded, flattened against posterior pygofer. Median uncus lobe curved ventrally at apex, with terminal notch. Lateral uncus lobe rounded ventrally and laterally, diverging ventrally. Aedeagus castaneous.

Female genitalia ( Figures 1H,I ). Abdominal segment 9 ochraceous with light tawny band on dorsolateral surface which can be reduced to a spot or expanded across dorsal surface in some paratypes, covered with long white pile. Sternite VII ochraceous with tawny spot laterally which expands anteriorly to margin in some paratypes, posterior margin with a single medial notch, expanding laterally. Gonapophysis VIII ochraceous with tawny base. Gonapophysis IX tawny. Gonapophysis X ochraceous extending to level of anal styles. Dorsal beak small, not reaching anal styles.

Biogeography

Beameria ansercollis sp. nov. is currently only known from several locations in San Juan County, Utah ( Figure 2 ). We were able to expand the known distribution of B. wheeleri from Otero County , New Mexico to include Dona Ana , Grant , Hidalgo, Luna , and Socorro Counties in New Mexico, Cochise County , Arizona, and Mexico . Beameria venosa has the greatest distribution, being found in nine states and Mexico.

No specific host plants for either B. ansercollis sp. nov. or B. venosa were able to be determined. Both species were associated with multiple grass species in different habitats. They will also call from larger plant species (e.g. Artemesia spp. , Larrea spp. ) in the immediate area.

We determined B. wheeleri to be associated with Sporobolus R. Br. grass. The distribution of B. wheeleri is dependent on the presence of Sporobolus in the environment. The cicadas disappear when the grass disappears from the environment.

Song analysis

Calling song parameters are summarized in Table 2. The song of B. ansercollis sp. nov. is a continuous train of sound pulses ( Figure 3 ) similar to B. venosa (Sanborn et al. 2009) . The song of B. wheeleri is a repeating series of syllables ( Figure 4 ). Peak song frequency differs significantly between the Beameria species (ANOVA F = 5.499, d.f. = 2, 31, P = 0.009). A Tukey–Kramer multiple comparison test shows there are no statistically significant differences between peak song frequency of B. venosa and B. ansercollis sp. nov. (q = 0.6238, P> 0.05) but there are differences between B. ansercollis sp. nov. and B. wheeleri (q = 4.671, P <0.01), and B. venosa and B. wheeleri (q = 4.345, P <0.05).

Pulse repetition rates show statistically significant differences between the Beameria species (ANOVA F = 12.507, d.f. = 2, 31, P = 0.0001). A Tukey–Kramer multiple comparison test shows there are statistically significant differences between B. venosa and B. ansercollis sp. nov. (q = 3.253, P <0.05), B. ansercollis sp. nov. and B. wheeleri (q = 6.735, P <0.001), and between B. venosa and B. wheeleri (q = 4.994, P <0.001).

The song power data also differ significantly between the Beameria species (ANOVA F = 9.358, d.f. = 2, 42, P = 0.0004). A Tukey–Kramer multiple comparison test suggests there are no statistically significant differences between B. venosa and B. ansercollis sp. nov. (q = 1.374, P> 0.05), but there are differences between B. venosa and B. wheeleri (q = 6.053, P <0.001), and between B. ansercollis sp. nov. and B. wheeleri (q = 4.148, P <0.05). The Bartlett’s statistic (8.967, P = 0.0113) and the Kolmogorov and Smirnov test (P = 0.0306 for B. venosa and P = 0.0197 for B. ansercollis sp. nov.) suggested performing a nonparametric Kruskal– Wallis test. The Kruskal– Wallis statistic, KW = 16.051, suggests statistically significant differences (P = 0.0003). The Dunn’s multiple comparisons test suggests there are no significant differences between B. venosa and B. ansercollis sp. nov. (mean difference = −2.208, P> 0.05), but there are differences between B. venosa and B. wheeleri (mean difference = −18.417, P <0.001) and between B. ansercollis sp. nov. and B. wheeleri (mean difference = −16.208, P <0.01).

Thermal response B. venosa B. ansercollis B. wheeleri ANOVA P sp. nov. F (d.f.)

Minimum flight temperature (

◦

C) 21.9 ± 1.62 21.8 ± 2.00 22.8 ± 1.87 2.377 0.0951

n = 179 n = 36 n = 19 (2, 231) Maximum voluntary tolerance 37.3 ± 3.00 37.9 ± 2.92 36.2 ± 2.35 3.988 0.0194 temperature (◦ C) n = 235 n = 60 n = 41 (2, 333) Heat torpor temperature (◦ C) 45.5 ± 1.75 46.3 ± 1.57 46.3 ± 1.71 9.112 0.0001

n = 239 n = 58 n = 58 (2, 352)

Thermal responses

The thermal responses of the three Beameria species are summarized in Table 3. The minimum flight temperature does not differ significantly between the Beameria species (ANOVA F = 2.377, d.f. = 2, 231, P = 0.0951). Maximum voluntary tolerance temperature, however, does differs significantly between the Beameria species (ANOVA F = 3.988, d.f. = 2, 333, P = 0.0194). A Tukey–Kramer multiple comparison test shows there are differences between B. ansercollis sp. nov. and B. wheeleri (q = 3.980, P <0.05) but not between B. venosa and B. ansercollis sp. nov. (q = 1.990, P> 0.05) or B. venosa and B. wheeleri (q = 3.064, P> 0.05). The heat torpor temperature shows a significant difference between Beameria species (ANOVA F = 9.112, d.f. = 2, 352, P = 0.0001). A Tukey–Kramer multiple comparison test suggests there are differences between B. venosa and B. ansercollis sp. nov. (q = 4.579, P <0.01) and B. venosa and B. wheeleri (q = 4.753, P <0.01), but not between B. ansercollis sp. nov. and B. wheeleri (q = 0.1375, P> 0.05).

Figures

Figure 1. Beameria ansercollis sp. nov. (A) Holotype male (top left) from USA, Utah, San Juan County, Goosenecks State Preserve; paratype female (below right) from USA, Utah, San Juan County, 1 mile east of US 163 on County 216A; (B) paratype, dorsum illustrating head and thoracic markings; (C) holotype male, timbal and timbal cover; (D) paratype male, operculum; (E) paratype female, operculum; (F) lateral view of paratype male genitalia; (G) posterior view of paratype male genitalia; (H) lateral view of paratype female genitalia; (I) ventral view of paratype female genitalia. Scale bars: (A), 2 cm; (B, H, I), 2 mm; (C–G) 1 mm.

Figure 2. Distribution of the Beameria species. Beameria ansercollis sp. nov. (red triangles) is currently only known from San Juan County, Utah. The known distribution of B. wheeleri (blue squares) was expanded from Otero County, New Mexico to include five additional New Mexico counties, and locations in Arizona and Mexico. Beameria venosa (green circles) is found in nine US states and northern Mexico. [This figure can be viewed in colour online].

Figure 3. Oscillogram (upper trace) and sonogram (lower trace) of the calling song produced by B. ansercollis sp. nov. The call is a series of syllables of constant frequency and amplitude, with a peak frequency of about 15.7 kHz, with individual sound pulses being produced at a rate of about 375 Hz.

Figure 4. Oscillogram (upper trace) and sonogram (lower trace) of the calling song produced by B. wheeleri. The call is a series of syllables of constant frequency and amplitude with a peak frequency of about 13.6 kHz. Syllable duration is about 86 ms with individual sound pulses being produced at a rate of about 450 Hz.