Pacarina shoemakeri Sanborn and M. Heath, 2012

Sanborn, Allen F., Heath, Maxine S., Phillips, Polly K. & Heath, James E., 2012, The genus Pacarina Distant, 1905 (Hemiptera: Cicadidae) with the description of a new species, Journal of Natural History 46 (15 - 16), pp. 923-941 : 927-936

Pacarina shoemakeri Sanborn and M. Heath sp. nov.

Synonymy

Pacarina puella Davis (juniper variety), Sanborn and Phillips, 1995: 481.

Pacarina puella Davis, Kondratieff et al., 2002: 7 , 13, 28.

Pacarina shoemakeri Sanborn and Heath, Sanborn et al., 2011a: 152 , 153. Type series

Holotype male. “ARIZ: Apache Co. / US 180 – 12 m (iles) [19.2 km] S of St Johns / 28 June 1979 / P. Heath, B. Murphy / coll. EX: Juniper // Pacarina / puella” ( INHS).

Paratypes. Three males and two females same data as holotype (all MSHC except one female INHS) ; two males “ARIZ: Coconino Co. / I-40 – Welsh Rd Ramp / 2 July 1981 / Pamela Heath Coll. / mite infected // Pacarina / puella / alcohol bleached / probably Juniper” ( MSHC) ; two males “ARIZ: Yavapai Co. / 11 / 2 mi [2.4 km] SW of Junction / I-17 and Ariz. 169 / 30 June 1986 / Al Sanborn coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; two males “AZ: Yavapai Co. / 18 July 2008 / B. Kondratieff, / D. Leatherman and / W. Cranshaw / Rd 5, Williamson Valley / Mp 53” ( CSUC) ; two males “CO: Las Animas Co. / 13 July 1991 / M. Kippenhan, B. Kondratieff, D. / and M. Leatherman and Paul Opler cols. // Iron Cyn / Steinman’s Ranch / (Rourke Ranch) / PCMS / in Juniper” ( CSUC) ; one male “CO Las Animas Co. / PCMS, Iron Canyon / Steinman Ranch , Purgarorie Ri. / 14-VII- 1991 / BC Kondratieff and PA Opler // Pacarina puella / Davis / Det: BC Kondratieff 1991” ( CSUC) ; one male “ Jeff Davis Co., TX / 15 June 1992 / Kondratieff and Baumann / Hwy 18, S of Ft. Davis ” ( CSUC) ; two males “ Sidona (sic), Coconino Co. / Ariz. VI-15-61 / S.G. Jewitt, Jr. // Pacarina / puella / Davis / Det. TEMoore 1963” ( UMMZ) ; five males “ARIZ: Navajo Co. / AZ 77- 10 mi [16 km] No of / Showlow 31 (sic) June 1980 / J. Heath coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; two males “ARIZ: Yavapai Co. / AZ 179 just So of / Sedona 29 June 1981 / JE and MS Heath coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; one male “ARIZ: Coconino Co. / Schnebly Hill Rd. / 1 mi. [1.6 km] E. of Sedona / 26 June 1991. M.S. / Heath coll. EX: Juniper // Pacarina / puella / Juniper var. // Sent to BM / did not / match type” ( MSHC) ; one male “ARIZ: Coconino Co. / AZ 179 - 1 mi [1.6 km] No of / Bell Rock 30 Jun / 1987. Al Sanborn Coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; one male “ARIZ: Coconino Co. / AZ 179 - 1 mi [1.6 km] No of / Bell Rock 30 Jun / 1987. Al Sanborn / Coll. EX: Juniper // Pacarina / puella” ( MSHC) ; one male “ARIZ: Cochise Co. / 1 mi [1.6 km] E Paradise / 25 June 1985 / J.E. and M.S. Heath Coll. / EX: Juniper // Pacarina / puella // Recorded” ( MSHC) ; one male “ARIZ: Cochise Co. / Chiricahua Mts. / 26 June 1983 Coll. / C. Heath-Smith // Pacarina / puella” ( MSHC) ; one male “ ARIZONA, Coconino Co. / Sedona / 12 JUL 1991 / A. Sanborn, P. Phillips coll. // recorded specimen // Compared to Type Specimen of P. puella ” ( AFSC) ; two males “ ARIZONA, Coconino Co. / Sedona / 12 JUL 1991 / A. Sanborn, P. Phillips coll. // recorded specimen” ( AFSC) ; one male “ ARIZONA, Pima Co. / 12.6 mi [20 km] S Mt. View on AZ 83 / 31 May 1995 / A. Sanborn coll.” ( AFSC) ; one male “ ARIZONA, Apache Co. / 9 mi [14.4 km] NE of St. Johns on US 180 / 26 June 1998 / A. Sanborn coll. // Pacarina puella (juniper)” ( AFSC) ; one male “ ARIZONA, Apache Co. / 9 mi [14.4 km] NE of St. Johns on US 180 / 26 June 1998 / P. Phillips coll. // Pacarina puella (juniper)” ( AFSC) ; one male “ ARIZONA, Apache Co. / 35 mi [56 km] SE of Holbrook on US 180 / 26 June 1998 / A. Sanborn and P. Phillips coll. // Pacarina puella (juniper)” ( AFSC) ; four males “ ARIZONA, Navajo Co. / 9.3 miles [14.9 km] NE White River on Apache 55 / 33 ◦ 49.65 ′ N 109 ◦ 51.59’ W 5570 ft. [1698 m] / 24 June 2001 / A. Sanborn coll.” ( AFSC) ; one male “ ARIZONA, Navajo Co. / 9.3 miles [14.9 km] NE White River on Apache 55 / 33 ◦ 49.65 ′ N 109 ◦ 51.59 ′ W 5570 ft. [1698 m] / 24 June 2001 / M. Nouzova coll.” ( AFSC) ; one female “ ARIZONA, Navajo Co. / 0.5 miles [0.8 km] North Ft. Apache on AZ 73 / 33 ◦ 48.03 ′ N 109 ◦ 59.68 ′ W 5189 ft. [1582 m] / 24 June 2001 / A. Sanborn coll.” ( AFSC) ; one male “ ARIZONA, Gila Co. / ∼ 21 miles [33.6 km] North of Globe on AZ 77 / 33 ◦ 39.90 ′ N 110 ◦ 34.38 ′ W 4656 ft. [1419 m] / 23 June 2001 / A. Sanborn coll.” ( AFSC) ; one male “ NEW MEXICO, Catron Co. / US 180 10.4 mi [16.6 km] N of Alma / 20 June 1998 / A. Sanborn coll. // Pacarina puella (juniper)” ( AFSC) ; one male “ TEXAS, Brewster County / 7.2 mi [11.5 km] S of Alpine on TX 118 / 28 May 1995 / A. Sanborn coll. ex. Juniper” ( AFSC) ; one male “ TEXAS, Brewster County / 8 mi [12.8 km] S of Alpine / 7 July 1982 / JE and MS Heath // Pacarina / puella” ( MSHC) ; one male “ TEXAS, Jeff Davis County / 1.7 mi [2.7 km] NW of Fort Davis on TX 118 / 19 June 1998 / A. Sanborn, M. Heath coll. // Pacarina puella (juniper)” ( AFSC) ; three males “ TEXAS, Jeff Davis Co. / 1.7 mi [2.7 km] N of Fort Davis / on Texas highway 118 / 18 June 1998 / Al Sanborn coll. // Pacarina ex. Juniper” (two MSHC, one AFSC) ; two males “ TEXAS: Jeff Davis Co. / 4 mi. [6.4 km] So. of Fort Davis / 25 June 1983. M.S. Heath Coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; two males “N. MEX: San Miguel / Co. 4.5 mi. [7.2 km] No. of / Dilia. 21 Jun. 1991 / M.S. and J.E. Heath Coll. / EX: Juniper // Pacarina / puella / Juniper” ( MSHC) ; one male “N. MEX: San Miguel Co. / 4.5 mi. [7.2 km] No. of Dilia / 21 Jun. 1991 / M.S. and J.E. Heath Coll. / EX: Juniper // Pacarina / puella / Juniper” ( MSHC) ; three males “N. MEX: Guadalupe Co. / 1 mi. [1.6 km] W of Cuervo / 5 July 1984 / ex: Juniper // Pacarina / puella” ( MSHC) ; two males “N. MEX: Guadalupe Co. / 5.3 mi. [8.5 km] E of Newkirk / 23 June 1987 / Al Sanborn Coll. / EX: Juniper // Pacarina / puella // Recorded / specimen” ( MSHC) ; one male “N. MEX: Guadalupe Co. / 5.3 mi. [8.5 km] E of Newkirk / 23 June 1987 / A. Sanborn Coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; two males and one female “N. MEX: Guadalupe Co. / 5.3 mi. [8.5 km] E of Newkirk / 23 June 1987 / J. Breitbarth, Coll. / EX: Juniper // Pacarina / puella” ( MSHC) ; one male “N. MEX: Guadalupe / Co. 5.3 mi. [8.5 km] E of / Newkirk . 23 June / 1987. M.S. Heath Coll. / EX: Juniper // Pacarina / puella” ( MSHC) .

Etymology

The species is named in honour of Eugene Shoemaker, the brother of M.S.H., known among his peers as the father of astrogeology, to whom the Colorado Plateau and its ecology were dear.

Measurements (mm)

Twenty males and five females (n = 25), mean (range). Length of body: male 13.36 (12.3–14.7), female 14.16 (13.0–15.9); length of forewing: male 16.84 (15.0–18.7), female 17.88 (17.0–19.0); width of forewing: male 5.59 (5.0–6.0), female 5.86 (5.6–6.1); length of head: male 2.20 (1.9–2.4), female 2.12 (2.0–2.3); width of head including eyes: male 5.60 (5.2–6.0), female 5.92 (5.6–6.4); width of pronotum including suprahumeral plates: male 5.38 (4.7–5.6), female 5.84 (5.4–6.4); width of mesonotum: male 4.40 (4.3–4.6), female 4.92 (4.4–5.4).

Comparative morphometric data for P. shoemakeri sp. nov. and P. puella are provided in Table 1. Some of the data sets did not pass the Kolmogorov and Smirnov test for Gaussian distributions so a non-parametric Mann–Whitney U -test was performed with those data sets. Male body length (U = 122.50, U ′ = 277.50, P = 0.0370), head length (U = 86.00, U ′ = 314.00, P = 0.0017) and wingspan (U = 70.50, U ′ = 321.50, P = 0.0005) show statistically significant differences using the Mann–Whitney test whereas female body length (U = 13.00, U ′ = 22.00, P = 0.5138), pronotum width (U = 184.50, U ′ = 215.50, P = 0.6671) and mesonotum width (U = 182.00, U ′ = 218.00, P = 0.6162) showed no statistically significant differences between the species. Male live mass data sets did not pass the assumption test for equal standard deviations so a Welch correction t -test was performed but the means were still not statistically different (Welch’s approximate t = 0.8455, d.f. = 34, P = 0.4037).

Diagnosis

The species most similar to P. shoemakeri sp. nov. is P. puella . Male body length, forewing length, forewing width, head length, and wingspan show statistically significant differences between the species ( Table 1). Pacarina shoemakeri sp. nov. can be distinguished from P. puella by the hourglass-shaped mark on the medial pronotum, the greater number of pronotal markings, the black abdominal markings, the angulate anteromedial margin of the timbal cover rather than being smoothly arched, the secondary spine being upright rather than angled, the ventral extension of the median uncus lobes being longer than the dorsal extension, and the posterior margin of sternite VII in the female with a V-shaped notch rather than a smooth, shallow depression. Finally, the habitat where a specimen was collected can be used to differentiate the species as the two species are associated with different plant communities (see below).

Pacarina championi can be distinguished from P. shoemakeri sp. nov. by its larger size (body length> 16 mm), the head generally lacks a complete transverse black band, the anterior margin of the forewing is angled more strongly at the node, the distal forewing is proportionately longer with a ratio of the length of the forewing beyond the node to the length of the base to the node of about 0.94 rather than about 0.81, the ventrolateral timbal cover conceals the timbal, the lateral operculum is angled medially rather than being parallel to the long body axis, the lateral uncus lobes are not recurved, and female sternite VII has a shallow, curved depression rather than a V-shaped notch.

Pacarina schumanni also has a greater angle on the anterior margin of the forewing and a ventrolateral timbal concealed by the timbal cover compared with P. shoemakeri sp. nov. In addition, P. schumanni can be differentiated from P. shoemakeri sp. nov. by reduced black markings on the head and thorax, the central black stripe on the medial male opercula, the median uncus lobes lack ventral extensions, and sternite VII of the female has a shallow, curved depression rather than a V-shaped notch.

Description

Coloration. Ground color of head and abdomen black marked with testaceous, thorax testaceous marked with black ( Figure 1A ).

Head ( Figure 1B ). Head 1.01–1.04 times wider than mesonotum. Black except angular testaceous area between anteriomedial margin of eye and posterior head and light testaceous mark along anterior epicranial suture anterior arm along supraantennal plate. Two small black spots between lateral ocellus and posterior eye. Transverse testaceous lines anteriorly between supra-antennal plate and ventrally between postclypeus and ventral angle of eye, neither line reaching medial eye margin. Long silvery pile posterior to and along hind margin of head. Sparse, golden pile on dorsal head. Gena testaceous. Lorum black medially, tawny laterally, with testaceous lateral and anterior margins. Ventral head covered with long silvery pile, golden lateral to antennae. Antennal segments black, proximal scape and pedicel tawny in some paratypes. Postclypeus testaceous except black dorsal surface and stripe along ventral midline, reduced in some paratypes. Anteclypeus black with testaceous line medially expanding laterally on anterior margin. Mentum testaceous medially, tawny laterally. Labium testaceous proximally, tawny distally with piceous tip reaching to middle of hind coxae. White pubescence on gena, lorum, postclypeus transverse grooves and midline, lateral anteclypeus, very dense in some paratypes.

Thorax ( Figure 1B ). Pronotum testaceous with black mark expanding anteriorly and posteriorly into hourglass shape on midline. Black marks in paramedian fissure, lateral fissure connecting to mark in ambient fissure both anteriorly and posteriorly. Light tawny mark in posterior ambient fissure between central stripe and line connecting to posterior lateral fissure. Longitudinal mark extending posteriorly from middle of paramedial fissure not reaching pronotal collar. Pronotal collar testaceous with anterior margin along midline stripe tawny, and fuscus lateral angle. White pile in fissures, dense posterior to eye, and sparsely along anterior margin and on notum in some paratypes. Mesonotum with black submedian sigilla, lateral sigilla and two elongated spots on lateral surface. Fuscus spot arising between submedian sigillae, expanding laterally to anterior arms of cruciform elevation with transverse mark intersecting between submedian sigilla and scutal depression which expands anteriorly to submedial sigilla, expanding posteriorly forming a point connecting to fuscus scutal depression in some paratypes. Irregular tawny spots in lateral sigilla, also in submedian sigilla in some paratypes. Cruciform elevation black centrally and on proximal arms. Spots in scutal depression extend across anterior portion of anterior arm of cruciform elevation in some paratypes. Metanotum fuscus laterally. Long silvery pile in posterior wing groove and depressions of cruciform elevation, golden medially between anterior arms of cruciform elevation. Sparse white pile on anterior mesonotum along pronotal collar in some paratypes. Ventral surface with fine, white pile and waxy pubescence. Plates testaceous except black medial basisternum 2, medial katepimeron 2, episternum 2, and fuscus medial katepimeron 2, basisternum 3 and episternum 3.

Legs. Testaceous striped with tawny. Coxae ochraceous. Fore coxae tawny proximally, distally and anteriorly, middle and hind coxae with triangular spot anteriorly. Waxy, white pubescence on coxae. Trochanters testaceous, fore and middle trochanters with tawny stripes. Femora with tawny stripes, hind and middle femora ochraceous distally. Fore femora primarily tawny, testaceous mark proximolateral to primary spine. Primary spine angled, secondary spine upright, both fuscus. Fore tibia fuscus and tawny, middle and hind tibiae tawny, hind tibia fuscus distally. Tibial spurs fuscus. Tarsi tawny with lightly fuscus distal pretarsus. Claws testaceous proximally, tips tawny.

Forewing and wings ( Figure 1A ). Forewing hyaline with eight apical cells, infuscation on bifurcation of radius anterior 1 and 2, radial, radiomedial and medial cross-veins, and distal radius anterior 2, radius posterior, median veins 1–4 and cubitus anterior 1. Venation ochraceous proximally, darkening past node to black distally. Proximal costal margin with small spines, black. Black anteriorly within basal cell. Posterior anal vein 2 + 3 black. Tawny mark on sides of arculus. Membrane greyish. Wings hyaline with six apical cells. Venation ochraceous proximally, darkening distally. Anal veins 2 and 3 tawny, greyish along each side in anal cells 1–3.

Operculum. Male operculum testaceous, lightly fuscus at base and lateral margin, reaching to middle of sternite II ( Figure 1D ). Medial margins barely touching, posterior border smoothly rounded, lateral margins straight with slight lateral expansion at base. Female operculum castaneous on anterior half, posterior tawny, lateral margin edged in black ( Figure 1E ). Posterior margin rounded reaching to middle of sternite II. Operculum covered with white pubescence anteriorly and laterally. Meracanthus slender, elongated triangle, testaceous with tawny base.

Abdomen ( Figure 1A ). Tergite 1 testaceous, fuscus posteriorly, lateral fuscous marks in some paratypes. Tergite 2 testaceous, black anteriorly between timbals. Tergites 3–6 black with testaceous posterior margin, tergite 7 black with posterior third and tergites 8 with posterior half testaceous, testaceous portion of tergites 7 and 8 expanded in some paratypes. Timbal cover testaceous, incomplete, triangular, exposing timbals dorsally and laterally, curved mediad ( Figure 1C ). Timbal with four ribs, margin fuscus. Fine silvery pile dorsally, thicker on lateral portions of tergites, 3, 4, 7 and 8 and lateral to midline on anterior tergites 7 and 8. Sternites fuscous, sternites variously marked with tawny or testaceous in paratypes. Epipleurites same coloration as sternites. Ventral abdomen covered with sparse, long, silvery pile.

Male genitalia ( Figure 1F,G ). Pygofer fuscus anteriorly, testaceous posteriorly, with white pubescence and silvery pile laterally. Dorsal beak absent. Distal shoulder with fuscus recurved point. Pygofer basal lobes small, rounded, flattened against posterior pygofer. Uncus fuscus, rectangular with medial lobe recurved dorsally and bifurcating posteriorly. Lateral lobes expanding ventrally and laterally, margin rounded, recurved laterally. Aedeagus tawny.

Female genitalia ( Figure 1H,I ). Sternite VII tawny marked with castaneous on midline, posterior margin and lateral spots, single shallow V-shaped notch posteriorly. Abdominal segment 9 tawny, piceous anterior margin expanding onto dorsolateral segment and extending on either side of midline to posterior margin. Dorsal beak and anal styles piceous. Lateral abdominal segment 9 lightly castaneous with piceous lateral spot. Posterior ventral margin marked with piceous. Dorsolateral surface with short silvery pile. Ventral surface with sparse, long, erect golden pile. Gonocoxite IX castaneous with piceous medial margin. Gonopophysis IX ferruginous with piceous tip. Gonapophysis X piceous with long, erect golden pile extending to level of anal styles.

Biological notes

Pacarina shoemakeri sp. nov. is associated with various species of juniper ( Juniperus spp. ). Females were observed ovipositing in larger interior branches. Pacarina shoemakeri sp. nov. will congregate on juniper even when mesquite ( Prosopis spp. ) is present in the environment with only an occasional individual basking on unshaded mesquite branches when the ambient temperature is low. Males and females tend to congregate together on a few particular trees rather than separating in the habitat individually. When calling, P. shoemakeri sp. nov. raises its abdomen dorsally between the wings which are spread and depressed toward the perch. The species is active from late May to the middle of July.

Biogeography

The distributions of P. shoemakeri sp. nov. and P. puella are illustrated in Figure 2 . Pacarina shoemakeri sp. nov. uses juniper as a host plant and its distribution follows the distribution of juniper. It has currently been recorded from Arizona, New Mexico, Colorado and Texas. We found P. puella associated with mesquite. Pacarina puella has

been reported from Oklahoma, Texas, Louisiana and Arizona in the USA but extends southward into Central America ( Sanborn 2006, 2007, 2010) .

Song analysis

Calling song parameters are summarized in Table 2. The song of P. shoemakeri sp. nov. is a repeating series of syllables ( Figure 3 ) similar to P. puella ( Figure 4 ). Peak song frequency does not differ significantly between P. shoemakeri sp. nov. and P. puella (t = 0.0077, d.f. = 19, P = 0.9939). Syllable duration (t = 2.605, d.f. = 16, P = 0.0191), inter-syllable interval (t = 2.350, d.f. = 16, P = 0.0319), and syllable repetition rate (t = 4.973, d.f. = 16, P = 0.0001) all differ significantly between P. shoemakeri sp. nov. and P. puella and there is almost a statistically significant difference in pulse repetition rate (t = 1.998, d.f. = 15, P = 0.0642).

The song power data can only be partially analysed statistically because of the low sample size of the calling values. The data suggest that P. shoemakeri sp. nov. produces a calling song of greater intensity than P. puella . The power values for alarm calls produced by P. puella and P. shoemakeri sp. nov. do not differ significantly (t = 0.3658, d.f. = 36, P = 0.7166).

Thermal responses

The thermal responses for P. shoemakeri sp. nov. and P. puella are summarized in Table 2. The minimum flight temperature is the only variable to show statistically significant differences (t = 2.187, d.f. = 100, P = 0.0311). Some of the data sets did not pass the Kolmogorov and Smirnov test for Gaussian distributions so a nonparametric Mann–Whitney U -test was performed with those data sets. Minimum flight temperature (U = 602.00, U ′ = 1158.00, P = 0.0240) showed a statistically significant difference using the Mann–Whitney U -test while heat torpor temperature (U = 899.50, U ′ = 1205.50, P = 0.2637) showed no statistically significant differences between the species.

Figures

Figure 1. Pacarina shoemakeri Sanborn and M. Heath sp. nov. (A) Holotype male and paratype female habitus; (B) holotype male dorsum; (C) holotype male timbal cover; (D) paratype male operculum; (E) paratype female operculum; (F) paratype male lateral view of genitalia, arrows point to extensions of median uncus lobes which are diagnostic for the species; (G) paratype male posterior view of genitalia; (H) paratype female lateral view of genitalia; (I) paratype female ventral view of genitalia. Scale bars: A, 2 cm; B–E 2 mm; F–H, 1 mm.

Figure 2. Distribution of the Pacarina shoemakeri sp. nov. and Pacarina puella. Pacarina shoemakeri sp. nov. (diamonds) is currently known from Arizona, New Mexico, Colorado and Texas and is associated with juniper. Pacarina puella (circles) is associated with mesquite and is reported from Texas, Oklahoma and Arizona with its distribution extending southward into Central America.

Figure 3. Oscillogram (upper trace) and sonogram (lower trace) of the calling song produced by Pacarina shoemakeri sp. nov. The call is a series of syllables of constant frequency and amplitude with a peak frequency of about 16.4 kHz with individual sound pulses being produced at a rate of about 442 Hz within the syllables.

Figure 4. Oscillogram (upper trace) and sonogram (lower trace) of the calling song produced by Pacarina puella. The call is a series of syllables of constant frequency and amplitude with a peak frequency of about 15.6 kHz. Syllable duration is about 14 ms with individual sound pulses being produced at a rate of about 575 Hz within the syllables.