Guyalna Boulard & Martinelli, 1996

Sanborn, Allen F., 2016, The cicada genus Guyalna Boulard & Martinelli, 1996 (Hemiptera: Cicadidae: Cicadinae: Fidicinini): generic description, twelve new combinations, and a key to species, Zootaxa 4105 (5), pp. 430-454 : 431-452

Genus Guyalna Boulard & Martinelli, 1996

Type species. Fidicina bonaerensis Berg 1879: 140 ( Argentina) .

Species included. The genus is represented currently by the following species: G. atalapae Boulard & Martinelli, 2011 , G. bicolor ( Olivier, 1790), G. bleuzeni Boulard & Martinelli, 2011, G. bogotana ( Distant, 1892) n. comb., G. bonaerensis ( Berg, 1879) , G. b r i s a ( Walker, 1850) n. comb., G. chlorogena ( Walker, 1850) , G. c o f f ea ( Sanborn, Moore & Young, 2008) n. comb., G. c u t a ( Walker, 1850), G. densusa Boulard & Martinelli, 2011 , G. distanti ( Goding, 1925) n. comb., G. flavipronotum ( Sanborn, 2007a) n. comb., G. g l a u c a ( Goding, 1925) n. comb., G. jamesi Sanborn, 2016 , G. jauffreti Boulard & Martinelli, 2011 , G. maxineae Sanborn, 2016 , G. nadae Gogala, Šporar, Sanborn & Maccagnan, 2015 , G. nigra Boulard, 1999 , G. panamensis ( Davis, 1939) n. comb., G. parvula ( Jacobi, 1904) , G. platyrhina Sanborn & Heath, 2014 , G. rufapicalis Boulard, 1998 , G. sublaqueata ( Uhler, 1903) , G. variegata ( Sanborn, 2005) n. comb., and G. viridifemur ( Walker, 1850) n. comb.

Diagnosis. Within the Fidicinini, Guylana can be distinguished from the three genera ( Fidicina Amyot & Audinet-Serville, 1843 , Fidicinoides Boulard & Martinelli, 1996 , and Berglana Boulard & Martinelli, 1996) of the Fidicinina by the presence of three-part tarsi. Hemisciera Amyot & Audinet-Serville, 1843 is simply distinguished from all other Fidicinini by the bright green basal area of the fore wings and bright red on the basal area of the hind wings. Majeorona Distant, 1905 , has a head width wider than the mesonotum, with the prominent eyes stylate and extended anteriorly. The adpressed pronotal margins, non-parallel radial and radiomedial crossveins, relatively small eyes, small anterolaterally extended timbal covers, and the sternite separating the male opercula distinguish the members of Beameria Davis, 1934 . Pacarina Distant, 1905 can be distinguished by the prominent eyes that project laterally beyond the anterior edges of pronotum, the non-parallel radial and radiomedial crossveins, the abrupt facial color change, and triangular timbal covers. Similary, the genus Ollanta Distant, 1905 can be distinguished by the prominent eyes that project laterally beyond the anterior edges of the pronotum, the lobate lateral angles of the pronotal collar, the non-parallel radial and radiomedial crossveins, and triangular timbal covers. Ariasa Distant, 1905 , also has prominent eyes that project laterally beyond the anterior angles of the pronotum, the head is wider than the mesonotum, the abdominal tergites expand laterally to tergite 4 from base, and the median uncus lobes extend posteriorly in a straight line rather than bending as in Guyalna . The genera Proarna Stål, 1864 , Prasinosoma Torres, 1963 , Tympanoterpes Stål, 1861 , all possess a head that is about as wide as the mesonotum with the eyes barely or not extending beyond the anterior angle of the pronotum, radial crossvein vertically oriented to the radius anterior 2 and radius posterior veins with a radiomedial crossvein that is obliquely oriented, large semicircular male opercula, and triangular or small, curved timbal covers. The single species of Elassoneura Torres, 1964 shares most of the characters in these genera but the head is wider than the mesonotum. The single species of the genus Pompanonia Boulard, 1982 can be distinguished by a head that is wider than the mesonotum, the radial crossvein vertically oriented to the radius anterior 2 and radius posterior veins with a radiomedial crossvein that is obliquely oriented, semicircular male opercula, triangular timbal covers, and the very wide base of the abdomen with conspicuous timbals. Dorisiana Metcalf, 1952 is the most similar genus to Guyalna but can be distinguished by the head being as wide or wider than the mesonotum with eyes not extending beyond the lateral edges of the pronotum, triangular timbal covers, ventral margin of the timbal cover angled dorsally not parallel to the long body axis, and the lateral lobe of the uncus does not extend to the same lengths as the species of Guyalna .

The shape of the timbal cover is of particular importance in separating species of Guyalna and Dorisiana . The various interpretations of curved apex and triangular have led to species being assigned to incorrect genera. The expanded definition of the timbal cover possessed by members of Guyalna presented here is supported by the interpretation of Boulard, an authority for the genus Guyalna . After assigning G. bonaerensis ( Berg, 1879) and G. chlorogena ( Walker, 1850) to the genus when it was erected ( Boulard & Martinelli 1996), the next species transferred to the genus, G. cu t a ( Walker, 1850), and the second new species, G. nigra Boulard, 1999 (both in Boulard 1999) possess a timbal cover with a concave anteromedial margin and approximately parallel dorsal and ventral margins on the anterior extension above the lateral timbal with the ventral margin parallel to the long body axis before reaching the rounded apex that conceals only the lateral aspect of the timbal. The rounded apex even on the smaller timbal cover of these species clearly shows the interpretation of the generic authority was to include those species with a rounded timbal cover apex regardless of the size or position of the timbal cover. This interpretation has been maintained by the generic authorities with the descriptions of four species of Guyalna in their latest paper ( Boulard & Martinelli 2011) all of which possess similar timbal covers that extend over the lateral timbal cavity. The timbal cover of Dorisiana species is triangular or pointed if the timbal cover extends anteriorly and curves ventrally (as in the type species Fidicina semilata Walker, 1850 ) but the ventral anterior margin of the timbal cover is always angled dorsally away from the base and does not extend anteriorly parallel to the long body axis.

Description. Body size variable (12.9–36.7 mm body length). Head as wide as or slightly wider than mesonotum, eyes slightly protruding beyond anterior pronotum, postclypeus centrally sulcate, rounded or transverse anteriorly, rostrum reaching from between middle and hind coxae to sternite III. Pronotum shorter than mesonotum, trapezoidal with anterior margin narrower than lateral margins of pronotal collar, metanotum partially visible laterally by wing groove, cruciform elevation smoothly arched on posterior. Fore wings and hind wings generally hyaline, with eight and six apical cells, respectively, radial and radiomedial crossveins generally parallel. Basal areas of wings may be infuscated, wing apices are bronzed in G. atalapae , G. distanti n. comb., and G. maxineae and ambient vein, radial crossvein and radiomedial crossvein infuscated in G. bonaerensis and G. flavipronotum n. comb. Fore femora with primary and secondary spines at various angles to one another, and a small tertiary spine, tarsi three-segmented. Male operculum generally forms an approximate right triangle covering the tympanal cavity reaching to sternite II, extended posteriorly to sternite III as more of an equilateral triangle in G. chlorogena , opercula do not meet along midline, meracanthus tapering to a point, female operculum and meracantus generally of similar shape to male for each species but smaller. Abdominal segments generally with parallel sides to tergite 4 where the abdomen begins narrowing posteriorly to the genitalia, epipleurites reflexed to ventral surface, posterior sternites translucent in males of some species. Female sternite VII with sinuate posterior margin, may possess medial notch. Timbal cover with rounded anterior apex, timbal cover may conceal the majority of the timbal or only the lateral aspect of the timbal, timbal cover may have concave anteromedial margin producing approximate parallel dorsal and ventral margins on anterior extension above lateral timbal before reaching the rounded apex, ventral margin is parallel to long body axis in timbal covers that extend over lateral timbal cavity, timbals extend below wing bases. Pygofer distal shoulder rounded, dorsal beak absent, pygofer basal lobe indistinct, pygofer upper lobe generally large and separated from pygofer but there are some species where it is only slightly larger than the pygofer basal lobe (e.g., G. brisa n. comb.) or adpressed against the pygofer (e.g., G. maxineae ), of variable shape ranging from pointed to paddle-like, uncal lobes of variable size generally forming a horse shoe-shaped structure when viewed from the posterior, and male aedeagus ranging from a simple tubular struture, those with a bulbous distal terminus, or those with multiple spines at the terminus. Female abdominal segment 9 with dorsal beak well defined and sinuate posterior margin, ovipositor sheath extends beyond dorsal beak.

Measurements (mm). Length of body: 12.9–36.7; length of fore wing: 18.9–50.2; width of fore wing: 5.9–14.7; length of head: 2.1–4.8; width of head including eyes: 5.6–14.3; width of pronotum including suprahumeral plates: 5.3–14.1; width of mesonotum: 4.9–12.1.

Distribution. The genus is found over much of the Neotropics including references to Argentina, Brazil, Ecuador, French Guiana, Guyana, Paraguay, and Uruguay ( Metcalf 1963; Sanborn 2011a; 2011b; 2013; Sanborn & Heath 2014; Gogala et al. 2015). The new combinations for the species below extend the distribution to Bolivia, Colombia, Costa Rica, Panama, Peru, and Venezuela.

Key to species of Guyalna

1 Apical cells of the fore wings bronzed.................................................................... 2

- Apical cells of fore wings not bronzed.................................................................... 4

2 Apical cells of hind wings bronzed, basal area of wings brown and red, body length about 32 mm .............. G. atalapae

- Apical cells of the hind wings not bronzed, basal area of wings brown and black or not infuscated, body length 22–26 mm … 3

3 Basal area of wings brown and black............................................................. G. maxineae

- Basal area of wings not infuscated......................................................... G. dis tanti n. comb.

4 Ambient vein, radial and radiomedial crossveins infuscated................................................... 5

- Ambient vein, radial and radiomedial crossveins not infuscated................................................ 6

5 Body coloration green and black, pronotum green, basal area of wings orange, timbal cover beginning anterior extension at dorsal opening to timbal cavity............................................................... G. bonaerensis

- Body coloration brown and black, pronotum yellow, basal area of wings brown, timbal cover beginning anterior extension lateral to dorsal opening to timbal cavity................................................ G. flavipronotum n. comb.

6 Basal area of hind wing infuscated...................................................................... 7

- Basal area of hind wing not infuscated.................................................................... 12

7 Basal area of fore wing and hind wing brown, head and thorax brown and black, abdomen black, light circular region covering most of abdominal sternites................................................................... G. r uf a pi c a li s

- Basal area of fore wing not infuscated, body coloration not as above............................................ 8

8 Body length greater than 30 mm ......................................................................... 9

- Body length less than 30 mm .......................................................................... 10

9 Basal area of hind wings brown, pronotal collar brown, abdominal tergites dark brown........................ G. j am e s i

- Basal area of hind wings green, pronotal collar green, abdominal tergites light brown with a black anterior margin................................................................................................... G. chlorogena

10 Abdominal tergites brown without black markings forming an arch on dorsal surface, head and thorax green without black markings..................................................................................... G. bicolor

- Abdominal tergites marked with black forming an arch on dorsal surface, prothorax and mesothorax marked with black.. 11

11 Basal area of hind wing marked with orange, postclypeus transverse anteriorly, timbal cover concealing about one quarter of timbal............................................................................. G. variegata n. comb.

- Basal area of hind wing marked with brown, postclypeus arched anteriorly, timbal cover concealing about half of timbal........................................................................................... G. brisa n. comb.

12 Body length greater than 19 mm ........................................................................ 13

- Body length less than 19 mm .......................................................................... 21

13 Mesothorax green or brown, black mark along parapsidal suture which may be absent, anterior mesothoracic margin may be marked with black................................................................................... 14

- Mesothorax with submedian sigillae black................................................................ 19

14 Body green (may fade to yellow) with transverse black markings on posterior abdominal tergites....... G. glauca n. comb.

- Body green or brown, black mark along parapsidal suture.................................................... 15

15 Body green marked with black.......................................................................... 16

- Body brown marked with black........................................................................ 18

16 Head and thorax greenish ochraceous, abdominal tergites black marked with transverse brown markings....... G. bleuzeni

- Head and thorax greenish, abdominal tergites brown marked with black forming an arch........................... 17

17 Ventral head green with transverse black band, lorum, gena and anteclypeus marked with black, postclypeus flattened........................................................................................... G. coffe a n. comb.

- Ventral head green, postclypeus swollen........................................................... G. jauffreti

18 Ventral head with transverse black mark across postclypeus, postclypeus bulbous, body length about 27 mm ................................................................................................. G. bogotana n. comb.

- Ventral head lacking transverse black mark, postclypeus flattened, body length about 24 mm ...... G. panamensis n. comb.

19 Body black marked with brown, body length 17–22 mm ................................................. G. nigra

- Body green or brown marked with black, body length 23–25 mm .............................................. 20

20 Body brown, abdominal tergites black with brown posterior, black pile forms an arch on dorsal tergites................................................................................................... G. viridifemur n. comb.

- Body green, abdominal tergites half black, half brown with white pubescent spots on lateral tergites 2 and 3...... G. densusa

21 Mesothorax with lateral and submedian sigillae entirely black................................................. 22

- Mesothorax with submedian sigillae black, lateral sigillae incompletely marked with black......................... 24

22 Dorsal head almost entirely black, longitudinal black fascia on midline of pronotum, lateral and paramedian fissures of pronotum without black marking, body length 16 mm ................................................. G. sublaqueata

- Dorsal head with a transverse black fascia, pronotum lacking fascia along dorsal midline, lateral and paramedian fissures of pronotum black..................................................................................... 23

23 Abdominal tergites black with brown posterior margin, lacking arching pattern on dorsal surface formed by black pile, body length 17–22 mm ................................................................................ G. nigra

- Abdominal tergites with arching pattern on dorsal surface formed by black pile, body length 12.9–13.6 mm ...... G. nadae

24 Complete transverse fascia on head, postclypeus rounded ventrally......................................... G. c ut a

- Transverse fascia on head incomplete, postclypeus flattened ventrally........................................... 25

25 Black marking only in posterior paramedian fissure, lateral sigilla marked anteriorly and small spot on posterior, basal membranes of wings grayish......................................................................... G. p ar v ul a

- Black marking through entire paramedian fissure connecting to anterior tranverse mark and expanded spot on midline of ambient fissure on pronotum, lateral sigillae marked along entire lateral surface, basal membranes of wings orange. G. platyrhina