Hemiptera: Cicadidae) and a Key to Their Species

Journal of Asia-Paciﬁc Biodiversity xxx (2016) 1e7

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Original article Description of three new genera, Paratibicen, Megatibicen, and Ameritibicen, of Cryptotympanini (Hemiptera: Cicadidae) and a key to their species

Young June Lee*

Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT, USA article info abstract

Article history: Three new genera, Paratibicen gen. nov., Megatibicen gen. nov., and Ameritibicen gen. nov., are erected for Received 18 August 2016 the former members of Neotibicen Hill & Moulds, 2015 from North America, designating Cicada similaris Received in revised form Smith & Grossbeck, 1907, Cicada auletes Germar, 1834, and Cicada dorsata Say, 1825 as their respective 4 September 2016 type species. Morphological comparisons are made among the three new genera, Lyristes Horváth, 1926 Accepted 7 September 2016 (¼ Tibicen Latreille, 1825), Auritibicen Lee, 2015, Neotibicen sensu stricto, and other related genera. A key to Available online xxx the species of the genera Neotibicen, Paratibicen, Megatibicen, and Ameritibicen is provided. Copyright Ó 2016, National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA). Keywords: Auritibicen Production and hosting by Elsevier. This is an open access article under the CC BY-NC-ND license (http:// Hadoa creativecommons.org/licenses/by-nc-nd/4.0/). Lyristes Neotibicen new combinations North America Tibicen

Introduction species: H. biﬁda (Davis, 1916), H. chihuahuaensis (Sanborn, 2007), H. chiricahua (Davis, 1923), H. chisosensis (Davis, 1934), H. distanti By the erections of Auritibicen Lee, 2015 by Lee (2015) and (Metcalf, 1963), H. duryi (Davis, 1917), H. fusca (Davis, 1934), Neotibicen Hill & Moulds, 2015 and Hadoa Moulds, 2015 in Hill et al H. hidalgoensis (Davis, 1941), H. inaudita (Davis, 1917), (2015), and by the generic reassignments by Sanborn (2015), there H. longiopercula (Davis, 1926), H. minor (Davis, 1934), H. montezuma have been relocations of many species of the former Lyristes Hor- (Distant, 1881), H. neomexicensis (Stucky, 2013), H. paralleloides váth, 1926 (type species: Cicada plebeja Scopoli, 1763 from (Davis, 1934), H. parallela (Davis, 1923), H. robusta (Distant, 1881), Europe; ¼ Tibicen Latreille, 1825 nomen nudum) into different H. simplex (Davis, 1941), H. sugdeni (Davis, 1941), H. texana (Metcalf, genera. After synonymization of Lyristes altaiensis Schmidt, 1932 1963), and H. townsendii (Uhler, 1905). with Neotibicen resh (Haldeman, 1852) by Wang et al (2015) (also However, it was found through an alpha taxonomic study that see Lee 2015 and Sanborn 2015), the genus Lyristes is now there still should exist multiple groups or genera within the current comprised of only ﬁve species from sub-Mediterranean and Med- concept of the genus Neotibicen. iterranean areas in the Western Palaearctic Region (in Europe). Among the species of the genus Neotibicen, the following 11 Most of the former “Tibicen” species from North America are species form a monophyletic group: N. auriferus (Say, 1825), now placed in the two genera Neotibicen and Hadoa. Neotibicen N. canicularis (Harris, 1841), N. davisi (Smith & Grossbeck, 1907), includes 21 species, excluding an extinct species [Neotibicen ber- N. latifasciatus (Davis, 1915), N. linnei (Smith & Grossbeck, 1907), mudianus (Verrill,1902)] and an invalid name [Neotibicen variegatus N. lyricen (Degeer, 1773), N. pruinosus (Say, 1825), N. robinsonianus (Fabricius, 1794) nomen nudum]. Hadoa includes the following 20 (Davis, 1922), N. superbus (Fitch, 1855), N. tibicen (Linnaeus, 1758), and N. winnemanna (Davis, 1912). These 11 species remain in Neotibicen. Neotibicen similaris (Smith & Grossbeck, 1907) has its own * Corresponding author. peculiar morphology as discussed later and thus should be placed E-mail address: [email protected]. Peer review under responsibility of National Science Museum of Korea (NSMK) and in a separate monotypic genus. A new genus, Paratibicen gen. nov., Korea National Arboretum (KNA). is described for this species. http://dx.doi.org/10.1016/j.japb.2016.09.002 pISSN2287-884X eISSN2287-9544/Copyright Ó 2016, National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA). Production and hosting by Elsevier. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Please cite this article in press as: Lee YJ, Description of three new genera, Paratibicen, Megatibicen, and Ameritibicen, of Cryptotympanini (Hemiptera: Cicadidae) and a key to their species, Journal of Asia-Paciﬁc Biodiversity (2016), http://dx.doi.org/10.1016/j.japb.2016.09.002 2 YJ Lee / Journal of Asia-Paciﬁc Biodiversity xxx (2016) 1e7

The following three species form another monophyletic group: Hadoa townsendii (Uhler, 1905): 1_, Sierra County, NM, USA, 15 VI N. auletes (Germar, 1834), N. resh (Haldeman, 1852), and N. resonans 2012 (UCONN). (Walker, 1850). A new genus, Megatibicen gen. nov., is described for Cacama valvata (Uhler, 1888): 1_, Bent County, CO, USA, 4 VII these three species. 2004 (UCONN). The following six species form another monophyletic group: Photographs and illustrations provided by Cole (2008), Davis N. cultriformis (Davis, 1915), N. dealbatus (Davis, 1915), N. dorsatus (1915, 1916, 1917, 1923, 1926, 1927, 1934, 1938, 1941), Lawson (Say, 1825), N. ﬁguratus (Walker, 1858), N. pronotalis (Davis, 1938), (1920), Stucky (2013), and others were also referenced. and N. tremulus (Cole, 2008). Another new genus, Ameritibicen gen. The digital images of the male genitalia were captured using a nov., is described for these six species. Leica stereo-zoom microscope attached to a computer-assisted im- A key to the males of all the 21 species of Neotibicen, Paratibicen, aging system, Auto-Montage by Syncroscopy (Cambridge, UK), at Megatibicen, and Ameritibicen is provided in subsequent sections. the collection facility of the University of Connecticut. Terminology for morphological features generally follows that of Moulds (2005).

Materials and methods Genus Neotibicen Hill & Moulds, 2015 Many specimens of the species of the new genera and their Type species. Cicada canicularis Harris, 1841 (type locality: Mas- closely related genera including Lyristes, Auritibicen, Neotibicen sachusetts; Figure 1). sensu stricto, Hadoa Moulds, 2015, and Cacama Distant, 1904 were Included species. N. auriferus (Say, 1825), N. canicularis (Harris, examined for this study, which are specimens from Kathy B.R. Hill 1841), N. davisi (Smith & Grossbeck, 1907) [with the subspecies of and David C. Marshall’s cicada collection at the collection facility of N. davisi davisi (Smith & Grossbeck, 1907) and N. davisi hardeni the University of Connecticut, Storrs, CT, USA (UCONN), and from (Davis, 1918)], N. latifasciatus (Davis, 1915), N. linnei (Smith & the present author’s private collection (YJL), including the speci- Grossbeck, 1907), N. lyricen (Degeer, 1773) [with the subspecies of mens listed below. N. lyricen engelhardti (Davis, 1910), N. lyricen lyricen (Degeer, 1773), Neotibicen auriferus (Say, 1825): 1_, Meade County, KS, USA, 05 and N. lyricen virescens (Davis, 1935)], N. pruinosus (Say, 1825) [with IX 2007 (UCONN). Neotibicen canicularis (Harris, 1841): 1_, Man- the subspecies of N. pruinosus fulvus (Beamer, 1924) and chester, CT, USA, 10 VIII 2008 (YJL); 1_, Mansfield, CT, USA, 07 VIII N. pruinosus pruinosus (Say, 1825)], N. robinsonianus (Davis, 1922), 2008 (YJL); 1_, Storrs, CT, USA, 19 VIII 2008 (YJL). Neotibicen davisi N. superbus (Fitch, 1855), N. tibicen (Linnaeus, 1758) [with the (Smith & Grossbeck, 1907): 1_, Citrus County, FL, USA, 22 IX 2011 subspecies of N. tibicen australis (Davis, 1912) and N. tibicen tibicen (UCONN). Neotibicen latifasciatus (Davis, 1915): 1_, Cape May (Linnaeus, 1758)], and N. winnemanna (Davis, 1912). County, NJ, USA, 06 VIII 2010 (UCONN). Neotibicen linnei (Smith & Diagnosis. Head including eyes wider than base of mesonotum. Grossbeck, 1907): 1_, Kent County, DE, USA, 28 VIII 2004 (UCONN). Pronotum with lateral margin weakly ampliated, not dentate. Male Neotibicen lyricen (Degeer, 1773): 1_, Storrs, CT, USA, 30 VII 2008 opercula about as long as wide or longer than wide, overlapped at (YJL); 1_, Storrs, CT, USA, 6 VIII 2009 (YJL). Neotibicen lyricen inner margins. Wings hyaline. Forewing comparatively long and engelhardti (Davis, 1910): 1_, Albemarle County, VA, USA, 17 VIII slender (ratio of forewing length to width 2.65e3.20). Forewing 2003 (UCONN). Neotibicen pruinosus (Say, 1825): 1_, Barber County, radial cell slightly slenderer than ulnar cell 3. Male abdomen about KS, USA, 5 IX 2007 (UCONN). Neotibicen robinsonianus (Davis, as long as or slightly longer than head and thorax together. Male 1922): 1_, Culpeper County, VA, USA, 17 VIII 2003 (UCONN). Neo- abdominal sternite VIII widely rounded or V-shaped in posterior tibicen superbus (Fitch, 1855): 1_, Freestone County, TX, USA, 23 VI view. Timbal cover completely covering timbal. Pygofer barrel- 2002 (UCONN). Neotibicen tibicen (Linnaeus, 1758): 1_, Bergen shaped (with a little widened distal part) in ventral view. Dorso- County, NJ, USA, 15 VIII 2015 (YJL); 1_, Culpeper County, VA, USA, 17 distal part (between distal shoulders, including dorsal beak and VIII 2003 (UCONN). inward curvatures on both sides of dorsal beak) of pygofer lower Paratibicen similaris (Smith & Grossbeck, 1907) comb. nov.: 1_, than distal shoulders but with long and acute dorsal beak, forming Alachua County, FL, USA, 1 IX 2008 (UCONN). a trident-shape in dorsal view, with middle branch longest Megatibicen auletes (Germar, 1834) comb. nov.: 1_, Warren (Figure 1E). Uncal lobes fused into one, not bifurcate; long, at least County, GA, USA, 16 IX 2009 (UCONN). Megatibicen resh (Haldeman, one-third the length of pygofer; simple, without projections (see 1852) comb. nov.: 1_,Leflore County, MS, USA, 15 VIII 2010 Diagnosis of Paratibicen for comparison); widely and convexly (UCONN). Megatibicen resonans (Walker, 1850) comb. nov.: 1_, Leon rounded or truncate at apical part in ventral view; curved County, FL, USA, 21 VII 2008 (UCONN). inwardly in lateral view. Aedeagus thick and with widely truncate Ameritibicen cultriformis (Davis, 1915) comb. nov.: 1_, Santa Cruz apex. Basal lobe of pygofer paramedian, apart from lateral wall of County, AZ, USA, 02 IX 2007 (UCONN). Ameritibicen dealbatus pygofer, not curved laterad at apices, and basally connected to (Davis, 1915) comb. nov.: 1_, KS, USA, 05 VII 2004 (UCONN). each other. Ameritibicen dorsatus (Say, 1825) comb. nov.: 1_, Montgomery County, KS, USA, 6 IX 2007 (UCONN). Ameritibicen figuratus (Walker, 1858) comb. nov.: 1_, Chambers County, AL, USA, 25 VIII 2005 Genus Paratibicen gen. nov. (UCONN). Ameritibicen pronotalis pronotalis (Davis, 1938) comb. nov.: 1_, IA, USA, 6 IX 2009 (UCONN). Ameritibicen pronotalis wal- LSID. urn:lsid:zoobank.org:act:78D46A19-401E-42CB-A761- keri (Metcalf, 1955) comb. nov.: 1_, Montgomery, KS, USA, 06 IX AA500E46EBDC 2007 (UCONN). Ameritibicen tremulus (Cole, 2008) comb. nov.: 1_, Type species. Cicada similaris Smith & Grossbeck, 1907 (type lo- Comanche County, KS, USA, 5 IX 2007 (UCONN). cality: Fernandina, FL; Figure 2). Hadoa chiricahua (Davis, 1923): 1_, Cochise County, AZ, USA, 25 Included species. P. similaris (Smith & Grossbeck, 1907) comb. VI 2007 (UCONN). Hadoa duryi (Davis, 1917): 1_, Gila County, AZ, nov. USA, 29 VI, 2003 (UCONN). Hadoa inaudita (Davis, 1917): 1_, Otero Etymology. The generic name refers to the morphological simi- County, NM, USA, 28 VI 2007 (UCONN). Hadoa neomexicensis larity of this genus to Tibicen, adding the Greek prefix para-to (Stucky, 2013): 1_, Otero County, NM, USA, 28 VI 2007 (UCONN). Tibicen. Although Tibicen is treated here as nomen nudum, the name Hadoa texana (Metcalf, 1963): 1_, TX, USA, 18 VI 2012 (UCONN). Tibicen has been mainly used for the American species.

Please cite this article in press as: Lee YJ, Description of three new genera, Paratibicen, Megatibicen, and Ameritibicen, of Cryptotympanini (Hemiptera: Cicadidae) and a key to their species, Journal of Asia-Paciﬁc Biodiversity (2016), http://dx.doi.org/10.1016/j.japb.2016.09.002 YJ Lee / Journal of Asia-Paciﬁc Biodiversity xxx (2016) 1e7 3

Figure 1. Neotibicen canicularis (Harris, 1841), male, Connecticut, USA: A, Dorsal habitus; B, Ventral habitus; C, Ventral view of the pygofer; D, Lateral view of the pygofer; E, Dorsal beak.

Diagnosis. Head including eyes wider than base of mesonotum. Included species. M. auletes (Germar, 1834) comb. nov., M. resh Pronotum with lateral margin very weakly ampliated, not dentate. (Haldeman, 1852) comb. nov., and M. resonans (Walker, 1850) comb. Male opercula longer than wide, overlapped at inner margins. Wings nov. hyaline. Forewing comparatively long and slender (ratio of forewing Etymology. The generic name is a Latin masculine noun meaning length to width about 2.9). Forewing radial cell slenderer than ulnar “large Tibicen”, having the preﬁxofmega-. Although Tibicen is cell 3. Male abdomen slightly longer than head and thorax together. treated here as nomen nudum, the name Tibicen has been mainly Male abdominal sternite VIII narrowly V-shaped in posterior view. used for the American species. Timbal cover completely covering timbal. Pygofer oblong in ventral Diagnosis. Head including eyes wider than base of mesonotum. view. Dorsodistal part of pygofer not lower than distal shoulders, Pronotum with lateral margin weakly ampliated, not dentate. Male forming a triangle in dorsal view with long and acute dorsal beak opercula about as long as wide, overlapped at inner margins. Wings (Figure 2E). Uncal lobes fused into one, not bifurcate; comparatively hyaline. Forewing comparatively long and slender (ratio of forewing short, less than one-third the length of pygofer; with a pair of sharp length to width 2.90e3.20). Forewing radial cell slenderer than ulnar projections below the uncus (Figure 2D); tapering into narrow and cell 3. Male abdomen about as long as head and thorax together. Male triangularly rounded apex in ventral view; curved inwardly in lateral abdominal sternite VIII rounded in posterior view. Timbal cover view. Aedeagus thick and with widely truncate apex. Basal lobe of completely covering timbal. Pygofer inverted curvilinear triangle in pygofer lateral, attached to lateral wall of pygofer [contradictory to ventral view, with distal part broadened, in ventral view. Dorsodistal what was said in Lee (2015) in regards to the basal lobes of the part of pygofer lower than distal shoulders, with very short and dull “America Tibicen”, which becomes not contradictory when dorsal beak, not higher than distal shoulders (Figure 3E). Uncal lobes excluding this species or Paratibicen from the “America Tibicen” in fused into one, not bifurcate; comparatively short, less than one- Lee (2015)], not curved laterad at apices. third the length of pygofer; simple, without projections (see Diag- nosis of Paratibicen for comparison); widely concave at apical part in ventral view; lightly curved inwardly in lateral view. Aedeagus thick Genus Megatibicen gen. nov. but tapering into thinner but truncate apex, usually hidden under uncus. Basal lobe of pygofer sublateral or paramedian, apart from LSID. urn:lsid:zoobank.org:act:647F7490-7746-4688-A428- lateral wall of pygofer, curved laterad at apices. D40CC77F1358 Type species. Cicada auletes Germar, 1834 (type locality: Penn- sylvania; Figure 3), by the present designation. Genus Ameritibicen gen. nov. The species is designated as the type species for its typical morphological characteristics, its wide distribution, and its LSID urn:lsid:zoobank.org:act:36C2BE3C-9388-4E60-93D3- abundances. 31A815ACF6EE

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Figure 2. Paratibicen similaris (Smith & Grossbeck, 1907) comb. nov., male, Florida, USA: A, Dorsal habitus; B, Ventral habitus; C, Ventral view of the pygofer; D, Lateral view of the pygofer; E, Dorsal beak.

Type species. Cicada dorsata Say, 1825 (type locality: Atchison inwardly in lateral view. Aedeagus comparatively thick but County, KS; Figure 4), by the present designation. tapering into thin but truncate apex, thinner than Megatibicen, The species is designated as the type species for its typical with apex usually disposed out of apex of uncus. Basal lobe of morphological characteristics, its moderately wide distribution, pygofer paramedian, apart from lateral wall of pygofer, not curved and its abundances. laterad at apices. Included species. A. cultriformis (Davis, 1915) comb. nov., A. dealbatus (Davis, 1915) comb. nov., A. dorsatus (Say, 1825) comb. Morphological comparisons among related genera nov., A. ﬁguratus (Walker, 1858) comb. nov., A. pronotalis (Davis, 1938) comb. nov. [with the subspecies of A. pronotalis pronotalis The former members of Neotibicen are distinguished from Lyr- (Davis, 1938) comb. nov. and A. pronotalis walkeri (Metcalf, 1955) istes sensu stricto (from the Western Palaearctic area) by the comb. nov.], and A. tremulus (Cole, 2008) comb. nov. following characters: pygofer thin and long in lateral view (plump Etymology. The generic name is a Latin masculine noun meaning and broad in lateral view in Lyristes); dorsodistal part of pygofer, “ ” ﬁ American Tibicen , having the pre xofameri-, derived from excluding dorsal beak, lower than distal shoulders (dorsodistal part “ ” America . Although Tibicen is treated here as nomen nudum, the of pygofer triangular in Lyristes); uncus large, about one-third the name Tibicen has been mainly used for the American species. length of pygofer or more (comparatively small in Lyristes); and Diagnosis. Head including eyes wider than base of mesonotum. uncus simple, without aedeagus-supporting structures (with a pair Pronotum with lateral margin weakly ampliated, not dentate. Male of cap-shaped structures protruding below uncal lobe, supporting opercula about as long as wide, overlapped at inner margins. aedeagus, in Lyristes). Wings hyaline. Forewing comparatively short and broad in some The former members of Neotibicen are distinguished from species but long and slender in other species (ratio of forewing Auritibicen (from eastern Asia) by the following characters: fore- e length to width about 2.60 3.20). Forewing radial cell slightly wing costa slightly convex before node, making radial cell slightly slenderer than ulnar cell 3. Male abdomen about as long as or slenderer than ulnar cell 3 (roundly convex before node, making slightly longer than head and thorax together. Male abdominal radial cell broad enough to be slightly wider or about as broad as sternite VIII V-shaped in posterior view. Timbal cover completely ulnar cell 3 in Auritibicen); and basal lobe of pygofer sublateral or covering timbal. Pygofer oblong in ventral view. Dorsodistal part of paramedian, apart from lateral wall of pygofer (attached to lateral pygofer lower than distal shoulders without dorsal beak wall of pygofer in Auritibicen). (Figure 4E). Uncal lobes fused into one, not bifurcate; compara- Hadoa is distinguished from the former members of Neotibicen tively short, less than one-third the length of pygofer; simple, by the narrower head (with exceptions), narrowly V-shaped male without projections (see Diagnosis of Paratibicen for comparison); abdominal sternite VIII (when excluding Paratibicen from the former tapering into narrow and rounded apex in ventral view; curved members of Neotibicen), and differently positioned basal lobes of

Please cite this article in press as: Lee YJ, Description of three new genera, Paratibicen, Megatibicen, and Ameritibicen, of Cryptotympanini (Hemiptera: Cicadidae) and a key to their species, Journal of Asia-Paciﬁc Biodiversity (2016), http://dx.doi.org/10.1016/j.japb.2016.09.002 YJ Lee / Journal of Asia-Paciﬁc Biodiversity xxx (2016) 1e7 5

Figure 3. Megatibicen auletes (Germar, 1834) comb. nov., male, Georgia, USA: A, Dorsal habitus; B, Ventral habitus; C, Ventral view of the pygofer; D, Lateral view of the pygofer; E, Dorsal beak.

pygofer (with some exceptions). The complexity or multigrouping of acute dorsal beak (Figure 1E) [without dorsal beak (Figure 4E) in Hadoa will be discussed separately in the forthcoming study. Ameritibicen]; uncus long, at least one-third the length of pygofer Paratibicen is distinguished from all the former members of (less than one-third the length of pygofer in Ameritibicen); uncus Neotibicen by the following characters: male abdominal sternite VIII widely and convexly rounded or truncate at apical part (tapering narrowly V-shaped in posterior view; uncus with a pair of sharp into narrow apex in Ameritibicen) in ventral view; aedeagus with projections below uncus; dorsodistal part of pygofer not lower than widely truncate apex (tapering into thin apex in Ameritibicen). distal shoulders, forming a triangle in dorsal view with long and Megatibicen gen. nov. is distinguished from Ameritibicen gen. acute dorsal beak (Figure 2E); basal lobe of pygofer lateral, attached nov. by the following characters: male abdominal sternite VIII to lateral wall of pygofer. Paratibicen is thought to be close to Hadoa rounded (V-shaped in Ameritibicen) in posterior view; pygofer when considering the similarities in the shape of the male inverted curvilinear triangle in ventral view, with distal part abdominal sternite VIII, the position and shape of the basal lobes of broadened, in ventral view (oblong in ventral view in Ameritibicen); pygofer, the shape of the dorsodistal part of the pygofer, and the dorsodistal part of pygofer with short and dull but distinct dorsal shape of the uncus in ventral view. beak (Figure 3E) [without distinct dorsal beak (Figure 4E) in Neotibicen is distinguished from Megatibicen gen. nov. by the Ameritibicen]; uncus widely concave at apical part (tapering into following characters: pygofer barrel-shaped in ventral view narrow apex in Ameritibicen) in ventral view; basal lobes curved (inverted curvilinear triangle in ventral view, with distal part laterad at apices (not so in Ameritibicen). broadened, in Megatibicen); dorsodistal part of pygofer with long and acute dorsal beak (Figure 1E) [with very short and dull dorsal beak, not higher than distal shoulders (Figure 3E) in Megatibicen]; Key to the males of the species of Neotibicen, Paratibicen, uncus long, at least one-third the length of pygofer (less than one- Megatibicen, and Ameritibicen third the length of pygofer in Megatibicen); uncus widely and convexly rounded or truncate (widely concave in Megatibicen)at 1. Dorsodistal part of pygofer with long and acute dorsal beak apical part in ventral view; aedeagus with widely truncate apex (Figures 1E and 2E); aedeagus with widely truncate apex . 2 (tapering into thin apex in Megatibicen). - Dorsodistal part of pygofer with or without very short and Neotibicen is distinguished from Ameritibicen gen. nov. by the dull dorsal beak, not higher than distal shoulders, if any following characters: dorsodistal part of pygofer with long and (Figures 3E and 4E); aedeagus tapering into thin apex ... 13

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Figure 4. Ameritibicen dorsatus (Say, 1825) comb. nov., male, Kansas, USA: A, Dorsal habitus; B, Ventral habitus; C, Ventral view of the pygofer; D, Lateral view of the pygofer; E, Dorsal beak.

2. Uncus simple, without projections (Figure 1D); basal lobe of tergite 2 and mostly exposed; white pollinosity on inner pygofer paramedian, apart from lateral wall of pygofer corner of timbal covers distinct and prominent ..... 6 (Figure 1C); dorsodistal part of pygofer lower than distal 6. Transverse black spots or fascia between posterior margin of shoulders (Figure 1E) ...3(Neotibicen Hill & Moulds, 2015) pronotal inner area and anterior margin of pronotal collar - Uncus with a pair of sharp projections below uncus short or indistinct, not extending to sublateral margin of (Figure 2D); basal lobe of pygofer lateral, attached to lateral pronotal collar; forewing comparatively broad (ratio of wall of pygofer (Figure 2C); dorsodistal part of pygofer not forewing length to width about 2.75e2.90) ...... 7 lower than distal shoulders (Figure 2E)...... - Transverse black fascia between posterior margin of pronotal ...... (Paratibicen gen. inner area and anterior margin of pronotal collar distinctly nov.) Paratibicen similaris (Smith & Grossbeck, 1907) comb. extending to sublateral margin of pronotal collar; forewing nov. comparatively slender (ratio of forewing length to width 3. Abdominal sternite VIII widely rounded in posterior view about 3.00) .....Neotibicen robinsonianus (Davis, 1922) ...... 4 7. Operculum generally symmetric ...... 8 - Abdominal sternite VIII V-shaped in posterior view ...10 - Operculum quite asymmetric, with apex biased laterad 4. Uncus not widened toward apex but with nearly straight ...... Neotibicen linnei (Smith & Grossbeck, 1907) lateral side up to apex in ventral view; pronotal collar mostly 8. Abdominal tergite 3 with or without a pair of short trans- light-colored (dull green or ochraceous) ...... 5 verse fasciae of white pollinosity on latero-anterior margin of - Uncus slightly widened toward apex, with slightly convex tergite 3...... 9 lateral side at apex in ventral view; pronotal collar mostly - Abdominal tergite 3 with a pair of transverse thick and long dark-colored (black to fuscous) ...... fasciae of white pollinosity along anterior margin of tergite 3, ...... Neotibicen lyricen (Degeer, 1773) discontinued only on median part ...... 5. Body comparatively small (about 27 mm long); abdominal ...... Neotbicen latifasciatus (Davis, 1915) tergite 1 very short, about one-quarter the median length of 9. Abdominal tergite 3 with a pair of small but distinct fasciae of tergite 2 and largely hidden under cruciform elevation; white pollinosity on latero-anterior margin of tergite 3 .. white pollinosity on inner corner of timbal covers very ...... Neotibicen pruinosus (Say, 1825) minimal or absent ... Neotibicen canicularis (Harris, 1841) - Abdominal tergite 3 with or without indistinct white polli- - Body comparatively large (longer than 31 mm); abdominal nosity on latero-anterior margin of tergite 3 ...... tergite 1 longer than one-quarter the median length of ...... Neotibicen winnemanna (Davis, 1912)

Please cite this article in press as: Lee YJ, Description of three new genera, Paratibicen, Megatibicen, and Ameritibicen, of Cryptotympanini (Hemiptera: Cicadidae) and a key to their species, Journal of Asia-Paciﬁc Biodiversity (2016), http://dx.doi.org/10.1016/j.japb.2016.09.002 YJ Lee / Journal of Asia-Paciﬁc Biodiversity xxx (2016) 1e7 7

10. Operculum comparatively short, not reaching posterior 19. Head including eyes wider than abdominal tergite 2 margin of abdominal sternite III; uncus not widened toward including timbal covers; mesonotum and abdomen covered apex but with nearly straight lateral side up to apex in ventral in large areas with white pollinosity ...... view; pronotal collar mostly light-colored (dull green or ...... Ameritibicen pronotalis (Davis, 1938) comb. nov. ochraceous) ...... 11 - Head including eyes about as wide as abdominal tergite 2 - Operculum long, reaching or extending beyond posterior including timbal covers; mesonotum and abdomen covered margin of abdominal sternite IV; uncus slightly widened in much reduced areas with white pollinosity ...... toward apex, with slightly convex lateral side at apex in ...... Ameritibicen dealbatus (Davis, 1915) comb. nov. ventral view; pronotal collar mostly dark-colored (black to 20. Timbal cover mostly ochraceous; basal lobes of pygofer very fuscous) ...... Neotibicen tibicen (Linnaeus, 1758) narrowly separated from each other and with blunt apices 11. Abdomen mostly black; operculum comparatively short, not ...... Ameritibicen dorsatus (Say, 1825) comb. nov. reaching posterior margin of abdominal sternite II ... 12 - Timbal cover mostly black; basal lobes of pygofer widely - Abdomen mostly ochraceous to brown; operculum compar- separated from each other and with more or less acute apices atively long, well extending beyond posterior margin of ...... Ameritibicen tremulus (Cole, 2008) comb. nov. abdominal sternite II ... Neotibicen superbus (Fitch, 1855) 12. Uncus in similar width until subapex; rostrum comparatively Acknowledgments long, reaching anterior margin of hind coxae ...... Neotibicen davisi (Smith & Grossbeck, 1907) I am indebted to Kathy B.R. Hill and David C. Marshall (University - Uncus slightly tapering into narrower subapex; rostrum of Connecticut) for access to their cicada collection at the collection comparatively short, extending slightly beyond posterior facility of the University of Connecticut. I am grateful to David C. .... margin of mid coxae Neotibicen auriferus (Say, 1825) Marshall for reading an earlier version of the manuscript, and to 13. Pygofer inverted curvilinear triangle in ventral view Stéphane Puissant (Museum National d’Histoire Naturelle, Paris, (Figure 3C), with distal part broadened; dorsodistal part of France) and an anonymous reviewer for their valuable comments. pygofer with short and dull dorsal beak; uncus widely concave at apical part in ventral view; basal lobe of pygofer curved laterad at apices .....14(Megatibicen gen. nov.) References - Pygofer oblong in ventral view (Figure 4C); dorsodistal part Cole JA. 2008. A new cryptic species of cicada resembling Tibicen dorsatus revealed of pygofer without dorsal beak; uncus tapering into narrow by calling song (Hemiptera: Auchenorrhyncha: Cicadidae). Annals of the Ento- apex in ventral view; basal lobe of pygofer not curved laterad mological Society of America 101:815e823. at apices ...... 16 (Ameritibicen gen. nov.) Davis WT. 1915. Notes on some cicadas from the eastern and central United States with a description of a new variety of Cicada pruinosa. Journal of the New York 14. Brownish marks on mesonotum not prominent on black Entomological Society 23:1e10. ground color; forewing basal membrane and hindwing Davis WT. 1916. Notes on cicadas from the United States with descriptions of several jugum grayish ...... 15 new species. Journal of the New York Entomological Society 24:42e65. Davis WT. 1917. Sonoran cicadas collected by Harry H. Knight, Dr. Joseph Bequaert - Greenish W-shaped mark and other marks on mesonotum and others, with descriptions of new species. Journal of the New York Entomo- well contrasted with black ground color; forewing basal logical Society 25:203e215. membrane and hindwing jugum orange-colored Davis WT. 1923. Notes on North American cicadas with descriptions of new species. e ...... Megatibicen resh (Haldeman, 1852) comb. nov. Journal of the New York Entomological Society 31:1 15. Davis WT. 1926. New cicadas from California and Arizona with notes on several other 15. Pronotal collar olive-colored; uncus with apex roundly or species. Journal of the New York Entomological Society 34:177e197. triangularly concave in ventral view ...... Davis WT. 1927. New cicadas from the western United States with notes on several e ...... Megatibicen auletes (Germar, 1834) comb. nov. other species. Journal of the New York Entomological Society 35:373 389. Davis WT. 1934. New cicadas from North America. Journal of the New York Ento- - Pronotal collar rust-colored or reddish brown; uncus with mological Society 42:37e62. apex medially even (in straight line) in ventral view Davis WT. 1938. New North American cicadas with notes on described species...... Megatibicen resonans (Walker, 1850) comb. nov. Journal of the New York Entomological Society 46:291e310. Davis WT. 1941. New cicadas from North America with notes. Journal of the New York 16. Forewing very slender (ratio of forewing length to width Entomological Society 49:85e96. >3.20); forewing basal cell dark-colored; mesonotal marks Hill KBR, Marshall DC, Moulds MS, et al. 2015. Molecular phylogenetics, diversifica- around submedian sigilla delicate ...... tion, and systematics of Tibicen Latreille 1825 and allied cicadas of the tribe .... . fi Cryptotympanini, with three new genera and emphasis on species from the USA ... Ameritibicen guratus (Walker, 1858) comb. nov. and Canada (Hemiptera: Auchenorrhyncha: Cicadidae). Zootaxa 3985:219e251. - Forewing comparatively broad (ratio of forewing length to Lawson PB. 1920. The Cicadidae of Kansas. Kansas University Science Bulletin 12: width <3.00); forewing basal cell tinged with green or 309e376. Lee YJ. 2015. Description of a new genus, Auritibicen gen. nov., of Cryptotympanini ochraceous; mesonotal marks around submedian sigilla (Hemiptera: Cicadidae) with redescriptions of Auritibicen pekinensis (Haupt, broad ...... 17 1924) comb. nov. and Auritibicen slocumi (Chen, 1943) comb. nov. from China 17. Basal lobe of uncus short, with its apex far from base of uncus and a key to the species of Auritibicen. Zootaxa 3980:241e254. fi ...... 18 Moulds MS. 2005. An appraisal of the higher classi cation of cicadas (Hemiptera: Cicadoidea) with special reference to the Australian fauna. Records of the - Basal lobe of uncus very long, almost reaching base of uncus Australian Museum 57:375e446. . Ameritibicen cultriformis (Davis, 1915) comb. ....nov. Sanborn AF. 2015. New combinations for six species belonging to Cryptotympanini 18. Head including eyes about as wide as or wider than Handlirsch (Hemiptera: Cicadidae), former members of the genus Tibicen Latreille, 1825. Zootaxa 4027:447e450. abdominal tergite 2 including timbal covers; forewing basal Stucky BJ. 2013. Morphology, bioacoustics, and ecology of Tibicen neomexicensis sp. cell not surrounded by fuscous fasciae ...... 19 n., a new species of cicada from the Sacramento Mountains in New Mexico, - Head including eyes distinctly narrower than abdominal U.S.A. (Hemiptera, Cicadidae, Tibicen). ZooKeys 337:49e71. Wang X, Hayashi M, Wei C. 2015. Taxonomic note on two Lyristes cicadas (Hemi- tergite 2 including timbal covers; forewing basal cell sur- ptera: Cicadidae) described by E. Schmidt from China. Entomotaxonomia 37: rounded by fuscous fascia anteriorly ...... 20 197e200.