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A revision of the endemic Australianspittlebug genus Anyllis Kirkaldy(Hemiptera: Aphrophoridae) withdescriptions of two new speciesAi-Ping Liang a & Rong-Rong Wang aa Key Laboratory of Zoological Systematics and Evolution, Instituteof Zoology, Chinese Academy of Sciences, 1 Beichen West Road,Chaoyang District, Beijing, 100101, China

Available online: 07 Mar 2012

To cite this article: Ai-Ping Liang & Rong-Rong Wang (2012): A revision of the endemic Australianspittlebug genus Anyllis Kirkaldy (Hemiptera: Aphrophoridae) with descriptions of two new species,Journal of Natural History, 46:15-16, 1005-1023

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Journal of Natural HistoryVol. 46, Nos. 15–16, April 2012, 1005–1023

A revision of the endemic Australian spittlebug genus Anyllis Kirkaldy(Hemiptera: Aphrophoridae) with descriptions of two new species

Ai-Ping Liang* and Rong-Rong Wang

Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academyof Sciences, 1 Beichen West Road, Chaoyang District, Beijing 100101, China

(Received 18 May 2011; final version received 17 December 2011; printed 3 March 2012)

The endemic Australian spittlebug genus Anyllis Kirkaldy (Hemiptera:Auchenorrhyncha: Cercopoidea: Aphrophoridae) is revised and five speciesincluding two new species are recognized: Anyllis gibbosus sp. nov. (Queensland),Anyllis leiala Kirkaldy, 1906 (type species, throughout the eastern part of Australiafrom north Queensland to Tasmania), Anyllis pseudotiegsi sp. nov. (Tasmania),Anyllis spinostylus Liang, 2005 (Tasmania) and Anyllis tiegsi (China, 1952)(Victoria and the higher parts of southeast New South Wales). Descriptions orredescriptions of the genus and its included species are provided together with thedorsal habitus of the adults and the structural illustrations of the head, pronotum,scutellum and male genitalia. A diagnostic key is given for differentiation of theknown species.

Keywords: Auchenorrhyncha; Cercopoidea; taxonomy; new species; Australia

Introduction

The Australian Cercopoidea (Hemiptera: Auchenorrhyncha), or spittlebugs, comprisesa small but distinctive component of the Auchenorrhyncha fauna of the isolated islandcontinent (Liang and Fletcher, 2003). Evans (1966) reviewed the Australian fauna andrecognized 16 species in 10 genera. Liang and Fletcher (2003) established two newgeneric synonyms, two new specific synonyms and five new specific combinations forthe Australian Aphrophoridae and recognized 13 valid species in seven genera. Thisnumber of described species probably represents only a small fraction of the actualdiversity of the whole Australian aphrophorid fauna considering its vast territory andvarious complex habitats (Liang and Fletcher 2003). Much alpha taxonomic work,including field surveys in this group, remains to be done in Australia.

This paper on the genus Anyllis Kirkaldy (Hemiptera: Auchenorrhyncha:Cercopoidea: Aphrophoridae) represents another in a continuing series of worksemphasizing the taxonomy of the spittlebug fauna of Australia (Liang and Fletcher2002, 2003; Liang et al. 2005; Liang et al. 2012).

The genus Anyllis was erected by Kirkaldy in 1906 for Anyllis leiala Kirkaldy 1906from Australia. Lallemand (1912) redescribed the genus and designated A. leiala as thetype species. Metcalf (1961) wrongly placed Anyllis in the tribe Locrisini Lallemand ofthe subfamily Cercopinae of the family Cercopidae. Evans (1966) placed Anyllis in

*Corresponding author. Email: liangap@ioz.ac.cn

ISSN 0022-2933 print/ISSN 1464-5262 online© 2012 Taylor & Francishttp://dx.doi.org/10.1080/00222933.2011.651646http://www.tandfonline.com

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Aphrophoridae and synonymised the monotypic genus Aphrophorinella Lallemand,1946 with Anyllis. Liang and Fletcher (2003) placed Anyllis in the Aphrophoridae andsynonymized Neoaphrophora China 1952 with Anyllis, but gave no tribal placement.Liang et al. (2005) redefined the genus and placed it in the tribe Aphrophorini. Amongseven aphrophorid genera known in Australia, Anyllis is the only member of the rel-atively derived tribe Aphrophorini (Liang and Fletcher 2003). Before this study, threespecies, namely Anyllis leiala Kirkaldy, 1906, Anyllis tiegsi (China, 1952) and Anyllisspinostylus Liang, 2005, were previously recorded in the genus. The genus appears tobe endemic to Australia.

While sorting and identifying Aphrophoridae from material from the AustralianNational Insect Collection (ANIC), we found two new species of Anyllis collectedfrom Queensland and Tasmania, respectively. The purpose of the present paper is toredescribe or describe the genus Anyllis and its included species currently known fromAustralia and to provide a key to the known species and describe the dorsal habitus ofthe adults, with structural illustrations of the head, pronotum, scutellum and the malegenitalia and distributional data for their identification.

Materials and methods

Dry pinned specimens were used for the description and illustration. External mor-phology was observed under a stereoscopic microscope and characters were measuredwith an ocular micrometer. Abdomens were removed and macerated in cold 10%potassium hydroxide overnight. Precise dissections and cleaning of genital structureswere finished in distilled water. Observations and drawings were made in glycer-ine under a compound light microscope. Photographs of the types were taken witha Nikon Coolpix 5400 digital camera. The digital images were then imported intoADOBE PHOTOSHOP 8.0 for labelling and plate composition. Line figures were drawnwith the aid of a camera lucida mounted on a Zeiss Stemi SV-11 stereomicroscope.

Specimens examined during the course of this study are deposited in the AmericanMuseum of Natural History, New York, USA (AMNH); the Australian NationalInsect Collection (ANIC); NSW Agricultural Scientific Collections Unit, Orange,NSW, Australia (ASCU); The Natural History Museum, London, UK (BMNH); andthe Bernice P. Bishop Museum, Honolulu, HI, USA (BPBM).

Morphological terminology follows that of Liang et al. (2005).

Checklist of species of Anyllis Kirkaldy

Anyllis Kirkaldy, 1906Neoaphrophora China, 1952

gibbosus sp. nov.; Australia (Queensland).leiala Kirkaldy, 1906; Australia (throughout the eastern part of Australia from

North Queensland to Tasmania).tonnoiri Lallemand 1946

pseudotiegsi sp. nov.; Australia (Tasmania).spinostylus Liang, 2005; Australia (Tasmania).tiegsi (China, 1952); Australia (Victoria and the higher parts of SE New South

Wales).

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Taxonomy

Genus Anyllis Kirkaldy

Anyllis Kirkaldy, 1906: 386; Lallemand, 1912: 74, 83; Metcalf, 1961: 63; Evans, 1966:316, 317; Liang and Fletcher, 2003: 86; Liang et al., 2005: 301, 302. Type species:Anyllis leiala Kirkaldy 1906, by monotypy.

Neoaphrophora China, 1952: 789; Metcalf, 1962: 431; Evans, 1966: 316, 317. Typespecies: Neoaphrophora tiegsi China 1952, by original designation. [Synonymized byLiang and Fletcher, 2003: 86.]

DescriptionSmall-sized aphrophorine species, body length (from apex of vertex to tip offorewings): male 4.5–6.9 mm, female, 4.8–9.0 mm.

General colour brown to dark brown (reddish brown in gibbosus sp. nov.).Vertex, pronotum and clavus of forewings usually much paler, giving pale strami-neous appearance; dorsum covered with fuscous punctures. Frons, thorax beneathand coxae and femora of legs sometimes suffused with dark brown or fuscous.Forewings dark brown to blackish, usually marked with whitish fasciae and spots,the extreme basal costal area, an oblique fascia running from middle costa to apexof scutellum, the subapical costal and subcostal area and an oblique fascia runningfrom the basal inner edge of the marking on the subapical costal and subcostalarea to the claval apex, the apex of clavus and the area beyond apex of clavus, andthe subapical corial area usually whitish; the fascia sometimes greatly reduced andwhitish-spotted (Figure 1).

Body with dorsum distinctly punctate. Head (Figures 1(A–L), 2A, 3A, 4A, 5A)relatively short and broad, shorter than pronotum, distinctly wider betweeneyes than long in middle, slightly narrower than pronotum at shoulders. Vertex(Figures 1A,C,E,G,I,K, 2A, 3A, 4A, 5A) somewhat triangular in shape, sloping ante-riorly, flat to undulate dorsally, usually with a median longitudinal carina (nearlyobsolete in A. pseudotiegsi sp. nov.) and a small callous fovea on posterior marginalarea between ocellus and eye, anterior margin clearly defined and carinate. Eyes dis-tinctly longer than wide in dorsal view. Tylus (Figures 1(A–L), 2A, 3A, 4A, 5A)short and transversely long, about three times as broad as median length, with pos-terior margin arched anteriorly in middle. Ocelli very close to each other. Frons(Figures 1B,D,F,H,J,L, 2B, 3B, 4B, 5B) strongly compressed and depressed laterally,narrow and keel-like medially or moderately bulbous in males, and usually swollen, notstrongly compressed laterally in females. Antennae with flagellar base expanded andvisible, bearing an apical, broadly conical process being next and parallel to the flag-ellar bristle and nine small coeloconic sensilla on ventral side (see Liang et al., 2005).Rostrum relatively long, nearly reaching hind coxae.

Pronotum (Figures 1A,C,E,G,I,K, 2A, 3A, 4A, 5A) usually with a median lon-gitudinal carina (nearly obsolete in A. pseudotiegsi sp. nov.), anterior margin evenlyconvexly rounded or slightly angulate in middle, anterior half distinctly callouscentrally, posterior half transversely rugose; anterior lateral margin relatively long,shorter than longitudinal axis of eyes. Scutellum nearly triangular, centrally slightly

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Figure 1. Anyllis species, dorsal and lateral habitus. (A–D) Anyllis gibbosus sp. nov.; (A, B) male,paratype; (C, D) female, paratype, Australia: Queensland; (E, F) Anyllis leiala, male, Australia:New South Wales; (G, H) Anyllis pseudotiegsi sp. nov., male paratype, Australia: Tasmania; (I, J)Anyllis spinostylus, male paratype, Australia: Tasmania; (K, L) Anyllis tiegsi, male, Australia:New South Wales.

depressed and transversely rugose. Forewings (Figure 1A–L) narrow, with very sparsepubescence, costal margin evenly convex from base to apex, venation reticulate api-cally. Hindwings translucent, veins brown. Hind tibia with one strong spur on lateraledge at distal two-fifths and 13–16 black-tipped spines apically, metatarsomeres I andII with 5–8 and 6–11 black-tipped spines apically, respectively.

Male genitalia with pygofer (Figures 2C, 3C, 4C, 5C) in lateral view rela-tively short, distinctly high (compared with typical Aphrophora species) and nearly

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subquadrate, slightly wider ventrally than dorsally. Anal segments (Figures 2C,E,3C,E, 4C,E, 5C,E) relatively short and small, basal anal segment with a pair ofposteriorly directed, spinous processes on ventral margin, the inner edge of thosespinous processes usually covered with fine spines. Subgenital plates (Figures 2C,D,3C,D, 4C,D, 5C,D) reduced, short and small (compared with typical Aphrophoraspecies), completely fused to pygofer, apically slightly bilobed or sometimes parabola-like in ventral view, right and left plates separated each other, forming a V-shapedempty space and making the base of aedeagus visible in ventral view. Genital styles(Figures (2–5)C, (2–5)D, (2–5)F, (2–5)G) relatively elongate and slender, sinuate inlateral view; mesially expanded outwards, narrowed to apex over distal half, apexbidentate; arched inward over distal half with extreme apex nearly touching each otherin ventral view. Aedeagal shaft (Figures 2C,G, 3C,G, 4C,G, 5C,G) short, tubular, ante-riorly arched in lateral view, anterior edge of the shaft covered with very fine spines,gonopore apical.

BiologyBiological data for the species Anyllis is scarce. Anyllis leiala is one of the mostcommon and widespread aphrophorids found in Australia. It is commonly attractedto mercury-vapour lights and is frequently swept from a wide range of native,shrubby, dicotyledonous plants (Fletcher 2010; Liang and Fletcher 2003; Liang et al.2005). Morrow (1977) reported that A. leiala is associated with Eucalyptus pauciflora,Eucalyptus perriniana and Eucalyptus stellulata and Ohmart et al. (1983) recorded theassociation of A. leiala with Eucalyptus delegatensis.

RemarksAnyllis is the only genus of the tribe Aphrophorini distributed in the Australian region.Members of Aphrophorini are widely distributed in the Palaearctic, Nearctic andOriental regions with generic diversification primarily in the Oriental region. Anyllisdiffers from other members of Aphrophorini in the following characters: frons in themales usually thin and strongly compressed laterally; antennae with expanded flagel-lar base having an apical, broadly conical process (a special bulbous sensillum) beingadjacent and parallel to the flagellar bristle and nine small coeloconic sensilla on ven-tral side, lacking placoid sensilla subapically (Aphrophorini usually have three largeplacoid sensilla on apical area of the expanded flagellar base, Liang, unpublisheddata; see also Liang and Fletcher 2002); hind tibiae with one lateral spur on outeredge (Aphrophorini usually have two lateral spurs on hind tibiae with exception ofSinophora with four to six lateral spurs); basal anal segment in males with a pair ofbasal anal processes; subgenital plates short and separated from each other; genitalstyles slender and elongate; aedeagal shaft relatively elongate and slender, coveredwith fine spines on anterior edge in lateral view. These characters are autapomor-phies of Anyllis and strongly support the monophyly of the genus. These may reflectthe long independent evolution of Anyllis because of its long isolation from otherAphrophorini, which are predominantly distributed in the Holarctic region (Lianget al. 2005).

Anyllis can be easily distinguished from other Australian aphrophorid genera byits punctate dorsum, strongly compressed and depressed frons in males, one lateral

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spur on hind tibiae and the presence of a pair of basal anal processes on the basal analsegment in males.

DistributionSpecies of Anyllis are currently known from New South Wales, Victoria, Queensland,Australian Capital Territory and Tasmania in Australia. No specimens of Anyllisspecies were collected from Western Australia and Northern Territory, althoughthis is probably an artefact of collection. It appears that the genus is endemic toAustralia.

Key to the species of Anyllis

1. Frons (Figures 2B, 3B) in males strongly compressed and depressed laterally,narrow and keel-like medially, with a distinct median longitudinal carina . . 2Frons (Figures 4B, 5B) in males moderately bulbous, with an indistinctmedian longitudinal carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Body more elongate and larger (length male 6.5–6.9 mm, female 8.7–9.0 mm),reddish brown (Figure 1A–D); pygofer with basal anal processes withinner edges in males without fine spines (Figure 2C,E); subgenital plates(Figure 2C,D) apically parabola-like; aedeagal shaft (Figure 2C,G) relativelystout and broad, straight in lateral view, with a gibba-like process on anterioredge subapically in lateral view; Queensland . . . . . . . . . . . . A. gibbosus sp. nov.Body smaller and relatively oval (length male 5.1–5.9 mm, female 5.8–6.8 mm), colour not as above (Figure 1E,F); pygofer with basal anal processwith inner edges in males covered with fine spines (Figure 3C,E); subgenitalplates (Figure 3C,D) apically bilobed, aedeagal shaft (Figure 3C,G) more slen-der, distinctly arched anteriorly in lateral view, without a gibba on anterioredge in lateral view; Queensland, New South Wales, Victoria and Tasmania. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. leiala Kirkaldy

3. Body relatively elongate and large, length male 5.8–6.0 mm, female 7.0–7.2 mm; genital styles more narrow and slender, apex hook-like in ventralview; aedeagal shaft elongate and slender (see Liang et al. 2005: 304, figs12–16); Tasmania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spinostylus LiangBody relatively smaller and more oval, length male 4.5–5.1 mm, female6.0 mm; genital styles relatively shorter and broader, apex bidentate in ventralview (Figures 4C,F,G, 5C,F,G); aedeagal shaft short and stout (Figures 4C,G,5C,G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Body slightly larger, length male 5.0–5.1 mm, female 6.0 mm; vertex with amedian longitudinal carina (Figures 1K, 5A); pronotum with median longitu-dinal carina obsolete on posterior half (Figures 1K, 5A); pygofer with basalanal processes with inner edges covered with fine spines on apical two-thirds(Figure 5E); New South Wales and Victoria . . . . . . . . . . . . . . . A. tiegsi (China)Body distinctly small, short and oval, length male 4.5–4.8 mm, femaleunknown; vertex and pronotum without median longitudinal carina(Figures 1G, 4A); pygofer with basal anal processes with inner edgescovered with fine spines on apical half (Figure 4C,E); Tasmania. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pseudotiegsi sp. nov.

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Anyllis gibbosus sp. nov.(Figures 1A–D, 2A–G)

Description

Body large, distinctly elongate and slender, length (from apex of vertex to tip offorewings): male (n = 9), 6.5–6.9 mm; female (n = 3), 8.7–9.0 mm.

General colour reddish brown, vertex, pronotum and scutellum covered with fus-cous punctures, anterior margin of tylus and median carina of pronotum black,scutellum with median line suffused with yellowish white; head beneath reddish brown,frons suffused with dark brown, antennal sockets, scape and pedicel reddish brown;rostrum with apical segment dark brown; thorax beneath pale yellowish with pleu-rae partly brownish; legs pale yellowish, claws brownish, tips of spines on hind tibiaeand tarsi black; forewings (Figure 1A–D) reddish brown, the extreme costal area, asmall costal marking beyond middle, a relatively large subtriangular costal markingat subapical one-third, whitish; some indistinct spots scattered on clavus and corium,the subapical area of calvus and the cellar areas at apical one-fifth very pale whitish;hindwings pale brown, covered with brown hairs, veins brown; abdomen fuscous withsegmental margins ochraceous.

External structure as in the generic description. Head (Figures 1A,C, 2B)maximum width (including eyes) 3.7–4.0 times as long as median length. Vertex(Figures 1A,C, 2B) with a distinct median carina, median length 0.4–0.5 times aslong as length of pronotum medially. Frons (Figures 1B,D, 2A) in the males verystrongly compressed and depressed laterally, thin in lateral view, median longitudinalcarina distinct. Pronotum (Figures 1A,C, 2B) with median longitudinal carina reach-ing to posterior margin of pronotum, median length 0.5–0.6 times as long as maximumwidth.

Male genitalia with pygofer (Figure 2C) high, wider ventrally than dorsally, ante-rior and posterior margins nearly straight and quadrate in lateral view. Basal analprocesses (Figure 2C,E) elongate, slender and curved, inner margins without finespines. Subgenital plates (Figure 2C,D) relatively elongate, apically parabola-like.Genital styles (Figure 2C,D,F,G) relatively elongate and slender, broad mesially, nar-rowed to apex over distal one-third, apically bidentate. Aedeagal shaft (Figure 2C,G)short and broad, straight, with a gibba-like process on anterior edge subapically inlateral view, the process with its semicircular outer edge covered with fine spines;gonopore dorsal.

Material examined

Holotype. Male, [Australia, Queensland] 6.4 km west of Paluma, Qld, 900 m, 25 April1969 (I.F.B. Common & M.S. Upton) (ANIC).

Paratypes. [Australia, Queensland] six males, three females, same data as holotype(ANIC); two males, Mt Edith, 29 km northeast of Atherton, Q., 1020 m above sealevel, 18 March 1964 (I.F.B. Common and M.S. Upton) (ANIC).

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Figure 2. Anyllis gibbosus sp. nov. (A) Head, pronotum and scutellum, dorsal view; (B) head,ventral view; (C) male genital capsule, lateral view; (D) male genital capsule, ventral view; (E)anal segment and basal anal processes, ventral view; (F) genital styles and base of aedeagus,ventral view; (G) aedeagus and left genital style, left view. Scale bars: A, B 0.5 mm; C–G0.125 mm.

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Etymology

This new species is named for the presence of a gibba-like process on the anterior edgeof the aedeagal shaft subapically in lateral view (Figures 2E–G).

Distribution

Australia (Queensland).

Remarks

This is the largest species currently known in the genus. It can be easily distinguishedfrom other known species in the genus by its largest size, more elongate and slen-der body (Figure 1A–D) and the male genital structure, especially the aedeagal shaftwith a gibba-like process on anterior edge subapically in lateral view (Figure 2C,G).Externally this species is similar to A. leiala Kirkaldy, but differs from the latter in itsvertex, pronotum and mesonotum covered with fuscous punctures, pronotum with themedian longitudinal carina black, and the scutellum with median line suffused withyellowish white, in addition to the differences in the body size and the male genitalcharacters mentioned above.

Anyllis leiala Kirkaldy(Figures 1E,F, 3A–G)

Anyllis leiala Kirkaldy, 1906: 386; Metcalf, 1961: 63; Liang and Fletcher, 2003: 86,fig. 2; Liang et al., 2005: 305. Syntype one male, Australia (Queensland: Kuranda)(BPBM) [examined].

Aphrophorinella tonnoiri Lallemand 1946: 195; Metcalf, 1962: 551. Syntypes six males,nine females, Australia (Tasmania) (BMNH) [examined]. [Synonymized by Evans1966: 317.]

Description

Length (from apex of vertex to tip of forewings): males (n = 36), 5.1–5.9 mm; females(n = 15), 5.8–6.8 mm.

General colour ochraceous brown, carinae on vertex and pronotum suffused withbrown, scutellum with brown suffusion medially; head beneath dark brown to black,frons suffused with black, and antennal sockets, scape and pedicel blackish; rostrumwith apical segment dark brown; thorax beneath dark brown with pleurae partlyochraceous; legs and claws dark brown, femora with apical part and hind tibiae andtarsi pale yellowish, fore and middle tibiae brown, tips of spines on hind tibiae andtarsi black; forewings (Figure 1E,F) dark brown, clavus ochraceous, apex of clavusmarked with hyaline white, with three hyaline white areas on costal area: the first verynarrow and small, situated at basal costal area; the second near middle, short andsub-rectangular; the third subapical, larger than the second, nearly oval; hindwingspale brown, covered with brown hairs, veins brown; abdomen fuscous with segmentalmargins ochraceous.

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Figure 3. Anyllis leiala. (A) Head, pronotum and scutellum, dorsal view; (B) head, ventral view;(C) male genital capsule, lateral view; (D) male genital capsule, ventral view; (E) anal segmentand basal anal processes, ventral view; (F) genital styles and base of aedeagus, ventral view; (G)aedeagus and left genital style, left view. Scale bars: A, B 0.25 mm; C–G 0.125 mm.

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External structure as in the generic description. Head (Figures 1E, 3A) maximumwidth (including eyes) 4.0–4.7 times as long as median length. Vertex (Figures 1E,3A) with median carina distinct, median length 0.3–0.4 times as long as length ofpronotum medially. Frons (Figures 1F, 3B) in the males strongly compressed anddepressed laterally, median longitudinal carina distinct. Pronotum (Figures 1E, 3A)with median longitudinal carina reaching to posterior margin of pronotum, medianlength 0.5–0.6 times as long as maximum width.

Male genitalia with pygofer (Figure 3C) high, slightly wider ventrally than dorsally,anterior margin concave submedially, posterior margin slightly produced posteriorlysubapically in lateral view. Basal anal processes (Figure 3C,E) elongate and curved,inner margins covered with distinct fine spines. Subgenital plates (Figure 3C,D) rel-atively long, apically bilobed, both plates separated from each other and leaving aV-shaped empty space between them. Genital styles (Figure 3C,D,F,G) relatively slen-der, with apical half narrowed towards apex, curved and apically bidentate. Aedeagalshaft (Figure 3C,G) short and sinuate, distinctly arched posteriorly medially in lateralview, anterior edge covered with many fine spines in lateral view.

Type material examined

Anyllis leiala Kirkaldy 1906, Syntype one male, Queensland: Kuranda, August 1904(Perkins) (BPBM). Aphrophorinella tonnoiri Lallemand 1946, syntypes six males,nine females, Tasmania: Cradle Mountain, Alpine Creek, Mt Hartz, A.L. Tonnoir;Margate, Muir; Mt Wellington, C. Davis; New South Wales: Mt Victoria, J.W. Evans(BMNH).

Anyllis leiala Kirkaldy 1906, Syntype one male, Queensland: Kuranda, August1904 (Perkins) (BPBM). Aphrophorinella tonnoiri Lallemand 1946, Syntypes six males,nine females, Tasmania: Cradle Mountain, Alpine Creek, Mt Hartz, A.L. Tonnoir;Margate, Muir; Mt Wellington, C. Davis; New South Wales: Mt Victoria, J.W. Evans(BMNH).

Other material examined

Australia, Tasmania: four males, Lake Leake, T., 600 m, 27 February 1963 (I.F.B.Common and M.S. Upton); two males, Eaglehawk Neck, T., 26 February 1963, I.F.B.Common and M.S. Upton; three males, Lyell Hwy. at Franklin R., 55 km, esast-southeast Queenstown, 400 m, 19 February 1980 (A. Newton and M. Thayer) (all inANIC), open Eucalyptus forest, window trap; [green label]. If designated as holotypespecimen must be returned to Australia (AMNH). Australia, Victoria: five males, MtWellington, T. 720 m, 25 February 1963 (I.F.B. Common and M.S. Upton) (ANIC).Australia, New South Wales: one male, 8 km northwest of Coffs Harbour, NSW,1 November 1965 (M.S. Upton); one male, same data as preceding but 240 m; onemale, one female, 35.34◦ S, 148.47◦ E, Mt Gingera, ACT, 23 February 1984 (A. Calderand M. Stevens); three males, one female, 36.12◦ S, 148.43◦ E, Dainers Gap, NSW,6 February 1974 (P. Morrow), Ex Eucalyptus pauciflora, 1585 m, E. pauciflora, E. stel-lulata and E. perriniana forest; one male, same data as preceding but 14 February 1973;two males, same data but 20 March 1974; one male, same data but February 1973; onemale, same data but 21 February 1973; one male, same data but 21 February 1974;

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one male, same data but 22 January 1974; one male, same data but 28 February 1973;one male, same data but 29 January 1973 (all in ANIC); two males, Illawarra, NSW,no date (H. Petersen) (USNM); one male, NSW, Kuringai Chase, 8 November 1967 (J.and M. Sedlacek) (BPBM); one male, NSW Roy. Nat. Park, 40 km south of Sydney,23 April 1960 (C.W. O’Brien), dry sclerophyll forestry (AMNH); one male, Sydney,November 1967 (J. and M. Sedlacek) (BPBM).

Distribution

Australia (throughout the eastern part of Australia from north Queensland toTasmania).

Remarks

Anyllis leiala is one of the most common species of Australian spittlebugs. It is fre-quently attracted to lights and can be swept from a wide range of shrubby native plants(Fletcher 2010; Liang and Fletcher 2003). This species can be distinguished from otherknown species in the genus by its strongly compressed and depressed frons with dis-tinct median carina and the minutiae of the male genital structure (Figure 3C–G). It issimilar to A. gibbosus sp. nov., but can be separated from the latter by its body beingdistinctly smaller and relatively oval in dorsal view (Figure 1E,F), the pygofer with thebasal anal processes with inner margins covered with distinct fine spines (Figure 3C,E)and the aedeagal shaft relatively narrow, sinuate, mesially distinctly arched posteri-orly in lateral view and the anterior edge without a gibba-like process subapically(Figure 3C,G).

Anyllis pseudotiegsi sp. nov.(Figures 1G–H, 4A–G)

Description

Small, oval species, length (from apex of vertex to tip of forewings): male (n = 3),4.5–4.8 mm.

General colour stramineous brown, frons pale yellowish with base black, clypeuspale yellowish, antennal sockets, scape and pedicel black; rostrum with apical seg-ment brown; thorax beneath brownish; legs and claws dark brown, hind coxae andtrochanters pale yellowish, hind tibiae and tarsi pale brown, tips of spines on hind tib-iae and tarsi black; forewings (Figure 1G,H) fuscous, the apical marginal area blackish,with the following areas or fasciae whitish: the extreme basal costal area, an obliquefascia running from middle costal margin to clavus beyond apex of scutellum, a sub-triangular marking on the subapical costal and subcostal area (the veins within themarking brown) and an indistinct, narrow, oblique fascia running from the inner basaledge of this whitish subapical costal and subcostal marking to the claval apex; the apexof clavus and the subapical corium whitish; hindwings hyaline, covered with brownhairs, veins brown; abdomen black with segmental margins white.

External structure as in the generic description. Head (Figures 1G, 4A) maximumwidth (including eyes) 3.3–3.4 times as long as median length. Vertex (Figures 1G, 4A)without median carina, median length 0.5–0.6 times as long as length of pronotummedially. Frons (Figures 1H, 4B) in the males moderately bulbous, not strongly

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Figure 4. Anyllis pseudotiegsi sp. nov. (A) Head, pronotum and scutellum, dorsal view; (B) head,ventral view; (C) male genital capsule, lateral view; (D) male genital capsule, ventral view; (E)anal segment and basal anal processes, ventral view; (F) genital styles and base of aedeagus,ventral view; (G) aedeagus and left genital style, left view. Scale bars: A, B 0.25 mm; C–G0.125 mm.

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compressed laterally, median longitudinal carina indistinct. Pronotum (Figures 1G,4A) without median longitudinal carina, median length 0.5–0.6 times as long asmaximum width.

Male genitalia with pygofer (Figure 4C) high, slightly narrow, width ventrallynearly as wide as dorsally, subquadrate in lateral view, anterior margin sinuate,dorsal posterior margin angularly produced in lateral view. Basal anal processes(Figure 4C,E) relatively short, directed posterodorsally in lateral view, inner edgecovered with fine spines on apical half. Subgenital plates (Figure 4C,D) short,apically bilobed, both plates widely separated from each other. Genital styles(Figure 4C,D,F,G) relatively short, narrow and slender, evenly tapered sub-basally toapex, apically bidentate. Aedeagal shaft (Figure 4C,G) short and stout, slightly archedanteriorly in lateral view, anterior edge covered with many tiny spines in lateral view;gonopore apical.

Material examined

Holotype. male, [Australia] 5 km east of Stanley, Tas., 15 November 1975, K.R. Norris(ANIC).

Paratypes. [Australia:] two males, same data as holotype (ANIC).

Etymology

This new species is named for its general appearance being similar to A. tiegsi (China,1952).

Distribution

Australia (Tasmania).

Remarks

This is the smallest species currently known in Anyllis. This species can be distin-guished from other known species in the genus by its small body size and distinctly ovalbody form (Figure 1G,H), vertex and pronotum without median carina (Figures 1G,4A), frons with indistinct median carina (Figure 4B) and the male genitalic structure(Figure 4C–G). In external appearance, this species is similar to A. tiegsi (China, 1952),but differs from the latter in the smaller body; vertex (Figure 4B) without mediancarina, median longitudinal carina on pronotum (Figure 4A) completely obsolete,and the pygofer with the basal anal process directed posterodorsally, with inner edgecovered with fine spines only on apical half (Figure 4C,E).

Anyllis spinostylus Liang(Figures 1I–J)

Anyllis spinostylus Liang in Liang et al., 2005: 301–307, figs. 10–16. Holotype male,Australia (Tasmania) (BPBM) [examined].

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Description

Length (from apex of vertex to tip of forewings): male (n = 4), 5.8–6.0 mm; female(n = 5), 7.0–7.2 mm.

General colour ochraceous brown, posterior area of pronotum pale fuscous,scutellum with brown suffusion medially; head beneath ochraceous, antennal socketsand flagellar base pale fuscous, rostrum with apical segment brown; thorax beneathochraceous with pleurae partly brownish; legs ochraceous, claws dark brown, fore andmiddle tarsi brown, tips of spines on hind tibiae and tarsi black; forewings (Figure 1I,J)fuscous, marked with some stramineous spots or irregular markings, with three hyalinewhite areas on costal area, the first basal and very narrow, the second near middle andbeing very short and small, and the third subapical, large and elongate, nearly triangu-lar; hindwings hyaline, covered with brown hairs, veins brown; abdomen dark brownto fuscous with segmental margins ochraceous.

External structure as in the generic description. Head (Figure 1I) maximum width(including eyes) 3.2–4.0 times as long as median length. Vertex (Figure 1I) with adistinct median carina, median length 0.4–0.5 times as long as length of pronotummedially. Frons (Figure 1J, also see Liang et al. 2005: 304; fig. 11) in the males moder-ately bulbous, not strongly compressed laterally, median longitudinal carina indistinct.Pronotum (Figure 1I, also see Liang et al. 2005: 304; fig. 10) with median longitudinalcarina obsolete on posterior half, median length 0.6–0.7 times as long as maximumwidth.

Male genitalia with pygofer high, wider ventrally than dorsally, upper posteriorend angularly posteriorly produced in lateral view. Anal segments very short andbroad, anal style small; basal anal processes robust and straight, with inner edgewithout fine spines. Subgenital plates very short and small, apically weakly bilobed,outer margin with one very small angular process. Genital styles slender and elon-gate, angularly expanded laterad medially, apex hook-like in ventral view. Aedeagalshaft distinctly slender and elongate, slightly arched anteriorly in lateral view, anterior–ventral margin with an angular process subapically, upper anterior edge covered withfine spines in lateral view (see Liang et al. 2005: 304; figs 12–16).

Material examined

Holotype male, Australia: Tasmania, Mt Field, Nat. Park, Dobson Lake, 1000 m,25 December 1960, collected by J.L. Gressitt, deposited in the Bernice P. BishopMuseum, Honolulu, Hawaii, USA (BPBM).

Paratypes. Australia: two males, Lake Fenton, Mt Field National Park, Tasmania,1000 m, 42◦40.6′ S, 146◦37.4′ E, 6 December 1999 (J. Keble-Williams), beatingNothofagus cunninghamii (ASCU); two females, same data but collected on 25 January2000 (ASCU); one male, one female, Mt Arthur, Wellington Range, Tasmania, 1080 m,42◦53.0′ S, 147◦13.1′ E, 10 February 2000, J. Keble-Williams, beating Nothofagus cun-ninghamii (ASCU); one female, Tasmania, Mt Field National Park, Lake DobsonRoad, 710 m, 2 February 1980 (A. Newton and M. Thayer), Eucalyptus, Nothofagusforest, window trap; [green label]. If designated as holotype specimen must be returnedto Australia (AMNH); one female, Tasmania, Hartz Mts NP, Hartz Road, 800 m,8–10 February [19]80 (A. Newton and M. Thayer), Eucalyptus Scrub, moor edge;

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beating Eucalyptus sp.; [green label]. If designated as holotype specimen must bereturned to Australia (AMNH).

Distribution

Australia (Tasmania).

Remarks

This species can be distinguished from another Tasmanian species A. pseudotiegsi sp.nov. by its relatively elongate body and large size (length male 5.8–6.0 mm, female7.0–7.2 mm); vertex with a median carina; pronotum with median carina obsolete onposterior half; forewings (Figure 1I,J) marked with more stramineous spots; genitalstyles more narrow and slender, apex hook-like in ventral view; and aedeagal shaftelongate and slender (see Liang et al. 2005: 304, figs 12–16).

Anyllis tiegsi (China)(Figures 1K–L, 5A–G)

Neoaphrophora tiegsi China, 1952: 791, figs. 1G,H, 2A–C; Metcalf, 1962: 431; Evans,1966: 316, 317; Liang et al., 2005: 305. Holotype male, Australia (Victoria) (BMNH)[examined].

Anyllis tiegsi (China); Liang and Fletcher, 2003: 86–87, fig. 3; Liang et al., 2005:301–305.

Description

Length (from apex of vertex to tip of forewings): male (n = 2), 5.0–5.1 mm; female(n = 1), 6.0 mm.

General colour pattern and external appearance similar to A. pseudotiegsi sp.nov. General colour stramineous brown, scutellum with brown suffusion medially;frons stramineous with base dark brown; antennal sockets, scape and pedicel blackor dark brown; clypeus stramineous; rostrum with apical segment pale yellowish; tho-rax beneath dark brown with pleurae partly yellowish; legs and claws dark brown,hind coxae and trochanters pale yellowish, hind tibiae and tarsi pale brown, tips ofspines on hind tibiae and tarsi black; forewings (Figure 1K,L) brown, apical marginalarea nearly black, clavus with basal two-thirds stramineous, corium with the followingareas or fasciae whitish: the extreme basal costal area, an oblique fascia running frommiddle costal margin to clavus beyond apex of scutellum, a subtriangular marking onthe subapical costal and subcostal area (the veins within the marking brown) and anoblique fascia running from the inner basal edge of this whitish subapical costal andsubcostal marking to the claval apex; the apex of clavus and two irregular spots onsubapical corium whitish. Abdomen black with segmental margins white. Female withblack punctures much reduced ventrally, legs with femora and pleura ochreous.

External structure as in the generic description. Head (Figures 1K, 5A) maximumwidth (including eyes) 2.9–3.5 times as long as median length. Vertex (Figures 1K, 5A)

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Figure 5. Anyllis tiegsi. (A) Head, pronotum and scutellum, dorsal view; (B) head, ventral view;(C) male genital capsule, lateral view; (D) male genital capsule, ventral view; (E) anal segmentand basal anal processes, ventral view; (F) genital styles and base of aedeagus, ventral view; (G)aedeagus and left genital style, left view. Scale bars: A, B 0.25 mm; C–G 0.125 mm.

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with a median carina, median length 0.6–0.7 times as long as length of pronotummedially. Frons (Figures 1L, 5B) in males moderately bulbous, not strongly com-pressed laterally, median longitudinal carina indistinct. Pronotum (Figures 1K,5A) with median longitudinal carina obsolete on posterior half, median length0.5–0.6 times as long as maximum width.

Male genitalia with pygofer (Figure 5C) high, wider ventrally than dorsally, ante-rior margin concave medially and posterior margin nearly straight in lateral view.Basal anal processes (Figure 5C,E) relatively short, the inner edges covered withfine spines on apical two-thirds. Subgenital plates (Figure 5C,D) relatively short andbroad, apically distinctly bilobed, both plates widely separated each other. Genitalstyles (Figure 5C,D,F,G) relatively stout, moderately broad mesially, evenly narrowedtowards apex over apical one-third in lateral view, apically bidentate. Aedeagal shaft(Figure 5C,G) relatively stout and broad, slightly arched anteriorly, anterior edgecovered with many fine spines in lateral view; gonopore dorsal.

Material examined

Holotype Male, [Australia:] Victoria, Melbourne, O.W. Tiegs (BMNH).Holotype ♂, [Australia:] Victoria, Melbourne, O.W. Tiegs (BMNH).

Other material examined

Australia, New South Wales: one male, New England National Park, NSW,11 February 1968 (D.H. Colless) (ANIC). Australia, Victoria: one female, 15 km westBallarat, Vic., 12 November 1969 (E.F. Riek) (ANIC).

Distribution

Australia (Victoria and the higher parts of southeast New South Wales).

Remarks

This species is similar to A. pseudotiegsi sp. nov. from Tasmania but can be distin-guished from the latter by the slightly larger body (length male 5.0–5.1 mm, female6.0 mm); vertex with a median longitudinal carina (Figures 1K, 5A); pronotum withmedian longitudinal carina obsolete on posterior half (Figures 1K, 5A); pygofer withbasal anal processes with inner edges covered with fine spines on apical two-thirds(Figure 5E) and the minutiae of the male genitalia (Figure 5A–G).

Acknowledgements

We are grateful to Dr John LaSalle and Mrs Nicole Fisher (ANIC), Mr David J. Preston andDr Scott Miller (BPBM), Dr Mick D. Webb (BMNH), Dr Murray J. Fletcher (ASCU) andDr Randall T. Schuh (AMNH) for loans of the specimens for study. This work resulted from thesenior author’s visit to the Australian National Insect Collection, CSIRO, Black Mountain,Canberra, Australia in April–May 2005, funded by the exchange programme between theChinese Academy of Science (CAS) and the CSIRO. The senior author wishes to thank Mr Ren-Hua Sun (CAS, Beijing), Dr Ta-Yan Leong (CSIRO, Canberra), Dr John LaSalle, Mrs Nicole

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Fisher, Mr Chris Manchester and Dr Lawrence L. Mound (ANIC), for various assistance dur-ing his stay in Canberra. This work was supported by the National Natural Science Foundationof China (grant nos. 31172128 and 39925006 (awarded to A-P.L.) and 30900145 (awarded toR-R.W.)).

References

China WE. 1952. A new genus and species of Aphrophorinae (Cercopidae, Hemiptera–Homoptera) from Australia. Ann Mag Nat Hist. 5(12):789–792.

Evans JW. 1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera:Cicadelloidea and Cercopoidea). Aust Mus Mem. 12:1–347.

Fletcher MJ. 2010. Identification Key and Checklists for the Froghoppers and Spittlebugs(Hemiptera: Cercopoidea) of Australia and neighbouring areas. [Cited 28 April 2011].Available from http://www1.dpi.nsw.gov.au/keys/cercopid/index.html

Kirkaldy GW. 1906. Leafhoppers and their natural enemies. Bull Hawaiian Sugar Plant AssocExp Stn Entomol Ser. 1(9):271–479.

Lallemand V. 1912. Homoptera Fam. Cercopidae. Gen Insect. 143:1–167.Lallemand V. 1946. Quatrième Note sur les Cercopides. Bull Ann Soc Entomol Belg 82:189–197.Liang A-P, Combéfis É, Jiang G-M. 2012. Revision of the spittlebug genus Euryaulax Kirkaldy

(Hemiptera: Cercopoidea: Cercopidae) from Australia. Ann Entomol Soc Am 105(2).Liang A-P, Fletcher MJ. 2002. Morphology of the antennal sensilla in four Australian spit-

tlebug species (Hemiptera: Cercopidae) with implications for phylogeny. Aust J Entomol.41:39–44.

Liang A-P, Fletcher MJ. 2003. Review of the Australian aphrophorid spittlebugs (Hemiptera:Aphrophoridae). Aust J Entomol 42(1):84–93.

Liang A-P, Fletcher MJ, Jiang G-M. 2005. The endemic Australian spittlebug genus AnyllisKirkaldy (Hemiptera: Aphrophoridae) with description of a new species from Tasmania.Journal of the Kansas Entomol Soc. 78(4):301–307.

Metcalf ZP. 1961. General Catalogue of the Homoptera. Fasc. VII. Cercopoidea, Part 2.Cercopidae. Baltimore (MD): Waverly Press, Inc. 607 pp.

Metcalf ZP. 1962. General Catalogue of the Homoptera. Fasc. VII. Cercopoidea, Part 3.Aphrophoridae. Baltimore (MD): Waverly Press, Inc. 600 pp.

Morrow P. 1977. Host specificity of insects in a community of three co-dominant Eucalyptusspecies. Aust J Ecol. 2:89–106.

Ohmart CP, Stewart LG, Thomas JR. 1983. Leaf consumption by insects in three Eucalyptusforest types in southeastern Australia and their role in short-term nutrient cycling.Oecologia (Berlin). 59:322–330.

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