Bioacoustics of Cicadidae

REMARKS.—The type genus was not included in the recent family level genomic study ( Marshall et al. 2018 ) but several authors have suggested an affinity of Plautilla to Zammara , beginning with Stål (1865) . Jacobi (1907a) also described the structural similarities of Plautilla and Zammara . Handlirsch (1925) placed the genus Plautilla near Zammara within the Cicadini Latreille, 1802 . The genus was classified within the Tibicininae Distant, 1905d in Metcalf (1963c) but separated in its own family by Boulard (1975) . Moulds (2005) provided a detailed morphological cladistic analysis and demonstrated that the Plautillini formed a clade with the Zammarini . However, he was cautious about changing more than the position of the Plautillini from the Tibicininae to the Cicadinae choosing to await molecular studies to clarify the situation further ( Moulds 2005 ). Goemans (2016) performed these molecular analyses and proposed the Plautillini as a junior synonym of the Zammarini in a reference not recognized by the Code as making a valid change to its status. The genetic analyses showed that Plautilla and Onoralna Boulard, 1996 form a distinct clade from the species of the subtribe Zammarina and Dazina Kato, 1932 ( Goemans 2016 ) . As a result, I am forming the subtribe Plautillina rev. stat. Species of the subtribe Plautillina rev. stat. are characterized by the small head with prominent antennae, posterior extensions on the metanotum that form the timbal covers in Plautilla but are reduced to a small extension over the anterior timbal cavity in Onoralna , fore wing ulnar cell 1 not as long as apical cell1 or 2, hind wing jugum very small, males possess large inflated opercula that extend laterally beyond the anterior abdomen, the abdomen is narrowed in segments 1 and 2 and expands to its widest at segment 3, and there is a well developed mesonotal stridulatory apparatus. Onoralna was originally assigned to the Dazini (Boulard 1996) and was moved along with other genera of the tribe to the subtribe Dazina when the subtribe was formed based on the primary structure used to distinguish Dazina from Zammarina , a curled ridge along the posterior timbal cavity margin ( Sanborn 2018a ). However, the species of Onoralna possesses the complete set of characters of the Plautillina rev. stat. and is reassigned to the Plautillina rev. stat. here. This morphologically based assignment is supported by the molecular phylogenies of Goemans (2016) .

1 Metanotum entirely concealed at dorsal midline, fore wing cubitus anterior 1 division such that proximal portion is longest, timbal covers present Cicadinae Latreille 1802 …........................................................... 2 - Metanotum partially concealed at dorsal midline, fore wing cubitus anterior 1 division such that proximal portion is shortest or fore wings reticulated, timbal covers absent or with a partial turned-back rim ….................................. 31 2 Body vertically compressed, pronotal collar with lateral margin strongly ampliate, head narrower than pronotum, fore leg primary spine adpressed to femur Platypleurini Schmidt 1918 ….................................................. 3 - Body not vertically compressed, pronotal collar with lateral margin not strongly ampliate, head narrower than pronotum, fore leg primary spine erect …............................................................................. 23 3 Head about as wide as or wider than mesonotum.......................................................... …4 - Head about two-thirds the width of mesonotum ….......................................................... 16 4 Fore wing costal margin not prominently dilated nor arched at base, lateral pronotum formed into spine-like extension Platypleura Amyot & Audinet-Serville, 1843 ….................................... Platypleura spiculata Distant, 1905b - Fore wing costal margin prominently dilated or arched at base, lateral pronotum without spine-like extension ….......... 5 5 Head not frontally produced with transverse posterior postclypeus, not as long as pronotum without pronotal collar Kongota Distant, 1904a …........................................................ Kongota handlirschi ( Distant, 1897 ) - Head frontally produced with posterior postclypeus anteriorly curved, about at long as pronotum without pronotal collar Yanga Distant, 1904a …..................................................................................... 6 6 Fore wings and hind wings hyaline.....................................................................… 7 - Fore wings and hind wings opaque …..................................................................... 8 7 Silver-gray marking on fore wing, male opercula rounded...................... … Y. argyrea ( Melichar, 1896 ) rev. stat. - No silver-gray marking on the fore wing, lateral male operculum margin obliquely rounded....................................................................................................… Yanga pulverea ( Distant, 1882 ) 8 Hind wings ochraceous marked with piceous-brown or castaneous …............................................ 9 - Hind wings primarily piceous or piceous-brown …......................................................... 13 9 Hind wings piceous brown basally….............................................. Yanga bouvieri Distant, 1905b - Hind piceous brown or castaneous distally..............................................................… 10 10 Hind wing apical cells 1–3 not completely piceous brown..................................................… 11 - Hind wings apical cells 1–3 completely piceous brown or castaneous.........................................… 12 11 Hind wing with central piceous brown spot and piceous brown distal apical cells...... … Yanga guttulata ( Signoret, 1860 ) - Hind wing piceous brown of light intensity forming a central spot, spots in distal apical cells and within anal cell 2 …............................................................................... Yanga grandidieri Distant, 1905b 12 Hind wing piceous brown restricted to distal apical cell 6, no piceous brown on hind wing margin or distal anal cell 2 ….............................................................................. Yanga andriana ( Distant, 1899 ) - Hind wing castaneous covering most of apical cell 6, piceous brown on hind wing margin and in distal anal cell 2 …................................................................................... Yanga mahajangaensis n. sp. 13 Hind wings piceous or piceous-brown with lighter areas in distal apical cells …............ Yanga antiopa ( Karsch, 1890 ) - Hind wings piceous with white margins or white apical area …................................................ 14 14 Hind wings piceous with white lateral and posterior margins …........................... Yanga heathi ( Distant, 1899 ) - Hind wings piceous basally with white apical areas …....................................................... 15 15 Hind wing piceous in medial cell incomplete along median vein, piceous in cubital cells, anal cells 1 and 2 incomplete distally, no piceous spots on distal apical cell veins...................................... … Yanga brancsiki ( Distant, 1893 ) - Hind wing piceous, white in distal apical cells, piceous spots on distal apical cell veins......… Yanga hova ( Distant, 1901 ) 16 Fore wing costal membrane as broad or broader than costal area … Pycna Amyot & Audinet-Serville, 1843 .......... …17 - Fore wing costal membrane much narrower than costal area … Umjaba Distant, 1904a ….......................... 22 17 Body length greater than 37 mm .............................................. … Pycna itremensis Boulard, 2008 - Body length less than 37 mm …........................................................................ 18 18 Wingspan less than 100 mm …......................................................................... 19 - Wingspan greater than 110 mm .......................................................................… 20 19 Pronotum width about 16 mm , anterior pronotal angles oblique, mesonotum unmarked, abdomen piceous with dorsal tergites 1–3 with hour-glass shaped fulvous mark, opercula broadly margined with piceous, external margin of the hind wings dark fuscous …............................................................ Pycna madagascarensis ( Distant, 1879 ) - Pronotum width about 14 mm , anterior pronotal angles almost rectangular, mesonotum with lateral fascia, abdominal tergites piceous, opercula lacking broad piceous margin, external margin of the hind wings piceous …. Pycna angusta ( Butler, 1882 ) 20 Wingspan less than 120 mm , mesonotum with piceous spot on posterior lateral sigillae, scutal depression ground color................................................................... … Pycna strix Amyot & Audinet-Serville, 1843 - Wingspan greater than 120 mm , mesonotum without piceous spot on posterior lateral sigillae or with piceous scutal depression…............................................................................................ 21 21 Wingspan about 122 mm , scutal depression ground color, timbal covers contrasting abdominal tergites, abdominal tergites not covered with dense pile …........................................................ Pycna gigas ( Distant, 1881 ) - Wingspan about 135 mm , scutal depression piceous, timbal covers same color as abdominal tergites, abdominal tergites covered with dense pile........................................................... …. Pycna rudis ( Karsch, 1890 ) 22 Fore wing width about one-third fore wing length, rostrum reaching to anterior opercular margin ….............................................................................................. Umjaba alluaudi Distant, 1905b - Fore wing width almost half fore wing length, rostrum reaching past anterior opercular margin ….............................................................................................. Umjaba evanescens ( Butler, 1882 ) 23 Large species, body length 38–40 mm , wingspan 121–123 mm , hyaline fore wings with macular infuscation on fore wing radial and radiomedial crossveins and distal veins between apical cells, head with an elongated vertex, distance between eye and supra-antennal plate much longer than supra-antennal plate, aedeagus with one or two subapical cerci Macrotristriini Marshall & Moulds, 2018 Mouldspsaltria n. gen. ......................... Mouldspsaltria madegassa ( Boulard, 1996 ) n. comb. - Not as above Antankariini n. tribe ...................................................................... 24 24 Globular eyes angled dorsally slightly wider than anterior pronotum, anterior postclypeus not extending much beyond supraantennal plates, lateral supra-antennal plates extended anteriorly from vertex Antankaria Distant, 1904b ......................................................................................... Antankaria signoreti Metcalf, 1955 - Ovoid eyes not angled dorsally, wider than anterior pronotum, anterior postclypeus extended beyond supra-antennal plates forming a triangular anterior head, lateral supra-antennal plates not extended anteriorly from vertex Orientafroinsularis n. gen. ............................................................................................… 25 25 Wingspan greater than 95 mm ….............................. Orientafroinsularis martini ( Distant, 1905b ) n. comb. - Wingspan less than 95 mm ….......................................................................... 26 26 Wingspan greater than 80 mm …............................... Orientafroinsularis matilei ( Boulard, 2000 ) n. comb. - Wingspan less than 80 mm ….......................................................................... 27 27 Wingspan less than 60 mm , song begins as a series of short syllables followed by a period of rapidly produced syllables of increasing intensity followed by a period of various amplitude modulation becoming separated into individual syllables with peak energy about 11 kHz, distributed over the western half of Madagascar ….......................................................................................... Orientafroinsularis pulverulenta ( Distant, 1905b ) n. comb. - Wingspan greater than 60 mm .......................................................................... 28 28 Wingspan less than 70 mm ......................... … Orientafroinsularis elenae ( Boulard, 2001 ) n. comb. , rev. stat. - Wingspan 70–80 mm ….............................................................................. 29 29 Fore wing radial and radiomedial crossveins infuscated ….......... Orientafroinsularis nigrans ( Distant, 1904c ) n. comb. - Fore wing radial, radiomedial and medial crossveins infuscated …............................................. 30 30 Infuscation on distal fore wing veins between apical cells …........... Orientafroinsularis loici ( Boulard, 2000 ) n. comb. - No infuscation on distal fore wing veins between apical cells …....... Orientafroinsularis hova ( Distant, 1905b ) n. comb. 31 Pygofer distal shoulder undeveloped, pygofer upper lobe present, uncus absent of small and duck-bill shaped, retractable, claspers present, aedeagus restrained by claspers Cicadettinae Buckton, 1890 …...................................... 32 - Pygofer distal shoulder developed or undeveloped (in Hovana ), pygofer upper lobe absent or present (in Hovana ), uncus dominant, not retractable, of moderate to long length, claspers absent, aedeagus restrained before or below uncus Tettigomyiinae Distant, 1905e ….................................................................................... 46 32 Body length 10 mm or less, aedeagus without pseudoparameres or conjunctival claws, pygofer basal lobes more than half pygofer length, small upper pygofer lobes associated with basal lobes, claspers narrowing distally Nelcyndanini Marshall & Moulds, 2018 …............................................................................................ 33 - Body length greater than 10 mm , aedeagus with pseudoparameres or conjunctival claws, basal pygofer lobes not half the length of pygofer, upper pygofer lobes separate from basal lobes, may be elaborate …................................... 34 33 Hind wings with four apical cells ….................................. Nelcyndana madagascariensis Distant, 1905d - Hind wings with five apical cells …........................................ Nelbroma betrokaensis n. gen. , n. sp. , 34 Eyes not distinctly projecting beyond anterior pronotal angles, more or less subquadrate pronotum, claspers are usually not widely separated, aedeagus lacking conjunctival claws Parnisini Distant, 1905c .................................… 35 - Eyes distinctly projecting beyond anterior pronotal angles, rectangular prothorax, claspers are usually widely separated, aedeagus with conjuntival claws Lamotialnini Boulard, 1976 …................................................. 38 35 Small, greenish species (body length about 16 mm , wingspan about 44 mm ) with an elongated, finger-like extension on anterior postclypeus, fore wings with seven apical cells, and fuscous marking in the distal fore wing apical cells and wing margin Rhinopsalta Melichar, 1908 …............................................... Rhinopsalta sicardi Melichar, 1908 - Species without elongated, finger-like extension on anterior postclypeus, fore wings with more than seven apical cells lacking fuscous distal markings …............................................................................ 36 36 Small, greenish species (body length about 16 mm , wingspan about 42 mm ) with an extended anterior postclypeus, antennae prominent, head about as long as pronotum, posterior pronotal lateral margins angled posteriorly, fore wings with eight apical cells, male abdomen globose, male opercula short, apices of tibiae and tarsi red Jafuna Distant, 1912b ….......................................................................................... Jafuna melichari Distant, 1912b - Not as above …..................................................................................... 37 37 Body length about 23 mm , fore wing length about 24.5 mm , dark brown species with dense silver pile on the dorsolateral abdomen, no infuscation on the distal fore wing veins between apical cells 1–3, large, domed abdominal sternite I, large, domed, curving opercula Crassisternalna Boulard, 1980a …......................... Crassisternalna pauliani Boulard, 1980a - Body length about 12 mm , fore wing length about 17 mm , significant piceous pronotal markings, hyaline wings, carinate abdominal tergites..............................................… Timbaltransversa pronotosigillata n. gen. , n. sp. 38 Fore wing radial crossvein distinctly angled to radius anterior 2 and radius posterior veins Trismarcha Karsch, 1891 …... 39 - Fore wing radial crossvein nearly perpendicular to radius anterior 2 and radius posterior veins Abroma Stål, 1866 .....… 41 39 Wingspan greater than 50 mm …............................................ Trismarcha sikorae ( Distant, 1905d ) - Wingspan less than 50 mm ….......................................................................... 40 40 Wingspan greater than 40 mm , eyes wider than lateral angles of pronotal collar …...... Trismarcha voeltzlowi Jacobi, 1917 - Wingspan less than 40 mm , eyes not as wide as lateral angles of pronotal collar …............... Trismarcha parva n. sp. 41 Wingspan greater than 55 mm , contrasting green costal margin..................... … Abroma guerinii ( Signoret, 1860 ) - Wingspan less than 55 mm ….......................................................................... 42 42 Wingspan greater 50 mm …................................................. Abroma antandroyae Boulard, 2008 - Wingspan less than 50 mm ….......................................................................... 43 43 Wingspan greater than 40 mm ….............................................. Abroma inaudibilis Boulard, 1999 - Wingspan less than 40 mm ….......................................................................... 44 44 Body length 15.5 mm , wingspan 35–40 mm , abdomen covered with long golden pile, long silvery pile posterior to timbal cavity, posterior dorsolateral tergites 6 and 7 and all of tergite 8 …........................... Abroma minor Jacobi, 1917 - Body length less than 15 mm , wingspan less than 35 mm ….................................................. 45 45 Timbal with 12 ribs, male operculum covering anteromedial sternite II, head primarily piceous, lateral pronotum piceous, claspers terminating posteriorly with recurved point …................................. Abroma bezamahafalyensis n. sp. - Timbal with 10 ribs, male operculum not reaching anteromedial sternite II, head with transverse piceous fascia, lateral pronotum castaneous, claspers with curved posterior terminus …................................... Abroma falcata n. sp. 46 Body leaf-like, fore wings reticulated, widely separated fore wing costal and radius & subcostal veins, mesonotum with triangular posterior extension of scutellum, proportionately wide pronotal collar, primary femoral spine adpressed to femur, opercula not covering tympanal cavity, tapering posterior abdominal sides, undeveloped pygofer distal shoulder, upper pygofer lobes present, uncus widening towards apex … Hovanini Sanborn, Marshall & Moulds, 2020 …. Hovana distanti ( Brancsik, 1893 ) - Body not leaf-like, fore wings not reticulated, fore wing costal and radius & subcostal veins not widely separated, mesonotum lacking triangular posterior extension of scutellum, pronotal collar of normal width, primary femoral spine elevated above femur, opercula covering tympanal cavity, parallel abdominal sides, development of the pygofer distal shoulder, lack of upper pygofer lobes, uncus not widening towards apex Malagasiini Marshall & Moulds, 2018 .......................... …47 47 Green species (may fade to ochraceous) with a head and pronotum forming a triangle, postclypeus inflated, head longer than pronotum, fore wing basal cell about six times longer than wide, hind wings with five apical cells, small triangular opercula Cephalalna Boulard, 2006a ….......................................................................... 48 - Body not as above …................................................................................. 49 48 Small species (male body length about 15.5 mm , female body length about 17.75 mm , wingspan 33-40 mm ), fore wing basal cell length about six times width, ventral postclypeus with obtuse angle in lateral view, known from northern Madagascar …................................................................. Cephalalna francimontanum Boulard, 2006a - Larger species (male body length about 19.4 mm , female body length about 18.5 mm , wingspan 51-57 mm ), fore wing basal cell length about five times width, ventral postclypeus smoothly curved in lateral view, known from southern Madagascar ............................................................................ … Cephalalna tulearensis n. sp. 49 Opercula absent Anopercalna Boulard, 2008 …............................................................ 50 - Opercula present ….................................................................................. 54 50 Large green species, body length greater than 38 mm , wingspan greater than 75 mm … Anopercalna vadoni ( Boulard, 1980b ) - Smaller species, variably colored, body length less than 38 mm , wingspan less than 75 mm …...................... 51 51 Small green species with transverse piceous fascia on head, body length 17–18 mm , wingspan 54–60 mm …................................................................................. Anopercalna virescens ( Distant, 1905d ) - Not as above …..................................................................................... 52 52 Small greenish-brown species with red on the dorsal head and costal margin, body length 17.5 mm , wingspan 51 mm ............................................................................ … Anopercalna viettii Boulard, 2008 , - Not as above …..................................................................................... 53 53 Tawny species marked with piceous, incomplete transverse fascia on head, median ocellus not surrounded by piceous, mesothorax with significant marking, body length 16 mm , wingspan 44–56 mm … Anopercalna distanti ( Karsch, 1890 ) - Brownish-yellow species marked with piceous spots and fascia, complete transverse fascia on head, median ocellus surrounded by piceous, mesothorax lacks significant marking, body length 15 mm , wingspan 45–47 mm ................................................................................................ Anopercalna distanti ( Karsch, 1890 ) 54 Small (body length 15–17 mm , wingspan 42–53 mm ) olivaceous green species with head and thorax striped with piceous, posterolateral operculum straight, angled mediad, medial operculum smoothly curved, distal shoulder of pygofer a finger-like extension Ligymolpa Karsch, 1890 …........................................ Ligymolpa madegassa Karsch, 1890 - Not as above …..................................................................................... 55 55 Eyes globular, angled anteriorly, infuscation on all fore wing crossveins and distal veins between apical cells Viettealna Boulard, 1980a …........................................................... Viettelana griveaudi Boulard, 1980a - Not as above.....................................................................................… 56 56 Body length about 21 mm , fore wing length about 28 mm , a black and green species with significant pile, head and prothorax approximately triangular, infuscation on the distal fore wing veins between apical cells 1–3, abdominal sternite I of normal appearance, short opercula not covering tympanal cavity Malgotilia Boulard, 1980b ….... Malgotilia sogai Boulard, 1980b - Not as above.....................................................................................… 57 57 Postclypeus extended anteriorly beyond supra-antennal plates, fore wing medial cell bent …........................................................................................... Deremeces andringitraensis n. gen. , n. sp. - Not as above …..................................................................................... 58 58 Anterior head margin truncated not extending beyond anterior supra-antennal plates, male abdomen tapering posteriorly, and small cruciform elevation Malgachialna Boulard, 1980b ….................................................. 59 - Anterior head margin extending anteriorly beyond supra-antennal plates, male abdomen cylindrical, cruciform elevation of normal size Malagasia Distant, 1882 .................................................................... 60 59 Small, body length about 15 mm , wingspan about 45.5 mm , tawny species with red fore wing costal margin........................................................................................ Malgachialna rubracosta n. sp. - Body length about 24.5 mm , wingspan about 65 mm , tawny with reddish head and hind wing venation..................................................................................... Malgachialna noualhieri Boulard, 1980b 60 Small, body length about 20 mm , wingspan about 53 mm , green species with red ankles.... Malagasia rubrastragala n. sp. - Not as above....................................................................................... 61 61 Small, body length about 15 mm , wingspan about 45.5 mm , tawny species with red fore wing costal margin........................................................................................ Malagasia italivianaensis n. sp. - Not as above....................................................................................... 62 62 Large species, body length about 40–42 mm , wingspan about 98–108 mm , male abdomen inflated................................................................................................ Malagasia inflata Distant, 1882 - Body length less than 38 mm , wingspan less than 90 mm .................................................... 63 63 Large tawny and piceous species, wingspan about 84 mm ................................. Malagasia argentea n. sp. - Wingspan less than 75 mm ............................................................................. 64 64 Wingspan greater than 68 mm , posterior and medial opercular margins forming an approximate right angle, male abdomen not inflated, lateral male abdomen parallel......................................... Malagasia aperta ( Signoret, 1860 ) - Wingspan less than 68 mm , posteromedial margin curved, male abdomen inflated, lateral male abdomen expanding to segment 4....................................................................... Malagasia mariae Boulard, 1980b

Description and Life History Cicadas ( Hemiptera : Cicadidae ) commonly infest coffee plantations in Brazil, and at least 10 species have been recorded. The most important species are Quesada gigas (Olivier) , Quesada sodalis (Walker) , Fidicina mannifera (Fabricius) , Fidicina pullata Berg , and Carineta fasciculata (Germar) . These species are pests in the states of Minas Gerais, Sao Paulo, and Parana. Besides being the most important of this group, Q. gigas is the largest in size and also a pest of many other plants. Q. gigas has a wide range of geographical distribution in South America, and it is the best studied species of this group in the South American continent. For most winged cicadas, membranous wings are held rooflike over the body. All have three ocelli and the antennae arise between, rather than beneath, the eyes. Cicada songs sound buzzy, raspy, or whiney, because their carrier frequencies are less pure and are higher than those of crickets. Cicadas call almost exclusively during daylight hours and at dusk, usually from trees and shrubs. On uncultivated land, the nymphal stage feeds on the roots of indigenous trees. When the bush is cleared and coffee is planted, the larvae move to the coffee roots and feed, causing damage. Nymphs of Q. gigas have been observed up to 50 cm below the ground on the roots. The nymphal stage goes through five instars. They suck the roots and form cavities in the soil around them, creating openings for infections by disease-causing organisms. The lifespan of the nymphal stage of these cicadas varies from 4 up to 17 years depending on the species, but the adult stage lasts 1-3 months. Adults are good fliers and rest on the tops of trees. The female adults mate upon emergence from the soil, attracted by the song of the males, and then lay eggs on the bark of trees. The nymph emerges and drops to the soil, where it searches for the coffee roots to feed upon and complete its development. Damage Cicadas are characterized by piercing-sucking mouthparts, and they feed on the sap of the plant root. Cicada nymphs damage the plants, causing chlorosis of the leaves in the apexes of branches, loss of leaves, low production, and death of plants under heavy infestations. Population Management Because of the cryptic (hidden) habits of nymphs feeding on the roots of coffee plants, insecticidal treatments are only effective when adults are emerging and have not laid eggs. Monitoring the adult flights should help to program the control activities. An alternative to be considered for nymphal control is the use of entomopathogenic nematodes.