Table Of ContentPROC. ENTOMOL. SOC. WASH.
97(1), 1995, pp. 1-16
TWO NEW NEOTROPICAL TREEHOPPER GENERA AND INVESTIGATION OF
THE PHYLOGENY OF THE SUBFAMILY MEMBRACINAE
(HOMOPTERA: MEMBRACIDAE)
Christopher H. Dietrich and Stuart H. McKamey
(CHD) Systematic Entomology Laboratory, USDA-ARS, % National Museum ofNat-
ural History, Washington, D.C. 20560*; (SHM) Department ofEcology & Evolutionary
Biology, U-43, University ofConnecticut, Storrs, Connecticut 06269.
Abstract.—Tv/o new genera are described in the membracid tribe Membracini: Havi-
landia, new genus, including H. pruinosa (Haviland), new combination, and H. hypso-
proroides, new species; and Lewdeitzia, new genus, including L. lunata, new species.
Havilandia is morphologically and behaviorally similarto Talipedini and some Leioscyta
Fowler (Membracini), but may be the sister-group ofHypsoprorini. Lewdeitzia is appar-
ently dimorphic fora feature ofthe forewingvenation usedto distinguishErechtia Walker
from Leioscyta. Preliminary cladistic analyses of27 taxa in the subfamily Membracinae
place the new genera among others in the tribe Membracini, but suggest that this tribe is
paraphyletic, havinggivenrisetoothertribesinthesubfamily. Theanalysesfurthersuggest
thatthetribeTalipedini isparaphyleticandthegeneraLeioscytaandMembracisFabricius
are para- orpolyphyletic. Reclassification ofthe Membracinae is needed, but shouldawait
elucidation ofthe limits ofsome genera ofMembracini by more extensive cladistic anal-
yses. A key to the tribes of Membracinae and the genera of Membracini is presented.
Nomenclatural changes include restoration ofthe original spelling ofParagara Coding,
1926, andofthecombinationP. tholoidea(fromParagargara)andtwonewcombinations,
Paragara nigra (Funkhouser) (from Paragargara) and Enchenopa beebei(Haviland) (from
Leioscyta). Two species most recently treated as Membracis are here considered Mem-
bracini incertae sedis.
Key Words: Cladistics, evolution, morphology, parsimony, paraphyly
The subfamily Membracinae constitutes some tropical genera (Loye 1992, Wood
a major component oftreehopper diversity 1993a) and may be explained partly by the
in the New World and is especially diverse habit in many species, but in Membracinae
in the tropics. The North American En- only, ofdepositing egg masses on the host
chenopa binotata species complex, whose plantsurface, therebyobviatingmanyofthe
females insert eggs into host plant tissue, plant's first-line structuraland chemical de-
hasattractedattentionasanexampleofhost- fenses against herbivore colonization,
mediated sympatric speciation (Wood Cladisticanalysesofthefamily-grouptaxa
1993b). Polyphagyhasbeenreportedwithin ofMembracidae (Dietrich and Deitz 1993;
Dietrich, Deitz, and McKamey, unpubl.)
pa*'rCt^omrernetsopfoMnidcer.nocbei:olorCg^huyn,stC,ooplhoureardouH.SrtaD^tiee.UtrnuiicvThe,r^si.Dtey-, ^sUutp*^,p*^ourntti.lthncowm,outohpe*rh^ey'l^hyasobfeMeenmbnroaecxipnlai.cei,.t
Ft. Collins, CO 80523. cladistic analysis ofrelationships among its
.
PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
five tribes(Deitz and Dietrich 1993): Acon- concepts of tribes and genera. Species of
ophorini, Hoplophorionini, Hypsoprorini, Havilandia, new genus, share morphologi-
Talipedini, and Membracini. Nonetheless, cal features with species of Hypsoprorini,
three are supported by apparent synapo- Hoplophorionini, Talipedini, and Acono-
morphies. The Aconophorini and Hoplo- phorini. Lewdeitzia lunata, new genus, new
phorionini are well-supported monophylet- species, may be sexually dimorphic for the
ic groups (Dietrich and Deitz 1991; forewing venation feature distinguishing
McKamey and Deitz, unpubl.)- The species Leioscyta Fowler from Erechtia Walker
ofHypsoprorini arealso united byapparent (tribe Membracini) and the female has a
synapomorphies that are unique among the broad, obtusely rounded, marginally com-
Membracinae: pronotum with lateral mar- pressed, anterior pronotal horn similar to
gins ofposterior process overlapping fore- that ofAconophorini and some Hoplopho-
wing at rest, and clavus of forewing acute rionini. The features ofthese new taxa ac-
and extensively associated with apical lim- centuateproblemswiththegenericandtrib-
bus. No synapomorphies have been found alclassificationoftheMembracinae.Tohelp
to support the monophyly ofTalipedini or place the new taxa, we analyzed the rela-
Membracini. tionships among the genera ofMembracini
The tribe Talipedini was erected (Deitz and representatives ofothertribes ofMem-
1975) to draw attention to the position of bracinae.
Trinarea (as Talipes) appendiculata (Fon- Giventheneedoffurthertaxonomicwork
seca) as morphologically intermediate be- revealed by the cladistic analysis, the nu-
tween the tribes Membracini and Hoplo- merous modifications to Membracini since
phorionini. Talipedini and Hoplophorionini thelastkey(Funkhouser 1951), andthedis-
both haveclavatehindtibiaeand small hind covery ofsome previously unknown or ne-
tarsi, a unique synapomorphy, but features glected intrageneric variation, we present a
used by Deitz (1975) to distinguish Tali- key to tribes ofMembracinae andgenera of
pedini from Hoplophorionini arepresent in Membracini to clarify current taxonomic
Membracini, suggestingthat Talipedini may concepts in the subfamily.
beparaphyletic. McKameyand Deitz(1991) Key TO THE Membracinae
transferred two more species into Talipe-
dini, but did not list additional diagnostic 1 ForewingveinRinitiallydividedintoR,+,+3
features for the tribe. andRj15orvenationreticulate,orwithboth
The most diverse tribe in the subfamily conditions 2
is Membracini, which, now with 17 genera 1'. RFosr(e=wRi:ng+,v+e4i+ns)R(Fiingitsi.al1l,y2d)i,vviedneadtiinotnonRo,tarned-
and nearly 200 described species, is twice ticulate 4
as rich in genera and species as the next 2. Metathoraciclegswithtibiaeclavateandwith
largest tribe, Hoplophorionini. Among the tarsi distinctly shorterthan anteriortarsi . .
Hoplophorionini'
dMieamgbnroasctiicnife,atnuorneeslairseteudnbiyquDeeittozt(h1a9t75t)rifbeo.r T. Metathoraciclegswithtibiaenotclavateand
Thus, Membracini (sensu Deitz 1975, ex- 3. wPirtohntoatrsuimaswiltohnglaastearnaltermiaorrgtianrssio(fFigp.os2tce)rio.r. 3
panded by McKamey 1992 and Sakakibara processoverlappingportionsofapicallimbus
1992), although defined by a unique com- andveinsofforewinginrepose;forewingcla-
bination offeatures, also may be paraphy- vusacuminate Hypsoprorini-
letic.
toTMweombnreawcitnaixa(wDeerietzdi1s9c7o5v)e,rebdutthwaetrkeeyneodt ' Key togenera in prep, by McKameyand Deitz.
- See Funkhouser's (1951) key to genera of"Noto-
assignable to any known genus and have cerini,"plusJibaritaRamosandHypsoprorachisFon-
combinations offeatures traversing current seca and Diringshofen.
. . .
VOLUME NUMBER
97, 1
3'. Pronotumevenlytaperedposterolaterally,at lateral carinae and a ventral lobe produced
mostconcealingpartofapicallimbusoffore- in front ofthe vertex ofthe head
wing in repose; forewingclavus obliqueapi- W PhyllotropisStil
cally Aconophorini' Pronotal metopidium without dorsolateral
4. Metathoraciclegswithtibiaeclavateandwith carinaeor without ventral lobe, orwith nei-
tarsidistinctlyshorterthananteriortarsi;ab- ther feature 12
domen with middorsal tuberosities 12. Pronotal dorsum strongly foliaceous, sides
Talipedini (monotypic: Trinarea Coding) extensively compressed into a single plate-
4'. Metathoraciclegswithtibiaenotclavate(Fig. like median carina Folicarina Sakakibara''
2c)or,ifclavate,thenabdomenwithoutmid- 12' Pronotaldorsumnotsostronglycompressed,
dorsaltuberosities;metathoracictarsiatleast sidesnotextensively fused
as longas anteriortarsi (Membracini) 5 . . Memhracis(in part;e.g. M. carinulataRichter)
5 Metathoracictibiaeclavate,withdorsaledges 13. Pronotumfoliaceousandsmooth(pitssmall,
flattened,and withcucullatesetaenumerous shallow,andindistinctabovehumeriatleast);
andsmall in row I, small orabsentin row II forewing surface coriaceous throughout (ar-
(Fig. Ij) Havilandia, n. gen. cuatechaetoids conspicuous)
5'. Metathoracic tibiae with dorsal edges not Membracis(in part; e.g. M.foliata [L.])
flattened, with largecucullatesetae in rows I 13' Pronotum depressed, at least posterolateral-
and II (Fig. 2c) 6 ly, and more coarsely punctate (pits larger,
6. Pronotal metopidium with 1 to several deeper, and distinct above humeri); distal
obliquecarinaeoneachsideandwithpairof forewing surface more glossy (arcuate chae-
dorsolateral carinae or dorsum strongly bi- toids absent or inconspicuous) 14
sinuate posteriorly, or with all three condi- 14. Anteromedial pronotal horn present (except
tions 7 in some $ Tritwpidia, which are indistin-
6'. Pronotal metopidium without oblique cari- guishable from Leioscyta) 15
nae, with or without dorsolateral carinae, 14' Anteromedial pronotal horn absent 16
dorsumsublinearoronce-sinuateposteriorly 15. Pronotal horn narrowin lateralaspect; clyp-
13 eus distally narrowed (as in Fig. Ih)
7. Pronotallateralcarinaeextendingtopostero- Tritwpidia Stdl
lateral margins or nearly so; pronotal integ- 15'. Pronotalhornbroadinlateralaspect;clypeus
ument coarsely punctate (pits distinct) and distally truncate (Fig. 2g) ...Lewdeitzia, n. gen.
thick 8 16. Forewingwith 2 or more r-m crossveins . . 17
7'. Pronotallateralcarinaeterminatingbeforeor 16'. Forewingwith only 1 r-m crossvein 18
above humeral angles; pronotal integument 17. Pronotum rugosely carinate, steeply decli-
sometimes smooth (pits indistinct), mem- vous posteriorly, humeri without transverse
brane-like 9 carinae BolbonotodesFowler
8. Pronotumbisinuate,decliningstepwisefrom 17' Pronotum evenly carinate, gradually decli-
above humeral angle to posteriorapex .... vous posteriorly, humeri often with trans-
Tylopelta Fowler versecarinae Erechtia
8'. Pronotum declining evenly from above hu- Pronotal dorsum with irregular gibbosities
meralangleto posteriorapex throughout, transverselycarinate orgibbous
. .. CampylenchiaStAl, EnchenopaAmy. & Serv. subapically,and withoutlongstraightlateral
9. Pronotum and its horn depressed, pronotal carinae; mesothoracic tibiae foliaceous ....
integument thickandcoarsely punctate(pits Bolbonota Amy. & Serv.
large, deep, and distinct above humeri) . . Pronotal dorsum without irregular gibbosi-
Kronides Kirkaldy ties, usually with 1 pair sublinear lateral ca-
9'. Pronotumanditshorn,ifpresent,foliaceous, rinaeextendingposteriorlyoverdorsum;me
thin, and smooth (pits small, shallow, and sothoracic tibiae cylindrical to foliaceous . 19
indistinct above humeri at least) 10 19. Pronotal contour in lateral aspect strongly
10. Pronotal horn present and distinct sinuate Paragara
Enchophyllum Amy. & Serv.
10'. Pronotal horn absent or indistinct 11
11 Pronotal metopidium with a pair ofdorso- ^ Newdistribution record forFolicarina nr. bicolor.
Trinidad, Arima Valley, St. Andrew's Trace, 16-24
June 1981, leg. S.H. McKamey.
' See Dietrich and Deitz (1991) for keys to genera 5 ParagaraCoding, 1926a; originalspellingherere-
and species. stored from Paragargara. Typespecies: Paragaratho-
5
PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
19'. Pronotal contour in lateral aspect linear or tex; frontoclypeus flat, diamond shaped,
nearly so 20 margins weakly produced, cibarial muscle
20. HindwingMwithoutr-mandm-cucrossveins, scarsparalleltoventrolateral margins; fron-
with vein free Euniisa Fonseca
20'. Hindwingwith 1 r-m and 1 m-cucrossvein toclypeus, clypellus, and rostrum, in lateral
Leioscyta view (Figs, la, i) forming continuous arc,
Descriptions of New Taxa rostrum extended to base of abdomen; in
anterior view with distance between lateral
Morphologicalterms,techniques,andthe margins of eyes slightly less than distance
convention for quoting labels follow Deitz between pronotal humeri. Thorax: Pro-
(1975), Dietrich (1989), and Dietrich and notum(Figs, la,h-i)elongateandwithmid-
Deitz (1991), except forewing crossvein s line strongly carinate and with 1 dorsolat-
(that connects veins R2+3 and R4+5) equals eral pair of strong carinae diverging from
"r"ofDeitz(1975). Specimensforthisstudy apex, extending to posterolateral margin of
were provided by The Natural History Mu- pronotum. Forewing (Figs, la, i) punctate
seum, London (BMNH); North Carolina in basal halfbetween veins C and M and in
State University, Raleigh (NCSU); the per- anal area, vein R initially divided into R,
sonalcollectionofS. H. McKamey(SHMC); andRs,crossveinssandr-m, absent, 2 m-cu
and the United States National Museum of crossveins present, apical limbus relatively
Natural History, Washington (USNM). narrow, contiguous with clavus for short
Character states for taxa other than those distance only. Hind wing with crossveins
describedbelowarebased on materialiden- r-m and m-cu present. Legs: Pro- and me-
tified and labeled in the NCSU and USNM sothoracic tibiae (Fig. li) compressed,
collections. translucent,marginsofprothoracictibianot
Tribe Membracini Rafinesque expanded, mesothoracic tibia with poste-
Havilandia, New Genus rior margin slightly expanded; mesothorac-
(Figs, la-k) ic coxa without acuteprocess; metathoracic
femur with pair ofdorsoapical cucullate se-
Type species: Tropidoscyta pruinosa
Haviland. tae; metathoracic tibia (Figs, lb,j) clavate,
Diagnosis.—Metathoracic tibia clavate posteriormargin compressed, rowI with 1
or more small cucullate setae, row II with
and compressed along dorsal edge, with se-
tal row I bearing numerous small cucullate 5 or fewer, row III absent; all tarsi subequal
in length; metathoracic tarsus relatively
setae and row II with few ornone; forewing
with cell membranes clothed by erect mac- slender, tarsomere I with small apical cu-
cullateseta.Abdomen: SternumIIIwithout
rotrichia, without crossvein s.
median tubercle, transverse carina indis-
Description.—Z/^'^afif." Vertex (Fig. Ih)
impressed mesad of ocelli, ventrolateral tinctorabsent; tergawithouttuberositiesor
margins weakly produced; ocelli approxi- fenestrae. Male: Pygofer(Fig. 1d)withweak
mately 2X as far from each other as from vertical lateral carina; lateral plate free, un-
mesal margins ofeyes and approximately 1 daerrm,edta;paeeridnegagaupsic(aFlilgys,.a1nft-egr)iwoirtfhacsehaoftfsalpeenx-
ocellardiameterfrom dorsal margin ofver-
evenlydenticulate; gonopore membrane, in
posteriorview,occupyingapicalhalfofshaft;
loidea Goding, 1926a, by original designation. Re- shank of style (Fig. le) slightly expanded
storedcombination: ParagaratholoideaGoding. New preapically, shank sparsely setose, apex re-
combination:Paragaranigra(Funkhouser 1940).Par- curved, acute, in posterior view oriented
agargara(Goding 1926b)wasan incorrectsubsequent
dorsolaterally; subgenital plate with preap-
spellingand has no availability. Paragargara Goding,
1929, was an unjustifiedemendationand is an objec- ical dorsal lobe. Female: Posterior margin
tivejunior synonym oiParagara. ofsternum VII with arcuate emargination;
VOLUME NUMBER
97, 1
Fig. 1. Havilandia, newgenus, a-g, H. pruinosa (Haviland). a-c, holotype9: a, habitus, lateral view; b, left
metathoracic tibia and tarsus, lateral view; c, second valvulae, lateral aspect; d-g, #7-90-337g$\ d, terminalia,
lateral view (genitalia removed); e, shank ofleft style, ventrolateral view; f-g, aedeagus, posterior and lateral
views, respectively, h-k, H. hypsoproroides, new species, holotype 9: h, head and pronotum, anterior view; i,
habitus, lateral view; j, left metathoracic tibia and tarsus, lateral view; k, second valvulae, lateral view, avc,
anteroventral carina; die, dorsolateral carina; dsc, dorsal submedial carina; p, pygofer; mc, median carina; Ip,
lateral plate; sp, subgenital plate; tc, transverse sternal carina.
PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
2nd valvulae (Figs. Ic, k) in lateral view Materialexamined.—Holotype2[BMNH].
graduallyexpanded toward midlength, then Otherspecimens: Nestor,Trinidad; San Mar-
taperingslightlytowardapex, dorsal margin tin and Shapajilla, Peru; and Culebra, Ama-
with arcuate emargination preapically, zonas, Venezuela [all USNM]; road to Foco
without distinct crenulae or teeth. Integu- Mine, 33 km SE El Dorado, Bolivar, Vene-
mental vestiture: Vertex and pronotum zuela [SHMC]. Datesofcollection: February,
evenly punctate; forewing membrane with April,May,August.Ateneral$specimenfrom
semi-erect pale setae. Abdominal terga Maroni, French Guiana [Figs. Id-g; Dietrich
coarselypunctate, most pitsassociatedwith Research #7-90-337g, NCSU] is probably
lateral seta; acanthae simple to dentate. conspecific with the holotype ofH. pruinosa
Notes and etymology.—Short-horned but lacks a pale macula on the forewing at
specimens ofthis genus resemble Leioscyta the apex of vein Cu that is present in the
Fowler, but are distinguishable by the pres- holotype (Fig. la).
—
ence ofa pair ofanteroventral longitudinal Notes. Females of H. pruinosa guard
carinae on the pronotum and the non-foli- their eggs and are not ant-attended (Havi-
aceousfronttibiae. Long-homed specimens land 1925), characteristics shared with Ho-
resemble Hypsoprora StSl (Hypsoprorini), plophorionini and some Aconophorini and
but lackapicolateral expansions on the pos- Membracini (Haviland 1925, Wood 1984,
terior pronotal process. Although Havilan- Dietrich and Deitz 1991). H. pruinosa fe-
dia best fits Deitz's (1975) concept of the males also deposit a pale waxlike material
tribeMembracini,certainfeaturesofthenew in spirals along the twig and leaf edges in
genus resemble those of other tribes. Like the vicinity of their egg masses (Haviland
species ofHoplophorionini and Talipedini, 1925; McKamey, unpubl.). Similardeposits
Havilandia has clavate hind tibiae with a areproducedbyLeioscytaspiralisHaviland
reduced number ofcuculate setae in row IL (Haviland 1925, Strumpel 1986) (tribe
Like some species ofHypsoprorini, the new Membracini), an additional (unidentified)
genus has a ventrolateral pair ofcarinae on Leioscyta species (McKamey, unpubl.),
the pronotal horn. Ochropeplatriangulum(Germar)(tribe Ho-
Genericrecognition isbasedprimarilyon plophorionini; Wood 1984 [as O. pallens
the tibial and abdominal features, but both St^l]), andAconophora mexicana St^l (tribe
species also have: vertex densely clothed Aconophorini; Wood 1984, Dietrich and
with pale setae; thoracic pleuron pilose, se- Deitz 1991). Nymphs of//, pruinosa, L.
tae clothed with white waxlike material; spiralis, and A. mexicana are also clothed
pronotum produced anterodorsally, with 1 with irregular waxlike exudates that facili-
ventrolateral pair ofstrong carinae diverg- tate crypsis among the spiral deposits (as in
ing from the apex; and forewings densely Strumpel 1986: Fig. 2). Many other species
clothed with arcuate chaetoids. of Membracinae have nymphs with white
This genus is named in honor of Maud waxlikeexudatesbutlackwaxydepositsad-
D. Haviland, a pioneer in the ecology and jacenttotheireggmasses. Suchfeatureshave
taxonomy ofNeotropical Membracidae. not been reported for species ofsubfamilies
other than Membracinae.
Havilandiapruinosa (Haviland),
New Combination Havilandia hypsoproroides,
(Figs, la-g) New Species
Synonymy: Tropidoscytapruinosa Hav- (Figs. Ih-k)
iland, 1925:237. Type locality: Fonteboa, Amazonas,
Typelocality: Kartabo, Cuyuni District, Brazil [BMNH].
Guyana [BMNH]. Description.—Head, pronotum, and ab-
VOLUME NUMBER
97, 1
domen reddish brown, pronolal horn darker compressed horn; forewing with or 2 r-m
1
thanrestofpronotum, legsyellow; headwith crossveins, with vein R initiallydivided into
vertex densely clothed with pale setae; tho- Ri and Rs.
—
racicpleuron pilose, setaeclothed withwhite Description. Head: Vertex(Fig. 2g)not
waxlike material; forewing membrane uni- impressed mesad of ocelli, ventrolateral
formly smoky hyaline. Pronotum (Fig. li) margins foliaceous and strongly produced;
produced into an elongate, slender antero- ocelli approximately 2x as far from each
medial horn, apex in lateral view expanded other as from mesal margins of eyes and
and obliquelyrounded, with 1 ventrolateral approximately 2 ocellar diameters from
pair ofstrong carinae diverging from apex; dorsal margin ofvertex; frontoclypeus flat,
posterior process approximately even with lateral comers rounded and apex truncate,
crossvein r-m of forewing at rest, apex margins foliaceous; rostrum extended to
slightly elevated. Forewing densely clothed base ofabdomen; frontoclypeus, in lateral
with arcuate chaetoids. Metathoracic tibia view, forming shelfabove clypellus; in an-
(Fig. Ij) with setal row I bearing 15-18 cu- terior view (Fig. 2g) with distance between
cullate setae. Dimensions (mm): Body lateral margins ofeyes slightly less than dis-
length (head toapex offorewingat rest) 5.1, tance between humeri. Thorax: Pronotum
width across humeri 1.7; vertex height 0.7, with broad, marginally compressed anter-
width 1.0;pronotumlength 5.3;headtohorn odorsal process, much larger in female (Fig.
apex 2.1; forewing length 4.5; prothoracic 2a) than in male (Fig. 2e), bearing pair of
tibia length 1.0; metathoracic tibia length lateral carinae extending ventrolaterally
1.5; ovipositor length 1.0. from apex but not attaining posterolateral
—
Material examined. Holotype 9 labeled: margins. Forewing (Figs. 2a, e) punctate in
M
"Amazon./ Fonteboa; Riksmuseum/Stock- basal half between veins C and and in
holm; Dietrich Res./ 7-90-33a9; HOLO- base of anal area, vein R initially divided
TYPE/ Havilandia/ hypsoproroides/ Die- into R, and Rs, crossveins s and r-m2 pres-
trich and McKamey" [BMNH]. ent, with crossvein r-m, present (Fig. 2a) or
Etymology.—The trivial name was se- absent (Fig. 2e), 2 m-cu crossveins present,
M
lected to indicate the similarity ofthis spe- m-cu, perpendicularto veins and Cu, 1st
M
cies to members of the genus Hypsopwra and 2nd cells subequal in length, apical
Stdl. limbus relatively wide, contiguous with cla-
vus for short distance only. Hind wing
Key to Species of Havilandia
crossveinsr-mandm-cupresent.Legs: Pro-
1. Anteriorpronotalprocess,inlateralview,short, and mesothoracic tibiae (Figs. 2h, i) com-
broad, and tapered, apex rounded (Fig. la) ... pressedand foliaceous; mesothoracic femur
pruinosa(Haviland) with anteroapical cucullate seta; metatho-
r. Anterior pronotal process, in lateral view,
elongate and slender, apex obliquely truncate racictibia(Fig. 2c)straight,notcompressed,
(Fig. li) hypsoproroides, n. sp. row I with 7-8 enlarged cucullate setae, row
II with 4-5 enlarged cucullate setae, row III
Lewdeitzia, New Genus withoutcucullate setae; metathoracic tarsus
(Figs. 2a-m) 4sliagphitclyallocnugceurlltahtaen osetthaeer.s,Atbadrosmomeenr:e ISwtietrh-
Type species: Lewdeitzia lunata, new num III without median tubercle, sternum
species. IV with distinct transverse carina; terga
Diagnosis.—Anteriorregion ofpronotum without tuberosities or fenestrae. Male:
in lateral viewelevated into broad, rounded Pygofer with vertical lateral carinae; lateral
processwith 1 carinaoneach side, in female plate free, with prominent rounded vertical
further developed into large, marginally ridge over entire height (Fig. 2j); aedeagus
PROCEEDINGSOFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON
Fig. 2. Lewdeitzialunata, newgenus, newspecies, a-d, holotype9: a, head, pronotum,and forewing,lateral
view(texture);b, headandpronotum,dorsalview(color);c, reversedrightmetathoracictibiaandtarsus,lateral
view, same scale as 2h-i; d, second valvulae, lateral view; e-m, <5; e, head, pronotum, and forewing, reversed
lateral view(texture); f-g, headand pronotum,dorsal (color)andanteriorviews, respectively; h-i, left pro-and
mesothoracictibiaeandtarsi,lateralviews;j,leftlateralplate,dorsalview(ant,anterior);k,leftstyle,ventrolateral
view; 1-m, aedeagus, anteriorand lateral views, respectively.
(Figs. 21, m) with shaft slender, in anterior ulate. Integumental vestiture: Vertex and
view subparallel, in lateral view tapering pronotum coarsely punctate, sparsely
apically, anterior face of apex evenly den- clothed with pale recumbent setae; thoracic
ticulate laterally; gonopore membrane, in pleuron sparsely setose, setae not clothed
posterior view, occupying nearly entire with waxlike material; forewing membrane
width ofshaft; shank ofstyle (Fig. 2k) with glabrous. Abdominal terga coarsely punc-
numerous long setae, apex recurved and tate, pits without lateral setae; acanthae
acute; subgenital plate evenly tapering api- dentate.
cally, without lobes. Female: Posterior Notesandetymology.—Thefemaleofthis
margin ofsternum VII with arcuate emar- genus resembles Aconophora Fairmaire
gination; 2nd valvulae, in lateral view (Fig. (Aconophorini), while the male resembles
2d), subparallel, apical '/? irregularly cren- Leioscyta. The genus is named to honor
1 , .
VOLUME NUMBER
97,
Lewis L. Deitz, whose higher classification mon in other species ofMembracinae (e.g.
and bibUographies have greatly facilitated see Sakakibara 1976 and Striimpel and
research on the Membracoidea. Striimpel 1978), we consider the nearly
identical color pattern and similar size of
Lewdeitzia lunata, New Species both Lewdeitzia specimens sufficient to re-
(Figs. 2a-m) gard them as a single species. The species
Tvpelocalitv: Sao Paulo, Sao PauloState, nameisderivedfrom Latin forthepalecres-
Brazil [NCSU]. cent-shaped band around the base of the
Description.—Ground color brown; female's horn,
pronotal carinae, crescent-shaped band
around posterior base ofhorn (female; Fig. Cladistics
2b)ormottlingbetweencarinaeand humeri Morphological characters
(male; Fig. 2f), 2 transverse bands farther
Head
posterad on dorsum, transverse band on
metopidiumbetweenhumeri, andlegspaler 1 Frontoclypeus: 0, apex in lateral view
orangebrown; forewingsmokyhyalinewith not forming a shelfover clypellus, margins
2 pale transverse bands (1 near base, 1 not expanded laterally (Fig. 1h); 1, apex in
aligned with end ofclavus). Female pron- lateral view not forming a shelf over cly-
otal horn with apically bifid lateral longi- pellus, margins expanded laterally; 2, apex
tudinal gibbosity, apex obtusely rounded, inlateral viewformingashelfoverclypellus
lateral carinae weak, terminating slightly (Sakakibara 1992: Figs. 1, 2, 4).
posterad of humeri; male horn short and
Pronotum
rounded, carinae distinct, marginal com-
pressed area narrower. Other morphology 2. Shape: 0, depressed or rounded (Figs,
as described for genus. la, i); 1, compressed (Deitz 1975: Fig. 15S);
Dimensions (mm).—Body length 9 6.3, 3 2, foliaceous {op. cit.: Fig. 15U).
5.3; maximum width 9 2.8, <5 2.4; pronotal 3. Anteromedial horn: 0, absent orweak
length 9 7.0, $ 4.5; head to horn apex 9 3.3; (Fig. la); 1, well developed (Fig. li). Al-
forewinglength 9 5.5, <3 4.8; prothoracic tib- though entirely enclosed by the foliaceous
ia length 9 1.5, 5 1.2; metathoracic tibia pronotal margins, the horns ofPhyllotropis
length 9 2.2, 6 1.9; ovipositor length 1.9. and Folicahna are nonetheless well indi-
Material examined.—Holotype 9 labeled: cated (Sakakibara 1992: Figs. 3, 4).
"Sao Paulo/ S. Paulo Brazil/ Nov. 1930; 4. Transverse metopidial carinae: 0, ab-
A.Mailer,Coll./Frankjohnson/Donor; 153 sent; 1, restricted to humeri; 2, complete,
9/ S. Paulo/ S. Paulo/ 10.30; HOLOTYPE/ two sides meeting medially.
Lewdeitzia/ lunata/ Dietrich and Mc- 5. Dorsolateral carinae: 0, absent; 1, one
Kamey" [NCSU]. Other material: $ from submedial pair (Figs, la, i); 2, two or more
Corupa, SantaCatarina, Brazil, XI-1929, A. pairs (Deitz 1975: Fig. 15R). The pronotal
Mailer, Coll. (Dietrich Research 15-91-40d carinae ofHoplophorionini converge at the
[NCSU]. base of the metopidium rather than at its
<3)
Notes and etymology.—The holotype apex, as in other Membracinae, and there-
lacks its left metathoracic leg. The male, fore may not be homologous.
which lacks its left hind tarsus and right 6. Posteriorextension ofdorsolateral ca-
hind tarsomeres II and III, was excluded rinae: 0, absent; 1, terminated above hu-
from type statusbecause it may not be con- meri; 2, extended beyond humeri.
specific, given the considerable differences 7. Dorsal submedial carinae: 0, absent;
compared to the female. Nonetheless, be- meetingmedialcarinaanteriorly(Fig. la);
1
cause extreme sexual dimorphism is com- 2, not meeting medial carina anteriorly.
. :
10 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
8. Anteroventral pair of carinae, which 17. Length ofmetathoracic tarsi relative
are contiguous with dorsolateral carinae: 0, to pro- and mesothoracic tarsi: 0, longer; 1,
absent; 1, present (Figs, la, h, i). subequal; 2, distinctly shorter (Deitz 1975:
9. Irregular, oblique metopidial carinae: Fig. 14A).
0, absent; 1, present (Sakakibara 1992: Fig.
Forewing
4).
10. Posterior transverse carina: 0, ab- 18. Vein R: 0, initially divided into R,
sent; 1, present (Haviland 1925: PI. II, Fig. and Rs (Fig. la); 1, initially divided into
6). R,+2+3 and R4+5 (Deitz 1975: Fig. 1 lA).
1 1 Posterior sublateral longitudinal ca- 19. Crossvein s: 0, absent; 1, present
rinae: 0, absent; 1, present(Fonsecaand Di- (Figs. 2a, e).
ringshofen 1969: Fig. 3). Our treatment of 20. Crossvein(s) r-m: 0, one (Figs, la, i);
this feature as a separate character from 1, two or more (Fig. 2a).
character 6 is supported by the presence of 21. Membrane, erect macrotrichia: 0,
both dorsolateral and sublateral carinae in absent; 1, present.
some taxa (e.g. Erechtia). 22. Membrane, arcuate chaetoids: 0, ab-
12. Punctation: 0, coarse—distance be- sent, surface glabrous; 1, present, surface
tween pits less than pit diameter, pits dis- shagreen(Dietrichand Deitz 1993: Fig. 27).
tinct above humeri; 1, fine—distance be- Abdomen
tween pits greater than pit diameter, pits
indistinctabovehumeri. Considerablevari- 23. Sternum III: 0, unarmed; 1, with me-
ation exists among taxa coded as state 0, dial tubercle; 2, with transverse carina (Fig.
but state 1 (e.g. Membracis) is distinctive la).
(compare Figs. 7 and 10 ofWood and Mor- 24. Sternum IV: 0, unarmed; 1, with
ris 1974). transverse carina (Deitz 1975: Fig. 3A).
25. Dorsaltuberositiesorfenestrae,terga
Pleuron and legs V-VIII: 0, absent; 1, paired {op. cit.\ Fig.
13. Pleural wax: 0, absent; 1, present. 3A); 2,unpairedmedial(Dietrich 1989: Fig.
State 1 is considered to represent a homo- 18). The presence, in Trinarea and some
log, although Havilandia has the pleuron Erechtia, of paired tuberosities on the an-
densely pilose with waxy setae while Hyp- terior terga and unpaired medial tuberosi-
soprora has thepleuron sparsely setosewith ties on the posterior terga suggests that one
waxlike deposits on the surface ofthe scler- feature was derived from fusion or separa-
ite. tion ofthe other.
14. Pro- and mesothoracic tibiae: 0, nar-
Second valvulae
row, semicylindrical; 1, subfoliaceous(Figs.
2h, i). 26. Length:0,elongate; 1,shortandbroad
15. Mesothoracic acute coxal process: 0, (Deitz 1975: Figs. 17H-J).
absent; 1, present. State 1 is a synapomor- 27. Dorsal teeth {op. cit.: Figs. 17C, M):
phy ofthe tribe Hoplophorionini. 0, absent; 1, one; 2, two. The serrations
16. Metathoracictibia:0,notclavate,row found in many membracid groups {op. cit.
II with numerous cucullate setae (Fig. 2a); Fig. 8) were not considered homologous.
1, distinctly clavate, row II with few or no 28. Dorsoapical emargination: 0, absent;
cucullate setae (Figs, lb, j). Some Erechtia 1, present (Figs. Ic, k).
andLeioscytaspecieshavethemetathoracic
tibiaweaklyclavate, but with numerouscu- Analysis
cullate setae in row II, perhaps representing Ourdatamatrix(Table 1)includedatleast
an intermediate state ofthis character. one representative ofeach genus of Mem-
