Review of the "red" Empoascini leafhoppers in the genera Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 (Hemiptera, Cicadellidae, Typhlocybinae) from Asia and the West Pacific

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u n:lsid:zoobank.o g:pub:27DA11CC-1A56-4E2A-BF0E-77F630407E7D
Webb M. D. & Xu Y. 2022. — Re iew o he “ ed” Empoascini lea hoppe s in he gene a Baguoidea Mahmood, 1967,
Dayus Mahmood, 1967 and Homa Dis an , 1908 (Hemip e a, Cicadellidae, Typhlocybinae) om Asia and he Wes
Paci ic. Zoosys ema 44 (22): 549-563. h ps://doi.o g/10.5252/zoosys ema2022 44a22. h p://zoosys ema.com/44/22
ABSTRACT
The “ ed” Empoascini lea hoppe s (Cicadellidae, Typhlocybinae), om Asia and he Wes Paci ic,
o he gene a Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Dis an , 1908 a e
e iewed. The ollowing species and subspecies a e ea ed: Baguoidea u a (Melicha , 1903) om
S i Lanka and Myanma , placed as a senio synonym o bo h Baguoidea ub a Mahmood, 1967
n.syn., om he Philippines and B. yunanensis Qin& Zhang, 2010 n.syn., om China; Dayus
eu yphaessus (Ki kaldy, 1907) om Aus alia(?) and Fiji; D. eu yphaessus ssp. ub ocinc a (Linna uo i,
1960a) om Fiji; D. o mosus Dwo akowska& Vi ak ama h, 1978 om China (Hong Kong, new
eco d; Taiwan) and India; D.upoluanus (Osbo n, 1934) om Samoa; Homa insignis Dis an , 1908
n.s a . (a species e alida ed om synonymy wi h H.haema op ila (Ki kaldy, 1906)) om S i Lanka,
Myanma (new eco d), China (new eco d) and Thailand (new eco d); H.haema op ila (Ki kaldy,
1906) om Aus alia (no S i Lanka, Philippines, Thailand and China as eco ded by Xu e al. 2022);
H.ka oi Dwo akowska, 1984 om Malaysia including Sabah (new eco d); H. ub odo sa a Ka o,
1933 om Taiwan; H.sinensis Qin& Zhang, 2011 om China. Checklis s o all he species o
Homa and Dayus a e gi en, as well as a key o Homa species. Images a e gi en o he i s ime o
he ypes o Baguoidea ub a, Dayus upoluanus, D. eu yphaessus, D.eu yphaessus ssp. ub ocinc a and
Homa haema op ila and geni alia igu es p o ided o he i s ime o Dayus upoluanus oge he
wi h e ised geni alia igu es o Homa insignis, he same as gi en o H.haema op ila by Xu e al.
(2022), in e o .
Michael D. WEBB
Depa men o Li e Sciences (Insec s), The Na u al His o y Museum,
London, SW7 5BD (Uni ed Kingdom)
[email p o ec ed] (co esponding au ho )
Ye XU
Ins i u e o En omology, Jiangxi Ag icul u al Uni e si y,
Nanchang 330045 (China)
[email p o ec ed]
Submi ed on 3 Feb ua y 2022 | Accep ed on 22 July 2022 | Published 7 Decembe 2022
Re iew o he “ ed” Empoascini lea hoppe s in he gene a
Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and
Homa Dis an , 1908 (Hemip e a, Cicadellidae, Typhlocybinae)
om Asia and he Wes Paci ic
KEY WORDS
Aucheno hyncha,
O ien al egion,
eigh h abdominal
s e ni e apodemes,
Kiwi ui ,
new eco ds.
550 ZOOSYSTEMA • 2022 • 44 (22)
Webb M. D.& Xu Y.
RÉSUMÉ
Ré ision des cicadelles Empoascini “ ouges” des gen es Baguoidea Mahmood, 1967, Dayus Mahmood,
1967 e Homa Dis an , 1908 (Hemip e a, Cicadellidae, Typhlocybinae) de l’Asie e du Paci ique oues .
Les cicadelles Empoascini “ ouges” de l’Asie e du Paci ique oues appa enan aux gen es Baguoi-
dea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Dis an , 1908 son é isées. Les espèces
e sous-espèces sui an es son ai ées: Baguoidea u a (Melicha , 1903) de S i Lanka e Myanma ,
placée comme synonyme senio de Baguoidea ub a Mahmood, 1967 n.syn., des Philippines e de
B. yunanensis Qin& Zhang, 2010 n.syn., de Chine ; Dayus eu yphaessus (Ki kaldy, 1907) d’Aus a-
lie(?) e des îles Fidji ; D. eu yphaessus ssp. ub ocinc a (Linna uo i, 1960a) des îles Fidji ; D. o mosus
Dwo akowska& Vi ak ama h, 1978 de Chine (Hong Kong, nou elle signalisa ion; de Taïwan)
e de l’Inde ; D.upoluanus (Osbo n, 1934) de Samoa ; Homa insignis Dis an , 1908 n. s a . (une
espèce ici so ie de sa synonymie a ec H.haema op ila (Ki kaldy, 1906) e e alidée) du S i Lanka,
de Myanma (nou elle signalisa ion), de Chine (nou elle signalisa ion) e de Thaïlande (nou elle
signalisa ion) ; H.haema op ila d’Aus alie (non du S i Lanka, des Philippines, de Thaïlande ou de
Chine comme l’on appo é Xu e al. [2022]) ; H.ka oi Dwo akowska, 1984 de Malaisie, y comp is
Sabah (nou elle signalisa ion) ; H. ub odo sa a Ka o, 1933 de Taiwan ; H.sinensis Qin& Zhang,
2011 de Chine. Le lis e des espèces de Homa e Dayus es ou nie ainsi qu’une clé d’iden i ica ion
e isée pou les espèces de Homa. Des pho os son p oposées pou la p emiè e ois pou les ypes
de Baguoidea ub a, Dayus upoluanus, D. eu yphaessus, D.eu yphaessus ssp. Rub ocinc a e Homa
haema op ila ; les geni alia son illus és pou la p emiè e ois pou Dayus upoluanus ; e des igu es
é isées des geni alia de Homa insignis son ou nies, iden iques à celles données pa e eu pou
H.haema op ila pa Xu e al. (2022).
INTRODUCTION
Species o he lea hoppe gene a Baguoidea Mahmood, 1967
and Homa Dis an , 1908 and some species o Dayus Mahmood,
1967 a e mainly ed in colou . This colo a ion makes hem
conspicuous elemen s o he Asian and Paci ic auna and ead-
ily no iceable in collec ions. All h ee gene a belong o he
“Usha ia g oup” o Empoascini Dis an , 1908. This g oup,
i s men ioned by Qin e al. (2011), also comp ised he Asian
gene a, Goi a Dwo akowska, 1977, I ugoa Dwo akowska&
Pawa , 1974, T eu alka Qin& Zhang, 2008 and Usha ia
Dwo akowska, 1977; la e Radica u cus Qin& Zhang, 2010
was also included (Xu e al. 2021b: ig.3). The g oup was de-
ined by Xu e al. (2017: 468) by he unb anched MP+CuA
ein in he hind wing (Fig.3D); a solidly a ached o used
aedeagus and connec i e (Fig.3J) and in Dayus, Homa and
I ugoa he connec i e a ms a e also highly modi ied (Figs4E;
5K). All gene a also ha e all apical cells o he o ewing a ising
om he m cell (Fig.3C) wi h he 3 d apical cell pe iola e in
Baguoidea and Dayus (Fig.3C) and he basal g oup mac ose ae
o he subgeni al pla e p esen , which a e unca e apically in
Baguoidea (Fig.3L) and some Homa (Fig.5F). In addi ion,
all h ee gene a ha e he en al abdominal basal apodemes
educed, being eplaced in Baguoidea and Dayus wi h long
di e gen do sal basal apodemes (Fig.3M), and in Dayus an
unusual en al apodeme o he 8
h
s e ni e is also p esen
(Fig.4F), see also Rema ks unde Baguoidea and Dayus. P e ious
igu es o he basal abdominal apodemes in hese gene a ha e
ei he inco ec ly s a ed hem as en al o ha e no indica ed
hei posi ion. Qin e al. (2014) p o ided a key including all
h ee gene a and Xu e al. (2021b) a phylogeny o Empoascini.
The cu en wo k a ose om iden i ying ed ma ked y-
phlocybine specimens om he Wes Paci ic in he Na u al
His o y Museum, London collec ion, specimens which
p o ed o be mainly new species o Dayus and Homa. All
species o Homa ha e dis inc i e colou pa e n (Fig.1J)
and he single species o Baguoidea and some species o
Dayus a e mainly ed in colou ; hese gene a and species
a e e iewed he e and a key p o ided o hei sepa a ion.
A key o all Empoascini, including he “Usha ia g oup”
om China is gi en by Qin e al. (2014). The abo e h ee
gene a we e included in he i s majo wo k on O ien al
Typhlocybinae Ki schbaum, 1868 by Mahmood (1967),
see ‘Discussion’. O he ex e nally simila , p edominan ly
eddish, Asian Empoascini include he ollowing: Aleb oides
ubicundus Ishiha a, 1953 (see Dwo akowska 1997: 311,
igs726-731), A. ub us Dwo akowska, 1994b: 98, Schi-
zand asca ub i ons (Ma sumu a, 1931) (Dwo akowska
1982: 53, igs259, 260), and males o Aleb asca ac ini-
diae Hayashi& Okado, 1994 (on Kiwi ui ), Rubipa us
bis igma Xu, Die ich& Qin, 2016 (see Xu e al. 2016:
585, igs1-4), Nikko e ix galloisi Ma sumu a, 1931 (see
Dwo akowska 1982, igs268-275) and N. aibaiensis
Qin& Zhang, 2003.
MATERIAL AND METHODS
“Wes Paci ic” in he ex e e s o he egion be ween and
including Aus alia and he Philippines. Excep whe e indi-
ca ed, dis ibu ion eco ds ending wi h (?) ela e o eco ds
conside ed dubious by he au ho s.
MOTS CLÉS
Aucheno hyncha,
égion o ien ale,
apodèmes du hui ième
s e ni e abdominal,
kiwi,
signalisa ions nou elles.
551
Re ue o he “ ed” Empoascini lea hoppe
ZOOSYSTEMA • 2022 • 44 (22)
Abb e iA ions
The specimens s udied o e e ed o a e deposi ed in he collec ions
abb e ia ed in he ex as ollows:
EIHU En omological Ins i u e o Hokkaido Uni e si y,
Sappo o;
INHS Illinois Na u al His o y Su ey, Champaign,
NHM The Na u al His o y Museum, London;
NWAFU En omological Museum, No hwes A&F Uni e si y,
Yangling, Shaanxi;
QSBG Queen Si iki Bo anical Ga den, Chiang Mai;
USNM Uni ed S a es Na ional Museum, Washing on.
TAXONOMY
CheCklis o he “ ed” Wes PACi iC gene A
And sPeCies o he “Usha ia g ouP” o emPoAsCini
Baguoidea u a (Melicha , 1903); S i Lanka, Myanma , Phil-
ippines, China.
Baguoidea ub a Mahmood, 1967 n. syn.
Baguoidea yunanensis Qin& Zhang, 2010 n. syn.
Dayus eu yphaessus (Ki kaldy, 1907); Aus alia(?), Fiji
Dayus eu yphaessus ssp. ub ocinc a (Linna uo i, 1960a), Fiji
Dayus o mosus Dwo akowska& Vi ak ama h, 1978, India,
China (Hainan, Hong Kong, new eco d and Taiwan)
Dayus upoluanus (Osbo n, 1934), Samoa
Homa haema op ila (Ki kaldy, 1906); Aus alia (no S i Lanka,
Philippines, Thailand and China as eco ded by Xu e al. [2022]).
Homa insignis Dis an , 1908, n. s a ., S i Lanka, Myanma
(new eco d), China (new eco d), Thailand (new eco d)
Homa ka oi Dwo akowska, 1984; Malaysia, including Sabah
(new eco d).
Homa ub odo sa a Ka o, 1933; Taiwan.
Homa sinensis Qin& Zhang, 2011; China.
Family CiCAdellidAe La eille, 1825
Genus Baguoidea Mahmood, 1967
Baguoidea Mahmood, 1967: 40.— Qin e al. 2010: 55; 2014:
1498 (key).
yPe sPeCies.— Baguoidea ub a Mahmood, 1967 by o iginal
designa ion.
dis ibu ion.— Mainland Asia (S i Lanka, Myanma and China),
Philippines and Japan(?), see Rema ks o B. u a below.
emA ks
This mono ypic genus is en a i ely ega ded as dis inc om
Dayus. I di e s only in ha ing he basal g oup se ae o he
subgeni al pla e blun - ipped (also ound in some Homa
species) and dis al mac ose ae mo e nume ous (Fig.3L) and
in lacking an unusual en al apodeme o he 8 h abdominal
s e ni e, ound in Dayus. In o he espec s, i is simila o
Dayus in i s o e all ed colou , ha ing he3 d apical cell in
he o ewing pe iola e (Fig.3C), male pygo e wi h p ocesses
(Fig.3G), o m o he subgeni al pla e and aedeagus and well
de eloped male basal do sal abdominal apodemes (Fig.3M).
The genus was edesc ibed by Yu& Yang (2013) as he e we e
some e o s in Mahmood’s o iginal desc ip ion and igu es,
which also had implica ions on subsequen s udies (see Re-
ma ks unde B. u a below). Yu& Yang (2013) also no ed
ha Dwo akowska (1973) had s a ed (and igu ed) ha he
acial la e o on al su u es we e absen in Baguoidea bu we e
p esen in hei specimens, and a e also p esen in he speci-
mens s udied he e. This appa en mis ake by Dwo akowska
may ha e been due o he su u es being obscu e agains he
ed acial colou .
Baguoidea u a (Melicha , 1903)
(Figs1A-F; 3)
Empoasca u a Melicha , 1903: 212, pla e i, ig.2a, b.— Dis an
1908: 402.— Me cal 1968: 351 (see Rema ks below).
Baguoidea ub a Mahmood, 1967: 42, pla e 9, ig.1. n. syn.
Baguoidea u a – Dwo akowska 1973: 49, igs1-12, 15; 1994a: 5.
Baguoidea yunnanensis Qin& Zhang in Qin e al., 2010: 55, igs15-
27.— Qin e al. 2014: 1495, igs12, 46, 65, 94. n. syn.
dis ibu ion.— Mainland Asia (S i Lanka, Myanma and China),
Philippines and Japan(?) see inal commen s in Rema ks below.
mA e iAl exAmined.— S i Lanka • 1 ♀ ; Pe adeniya; IV.1906;
Dis an Coll; NHM.
Myanma • 1 ♂ ; Myi a, Dohe y; coll. Dis an ; NHM; pa asi ized
• 1 ♂ ; Myi a, Dohe y; coll. Dis an ; NHMUK 013588830.
Philippines • 1 ♂ ; I ugao P o ., Luzon, Banaue; 20.VII.1980; NHM.
emA ks
B. u a was desc ibed om a single specimen om S i Lanka
wi h he ollowing da a ( ansla ed om he Ge man): “Pe -
adeniya. This nice Cicadine (1 ♂) was cap u ed by D Uzel on
2 May 1903 in he Bo anical Ga den on he sh ub Dichopsis
lae i olia Ben h. [=Palaquium lae i olium (Thwai es) Engl.
(Sapo aceae)]”. As he eco ded hos plan in S i Lanka is
an endemic (c i ically endange ed) species and as B. u a is
known om ou side S i Lanka i clea ly eeds on o he hos s.
The specimens eco ded om Myanma by Dis an (1908) a e
p obably he same as examined he e (Fig.1E, F). Baguoidea
ub a was desc ibed om he holo ype male (Fig.1C, D) and
i e pa a ypes (Fig.1A, B) om he Philippines wi h da a:
“Baguio, Bengue , Bake ” (USNM). The new synonymy o
B. u a and B. ub a is based on he ype igu es o he o -
me gi en by Dwo akowska (1973), he o iginal desc ip ion
o he la e and images o i s holo ype sen by J. Zahnise
(USNM) and he specimens s udied. The di e ences be-
ween he wo species, no ed by Dwo akowska (1973: 49),
a e ei he e o s in he o iginal desc ip ion, i.e., Mahmood’s
inco ec s a emen o o ewings “mo led wi h ed pa ches”,
which a e no p esen in he holo ype images seen (see abo e)
o an accep able ange o species a ia ion, i.e., posi ion o
dis al aedeagal p ocesses; while he long pygo e p ocesses
igu ed by Mahmood o B. ub a is also p obably an e o .
A specimen om Myanma examined di e s sligh ly in he
male geni alia om he S i Lanka ype o B. u a ( igu ed by
Dwo akowska 1973) and he examined Philippine specimen

552 ZOOSYSTEMA • 2022 • 44 (22)
Webb M. D.& Xu Y.
in ha ing he pygo e p ocess sligh ly mo e sinua e apically
and in ha ing he la e al ine se ae adjacen o he mac ose al
ow sho e . The same se ae a e shown longe and g ea e in
numbe in Qin e al.’s (2010) ig.26 o he junio synonym
B.yunnanensis (see ep oduced igu e he e, Fig.3L). The la -
e species was desc ibed om a single specimen om China
and dis inguished om B. u a (and B. ub a, he o he
junio synonym o B. u a) by he o ewing colou (which
acco ding o all specimens seen is e oneous) and di e ences
in pygo e and subgeni al pla e se ae and spines a he apex
o he aedeagal p ocesses, all di e ences which all wi hin he
accep ed ange o species a ia ion. Geni alia igu es d awn
by Dwo akowska (1973) we e p esumably aken om he
holo ype, as he only specimen examined, and as shown by
Dwo akowska’s igs7-9 he base o he aedeagus was damaged
when dissec ed. Howe e , he co ec aedeagal base is shown in
Fig.3I (la e al iew) and Fig.3J (do sal iew) which ma ches
he specimens examined he e and which is ema kably simila
o ha o some Dayus species (see Fig.4D). I should also be
no ed, ha he subgeni al pla e basal g oup se ae a e do sal
(Fig.3L) a he han en al as shown in Dwo akowska’s (1973)
igs3, 4 and ha he abdominal apodemes desc ibed by Qin
e al. (2010), and shown in hei igu e 27 (and ep oduced
he e, Fig.3M), a e do sal, and a e a ea u e o he genus (see
gene ic Rema ks). Finally, he e e ences o Japan o his
species by Esaki (1932, 1950), Esaki& I o (1954) and Ka o
(1933b) need o be con i med due o he simila i y o some
o he ed ma ked Empoascini (see In oduc ion). The e e -
ence o he species om Japan (Ma sumu a 1934) p esumably
e e s o Dayus akagii Dwo akowska, 1971, as his species
was desc ibed om ma e ial in Ma sumu a’s collec ion om
Japan and also Hong Kong (see Rema ks unde D. akagii).
Genus Dayus Mahmood, 1967
Dayus Mahmood, 1967: 39.— Qin& Zhang 2007: 43.— Yu&
Yang 2013: 2.— Qin e al. 2014: 1498 (key); 2021a: 229.
yPe
sPeCies
.— D. elonga us Mahmood, 1967, by o iginal designa ion.
dis ibu ion.— Widesp ead in Asia and he Paci ic.
emA ks
This genus is en a i ely ega ded as dis inc om Baguoidea
(see Rema ks unde ha genus). I di e s only in ha ing he
basal g oup se ae o he subgeni al pla e no blun - ipped and
in ha ing an unusual en al apodeme o he 8 h abdominal
s e ni e wi h a pai o sho an e io lobes (Fig.4F), and also
ound he e wi h a se ies o sho se ae a he pos e io la e al
co ne o he 8
h
s e ni e. The e is conside able a ia ion among
species in p opo ion o he e ex, size and colou , o exam-
ple, in Dayus o mosus he e ex is dis inc ly longe medially
han nex o eye bu mo e o less he same in D. i u ca us
Yu& Yang, 2013 while he la e is yellow and no ed as in
some o he congene s and e y much bigge ( wopa a ypes
o D. i u ca us [♂ and ♀] examined, NHM). A key o he
Chinese species o he genus was gi en by Yu& Yang (2013).
CheCklis And dis ibu ion o DayUs sPeCies
D.bi u ca us Yu& Yang, 2013: 3, igs1-9. China (Zhejiang).
D.elonga us Mahmood, 1967: 39, pla e 8, ig.1. Malaysia
(Singapo e).
D.eu yphaessus (Ki kaldy, 1907): 68. Fiji, Aus alia(?).
D. o mosus Dwo akowska& Vi ak ama h, 1978: 544, igs33-
41. India, China (Hainan, Hong Kong new eco d, Taiwan).
D. u ca us Xu, Die ich& Qin, 2021a: 231, igs10-17.
Thailand.
D.lamella us Qin& Zhang, 2007: 48, igs22-31. China
(Fujian, Zhejiang).
D.lii Qin& Zhang, 2007: 45, igs1-12. China (Fujian).
D.memb anaceus Qin& Zhang, 2007: 45, igs13-21. China
(Fujian, Jiangxi).
D.se a us Yu& Yang, 2013: 4, igs24-30. China (Hainan).
D. akagii Dwo akowska, 1971: 501, igs1-11. China (Si-
chuan, Hong Kong, Taiwan), Japan.
D. i u ca us Yu& Yang, 2013: 3, igs10-23. China (Chong-
qing).
D.upoluanus (Osbo n, 1934): 190, ig.15. Wes e n Samoa.
Dayus eu yphaessus (Ki kaldy, 1907)
(Fig.2A-F)
Cicadula eu yphaessa Ki kaldy, 1907: 68; 1908: 383.
Empoasca eu yphaessa – Linna uo i 1960a: 17, Fig.5 , h-j.— E ans
1966: 266 (misiden i ica ion(?) see Rema ks below).— Wilson
2009: 46.
Empoasca eu yphaessa ub ocinc a Linna uo i, 1960a: 18.— Dwo a-
kowska 1971: 501.— Wilson 2009: 46.
Dayus eu yphaessus – Dwo akowska 1971: 501.
dis ibu ion.— Fiji, Aus alia(?) see Rema ks below.
mA e iAl exAmined.— Fiji • 1 ♂ ; Labasa; R. Vei ch; VII.1921;
NHM • 1 ♀; Lau oka; W. G eenwood; 6.XII.1921; b eeding on
lea es o Glochidion sp.; NHM • 1 ♀; Lolo i; W. G eenwood;
19.XII.1920; NHM.
emA ks
This species was desc ibed om an unknown numbe o
specimens (syn ypic) om Fiji wi h da a: “Vi i Le u, Rewa
(Ma .-Ap .) Na na (Feb. Mui ’s No. 53) on a na i e ee, also
on Sacca um o icina um” (Suga cane) (BPBM). The da a
label on an imaged specimen seen (BPBM) labelled bo h
“ ype” and “Holo ype”, is as ollows: “Fiji Is 1905 [p in ed]
53[handw i en]” (Fig.2A, B). An image o ou o he syn ype
se ies specimens (labelled “Pa a ype”) has been seen (BPBM).
In i s desc ip ion Ki kaldy no ed he ollowing: “Allied o
C. u a (Melicha ) [=Dayus u us] bu he head is longe ,
p ono um sho e , legs pale, e c. B igh sca le , e ex pale
es aceous wi h a medio-longi udinal su used sca le s ipe,
which o ks a he base o he ons and ex ends all o e he
ace su usedly (some imes he ed s ipe is obsole e a he
o k, he ons hen being en i ely pale, only he genae and
he clypeus, e c., ed.)”.
553
Re ue o he “ ed” Empoascini lea hoppe
ZOOSYSTEMA • 2022 • 44 (22)
Fig. 1. — Baguoidea Mahmood, 1967 and Homa Dis an , 1908 species: A-F, Baguoidea u a (Melicha , 1903), A, B, pa a ype o B. ub a Mahmood, 1967, A, do sal
habi us, B, labels; C, D, holo ype o B. ub a Mahmood, C, male geni alia (on moun ), D, o e and hind wings on moun ; E, do sal habi us (non- ype: Myanma );
F, labels o same; G-I, Homa insignis Dis an , 1908 (lec o ype), G, do sal habi us; H, o e and hind wings on moun ; I, labels; J-K, Homa ka oi Dwo akowska, 1984
(pa a ype), J, do sal habi us, K, labels; L-M, Homa haema op ila (Ki kaldy, 1906) (holo ype), L, do sal habi us, M, labels. Scale ba s: A, L, 1 mm; E, G, J, 2 mm.
A
B
E
F
G
J
K
L
M
HI
CD
554 ZOOSYSTEMA • 2022 • 44 (22)
Webb M. D.& Xu Y.
Fig. 2. — Dayus Mahmood, 1967 species: A-F, Dayus eu yphaessus (Ki kaldy, 1907), A, B, holo ype, do sal habi us and labels espec i ely; C-F, ypes Empoasca
eu yphaessa ub ocinc a Linna uo i, 1960, C, D, allo ype, do sal habi us and labels espec i ely, E, F, holo ype, do sal habi us and labels espec i ely; G-N, Dayus
upoluanus (Osbo n, 1934), G, do sal habi us o emale pa a ype; H, labels o same; I, male basal do sal apodemes; J, male pygo e , la e al iew; K subgeni al pla e,
en al iew; L, X h segmen , en al iew; M, aedeagus, la e al iew; N, aedeagus, connec i e and s yle, en al iew. Scale ba s: A, C, E, 0,75 mm; G, 2 mm.
A
B
C
GH
J
K
L
M
N
I
D
E
F
555
Re ue o he “ ed” Empoascini lea hoppe
ZOOSYSTEMA • 2022 • 44 (22)
Dwo akowska (1971: 501) no ed he placemen o D.eu y-
phaessus in Dayus was based on he o iginal desc ip ion,
p esumably i s colou , p opo ions o he head and ho ax
and ena ion. In he o iginal desc ip ion i was also no ed
ha he ena ion was simila o Baguoidea u a and his is
con i med by Linna uo i’s (1960a) edesc ip ion, which
eads “ hi d apical cell o ely a iangula and s alked” and
he ma e ial examined he e. Howe e , he species is only
en a i ely e ained in Dayus as he male geni alia show
ce ain di e ences o ha genus, i.e., he pygo e lacks a
caudo-do sal lobe, he subgeni al pla e lacks a basal se al
g oup and he aedeagal sha is e y sho wi hou p o-
cesses, all cha ac e s also ound in he simila D.upoluana,
al hough he aedeagus and connec i e a e used (Fig.2N)
and do sal abdominal apodemes a e p esen (Fig.2I), as in
o he congene s.
Linna uo i (1960a) iden i ied (and igu ed) D.eu yphaessus
om 29 specimens om Fiji, Lami, as pa o a collec ion
om Fiji sen o him by BPBM (see In oduc ion in Linna-
uo i 1960a) and no he ype se ies, also BPBM (see abo e).
The iden i ica ion is howe e conside ed co ec based on he
dis inc i e ma king o he species. I is also wo h no ing ha
Linna uo i’s igu e o he aedeagus is in la e al iew, bu i s
a ached connec i e is shown in do sal iew, while his igu e
o long s e nal abdominal apodemes a e do sal, as in o he
Dayus species.
Linna uo i (1960a) also desc ibed a new subspecies (Em-
poasca eu yphaessa ub ocinc a) om he holo ype and allo ype
(BPBM) (Fig.2C-F) and one pa a ype in his own collec ion,
om he same locali y as he nomina e subspecies, Fiji, Lami.
I seems s ange ha Linna uo i (1960a), when e e ing o
his new subspecies, should say “As he nomina e o m…”
as he desc ibed he ed ma king o he wo subspecies di -
e en ly, con i med by he images ep oduced he e (Fig.2A,
C, E). Howe e , his obse a ion ha he male geni alia we e
he same in he wo subspecies, and hei same ype locali y,
sugges s hey a e he same axon wi h unaccoun ably di e en
ma kings. In synonymising he wo, Dwo akowska (1971)
no ed he ype-se ies o Empoasca eu yphaessa ub ocinc a
had been s udied bu om pe sonal communica ion a single
specimen was s udied.
The eco d o he species om Aus alia (Ku anda) by E ans
(1966: 266), could be inco ec and could be he specimen o
Homa haema op ila, om Ku anda, no ed below.
Dayus o mosus
Dwo akowska& Vi ak ama h, 1978
Dayus o mosus Dwo akowska& Vi ak ama h, 1978: 544, igs33-
41.— Qin& Zhang 2007: 51.
dis ibu ion.— India, China (Hainan, Hong Kong new eco d),
Taiwan.
mA e iAl exAmined.— India • 2 ♀ ; T. Nadu; Ye caud; I. Dwo a-
kowska de .; NHM.
China • 1 ♂ ; Hong Kong (new eco d); NHM.
emA k
This species was desc ibed om he holo ype male and onemale,
h ee emale pa a ypes om India. An addi ional male om
Taiwan was also eco ded.
Dayus akagii Dwo akowska, 1971
Empoasca u a – Ma sumu a 1934: 5 (nec Melicha 1903, misi-
den i ica ion).
Dayus akagii Dwo akowska, 1971: 501, igs1-11.— Lee 1979:
565-566, pla e 48, ig.264, igs528-530 (Ko ea).— Dwo akowska
1982: 54, igs282-288 (Japan).— Qin& Zhang 2007: 48, igs32-
40, China.
dis ibu ion.— Japan, Ko ea, China, Hong Kong(?) see Re-
ma ks below.
mA e iAl exAmined.— Japan • 1 ♀ ; Kyushu; Fukuoka; 10.I.1958;
NHM.
emA ks
This species was desc ibed om he holo ype male and se e al
pa a ypes (male and emale) om Kyushu (Japan) and wo
emale pa a ypes om Hong Kong. Al hough no deposi o y
o he ypes is indica ed in he o iginal desc ip ion, i s a es
in he acknowledgemen s o he pape ha he ype se ies was
bo owed om EIHU, and his is p esumed o be Ma sumu a’s
collec ion. The e o e, Ma sumu a’s misiden i ica ion gi en
in he o iginal desc ip ion (see abo e) e e s o Ma sumu a
(1934) whe e Empoasca u a Melicha is lis ed om Japan and
no (as s a ed) Ma sumu a (1931) whe e i is lis ed om Fo -
mosa. The emale pa a ypes om Hong Kong could equally
be D. o mosus examined he e om a male om Hong Kong
and hence he ques ion ma k unde Dis ibu ion.The colou
o D. akagii was no o iginally desc ibed excep ha he spe-
cies esembled D. eu yphaessus bu wi h pale colo a ion. The
emale en a i ely iden i ied he e as his species (based on i s
locali y Japan), is mainly eddish in colou .
Dayus upoluanus (Osbo n, 1934)
(Fig.2G-N)
Homa upoluana Osbo n, 1934: 190, ig.15.
Empoasca (Homa) upoluana – Me cal 1968: 442.
Dayus upoluanus – Dwo akowska 1971: 501.— Qin& Zhang
2007: 44.
dis ibu ion.— Samoa.
mA e iAl exAmined.— Pa a ypes • 2 ♂ , 1 ♀ , 1 specimen missing
om moun ; “Upolu Samoa”; “Apia, 9-15-23”; “Swezey& Wilde ”;
“B i . Mus. 1930-467”; NHMUK 013588829 • 1 ♂ ; “Sa aii, Samoa”
“Sa une, .2.24”; “ ain o es ; 2000-4000”; “H. B yan J ”; “B i .
Mus. 1930-467”; NHM.
emA ks
This species was desc ibed om 16 specimens o bo h sexes
(holo ype, allo ype and pa a ypes) wi h o iginal da a: “Apia,
562 ZOOSYSTEMA • 2022 • 44 (22)
Webb M. D.& Xu Y.
Acknowledgemen s
The au ho s hank Valé ie A.Lemaî e (NHM), Chand a
Vi ak ama h (Ag icul u al Uni e si y, Bangalo e, India),
Ch is Die ich and Dmi y Dmi ie (INHS) and Mu ay
Fle che ( o me ly o he O ange Ag icul u al Ins i u e, New
Sou h Wales, Aus alia) o cons uc i e commen s on ea lie
d a s o he manusc ip , James Zahnise (USNM) o imaging
ypes o Baguoidea ub a, D I. Dwo akowska ( o me ly o
he Museum and Ins i u e o Zoology o he Polish Academy
o Sciences, Wa saw, Poland and o he Fo es Sciences De-
pa men o he Uni e si y o B i ish Columbia, Vancou e ,
Canada) and Masami Hayashi ( o me ly o he Depa men
o Biology, Facul y o Educa ion, Sai ama Uni e si y, U awa
338, Japan) o help ul discussion and Rad ile Ma ke iciu e
(Na u e Resea ch Cen e, Li huania) o p epa ing he pla es.
Special men ion is made o Jim Boone (BPBM) who p ema-
u ely passed away, soon a e sending images o BPBM ypes.
The wo k by he second au ho was unded by he Na ional
Na u al Science Founda ion o China (32160121).
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wi hou a dis inc lobe (Figs 3L; 4I); aedeagal sha elonga e o e y sho .................................................... 2
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la e al lobe (Figs 5F; 6A); aedeagal sha elonga e (Homa Dis an , 1908) ..................................................... 5
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basal wo hi ds, i egula ly a anged and nume ous in dis al hi d (Fig. 3L); aedeagal sha elonga e (Fig.3I);
male 8 h abdominal s e ni e wi hou apodeme .............................................. Baguoidea u a (Melicha , 1903)
— Subgeni al pla e basal g oup mac ose ae acu e apically, la e al mac ose ae unise ia e h oughou and ew in
numbe (Fig.4I); male 8 h abdominal s e ni e wi h apodeme (Fig.4F) (Dayus Mahmood, 1967) ............... 3
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India, China (Hainan, Taiwan and Hong Kong) ................. Dayus o mosus Dwo akowska & Vi ak ama h, 1978
— Aedeagal sha e y sho wi hou a pai o apical p ocesses ......................................................................... 4
3. Aedeagal sha e enly cu ed in la e al iew (Fig. 2M); Samoa ..................... Dayus upoluanus (Osbo n, 1934)
— Aedeagal sha angula ly cu ed in la e al iew (see Linna uo i 1960: ig.5i); Fiji ..........................................
................................................................................................................ Dayus eu yphaessus (Ki kaldy, 1907)
4. Subgeni al pla e wi h la e o-basal lobe na ow, wi h g oup o la e al mac ose ae nea midleng h (Fig. 5M);
China ....................................................................................................... Homa sinensis Qin & Zhang, 2011
— Subgeni al pla e wi h la e o-basal lobe b oad, wi hou g oup o la e al mac ose ae nea midleng h (Fig. 6G) .... 6
5. Aedeagal sha ela i ely b oad in la e al iew (Fig. 6A); mainland Asia .............. Homa insignis Dis an , 1908
— Aedeagal sha e y na ow in la e al iew (see Dwo akowska 1978: ig.54); Malaysia ....................................
.................................................................................................................... Homa ka oi Dwo akowska, 1984

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Submi ed on3Feb ua y 2022;
accep ed on 22July 2022;
published on 7 Decembe 2022.