83
A new species o he genus Donacia Fab icius (Coleop e a,
Ch ysomelidae, Donaciinae) om Sou h Ko ea: mo phology,
ecology, and COI gene analysis
Joong Youb Kim1, Yeong-Deok Han1, Jong Eun Lee2
1 Resea ch Cen e o Endange ed Species, Na ional Ins i u e o Ecology, Yeongyang, Republic o Ko ea
2 Depa men o Biological Science, Gyeongkuk Na ional Uni e si y, Andong, Republic o Ko ea
Co esponding au ho : Jong Eun Lee ([email p o ec ed])
Copy igh : © Joong Youb Kim e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Donacia ko eana sp. no . is desc ibed om Sou h Ko ea, wi h de ailed desc ip ions and
illus a ions o bo h he adul and la al s ages. A key o species iden i ica ion, dis i-
bu ion map, hos plan associa ions, o iposi ion beha io s, eeding pa e ns, and o he
ecological in o ma ion, along wi h pa ial sequences o mi ochond ial cy och ome c ox-
idase subuni I, a e also p esen ed.
Key wo ds: La a, new species, eed bee les, axonomy
In oduc ion
The genus Donacia Fab icius, commonly known as eed bee les o pond lily lea
bee les, belongs o he amily Ch ysomelidae. In BugGuide.ne and iNa u alis .
com, Donaciinae a e e e ed o as aqua ic lea bee les, while Donacia as a genus
has no been assigned a mo e speci ic common name because he genus is oo
di e se. I is widely dis ibu ed ac oss he wo ld, wi h many species inhabi ing
eshwa e en i onmen s, and plays a signi ican ole in aqua ic ecosys ems.
Mos Donacia species occu in di e se eshwa e habi a s, such as ponds,
we lands, swamps, lakes, and s eams. Thei unique li e cycles and ecological
cha ac e is ics make hem impo an en i onmen al indica o species (Hayashi
2020; Geise 2023; Aske old 2024). Adul s eed on lea es and lowe s abo e he
wa e su ace, whe eas la ae li e unde wa e and eed on sap om plan oo s.
The la ae b ea he by apping he ae enchyma o he plan using wo hollow ab-
dominal spi acle hooks connec ed o hei acheal sys em (Geise 2023). Aske -
old (in p ess) summa ized he known esea ch on his subjec , as how donaci-
ines ob ain oxygen is no de ini i ely demons a ed o be jus one mechanism.
In Sou h Ko ea, se en species belonging o he genus Donacia ha e been
epo ed (Lee and An 2001; An 2019; Cho and An 2020; KSAE and ESK 2021;
Jeong e al. 2024). Wi hin his species lis , D. simplex Fab icius was misiden-
i ied as D. bicolo ico nis Chen, making hei accu a e di e en ia ion c ucial.
In pa icula , Eas e n Palea c ic Donacia species a e cha ac e ized by dis inc
di e ences in he p esence o absence o hai s on he p ono um and ely a,
Academic edi o :
Alexande Kons an ino
Recei ed:
18 Augus 2025
Accep ed:
20 Oc obe 2025
Published:
21 No embe 2025
ZooBank: h ps://zoobank.o g/
F7B1C36F-58E6-41EF-A82C-
DAC2DD6392BC
Ci a ion: Kim JY, Han Y-D, Lee JE
(2025) A new species o he genus
Donacia Fab icius (Coleop e a,
Ch ysomelidae, Donaciinae) om
Sou h Ko ea: mo phology, ecology,
and COI gene analysis. ZooKeys 1261:
83–99. h ps://doi.o g/10.3897/
zookeys.1261.169034
ZooKeys 1261: 83–99 (2025)
DOI: 10.3897/zookeys.1261.169034
84
ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034
Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
which has led in pa o con usion in classi ica ion (Chen 1966; Aske old 1990).
Kippenbe g (2015) sugges ed a e ised a angemen o Donacia species, p o-
posing se e al new subgene a and cla i ying some exis ing subgene ic names.
No ably, he subgenus name Donacocia Gis el applies o he as majo i y o
membe s o Donacia sensu la o. He e, he new species would be placed in he
D. (Donacocia) subgenus. The wo o he species o Donacia known o be in
Sou h Ko ea a e bo h in he subgenus D. (Cyphogas e ) Goecke.
Adul D. (Donacocia) a e ypically cha ac e ized wi hin he Donaciini ibe
by he coa sely punc a e- ugose p ono um, dis inc ly ugose o ugulose and
punc ula e ely a, ypically en i ely me allic colo o ely a, p ono um and ap-
pendages. Some excep ion exis in which species a e no me allic (e.g. D en-
nica), o wi h base o emo a eddish (e.g. D. bicolo ico nis and se e al o he s).
Species o D. (Donacocia) a e ne e en i ely pubescen do sally (D. pubescens
o No h Ame ica may no belong in his subgenus; Aske old in p ep.)
The main mo phological cha ac e is ics o Donacia la ae include: a sligh ly
con ex body wi h a e y small, scle o ized, p ogna hous head; dis inc ons,
clypeus, and lab um; mandible bea ing wo p ominen ee h; sho , sligh ly
cu ed, and awl-shaped legs lacking pul illi; i e s emma a on each side o he
head; a ho ax ha is unscle o ized and co e ed wi h nume ous se ae; abdomi-
nal segmen s lacking ube cles; and a la ge, scle o ized spi acle hook on he 8 h
abdominal segmen (Lee 1991).
In he p esen s udy, de ailed desc ip ions, illus a ions, and axonomic keys
we e de eloped o bo h adul and la al s ages o he new species. Fu he -
mo e, ecological da a, dis ibu ion maps, hos -plan associa ions, o iposi ion
beha io s, and eeding beha io s o he genus Donacia we e syn hesized, and
cy och ome c oxidase subuni I (COI) gene sequences we e analyzed.
Ma e ials and me hods
Type specimens o he new species desc ibed ha e been deposi ed in he Na-
ional Ins i u e o Biological Resou ces, Sou h Ko ea (NIBR) and J.Y. Kim’s p i-
a e collec ion, Sou h Ko ea (JYK).
Biological obse a ions we e conduc ed om May o June o 2022–2023 a he
ype locali y and unde labo a o y condi ions. Adul s we e collec ed by sweeping
and handpicking, whe eas la ae we e ob ained om oo s o hos plan s. Cap-
u ed adul s we e placed in Ziplock bags (20 cm wide, 15 cm long, and 8 cm deep)
wi h hos plan s and moni o ed daily, whe eas la ae we e p ese ed in 70% e hyl
alcohol in he ield. Fo mo phological analysis o minu e s uc u es, some la ae
we e dissec ed, clea ed in 10% KOH solu ion o 5 h, and insed in dis illed wa e .
Mic oscopic issues we e moun ed on slides, and specimens we e p epa ed us-
ing Neo-Shiga al solu ion (Shiga Konchu Fukyusha, Tokyo, Japan). To examine
geni alia, specimens we e boiled in ho wa e o app oxima ely 3 min o so en
issues. Geni alia we e hen ca e ully dissec ed using en omological needles and
subsequen ly imme sed in 10% KOH solu ion o 1–2 h o acili a e clea ing. De-
sc ip ions and illus a ions we e p epa ed using a s e eoscopic mic oscope (LEI-
CA M125, Olympus SZX-12) and an op ical mic oscope (Olympus BX50) equipped
wi h a d awing ube and Adobe Pho oshop CS6 so wa e (Adobe Sys ems, Inc.,
San Jose, CA, USA) using able s (WACOM In uos CLT-4100). Habi us images
we e cap u ed using a a Canon EOS 5D Ma k 5D IV digi al came a a ached o a
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ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034
Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
Canon EF 100 mm F2.8 Mac o USM. Mo phological e minology o adul s la gely
ollowed G essi and Kimo o (1961), Kimo o and G essi (1979), Mann and C ow-
son (1983), Aske old (1987, 1988, 1990, 1991), and Hayashi (2020), whe eas e -
minology o la ae was based p ima ily on Ande son (1947) and LeSage (2012).
Genomic DNA was ex ac ed om he legs o wo specimens using he Labo-
Pas Tissue Genomic DNA Isola ion Ki Mini (Cosmo Gene ech Inc., Seoul, Ko ea)
acco ding o he manu ac u e ’s ins uc ions. The COI ba code agmen was am-
pli ied using he uni e sal p ime se (LepF1 and HCO2198) unde he ollowing
cycling condi ions: ini ial dena u a ion a 94 °C o 2 min; ollowed by 35 cycles
o 98 °C o 15 s, 53 °C o 30 s, and 68 °C o 60 s; and a inal ex ension a 68
°C o 5 min. Ampli ied p oduc s we e hen sequenced using an ABI 3100 au o-
ma ed sequence (Pe kin-Elme , CA, USA). Sequence assembly, alignmen , and
imming we e pe o med using he Geneious 8.1.9 so wa e (Kea se e al. 2012).
COI sequences ampli ied om he newly iden i ied Donacia species we e
aligned wi h hose o 19 o he Donacia species e ie ed om he Na ional Cen-
e o Bio echnology In o ma ion da abase, using Geneious . 10.2.6 (Table 1).
Pai wise gene ic dis ances we e calcula ed using he Kimu a 2-pa ame e sub-
s i u ion model in MEGA X . 12.0.11 (Kuma e al. 2024). Phylogene ic ela ion-
ships we e in e ed using maximum likelihood analysis implemen ed in PhyML
. 3.1, based on he GTR + I + G model selec ed using jModelTes . 2.1.10, wi h
1000 boo s ap eplica es. The COI sequence o Pla euma is b acca a (Scopoli)
was used as an ou g oup o his analysis.
Resul s
Donacia ko eana Kim & Lee sp. no .
h ps://zoobank.o g/C52E8C52-E941-4A82-B007-2242FC6BB524
Figs 1–4
Type locali y. Sou h Ko ea, Kangwon S a e, Hyeoncheon- i, Dunnae-myeon,
Hoengseong-gun, swamp, 37°30'9.37"N, 128°9'53.57"E.
Type ma e ials. Holo ype (NIBR no. NIBRIN0001069714): male, Sou h Ko-
ea • Kangwon S a e, Hyeoncheon- i, Dunnae-myeon, Hoengseong-gun, swamp,
37°30'9.37"N, 128°9'53.57"E, 520 m, 29.V.2022, J.Y. Kim. Pa a ypes (NIBR no.
NIBRIN0001069715): • i e males and se en emales, same da a as he holo-
ype, excep o 6.VI.2023, J.Y. Kim.
O he ma e ials. Twen y la ae we e collec ed om he oo s o Sci pus wich-
u ae Boeckele wi h he same da a as o he pa a ype, excep o 6.VI.2023.
Diagnosis, adul (Figs 1, 2, 4). The new species closely esembles Donacia bi-
colo a Zschach, bu can be dis inguished by he ollowing diagnos ic cha ac e is-
ics: he ows o punc u es a e s ill clea ly isible e en in he las hi d o he ely a
(D. bicolo a: hese punc u es a e almos comple ely obscu ed by he silky, shiny,
ine w inkles in he las hi d o he ely a).; and i gene ally occu s in wo dis inc
colo mo phs: cup eous and g eenish (D. bicolo a exhibi s a a ie y o body col-
o a ions, commonly g eenish, bluish, o blackish). The legs a e en i ely me allic.
Desc ip ion, adul (Figs 1, 2, 4). Male. Leng h 7.9–8.1 mm, wid h 2.7–2.8
mm. Body na owly elonga e; gene ally cup eous o g eenish; head cup eous
o g eenish; an ennome es 1–4 coppe y o g eenish, 5 pa ially da kened, 6–11
da k; p ono um cup eous o g eenish; legs cup eous o g eenish.
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ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034
Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
Head. Na owe han p o ho ax; eyes small and con ex; sup aocula u ow
p esen bu shallow; e ex sligh ly swollen, pubescen wi h a ine median line.
An enna ili o m and pubescen ; an ennome e 5 longes amongs 2nd o 6 h; an-
ennome e 4 sligh ly longe han 3 d; an ennome e 5 2.5 imes longe han 2nd.
P ono um. Gene ally quad a e, wid h as long as leng h, pos e io and an e io
co ne s p ominen ; an e ola e al calli p esen , callosal sulcus p esen bu shal-
low; median line deep; disc coa sely punc a e and ans e se deep ugae; basal
sulcus p esen bu shallow, coa sely punc a e.
Scu ellum. Sub iangula , closely co e ed wi h sil e y pubescen .
Ely on. Subpa allel-sided, g adually na owed apically; su ace dis inc ly 10
longi udinal egula s ia e, s iae egula and deeply punc a e; wi h a shallow
dep ession a he base nea he su u e (1s –4 h su u al in e als), and a deep
dep ession a he base o he median a ea (5 h–8 h su u al in e als); apex un-
ca e, ou e apical angle ob usely ounded, inne apical angle ob use.
Legs. Slende ; me a emu obus in shape, ape ing and no dis inc ly cla -
a e, wi h a p ominen la ge oo h.
Pygidium. T apezoidal and apex pubescen , ema gina e in bo h sexes.
Table 1. Lis o Donacia species used in molecula analysis and he espec i e e e ences.
Species Collec ion locali y GenBank accession No. Re e ence
Donacia ko eana sp. no . Ko ea This s udy
Donacia aqua ica Ge many KU912172 Rulik and Ah ens 2017
Finland MZ659808 Roslin e al. 2022
Donacia au eocinc a Finland MZ658052 Roslin e al. 2022
Donacia bicolo a Finland MZ656234, MZ658244 Roslin e al. 2022
Donacia cinc ico nis Canada KM844344, HM411995 –
Donacia cine ea Ge many KU917042 Rulik e al. 2017
Finland MZ658078 Roslin e al. 2022
Donacia lemola China MK049870 Nie e al. 2020
Donacia hypoleuca Canada JF887697 –
Donacia ma gina a Ge many KU919476 Rulik e al. 2017
Finland KJ964703 Pen insaa i e al. 2014
Donacia obscu a I aly MH323099 Magoga e al. 2018
Finland KJ962943 Pen insaa i e al. 2014
Donacia pisca ix Canada KM844563 –
Donacia semicup ea Ge many KM443995 Rulik e al. 2017
Sweden KJ965000 Pen insaa i e al. 2014
Donacia simplex Ge many KU913219, KU906925 Rulik e al. 2017
Donacia spa ganii Ge many KU910558 Rulik e al. 2017
Donacia sub ilis Canada KR483642, KR486904 Hebe e al. 2016
Donacia halassina Ge many KU912621 Rulik e al. 2017
Finland KJ967484 Pen insaa i e al. 2014
Donacia omen osa F ance KM451348 Hend ich e al. 2015
Finland MZ631647 Roslin e al. 2022
Donacia e sicolo ea Ge many KU917168, KU919222 Rulik e al. 2017
Donacia ulga is Ge many KU915372 Rulik e al. 2017
Finland MZ631910 Roslin e al. 2022
Donacia p o os ii Ko ea OL663187 –
Pla euma is b acca a Finland MZ633983 Roslin e al. 2022
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Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
Male geni alia. Median lob o geni alia na owed apically wi h median lip and
bo h sides ema gina e subapically; a cap o egmen slende , apex wi h shallow
dep ession; endophallus elonga e, median p ocess sho and cu ing do sal,
BSB elonga e.
S e na. En i ely g eenish and pubescen ; apical shape o las s e ni e unca e.
Female. Leng h 9.0–9.3 mm, wid h 2.8–3.1 mm. Body la ge han ha o
males; p ono um on al ou line quad a e, wid h longe han leng h; apical
shape o las s e ni e poin ed.
Spe ma heca. Sho and b oad, s ongly C-shaped; apex ape ed; collum and
amus dis inc ; co nu and nodulus used in o a globose basal swelling; ho ns
widely sepa a ed.
Desc ip ion, i s ins a la a. Leng h 1.66–1.78 mm, wid h 0.40–0.41 mm,
spi acle-hook leng h (0.213 mm), and spi acle-hook wid h (0.033 mm (n = 4).
C eamy whi e, body sligh ly con ex, ho ax and abdomen wi h nume ous se ae,
bu no as se ae as he las ins a la a; egg bus e absen .
Diagnosis, las ins a la a (Figs 3, 4C). The las ins a la a is easily dis in-
guished om he o he Donacia la ae by he ollowing cha ac e is ics: epic a-
nium wi h six pai s o do sal se ae; wo pai s o sensilla; lab um wi h h ee pai s
o lab al se ae and one pai o sensilla; and p emen um wi h one pai o se ae
and wo pai s o sensilla.
Desc ip ion, las ins a la a (Figs 3, 4C). Leng h 10.5–12.3 mm, wid h
3.7–4.3 mm, head leng h 0.78–0.84 mm, spi acle hook leng h 0.73–0.77 mm,
spi acle hook wid h 0.42–0.44 mm (n = 5). Body yellowish, sligh ly con ex, and
C-shaped. Head, p o ho ax, and legs pale b own; se ae and spi acles b own.
Head. P ogna hous, small, ounded, well scle o ized, s ongly e ac ile in o
p o ho ax; hind co ne o epic anium sligh ly pos e io ly p oduced. Epic anial
su u e Y-shaped; on al su u e nea ly s aigh . S emma a pigmen ed, i e in
numbe . Epic anium wi h six pai s o do sal se ae, wo pai s o sensilla, and
h ee pai s o la e al se ae. F ons wi h ou pai s o on al se ae and one pai
o sensilla. An enna h ee-segmen ed: segmen 1 wi h wo sensilla, segmen
Figu e 1. Donacia ko eana sp. no . A, B. Male (A. Holo ype); C, D. Female (C. Pa a ype). Scale ba s: 1.0 mm (A–D).
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Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
2 wi h conical senso y papilla and wo sensilla basiconica, and segmen 3
wi h wo sensilla basiconica and one se a. Endoca ina absen ; epis omal su-
u es dis inc . Clypeus wi h wo pai s o clypeal se ae and one pai o sensil-
la. Lab um sligh ly no ched an e io ma gin wi h h ee pai s o lab al se ae
and one pai o sensilla. Epipha ynx wi h wo pai s o spini o m se ae. Man-
dible conical, s ongly scle o ized, wi h wo ee h, 2nd se a ed, and one man-
dibula se a. Maxilla palp h ee-segmen ed; palpi e wi h wo se ae; s ipes,
Figu e 2. Adul : Donacia ko eana sp. no . A. Leg (l. .); B, C. Pygidium (d. .); D, E. Abdo-
men (d. .); F. An enna (d. .); G. Median lobe (l. .); H. Apex o median lobe (d. .); I. A cap
o egmen; J, K. Endophallus (d. . and . .); L. Spe ma heca. male: A, B, D, F–K; emale:
C, E, L. Scale ba : 1.0 mm (A, D, E, F); 0.5 mm (B, C, G, H); 0.2 mm (I–L).
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Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
wo se ae and one sensillum; galea used wi h lacinia, spea -like; ca do la ge,
apezi o m, wi h one se a. Labial palp one-segmen ed wi h one sensillum;
ligula wi h nume ous se ae; p emen um wi h one pai o se ae and wo pai s
o sensilla; pos men um wi h wo pai s o se ae and one pai o sensilla.
Tho ax. P ono um pale b own, sligh ly scle o ized, wi h nume ous se ae.
Meso-and me ano as no scle o ized. Tho acic spi acles annuli o m, si ua ed
on he EPa; spi acula opening ounded. Legs sho and s ou ; emu wi h en
se ae and ou sensilla; ibio- a sus wi h h ee se ae; claws sligh ly cu ed, awl-
shaped, based enla ged, wi h one se a; pul illus absen .
Figu e 3. La a: Donacia ko eana sp. no . A. Las ins a la a (l. .); B. An enna (d. .);
C. Head (d. .); D. Clypeus, lab um and epipha ynx (d. . and . .); E. Mandible (b. .); F.
Spi acle (d. .); G. Maxilla and labium; H. Leg ( . .); I. 8 h abdominal spi acle. Scale ba :
1.0 mm (A); 0.5 mm (C, H, G, I); 0.1 mm (B, E, F).
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Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
Abdomen. Nine segmen ed, unscle o ized. Typical abdominal segmen s
wi h wo olds. Abdominal spi acles p esen on segmen s 1–8 simila o meso-
ho acic spi acles excep o 8 h spi acle. Spi acle hook leng h abou 2.3 imes
as long as wid h; p ojec ed, la ge, peg-like, s ongly scle o ized.
Dis ibu ion. Sou h Ko ea.
Hos plan s. Cype aceae: Sci pus wichu ae Boeckele , Eleocha is ussu iensis
G. Zinse l.
Key o he adul s o Sou h Ko ean Donacia species
1 En i e do sum o he bee le no eally me allic a all; appendages p edominan -
ly bicolo ed o u escen ; p ono al disc wi h a mos ine punc u es and ugu-
lae; ely a wi h a mos ew ine ugulae and minu e punc ulae in in e als ... 2
– En i e do sum o bee le me allic; appendages p edominan ly da k; p ono-
al disc coa sely punc u ed and o en ans e sely coa sely ugose; ely al
disc gene ally coa sely ugose be ween s iae ...........................................3
2 Thi d an ennal segmen sligh ly longe han, o as long as second ............
.......................................................................................... D. lenzi Schön eld
– Thi d an ennal segmen much longe han second .......D. p o os ii Fai mai e
3 Femo a and ibia en i ely me allic-colo ed ..................................................4
– Femo a and ibia pa ly me allic-colo ed ( emo al base usually u ous) ... 5
Figu e 4. Donacia ko eana sp. no . A, B. Adul s; C. La a; D. Pupal case; (C, D ed ci cle).
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Joong Youb Kim e al.: Donacia ko eana sp. no . om Sou h Ko ea
4 Ely on wi h ed longi udinal s ipe ........................... D. aqua ica (Linnaeus)
– Ely on wi hou ed longi udinal s ipe .........................................................6
5 P ono al disc wi h dis inc ine um wi h sho hai s (mos specimens) .....
...................................................................................D. cla a eaui Jacobson
– P ono al disc wi hou dis inc sho hai s .................D. bicolo ico nis Chen
6 Body and legs en i ely black, unicolo ed ......................... D. lemola Goecke
– Body and legs en i ely cup eous o g een ......................D. ko eana sp. no .
Key o he la ae o Sou h Ko ean Donacia species
1 Head inside he p o ho ax .............................................................................2
– Head ou side he p o ho ax ............................................. D. lemola Goecke
2 Hind co ne s o epic anium mode a ely p oduced .....................................3
– Hind co ne s o epic anium sligh ly p oduced ............................................4
3 Ephipha ynx wi h h ee pai s o spini o m se ae ....D. cla a eaui Jacobson
– Epipha ynx wi h wo pai s o spini o m se ae ............... D. ko eana sp. no .
4 Head o oid .....................................................................................................5
– Head ounded ................................................................................................6
5 Pos men um wi h wo pai s o se ae ............................. D. lenzi Schön eld
– Pos men um wi h h ee pai s o se ae..................... D. aqua ica (Linnaeus)
6 Clypeus wi h wo pai s o clypeal se ae....................D. bicolo ico nis Chen
– Clypeus wi h h ee pai s o clypeal se ae ..................D. p o os ii Fai mai e
Biological no e
Feeding pa e n
To in es iga e he eeding beha io o he Ko ean Donaciinae, he classi ica ion
sys em p oposed by Bieńkowski (2015) was adop ed. Acco ding o his ame-
wo k, h ee dis inc eeding pa e ns we e iden i ied.
• Type I: whe e he insec inclines i s head, u ns i o he igh o le , and
inse s bo h mandibles in o he lea issues.
• Type II: cha ac e ized by he inse ion o one mandible in o he lea , whe e-
as he o he emains on he lea su ace.
• Type III: he head mo emen o he insec esembles ha o Type II, bu
bo h mandibles cap u e he uppe and lowe su aces o he lea .
Based on he obse a ions ollowing his classi ica ion, D. lenzi and D. p o os ii
exhibi ed Type I eeding beha io . Donacia aqua ica, D. bicolo ico nis, and D. cla-
a eaui displayed Type II eeding beha io , whe eas D. lemola and D. ko eana
sp. no . exhibi ed Type III beha io (Fig. 5). No ably, al hough D. ko eana sp. no .
engaged in lea eeding, i s p ima y eeding si es we e he lowe s o Sci pus
wichu ae and Eleocha is ussu iensis a he han lea issues.
O iposi ion pa e ns
Adul Donaciinae exhibi di e se o iposi ion beha io s ha a y depending on
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