In e eb a e Zoology, 2025, 22(1): 30–43 © INVERTEBRATE ZOOLOGY, 2025
P oboscis appa a us: Wha do we know (and no know)
abou neme eans?
A.V. Che nyshe 1*, T.Yu. Maga lamo 1
1 A.V. Zhi munsky Na ional Scien i ic Cen e o Ma ine Biology, Fa Eas e n B anch, Russian Academy
o Sciences, ul. Palche skogo 17, Vladi os ok 690041 Russia.
Alexey Che nyshe : [email p o ec ed] ORCID h ps://o cid.o g/0000-0002-2203-3001
Timu Maga lamo : [email p o ec ed] ORCID h ps://o cid.o g/0000-0003-4175-5007
* Co esponding au ho
ABSTRACT: This pape p o ides an o e iew o ecen da a on he mo phology and unc-
ions o he p oboscis in neme eans. I is s ill unclea whe he hynchocoel and p oboscis
a e synapomo phies o he phylum Neme ea and whe he hynchocoel is homologous o
coelom in o he Spi alia. The p oboscis ini ially had a s uc u e simila o he body wall,
which is ypical o he class Palaeoneme ea. Howe e , subsequen ly, he p oboscis e ol ed
in wo di ec ions: (1) he de elopmen o bila e al symme y in palaeoneme eans om he
amily Cephalo ichellidae and Pilidiopho a (excep some Valenciniidae species) and (2)
he de elopmen o adial symme y in Hoploneme ea. Pilidiopho ans a e cha ac e ized
by a wide a ie y o p oboscis s uc u es, while hoploneme eans ha e he mos di e se
hynchocoel mo phology. The eme gence o he s yle appa a us in hoploneme eans led o
he loss o diagonal muscula u e and pseudocnidae in he p oboscis, as well as o a dec ease
in he numbe o he amily o pep ide oxins.
How o ci e his a icle: Che nyshe A.V., Maga lamo T.Yu. 2025. P oboscis appa a us:
Wha do we know (and no know) abou neme eans? // In e . Zool. Vol.22. No.1. P.30–43.
doi: 10.15298/in e zool.22.1.03
KEY WORDS: p oboscis, hynchocoel, coelom, endo helium, pseudocnidae, a ma u e,
e olu ion.
Хоботной аппарат: что нам известно (и не известно)
о немертинах?
А.В. Чернышев1*, Т.Ю. Магарламов1
1 Национальный научный центр морской биологии им. А.В. Жирмунского Дальневосточного
отделения Российской академии наук, Пальчевского 17, Владивосток, 690041 Россия.
* Ответственный за переписку: [email p o ec ed]
РЕЗЮМЕ: Данная работа посвящена обзору морфологии и функционированию
хоботного аппарата немертин. До сих пор нет однозначного представления является
ли наличие ринхоцеля и хобота синапоморфиями типа Neme ea и гомологичен ли
ринхоцель целому других Spi alia. Первоначально хобот немертин имел строение
сходное со стенкой тела, что типично для представителей класса Palaeoneme ea.
Однако в дальнейшем хобот эволюционировал в двух направления: (1) развитие
билатеральной симметрии у палеонемертин из семейства Cephalo ichellidae и у
Pilidiopho a (за исключением некоторых Valenciniidae) и (2) формирование радиальной
симметрии у представителей Hoploneme ea. Если пилидиофоры характеризуются
большим разнообразием в организации хобота, то у гоплонемертин наблюдается
De o ed o memo y o Claus Nielsen.
P oboscis appa a us: Wha do we know (and no know) abou neme eans? 31
In oduc ion
The p oboscis appa a us, a o ma ion consis -
ing o wo gene al componen s, he hynchocoel
(o p oboscis shea h) and he p oboscis p ope ,
is speci ic o Neme ea (Fig. 1A). Wi h con ac-
ion o he hynchocoel walls, he p oboscis is
e e ed ia hynchopo e and is in e ed back by
he e ac o muscle. The p oboscis is an o gan
o a ack and de ense; in some neme eans (p i-
ma ily e es ial and semi e es ial ones) (Fig.
1B), i is also used o locomo ion (Gibson, 1972;
Moo e, Gibson, 1981). The hynchocoel, in ad-
di ion o being a p oboscis ese oi , pe o ms
a suppo ing unc ion, especially well exp essed
in ben hic polys ili e ous neme eans (Che ny-
she , 2011). The neme ean b ain su ounds
he hynchocoel like a ing, bu no he o egu
as in many o he Spi alia. In mos neme eans,
one o wo essels o he ci cula o y sys em
en e he hynchocoel o e en he p oboscis
(Kajiha a, 2010; Che nyshe , 2011). In mos
hoploneme eans, he mou h me ges wi h he
hynchopo e o o m he hynchos omadaeum o
he a ium. All he abo e ac s sugges ha he
p oboscis appa a us is an essen ial componen in
he ‘Bauplan’ o neme eans. Thus, ano he name
o he phylum Neme ea is Rhynchocoela, which
is p e e ed by some au ho s (Hyman, 1951).
On he basis o p oboscis s uc u e, neme eans
we e di ided in o wo g oups: Anopla (una med
neme eans) and Enopla (a med neme eans). To
da e, a high classi ica ion has been adop ed ha
di ides neme eans in o h ee classes and se en
o de s: Palaeoneme ea (o de s Ca inomi o mes,
A chineme ea, and Tubulani o mes), Pilid-
iopho a (o de s Hub ech ii o mes and He e one-
me ea), and Hoploneme ea (o de s Polys ili e a
and Monos ili e a) (Che nyshe , 2021).
The only known neme ean species ha lacks
he p oboscis appa a us is he enigma ic A hyn-
choneme es axi desc ibed om New Zealand
(Rise , 1988). I s sys ema ic posi ion emains
un esol ed: i is ei he a ep esen a i e o an
ancien b anch o neme eans ha ha e no ye
acqui ed he p oboscis o a specialized neme ean
ha has los i s p oboscis and hynchocoel (Rise ,
1989). I he o me assump ion is co ec , hen
he p oboscis appa a us is no a synapomo phy
o Neme ea. Howe e , e en i he molecula
phylogene ic analysis shows he basal posi ion
o A. axi, i s ill does no answe he ques ion as
o whe he ances o s o his neme ean had he
p oboscis appa a us.
In he p esen e iew, we conside he mos
deba able issues o he mo phology and unc ions
o he p oboscis appa a us. These a e, i s , he
o igin o he p oboscis appa a us and also he
e olu ion o i s muscula u e, ne ous sys em,
and some epi helial s uc u es.
Do neme eans ha e ue coelom?
The p oboscis appa a us was well s udied
a he ligh mic oscopy le el in he la e 19 h
cen u y. The magni icen d awings o he p o-
boscis, especially he s yle appa a us, in he
books by McIn osh (1873–1894) and Bü ge
(1895, 1897–1907), s ill emain unsu passed. A
numbe o s udies wi h elec on mic oscopy o
he p oboscis we e published in he second hal
o he 20 h cen u y (Ling, 1971; S icke , Cloney,
1983; S icke , 1985; Mon al o e al., 1996, 1998;
Junoy e al., 2000). The mos impo an we e he
compa a i e analyses o mic oscopic ana omies
o neme eans om di e en g oups, based on
which he au ho s concluded ha he neme ean
hynchocoel is a coelomic ca i y (Tu be ille,
Ruppe , 1985; Tu be ille, 1991). The s a emen
ha he hynchocoel is a coelom was made by
Hyman (1951) and S a oboga o (1983), bu
Tu be ille & Rupe (1985) showed ha he en-
do helium o he hynchocoel and p oboscis has
all he mo phological ea u es o coelo helium.
высокое разнообразие в морфологии ринхоцеля. Появление стилетного аппарата у
гоплонемертин привело к потери диагональной мускулатуры и псевдокнид в хоботе,
а также к уменьшению числа семейств пептидных токсинов, которые используются
при нападении на добычу.
Как цитировать эту статью: Che nyshe A.V., Maga lamo T.Yu. 2025. P oboscis appa a us:
Wha do we know (and no know) abou neme eans? // In e . Zool. Vol.22. No.1. P.30–43.
doi: 10.15298/in e zool.22.1.03
КЛЮЧЕВЫЕ СЛОВА: хобот, ринхоцель, целом, эндотелий, псевдокниды, вооруже-
ние, эволюция.
A.V. Che nyshe , T.Yu. Maga lamo 32
Tu be ille & Rupe (1985) conside ed no only
he hynchocoel bu also he ci cula o y sys em
and gonadal sacs as de i a i es o coelom.
This in e p e a ion has no ound wide suppo
(especially as ega ds gonadal sacs), al hough
i has changed he opinion abou neme eans as
exclusi ely pa enchyma ous wo ms (howe e , in
many Russian uni e si ies, neme eans a e s ill
conside ed as acoelomic in e eb a es).
Ax (1996) insis ed on he o igin o he coelom
( hynchocoel and blood essels) in neme eans
independen om ha in Spi alia (he did no
a ibu e neme eans o Spi alia). Nielsen (2001)
exp essed a simila opinion by placing neme -
eans close o Pla yhelmin hes. Subsequen ly,
he (Nielsen, 2012) ended o assume ha blood
essels can be in e p e ed as modi ied coelomic
ca i ies, bu hynchocoel should unambiguously
be conside ed as an independen ly o igina ed sec-
onda y body ca i y. Malakho & Bogomolo a
(2016), in con as , s a ed ha hynchocoel is
he same homologue o coelom as blood essels.
Che nyshe (1999, 2011) p oposed a hypo he i-
cal scena io o he ans o ma ion o he pai ed
coelom in o he hynchocoel, blood essels,
and gonadal sacs in he neme ean ances o . I
opposed he hypo hesis by S a oboga o (1983)
who conside ed he hynchocoel homologous
o he unpai ed ac ocoelom (head coelom), and
he gonadal unnels homologous o he pai ed
unk coeloms.
The ques ion as o whe he he hynchocoel is
homologous o he ue coelom o o he Spi alia
o no emains open. I is also unclea which
one was o med ea lie , he p oboscis o he
hynchocoel. Wijnho (1914) and Senz (1997)
Fig. 1. Neme eans wi h in e ed (A) and e e ed (B) p oboscis appa a us. A — diag am o in e nal mo phol-
ogy o a hoploneme ean (d awn by Oleg Dob o olsky); B — palaeoneme ean Cephalo h ix c . simula wi h
e e ed p oboscis a acking a polychae e.
P oboscis appa a us: Wha do we know (and no know) abou neme eans? 33
sugges ed he p oboscis as an in agina ion o he
an e io end o he head ha was o med i s , and
hen he hynchocoel appea ed a ound i . This
hypo hesis does no explain why he coelomic
epi helium lines bo h he hynchocoel and he
in e ed p oboscis. Ano he hypo hesis seems
mo e plausible: he elonga ed an e io end o he
head in neme ean ances o s se ed o cap u e
ood (Fig. 2A) and subsequen ly began o in e
in o he al eady exis ing coelomic ca i y ( he
u u e hynchocoel) (Che nyshe , 2011). As a
esul , he p oboscis became lined inside by he
coelomic epi helium o his ca i y (Fig. 2B–E).
I is s ill impossible o con i m he co ec ness o
such a pa e n, and i emains pu ely hypo he ical.
The endo helium o he hynchocoel and
p oboscis has cilia cha ac e is ic o coelo he-
lium. These a e mos ly a e and sho , bu in
some neme eans hey can be long and nume -
ous (Maga lamo , Che nyshe , 2015) (Fig. 3D,
E). The endo helium s uc u es in he p oboscis
and he hynchocoel di e . The p oboscis endo-
helium is pseudos a i ied coelo helium, whe e
pe i oneal cells o e lap myocy es ha o m
he ou e (endo helial) ci cula muscula u e
(Maga lamo , Che nyshe , 2015). Ini ially,
myocy es a e ew in numbe and sca e ed, and
can only be ound in ul a hin sec ions o by
con ocal lase scanning mic oscopy (CLSM)
wi h phalloidin labeling (Fig. 3A). Howe e ,
in many neme eans, myocy es a e nume ous
and a anged in o se e al laye s (Fig. 3B); in
his case, ou e ci cula muscula u e becomes
clea ly isible in his ological sec ions. This has
c ea ed an e oneous imp ession ha some o
neme eans ha e he endo helial muscula u e
in he p oboscis while o he s do no .
The endo helium o he hynchocoel is ue
pe i oneum, and myocy es a e sepa a ed om he
endo helium by basal lamina and ex acellula
ma ix (Fig. 3C). I we assume ha myoepi he-
lium is he ini ial s a e o coelo helium (Kuzmina
e al., 2018), hen he p oboscis endo helium
has e ained mo e a chaic ea u es han he
hynchocoel endo helium.
The p oboscis mo phology as a
e lec ion o he an e io body end
While he o igin o he hynchocoel emains
unclea , he e a e almos no disag eemen s as
ega ds he o igin o he p oboscis: i is a modi ied
an e io ou g ow h o he head in e ed inside.
As Hyman (1951) no ed, his explains why he
s uc u e o he neme ean p oboscis epea s he
s uc u e o he neme ean’s body wall. Is i so?
Se en y yea s ago, he loca ions o he muscula
Fig. 2. O igin and ea ly e olu ion o Neme ea. A — hypo he ical ances o o neme eans (d awn by Oleg
Dob o olsky); B–E — diag ams o p esump i e o igin o p oboscis appa a us (yellow — hynchocoel,
ed — endo helium; blue — diges i e ac , g een — muscle- e ac o ).
A.V. Che nyshe , T.Yu. Maga lamo 34
laye s o he p oboscis and he body wall we e
examined only in his ological sec ions. The
CLSM and phalloidin labeling me hods ha e
e ealed new de ails in he s uc u e o bo h
he p oboscis and he body walls in neme eans
(Che nyshe , 2010, 2011, 2015). The p oboscis
s uc u e ully epea s he s uc u e o he body
wall only in palaeoneme eans ha ha e ou
laye s o muscula u e (ou e ci cula , diagonal,
longi udinal, and inne ci cula ) and a pai o
in aepi helial o subepi helial ne e co ds bo h
in he p oboscis and in he body wall (Fig. 4A,
B). The excep ion is ep esen a i es o he o de
A chineme ea whose diagonal muscula u e is
loca ed unde he ou e ci cula muscula u e
in he p oboscis and unde he epide mis in
he body wall (Che nyshe , Kajiha a, 2019).
Fu he mo e, he p oboscis o a chineme eans
Cephalo ichella (Fig. 5A) and Balioneme es
has he ou e longi udinal muscula u e ha is
absen om he body wall (Che nyshe , 2015;
Che nyshe , Kajiha a, 2019).
In a g ea numbe o species in he class Pi-
lidiopho a, bo h he p oboscis and he body wall
ha e he ou e longi udinal muscula u e (Fig.
4C). Howe e , he e is no comple e simila i y
he e because he diagonal muscula u e in he p o-
boscis is loca ed be ween he ou e ci cula and
inne longi udinal muscula u es (Fig. 4C), while
he diagonal muscula u e in he e oneme eans is
Fig. 3. Endo helium o p oboscis appa a us in neme eans. A — diag am o p oboscis endo helium o
Cephalo h ix c . simula, longi udinal sec ion; B — diag am o p oboscis endo helium in To us okmako ae,
longi udinal sec ion; C — diag am o hynchocoel endo helium in Hub ech ella juliae, ans e se sec ion;
D, E — CLSM mic og aphs o p oboscis endo helium labeled wi h α- ubulin an ibodies (D — Baseodiscus
c . p inceps; E — Ce eb a ulus sp.). Scale — 10 µm.
P oboscis appa a us: Wha do we know (and no know) abou neme eans? 35
Fig. 4. Diag ams o ans e se sec ions o e e ed p oboscis (A, C, E) and body (B, D, F). A, B — Palaeone-
me ea; C, D — He e oneme ea; E, F — Hoploneme ea.
siomo phic posi ion be ween he ou e ci cula
and inne longi udinal muscle laye s (Schwa z,
No enbu g, 2005; Che nyshe , 2010). Many
pilidiopho ans lack he ou e longi udinal mus-
cula u e in he p oboscis: in hub ech iids, his
lack is p ima y, while in he e oneme eans, i is,
loca ed in he cu is, and also be ween he ou e
longi udinal and ou e ci cula muscula u es o
he body wall in some species (Che nyshe , 2011;
Hookabe, Kajiha a, 2020). Only hub ech iids
and he he e oneme ean A chimic u a ignae
ha e he body diagonal muscula u e wi h a ple-
A.V. Che nyshe , T.Yu. Maga lamo 36
Fig. 5. Ligh (A) and CLSM (B–G) mic og aphs o p oboscis labeled wi h phalloidin and 5-HT an ibodies.
A — Cephalo ichella echinicola ( ans e se sec ion); B — Lineus i idis ( ans e se sec ion); C — Ca i-
noma sp. ( ans e se sec ion, da k a ows show muscle c osses); D — Cephalo h ix c . simula (longi udinal
sec ion); E — Ce eb a ulus sp. (longi udinal sec ion); F — Baseodiscus sp. (longi udinal sec ion); G — Ku-
iloneme es dilu ebasisae (longi udinal sec ion).
Abb e ia ions: e — epi helial idge, ilm — inne longi udinal muscula u e, mc — muscle c osses, ocm — ou e ci cula
muscula u e. Scale: A, C — 50 µm, B, D–G — 100 µm.
appa en ly, seconda y. In species o he gene a
Hub ech ella and Baseodiscus, he p oboscis
diagonal muscula u e is loca ed be ween he
inne ci cula and longi udinal muscula u es,
while he ou e ci cula muscula u e is absen , i.e.,
he p oboscis muscula u e di e s signi ican ly
om he body wall muscula u e (Che nyshe
e al., 2013). A o al o a leas eigh di e en
a ian s o a angemen o he muscula laye s
in he p oboscis ha e been eco ded among
Pilidiopho a, which is wice as many as in he
o he wo classes combined (Che nyshe , 2015).
Ano he ea u e o he p oboscis muscula-
u e in pilidiopho ans is he p esence o one o
wo muscle c osses, connec ed o he diagonal
muscula u e (Fig. 5B) (Che nyshe , 2010, 2015).
Muscle c osses a e p esen in mos he e one-
me eans and ecen ly ha e been desc ibed in
Hub ech ella ijimai (Kajiha a, 2006; Che ny-
she e al., 2017). In he body wall o some o
he e oneme eans, a do sal “c oss” is p esen
be ween he ou e ci cula muscle laye and he
hynchocoel muscula u e. The do sal and en al
muscle c osses a e cha ac e is ic o he body wall
in many palaeoneme eans which, howe e , lack
any simila s uc u es in hei p oboscises, excep
hose in Ca inoma (Fig. 5C) and Pa ahub ech ia
(see Che nyshe , 2010; Che nyshe e al., 2017).
In neme eans o he class Hoploneme ea, he
p oboscis epea s he body wall o an e en lesse
ex en : he p oboscis lacks diagonal muscula u e,
and se e al ne e co ds a e o ien ed adially in
he longi udinal muscula u e (Fig. 3E, F). Thus,
he muscula u e s uc u e and he loca ion o
he ne ous sys em in he an e io pa o he
p oboscis in hoploneme eans is e y conse ed,
wi h, howe e , some de ia ions in Malacobdella
and O o yphloneme es alen inae (Maga lamo ,
Che nyshe , 2010; Che nyshe , 2015).
The hoploneme eans a e cha ac e ized by
a di e si y o hynchocoel mo phologies. In a
numbe o hoploneme eans, he hynchocoel
has la e al, en al, o do sal pouches; in spe-
cies o he genus Unipo us, he la e al pouches
a e b anched. A leas nine di e en a ian s o
he hynchocoel muscula wall s uc u e ha e
P oboscis appa a us: Wha do we know (and no know) abou neme eans? 37
been dis inguished wi hin he class ( s. only
ou a ian s ound in palaeoneme eans and
pilidiopho ans), and i e, whe e he ci cula
muscula u e is ans o med in o he spi al one,
a e unique o pelagic neme eans (No enbu g,
Roe, 1998; Che nyshe , 2011; Che nyshe ,
Polyako a, 2018, 2019). The e is s ill no consen-
sus as o which a angemen o he hynchocoel
muscula u e is p ima y o Hoploneme ea:
om he ou e ci cula and inne longi udinal
muscula u es as in a sis e g oup, Pilidiopho a,
o om he in e wined longi udinal and ci cula
muscles as in ben hic Polys ili e a (see Kajiha a,
2021). Some au ho s sugges ed ha ini ially he
hynchocoel wall should ha e also e lec ed
he body wall, i.e., consis ed o sepa a e laye s
(Wijnho , 1914; Senz, 1997).
The p oboscis ne ous sys em in palaeone-
me eans consis s o wo ne e co ds loca ed
in aepi helially o subepi helially, epea ing
he ne ous sys em o he body wall (Figs 4A,
B; 5D). He e oneme eans usually ha e wo o
mo e pai s o main ne e co ds ha a e andomly
connec ed ia seconda y ne es, o ming a ne e
plexus wi h a p onounced bila e al symme y
(Figs 4C; 5E). In Baseodiscus, Sonneneme es,
and Hub ech ella, he ne e plexus o he p o-
boscis los i s bila e al symme y (Fig. 5F) and
became simila o he p oboscis ne ous sys em
o hoploneme ean Malacobdella (Maga lamo
e al., 2011; Che nyshe e al., 2013; Che nyshe ,
Polyako a, 2018). In hoploneme eans, se en
o 36 ne e co ds embedded in he longi udinal
muscula u e and nume ous connec i es be ween
hem o m a mo e o less o de ed adially sym-
me ical ne e plexus (Fig. 5G) (Che nyshe ,
2011; Maga lamo e al., 2011) (excep O o-
yphloneme es alen inae wi h i s seconda ily
bila e ally symme ical p oboscis — see Che -
nyshe , 2015).
P oboscis a ma u e
The main dis inguishing ea u e o he hop-
loneme ean p oboscis is a ma u e, which is also
ound, howe e , in some o Palaeoneme ea and
Pilidiopho a. The owel-like a ma u e is p esen
in he p oboscis o palaeoneme eans o he genus
Calline a (Kajiha a, 2006; Che nyshe , 2011,
2015) (Fig. 6A). I was i s econs uc ed and
desc ibed by Hi oshi Kajiha a and, he e o e, we
sugges e e ing o i as Kajiha a’s s yle . This
s uc u e is appa en ly composed o chi in and is
lexible. Func ions o he Kajiha a’s s yle emain
unknown, bu , when he p oboscis is e e ed,
i is loca ed subapically and can po en ially
cause wounds o p ey. An a ma u e consis ing
o a mul i ude o hooked pa as yle s has been
ound in he p oboscis o he he e oneme ean
He e oenopleus enigma icus (We n, 1998). Thus,
he e a e a med “una med” neme eans. Mo e-
o e , he e a e also una med hoploneme eans
such as species o he genus Malacobdella ha
comple ely los he s yle appa a us.
The ue s yle appa a us is p esen only in
hoploneme eans and is loca ed in he middle
p oboscis chambe . Two ypes o his appa a us
a e dis inguished: polys ili e ous (wi h nume ous
sho s yle s on a alci o m basis) (Fig. 6B) and
monos ili e ous (wi h a single long s yle on he
longi udinally s e ched basis) (Fig. 6C).
Fig. 6. P oboscis a ma u e. A — Calline a sp. (a ow shows Kajiha a’s s yle ); B — polys ili e ous a ma u e
o D epanopho idae sp. (inse shows s yle s a high magni ica ion); C — monos ili e ous a ma u e o Am-
phipo us sp. (a ows show accesso y s yle s); D — monos ili e ous a ma u e o Nipponneme es c . ubella
(a ow shows accesso y s yle inside basis); E — “bis ili e ous” a ma u e o C a eneme idae sp. IZ-45644.
Scale: A, B, D — 50 µm, C — 100 µm.
A.V. Che nyshe , T.Yu. Maga lamo 38
Fig. 7. Monocilia ed senso y cells (A–E) and pseudocnidae (F–I). A — diag am o monocilia ed senso y cell
o Lineus i idis; B — diag am o ans e se sec ion o he colla pa ; C — agmen o e e ed p oboscis
in Cephalo h ix c . simula wi h monocilia ed senso y cells (a ows show senso y p ocesses); D — CLSM
mic og aph o p oboscis epi helium in Nipponomic u a uchidai labeled wi h phalloidin (wi h colla mic o-
illi isible); E — SEM mic og aph o p oboscis epi helium o monocilia ed senso y cells in Hub ech ella
juliae (whi e a ow shows bulb-like s uc u e, black a ow shows mic o illi); F — diag am o pseudocnida
s uc u e in Lineus i idis; G — CLSM mic og aph o p oboscis epi helium in Cephalo h ix c . simula wi h
la ge (a ows) and small (a owheads) pseudocnidae (au o luo escence); H — pseudocnida o Hub ech ella
juliae wi h ex uded co e (a ow); I — SEM mic og aph o e e ed p oboscis in Mic u a kuliko ae showing
epi helial idge (a ows) wi h pseudocnida clus e s (inse shows pseudocnidae clus e a high magni ica ion).
Abb e ia ion: a — axial oo le . Scale: C, D, G, H — 10 µm, E — 2 µm, I — 50 µm.
