© INVERTEBRATE ZOOLOGY, 2023In e eb a e Zoology, 2023, 20(1): 57–89
Rossellidae (Po i e a: Hexac inellida) om he Be ing Sea
and o Be ing Island
K.R. Tabachnick1*, L.L. Menshenina2, H. Eh lich3
1 P.P.Shi sho Ins i u e o Oceanology, Russian Academy o Sciences, Moscow, Nahimo sky 36,
117997 Russia.
2 Biophysical Depa men , Physical Facul y, Moscow S a e Uni e si y, Moscow, 119992, Russia.
3 Ins i u e o Elec onics and Senso Ma e ials, Technische Uni e si ä Be gakademie F eibe g,
F eibe g 09599, Ge many.
* Co esponding au ho : [email p o ec ed]
Kons an in Tabachnick ORCID 0000–0002–7669–7233
La isa Menshenina ORCID 0000–0002–7668–9778
He mann Eh lich ORCID 0000–0003–4951–3555
ABSTRACT: Membe s o he amily Rossellidae a e he main con ibu o s o he hexac i-
nellid auna and e en o he mac oben hic auna in he deep wa e s o he No h Paci ic.
Rep esen a i es o his amily o en each a la ge size han o he Hexac inellida, Po i e a
and e en o he ep esen a i es o ben hic in e eb a es.
Desc ip ion o Rossellidae om he Be ing Sea collec ed by awls, subme sibles and
emo ely ope a ed ehicles (ROVs) du ing he expedi ions o RVs ‘Akademik M.A.
La en ye ’ and ‘Akademik Ms isla Keldysh’ a e gi en. Subgene a, species and subspe-
cies in i e gene a a e desc ibed and include: Acan hascus (Acan hascus) alani p o undus;
A. (Acan hascus) alani mic odiscoc as e us ssp.n.; Acan hascus (Rhabdocalyp us) bo e-
alis; Acan hascus (Rhabdocalyp us) he e as e ; Acan hascus (Rhabdocalyp us) mi abilis;
Acan hascus (S au ocalyp us) ylo us; Aulosaccus ijimai; Ba hydo us lanige ; B. spino-
sus; B. spinosissimus; B. sp.; Caulophacus (Caulophacus) elegans; C. (Caulophacus)
hype bo eus; C. (Caulophacus) mi i sp.n.; C. (Caulophacus) suba c icus sp.n.; C. (Cau-
lophacus) sp.; Hyalascus gigan eus; H. keldishi sp.n.
We sugges a new name B. le is neospinosus ssp.n. o B. le is spinosus o Wilson o a oid
he homonymy wi h B. spinosus Schulze and simul aneously we o e i as a lowe synonym
o he la e . Mean ime B. le is and B. spinosus a e conside ed o be di e en species.
The subdi ision o Rossellidae in o Rossellinae and Lanuginellinae has no easonable
mo phological g ounds in hei diagnoses anymo e and suppo o uni ing hese wo
sub amilies is p o ided.
How o ci e his a icle: Tabachnick K.R., Menshenina L.L., Eh lich H. 2023. Rossellidae
(Po i e a: Hexac inellida) om he Be ing Sea and o he Be ing Island // In e . Zool.
Vol.20. No.1. P.57–89, Suppl. Tables. doi: 10.15298/in e zool.20.1.03
KEY WORDS: Rossellidae, Be ing Sea, species and subspecies desc ip ions, Hexac inellida.
Rossellidae (Po i e a: Hexac inellida) из Берингова моря
и у острова Беринга
К.Р. Табачник1*, Л.Л. Меньшенина2, Г. Эрлих3
1 Институт океанологии им. П.П. Ширшова, РАН, Москва, Нахимовский проспект 36,
117997 Россия.
2 Кафедра биофизики, Физический факультет, Московский государственный университет
им. М.В. Ломоносова, 119992 Россия.
3 Ins i u e o Elec onics and Senso Ma e ials, Technische Uni e si д Be gakademie F eibe g,
F eibe g 09599, Ge many.
* Co esponding au ho : [email p o ec ed]
58 K.R. Tabachnick e al.
РЕЗЮМЕ: Губки семейства Rossellidae являются основными представителями Ше-
стилучевых губок и вцелом макробентоса в глубоководной зоне северной части
Тихого океана. Эти губки часто достигают значительных размеров среди других
Шестилучевых губок, представителей других классов губок и даже других бентос-
ных беспозвоночных.
Приводятся описания представителей Rossellidae из Берингова моря и тихоокеанс-
кой части острова Беринга собранные тралами, обитаемыми и необитаемыми под-
водными аппаратами в ходе экспедиций на кораблях “Академик Мстислав Келдыш”
и “Академик М.А.Лаврентьев”. Пять родов включают ряд видов, подвидов и подро-
дов: Acan hascus (Acan hascus) alani p o undus; A. (Acan hascus) alani mic o-
discoc as e us ssp.n.; Acan hascus (Rhabdocalyp us) bo ealis; Acan hascus (Rhabdo-
calyp us) he e as e ; Acan hascus (Rhabdocalyp us) mi abilis; Acan hascus (S au o-
calyp us) ylo us; Aulosaccus ijimai; Ba hydo us lanige ; B. spinosus; B. spinosissimus; B.
sp.; Caulophacus (Caulophacus) elegans; C. (Caulophacus) hype bo eus; C. (Caulophacus)
mi i sp.n.; C. (Caulophacus) suba c icus sp.n.; C. (Caulophacus) sp.; Hyalascus gigan eus;
H. keldishi sp.n.
Новое название B. le is neospinosus ssp.n. предложено для B. le is spinosus Wilson для
избегания ононимии с B. spinosus Schulze и одновременно назначается младшим
синонимом последнего. В тоже время B. le is и B. spinosus различаются как разные
виды.
Подразделение Rossellidae на Rossellinae и Lanuginellinae в настоящее время лишено
достоверных морфологических оснований, их диагнозы неразличимы, приводятся
доводы по ликвидации этих двух подсемейств.
Как цитировать эту статью: Tabachnick K.R., Menshenina L.L., Eh lich H. 2023.
Rossellidae (Po i e a; Hexac inellida) om he Be ing Sea and o he Be ing Island //
In e . Zool. Vol.20. No.1. P.57–89, Suppl. Tables. doi: 10.15298/in e zool.20.1.03
КЛЮЧЕВЫЕ СЛОВА: Rossellidae, Берингово море, видовые и подвидовые описа-
ния, Hexac inellida.
sampled by he ROVs di es du ing US expedi-
ions. We con inue he e he desc ip ion o he
Be ing Sea hexac inellids collec ed in i s wes -
e n pa , gene ally om he Piip Volcano, Ko-
mmando sk Basin and o Be ing Island. Sponges
we e sampled by awls, HOVs ‘Mi -1’ and
‘Mi -2’ du ing he 22nd c uise o he RV ‘Aka-
demik Ms isla Keldysh’and by he ROV ‘Co-
manche-8’ du ing he 75 h and 82nd c uises o
he RV ‘Akademik M.A. La en ye ’. A single
hexac inellid ep esen a i e belonging o a new
genus and species — Ijimaiella be ingiana om
he amily Euplec ellidae — was desc ibed ea -
lie by Tabachnick (2002a). Con inued desc ip-
ions o new axa show ha ou knowledge o
hese la ge-sized conspicuous animals is s ill a
om being comple e. Ree building hexac inel-
lids o amilies Fa eidae and Eu e idae om
he Be ing Sea a e desc ibed sepa a ely (Tabach-
nick e al., in p ess).
The ep esen a i es o he amily Rossel-
lidae, which a e he main con ibu o s o he
In oduc ion
In es iga ion o hexac inellid sponges om
he Be ing Sea s a ed wi h he desc ip ion o he
US ‘Alba oss’ collec ion in 1888–1890 om
he Aleu ian Islands ( o mally ou side he Be -
ing Sea (Schulze, 1899). Se e al o he ep e-
sen a i es o Hexac inellida om his a ea we e
subsequen ly desc ibed by Schulze (1899). In
1906 a se ies o awl s a ions we e made by a
subsequen ‘Alba oss’ expedi ion in he Be ing
Sea, o Be ing Island, o he Ku il Islands, and
o Japan. Specimens collec ed om hese s a-
ions we e examined and desc ibed by Okada
(1932). La e in es iga ion o he Be ing Sea
hexac inellids was connec ed wi h he SU expe-
di ions o he RV ‘Vi yaz’ and o he ships. The
esul s o hese expedi ions we e published by
Kol un (1967). Desc ip ion o hexac inellid
sponges om he eas e n pa o he Be ing Sea
we e published by S one e al. (2011) and Re-
iswig and S one (2013), whose ma e ials we e
59Rossellidae om he Be ing Sea
hexac inellid auna and e en o he en i e mac-
o auna in he No h Paci ic a e desc ibed be-
low. Rep esen a i es o Rossellidae in he gen-
e a Acan hascus and Aulosaccus o en each
abou hal a me e in leng h. O he species in he
genus Caulophacus, ha e peduncles o e 1–1.5
m long.
Recen ly, di e se s uc u al pecula i ies o
Rossellidae (Eh lich e al., 2008) emain o be
also in ocus o expe s in bioma e ials science.
Hexac inellids p incipally ha e been accep ed
wi hin his scien i ic di ec ion as unique sou ces
o biosilica inspi ed ma e ials chemis y (Eh -
lich e al., 2011, 2016) as well as po i e an
sca olding s a egies (Tsu kan e al., 2020;
Kh unyk e al., 2020). In addi ion, he ecen
disco e y o ac in wi hin skele al s uc u es o
ossellids (Eh lich e al., 2022) s imula es u -
he esea ch wi h espec o hei mo phology,
skele ogenesis and s uc u al biology.
Ma e ial and me hods
The specimens examined o his pape we e
collec ed du ing he c uise o he RV ‘Akademik
Ms isla Keldysh’ oyage 22 wi h Sigsbee awls
and HOVs ‘Mi ’; RV ‘Akademik M.A. La en ye ’
oyages 75 and 82 wi h he ROV ‘Comanche’. The
collec ed ma e ials we e p ese ed in 80–96% alco-
hol and s o ed a oom empe a u e. Fo ligh mic os-
copy, spicule p epa a ions we e made by he me hod
desc ibed by Janussen e al. (2004): a K2C 2O7
solu ion was made wi h wa e (K2C 2O7 powde :
wa e ˜1:1 ol%) and H2SO4 (96% conc.) was added
(K2C 2O7 solu ion: H2SO4 ˜1:1 ol%). A d y sponge
sample was placed on he mic oscopic slide; 1–2
d ops (depending on sample size) o he K2C 2O7
solu ion we e added. The mic oscopic slide was
hea ed (ca. 50–70 °C) o a ew minu es o le he
solu ion eac . A e e apo a ion o he luid, he
slide was emo ed om he hea and placed on a cold
su ace (ca. 20 °C) and a ew d ops o wa e we e
added. The wa e solu ion was emo ed by one o
se e al small pieces o no mal il e pape . Wa e
was added again and he spicule ca e ully s i ed by
needles, and again il e pape was used o emo e
excess wa e (occasionally i was necessa y o epea
his p ocedu e se e al imes). The d y p epa a ions
we e co e ed by Canada balsam and co e glass. The
p epa a ions we e examined wi h a BIOLAR op ical
mic oscope wi h PA–7 came a lucida adop ed o i .
Abb e ia ions: a g — a e age; D, d — diame-
e ; HOV — human-occupied ehicles; IORAS —
P.P. Shi sho Ins i u e o Oceanology o Russian
Academy o Sciences; L, l — leng h; max — maxi-
mum; min — minimum; n — numbe o measu es;
ROV — emo ely ope a ed ehicle; RV — esea ch
essel; s a. — s a ion; s d — s anda d de ia ion.
Sys ema ics
Rossellidae Schulze, 1885
The la es summa y o he ea ly his o y o Ros-
sellidae and i s scope is gi en by Tabachnick (2002b).
This amily is in he o de Lysaccinosida and was
o iginally subdi ided in o 3 sub amilies, Rosselli-
nae (cha ac e ized by ha ing nei he discoc as e s
no s obiloplumicomes), Lanuginellinae (cha ac-
e ized by he p esence o s obiloplumicomes bu no
discoc as e s), and Acan hascinae (cha ac e ized by
ha ing discoc as e s bu no s obiloplumicomes).
Tabacknick (2002b) abolished he la e sub amily
due o he p esence o spicules esembling discoc-
as e s in he holo ype o he ype species o he
genus Caulophacus — C. (Caulophacus) la us. This
genus oge he wi h Caulophacella was la e mo ed
om he Rossellinae o he Lanuginellinae (Doh -
mann e al., 2017; Reiswig e al., 2021). Reiswig and
S one (2013) esu ec ed he sub amily Acan hasci-
nae, a guing ha discoc as e s we e only ound in he
h ee gene a wi hin ha g oup, wi hou commen on
he possible p esence o hese dis inc i e spicules in
he genus Caulophacus. Howe e , we con inue o
adhe e o con end ha he abolishmen o he sub-
amily Acan hascinae is app op ia e and sugges
conside a ion o i s ee gene a s a us in he lowe
subgene ic le el. As o he wo emnan sub amilies
o Rossellidae, he only mo phological eason o
he subdi ision o his amily was absence in Rossel-
linae and p esence in Lanuginellinae o speci ic
spicules — s obiloplumicomes. The gene ic mono-
phyle ic cha ec e s o hese g poops sudges ed by
Doh mann e al. (2017) a e no suppo ed by any
common mo phological ea u es su able o he sep-
a a e diagnoses o hese wo sub amilies. Inspi e ha
he gene ic analysis o Lanuginellinae suppo s keep-
ing his sub amily no elible mo phological ea u es
could be es ablished o be alid o he diagnosis,
which p o ides i s de ini ion om Rossellinae. A
weak ea u e o his subdi ision is connec ed wi h
wo ac s. The i s : absence o a peculia i y (in his
case s obiloplumicomes in Rossellinae) is a p inci-
pally weak axonomic ea u e, being alone i does no
p o ide eliable g ound o he axon’s cha ac e iza-
ion – absence o a ea u e may be a esul o mul iple
loses o i , o he ea u e has no appea ed a all, hus
he possibili y o a polyphyle ic o igin o he axon
wi h such cha ac e is highly likely. And he second:
s obiloplumicomes we e no ound in some speci-
mens o monospeci ic Mellonympha — M. ella a (a
doub ul Lanuginellinae ep esen a i e) (Tabach-
60 K.R. Tabachnick e al.
Fig. 1. Acan hascus (Acan hascus) alani mic odiscoc as e us ssp.n. Comple e specimen be o e ROV
‘Comanche’ sampling.
Рис. 1. Acan hascus (Acan hascus) alani mic odiscoc as e us ssp.n. Целый экземпляр перед взятием
образца подводным роботом “Comanche”.
high wi h 2 oscula abou 80 mm in diame e . The
sponge was a ached o ocky subs a um; i s basal
pa is abou 100 mm in diame e . Some i egula
p os alia diac ins a e obse ed, hey a e gene ally
oscula ia bu some o hese spicules a e la e alia. The
walls o his specimen a e up o 25–30 mm in
hickness. Only a po ion, abou 100x100 mm and
10–25 mm in hickness, om he uppe pa o his
sponge was sampled, a la ges po ion o he holo-
ype specimen was le on he bo om ali e. The
collec ed ma e ial was educed o many lamella e
agmen s du ing he ixa ion and anspo a ion.
SPICULES. MEGASCLERES. Choanosomal
spicules a e diac ins wi h s ou , smoo h sha s, and
ounded, ough ou e ends. These diac ins a e 1.4–4/
0.004–0.013 mm. Se e al hypode mal pen ac ins
wi h smoo h and spiny angen ial ays we e ound
among he spicule p epa a ions, hey likely belong o
o he ep esen a i es o Acan hascus nume ously
dis ibu ed a ound his specimen. De malia and a i-
alia a e e y simila , bo h consis ing o pen ac ins,
hexac ins, some s au ac ins and a e diac ins wi h
sho -spiny ays and ounded ou e ends. Mos pen-
ac ins and some s au ac ins ha e ube cula ed udi-
men s o he absen ays. The only di e ence be-
ween de mal and a ial spicules is he dominance o
pen ac ins in he de mal sides o he colony and
hexac ins in he a ial sides. De mal pen ac ins ha e
angen ial ays 0.081–0.196 mm long (n=25, a g:
0.150 mm, s d: 0.029 mm), he p oximal ays 0.104–
0.233 mm long (n=25, a g: 0.169 mm, s d: 0.032
mm), and udimen s o he dis al ay i p esen
0.007–0.019 mm long (n=19, a g: 0.011 mm, s d:
nick, 2002b). Now he new phylogene ic da a led o
he ans e ing o he genus Caulophacus (i is
unclea comple e (wi h nume ous subgene a) o only
a pa o i ) and sugges ion o a new diagnosis o
Lanuginellinae (Reiswig e al., 2021). This ope a-
ion is a inal poin in he disappea ance o he las
eliable di e ences be ween he wo emaining sub-
amilies and his ac equi es he inal o mal ac ion
o he en i e abolishmen o discussed sub amilies.
Thus Rossellidae now is an in eg al amily, which
has no subdi ision in o sub amilies. Mean ime he
ac o p esence o absence o speci ic mic oscle es:
discoc as e s ( o me speci ic ea u e o Acan hasci-
nae abolished by Tabachnick (2002b)) and p esence
o s obiloplumicomes (speci ic ea u e o o me
Lanuginellinae) can be de ini ely used in keys o
gene a and subgene a o nume ous ossellid axa in
co esponding chap e s sugges ed by Tabachnick
(2002b) ollowing he insubs an ial (as i is ob ious
now) subdi ision in o g oups.
Acan hascus Schulze, 1886
Acan hascus (Acan hascus) alani Ijima, 1898
Acan hascus (Acan hascus) alani
mic odiscoc as e us ssp.n.
Figs 1, 2; Suppl. Tab. 1.
MATERIAL. Holo ype: IORAS 5/2/3805. RV
‘Akademik M.A. La en ye ’ – 75, ROV ‘Co-
manche’, s a. 18, spec. 5-2, 55.4352° N 167.2668°
E, 1420 m.
DESCRIPTION. BODY. Funnel-like body wi h
smoo h de mal and a ial su aces, abou 500 mm
61Rossellidae om he Be ing Sea
Fig. 2. Spicules o Acan hascus (Acan hascus) alani mic odiscoc as e us ssp.n., holo ype. A — de mal
s au ac in. B — de mal hexac in. C — de mal pen ac in. D — de mal diac in. E–F — choanosomal diac ins
and hei ou e ends. G–J — discoc as e s. K — p ima y ose e o discoc as e , seconda y ays no
de eloped. L — seconda y ay o discoc as e . M–N — sphe ical mic odiscohexas e s. O — s ella e
mic odiscohexas e . P–T — oxyoidal mic oscle es. U — p ima y ose e and seconda y ay o oxyoidal
mic oscle e.
Рис. 2. Спикулы Acan hascus (Acan hascus) alani mic odiscoc as e us ssp.n., голотип. A — дермальная
стаурактина. B — дермальная гексактина. C — дермальная пентактина. D — дермальная диактина.
E–F — хоаносомальные диактины и их концевые элементы. G–J — дискоктастры. K — первичная
розетка дисоктастра, вторичные лучи не развиты. L— вторичные лучи дискоктастра. M–N —
сферические микродискогексастры. O — звездчатый микродискогексастр. P–T — оксиоидные
микросклеры. U — первичная розетка и вторичные лучи оксиоидной микросклеры.
62 K.R. Tabachnick e al.
0.002 mm). The diame e o hese ays is 0.009–
0.015 mm. De mal hexac ins ha e ays o equal
leng h 0.159–0.252 mm long (n=3, a g: 0.201 mm,
s d: 0.047 mm). S au ac ins ha e angen ial ays
0.137–0.189 mm long (n=3, a g: 0.158 mm, s d:
0.027 mm), hei udimen al ube cles i p esen a e
0.007–0.015 mm long (n=3, a g: 0.012 mm, s d:
0.004 mm). Some unique diac ins ha e ays abou
0.222 mm long wi h a widening in he middle. A ial
pen ac ins ha e angen ial ays 0.085–0.178 mm
long (n=13, a g: 0.144 mm, s d: 0.024 mm), he ay
di ec ed inside he body is 0.074–0.204 mm long
(n=13, a g: 0.145 mm, s d: 0.033 mm), he udimen
o he dis al ay i p esen is 0.007–0.019 mm long
(n=13, a g: 0.010 mm, s d: 0.003 mm), he diame e
o hese ays is 0.010–0.011 mm. A ial hexac ins
ha e ays 0.104–0.326 mm long (n=26, a g: 0.170
mm, s d: 0.046 mm). A ial s au ac ins ha e angen-
ial ays 0.111–0.144 mm long (n=3, a g: 0.132 mm,
s d: 0.018 mm), hei udimen al ube cles i p esen
a e 0.011–0.015 mm long (n=4, a g: 0.013 mm, s d:
0.002 mm).
MICROSCLERES. Discoc as e s a e common
and a e discomul ias e s a e p esen in his species.
Unlike discohexas e s, discomul ias e s ha e mo e
han 8 u s o seconda y ays — up o 11, some imes
hese addi ional seconda y ays ha e a single sec-
onda y ay, bu usually hey ha e 2–8 seconda y ays
in a u . The seconda y ays o discoc as e s ha e
spiny sha s. Some discoc as e s ha e a speci ic
shape wi h 4 seconda y ay u s loca ed in one plane
dis ibu ed a abou 90 deg ee o each 4 o he second-
a y ay u s which a e simila o he o me ones a e
si ua ed below i . Some discoc as e s wi hou de el-
oped seconda y ays we e ound. The discohexas e s
a e 0.036–0,079 mm in diame e (n=25, a g: 0.061
mm, s d: 0.010 mm), hei p ima y ose e is 0.025–
0.050 mm in diame e (n=25, a g: 0.038 mm, s d:
0.007 mm). Sphe ical discohexas e s wi h nume ous
seconda y ays a e 0.023–0.036 mm in diame e
(n=20, a g: 0.028 mm, s d: 0.003 mm), hei p ima y
ose e is 0.006–0.014 mm in diame e (n=20, a g:
0.009 mm, s d: 0.002 mm). An occasional sphe ical
discohexas e is 0.065 mm in diame e wi h p ima y
ose e 0.011 mm in diame e . A unique s ella e
discohexas e was also ound wi h 3–4 seconda y
ays 0.054 mm in diame e wi h he p ima y ose e
0.027 mm in diame e . Oxyoidal mic oscle es a e
e y agile and in he spicule p epa a ions hey we e
usually b oken wi h hei p ima y ose es and sec-
onda y ays sepa a ed. The seconda y ays (1–3 in
numbe , bu some imes up o 7) ha e ines pa a
base wha p o ides he e ec desc ibed abo e, hey
a e co e ed by nume ous spines di ec ed owa ds he
spicula cen e especially a hei basal pa . Many
oxyoidal mic oscle es ha e seconda y ays o di e -
en leng h. The oxyoidal mic oscle es included oxy-
hexas e s, oxyhemihexas e s (mos o hese wo o ms
should be a he called anysooxyhexas e s and
anysooxyhemihexas e s) and hexac ins. The oxy-
hexas e s and oxyhemihexas e s a e 0.067–0.281
mm in diame e (n=26, a g: 0.139 mm, s d: 0.035
mm), hei p ima y ose e is 0.008–0.022 mm in
diame e (n=20, a g: 0.013 mm, s d: 0.003 mm). A
unique oxyhexas e was 0.027 mm in diame e wi h
p ima y ose e 0.009 mm in diame e . The oxy-
hexac ins a e 0.126–0.154 mm in diame e (n=7,
a g: 0.154 mm, s d: 0.025 mm).
REMARKS. In he p esence o he se o de mal
spicules and agile oxyoidal mic oscle es when
such spicules easily spli o p ima y and seconda y
ays he new subspecies is simila o wo p e iously
desc ibed subspecies: A. (Acan hascus) alani alani
Ijima, 1898 and A. (Acan hascus) alani p o undus
Kol un, 1967. The di e ences a e in he sizes o
discoc as e s, he p esence o discomul ias e s and
in he simila i y be ween de mal and a ial spicules
in he new subspecies (in p e iously desc ibed sub-
species a ialia a e gene ally hexac ins). Hypode -
mal pen ac ins wi h smoo h and spiny angen ial ays
ound in he new subspecies a e conside ed o ha e
alloch honous o igin bu mean ime hey poin he
close a ini ies be ween ee subgene a o Acan has-
cus which axonomic s a us was downg aded by
Tabachnick (2002b). I is e y likely, ha hese
subspecies should be aised o he species le el bu
due o he his o ical adi ion and com o in iden i-
ica ion o mul ispeci ic axon Acan hascus i s axo-
nomic posi ion is p ese ed a he momen . A ques-
ion o ising o he s a us o all he desc ibed
subspecies o speci ic le el equi es u he in es i-
ga ions.
ETYMOLOGY. The subspecies name e lec s
he cha ac e is ically small size o discoc as e s.
DISTRIBUTION. Cu en ly ound only a he
Piip Volcano slope, Be ing Sea, a 1420 m dep h.
Acan hascus (Acan hascus) alani p o undus
Kol un, 1967.
Figs 3, 4; Suppl. Tab. 2.
SYNONYMY. Acan hascus alani p o undum Kol-
un, 1967: 93.
MATERIAL. IORAS 5/2/3830, 3831: RV ‘Ak-
ademik M.A. La en ye ’ – 75, ROV ‘Comanche’,
s a. 9, spec. 2-3 and 2-4 co espondingly, 55.4282°
N 167.2772° E, 986 m. IORAS 5/2/3835: RV ‘Ak-
ademik M.A. La en ye ’ – 75, ROV ‘Comanche’,
s a. 9, spec. 1, 55.4282° N 167.2772° E, 984 m.
DESCRIPTION. BODY. The specimens a e sac-
cula : IORAS 5/2/3835 — 600 mm high, diame e o
he osculum is app oxima ely 250 mm (a agmen o
his specimen was sampled, i measu es 180 mm
high, 100 mm in maximal diame e o he osculum,
63Rossellidae om he Be ing Sea
Fig. 3. Acan hascus (Acan hascus) alani p o undus,
a ROV ‘Comanche’ pho o o he specimen IORAS/
5/2/3826. The a ow shows sampled agmen .
Рис. 3. Acan hascus (Acan hascus) alani p o undus,
фотография выполненая подводным роботом
“Comanche” образца IORAS/5/2/3826. Стрелка
показывает на взятый фрагмент.
Fig. 4. Acan hascus (Acan hascus) alani p o undus, ex e nal shape. Scale 50 mm. A — wo specimens:
la ge — IORAS/5/2/3830: small — IORAS/5/2/3831. B-C — iew om di e en sides IORAS/5/2/3826.
Рис. 4. Acan hascus (Acan hascus) alani p o undus, внешний вид. Масштаб 50 мм. A — два экземпляра:
большой — IORAS/5/2/3830; маленький — IORAS/5/2/3831. B-C — вид с разных сторон IORAS/5/
2/3826.
he walls a e 10–15 mm in hickness); 5/2/3830 —
180 mm high, 100 mm in diame e , he walls a e 8–
10 mm in hickness; 5/2/3831 — 110 mm high, 40
mm in diame e , he walls a e 5 mm in hickness. The
la e wo specimens a e a ached o each o he by
hei basal pa s; hei a ial ca i ies a e no common.
P os alia la e alia a e diac ins which p o ude a 10–
45 mm o e he de mal su ace.
REMARKS. Despi e some no able di e ences
in spicule sizes o de mal and a ial spicules and he
p esence o mic odiscohexas e s he newly ound
specimens a e assigned o Acan hascus (Acan has-
cus) alani p o undus Kol un, 1967 i s o all owing
o he p esence o wo size classes o discoc as e s.
Ano he ype o spicules ound in he new specimens
is mic odiscohexas e . Qui e p obably ha he di -
e ences be ween ou specimens and he o iginal
desc ip ion o he subspecies is a esul o ha only
a small agmen o a sponge was s udied ini ially
(Kol un, 1967). All o he spicules show a simila
ype in he a ia ions desc ibed ea lie and in newly
ound specimens. I is e y likely ha A. (Acan has-
cus) alani p o undus should be synonymized wi h A.
(Acan hascus) pla ei Schulze, 1899, bu i equi es a
signi ican e ision since he la ges discoc as e s
a e de mal and he smalles a e a ial (unlike A.
(Acan hascus) alani p o undus). Ano he species
wi h a no able di e ence in de mal and a ial discoc-
as e s is A. (Acan hascus) cac us, bu unlike he
specimens discussed abo e his species has gene al-
ly de mal s au ac ins and a ial pen ac ins.
DISTRIBUTION. Be ing Sea, a 984–2440 m
dep h.
64 K.R. Tabachnick e al.
mal pen ac ins ( hei angen ial ays) a e 10–15 mm
o e he de mal su ace. The hypode mal pen ac ins
a e mos pa a h opal wi h spiny angen ial ays.
REMARKS. Di e ences be ween he new and
p e iousely desc ibed specimens o A. (Rhabdoca-
lyp us) bo ealis (Okada, 1932; Kol un, 1967) a e
gi en in he Suppl. Tab. 3. The mos impo an o
hem: no mic odiscohexas e s in all in es iga ed
specimens we e obse ed; smalle diame e o he
oxyoidal mic oscle es; he ay o a ial hexac in may
be no ably smalle . Ne e heless hese ea u es a e
conside ed o be in aspeci ic, hey a e widening he
species limi s. Mo e e , he specimen desc ibed by
Kol un (1967) has some addi ional in e media e
pa ame e s.
DISTRIBUTION. Be ing Sea, Paci ic side o
Ku il Islands, a 440–2849 m dep h.
Acan hascus (Rhabdocalyp us) he e as e
Okada, 1932
Fig. 6; Suppl. Tab. 4.
MATERIAL. IORAS 5/2/3824: RV ‘Akademik
M.A. La en ye ’ – 75, ROV ‘Comanche’, s a. 21,
spec. 4, 55°28′58.8″ N 167°15′30.6″ E, 2849 m.
DESCRIPTION. BODY. The p ese ed speci-
men is ubula , abou 90 mm high, 50 mm in maximal
diame e ; he walls a e 1–3 mm hick. P os alia
la e alia a e hypode mal o ho opal ( a ely pa a h-
Fig. 5. Acan hascus (Rhabdocalyp us) bo ealis,
ex e nal shape. Scale 50 mm. A — IORAS/5/2/3827.
B — IORAS/5/2/3814. C — IORAS/5/2/3808.
Рис. 5. Acan hascus (Rhabdocalyp us) bo ealis, вне-
шний вид. Масштаб 50 мм. A — IORAS/5/2/3827.
B — IORAS/5/2/3814. C — IORAS/5/2/3808.
Acan hascus (Rhabdocalyp us) Schulze, 1886
Acan hascus (Rhabdocalyp us) bo ealis
Okada, 1932
Fig. 5; Suppl. Tab. 3.
MATERIAL. IORAS 5/2/3827: RV ‘Akademik
M.A. La en ye ’ – 75, ROV ‘Comanche’, s a. 18,
spec. 3-1, 55.4382° N 167.2652° E, 1569 m. IORAS
5/2/3808; 3814: RV ‘Akademik M.A. La en ye ’ –
75, ROV ‘Comanche’, s a. 9, spec. 2-5 and 2-2
co espondingly, 55°28′58.8″ N 167°15′30.6″ E,
2849 m.
DESCRIPTION. BODY. The bigges specimen
is IORAS 5/2/3827: 160 mm high, 60 mm in maxi-
mal diame e (close o he osculum); walls a e 2–6
mm hick. P os alia oscula ia a e diac ins which
p o ude up o 20 mm o e he osculum, p os alia
la e alia a e hypode mal pen ac ins which make a
dense co e up o 15 mm abo e he de mal su ace.
O he specimens a e smalle : IORAS 5/2/3814: 60
mm high, 20 mm in diame e in he middle and 15
mm in diame e o he osculum, he walls a e 1–1.5
mm hick; IORAS 5/2/3808: 50 mm high, 25 mm in
diame e in he middle and 20 mm in diame e o he
osculum, he walls a e 2–3 mm hick. P os alia in he
small specimens is smalle : oscula ia diac ins p o-
ude up o 15–20 mm, p os alia la e alia o hypode -
Fig. 6. Acan hascus (Rhabdocalyp us) he e as e ,
ex e nal shape. Scale 10 mm. A — iew om de mal
side. B — iew om a ial side.
Рис. 6. Acan hascus (Rhabdocalyp us) he e as e ,
внешний вид. Шкала 10 мм. A — вид с дермаль-
ной стороны. B — вид с атриальной стороны.
65Rossellidae om he Be ing Sea
opal) spiny pen ac ins i egula ly dis ibu ed and
p o uding 1–4 mm o e he de mal su ace.
REMARKS. The newly ound specimen has spi-
cules size pa ame e di e en om hose o he ype
specimen desc ibed by Okada (1932) (see he Suppl.
Tab. 4). The long ay o a ial hexac ins (di ec ed in o
he a ial ca i y) in he new specimen is no ably longe
hen hose in p e iouse desc ip ions; discoc as e s a e
o a single ype and hei minimal size is less han ha
in he ype specimen and la ge han he minimal size
o his ype o mic oscle e in a specimen desc ibed by
Kol un (1967). The discoc as e s in he new specimen
a e no ably la ge han he same ype o discoc as e s
in he ini ial desc ip ion, mic odiscohexas e s a e
absen and oxyoidal mic oscle es each la ge sizes.
Ne e heless mos spicule pa ame e s o his new
specimen co espond o A. (Rhabdocalyp us) he -
e as e much mo e hen o o he species o he
subgenus and i seem mo e easonable o expand he
spicule pa ame e s in he p e ious species hen o
c ea e a new one.
DISTRIBUTION. Be ing Sea, o Paci ic coas
o Canada, 445–2849 m dep h.
Acan hascus (Rhabdocalyp us) mi abilis
Schulze, 1899
Suppl. Tab. 5.
MATERIAL. IORAS 5/2/3826: RV ‘Akademik
M.A. La en ye ’ – 75, ROV ‘Comanche’, s a. 17,
spec. 3, 55.4599° N 167.269° E, 2279 m.
DESCRIPTION. BODY. The specimen IORAS
5/2/3826 is a agmen o he 5–10 mm hick uppe
wall o he sponge. P os alia oscula ia and la e alia
o la ge diac ins p o ude a 20–35 mm o e he
osculum ma gin. P os alia o hypode mal pen ac ins
p o ude a abou 10 mm o e he de mal su ace.
The diame e o he osculum is abou 80 mm in
diame e ( econs uc ed). Hypode mal pen ac ins o
bo h o ho opal and pa a h opal ypes ha e spiny o
smoo h angen ial ays.
REMARKS. The new specimen ag ees wi h he
desc ip ion o specimens o Rhabdocalyp us mi abilis
(Reiswig e al., 2013) excep o he absence o mic o-
discohexas e s. Bo h hese specimens ha e lo icoidal
discoc as e s ha di e om hose in he o iginal
desc ip ion o his species (Schulze, 1899) while hey
ha e lo icoidal discoc as e s. The discs o lo icoidal
discoc as e s a e assyme ically a ached o he second-
a y ay, while in he o iginal desc ip ion hey a e
comple ely symme ical. Spicules wi h assyme ical
ea u es a e also known in: A. (Rhabdocalyp us) un-
guicula us (Ijima, 1904) — a simila species wi h
p edomina ion o oxyhexas e s and oxyhemihexas-
e s o e he oxyhexac ins. This specimen also di e s
om A. (Rhabdocalyp us) gomezei (Tabachnick e
al., 2019), a species wi h no able amoun o de mal
s au ac ins, and A. (Rhabdocalyp us) biden a us —
s au ac ins p edomina e in his species and seconda y
ays o discoc as e s ha e 2 ee h (Okada, 1932).
DISTRIBUTION. O S o Alaska Peninsula,
Be ing Sea, 1143–2311 m dep h.
Fig. 7. Acan hascus (S au ocalyp us) ylo us, a ROV ‘Comanche’ pho os. A — wo specimens be o e
cap u ing su ounded by nume ous specimens o Fa ea sp. A — IORAS 5/2/3832 – a smalle specimen,
o he le ; IORAS 5/2/3807 – a bigge specimen, o he igh . B — magni ied po ion o he specimen IORAS
5/2/3807).
Рис. 7. Acan hascus (S au ocalyp us) ylo us, фотография подводного робота “Comanche”. A — два
экземпляра перед поимкой, окруженные многочисленными представителями Fa ea sp. A — IORAS
5/2/3832 – маленький экземпляр, слева; IORAS 5/2/3807 – больший экземпляр, справа. B —
увеличенный фрагмент образца IORAS 5/2/3807).
72 K.R. Tabachnick e al.
Fig. 12. Spicules o Ba hydo us spinosus, IORAS 5/2/3809 (addi ional ypes o p e ious desc ip ions).
A — small oxyhexas e . B — la ge oxyhexas e . C — oxyhexac in. D, E, G — abno mal oxyoidal
mic oscle es. F — oxyhemihexas e .
Рис. 12. Спикулы Ba hydo us spinosus, IORAS 5/2/3809 (в дополнение к предыдущим описаниям).
A — маленикий оксигексастр. B — большой оксигексастр. C — оксигексактина. D, E, G —
анормальные оксиоидные микросклеры. F — оксигемигексастр.
Fig. 13. Spicules o Ba hydo us sp., IORAS 5/2/
3838. A–B — de mal s au ac ins. C — oxyhemi-
hexas e . D — oxyhexac in.
Рис. 13. Спикулы Ba hydo us sp., IORAS 5/2/
3838. A–B — дермальные стаурактины. C —
оксигемигексастр. D — оксигексактина.
SPICULES. MEGASCLERES. Hypode mal
(and hypoa ial pen ac ins as well), a ely s au ac ins
wi h conically poin ed, ough ou e ends. The an-
gen ial ays o hese pen ac ins a e 0.3–1.0 mm long,
he ay di ec ed inside he body is 0.5–1.0 mm long,
hese ays a e 0.009–0.034 mm in diame e . Choano-
somal diac ins 2.3–3.7/0.008–0.014 mm ha e s ou
sha s o wi h a widening in he middle wi h ounded,
ough ou e ends.
De malia and possibly a ialia a e s au ac ins,
a ely auac ins. They ha e sligh ly ough ays wi h
conically poin ed o ounded ou e ends. The ays o
hese s au ac ins a e 0.085–0.344 mm in diame e
(n=27, a g: 0.192 mm, s d: 0.065 mm), he diame e
o hese ays is 0.004–0.013 mm.
MICROSCLERES. Oxyoidal mic oscle es ha e
e y hin ays. They a e usually oxyhexas e s and
oxyhemihexas e s wi h 1–3 seconda y ays, o en
cu ed o some imes oxyhexac ins. The diame e o
oxyhexas e s is 0.144–0.237 mm (n=25, a g: 0.170
mm, s d: 0.020 mm), he diame e o hei p ima y
ose e is 0.007–0.015 mm (n=25, a g: 0.008 mm, s d:
0.002 mm). The diame e o oxyhexac ins is 0.148–
0.215 mm (n=8, a g: 0.175 mm, s d: 0.021 mm).
73Rossellidae om he Be ing Sea
Fig. 14. Caulophacus (Caulophacus) elegans, ex e nal shape om di e en sizes. Scale 50 mm. A —
IORAS 5/2/2152. B — IORAS 5/2/2147. C — IORAS 5/2/2216. D — IORAS 5/2/2187.
Рис. 14. Caulophacus (Caulophacus) elegans, внешний вид с разных сторон. Масштаб 50 мм. A —
IORAS 5/2/2152. B — IORAS 5/2/2147. C — IORAS 5/2/2216. D — IORAS 5/2/2187.
Caulophacus (Caulophacus) elegans
Schulze, 1886
Figs 14, 15; Suppl. Tab. 11.
SYNONYMY: Caulophacus elegans Schulze, 1885:
438, ig. 159 — nomen nudum in he ig. oo no e; 1886:
46; 1887: 126; Kol un, 1967: 118.
MATERIAL. IORAS 5/2/2147: RV ‘Akademik
Ms isla Keldysh’ – 22, s a. 2309, awl, N o he
Be ing Island, 55°13.24–12.02′ N 167°29.07–26.7′
E, dep h 3957–3978 m. IORAS 5/2/2151; 2152;
2155; 2157; 2158.1; 2162; 2164; 2167; 2168; 2169;
2172; 2173; 2174; 2175; 2179; 2180; 2180.1; 2184;
2185; 2186; 2187; 2191; 2194; 2195; 2199; 2202;
2204; 2207; 2208; 2210; 2211; 2212; 2214; 2216;
2218; 2236; 2245; 2260; 2261; 2269; 2275; 2296;
2299; 2306; 2308: RV ‘Akademik Ms isla Keldy-
REMARKS. Among he species o Ba hydo us
only B. se a us Topsen , 1927 (1928) has simila
oxyoidal mic oscle es wi h such hin ays. Bu de -
mal s au ac ins a e much smalle as well as oxyoidal
mic oscle es a e also smalle in he newly ound
ma e ial, besides de mal diac ins a e absen (unlike
B. se a us). Based on a small agmen , desc ip ion
o a new species does no seem easonable, mo e-
o e , he spicule con en may be no comple e and
hus he gene ic iden i ica ion is also no pe ec .
Spicules o o he hexac inellids — Fa ea and
Pe iph agella we e also ound in in es iga ed spi-
cule p epa a ions so he specimen is ob iously con-
amina ed by o eign spicules.
Caulophacus Schulze, 1886
Caulophacus (Caulophacus) Schulze, 1886
74 K.R. Tabachnick e al.
Fig. 15. Spicules o Caulophacus (Caulophacus) elegans. A — de mal hexac in. B — a ial hexac in. C–
D — de mal o a ial pen ac ins: C — smoo h; D — wi h spines on he ays close o he spicule cen e . E —
choanosomal hexac in. F–H — choanosomal diac ins. I — spiny sphe ical discohexas e . J — abno mal
spiny discohexas e . K–O — lophodiscohexas e s: O — anysolophodiscohexas e . P — lophodiscodias e .
A–J; L; N–O — IORAS 5/2/2152. K; M; P — IORAS 5/2/2147.
Рис. 15. Спикулы Caulophacus (Caulophacus) elegans. A — дермальная гексактина. B — атриальная
гексактина. C–D — дермальные или атриальные пентактины: C — гладкая; D — с шипами на
основаниях лучей. E — хоаносомальная гексактина. F–H — хоаносомальные диактины. I —
шиповатый сферический дискогексастр. J — аномальный шиповатый дискогексастер. K–O —
лофодискогексастеры. O — анизолофодискогексастер. P — лофодискодиастер. A–J; L; N–O —
IORAS 5/2/2152. K; M; P — IORAS 5/2/2147.
75Rossellidae om he Be ing Sea
sh’ – 22, s a. 2316, awl, N o he Be ing Island,
55°36.08–35.00′ N 167°23.04–24.46′ E, dep h 4200–
4294 m.
DESCRIPTION. BODY. The specimens ha e a
mush oom shaped body ypical o Caulophacus.
Some specimens howe e a e jus agmen s. Dis-
coidal bodies a e om 33 o 200 mm in diame e ,
1.5–10 mm in hickness, wi h undula ing edges o
discs. Peduncles a e ubula 2.5–20 mm in diame e .
SPICULES. MEGASCLERES. Hypode malia
and hypoa ialia a e pen ac ins wi h ays ha ing
smoo h and ounded ou e ends o hey ha e ube -
cles o nume ous ube cles on he sha s close o he
spicula cen e . The angen ial ays o hypode mal
and hypoa ial pen ac ins a e 0.3–0.6 mm long, he
ay di ec ed inside he body is 0.5–0.7 mm long,
hese ays a e 0.017–0.038 mm in diame e . Choano-
somal spicules a e diac ins and hexac ins. Choano-
somal diac ins usually ha e a widening in he mid-
dle, some imes ou udimen al ube cles o hey a e
s ou , hei ou e ends a e ounded and ough. Cho-
anosomal diac ins a e 1.1–1.5/0.005–0.011 mm. The
diac ins o he peduncle a e 0.019–0.022 mm in
diame e ; hey a e connec ed by nume ous synap ic-
ula junc ions. Choanosomal hexac ins ha e smoo h
ays wi h ounded ou e ends, ha a e 0.45–0.9/
0.023–0.044 mm in leng h.
De malia and a ialia a e simila o each o he in
shape and gene ally in size. They a e hexac ins
which ha e ays di ec ed ou side he body. Rays a e
pinula and smoo h o sligh ly co e ed by ube cles;
o he ays ha e ounded o conically poin ed, ough
ou e ends. Ra ely de mal and a ial spicules a e
pen ac ins wi h a udimen al ube cle si ua ed in
place o he ay di ec ed inside he body. The pinula
ay o de mal hexac ins is 0.096–0.248 mm long,
angen ial ays a e 0.067–0.141 mm long, he ay
di ec ed inside he body is 0.048–0.137 mm long, he
diame e o hese ays is 0.007–0.011 mm; he diam-
e e o he pinula pa is 0.037–0.044 mm. The
pinula ay o a ial hexac ins is 0.111–0.340 mm
long, angen ial ays a e 0.067–0.144 mm long, he
ay di ec ed inside he body is 0.037–0.118 mm
long, he diame e o hese ays is 0.007–0.009 mm;
he diame e o he pinula pa is 0.026–0.037 mm.
MICROSCLERES. Ob iously wo ypes o dis-
coidal mic oscle es a e known in hese specimens.
Some oxyoidal mic oscle es bu likely hey belong
o o he hexac inellids om close loca ions. Disco-
hexas e s ha e sho p incipal ays, he ays a e spiny
and usually hey ha e 5–6 seconda y ays, a ely 1.
Abno mal discohexas e s a e a e. The discohexas-
e s a e 0.086–0.151 mm in diame e wi h p ima y
ose e 0.011–0.036 mm in diame e . Lophodisco-
hexas e s o en ha e ube cles ca ying some addi-
ional discoidal ays on he smoo h sha s; some-
imes he seconda y ays a e o di e en sizes (lo-
phoanysodiscohexas e s). The seconda y ays o la ge
lophodiscohexas e s a e sho –spiny, in small spi-
cules hey look o be smoo h and e en some imes
yloidal in shape (no comple ely de eloped). The
numbe o seconda y ays is om 4 o abou 30.
Ra ely lophodiscodias e s we e ound. Lophodisco-
hexas e s a e e e ed o a single size class o spi-
cules in spi e he signi ican size a ia ion. The gap
o hei size classes was obse ed in some specimens
bu in o he he gap is di e en and in some com-
ple ely absen . The diame e o lophodiscohexas e s
is 0.047–0.324 mm wi h p ima y ose e 0.022–
0.144 mm in diame e . The la ges lophodiscohexas-
e was ound in he specimen 5/2/2168 — 0.324 mm
in diame e wi h he p ima y ose e 0.108 mm in
diame e wha is close o ha desc ibed by Kol un
(1967) (0.330 mm in diame e ) which he conside ed
o ha e alloch honic o igin.
REMARKS. As i was obse ed, he specimens
o di e en sizes: small and big (Suppl. Tab. 11)
ha e e y simila sizes o spicules. Mo eo e , Table
11 shows ha he smalles specimen 5/2/2187 has
he mos spicules o la ge sizes han la ge speci-
mens.
Among nume ous species o Caulophacus (Cau-
lophacus) he newly ound ma e ials a e mos o all
simila o C. (Caulophacus) elegans in he p esence
o discohexas e s (unlike discohexac ins), shape o
mos spicules, bu unlike he p e iously known spe-
cies i has dissimila de mal and a ial pinula hexac-
ins. The de mal hexac ins a e smalle han he a ial
ones in C. (Caulophacus) elegans and gene ally
equal in he new specimens om he Be ing Sea.
DISTRIBUTION. Be ing Sea, E o Japan, 3680–
4220 m.
Caulophacus (Caulophacus) hype bo eus
Kol un, 1967
Figs 16–18; Suppl. Tab. 12.
MATERIAL. IORAS 5/2/2084; IORAS 5/2/
2153; 2154; 2161.1; 2228.1; 2229; 2231; 2232;
2233; 2234: RV ‘Akademik Ms isla Keldysh’ – 22,
s a. 2309, awl, N o he Be ing Island, 55°13.24–
12.02′ N 167°29.07–26.7′ E, 3957–3978 m. IORAS
5/2/2150; 2158; 2177; 2178; 2182; 2183; 2183.1;
2217: RV ‘Akademik Ms isla Keldysh’ – 22, s a.
2316, awl, N o he Be ing Island, 55°36.08–
35.00′ N 167°23.04–24.46′ E, 4200–4294 m. IO-
RAS 5/2/3811; 3811.2–3811.9; 3820, 3821: RV
‘Akademik M.A. La en ye ’ – 75, s a. 22, ROV
‘Comanche’, 55°30′36.1″ N 167°19′27.1″ E, 3879–
3578 m.
DESCRIPTION. BODY. The specimens had
mush oom-like bodies o agmen s o en a ached
o a peduncle b anching in o many sepa a e bodies.
I is no ob ious i hey ep esen a single specimen
76 K.R. Tabachnick e al.
Fig. 16. Specimens o Caulophacus (Caulophacus)
hype bo eus a ached o a dead skele on o Pe i-
ph agella (no isible he e). A ROV ‘Comanche’
pho o. RV ‘Akademik M.A. La en ye ’ – 75, s a.
22, 55°30′36.1″ N 167°19′27.1″ E, dep h 3879–
3578 m.
Рис. 16. Экземпляры Caulophacus (Caulophacus)
hype bo eus прикрепленные к мертвому скелету
Pe i agella (не виден). Фотография подводного
робота “Comanche”; НИС “Академик М.А. Лав-
рентьев” – 75, станция 22, 55°30′36.1″ N 167°
19′27.1″ E, глубина 3879–3578 м.
Fig. 17. Specimens o Caulophacus (Caulophacus) hype bo eus: A — IORAS 5/2/2232. B — IORAS 5/
2/2250, iew om di e en sides. C — IORAS 5/2/2661, iew om di e en sides. Scale 10 mm.
Рис. 17. Экземпляры Caulophacus (Caulophacus) hype bo eus: A — IORAS 5/2/2232. B — IORAS 5/
2/2250, вид с разных сторон. C — IORAS 5/2/2661, вид с разных сторон. Масштаб 10 мм.
77Rossellidae om he Be ing Sea
Fig. 18. Spicules o Caulophacus (Caulophacus) hype bo eus, IORAS 5/2/2229 (addi ional ypes o
p e ious desc ip ions). A — de mal pinula hexac in. B — de mal pinula pen ac in. C — a ial pinula
hexac in. D — a ial pinula pen ac in.
Рис. 18. Спикулы Caulophacus (Caulophacus) hype bo eus, IORAS 5/2/2229 (в дополнение к преды-
дущим описаниям). A — дермальная пинулярная гексактина. B — дермальная пинулярная пентак-
тина. C — атриальная пинулярная гексактина. D — атриальная пинулярная пентактина.
wi h s olonial b anching o mul iple o ganisms o
same species ha se led on he peduncle o one
colony. The specimen sizes anged om hose wi h
he disc 3 mm in diame e and 1 mm in hickness wi h
peduncle 1 mm in diame e o he bigges ones 55
mm in diame e , 3–4 mm in hickness and he pedun-
cle abou 5 mm in diame e . Bu some o hem each
e en la ge sizes — a single lamella e agmen o he
discoidal body 90x70 mm and 4–6 mm in hickness
was ound.
REMARKS. The species was o iginally desc ibed
as a subspecies: Caulophacus schulzei hype bo eus
Kol un, 1967. Bo h species om he No h Paci ic,
C. (Caulophacus) schulzei and C. (Caulophacus)
elegans, a e comple ely di e en om he subspe-
cies desc ibed by Kol un (1967).
Among he newly ound specimens o Caulo-
phacus hype bo eus many had pinula pen ac ins
among bo h de mal and a ial hexac ins some imes
in la ge amoun s. As i is seen om he Suppl. Tab.
12, i can be assumed ha spicule combina ion and
hei sizes a e e y simila be ween in es iga ed
specimens and dissimila i ies a e conside ed as an
ine speci ic a ia ion. Unlike specimens desc ibed
by Kol un, some specimens om he new se ies om
he Be ing Sea ha e de mal and a ial pinula pen ac-
ins wi h hei udimen a y ay di ec ed inside he
body, in some specimens hey a e e en mo e nume -
ous hen co esponding hexac ins ( o ins ance, IO-
RAS5/2/2229; 2232 and some o he specimens).
Mean ime, he o he spicules co espond in shape
and size o ypical ep esen a i es and hese speci-
mens a e undoub edly belonging o he same species.
DISTRIBUTION. Be ing and Okho sk Seas,
3400–4294 m dep h.
Caulophacus (Caulophacus) mi i sp.n.
Figs 19, 20.
MATERIAL. HOLOTYPE. IORAS 5/2/2054.
RV ‘Akademik Ms isla Keldysh’ – 22, s a. 2296,
HOV ‘Mi -1’, Paci ic side o Be ing Island, 54°57.7′
N 165°42.85′ E, 5952 m.
DESCRIPTION. BODY. The specimen is un-
gus-like wi h a discoidal o oid body 140x190 mm
and 20 mm hick, he edge o he disc is e e ed down
so ha he a ial su ace p esen also on he la e al
edge o he discoidal body. The ubula peduncle was
b oken in o se e al pieces du ing collec ion bu was
180 mm long wi h walls 5 mm in hickness. The
uppe pa o he peduncle is 20x25 mm in diame e
while he lowe pa is abou 20 mm in diame e .
SPICULES. MEGASCLERES. Choanosomal
spicules a e diac ins and hexac ins. The diac ins a e
some imes hick bu mos we e hin. The hick diac-
ins ha e s ou sha s and ounded ou e ends, hey
a e 2–2.7/0.032–0.110 mm. The hin diac ins ha e
s ou sha s, some imes wi h a widening in he mid-
dle o ou ube cle udimen s, hei ou e ends a e
ounded, smoo h o ough, hey a e 1.5–1.9/0.008–
0.024 mm. Choanosomal hexac ins ha e ays 0.4–
1.6/0.015–0.1 mm wi h conically poin ed ou e ends.
Hypode malia and hypoa ialia a e pen ac ins wi h
conically poin ed ou e ends. Tangen ial ays o
hese pen ac ins a e 0.2–1.4 mm long, he ay di ec -
ed inside he body 0.1–0.8 mm long, hei diame e
78 K.R. Tabachnick e al.
Fig. 19. Caulophacus (Caulophacus) mi i sp.n., holo ype, ex e nal shape om di e en sides. Scale 50 mm.
Рис. 19. Caulophacus (Caulophacus) mi i sp.n., голотип, внешний вид сразных сторон. Масштаб 50 мм.
mm), he diame e o pinula ay is 0.019–0.033 mm
a base and 0.041–0.063 mm in maximal media e
pa , he diame e o o he ays is 0.015–0.019 mm a
base. The pinula ay in a ial hexac ins may also be
sphe ical bu o en hey a e elonga e, o en wi h
i egula sha ha ing a widening some dis ance
om he base. Pinula ay o a ial hexacins is 0.074–
0.463 mm in diame e (n=30, a g: 0.226 mm, s d:
0.135 mm), angen ial ays a e 0.078–0.152 mm in
diame e (n=30, a g: 0.114 mm, s d: 0.018 mm), he
ay di ec ed inside he body is 0.078–0.152 mm in
diame e (n=30, a g: 0.105 mm, s d: 0.014 mm), he
diame e o pinula ay is 0.019–0.026 mm a base
and 0.022–0.052 mm in maximal media e pa , he
diame e o o he ays is abou 0.015 mm a base.
MICROSCLERES. Mos mic oscle es a e hick-
ayed discohexac ins, bu he e a e a e discohemi-
hexas e s and some imes discohexas e s wi h 2–4
seconda y ays. Lophodiscohexas e s wi h 4–12 sec-
onda y ays a e common close o he de mal and
a ial su aces. Some o hese spicules ha e s ou
sha s, some ha e some seconda y ays aised om
he side o he p ima y ays, a ely lophodiscos au-
is 0.02–0.10 mm. Speci ic ypes o la ge hexac ins
a e loca ed benea h he a ial su ace, each ha ing a
sho ened ay di ec ed ou side he body wi h ough
o sho spiny ounded dis al ends. Some imes his
ay is lanceola e in shape, and hey do no p o ude
o e he en i e a ial su ace. This ou wa dly p o-
uding ay is abou 0.6 mm long, whe eas angen ial
ays a e 1,3 mm long. The ay di ec ed inside he
body is abou 0.8 mm long and he diame e o hese
ays is 0.04–0.05 mm. The diac ins o he peduncle
ha e a diame e o 0.015–0.053 mm and a e connec -
ed o each o he by nume ous synap icula junc ions.
De malia and a ialia a e pinula hexac ins, hey
ha e ounded o conically poin ed ou e ends, an-
gen ial ays and he ay di ec ed inside he body a e
sho -spiny. Some a e de mal pen ac ins ha e no
pinula ay; he unpai ed ay is equal in shape o
o he ays. Pinula ay o de mal hexacins is sphe -
ical, 0.067–0.107 mm in diame e (n=25, a g: 0.085
mm, s d: 0.011 mm), angen ial ays a e 0.074–0.141
mm in diame e (n=25, a g: 0.104 mm, s d: 0.016
mm), he ay di ec ed inside he body is 0.056–0.111
mm in diame e (n=25, a g: 0.089 mm, s d: 0.015
79Rossellidae om he Be ing Sea
Fig. 20. Spicules o Caulophacus (Caulophacus) mi i sp.n., holo ype. A — hick choanosomal diac in. B–
D — choanosomal diac ins. E — choanosomal hexac in. F — hypode mal o hypoa ial pen ac in. G —
hypoa ial hexac in. H — de mal pinula hexa in. I — de mal hexac in. J–M — a ial pinula hexac ins. N–
O — discohexas e s. P — abno mal discohexas e . Q–R — discohemihexas e s. S — abno mal discohexas e .
T–V — lophodiscohexas e s. W — lophodiscos au as e . X — abno mal s au ac in.
Рис. 20. Спикулы Caulophacus (Caulophacus) mi i sp.n., голотип. A — толстая хоаносомальная
диактина. B–D — хоаносомальные диактины. E — хоаносомальная гексактина. F — гиподермальная
или гипоатриальная пентактина. G — гипоатриальная гексактина. H — дермальная пинулярная
гексактина. I — дермальная гексактина. J–M — атриальные пинулярные гексактины. N–O — дискогек-
састры. P — аномальный дискогексастр. Q–R — дискогемигексастры. S — аномальный дискогексастр.
T–V — лофодискогексастры. W — лофодискостаурастер. X — аномальная стаурактина.
wi h he diame e o hei p ima y ose e being
0.023–0.063 mm (n=25, a g: 0.037 mm, s d: 0.010
mm).
REMARKS. In compa ison o o he species in
his subgenus, he new species has a no able and
signi ican di e ence in he sizes be ween de mal
and a ial spicules. This ea u e is obse ed also in
he ype species — Caulophacus (Caulophacus)
as e s a e ound. The diame e o discohexac in is
0.089–0.185 mm (n=27, a g: 0.135 mm, s d: 0.032
mm). The hick– ayed discohexas e s a e 0.059–
0.104 mm in diame e (n=9, a g: 0.078 mm, s d:
0.013 mm), he diame e o he p ima y ose e is
0.036–0.044 mm (n=9, a g: 0.039 mm, s d: 0.003
mm). The lophodiscohexas e s a e 0.036–0.101 mm
in diame e (n=25, a g: 0.067 mm, s d: 0.016 mm),
80 K.R. Tabachnick e al.
la us Schulze, 1886 ( e-desc ibed la e : Schulze,
1887; Tabachnick, 2002b). Unlike i , he new spe-
cies has de mal hexac ins wi h sphe ical shape o
pinula heads; a ial hexac ins wi h widened pinula
ays and o en hey ha e i egula o e all shape o
he pinula ay when i is hickes a base, close o he
cen e o he spicule. Mic oscle es sizes a e also
qui e di e en be ween hese species. Caulophacus
(Caulophacus) la us has calycocomes o lophodis-
cohexas e s which a e ex emely la ge. O he spe-
cies o Caulophacus (Caulophacus) which also ha e
la ge pinula ay in a ial hexac ins and pen ac ins
ca y sho spines on his ay.
ETYMOLOGY. The species is named a e he
HOV ‘Mi ’ which collec ed his specimen.
DISTRIBUTION. Cu en ly ound only o he
Paci ic side o Be ing Island, a 5952 m dep h.
Caulophacus (Caulophacus) suba c icus
sp.n.
Figs 21, 22; Suppl. Tab. 13.
MATERIAL. HOLOTYPE: IORAS 5/2/2274:
RV ‘Akademik Ms isla Keldysh’ – 22, s a. 2306,
awl, Paci ic sine o he Be ing Island, 54°57.1–
57.7′ N 165°49.9–51.0′ E, 4401–3797 m.
DESCRIPTION. BODY. The specimen is un-
gus–like wi h a discoidal body 25 mm in diame e
and 1–2 mm in hickness. The peduncle is abou 1.5
mm in diame e and 35 mm long.
SPICULES. MEGASCLERES. Choanosomal
spicules a e diac ins and hexac ins. The diac ins a e
di ided in o wo ypes: la ge and small. The o me
(abou 1.5/0.02 mm) ha e s ou sha s, conically
poin ed ou e ends. Common diac ins a e hin wi h a
widening in he middle, he ou e ends a e ough,
ounded o cla a e, hese diac ins a e 0.9–2.4/0.006–
0.009 mm. Choanosomal hexac ins ha e conically
poin ed ou e ends, hei ays a e 0.6–0.8/0.02–0.04
mm. Hypode mal and hypoa ial pen ac ins ha e
smoo h angen ial ays o hey a e spiny a base, he
ay di ec ed inside he body may be spiny, he ou e
ends a e smoo h, conically poin ed. The angen ial
ays o hypode mal and hypoa ial pen ac ins a e
0.3–0.5 mm long, he ay di ec ed inside he body is
0.6–1.0 mm, he diame e o hese ays is 0.02–0.03
mm. Spicules o he peduncle a e diac ins abou 8/
0.006–0.023 mm wi h s ou sha s, ounded o cla -
a e ou e ends. Unlike he penduncula diac ins o
o he species, hese do no use o each o he .
De malia a e hexac ins as well as a e pen ac ins
wi h a udimen al ube cle ins ead o a six h ay
di ec ed inside he body. De mal hexac ins include
hose wi h cla a e pinula ays and long angen ial
ays as well as hose wi h a lanceola e pinula ay.
The angen ial ays and he ay di ec ed inside he
body a e ough o sho -spiny a hei dis al hal wi h
conically poin ed ou e ends. The cla a e pinula ay
o de mal hexac in is 0.037–0.141 mm long (n=25,
a g: 0.079 mm, s d: 0.023 mm), maximal diame e o
his ay is 0.011 mm, angen ial ays a e 0.041–0.107
mm long (n=25, a g: 0.087 mm, s d: 0.015 mm), he
ay di ec ed inside he body is 0.030–0.093 mm long
(n=25, a g: 0.066 mm, s d: 0.018 mm), he diame e
o hese ays is 0.006 mm. Lanceola e pinula ay o
de mal hexac in is 0.070–0.159 mm long (n=25,
a g: 0.123 mm, s d: 0.020 mm), he maximal diam-
e e o his ay is 0.004 mm, angen ial ays o hese
spicules a e 0.052–0.081 mm long (n=25, a g: 0.064
mm, s d: 0.010 mm), he ay di ec ed inside he body
is 0.033–0.078 mm long (n=25, a g: 0.059 mm, s d:
0.011 mm), he diame e o hese ays is abou 0.003
mm. A ialia a e pen ac ins wi h udimen al ube cu-
la ed ay di ec ed inside he body o some imes
hexac ins, hese spicules ha e pinula ay elonga e
Fig. 21. Caulophacus (Caulophacus) suba c icus
sp.n., holo ype, ex e nal shape. Scale 10 mm.
Рис. 21. Caulophacus (Caulophacus) suba c icus
sp.n., голотип, внешний вид. Масштаб 10 мм.
81Rossellidae om he Be ing Sea
Fig. 22. Spicules o Caulophacus (Caulophacus) suba c icus sp.n., holo ype. A–B — de mal pinula
hexac ins. C — de mal pinula pen ac in. D–E — a ial pinula pen ac ins. F — hypode mal o hypoa ial
pen ac in. G — choanosomal hexac in. H–I — choanosomal diac ins. J — hick- ayed discohexac in. K —
hin– ayed discohexac in. L — discohexas e . M — in e medium spicule be ween discohexas e and
lophodiscohexas e . N — lophodiscohexas e .
Рис. 22. Спикулы Caulophacus (Caulophacus) suba c icus sp.n., голотип. A–B — дермальные пинуляр-
ные гексактины. C — дермальная пинулярная пентактина. D–E — атриальные пинулярные пентак-
тины. F — гиподермальная или гипоатриальная пентактина. G — хоаносомальная гексактина. H–I —
хоаносомальные диактины. J — дискогексактина с толстыми лучами. K — дискогексактина с
тонкими лучами. L — дискогексастер. M — спикула промежуточная между дискогесастером и
лофодискогексастером. N — лофодискогексастер.
88 K.R. Tabachnick e al.
icipa ed in he collec ion o he desc ibed ma e ials.
We highly app ecia e he e iewe s’ insigh ul and
help ul commen s on ou manusc ip .
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Responsible edi o A.V. E esko sky
