On the taxonomic position of Phaenomenella Fraussen & Hadorn, 2006 (Neogastropoda, Buccinoidea) with description of two new species

Di ec eu De la publica ion : B uno Da id
P ésiden du Muséum na ional d’His oi e na u elle
éDac ice en che / Edi o -in-chiE : Lau e Desu e -G andcolas
assis an s De éDac ion / Assis An Edi o s : Anne Mabille ([email p o ec ed])
Mise en page / PAgE lAyou : Anne Mabille
coMi é scien i ique / sciEn i ic boA d :
James Ca pen e (AMNH, New Yo k, É a s-Unis)
Ma ia Ma a Cigliano (Museo de La Pla a, La Pla a, A gen ine)
Hen ik Engho (NHMD, Copenhague, Danema k)
Ra ael Ma quez (CSIC, Mad id, Espagne)
Pe e Ng (Uni e si y o Singapo e)
No man I. Pla nick (AMNH, New Yo k, É a s-Unis)
Jean-Y es Rasplus (INRA, Mon e ie -su -Lez, F ance)
Jean-F ançois Sil ain (IRD, Gi -su -Y e e, F ance)
Wanda M. Weine (Polish Academy o Sciences, C aco ie, Pologne)
John Wenzel (The Ohio S a e Uni e si y, Columbus, É a s-Unis)
cou e u e / co E :
Shells o Phaenomenella nicoi n. sp.
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Kan o Y., Kosyan A., So okin P. & Fedoso A. 2020— On he axonomic posi ion o Phaenomenella F aussen &
Hado n, 2006 (Neogas opoda, Buccinoidea) wi h desc ip ion o wo new species. Zoosys ema 42 (3): 33-55. h ps://
doi.o g/10.5252/zoosys ema2020 42a3. h p://zoosys ema.com/42/3
Abs Ac
This con ibu ion p o ides no el in o ma ion on he ana omy, adula and phylogeny o se e al species
o Phaenomenella F aussen& Hado n, 2006, a genus o Buccinoidea Ra inesque, 1815 wi h unclea
a ini ies. Molecula phylogene ic analysis based on sequences o mi ochond ial COI and nuclea 28S
RNA genes o di e en ep esen a i es o Buccinoidea e ealed close ela ionships o Phaenomenella
wi h Siphonalia A.Adams, 1863 bo h axa o ming a clade wi h maximal suppo . The ana omy o
wo species o he la e genus was examined o he i s ime o compa a i e pu poses. The sub amily
Siphonaliinae Finlay, 1928 was e ec ed o se e al Recen and ossil gene a o Sou he n Hemisphe e
Buccinidae Ra inesque, 1815, and is s ill ecognized by cu en axonomis s (Bouche e al. 2017).
Species o all Recen gene a o Siphonaliinae we e included in ou analysis and he monophyly o he
sub amily Siphonaliinae in i s o iginal scope is ejec ed. Molecula and mo phological da a e ealed
wo s ill unnamed species o Phaenomenella om he lowe ba hyal zone o he Sou h China Sea.
These species, Phaenomenella nicoi n.sp. and P. samadiae n.sp. a e desc ibed in he p esen s udy.
Yu i KANTOR
Alisa KOSYAN
Pa el SOROKIN
Alexande FEDOSOV
A.N. Se e so Ins i u e o Ecology and E olu ion o Russian Academy o Sciences,
Leninski p ospec 33, 119071 Moscow, Russian Fede a ion (Russia)
kan o [email p o ec ed]
[email p o ec ed]
[email p o ec ed]
[email p o ec ed]
Submi ed on 28 Janua y 2019 | Accep ed on 8 Ap il 2019 | Published on 4 Feb ua y 2020
On he axonomic posi ion o Phaenomenella F aussen &
Hado n, 2006 (Neogas opoda, Buccinoidea) wi h desc ip ion
o wo new species
Key wo ds
Molecula phylogeny,
COI,
28S,
classi ica ion,
Siphonaliinae,
new species.
34 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
IN ROduC ION
The genus Phaenomenella F aussen& Hado n, 2006, wi h
he ype species Mana ia in la a Shikama, 1971, was es abli-
shed o h ee species o “Buccinidae” Ra inesque, 1815 om
aiwan and Sou h China Sea (F aussen& Hado n 2006).
Se e al addi ional species we e desc ibed since and p esen ly
he genus includes nine species, all excep one om o Sou h
Eas Asia (F aussen 2008; F aussen& S ahlschmid 2012;
F aussen& S ahlschmid 2013). The ana omy o he genus
has ne e been examined, bu he adula was illus a ed o
wo species, Phaenomenella in la a (Shikama, 1971) and Phae-
nomenella angus a F aussen& Hado n, 2006. The adula is
o gene al buccinid appea ance, which gi es no clues o he
ela ionships o Phaenomenella o o he buccinid gene a. The
posi ion o he genus wi hin Buccinidae was no add essed
in p e ious publica ions, p obably due o s ill unesol ed in-
a amilia classi ica ion o he amily. The in agene ic shell
a iabili y o Phaenomenella is high ha is hampe ing p o id-
ing eliable gene ic diagnosis.
Among Sou h-Eas Asian buccinids se e al gene a bea
some conchological esemblance o Phaenomenella, ie.,
Mana ia Smi h, 1906, Eosipho Thiele, 1929, Gaillea Kan-
o , Puilland e, F aussen, Fedoso & Bouche , 2013 (all
h ee gene a dwelling on biogenic subs a es; Kan o e al.
2013) and Siphonalia A.Adams, 1863. Buccinidae om
biogenic subs a es cons i u e a clade ha is well-suppo ed
by molecula da a, and a e cha ac e ized by bicuspid la e al
ee h (Bouche & Wa én 1986; Kan o e al. 2013), while
he adula o bo h Phaenomenella and Siphonalia is simila
and has icuspid la e al ee h.
Rela ionships o Siphonalia wi h o he Buccinidae a e no
clea . I s isola ed posi ion was ecognized by Finlay (1928),
who p oposed a new sub amily Siphonaliinae in he newly
es ablished amily Buccinulidae. Ha ing been p oposed wi h-
ou diagnosis o discussion, he sub amily o iginally included
se e al Recen and ossil gene a, which we e la e synonymized
wi h Penion Fische , 1884 (=†Aus osipho Cossmann, 1906,
Ve conella I edale, 1914, Be ylsma I edale, 1924), Aenea-
o (=†Ellicea Finlay in Ma wick 1928, †Pi ella Ma wick,
1928), as well as Glaphy ina Finlay, 1926 (p esen ly a ib-
u ed o Fasciola iidae G ay, 1853; Cou o e al. 2016) and
†Pomahakia Finlay, 1927. All gene a excep he ype one a e
con ined o sou he n hemisphe e, mos ly o Aus alian-New
Zealand egion. The alidi y and scope o he sub amily ha e
no been e ised, al hough i is ecognized in he cu en ax-
onomy (Bouche e al. 2017). A ecen molecula phylogeny
o some sou he n hemisphe e Buccinulinae based on whole
mi ochond ial genome and nuclea ibosomal sequence da a
(Vaux e al. 2017) e ealed ha Recen gene a included by
Finlay in o Siphonaliinae do no cons i u e a monophyle ic
g oup. Bu Siphonalia i sel was no included in he analysis
and he e o e i s ela ionships emained un esol ed.
In he cou se o expedi ions o ganized by he Muséum na-
ional d’His oi e na u elle, Pa is (MNHN) o he Sou h China
Sea se e al specimens o di e en species o Phaenomenella
and one species o Siphonalia we e collec ed and p ese ed o
dNA sequencing. The examina ion o his ma e ial e ealed
wo s ill unnamed species. The pu pose o he p esen pape
is o p o ide o mal desc ip ion o he e ealed new species,
and o cla i y ela ionships o Phaenomenella based bo h on
ana omy and on molecula da a o ex ended da ase o Buc-
cinoidea, including g oups ha a e conchologically simila
o Phaenomenella.
MA ERIAL ANd ME HOdS
The ma e ial was collec ed mos ly in he esea ch c uises in he
Sou h China Sea (dONGSHA 2014, ZHONGSHA 2015)
and in he Philippines (AuRORA 2007) ha we e o ganized
espec i ely by he Na ional aiwan uni e si y and he Mu-
ÉsUMÉ
Posi ion axonomique de Phaenomenella F aussen& Hado n, 2006 (Neogas opoda: Buccinoidea) e
desc ip ion de deux nou elles espèces.
Ce e con ibu ion ou ni de nou elles in o ma ions su l’ana omie, la adula e la phylogénie de
plusieu s espèces de Phaenomenella F aussen& Hado n, 2006, un gen e de Buccinoidea Ra inesque,
1815 aux a ini és peu clai es. une analyse phylogéné ique moléculai e basée su des séquences des
gènes mi ochond iaux COI e d’ARN 28S nucléai de di é en s ep ésen an s de Buccinoidea a
é élé des ela ions é oi es de Phaenomenella a ec Siphonalia A.Adams, 1863 qui o men un clade
a ec un sou ien maximal. L’ana omie de deux espèces de ce de nie gen e a é é examinée pou la
p emiè e ois à des ins de compa aison. La sous- amille Siphonaliinae Finlay, 1928 a é é é igée pou
plusieu s gen es écen s e ossiles de Buccinidae Ra inesque, 1815 de l’hémisphè e sud, e es oujou s
econnue pa les axonomis es ac uels (Bouche e al. 2017). Les espèces de ous les gen es écen s de
Siphonaliinae on é é incluses dans no e analyse e la monophylie de la sous- amille Siphonaliinae
dans sa dé ini ion o iginelle es eje ée. Les données moléculai es e mo phologiques on é élé deux
espèces enco e inconnues de Phaenomenella de la zone ba hyale in é ieu e du sud de la me de Chine.
Ces espèces, Phaenomenella nicoi n.sp. e P. samadiae n.sp. son déc i es dans la p ésen e é ude.
Mo s clÉs
Phylogénie moléculai e,
COI,
28S,
classi ica ion,
Siphonaliinae,
espèces nou elles.
35
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
séum na ional d’His oi e na u elle, Pa is (MNHN) wi h he
Philippines Bu eau o Fishe ies and Aqua ic Resou ces (BFAR).
Specimens collec ed we e p ocessed wi h a mic owa e o en
(Galindo e al. 2014): he li ing molluscs in small olumes o
sea wa e we e exposed o mic owa es o 10-30 s, depending
on specimen size. Bodies we e immedia ely emo ed om shells
and d opped in 96% e hanol. Specimens p ocessed in his way
a e sui able o u he ana omical s udies a e soaking hem
in 70% e hanol. Specimens a e egis e ed in he MNHN col-
lec ion and specimens and sequences a e deposi ed in BOLd
(Ba code o Li e da asys em) and GenBank ( able1).
due o echnical easons, he sequenced specimens o Sipho-
nalia spadicea (Ree e, 1847) we e no a ailable o ana omical
examina ion and he ma e ial (no sui able o sequencing) on
wo o he species o Siphonalia was used o ana omical s udy.
Dna ex ac ion and pc
o al dNA was ex ac ed om he piece o oo using ei he
he dNeasy96 issue ki o In es iga o Ki (Qiagen), ol-
lowing he manu ac u e ’s ecommenda ions. The ba code
agmen o he Cy och ome Oxidase I (COI) gene (658bp)
and a 28S RNA agmen we e ampli ied using he uni e sal
p ime s LCO1490 and HCO2198 (Folme e al. 1994) and
C1 and d2 (Jo elin& Jus ine 2001), espec i ely. PCRs we e
pe o med in 20µl inal olume con aining app oxima ely
3ng empla e dNA, 1.5mM MgCl2, 0.26mM o each
nucleo ide, 0.3µl o each p ime , 5% dMSO and 0.75µl
o ei he aq Polyme ase (Qbiogene) o BioHY aq dNA
polyme ase (diala ).
The PCR p o ile o he COI s a ed wi h 5min a 95°C ol-
lowed by 40cycles wi h he dena u a ion a 95°C (35sec),
annealing a 50°C (35s) and elonga ion a 72°C (1min), wi h
inal elonga ion phase a 72°C (10min). Simila PCR p o iles
we e se o 28S (annealing a 56°C). COI and 28S genes we e
sequenced in bo h di ec ions o con i m accu acy o each se-
quence. The sequencing was pe o med by Eu o ins o in he
SIEE RAS molecula acili y on an ABI 3500 Gene ic analyse .
Mo phology and adula
Radulae we e cleaned using dilu ed bleach (NaOCl), ai –
d ied, coa ed wi h gold and examined by scanning elec on
mic oscope eScan S5130MM in he Ins i u e o Ecology
and E olu ion o Russian Academy o Sciences, Moscow
(IEE RAS). Ana omy was examined on manual dissec ions.
phylogene ic analysis
COI and 28S sequences we e aligned using Clus alW im-
plemen ed in BioEdi . 7.0.9.0 (Hall 1999); he accu acy o
each alignmen was checked by eye and i needed modi ied.
COI and 28S sequences o addi ional 32 buccinoidean axa,
mainly om he da ase s o Kan o e al. (2013) and Vaux
e al. (2017) we e accessed om GenBank.
Th ee da ase s we e analyzed, a single-gene COI da ase
(57 axa) wi h h ee codon posi ions coded as h ee indepen-
den pa i ions, a nuclea 28S (45 axa) as single pa i ion,
and a conca ena ed COI–28S da ase (45 axa), whe e ou
pa i ions co esponded o h ee codon posi ions o COI and
o he 28S agmen espec i ely. The single gene da ase s we e
mainly used o e alua e p ima y species hypo heses (PSHs),
p oposed based on he shell mo phology, whe eas he analysis
o conca ena ed da ase allowed us o es ima e ela ionships
o Siphonalia and emaining gene a o iginally included in
Siphonaliinae. The lineage comp ising buccinids om he
biogenic subs a es (Kan o e al. 2013) was used o oo he
ee based on he opology o Buccinidae ee eco e ed by
Galindo e al. (2016). In he RAxML analyses (S ama akis
2006) obus ness o nodes was assessed using he Tho ough
Boo s apping algo i hm (Felsens ein 1985) wi h 1000i e a-
ions. The Bayesian in e ence analyses (BI) we e pe o med
using M Bayes (Huelsenbeck& Ronquis 2001), unning
wo pa allel analyses, consis ing each o six Ma ko chains
o 20 000 000gene a ions wi h de aul pa ame e s. Pa a-
me e s o he subs i u ion model we e es ima ed du ing he
analysis (sixsubs i u ion ca ego ies, a gamma-dis ibu ed a e
a ia ion ac oss si es app oxima ed in ou disc e e ca ego ies
and a p opo ion o in a iable si es). The ees om he i s
5 000 000gene a ions (25% om o al numbe o gene a-
ions) we e disca ded as bu n-in p io o he calcula ion o
consensus ees. Con e gence o each analysis was e alua ed
using ace 1.4.1 (Rambau e al. 2014) o check ha all
ESS alues exceeded 200 (wi h de aul bu ning). All analyses
we e pe o med on he Cip es Science Ga eway (h p://www.
phylo.o g/po al2), using M Bayes on XSEdE (3.2.6) and
RAxML-HPC2 on XSEdE (8.2.10) (Mille e al. 2010). The
ma ix o K2P pai wise gene ic dis ances o COI, was calcu-
la ed o he Phaenomenella and Siphonalia axa in MEGA 6
( amu a e al. 2013) ( able2).
abb e ia ions and con en ions
adg opening o an e io duc o diges i e gland;
AL ape u e leng h;
ao an e io ao a;
aoe an e io oesophagus;
ba buccal a e y;
bm buccal mass;
bn buccal ne es;
cep. cephalic en acles;
cg capsule gland;
cm columella muscle;
c c enidium;
dd dead shell;
dg diges i e gland;
eye eye;
oo ;
gl gland o Leiblein;
gon gonad;
hd head;
hg hypob anchial gland;
in in es ine;
kd kidney;
l l longi udinal old on inne s omach wall;
l li e collec ed specimen;
m medial e ac o o adula;
n ne es;
n ne e ing;
od odon opho e;
odn odon opho e ne es;
od odon opho e e ac o s;
oeo oesophageal opening;

36 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
Table 1
. — Lis o sequenced ma e ial used in phylogene ic econs uc ions. Exac collec ion locali ies o specimens om MNHN (specimen code s a s wi h MNHN-
IM-) can be ound a MNHN si e a h ps://science.mnhn. /ins i u ion/mnhn/collec ion/im/i em/****-***** (as e isks co espond o he digi al numbe o specimen).
Accession numbe s
Specimen
code Genus species Collec ion da a BOLD
Genbank
(COI)
Genbank
(28S) Sou ce
IM-2007-32673 Siphonalia spadicea TAIWAN 2004, s . CP264 NEOGA544-10 MK567642 – P esen s udy
IM-2007-32674 spadicea TAIWAN 2004, s . CP264 NEOGA545-10 MK567658 – P esen s udy
IM-2007-32762 spadicea TAIWAN 2001 s , CP79 NEOGA589-10 MK567643 – P esen s udy
IM-2007-32856 spadicea TAIWAN 2004, s . CP244 NEOGA601-10 MK567646 – P esen s udy
IM-2007-32995 spadicea TAIWAN 2001 s , CP109 NEOGA666-10 MK567652 – P esen s udy
IM-2007-32997 spadicea TAIWAN 2001 s , CP109 NEOGA668-10 MK567664 – P esen s udy
IM-2007-32996 spadicea TAIWAN 2001 s . CP109 NEOGA667-10 MK567656 MK567635 P esen s udy
IM-2009-11271 spadicea TAIWAN 2004 s . CH257 BUCC001-19 MK567649 MK567632 P esen s udy
IM-2013-41068 Phaenomenella c hachi TAIWAN 2013 s . CP4090 BUCC007-19 MK567653 – P esen s udy
IM-2013-41072 in la a TAIWAN 2013 s . CP4090 BUCC008-19 MK567650 – P esen s udy
IM-2013-41073 in la a TAIWAN 2013 s . CP4090 BUCC009-19 MK567660 – P esen s udy
IM-2013-50012 c hachi DONGSHA 2014 s . CP4120 BUCC006-19 MK567654 MK567634 P esen s udy
IM-2013-50203 insulap a asensis DONGSHA 2014 s . CP4129 BUCC002-19 MK567648 MK567631 P esen s udy
IM-2013-50204 insulap a asensis DONGSHA 2014 s . CP4129 BUCC003-19 MK567661 – P esen s udy
IM-2013-50205 insulap a asensis DONGSHA 2014 s . CP4129 BUCC004-19 MK567655 – P esen s udy
IM-2013-50260 c callopho ella DONGSHA 2014 s . CP4130 BUCC005-19 MK567665 MK567639 P esen s udy
IM-2013-61674 samadiae n. sp.
ZHONGSHA 2015 s . CP4134
BUCC013-19 MK567662 MK567638 P esen s udy
IM-2013-61617 samadiae n. sp.
ZHONGSHA 2015 s . CP4133
BUCC015-19 MK567644 MK567628 P esen s udy
IM-2007-34644 nicoi n. sp.
AURORA 2007 s . CP2685
NEOGA783-10 MK567663 – P esen s udy
IM-2013-59398 nicoi n. sp.
ZHONGSHA 2015 s . CP4157
BUCC016-19 MK567645 MK567629 P esen s udy
IM-2013-61585 nicoi n. sp.
ZHONGSHA 2015 s . CP4132
BUCC014-19 MK567641 MK567627 P esen s udy
IM-2013-61637 nicoi n. sp.
ZHONGSHA 2015 s . CP4133
BUCC010-19 MK567647 MK567630 P esen s udy
IM-2013-61639 nicoi n. sp.
ZHONGSHA 2015 s . CP4133
BUCC011-19 MK567651 MK567633 P esen s udy
IM-2013-61673 nicoi n. sp.
ZHONGSHA 2015 s . CP4134
BUCC012-19 MK567659 MK567637 P esen s udy
IM-2007-34639 nicoi n. sp. AURORA 2007 s . CP2685 NEOGA781-10 MK567657 MK567636 P esen s udy
IM-2007-34635 Calag asso alde menensis AURORA 2007 s . CP2673 KC756037 KC755997
Kan o e al. 2013
IM-2007-32864 Mana ia clandes ina PANGLAO 2005 s . CP2389 KC756058 KC756004
Kan o e al. 2013
IM-2007-32952 b e icauda SALOMON 2 s . CP2219 KC756055 KC756003
Kan o e al. 2013
IM-2009-7079 Enigma icolus monnie i MIRIKY s . CP3279 KC756034 KC755996
Kan o e al. 2013
20140783 Buccinum unda um Reykjanesskagi, Iceland MK558051 MK543285 Vaux e al. 2017
KK12 Kelle ia kelle ii
San a Ba ba a, Cali o nia, USA
MH198161 MH277543 Vaux e al. 2017
KL2 lischkei
Kansai, Mie P e ec u e, Japan
MH198160 MH277544 Vaux e al. 2017
SFKH-TMP005 Pa a e i usus ca ina us Cha ham Rise, NZ MK583342 MK543289 Vaux e al. 2017
M.190082/2 Penion cha hamensis Cha ham Rise, NZ MH140429 MH277545 Vaux e al. 2017
M.190085 cha hamensis Cha ham Rise, NZ MH140428 MH277546 Vaux e al. 2017
M.183792/1 cu ie ianus Red Me cu y Island, NZ MH140431 MH277548 Vaux e al. 2017
M.183927 cu ie ianus Co omandel, NZ MH140432 MH277549 Vaux e al. 2017
C.456980 manda inus
Gabo Island, Vic o ia, Aus alia
MG211145 MH277553 Vaux e al. 2017
C.487648 maximus Te igal, NSW, Aus alia MG211144 MH277554 Vaux e al. 2017
Phoenix1 ai ieldae O ago Peninsula, NZ MH198165 MH277547 Vaux e al. 2017
Phoenix9 sulca us Auckland, NZ MG098232 MG194426 Vaux e al. 2017
M.274111 Aenea o ben hicolus Cape Pallise , NZ MK577960 MK543272 Vaux e al. 2017
SFKH-TMP015 elegans Cha ham Rise, NZ MH198157 MH277534 Vaux e al. 2017
M.279437 o agoensis Tasman Bay, NZ MK577961 MK543274 Vaux e al. 2017
M.190119 ecens Cape Tu nagain, NZ MH198159 MH277535 Vaux e al. 2017
SFKH-TMP013 aledic us TAN 616/83, NZ MK577962 MK543276 Vaux e al. 2017
M.183832
An a c onep unea
ben hicola Cha ham Rise, NZ MH198156 MH277537 Vaux e al. 2017
SFKH-TMP014 Aus o usus glans Island Bay, Welling on, NZ MK558053 MK543277 Vaux e al. 2017
M.302907/2 Buccinulum uscozona um A iel Bank, Gisbo ne, NZ MH198158 MH277540 Vaux e al. 2017
M.258277/6 pallidum S ewa Island, NZ MK577963 MK543280 Vaux e al. 2017
M.302870/2 pe inax inlayi Poin Gibson, NZ MH198162 MH277541 Vaux e al. 2017
M.314755/1 obus um
One oa Bay, Bay o Islands, NZ
MK577965 MK543282 Vaux e al. 2017
SFKH-TMP012 i a um i a um Mahia Peninsula, NZ MK577964 MK543284 Vaux e al. 2017
SFKH-TMP009 Cominella adpse sa U upukapuka Bay, NZ MH198163 MH277542 Vaux e al. 2017
SFKH-TMP010 i ga a Spi i s Bay, NZ MK558054 MK543287 Vaux e al. 2017
SFKH-TMP004 Glaphy ina cauda a
Fa well Spi , Golden Bay, NZ
MK558055 MK543288 Vaux e al. 2017
37
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
op ope culum;
os osph adium;
p penis;
pdg opening o pos e io duc o diges i e gland;
pma pos e io mixing a ea;
poe pos e io oesophagus;
p p oboscis;
p p p opodium;
p pg p opodial g oo e;
p p oboscis e ac o s;
pw p oboscis wall;
adula;
d hynchodaeum;
e ec um;
s siphon;
sd sali a y duc ;
sg sali a y gland;
SL shell leng h;
sp seminal papilla;
s s omach;
s . s a ion;
SW shell wid h;
a agina;
d as de e ens;
l al e o Leiblein.
Ins i u ion
MNHN Muséum na ional d’His oi e na u elle, Pa is.
RESuL S
phylogene ic analysis
Phylogene ic analysis o bo h he he COI and 28S da ase s
eco e ed a well-suppo ed clade comp ising sequences o
Siphonalia and Phaenomenella (
Figs1; 2
). The K2P pai wise
dis ances dis ibu ion o Phaenomenella and Siphonalia e ealed
wo modes: below 1.1% and abo e 7.2% ha we in e p e
as co esponding o he in a- and in e speci ic compa isons
espec i ely (
able2
). The se en e ealed clus e s also co e-
sponded o he well-suppo ed monophyle ic o single specimen
lineages on he COI-based ee bu ela ionships among hem
a e mos ly poo ly esol ed. Specimens o only six clus e ecog-
nized based on he COI we e p esen in he 28S da a se , and
hey o med i e ecip ocally monophyle ic lineages (
Fig.2
).
Thus he 28S-based clus e s mos ly co esponded o hose e-
ealed wi h COI, excep o he un esol ed ela ionship o he
specimen MNHN-IM-2013-50260. Based on he analysis o
single-gene da ase s, we conclude ha he analyzed specimens
o Phaenomenella and Siphonalia comp ise se en MO us.
wo o he MO u can be iden i ied as Phaenomenella
in la a Shikama, 1971 (Fig.1 (1)) and Phaenomenella insul-
ap a asensis (Oku ani& Lan, 1994) (Fig.1(5)). One MO u
was ep esen ed by a single specimen only in he COI da a
se (Fig.1(3), MNHN-IM-2013-50260). I has s ong e-
semblance o Phaenomenella callopho ella (F aussen, 2004),
desc ibed o aiwan based on a single specimen om simila
dep hs (500-900m s 795-822m in ou specimen) (F aus-
sen 2004), al hough i di e s in ha ing highe spi e and less
in la ed who ls. Since we a e no able o iden i y he a iabili y
o his species we a ibu e ou specimen o P.callopho ella
wi h some ese a ion. wo specimens (Fig.1(2), MNHN-
IM-2013-50012 and MNHN-IM-2013-41068) a e subadul s
(wi h shell leng h less han 15mm) and ha e some esem-
blance o P. hachi F aussen& S ahlschmid , 2012. Al hough
hey may ep esen a di e en species, he limi ed ma e ial
a ailable o us p e en s us p esen ly om u he axonomic
conside a ion o he species. wo MO us we e ep esen ed
by se e al well-p ese ed adul specimens and canno be a -
ibu ed o any exis ing species o Phaenomenella. They a e
desc ibed he ein as new species, Phaenomenella samadiae n.sp.
and P. nicoi n.sp. Finally one MO u can be unambiguously
iden i ied as Siphonalia spadicea (Ree e, 1847).
The analysis o he COI–28S da a se (Fig.3) eco e ed a
well-suppo ed Phaenomenella-Siphonalia clus e consis en
wi h he one in he COI ee, excep o P.in la a, which
was no ep esen ed in he conca ena ed da a se . Whe eas
Phaenomenella c . hachi, P. c . callopo ella and P. nicoi n.sp.
o med a well-suppo ed subclade (BI pos e io p obabili y
= 0.97), P. samadiae n.sp. showed weakly suppo ed a in-
i y o Siphonalia spadicea (BI pos e io p obabili y = 0.75).
This esul ques ions monophyly o Phaenomenella in ela-
ion o Siphonalia. In he absence o he da a on Siphonalia
cassida iae o mis, he ype species o he genus, and bea ing
in mind he conchological di e ences be ween he gene a,
we accep he conse a i e app oach and do no synonymize
Phaenomanella wi h Siphonalia.
The wo majo clades o he Sou he n hemisphe e buccinids
included in he p esen s udy o med a weakly suppo ed
clade comp ising a well-suppo ed Buccinulum – Aenea o
g ouping (BI pos e io p obabili y = 1.00), and a weakly sup-
po ed Penion-Kelle ia-An a c onep unea clade (BI pos e io
p obabili y = 0.73). This clade showed no suppo ed ela ion-
ship o he Phaenomenella-Siphonalia clus e . As men ioned
abo e, Penion and Aenea o we e o iginally included by Finlay
(1928) in Siphonaliinae.
SYS EMA IC ACCOuN
O de NEOGAS ROPOdA Wenz, 1938
Family buccinidae Ra inesque, 1815
Genus Phaenomenella F aussen& Hado n, 2006
ype species.— Mana ia (?) in la a Shikama, 1971 (Od).
eMa ks
The in agene ic shell a iabili y o Phaenomenella is high
and a ew gene al cha ac e s can be men ioned – “b oade
han high p o oconch wi h la ened ip and (…) a a he
sha p angula ion jus abo e he su u e. The uppe eleoconch
who ls a e shoulde ed, a shape which is accen ua ed by he
axial knobs o ibs, o ha e he appea ance o being by he
p esence o ob iously con ex axial ibs” (F aussen& S ahls-
chmid 2013:82). Radula wi h a icuspid cen al oo h wi h
ec angula base and la e als wi h 3 o 4cusps. An e io o egu
wi h well-de ined al e o Leiblein and la ge gland o Leiblein.
38 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
Table 2. — The ma ix o K2P pai wise gene ic dis ances in % (e.g. 133 = 0.133) o COI o he Phaenomenella F aussen & Hado n, 2006 and Siphonalia
A. Adams, 1863 species. Wi hin species: min = 0.000, mean = 0.018, max = 0.011. Be ween species: min = 0.072, mean = 0.157, max = 0.198.
IM-2013-41072
Ph. in la a
IM-2013-41073
Ph. in la a
IM-2013-50203
Ph. insulap a asensis
IM-2013-50204
Ph. insulap a asensis
IM-2013-50205
Ph. insulap a asensis
IM-2013-41068
Ph. c hachi
IM-2013-50012
Ph. c hachi
IM-2013-50260
Ph. c callopho ella
IM-2007-34639
Ph. nicoi
IM-2007-34644
Ph. nicoi
IM-2013-59398
Ph. nicoi
IM-2013-61585
Ph. nicoi
IM-2013-61637
Ph. nicoi
IM-2013-61639
Ph. nicoi
IM-2013-61673
Ph. nicoi
IM-2007-32673
S. spadicea
IM-2007-32674
S. spadicea
IM-2007-32762
S. spadicea
IM-2007-32856
S. spadicea
IM-2007-32995
S. spadicea
IM-2007-32996
S. spadicea
IM-2009-11271
S. spadicea
IM-2007-32997
S. spadicea
IM-2013-61674
Ph. samadiae
IM-2013-61617
Ph. samadiae
IM-2013-41072
Ph. in la a
IM-2013-41073
Ph. in la a
000
IM-2013-50203
Ph. insulap a asensis
133 133
IM-2013-50204
Ph. insulap a asensis
133 133 000
IM-2013-50205
Ph. insulap a asensis
139 139 006 006
IM-2013-41068
Ph. c hachi
130 130 127 127 124
IM-2013-50012
Ph. c hachi
127 127 127 127 127 002
IM-2013-50260
Ph. c callopho ella
130 130 120 120 123 077 080
IM-2007-34639
Ph. nicoi n. sp.
140 140 130 130 136 072 072 079
IM-2007-34644
Ph. nicoi n. sp.
140 140 130 130 136 072 072 079 000
IM-2013-59398
Ph. nicoi n. sp.
140 140 131 131 136 081 082 079 011 011
IM-2013-61585
Ph. nicoi n. sp.
140 140 131 131 136 081 082 079 011 011 000
IM-2013-61637
Ph. nicoi n. sp.
140 140 130 130 136 076 077 079 004 004 006 006
IM-2013-61639
Ph. nicoi n. sp.
143 143 133 133 139 079 079 077 009 009 002 002 004
IM-2013-61673
Ph. nicoi n. sp.
143 143 133 133 139 079 079 077 009 009 002 002 004 000
IM-2007-32673
S. spadicea
195 195 194 194 191 171 174 165 172 172 170 170 169 173 173
IM-2007-32674
S. spadicea
195 195 194 194 191 171 174 165 172 172 170 170 169 173 173 000
IM-2007-32762
S. spadicea
195 195 194 194 191 171 174 165 172 172 170 170 169 173 173 000 000
IM-2007-32856
S. spadicea
185 185 185 185 182 168 171 162 169 169 166 166 166 169 169 006 006 006
IM-2007-32995
S. spadicea
198 198 198 198 194 174 178 168 176 176 173 173 173 176 176 002 002 002 009
IM-2007-32996
S. spadicea
192 192 191 191 188 168 171 162 169 169 167 167 166 170 170 002 002 002 004 004
IM-2009-11271
S. spadicea
195 195 194 194 191 171 174 165 172 172 170 170 169 173 173 000 000 000 006 002 002
IM-2007-32997
S. spadicea
192 192 191 191 188 168 171 162 169 169 167 167 166 170 170 002 002 002 004 004 000 002
IM-2013-61674
Ph. samadiae n. sp.
157 157 157 157 163 172 171 156 176 176 173 173 179 176 176 186 186 186 176 182 183 186 183
IM-2013-61617
Ph. samadiae n. sp.
160 160 159 159 166 169 168 152 167 167 164 164 170 167 167 179 179 179 170 183 176 179 176 006
39
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
1.00
Phaenomenella c hachi
IM-2013-50012 (2)
Aenea o aledic us S
FKH-TMP013
Glaphy ina cauda a S
FKH-TMP004
Kelle ia kelle ii
KK12
Aenea o elegans
SFKH-TMP015
Volu opsius no wegicus
20140783
Pa a e i usus ca ina us
SFKH-TMP005
Phaenomenella insulap a asensis
IM-2013-50205
Penion sulca us
Phoenix1
IM-2013-61639
Aus o usus glans
SFKH-TMP014
IM-2013-61673
Mana ia b e icauda
Siphonalia spadicea
IM-2007-32997
Siphonalia spadicea
IM-2007-32996
IM-2007-34639
Aenea o ecens
M.190119
Siphonalia spadicea
IM-2007-32856
(6)
IM-2013-61674
Penion cu ie ianus
M.183792
Siphonalia spadicea
IM-2007-32762
IM-2007-34644
Calag asso alde menensis IM-2007-34635
Penion manda inus
C.456980
IM-2013-61585 (4)
Penion cha hamensis
M.190082/2
Enigma icolus monnie i
IM-2009-7079
Penion cu ie ianus
M.183927
Phaenomenella insulap a asensis
IM-2013-50204
Buccinum unda um
20070640
Aenea o o agoensis
M.279437
Cominella adspe sa S
FKH-TMP009
Penion cha hamensis
M.190085/3
Buccinulum pe inax inlayi
M.302870/2
Buccinulum i a um i a um
SFKH-TMP012
An a c onep unea ben hicola
M.183832
Phaenomenella c callopho ella
IM-2013-50260 (3
)
Kelle ia lischkei
KL2
Penion sulca us
Phoenix9
Phaenomenella insulap a asensis IM-2013-50203 (5)
Phaenomenella c hachi
IM-2013-41068
Siphonalia spadicea
IM-2007-32673
Cominella i ga a b ookesi
SFKH-TMP010
Phaenomenella in la a
IM-2013-41072 (1)
IM-2013-61637
Siphonalia spadicea
IM-2007-32674
Siphonalia spadicea
IM-2009-11271
Buccinulum uscozona um
M.302907/2
Phaenomenella in la a
IM-2013-41073
Buccinulum obus um
M.314755/1
IM-2013-59398
Mana ia clandes ina
IM-2007-32864
Buccinulum pallidum
M.258277_6
Siphonalia spadicea
IM-2007-32995
Aenea o ben hicolus
M.274111
Penion maximus
C.487648
IM-2013-61617 (7)
1.00
1.00
0.99 0.99
0.99
0.96
0.94
0.96
0.75
0.94
0.84
100
1.00
1.00
0.89
0.87
1.00
0.98
0.78
1.00 0.97
Phaenomenella nicoi
n.sp.
1.00
1.00
1.00
Phaenomenella samadiae
n.sp.
0.82
0.92
0.96
0.4
M
a
n
a
i
a
b
e
i
cauda
g
M
a
n
a
i
a
c
l
a
n
de
s
in
a
IM-2
00
7-
3
2
86
4
0.82
0
.
4
(1) (2) (3) (4) (5) (6) (7)
Fig. 1. — Phylogene ic ee o a da ase o Buccinoidea Ra inesque, 1815 ob ained wi h Bayesian analysis o COI gene sequences. Suppo alues shown o he
suppo ed nodes wi h pos e io p obabili y alues be ween 0.70 and 1.00 only. The numbe s in b acke s ollowing he species names co espond o illus a ed
specimens a he bo om o he igu e.
46 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
AB
CE
D
F
H
G
K
L
JI
Fig. 7. — Shells o Phaenomenella nicoi n. sp.: A-C, holo ype, MNHN-IM-2013-61585, SL 41.1 mm; D, ZHONGSHA 2015, s . CP4157, sequenced spm, MNHN-
IM-2013-59398, SL 37.5 mm; E, ZHONGSHA 2015, s . CP4133, sequenced spm, MNHN-IM-2013-61639, SL 43.2 mm; F, AURORA 2007, s . CP2685, sequenced
spm, MNHN-IM-2007-34639, SL 34.7 mm; G, AURORA 2007, s . CP2685, no -sequenced spm, MNHN-IM-2007-34644, SL 38.7 mm; H, ZHONGSHA 2015, s .
CP4133, sequenced spm, MNHN-IM-2013-61637, SL 30.6 mm (ana omy s udied); I, ZHONGSHA 2015, s . CP4134, sequenced spm, MNHN-IM-2013-61673,
SL 42 mm; J, ZHONGSHA 2015, s . CP4133, no -sequenced spm, MNHN-IM-2013-61636, SL 40.2 mm (ana omy s udied); K, L, ZHONGSHA 2015, s . CP4132,
no -sequenced spm, MNHN-IM-2013-61592, 39.1 mm; L, enla ged p o oconch. Scale ba s: A-K, 20 cm; L, 1 mm.

47
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
A
B
C
D
G
E
F
p
p pg
pma
eye
scm
s
gon
p
d
l
aoe
p
sd
gl
poe
ao
n
p
sd
ba
l
aoe
poe
p
m
bm
aoe
od d
bn
pw
gl
sg
sg
sg
n
poe
gl
sd
pma
s
poe
dg
cep.
hd os sp
p
c
h
poe
kd
in
p
dg
Fig. 8. — Ana omy o Phaenomenella nicoi n. sp.: A-D, MNHN-IM-2013-61636, male; E-G, MNHN-IM-2013-61637, emale: A, so body emo ed om he shell;
B, penis; C, p oboscis, opened do sally; D, o egu , igh la e al iew; E, o egu , igh la e al iew, igh sali a y gland displaced; F, o egu , igh la e al iew, igh
sali a y gland in o iginal posi ion; G, s omach, ou e iew. Abb e ia ions: see ma e ial and me hods. Scale ba s: 5 mm.
48 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
penul ima e who l, on las who l 55 co ds, o which abou 20
on canal. Co ds di e ing sligh ly in wid h, wi h mos na ow
on subsu u al amp, in e spaces be ween co ds abou 1/3-½ o
co d’s wid h. S ong axial ibs p esen on en i e shell, su u e
o su u e on uppe mos eleoconch who ls, g adually becom
-
ing obsole e on subsu u al amp and absen on amp o las
and penul ima e who ls. On las who l ibs dis inc only on
shoulde , p oducing i s angula ed appea ance. Numbe o
ibs s able, i.e., 15-16pe who l.
Ape u e o a e, whi e inside, angula ed pos e io ly, ou e
lip hin, e enly con ex, conca e a ansi ion o canal. Pa i-
e al wall and columella wi h na ow and hin glossy callus.
Shell co e ed wi h e y ligh oli e adhe ing pe ios acum,
o ming densely spaced low axial lamellae isible in in e spaces
be ween co ds.
Ope culum pa ially ab aded, when in ac (Fig.7 d) span-
ning mos o ape u e wi h dis inc ly u ned le wa ds e minal
nucleus and ounded uppe pa .
Radula (Fig.5 C-F)
Examined in i e specimens, including holo ype. Ra he
simila in all specimens; cen al oo h wi h ec angula basal
pa wi h weakly a cua e an e io ma gin and h ee medium
long iangula b oad cusps, cen al one sho e and na owe
han la e al ones. La e al ee h no mally icuspa e wi h weakly
cu ed basal side, a ached o memb ane. Ou e mos cusp
ecu ed, medium long, inne cusp weakly ecu ed, abou
2/3 o ou e cusp leng h. In e media e cusp sho es , si ua ed
sligh ly close o inne cusp. In one sequenced specimen
(MNHN-IM-2013-61673, Fig.5F) in e media e cusps o
he le la e al ee h o he adula pai ed, nea ly equal in size,
while la e al ee h on igh side ha e b oade in e media e
cusp subdi ided on op.
Measu emen s (holo ype)
Shell leng h 41.3mm, las who l leng h 26.9mm, ape u e
leng h (wi hou canal) 16.0mm, diame e 16.5mm. In he
la ges specimen s udied, shell leng h eaching 43 mm.
Ana omy
wo specimens examined: MNHN-IM-2013-61636, male,
and MNHN-IM-2013-61637, emale, sequenced pa a ype
(simila in bo h s udied specimens). So body pa ly ex ac ed
om he shell. Head medium la ge, wi h wo long en acles
and la ge black eyes on lobes a bases o en acles. Man le
simila o ha o Phaenomenella samadiae n. sp. Penis la -
ened, wi h seminal papilla si ua ed on i s op and su ounded
by ci cle old o skin (Fig.8B).
diges i e sys em. P oboscis almos comple ely in e ed in o
hynchodaeum (Fig.8d, p ). Se e al bands o p oboscis
e ac o s muscles a ached o middle pa on bo h sides o
hynchodaeum (Fig.8d-F, p ). Buccal mass sligh ly sho e
han e ac ed p oboscis (Fig.8C, bm), a ached o i s walls
by mul iple odon opho al e ac o s (Fig.8C, od ). Radula
lying in middle o buccal mass and a ached o p oboscis walls
by median e ac o (Fig.8C, m ).
An e io oesophagus s aigh , ollowing along en al side
o p oboscis (Fig.8E, aoe). Val e o Leiblein (Fig.8d, E, l)
medium la ge, coni o m, si ua ed immedia ely be o e ne e
ing (Fig.8d, n ). Sali a y glands medium-la ge (abou
0.3p oboscis leng h), used en ally benea h ne e ing
(Fig.8E, F, sg), wi h e y hin s ongly con olu ed sali a y
duc s ollowing along an e io oesophagus. Gland o Leiblein
medium in size (Fig.8d, E, gl), ollowing along pos e io oe-
sophagus and an e io ao a. S omach a he la ge, spanning
abou 0.4who l (Fig.8G). Pos e io mixing a ea (pma) wice
la ge han in Phaenomenella samadiae, pos e io oesophagus
and in es ine medium wide.
eMa ks
The new species is highly a iable in shell shape. Some o he
specimens a e much mo e slende (Fig.7d, I, G) and he axial
ibs a e ei he e y weak o obsole e. The specimen wi h no
axial ibs was collec ed a a maximal dep h (1634-1683m),
bu he e is no clea co ela ion wi h dep h, since syn opic
specimens can ha e s ong o weak axial ibs. Ne e heless he
molecula da a clea ly indica es he conspeci i y o “ ypical”
angula ed specimens wi h well-de eloped ibs and smoo h
ones. The specimens collec ed a g ea e dep h ha e he p o-
oconch and uppe eleoconch who ls mo e e oded o missing.
In gene al shape he new species has some esemblance o
P.mokeno um F aussen, 2008 om he Andaman Sea, di e -
ing in be e p onounced axial ibs in la e eleoconch who ls
and mo e a enua ed na ow siphonal canal. Also P.nicoi
n. sp. has smalle size (maximal shell leng h 42mm e sus
55.6mm in P.mokeno um).
Genus Siphonalia A.Adams, 1863
ype species. — Buccinum cassida iae o me Ree e, 1846
(Subsequen designa ion by Cossmann 1889).
Siphonalia cassida iae o mis (Ree e, 1846)
(Figs9A-C; 10; 11A)
Buccinum cassida iae o mis Ree e, 1846: pl. 2, sp. 11..
M
a e ial
exaMined
.— Japan • 2lo s, 3specimens; O Hashima,
Miyazaki P e ec u e, Kyushu; 10.V.1996 (no.1, Fig.9A); O A sumi
Peninsula, Aichi P e ec u e; 30m; 13.V.2001 (nos. 2, 3, Figs9B, C).
coMpleMen o desc ip ion
Radula
Radula s udied in h ee specimens. Ra he simila in all spec-
imens (Fig.11A, B); cen al oo h wi h ec angula basal pa
and weakly a cua e an e io ma gin and h ee medium long
iangula b oad cusps, cen al one equal in leng h bu sligh ly
na owe han la e al ones. La e al ee h icuspa e wi h weak-
ly cu ed, nea ly s aigh basal side, a ached o memb ane.
Ou e mos cusp ecu ed, medium long, inne cusp weakly
ecu ed, abou 2/3 o ou e cusp leng h. In e media e cusp
sho es , si ua ed close o inne cusp.
49
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
Fig. 9. — Shells o s udied Siphonalia spp.: A-C, S. cassida iae o mis (Ree e, 1846): specimen no. 1 (A), specimen no. 2 (B), specimen no. 3 (C); D, E, S. p e e i
G. B. Sowe by III, 1900; D, specimen no. 1; E, specimen no. 2. Scale ba : 10 mm.
A
B
C
D
E
50 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
A
A-C
E, F
D
G
H
I
B
C
E
F
D
G
I
H
dg
kd
p
s
sg
p
aoe poe
l l
adg
in
poe
oeo
gl
p
n
n
p
l
p
sg
sd dg
s pma
pma
gl
poe
n
aoe
adg
in
poe
oeo pdg
poe
op
s
dg
m
m
hd
eyes
cep.
hd
s
s
p
sp
eye
Fig. 10
. — Ana omy o Siphonalia cassida iae o mis (Ree e, 1846): A, B; en al (A) and do sal (B) iews o body o specimen no. 1, emo ed om he shell;
C, cephalopodium o no. 2, on iew; D, penis o no. 3; E, F, o egu o no. 1, wi h igh sali a y gland in o iginal posi ion (E) and emo ed (F); G, s omach o
no. 2, ex e nal iew; H, s omach o no. 1, opened do sally; I, s omach o no. 2, opened do sally. Abb e ia ions: see ma e ial and me hods. Scale ba s: A-F,
10 mm; G, I, 5 mm; H, 1 mm
51
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
Ana omy
So body (no.1, emale, Fig.10A, B, E, F, no.2, emale,
Fig.10C) wi h app oxima ely 3who ls. Head sho and
b oad, wi h sho con ac ed en acles. Eyes small, si ua ed
a small lobes in he middle o en acles (Fig.10B, C, eyes).
Foo con ac ed, wi h e y na ow p opodium and la ge ope -
culum wi h e minal nucleus. Penis o spm. no.3 (Fig.10d)
medium long, la ened, con ac ed, wi h small (con ac ed)
ounded seminal papilla a he op, su ounded by ci cula old
o skin. Man le wi h e y long siphon in dissec ed specimens
(longe han hal man le wid h).
diges i e sys em. P oboscis hal e e ed ou o hynchodaeum,
hick, con ac ed (Fig.10E, F, p ). P oboscis e ac o s (p )
a ached o hynchodaeum along bo h sides o an e io oe-
sophagus (mos ly on i s igh side), connec ing hynchodaeum
o la e al walls o body haemocoel. An e io oesophagus
sho and wide, do so- en ally la ened, along en al side
o p oboscis (Fig.10E, F, aoe) in o ela i ely small ounded
al e o Leiblein (Fig.10F, l), si ua ed immedia ely an e io
o ne e ing (Fig.10F, n ). Sali a y glands medium small
(abou 0.25p oboscis leng h) (Fig.10E, F, sg), wi h e y hin
s ongly con olu ed sali a y duc s (Fig.10F, sd) ollowing along
an e io oesophagus. Gland o Leiblein la ge (Fig.10E, F, gl),
ollowing along hick, ound in sec ion pos e io oesopha-
gus (Fig.10E, F, poe). S omach spanning abou 0.3who l
(Fig.10G). Pos e io mixing a ea no la ge in spm. no.1,
la ge in spm. no.2 (Fig.10G, I, pma). Pos e io oesophagus
and in es ine medium wide in bo h specimens. Opening o
pos e io duc o diges i e gland ( ound in spm. no.2) loca ed
nea oesophageal opening (Fig.10I, pdg), opening o an e io
duc ( ound in spms. nos. 1 and 2) loca ed nea beginning
AB
CD
Fig. 11. — Radulae o Siphonalia spp. A, S. cassida iae o mis (Ree e, 1846) no. 1; B, S. cassida iae o mis no. 2; C, S. p e e i G. B. Sowe by III, 1900 no. 1;
D, S. p e e i no. 2. Scale ba s: 200 µm.

52 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
o in es ine. Inne s omach wall be ween wo openings wi h
longi udinal old (Fig.10H, l l), lined wi h low oblique olds,
emaining pa o inne and ou e s omach walls lined wi h
mode a ely high ans e se olds.
Siphonalia p e e i G. B. Sowe by III, 1900
(Figs9d, E; 11C, d; 12)
Siphonalia p e e i G. B. Sowe by III, 1900: 440, pl. 11, ig. 3.
Ma e ial exaMined.— Japan • 1lo , 2specimens; O Hashima,
Miyazaki P e ec u e, Kyushu; 10.V.1996 (nos. 1, 2, igs8d, E).
coMpleMen o desc ip ion
Radula
Radula a he simila in bo h specimens (Fig.11C, d); cen al
oo h wi h ec angula basal pa and weakly a cua e an e io
ma gin and h ee medium long iangula b oad cusps, cen al
one equal in leng h bu sligh ly na owe han la e al ones.
La e al ee h icuspa e wi h weakly cu ed basal side, a ached
o memb ane. Ou e mos cusp ecu ed, medium long, inne
cusp weakly ecu ed, abou 2/3o ou e cusp leng h; inne
cusp in igh longi udinal ow o specimen spm. no.2pa ially
subdi ided (Fig.11d). In e media e cusp sho es , si ua ed
close o inne cusp; inne cusp o spm. no.2pa ially sub-
di ided in le longi udinal ow.
Ana omy (spm. no.1, male, Fig.12)
Head e y sho and b oad, en acles sho , con ac ed, wi h
small eyes a lobes. Foo con ac ed, p opodium mode a ely
wide, ope culum o al wi h e minal nucleus. Penis a he
la ge (Fig.12C), la ened, wi h long na ow seminal papilla
in deepening a he op. Man le wi h medium long siphon.
diges i e sys em. P oboscis pa ly e e ed ou o hyn-
chodaeum, wi h con ac ed walls. Mul iple p oboscis e ac-
o s a aching mos ly along igh side o an e io oesophagus
(Fig.12d, E, p ), connec ing hynchodaeum and la e al walls
o body haemocoel. Buccal mass sligh ly sho e han e ac ed
p oboscis (Fig.12F, bm), a aching o i s walls by mul iple
odon opho al e ac o s (od ). Radula lying in middle o buc-
cal mass and a ached o p oboscis walls by median e ac o
(Fig.12F, m ). Sali a y glands (Fig.12d, E, sg) medium la ge
(0.4p oboscis leng h), o al, wi h sali a y duc s ollowing on
bo h sides o an e io oesophagus. An e io oesophagus wide,
do so- en ally la ened (Fig.12E, aoe), al e o Leiblein
ounded, medium la ge. Pos e io oesophagus (poe) ela i ely
na ow. Gland o Leiblein la ge, olded benea h ne e ing
(Fig.12E, gl). S omach spanning abou 0.3who l (Fig.12G).
Pos e io mixing a ea no la ge (Fig.12G, H, pma). In es ine
medium wide. Opening o pos e io duc o diges i e gland
loca ed nea oesophageal opening (Fig.12I, pdg), opening o
an e io duc loca ed close o beginning o in es ine. Inne
s omach wall be ween wo openings con ains longi udinal
old (Fig.12H, l l), lined wi h low oblique olds, es pa
o inne and ou e s omach wall lined wi h mode a ely high
ans e se olds.
dISCuSSION
Resul s o he phylogene ic analysis sugges close a ini ies
o Siphonalia and Phaenomenella ha emained unno iced
p e iously. F aussen& Hado n (2006), while desc ibing
Phaenomenella, compa ed i o Mana ia and Eosipho, bu no
o Siphonalia. The shell ou line o some Phaenomenella (e.g.
Phaenomenella insulap a asensis) is a he simila o Siphonalia:
he shell is s ou , wi h s ongly con ex who ls and a ecu ed
siphonal canal. Species o Phaenomenella hough ha e a much
la ge (abou wice) p o oconch in compa ison wi h Siphonalia.
The in agene ic a iabili y o shell shape in Phaenomenella in
i s cu en de ini ion is e y high (F aussen& S ahlschmid
2013) and in i s ex emes he e is no esemblance be ween
he wo gene a. I should also be bo n in mind ha some
o he mos di e ging species o Phaenomenella we e no ye
sequenced and may all in o o he lineages.
Rep esen a i es o bo h gene a a e also ana omically simila ,
pa icula ly in he diges i e sys em cha ac e s. Bo h Phaeno-
menella and Siphonalia ha e a b oad, medium long in he
con ac ed s a e p oboscis, medium la ge sali a y glands and
a la ge gland o Leiblein. I should be men ioned ha despi e
hese gene al simila i ies, he e a e no unique mo phological
cha ac e s uni ing bo h gene a. The adula mo phology is
e y simila in bo h gene a ( adula o one mo e species, S.ma-
ybe hi Pa h, 1996 was illus a ed in Zhang& Zhang 2018),
howe e , as in he case wi h he body ana omy, i is o a he
gene alized buccinid appea ance; simila adula mo phology
can be ound in many un ela ed gene a – eg. La isipho dall,
1916 (Kosyan 2006), Plici usus dall, 1902 (Kosyan& Kan-
o 2012), Aulaco usus dall, 1918 (Kosyan& Kan o 2013).
Ou molecula analysis did no eco e Phaenomenella as
monophyle ic. In bo h COI and combined COI+28S analyses
he in e nal ela ionships wi hin Phaenomenella – Siphonalia
clade a e no esol ed. Siphonalia spadicea clus e wi hou sig-
ni ican suppo wi h P.samadiae n.sp. We ha e only a single
species o Siphonalia in ou analyses so i is oo p elimina y o
change he classi ica ion on he basis o he incomple e da ase .
The e o e we p esen ly e ain he alidi y o Phaenomenella,
al hough i is possible ha Phaenomenella and Siphonalia can
belong o a single genus. One o he dis inc ions be ween
he gene a is he dep h ange o known species. Gene ally,
species o Siphonalia dwell a shallowe dep hs – om 10 o
300m (Oku ani 2000), while Phaenomenella is eco ded a
190-1389m (F aussen& S ahlschmid 2013; he ein). The
new species a e a ibu ed o Phaenomenella based on s onge
conchological simila i y o o he species o he genus a he
han o species o Siphonalia. un o una ely he p o oconch
o P.samadiae n.sp. was decolla ed in all a ailable specimens,
bu he p o oconch o P.nicoi n.sp. is la ge globose, simila
o o he species o Phaenomenella.
The analysis o a b oade da ase o Buccinoidea ejec ed
he monophyly o Siphonaliinae in i s o iginal scope. None
o he Recen gene a, o iginally included by Finlay (1928) in
he sub amily, ha a e Penion, Aenea o and Glaphy ina, a e
closely ela ed nei he o each o he , no o Siphonalia. The
sys em o Buccinidae and Buccinoidea in gene al is s ill a
53
Taxonomic posi ion o Phaenomenella
ZOOSYSTEMA • 2020 • 42 (3)
Fig. 12
. — Ana omy o Siphonalia p e e i G. B. Sowe by III, 1900, specimen no. 1: A, cephalopodium, en al iew; B, cephalopodium, do sal iew, man le emo ed;
C, penis; D, o egu , do sal iew; E, o egu , en al iew; F, p oboscis opened do sally; G, s omach, ex e nal iew; H, s omach, opened do sally. Abb e ia ions:
see ma e ial and me hods. Scale ba s: A-F, 10 mm; G, H, 5 mm.
A
A, B
D-F
G
H
C
kd
c
os
cm
s
op
sp sp
p
eye p pg
od
n
gl
poe
sd
od
aoe
pw
bm
d
m
pdg
l l
adg
in
poe
oeo
pma
aoe
p
dg
pma
s
in
poe
d
p
sg
cep.
hd
DEF
GH
BC
54 ZOOSYSTEMA • 2020 • 42 (3)
Kan o Y. e al.
om being esol ed, wi h many p oblema ic buccinoidean
lineages (see e.g. Cou o e al. 2016; Ha asewych 2018). The e-
o e he ank o he in e ed clade Siphonalia + Phaenomenella
can be esol ed only a e ob aining he obus phylogeny o
he en i e supe amily Buccinoidea.
Acknowledgemen s
The ma e ial was collec ed mos ly in esea ch c uises in he
Sou h China Sea and he Philippines Sea o ganized espec i ely
by Na ional aiwan uni e si y (dONGSHA 2014, NANHAI
2014, ZHONGSHA 2015; PI Wei-Jen Chen), and Muséum
na ional d’His oi e na u elle, Pa is (MNHN) wi h he Phil-
ippines Bu eau o Fishe ies and Aqua ic Resou ces (BFAR)
(AuRORA 2007; PI Philippe Bouche ) The ma e ial in his
pape o igina es om se e al deep sea c uises, conduc ed by
MNHN, Ins i u de Reche che pou le dé eloppemen (IRd)
and o he pa ne s as pa o he T opical Deep-Sea Ben hos
p og amme. The aiwan and Sou h China Sea c uises we e
suppo ed by bila e al coope a ion esea ch unding om he
aiwan Minis y o Science and echnology (MOS 102-
2923–B–002–001–MY3, PI Wei-Jen Chen) and he F ench
Na ional Resea ch Agency (ANR 12-ISV7-0005- 01, PI Sa ah
Samadi). All expedi ions ope a ed unde he egula ions hen
in o ce in he coun ies in ques ion and sa is y he condi ions
se by he Nagoya P o ocol o access o gene ic esou ces. The
s udy was conduc ed using Join usage Cen e “Ins umen al
me hods in ecology” a he IEE RAS. We hank A. Ne e ina
and A. Nek aso o hei iendly and help ul assis ance wi h
he SEM acili ies. The s udies we e suppo ed by he g an
om he Russian Science Founda ion RSF 16–14– 10118. This
p ojec was pa ly suppo ed by he Se ice de Sys éma ique
moléculai e (uMS 2700 CNRS-MNHN) and he bila e al
coope a ion esea ch unding om he Minis y o Science and
echnology, aiwan (g an numbe MOS 102-2923-B-002-
001-MY3) and he F ench Na ional Resea ch Agency (g an
numbe ANR 12-ISV7-0005-01). We hank Philippe Bouche
o he possibili y o wo k wi h he ma e ial om MNHN,
Vi ginie Hé os, Philippe Maes a i, Ba ba a Buge, Jose u ge,
Cy il Chamba , and Nicolas Puilland e o hei ole in speci-
mens p ocessing and hei help in cu a ing and sequencing he
specimens. We hank he e e ees, N. Puilland e and anony-
mous e e ee o aluable ema ks and co ec ions.
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Submi ed on28 Janua y 2019;
accep ed on 8 Ap il 2019;
published on 4 Feb ua y 2020.