Mo phology and phylogeny o wo new species o deep-sea
mush oom so co als (Oc oco allia, Co allidae, An homas inae)
om he Sou he n Mid-A lan ic Ridge
Xinlong Li1,2, Ningxia Xu1,2, Meiling Ge1,2, Xuying Hu1,2, Mengna Li3, Zongling Wang1,2,
Xuelei Zhang1,2, Qinzeng Xu1,2
1 Key Labo a o y o Ma ine Eco-En i onmen al Science and Technology, Fi s Ins i u e o Oceanog aphy, MNR, Qingdao 266061, China
2 Labo a o y o Ma ine Ecology and En i onmen al Science, Qingdao Ma ine Science and Technology Cen e , Qingdao 266100, China
3 Na ional Deep Sea Cen e , MNR, Qingdao 266237, China
h ps://zoobank.o g/BBB6DED7-18E3-4706-A9EA-D31797EA00EF
Co esponding au ho : Qinzeng Xu ([email p o ec ed])
Academic edi o : Pa el S oe ♦
Recei ed
29 June 2025 ♦
Accep ed
19 Oc obe 2025 ♦
Published
5 No embe 2025
Abs ac
Membe s o he sub amily An homas inae Ve ill, 1922, commonly known as mush oom so co als, a e cha ac e ized by hei
capi a e o mush oom-shaped ed colonies and la ge au ozooids. Deep-sea mush oom co als o his sub amily emain poo ly doc-
umen ed in he Sou h A lan ic. This s udy desc ibes wo new An homas inae species, Neoan homas us longis ylus sp. no . and
An homas us mi abilis sp. no ., om he Sou he n Mid-A lan ic Ridge a dep hs o 1,553–2,145 m. Neoan homas us longis ylus sp.
no . is cha ac e ized by a slende s alk, siphonozooids dis ibu ed on he capi ulum and he uppe hi d o he s alk, and en acula
scle i es ha a e p edominan ly s aigh ods and ods wi h one end cu ed. An homas us mi abilis sp. no . is dis inguished by he
p esence o dumbbells in i s en acles, clubs and dumbbells in he an hocodial wall, and he absence o spindles. The mi ochond ial
genome o N. longis ylus sp. no . con ains 14 PCGs, 2 RNAs, and 1 RNA, while ha o A. mi abilis sp. no . comp ises 17 PCGs,
2 RNAs, and 1 RNA, wi h h ee duplica ed genes – ND3, ND6, and ND4L – ha a e iden ical in sequence. Phylogene ic analyses
based on he pa ial m Mu S gene and 13 PCGs om he mi ochond ial genomes (m Mu S no included) e eal he phylogene ic
ela ionships wi hin he sub amily An homas inae. These a e he i s eco ds o bo h gene a om he Sou he n Mid-A lan ic Ridge,
en iching he known species di e si y and p o iding c i ical baseline da a o u u e biodi e si y assessmen s.
Key Wo ds
An homas us, mi ochond ial genome, Neoan homas us, phylogene ic analysis, Sou he n Mid-A lan ic Ridge
In oduc ion
The amily Co alliidae Lamou oux, 1812 exhibi s e-
ma kable mo phological di e si y, wi h species displaying
di e se axial s uc u es – om absen axes o hose com-
posed o un used scle i es, and inally o ully used scle i -
ic axes (Baye 1993; Deb euil e al. 2011; McFadden e al.
2022). Wi hin his amily, he sub amily An homas inae is
dis inguished by i s lack o an axial skele on. Unlike Co al-
liidae and Pa ago giidae, in which au ozooids occu along
he axial b anches, mos An homas inae ha e he au ozo-
oid an hocodia concen a ed o e he capi ulum; in a ew
species, siphonozooids also occu on he s alk. (Molod so-
a 2013; Li e al. 2025). Thei mush oom-like appea ance
makes hem easily dis inguishable om o he co al g oups.
The classi ica ion o An homas inae has unde gone
signi ican e ision in ecen yea s. An homas inae was
his o ically placed wi hin he amily Alcyoniidae due
o mo phological simila i ies wi h gene a such as Al-
cyonium Linnaeus, 1758, Bellonella G ay, 1862, and
Eleu he obia Pü e , 1900 (Thomson 1911, Ve se eld
and Baye 1988; Williams 2003). Be n son e al. (2001)
Zoosys . E ol. 101 (4) 2025, 2103–2121|DOI 10.3897/zse.101.163559
Copy igh Li, X. e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use, dis ibu-
ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2104
e ealed ha An homas us is gene ically close o Co -
allium based on nuclea 18S RNA. An homas inae was
ans e ed o Co alliidae based on an ul a-conse ed
elemen s/exon phylogenomic analysis by McFadden e
al. (2022), which econs uc ed maximum likelihood
ees om 739 loci (60% axon occupancy). Li e al.
(2025) econs uc ed he phylogene ic ela ionships
o he i e gene a wi hin he sub amily An homas i-
nae, which a e An homas us Ve ill, 1878; Ba hyalcyon
Ve sluys, 1906; He e opolypus Tixie -Du i aul , 1964;
Pseudoan homas us Tixie -Du i aul & d’Hond , 1974;
and Neoan homas us Li, Li & Xu, 2025, a genus newly
es ablished by he au ho s.
The sub amily An homas inae is widely dis ibu ed
ac oss he No h A lan ic, Indian, Paci ic, and he Sou h-
e n Ocean, wi h a p edominan p esence in deep-sea en-
i onmen s (>200 m dep h; Molod so a 2013; Li and Xu
2023). The i s desc ibed species, An homas us g andi-
lo us Ve ill, 1878, was epo ed om he No h A lan ic
(Ve ill 1879). Subsequen ly, An homas inae species, es-
pecially An homas us, Pseudoan homas us, and Neoan-
homas us, ha e been documen ed ac oss he No h A -
lan ic, including coas al wa e s nea con inen al ma gins
and islands, as well as along he No he n Mid-A lan ic
Ridge. The Sou h A lan ic ea u es complex opog aphy,
including he Sou he n Mid-A lan ic Ridge (SMAR),
and a ich co al auna. Shallow-wa e ee s (e.g., B a-
zil’s Ab olhos Bank) and deep-sea co als (e.g., Desmo-
phyllum pe usum) a e well-documen ed in his egion
(Leão and Kikuchi 2001; Ca alho e al. 2023). Da a
om OBIS and GBIF e eal occu ences o An homas-
inae in he coas al Sou h A lan ic and nea i s sou he n
islands. Howe e , he deep seamoun s o he SMAR lack
any con i med eco ds o An homas inae. Recen disco -
e ies o new co al species a he Ca lsbe g Ridge (Hu e
al. 2025) u he sugges ha deep-sea ecosys ems may
ha bo nume ous undesc ibed axa, highligh ing he need
o a ge ed explo a ion.
Du ing a ben hic auna su ey o he Sou h A lan ic
Mid-Ocean Ridge, we collec ed samples o An hom-
as inae and disco e ed wo new species. By combining
mo phological axonomic iden i ica ion wi h molecula
phylogene ic analysis, his s udy aims o enhance ou
unde s anding o deep-sea co al species in he Sou he n
Mid-A lan ic Ridge.
Ma e ials and me hods
Specimen collec ion and mo phological
analyses
Eigh specimens we e collec ed in 2024 using he Ji-
aolong Human-Ope a ed Vehicle (HOV) du ing China's
Ocean 83 C uise o ganized by he Na ional Deep Sea
Cen e (NDSC) aboa d he R/V Shenhai Yihao om he
Sou he n Mid-A lan ic Ridge (Table 1). These specimens
we e pho og aphed in si u be o e being sampled and pho-
og aphed again on boa d. The collec ed samples we e
p ese ed using h ee me hods: alcohol p ese a ion,
RNAla e S abiliza ion Solu ion, and apid eezing in
liquid ni ogen. All ype ma e ials ha e been pe manen -
ly deposi ed in he Key Labo a o y o Ma ine Eco-En-
i onmen al Science and Technology, Fi s Ins i u e o
Oceanog aphy loca ed in Qingdao, China. The ouche
numbe s o hese specimens a e p o ided in Table 1.
The shape and colo o he capi ulum and s alk we e
obse ed, he numbe o au ozooids was coun ed, and
hei size was measu ed. Using a Nikon SMZ1270 s e-
eomic oscope, we examined and pho og aphed he ex-
e nal su aces o he capi ulum and au ozooid en acles.
We hen dissec ed he capi ulum and s alk o inspec he
in e io issues. The specimens we e dissec ed using dis-
sec ing needles and scalpels in o six pa s: au ozooid
en acles, pha ynx, an hocodial wall, capi ulum su ace,
s alk, and hold as . These pa s we e sepa a ely soaked in
a 10% sodium hypochlo i e solu ion o 1–2 hou s un il
he so issues we e comple ely dissol ed. The esul ing
scle i es we e cleaned wi h dis illed wa e and anhyd ous
e hanol, hen placed on double-sided insula ing ape. A -
e d ying, he samples we e gold-sp ayed and obse ed
unde a Regulus8100 scanning elec on mic oscope. The
scle i es we e classi ied acco ding o s anda d axonomic
con en ions (Baye 1956; Baye e al. 1983).
DNA ex ac ion and sequencing
Polyps we e collec ed in o 1.5 mL s e ile cen i uge ubes.
To al genomic DNA was ex ac ed using he Mic oElu e
Genomic DNA Ki (OMEGA, USA). DNA concen a ion
and pu i y we e measu ed using a spec opho ome e . Ac-
Table 1. Geog aphical da a o eigh mush oom so co al specimens collec ed om he Sou he n Mid-A lan ic Ridge.
Vouche numbe La i ude, Longi ude Dep h (m) Species
FIO-ATL83-JLBEN26602 22°58'45"S, 13°32'34"W 1929 Neoan homas us longis ylus sp. no .
FIO-ATL83-JLBEN26603 22°58'44"S, 13°32'35"W 1928 Neoan homas us longis ylus sp. no .
FIO-ATL83-JLBEN26604 22°59'2"S, 13°32'14"W 1860 Neoan homas us longis ylus sp. no .
FIO-ATL83-JLBEN26704 22°59'3"S, 13°29'31"W 2151 Neoan homas us longis ylus sp. no .
FIO-ATL83-JLBEN26705 22°58'56"S, 13°29'50"W 2145 Neoan homas us longis ylus sp. no .
FIO-ATL83-JLBEN27711 14°3'28"S, 14°20'53"W 1649 An homas us mi abilis sp. no .
FIO-ATL83-JLBEN27712 14°3'31"S, 14°20'50"W 1649 An homas us mi abilis sp. no .
FIO-ATL83-JLBEN27713 14°3'30"S, 14°20'48"W 1649 An homas us mi abilis sp. no .
Zoosys . E ol. 101 (4) 2025, 2103–2121
zse.penso .ne
2105
cep able c i e ia included an A260/A280 a io be ween
1.8 and 2.0, an A260/A230 a io g ea e han 1.8, and a
concen a ion abo e 100 ng/μL. The ex ac ed DNA was
sequenced on he Illumina pla o m by No ogene Tech-
nology, Inc. (Beijing, China). Addi ionally, he m Mu S
gene was ampli ied using p ime s An hoCo MSH (5’-AG-
GAGAATTATTCTAAGTATGG-3’; He e a e al., 2010)
and Mu -3458R (5’-TSGAGCAAAAGCCACTCC-3’;
Sánchez e al., 2003). The PCR eac ion mix u e (25 μL)
con ained: 1 μL o empla e DNA, 1 μL o each p ime
(0.4 μM), 12.5 μL o 2× Taq Plus Mas e Mix, and 9.5 μL
o ddH2O. The PCR ampli ica ion p o ocol included an
ini ial dena u a ion a 98 °C o 2 minu es, ollowed by 32
cycles (dena u a ion a 98 °C o 15 seconds, annealing a
50 °C o 15 seconds, ex ension a 72 °C o 60 seconds),
wi h a inal ex ension a 72 °C o 2 minu es. Each sample
was ampli ied in iplica e. PCR p oduc s we e isualized
using 2% aga ose gel elec opho esis and sequenced bidi-
ec ionally using Sange sequencing a Sangon Bio ech,
Inc. (Qingdao, China), wi h he same p ime s.
Mi ogenome da a assembly and anno a ion
The mi ochond ial genome was assembled using Ge O -
ganelle 1.7.5 (Jin e al. 2020). Gene anno a ion, includ-
ing p o ein-coding genes (PCGs), RNAs, and RNAs,
was conduc ed using MITOS2 (h ps://usegalaxy.eu). To
enhance he p ecision o RNA gene iden i ica ion, R-
NAscan-SE (h ps:// na.ucsc.edu/ RNAscan-SE/) was
employed. PCGs we e ansla ed in o amino acid se-
quences based on he mold, p o ozoan, and coelen e a e
mi ochond ial gene ic code ( ansla ion able 4). Ini ial
gene p edic ions gene a ed by MITOS we e u he e-
ined by emo ing edundan anno a ions and manually
adjus ing s a and s op codon posi ions. Finally, he com-
ple e ci cula mi ogenome was isualized using P oksee
(h ps://p oksee.ca/). To e i y he p esence o duplica ed
genes in he mi ochond ial genome o An homas us mi-
abilis sp. no ., p ime s we e designed using P ime 5,
and PCR ampli ica ion and sequencing we e pe o med
ollowing he same p o ocol desc ibed abo e (Suppl.
ma e ial 1: able S1). All sequence da a gene a ed in his
s udy ha e been deposi ed in he NCBI da abase (Suppl.
ma e ial 1: able S3).
Phylogene ic analyses
A o al o 47 pa ial m Mu S sequences we e analyzed,
including 8 om his s udy (N. longis ylus sp. no . and
A. mi abilis sp. no .), 35 om he NCBI da abase o he
sub amily An homas inae, and 4 sequences om gene a
ou side o An homas inae wi hin Co alliidae (ou g oups).
PhyloSui e was u ilized o conduc phylogene ic analyses
using bo h Maximum Likelihood and Bayesian me h-
ods. All sequences we e aligned in ba ches wi h MAFFT
7.505 (Ka oh and S andley 2013), and hen immed
wi h Gblocks 0.91. ModelFinde 2.2.0 (Kalyaana-
moo hy e al. 2017) was used o selec he bes - i mod-
el. Maximum likelihood phylogenies we e in e ed using
IQ-TREE 2.2.0 (Nguyen e al. 2015) unde he HKY+F
model o 5000 ul a as boo s aps. Bayesian in e ence
(BI) phylogenies we e econs uc ed using M Bayes
3.2.7a (Ronquis e al. 2012) unde HKY+F model. Fou
independen uns we e conduc ed, each consis ing o ou
Ma ko Chain Mon e Ca lo (MCMC) chains, un o 2
million gene a ions. The i s 25% o sampled ees om
each un we e disca ded as bu n-in o ensu e con e gence.
A o al o 35 comple e mi ochond ial genome sequences
we e analyzed, including 6 om his s udy, 23 om he
NCBI da abase o he sub amily An homas inae, and 6
sequences om gene a ou side o An homas inae wi hin
Co alliidae (ou g oups). Thi een p o ein-coding genes
(PCGs) we e ex ac ed using PhyloSui e 1.2.3 (Zhang e
al. 2020; Xiang e al. 2023), hen aligned in ba ches and
immed using MAFFT 7.505 and Gblocks 0.91. Sub-
s i u ion models we e selec ed o each gene using Mod-
elFinde 2.2.0 (Kalyaanamoo hy e al. 2017) unde he
Bayesian In o ma ion C i e ion (BIC). An edge-linked
pa i ion scheme was employed, and he selec ed models
we e subsequen ly used o phylogene ic in e ence. The
emaining s eps o ee cons uc ion ollowed he same
app oach as hose used o he pa ial m Mu S da ase .
The bes - i models and pa i ioning schemes o BI and
ML a e shown in Suppl. ma e ial 1: able S2. The phylo-
gene ic ee was isualized using he In e ac i e T ee o
Li e (iTOL) websi e (h ps://i ol.embl.de/). To assess he
gene ic di e gence among and wi hin di e en species,
we calcula ed gene ic dis ances wi h Mega 7, employing
he Kimu a 2-Pa ame e model on he conca ena ed 14
PCGs o mi ochond ial genomes.
Resul s
Mo phological axonomy
Class Oc oco allia Haeckel, 1866
O de Scle alcyonacea McFadden, an O wegen &
Qua ini, 2022
Family Co alliidae Lamou oux, 1812
Sub amily An homas inae Ve ill, 1922
Genus Neoan homas us Li, Li & Xu, 2025
Neoan homas us Li, Li & Xu, 2025: 12.
Diagnosis. (Modi ied om Li e al. 2025, wi h bold i al-
ics indica ing modi ica ions.) Mush oom-shaped, capi a e
colonies wi h dome-shaped capi ulum sepa a ed om a
conspicuous and o en cylind ical s alk. Polyps dimo -
phic, wi h scle i es. Au ozooids a anged o e capi u-
lum, s e ile, la ge, no e y nume ous, e ac ile bu no
comple ely e ac ed. Scle i es equally de eloped a ound
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2106
au ozooids. An hocodial a ma u e mo e de eloped nea
base o en acles. Siphonozooids sca e ed o e bo h ca-
pi ulum and s alk, e ile, conspicuous, eebly a mo ed,
and nume ous. Scle i es mul i adia es, ods, spindles,
needles, clubs, and c osses. Pha yngeal scle i es p e-
dominan ly odle s, some imes wi h c osses. Ten acula
scle i es p edomina ely ods, spindles, and mul i adia es.
Type species. Neoan homas us s ella us Li, Li & Xu,
2025.
Neoan homas us longis ylus Li & Xu, sp. no .
h ps://zoobank.o g/CF57044D-6FB2-43F5-9E3F-7219BD41ADAD
Figs 2–5
Ma e ial examined. Holo ype • FIO-ATL83-JL-
BEN26604, collec ed by Qinzeng Xu using he HOV Ji-
aolong om he R/V Shenhai Yihao on 14 Feb ua y 2024
a s a ion JL266 (22°59'2"S, 13°32'14"W), Sou he n
Mid-A lan ic Ridge, a a dep h o 1,860 m.
Pa a ypes • Two specimens (FIO-ATL83-JL-
BEN26602, pa o a colony; FIO-ATL83-JLBEN26603),
same s a ion as o holo ype • Two specimens (FIO-
ATL83-JLBEN26704; FIO-ATL83-JLBEN26705, pa
o a colony), collec ed wi h he same essel, gea , and by
he same collec o s as he holo ype, on 15 Feb ua y 2024
a s a ion JL267 (22°58'56"S, 13°29'50"W), Sou he n
Mid-A lan ic Ridge, a a dep h o 2,150 m.
Diagnosis. Mush oom-shaped, capi a e colony wi h
a sphe ical capi ulum sepa a ed om a cylind ical s alk.
S alk is long, ypically a leas h ee imes longe han
i s maximum wid h, and app oxima ely 2.5 imes longe
han he capi ulum. Au ozooids a e la ge, e ac ile bu
no comple ely e ac ed. Siphonozooids a e ed, small,
isible, and dis ibu ed ac oss he su ace o he capi u-
lum and es ic ed o he uppe 1/3 o he s alk. Scle i es
include ods, clubs, spindles, mul i adia es, and c osses;
scle i e densi y on he capi ulum su ace, en acles, and
pha ynx is abundan . Scle i es om he in e io o he
capi ulum a e absen . An hocodial walls con ain bo h
cu ed ods and clubs. Pha yngeal scle i es a e odle s
and c osses. Ten acula scle i es consis exclusi ely o
s aigh ods and ods wi h one end unila e ally cu ed,
lacking spindles and mul i adia es.
Desc ip ion o holo ype. Colony o m and size.
The specimen FIO-ATL83-JLBEN26604 is a mush-
oom-shaped, capi a e colony wi h a sphe oidal capi -
ulum suppo ed by a conspicuous s alk (Fig. 1B). The
capi ulum is 17 mm in diame e and 15 mm high. The
ansi ion be ween he capi ulum and he s alk is smoo h.
The conspicuous, longi udinally s ia ed s alk is long (45
mm in heigh , 75% o he colony), so , and g adually a-
pe s owa d he capi ulum (6 mm in diame e ), while he
base o he s alk is hicke (14 mm in diame e ). In li e,
he specimen was a ached o s one (Fig. 1B).
Polyps. The polyps a e dimo phic. 28 au ozooids a e
e enly a anged on he capi ulum, e ac ile bu no com-
ple ely e ac ed. Upon e ac ion, he en acles i s con-
ac in o a sphe ical shape, ollowed by he au ozooids
e ac ing in o he capi ulum, lea ing a p o usion on he
su ace o he capi ulum. The la ges au ozooid has a di-
ame e o 5 mm and a leng h o 33 mm, including he
en acles. The en acles each up o 12 mm wi h mos ly
14–18 pai s o pinnules. The longes pinnules a e loca ed
a he dis al hi d o he en acles. Siphonozooids a e dis-
ibu ed on he capi ulum and he uppe hi d o he s alk,
isible as ed p o usions, and a e usually 0.4–0.6 mm
wide, spaced 0.15–0.6 mm apa (Fig. 1D).
Scle i es. Ten acula scle i es (Fig. 2A) a e exclusi ely
composed o ods, including s aigh ods (0.17–0.35 mm
long) and ods wi h one end cu ed (0.18–0.34 mm long).
Pha yngeal scle i es (Fig. 2B) a e mos ly odle s (0.06–
0.09 mm long) and a ew c osses. An hocodial scle i es
(Fig. 3A) a e s aigh ods (0.16–0.36 mm long), ods
wi h one end cu ed (0.16–0.37 mm long), and a ew
clubs (0.18–0.25 mm long). Scle i es om he su ace
o capi ulum (Fig. 3B) a e s aigh ods (0.31–0.35 mm
long), ods wi h one end cu ed (0.23–0.30 mm long),
p ominen ube cula ed spindles (0.17–0.24 mm long),
p ominen ube cula ed clubs (0.18–0.23 mm long), and
mul i adia es (0.12–0.15 mm long). Scle i es om su ace
o he s alk (Fig. 4A) a e mos ly mul i adia es (0.11–0.14
mm long), a ew ods (0.31–0.34 mm long), and p omi-
nen ube cula ed spindles (0.15–0.28 mm long). Scle -
i es om in e io o he s alk (Fig. 4B) a e mos ly ods
(0.15–0.23 mm long) and a ew ods wi h a sho ans-
e se axis (0.18–0.20 mm long). Scle i es o he hold as
(Fig. 4C) a e ods (0.23–0.31 mm long), mul i adia es
(0.12–0.15 mm long), p ominen ube cula ed spindles
(0.15–0.25 mm long), a ew ods wi h a sho ans e se
axis (0.22–0.27 mm long), and c osses. Scle i es om he
in e io o he capi ulum a e absen .
Colo . In alcohol, bo h he capi ulum and he s alk ap-
pea ed, wi h he s alk being sligh ly ligh e in colo han
he capi ulum. The en acle ips and pinnules a e pale yel-
low, and he scle i es a e ed.
Va ia ion. Pa a ype FIO-ATL83-JLBEN26602 is
la ge, measu ing 120 mm in o al heigh , wi h a s alk
heigh o 95 mm and bea ing 28 au ozooids. Pa a ype
FIO-ATL83-JLBEN26603 is a ached o a dead co al
b anch, wi h a o al heigh o 50 mm. The s alk measu es
35 mm, app oxima ely wice he heigh o he capi ulum,
and bea s 12 au ozooids. FIO-ATL83-JLBEN26704 mea-
su es 75 mm in o al heigh , wi h a s alk heigh o 55 mm
and bea s 18 au ozooids. FIO-ATL83-JLBEN26705 has a
o al heigh o 90 mm, wi h a s alk heigh o 72 mm and a
ela i ely small capi ulum (18 mm) bea ing 9 au ozooids.
E ymology. The speci ic name longis ylus e e s o he
long s alk, one o he cha ac e is ics o his species.
Dis ibu ion and habi a . Known only om he
Sou he n Mid-A lan ic Ridge, 1,860–2,151 m.
Rema ks. N. longis ylus sp. no . has siphonozooids dis-
ibu ed ac oss he capi ulum’s su ace and ex ending on o
he s alk, wi h pha yngeal scle i es ha a e mos ly odle s.
These cha ac e is ics dis inguish i om o he gene a in he
sub amily, placing i wi hin he genus Neoan homas us.
Zoosys . E ol. 101 (4) 2025, 2103–2121
zse.penso .ne
2107
Figu e 1. Mo phology o Neoan homas us longis ylus sp. no . A–D. The holo ype FIO-ATL83-JLBEN26604: A. In si u iew;
B. Colony; C. The en acles o au ozooids, indica ed by a ows; D. The siphonozooids, indica ed by a ows. E–H. Pa a ypes
showing colony mo phology: E. FIO-ATL83-JLBEN26602; F. FIO-ATL83-JLBEN26603; G. FIO-ATL83-JLBEN26704; H. FIO-
ATL83-JLBEN26705. Scale ba s: 10 mm (B); 2 mm (C, E).
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2108
Figu e 2. Scle i es o Neoan homas us longis ylus sp. no . (FIO-ATL83-JLBEN26604). A. Scle i es om au ozooid en acles;
B. Scle i es om pha ynx. Scale ba s: 0.10 mm (A); 0.05 mm (B).
N. longis ylus sp. no . esembles N. gigan eus in ha ing
an elonga ed s alk, siphonozooids sca e ed on he uppe
1/3 o s alk. Howe e , hei scle i e cha ac e is ics di e . N.
longis ylus sp. no . has abundan scle i es on i s en acles,
pha ynx, and capi ulum su ace, in con as o N. gigan-
eus (Tixie -Du i aul , 1954), which has spa se scle i es.
Zoosys . E ol. 101 (4) 2025, 2103–2121
zse.penso .ne
2109
Figu e 3. Scle i es o Neoan homas us longis ylus sp. no . (FIO-ATL83-JLBEN26604). A. Scle i es om an hocodial wall;
B. Scle i es om su ace o capi ulum. Scale ba s: 0.10 mm.
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2110
Figu e 4. Scle i es o Neoan homas us longis ylus sp. no . (FIO-ATL83-JLBEN26604). A. Scle i es om su ace o s alk; B. Scle -
i es om in e io o s alk; C. Scle i es om hold as . Scale ba s: 0.10 mm.
Zoosys . E ol. 101 (4) 2025, 2103–2121
zse.penso .ne
2111
The au ozooid en acles o N. longis ylus sp. no . con ain
unila e ally cu ed ods, and i s an hocodial walls ha e
bo h cu ed ods and clubs, ea u es absen in N. gigan eus.
The su ace o N. longis ylus sp. no . lacks he spiny dumb-
bells ound on N. gigan eus. In addi ion, N. longis ylus sp.
no . colonies a e so and mos ly cu ed, while N. gigan e-
us colonies a e igid.
Beyond N. gigan eus, he new species is also clea ly
sepa a ed om all o he membe s o he genus. I is dis-
inguished om N. pu pu eus and N. hicksoni by possess-
ing a dis inc ly elonga ed s alk (a leas h ee imes longe
han wide), as opposed o hei sho s alks. Fu he mo e,
N. longis ylus sp. no . di e s om N. s ella us, N. ahino-
dus, and N. elonga us, which ha e siphonozooids co e -
ing he en i e s alk, whe eas he siphonozooids o he new
species a e es ic ed o he uppe one- hi d.
An homas us Ve ill, 1878
An homas us mi abilis Li & Xu, sp. no .
h ps://zoobank.o g/71F758BF-6C8E-4F9E-B9C6-EABAD67A781C
Figs 5–8
Ma e ial examined. Holo ype • FIO-ATL83-JL-
BEN27711, collec ed by Qinzeng Xu using he HOV Ji-
aolong om he R/V Shenhai Yihao on 28 Feb ua y 2024
a s a ion JL277 (14°3'28"S, 14°20'53"W), Sou he n
Mid-A lan ic Ridge, a a dep h o 1,553 m.
Pa a ypes • FIO-ATL83-JLBEN27712 and FIO-ATL83-
JLBEN27713 we e collec ed oge he wi h holo ype.
Diagnosis. Mush oom-shaped, wi h a sphe ical ca-
pi ulum and a sho and inconspicuous s alk. Polyps a e
dimo phic. Au ozooids a e la ge, e ac ile, and e enly
a anged o e he capi ulum. Siphonozooids a e in isi-
ble, nume ous, densely sca e ed among au ozooids. The
scle i e p o ile is di e se, bu cha ac e ized by he ab-
sence o spindles in all pa s o he colony. Scle i es o he
capi ulum su ace and s alk p edominan ly include mul-
i adia es and pla es. Ten acula scle i es a e composed
p edominan ly o dumbbells and pla es; an hocodial
walls con ain clubs and dumbbells. Pha yngeal scle i es
a e pla ele s wi h a dis inc median wais and c osses.
Desc ip ion o holo ype. Colony o m and size. The
specimen FIO-ATL83-JLBEN27711 has a p ominen
capi ulum suppo ed by an inconspicuous s alk, wi h an
app oxima e 1:1 heigh a io be ween he capi ulum and
s alk, and a o al heigh o 42 mm (Fig. 6A, G). The ca-
pi ulum is dome-shaped, wi h a diame e o 33 mm and a
heigh o 20 mm. The s alk is sho , hick, and cylind ical,
wi h a diame e o 20 mm and a heigh o 22 mm.
Polyps. The e a e 22 la ge au ozooids e enly a anged
o e he capi ulum, ypically spaced abou 8 mm apa .
The au ozooids a e e ac ile, wi h some being ully e-
ac ed while o he s a e con ac ed wi h hei en acles
cu ed inwa d, o ming small sphe ical p o usions on
he su ace o capi ulum (Fig. 5C). The au ozooids a e
la ge, wi h a maximum leng h o 51 mm (including en-
acles, which a e 17 mm long). The pinnules a e a anged
ei he opposi ely o al e na ely, wi h 12–18 pai s. Sipho-
nozooids a e small, nume ous, in isible, and sca e ed
among he au ozooids a in e als o 1.6–2.0 mm.
Scle i es. Ten acula scle i es (Fig. 6A) a e pla es
(0.12–0.21 mm long), ods (0.13–0.23 mm long), dumb-
bells (0.11–0.22 mm long), and a ew ods wi h a sho
ans e se axis (0.14–0.23 mm long). Pha yngeal scle -
i es (Fig. 6B) a e pla ele s wi h a median wais (0.05–
0.08 mm long) and a ew c osses de i ed om small
pla es. Scle i es o he an hocodial wall (Fig. 7A) a e ods
(0.17–0.35 mm long), clubs (0.14–0.19 mm long), dumb-
bells (0.13–0.18 mm long), mul i adia es (0.10–0.14 mm
long), and a ew c osses. Scle i es om he su ace o ca-
pi ulum (Fig. 7B) a e ods (0.18–0.37 mm long), clubs
(0.17–0.23 mm long), mul i adia es (0.11–0.14 mm
long), and a ew c osses. Scle i es om in e io o capi u-
lum (Fig. 7C) a e long ods (0.27–0.49 mm long). Scle -
i es om he su ace o s alk (Fig. 8A) a e mul i adia es
(0.10–0.14 mm long), ods (0.15–0.23 mm long), and a
ew c osses. Scle i es om he in e io o he s alk (Fig.
8B) a e long ods (0.32–0.40 mm long). Scle i es o he
hold as (Fig. 8C) a e ods (0.17–0.31 mm long), mul i a-
dia es (0.09–0.14 mm long), and a ew c osses.
Colo . Li e specimens display a b igh ed colo , while
appea ing e milion in e hanol. The dis al ends o he en-
acles a e yellowish-whi e.
Va ia ion. Pa a ype FIO-ATL83-JLBEN27712 was
a ached o a dead co al skele on and has 20 au ozooids.
The capi ulum is 16 mm high wi h a diame e o 23 mm.
The s alk is sho and inconspicuous, wi h a leng h o 6
mm. Pa a ype FIO-ATL83-JLBEN27713 was also a -
ached o a dead co al skele on and has 28 au ozooids.
The capi ulum is 22 mm high wi h a diame e o 33 mm,
and he s alk is 16 mm high.
E ymology. The La in mi abilis, meaning “wonde -
ul” and “as onishing,” concisely e lec s how his deep-
sea co al’s s iking ed, sphe ical o m wi h long au ozo-
oids cap i a es obse e s wi h i s beau y.
Dis ibu ion and habi a . Known only om he ype
locali y. Sou he n Mid-A lan ic Ridge, 1,553–1,649 m.
Rema ks. The mo phological cha ac e is ics o An-
homas us mi abilis sp. no . ma ch well wi h he ge-
ne ic de ini ion o An homas us. Mo phologically, A.
mi abilis sp. no . is mos closely allied wi h A. g an-
di lo us Ve ill, 1878, showing no ob ious di e ences
in he ex e nal mo phology o he capi ulum, s alk, and
polyps. Bo h species a e g ouped by ha ing pla ele s
as he p edominan pha yngeal scle i es and by ha -
ing au ozooids dis ibu ed ac oss he whole capi ulum.
Howe e , hey a e undamen ally sepa a ed by hei
scle i e composi ion: A. mi abilis sp. no . possesses
dumbbells and mo e pla es in i s en acles, and clubs
and dumbbells in i s an hocodial wall. C ucially, A.
mi abilis sp. no . lacks spindles h oughou he colony
(capi ulum, s alk, and hold as ), which is a key dis inc-
ion om A. g andi lo us, o which spindles a e a cen-
al diagnos ic ea u e.
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2118
and could p o ide edundan o backup unc ions. The
mi ochond ial gene o de o A. mi abilis sp. no . and N.
longis ylus sp. no . is iden ical, excep o h ee duplica ed
genes o A. mi abilis sp. no .. The mi ochond ial genome
a angemen s o hese wo species a e pe ec ly consis en
wi h he japonicum mi ochond ial gene a angemen de-
sc ibed in Figue oa and Baco (2015, shown in hei ig. 1).
This japonicum mi ochond ial gene a angemen is widely
p esen in he amily Co alliidae, se ing as a cha ac e is ic
gene o de o he gene a Co allium and Hemico allium
wi hin his amily, and di e s om he konojoi a ange-
men ound in he genus Pleu oco allium.
Figu e 10. The phylogene ic ee o species wi hin An homas inae based on he pa ial m Mu S gene. Fou gene a ou side o An ho-
mas inae wi hin Co alliidae a e used as ou g oups. The Maximum Likelihood (ML) ee sha es he same opology as he Bayesian
In e ence (BI) ee, wi h node suppo alues p esen ed as: ML boo s ap alues / BI pos e io p obabili ies.
Pa ago gia egalis KP714007
Pleu oco allium secundum MT655220
Co allium o uosum KP714002
Hemico allium egale MW514671
He e opolypus sp. PV153464
He e opolypus sp. PV153465
He e opolypus sp. PV153463
He e opolypus sp. PV153462
He e opolypus sp. KX008404
He e opolypus sp. KX008403
He e opolypus sp. KX008402
He e opolypus sol ON109690
He e opolypus i e i DQ302816
An homas inae sp. OL616208
Neoan homas us elonga us PV175375
Neoan homas us s ella us PV175376
Neoan homas us longis ylus sp. no . PV820546
Neoan homas us longis ylus sp. no . PV820544
Neoan homas us longis ylus sp. no . PV820543
Neoan homas us longis ylus sp. no . PV166046
Neoan homas us longis ylus sp. no . PV820545
An homas us sp. OL616209
Pseudoan homas us o na us PV175378
Pseudoan homas us sp. OL616265
Pseudoan homas us applana us PV175377
An homas us sp. MT655177
An homas us sp. MT655176
An homas us sp. KP324390
An homas us c . g andi lo us KC984603
Pseudoan homas us sp. OM798843
Pseudoan homas us sp. OM798844
Pseudoan homas us sp. OM798842
Ba hyalcyon obus um PV175346
Ba hyalcyon obus um PV175347
Ba hyalcyon obus um PV173768
Ba hyalcyon obus um PV173767
Ba hyalcyon obus um KC984604
An homas us mi abilis sp. no . PV820547
An homas us mi abilis sp. no . PV166047
An homas us g andi lo us OL616289
An homas us c . cana iensis ON109700
An homas us sp. KX008392
An homas us sphae icus PV153460
An homas us sphae icus PV153461
An homas us sphae icus PV153459
An homas us nanhaiensis PV175348
An homas us
Neoan homas us
He e opolypus
Ba hyalcyon
Pseudoan homas us
99/1
93/1
91/1
94/1
96/1
95/1
92/1
85/1
97/0.99
89/0.92
45/0.63
99/1 Ou g oup
T ee scale: 0.1
ML / BI
An homas us mi abilis sp. no . PV820548
Zoosys . E ol. 101 (4) 2025, 2103–2121
zse.penso .ne
2119
Ou phylogene ic analysis p o ides obus suppo o
he di ision o An homas inae in o i e majo clades, and
he clea placemen o ou new species wi hin he ame-
wo k de ined by Li e al. (2025) u he s abilizes he
sub amily’s axonomic sys em. Howe e , b oade ax-
onomic e isions emain challenging, p ima ily due o
he sca ci y o a ailable molecula da a o mos An ho-
mas inae species. Gi en ha his issue is compounded
by po en ially inaccu a e anno a ions in public da abas-
es NCBI, we p io i ized using such sequences o assess
phylogene ic opology and b anching pa e ns, a he
han o de ini i e axonomic alida ion. Fu u e e o s
should ocus on illing da a gaps and desc ibing he de-
ailed cha ac e is ics o e ine he axonomy o An hom-
as inae. Taxonomic s udies should in eg a e bo h molec-
ula da a and mo phological cha ac e is ics o es ablish a
mo e obus classi ica ion amewo k (McFadden e al.
2010; Hu e al. 2025).
In conclusion, his s udy desc ibes wo new deep-
sea mush oom so co al species, Neoan homas us
longis ylus sp. no . and An homas us mi abilis sp.
no ., om he Sou he n Mid-A lan ic Ridge. The mi-
ochond ial genomes o bo h species we e cha ac e -
ized, e ealing a no el duplica ion o h ee iden ical
genes (ND6, ND3, and ND4L) in A. mi abilis sp. no .
Phylogene ic analyses using bo h pa ial m Mu S and
14 PCGs i mly es ablished he e olu iona y posi ions
o hese species wi hin he sub amily An homas inae.
Ou indings signi ican ly en ich he known biodi e -
si y o he Sou he n Mid-A lan ic Ridge and p o ide
c i ical baseline da a o u u e ecological assessmen s
and conse a ion e o s.
Figu e 11. The phylogene ic ee o species wi hin An homas inae in e ed om 13 PCGs in mi ochond ial genomes (m Mu S no
included). Fou gene a ou side o An homas inae wi hin Co alliidae a e used as ou g oups. The Maximum Likelihood (ML) ee
sha es he same opology as he Bayesian In e ence (BI) ee, wi h node suppo alues p esen ed as: ML boo s ap alues / BI
pos e io p obabili ies.
Co allium japonicum AB595189
Co allium ub um AB700136
Pleu oco allium ela ius AB700134
Pleu oco allium konojoi AB595190
Sibogago gia cauli lo a KM015354
Pa ago gia co alloides KF785800
An homas us ongin PV795867
Pseudoan homas us sp. OL616264
Neoan homas us longis ylus sp. no . PV795866
Neoan homas us longis ylus sp. no . PV158192
Neoan homas us longis ylus sp. no . PV795865
Neoan homas us s ella us PV175376
An homas us sp. ANT22 OL616209
Neoan homas us elonga us PV175375
Pseudoan homas us sp. ANT287 OL616265
Pseudoan homas us o na us PV175378
An homas us sp. USNM1171062 KM015352
Pseudoan homas us applana us PV175377
An homas us sp. USNM1081145 KM015353
Ba hyalcyon obus um PV173767
Ba hyalcyon obus um PV173768
Ba hyalcyon obus um PV175346
Ba hyalcyon obus um PV175347
He e opolypus sp. PV153464
He e opolypus sp. PV153465
He e opolypus sp. PV153463
He e opolypus sp. PV153462
An homas us sphae icus PV153459
An homas us sphae icus PV153461
An homas us sphae icus PV153460
An homas us nanhaiensis PV175348
An homas us mi abilis sp. no . PV795869
An homas us mi abilis sp. no . PV795868
An homas us mi abilis sp. no . PV795867
An homas us g andi lo us OL616289
91/0.97
100/1
100/1
100/0.97
93/0.98
100/1
100/1
99/0.97
100/1
98/1
100/0.97
99/0.97
0.34
0.16
100/0.97
99/1
100/1
100/1
./0.84
100/1
100/1
100/0.97
100/1
100/1
98/1
96/0.82
100/0.97
T ee scale: 0.01
ML / BI
An homas us
Neoan homas us
He e opolypus
Ba hyalcyon
Pseudoan homas us
Ou g oup
zse.penso .ne
Li, X. e al.: New species o deep-sea mush oom so co als2120
Au ho con ibu ions
LXL conduc ed he analyses and d a ed he manusc ip .
XQZ designed and supe ised he p ojec . XQZ and
LMN collec ed he samples. LXL, XNX, GML and HXY
pe o med he labo a o y wo k. XQZ, XNX, GML, HXY,
LMN, WZL, and ZXL e iewed and edi ed he manu-
sc ip , wi h all au ho s con ibu ing o i s e ision.
Da a a ailabili y s a emen
The da a o his s udy a e a ailable om he co espond-
ing au ho upon easonable eques .
Acknowledgemen s
We app ecia e he in aluable suppo p o ided by he Na-
ional Key R&D P og am o China (No. 2023YFC2812903)
and Digi al Deep-sea Typical Habi a s P og amme o Chi-
na Deep Ocean A ai s Adminis a ion. We a e g a e ul o
all scien is and c ew membe s o ShenhaiYihao and he
pilo s o HOV Jiaolong du ing he c uise o DY83 o he
sample collec ion, especially Yue Dong (Fi s Ins i u e o
Oceanog aphy, MNR), Yadong Zhou (Second Ins i u e o
Oceanog aphy, MNR), Chengbing Song (Na ional Deep
Sea Cen e , MNR). We a e g a e ul o Wenle Du (In e na-
ional Depa men , Qingdao Academy) o he expe imen-
al wo k and image p ocessing e o s.
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Supplemen a y ma e ial 1
Supplemen a y ables
Au ho s: Xinlong Li, Ningxia Xu, Meiling Ge, Xuying Hu,
Mengna Li, Zongling Wang, Xuelei Zhang, Qinzeng Xu
Da a ype: xlsx
Explana ion no e: able S1. P ime lis and PCR he mal
p o iles; able S2. The bes models o Bayesian and
Maximum Likelihood analysis; able S3. NCBI acces-
sion numbe s o all samples; able S4. Es ima es o
e olu iona y di e gence be ween he species in Neoan-
homas us measu ed by Kimu a 2-pa ame e dis ances
(%) based on 14 p o ein coding genes o mi ochond ial
genomes; able S5. Es ima es o e olu iona y di e -
gence be ween he species in An homas us measu ed
by Kimu a 2-pa ame e dis ances (%) based on 14
p o ein coding genes o mi ochond ial genomes; able
S6. Gene agmen s o An homas us mi abilis sp. no .
ob ained h ough Sange sequencing.
Copy igh no ice: This da ase is made a ailable unde
he Open Da abase License (h p://openda acommons.
o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow us-
e s o eely sha e, modi y, and use his Da ase while
main aining his same eedom o o he s, p o ided
ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/zse.101.163559.suppl1
