Late Permian nautiloids from Baghuk Mountain (Central Iran)

La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
Die e KORN 1,* & Vachik HAIRAPETIAN 2,*
1 Museum ü Na u kunde, Leibniz Ins i u o Resea ch on E olu ion and Biodi e si y,
In alidens aße 43, 10115 Be lin, Ge many.
2 Depa men o Geology, Is ahan (Kho asgan) B anch, Islamic Azad Uni e si y, Is ahan, I an.
* Co esponding au ho s: die e [email p o ec ed], [email p o ec ed]
Abs ac . The Wuchiapingian o Changhsingian (La e Pe mian) Hambas Fo ma ion o sec ions a
Baghuk Moun ain and Shah eza (Cen al I an) has yielded di e se nau iloid assemblages om he
ea ly Wuchiapingian A axoce as beds (nine species), he ea ly Wuchiapingian P o o oce as beds
(one species), he la e Wuchiapingian Vedioce as beds ( wel e species) and he la e Changhsingian
Pa a i oli es beds ( wo species). These species belong o 14 gene a, six o which a e new: Lu onau ilus
gen. no ., Epi ainoce as gen. no ., Ocunau ilus gen. no . in Ocunau ilidae am. no ., Ai inau ilus
gen. no ., Baghuknau ilus gen. no . and Shah ezanau ilus gen. no . A o al o 24 species a e desc ibed,
all o which a e new: Se ome acoce as pen agonum sp. no ., Lu onau ilus c a us gen. e sp. no .,
Lu onau ilus elachus gen. e sp. no ., Lu onau ilus cymus gen. e sp. no ., Tainoce as hys a um sp. no .,
Epi ainoce as lu ense gen. e sp. no ., Foo dice as eicosacan hum sp. no ., Foo dice as decacan hum
sp. no ., Foo dice as asce um sp. no ., Ta dunau ilus ape imos sp. no ., Doma oce as myloide sp. no .,
Doma oce as canonium sp. no ., Doma oce as ocomphalum sp. no ., Ocunau ilus diplodocus gen. e
sp. no ., Ocunau ilus coelodesmus gen. e sp. no ., Ocunau ilus achy ephus gen. e sp. no ., Ai inau ilus
icanus gen. e sp. no ., Aza inau ilus pho minx sp. no ., Li oce as lep um sp. no ., Pa ali oce as
mac ogas e sp. no ., Pe monau ilus adelphidus sp. no ., Baghuknau ilus aplomo phus gen. e sp. no .,
Shah ezanau ilus weye i gen. e sp. no . and Shah ezanau ilus ghade ii gen. e sp. no . This nau iloid
assemblage is ema kable o i s high endemism. Al hough many gene a known om he T anscaucasian
egion a e ep esen ed, he e is no co espondence a he species le el.
Keywo ds. Nau iloidea, Nau ilida, Pe mian, I an, mo phology.
Ko n D. & Hai ape ian V. 2025. La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an). Eu opean Jou nal
o Taxonomy 1019: 1–76. h ps://doi.o g/10.5852/ej .1019.3071
In oduc ion
The aunal ansi ion du ing he se e e mass ex inc ion e en a he ansi ion o he Palaeozoic in o
he Mesozoic is one o he majo opics in he s udy o he e olu iona y his o y o he biosphe e, bu
nau iloids ha e so a played only a mino ole in hese s udies. This unde - ep esen a ion o nau iloids
may be due o he a he pa chy s a e o knowledge; compa ed o o he g oups o ossil o ganisms,
1
Eu opean Jou nal o Taxonomy 1019: 1–76
h ps://doi.o g/10.5852/ej .2025.1019.3071
eu opeanjou nalo axonomy.eu
ISSN 2118-9773
2025 · Ko n D. & Hai ape ian V.
This wo k is licensed unde a C ea i e Commons A ibu ion License (CC BY 4.0)
Recei ed: 25 Sep embe 2024 • Accep ed: 7 Ap il 2025 • Published: 24 Sep embe 2025
Topic edi o : Ma ie-Béa ice Fo el • Desk edi o : K is iaan Hoedemake s
Monog aph
u n:lsid:zoobank.o g:pub:E24EBF4A-9FE2-47E4-A656-E1698F88BB41
nau iloids ha e a ely been monog aphically desc ibed and ha e he e o e had li le o con ibu e o he
in es iga ion o he la ges mass ex inc ion in he Phane ozoic.
I has been shown in ecen yea s ha he disco e y o new Ca boni e ous and Pe mian nau iloid
assemblages has led o a signi ican inc ease in species numbe s (Ba sko e al. 2014; Niko & Mapes
2016a; Leono a & Shchedukhin 2020; Miao e al. 2021; Ko n & Bockwinkel 2022; Ko n e al. 2022;
Ko n & Klug 2023). Almos e e y new nau iloid assemblage has yielded p edominan ly new, p e iously
unknown species. These ecen s udies ha e also shown ha a s ic e applica ion o mo phome ic
me hods and mo e p ecise s a ig aphic assignmen can p o ide a much be e insigh in o he
e olu iona y his o y o nau iloids. These s udies ha e also led o imp o ed insigh s in o ques ions o
on ogene ic de elopmen , in aspeci ic a ia ion, and s a ig aphic and geog aphic ange.
La e Pe mian coiled ammonoids we e occasionally desc ibed in pionee ing s udies om a ious egions
such as Saxony (Geini z 1841), England (King 1850) and he Dolomi es (Mojsiso ics 1869; S ache
1877), bu mo e di e se assemblages we e known om only a ew egions. Those om he o me
A menian pa o he T anscaucasus we e he i s e e desc ibed (Abich 1878; on A habe 1900),
ollowed by he Sal Range in Pakis an (Waagen 1879) and Sou h China (Kayse 1883). These pionee ing
s udies ha e been complemen ed in a la ge numbe o subsequen publica ions o he T anscaucasus
(Shimansky 1965; Teiche & Kummel 1973), he Sal Range (Reed 1931, 1944), he Dolomi es (Me la
1930; P ino h & Posena o 2007) and Sou h China (Zhao e al. 1978; Liang 1984; Zheng 1984; Miao
e al. 2021). These s udies ha e signi ican ly imp o ed ou knowledge o nau iloids be o e he mass
ex inc ion a he Pe mian-T iassic bounda y. An o e iew o hei mo phological ange and mo phospace
occupa ion has al eady been ob ained (Ko n e al. 2020).
I was no un il he 1970s ha La e Pe mian nau iloids om he e i o y o I an became known.
Teiche & Kummel (1973) desc ibed a numbe o axa, mos ly in open nomencla u e, om a ound Jul a
(NW I an) and Ta az e al. (1981) lis ed some nau iloids om he Pe mian–T iassic bounda y sec ions
nea Abadeh (cen al I an). New inds in he Jul a a ea allowed Ko n & Ghade i (2025) o ca y ou a
Fig. 1. Geog aphic posi ion o Pe mian–T iassic bounda y sec ions, including Baghuk Moun ain, in
Cen al I an ( om Ko n e al. 2021b).
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
2
monog aphic s udy; 30 species om he Wuchiapingian and Changhsingian in e als we e desc ibed in
ha monog aph, 24 o hem new.
He e, we p esen a monog aphic desc ip ion o he coiled nau iloids om he Wuchiapingian and
Changhsingian o ma ions o Baghuk Moun ain. This monog aph complemen s he p e iously
published monog aph on he ish emains (Hampe e al. 2013) and he monog aph on he Changhsingian
ammonoids (Ko n e al. 2021a). In a pa allel a emp , he coiled nau iloids om he Jul a egion a e
desc ibed in ano he monog aph (Ko n & Ghade i 2025).
Ma e ial and me hods
A o al o 55 nau ilid specimens we e s udied. All bu one o he specimens desc ibed he e a e om he
Baghuk Moun ain a ea 140 km SSE o Es ahan and 50 km NNW o Abadeh (Fig. 1). The indi idual
sec ions o Baghuk Moun ain ha e been desc ibed by Ko n e al. (2021b) and Heue e al. (2022). All
he ma e ial comes om he Hambas Fo ma ion, which spans almos he en i e Wuchiapingian and
Changhsingian s ages (Ta az e al. 1981).
The majo i y o he specimens we e collec ed om loa ma e ial below he ou c ops. Howe e , all o
he specimens desc ibed he e can be assigned wi hou doub o a membe o he Hambas Fo ma ion on
he basis o hei li hology. Fou membe s we e dis inguished om bo om o op (Fig. 2):
(1) A axoce as beds (ea ly Wuchiapingian): da k g ey mic i ic nodula limes one (25 specimens).
• Doma oce as myloide sp. no . – 4 specimens
• Ocunau ilus diplodocus gen. e sp. no . – 3 specimens
Fig. 2. S a ig aphical posi ions o he La e Pe mian nau iloid axa om Baghuk Moun ain and Shah eza.
A ax = A axoce as beds; P o = P o o oce as beds; Vedi = Vedioce as beds; Pa a = Pa a i oli es beds.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
3
• Ocunau ilus coelodesmus gen. e sp. no . – 3 specimens
• Ocunau ilus achy ephus gen. e sp. no . – 1 specimen
• Ai inau ilus icanus gen. e sp. no . – 5 specimens
• Se ome acoce as pen agonum sp. no . – 1 specimen
• Epi ainoce as lu ense gen. e sp. no . – 2 specimens
• Pa ali oce as mac ogas e sp. no . – 2 specimens
• Pe monau ilus adelphidus sp. no . – 4 specimens
(2) P o o oce as beds (ea ly Wuchiapingian): ligh g ey pla y limes one wi h nume ous py i e c ys als
(one specimen).
• Ta dunau ilus ape imos sp. no . – 1 specimen
(3) Vedioce as beds (la e Wuchiapingian): pink o ligh ed nodula limes one (26 specimens).
• Doma oce as canonium sp. no . – 5 specimens
• Doma oce as ocomphalum sp. no . – 1 specimen
• Foo dice as eicosacan hum sp. no . – 2 specimens
• Foo dice as decacan hum sp. no . – 1 specimen
• Foo dice as asce um sp. no . – 1 specimen
• Lu onau ilus c a us gen. e sp. no . – 4 specimens
• Lu onau ilus elachus gen. e sp. no . – 2 specimens
• Lu onau ilus cymus gen. e sp. no . – 3 specimens
• Li oce as lep um sp. no . – 3 specimens
• Baghuknau ilus aplomo phus gen. e sp. no . – 2 specimens
• Shah ezanau ilus weye i gen. e sp. no . – 1 specimen
• Shah ezanau ilus ghade ii gen. e sp. no . – 1 specimen
(4) Pa a i oli es beds (la e Changhsingian): ed-g ey o g ey- iole nodula limes one ( h ee specimens).
• Aza inau ilus pho minx sp. no . – 1 specimen
• Tainoce as hys a um sp. no . – 2 specimens
Taxonomic concep
Fo he o de Nau ilida in cu en unde s anding, se e al pa ially con lic ing concep s o sys ema ics
ha e been published and modi ied o e he decades. While Flowe & Kummel (1950) and Kummel
(1953, 1964) ook a mo e conse a i e app oach, Shimansky (1962a, 1962b, 1965, 1967) p oposed a
Fig. 3. Conch and su u e line pa ame e s used in he axonomic desc ip ions. A. Conch pa ame e s.
B. Desc ip i e e ms o who l p o iles. C. Su u e line e minology.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
4
much mo e de ailed scheme. Dzik (1984) discussed he phylogeny and i s implica ions o he sys ema ic
scheme in g ea de ail and d ew a much mo e complex e olu iona y his o y o he coiled nau iloids.
In a e ision o he Ca boni e ous and Pe mian coiled nau iloids by Ko n (2025), a new sys ema ic
scheme was p esen ed. This scheme is la gely based on he phylogene ic conside a ions o Ruzhence &
Shimansky (1954) as well as Dzik (1984) and is la gely based, wi h some modi ica ions, on he sys ema ic
scheme o Shimansky (1967, 1979). This also means ha i di e s om he o en used scheme o he
T ea ise on In e eb a e Paleon ology (Kummel 1964).
The desc ip ion o he specimens ollows he e minology o conch, o namen and su u e line p oposed
by Ko n (2010) and Klug e al. (2015) o he cha ac e isa ion o ammonoids (Fig. 3). The e minology o
conch geome y used he e la gely co esponds o ha p oposed by Teiche (1964). The only di e ences
conce n he ollowing e ms: umbilical angle o shoulde (= umbilical ma gin) and umbilical a ea
(= umbilical wid h in ou desc ip ions).
Pho og aphy echnique
The pho og aphs o his monog aph we e aken using Re lec ance T ans o ma ion Imaging (RTI)
echniques wi h he B oncolo Scope D50 dome o c ea e su ace models wi hou p e iously whi ening
he specimens. P ocessing o he images was pe o med using he T u is Au hen ica C ea o p og am.
The expo ed images we e p ocessed in Adobe Pho oshop.
Abb e ia ions used in he species desc ip ions
ah = ape u al heigh
CLI = chambe leng h index (in deg ees)
dm = conch diame e
IZR = imp in zone a e
uw = umbilical wid h
WER = who l expansion a e
wh = who l heigh
ww = who l wid h
Abb e ia ion o he eposi o y
MB.C. = Cephalopod collec ion o he Museum ü Na u kunde, Be lin
Resul s
Class Cephalopoda Cu ie , 1795
Subclass Nau iloidea Agassiz, 1847
O de Nau ilida Agassiz, 1847
Diagnosis
Exogas ically cu ed o coiled nau iloids wi h a conch shape anging om gy oconic o cy oconic o
mo e o less igh ly coiled. Shell su ace smoo h o sculp u ed wi h a a ie y o elemen s ( ibs, nodes,
spines, longi udinal idges o lines). Sep a simply domed in mos species, wi h he shape o he who l
p o ile p oducing su u e lines wi h a iable lobes and saddles. Va ia ions in sep al shape wi h in lexions
p oducing deep lobes in some gene a. Sep al necks sho and s aigh , a ely sligh ly widened. Connec ing
ings cylind ical o beaded. Siphuncula o came al deposi s absen . Ju enile conch wi h cup-shaped
ini ial chambe and na ow siphuncle. Mo phological e olu ion includes he deg ee o coiling, he shape
and size o he ju enile and adul conch and he su u e line (a e Shimansky 1962b; emended).
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
5

Included subo de s
Nau ilina Agassiz, 1847 (Ju assic o Recen ); Solenochilina Flowe , 1950 (Ca boni e ous o Pe mian);
Li oce a ina Flowe , 1955 (Ca boni e ous o Ju assic); Ru oce a ina Shimansky, 1957 (De onian);
Tainoce a ina Shimansky, 1957 (Ca boni e ous o T iassic); Temnocheilina Flowe , 1963 (De onian o
Pe mian); Doma oce a ina Ko n, 2025 (Ca boni e ous o T iassic).
Subo de Doma oce a ina Ko n, 2025
Diagnosis
Subo de o he o de Nau ilida, in which a en ola e al shoulde and an umbilical ma gin a e o med
ea ly in on ogeny. Conch usually discoidal, subin olu e o e olu e. Ju enile who l p o ile ci cula . Adul
who l p o ile subquad a e o in e ed apezoidal wi h a dis inc en ola e al shoulde and a dis inc
umbilical ma gin in he ea ly species, showing modi ica ions du ing e olu ion including a conca e en e
in some de i ed species. Do sal who l zone always p esen , bu usually e y small excep o some
de i ed species. Ju enile sculp u e some imes wi h adial ibs on he lank; adul sculp u e is usually
lacking excep o elonga e en ola e al ube cles in de i ed species. Sep a simply domed in mos o
he species; wi h sep al in lexion and co uga ed sep a in some lineages. Su u e line usually depending
on he who l p o ile, usually wi h shallow lobes and low saddles; wi h dis inc lobes in some de i ed
lineages ( om Ko n 2025).
Included supe amilies
G ypoce a oidea Hya , 1900 (Ca boni e ous o T iassic); Pe moce a oidea Mille & Collinson, 1953
(Pe mian); Subclymenioidea Shimansky, 1962 (Ca boni e ous).
Rema ks
A de ailed discussion o he Doma oce a ina has been gi en by Ko n (2025).
Supe amily G ypoce a oidea Hya , 1900
Diagnosis
Supe amily o he subo de Doma oce a ina wi h a discoidal, subin olu e o e olu e conch. Who l
p o ile usually in e ed apezoidal wi h a dis inc en ola e al shoulde and a dis inc umbilical ma gin.
De i ed species show a a ia ion o modi ica ions including a conca e en e , a skid-like en ola e al
shoulde and an angula umbilical ma gin. Who l o e lap ex emely small o mode a e. Sculp u e in mos
species lacking, in some species wi h sho la e al ibs o en ola e al nodes. Sep a simply domed; su u e
line s ongly dependen on he who l p o ile, usually wi h b oadly ounded lobes and na owly ounded
o subangula saddles ( om Ko n 2025).
Included amilies
G ypoce a idae Hya , 1900; Doma oce a idae Mille & Youngquis , 1949; Ocunau ilidae am. no .
Rema ks
A discussion o he G ypoce a oidea has been gi en by Ko n (2025).
Family Doma oce a idae Mille & Youngquis , 1949
Diagnosis
Family o he supe amily G ypoce a oidea wi h a hinly o hickly discoidal, subin olu e o e olu e
conch. Who l p o ile in he adul s age usually comp essed subquad a e o in e ed apezoidal. Umbilical
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
6
ma gin dis inc o sha p; en ola e al shoulde nea ly ec angula o b oadly ounded, a ely skid-like.
O namen consis ing o ine g ow h lines; some species ha e ube cles on he en ola e al shoulde .
Su u e line always wi h ounded bu dis inc ex e nal, la e al and in e nal lobes sepa a ed by a na owly
ounded o subacu e saddles; wi hou annula p ocess ( om Ko n 2025).
Included gene a
Pselioce as Hya , 1884 (Pe mian); Ti anoce as Hya , 1884 (Ca boni e ous); Doma oce as Hya ,
1891 (Ca boni e ous o Pe mian); Pseudome acoce as Mille , Dunba & Cond a, 1933 [synonym
o Doma oce as Hya , 1891]; Pa adoma oce as Delépine, 1937 (Ca boni e ous); Plumme oce as
Kummel, 1953 (Pe mian); Neodoma oce as Ruzhence & Shimansky, 1954 (Pe mian); Neos enopoce as
Zhao, Liang & Zheng, 1978 (Pe mian); Pa apenascoce as Ruzhence & Shimansky, 1954 (Pe mian);
Pa as enopoce as Ruzhence & Shimansky, 1954 (Pe mian); Penascoce as Ruzhence & Shimansky,
1954 (Pe mian); Pe modoma oce as Ruzhence & Shimansky, 1954 (Pe mian); S enodoma oce as
Ruzhence & Shimansky, 1954 (Pe mian); Vi galoce as Schindewol , 1954 (Pe mian); Neos enopoce as
Zhao, Liang & Zheng, 1978 (Pe mian); Sha oce as Leono a & Shchedukhin, 2020 (Pe mian);
Omo phoce as Leono a & Shchedukhin, 2023 (Pe mian); Fididoma oce as Ko n & Ghade i, 2025
(Pe mian).
Rema ks
A de ailed discussion o he Doma oce a idae has been gi en by Ko n (2025).
Genus Doma oce as Hya , 1891
Type species
Doma oce as umbilica um Hya , 1891; o iginal designa ion.
Diagnosis
Genus o he amily Doma oce a idae wi h a subin olu e o e olu e conch. High o ex emely high
coiling a e; who l p o ile usually weakly comp essed. Ven e la ened o weakly conca e, lanks usually
la ened and sligh ly con e gen ; umbilical ma gin ounded o angula . Wi hou sculp u e excep o
small en ola e al ube cles in some species. Su u e line wi h small and shallow ex e nal lobe and
b oadly ounded la e al lobe; wi hou annula p ocess.
Included Ca boni e ous species
No h Ame ica (Meek & Wo hen 1865; Wo hen & Meek 1875; Gu ley 1883; Hya 1891; Mille &
Owen 1934; S u geon & Mille 1948; Tucke 1976; Tucke & Mapes 1978; S u geon e al. 1982;
Niko & Mapes 2016b; Niko e al. 2022): Nau ilus Lasallensis Meek & Wo hen, 1865, Kasimo ian,
Illinois; Nau ilus (Disci es) highlandensis Wo hen in Wo hen & Meek, 1875, Mosco ian, Ohio;
Disci es Toddanus Gu ley, 1883, Kasimo ian, Missou i; Doma oce as umbilica um Hya , 1891,
Mosco ian, Kansas; Doma oce as williamsi Mille & Owen, 1934, Mosco ian, Ohio; Doma oce as
obsole um S u geon, 1946, Mosco ian, Ohio; Doma oce as shephe di S u geon, 1948, Mosco ian,
Ohio; Doma oce as wo heni Tucke , 1976, Kasimo ian, Illinois; Doma oce as exanum Tucke &
Mapes, 1978, Kasimo ian, Texas; Doma oce as o esko ichi S u geon, Windle, Mapes & Hoa e,
1982, Mosco ian, Ohio; Doma oce as collins illense Niko & Mapes, 2016, Mosco ian, Oklahoma;
Doma oce as ucke i Niko, Mapes & Seuss, 2022, Kasimo ian, Texas.
Sou h China (Ruan & Zhou 1987): Doma oce as quad a um Ruan & Zhou, 1987, Bashki ian, Ningxia.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
7
Wes Russia (Tzwe ae 1888, 1898; Shimansky 1967): Nau ilus podolskensis Tzwe ae , 1888,
Mosco ian, Moscow Basin; Nau ilus mosquensis Tzwe ae , 1898, Mosco ian, Moscow Basin;
Doma oce as (Doma oce as) magis e Shimansky, 1967, Mosco ian, Moscow Basin.
Included Pe mian species
U als (Leono a & Shchedukhin 2020): Doma oce as bashki icum Leono a & Shchedukhin, 2020,
Asselian o Sakma ian, Sou h U als; Doma oce as s e li amakense Leono a & Shchedukhin, 2020,
Asselian o Sakma ian, Sou h U als.
T anscaucasia and I an (Abich 1878; Shimansky 1965; his pape ): Nau ilus pa allelus Abich, 1878,
Wuchiapingian, Aze baijan; Nau ilus con e gens Abich, 1878, Wuchiapingian, Aze baijan; Doma oce as
g acile Shimansky, 1965, Wuchiapingian, Aze baijan; Doma oce as a ypicum Shimansky, 1965,
Wuchiapingian, Aze baijan; Doma oce as mul i ube cula um Ko n & Ghade i, 2025, Wuchiapingian,
NW I an; Doma oce as canonium sp. no ., Wuchiapingian, Cen al I an; Doma oce as ocomphalum
sp. no ., Wuchiapingian, Cen al I an; Doma oce as myloide sp. no ., Wuchiapingian, Cen al I an.
Sou h China (Zhao e al. 1978): Doma oce as guangxiense Zhao, Liang & Zheng, 1978, Changhsingian,
Guangxi.
Japan (Ehi o & Takizawa 1989): Doma oce as oga suense Ehi o & Takizawa, 1989, Wuchiapingian,
Ki akami Massi .
Rema ks
The species o he genus Doma oce as ha e a e y simila conch mo phology om he Mosco ian o he
Changhsingian, sugges ing a e y conse a i e e olu iona y lineage. Shimansky (1967) also included
some Se pukho ian species in Doma oce as, bu hese di e om he ypical ep esen a i es o he
genus in ha hey ha e a wide conch and lack a subangula o angula en ola e al shoulde . These
s a ig aphically olde species may belong o Epidoma oce as Tu ne , 1954.
The simila i y in he mo phology o he conchs, he i ual absence o sculp u al elemen s and he ac
ha he su u e line depends essen ially on he shape o he who l p o ile make i di icul o dis inguish
he species wi hin Doma oce as. Ano he di icul y is ha he species o he genus occu in wo phases:
he i s phase ex ends om he Mosco ian o he Asselian–Sakma ian bounda y, and he second phase
ex ends om he Wuchiapingian o he Changhsingian. I should be no ed, howe e , ha o he La e
Pe mian species, abou en a e known om he Wuchiapingian and only one om he Changhsingian.
Appa en ly, no species a e known om he long in e al be ween he Sakma ian and he Capi anian.
Some species om he Kungu ian and Roadian, which we e placed unde Doma oce as by Mille &
Unklesbay (1942) and Mille & Youngquis (1949), ha e been assigned o o he gene a such as
Penascoce as by Ruzhence & Shimansky (1954).
Doma oce as myloide sp. no .
u n:lsid:zoobank.o g:ac :78A31C35-C06A-4B4C-9A86-137219AC51C5
Fig. 4; Table 1
Diagnosis
Species o Doma oce as wi h hinly discoidal, sube olu e conch (ww/dm ~ 0.35; uw/dm ~ 0.35), weakly
comp essed who l p o ile (ww/wh ~ 0.90) and e y high coiling a e (WER ~ 2.45) a a conch diame e o
70 mm. Who l p o ile wi h weakly la ened en e , ounded en ola e al shoulde , la ened and nea ly
pa allel lanks, subangula umbilical ma gin and s eep and la umbilical wall. Who l o e lap e y small.
Wi hou sculp u e. Su u e line wi h a small and e y shallow ex e nal lobe and a much la ge and deepe ,
b oadly ounded la e al lobe.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
8
E ymology
F om he G eek ‘μυλοειδής’ = ‘mills one-shaped’; because o he conch shape.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 4; MB.C.32101.
Pa a ypes
IRAN – Es ahan P o ince • 3 specimens; same da a as o holo ype; 2010; Ko n e al. leg.; MB.C.32102
o MB.C.32104.
Desc ip ion
Holo ype MB.C.32101 is a ully chambe ed in e nal mould specimen wi h a conch diame e o 72 mm,
which allows he s udy o one who l (Fig. 4B). The conch is hinly discoidal and sube olu e (ww/
dm = 0.35; uw/dm = 0.34) wi h an ex emely high coiling a e (WER = 2.47); he who l p o ile is weakly
comp essed (ww/wh = 0.90) wi h a weakly la ened en e , a ounded en ola e al shoulde , la ened
Fig. 4. Doma oce as myloide sp. no ., holo ype MB.C.32101 (Ko n e al. 2011 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. Do sal, la e al and en al iews. C. Su u e line a dm = 68.5 mm, ww = 22.5 mm, wh = 26.5 mm. Scale
ba uni s = 1 mm.
Table 1. Conch dimensions (in mm) and a ios o Doma oce as myloide sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32101 72.1 25.0 27.8 24.6 26.2 0.35 0.90 0.34 2.47 0.06
MB.C.32101 47.2 17.3 19.2 16.0 18.2 0.37 0.90 0.34 2.65 0.05
MB.C.32102 56.3 20.2 21.5 20.5 20.5 0.36 0.94 0.36 2.47 0.05
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
9
Rema ks
Ocunau ilus gen. no . can ha dly be con used wi h any o he genus o La e Pe mian nau ilids
because o i s e y cha ac e is ic conch shape. The a he sha p umbilical ma gin, he la ened and
apidly con e ging lanks, he angula , skid-like en ola e al shoulde and he mo e o less clea ly
conca e en e allow a clea sepa a ion. Pseudo i anoce as is simila o Ocunau ilus, bu his genus is
dis inguished by he p esence o ube cles on he en ola e al shoulde . Shimansky (1965) men ioned
ha among his P. a meniacum ma e ial he e we e also specimens wi hou en ola e al ube cles.
Pe haps hese do belong o Ocunau ilus a he han Pseudo i anoce as.
Ocunau ilus diplodocus gen. e sp. no .
u n:lsid:zoobank.o g:ac :46279074-DA8A-42D6-B45C-8C639250C01A
Fig. 8; Table 4
Diagnosis
Species o Ocunau ilus gen. no . wi h hinly discoidal, subin olu e conch (ww/dm ~ 0.42; uw/dm ~ 0.20),
weakly comp essed who l p o ile (ww/wh ~ 0.85) and ex emely high coiling a e (WER ~ 2.95) a a
conch diame e o 100 mm. Who l p o ile in e ed apezoidal wi h deeply conca e en e , angula
en ola e al shoulde , la ened and weakly con e gen lanks, angula umbilical ma gin and s eep and
weakly conca e umbilical wall. Who l o e lap small (IZR ~ 0.12). Wi hou sculp u e. Su u e line wi h
a e y deep, ounded V-shaped ex e nal lobe, an angula en ola e al saddle, a shallow and b oadly
ounded la e al lobe and a e y small umbilical lobe.
E ymology
F om he G eek ‘διπλόος’ = ‘double’ and ‘ὀξύς’ = ‘sha p’; because o he double-keeled en e .
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2013; Ko n e al. leg.; illus a ed in Fig. 8; MB.C.32110.
Pa a ypes
IRAN – Es ahan P o ince • 2 specimens; same da a as o holo ype; 2010; Ko n e al. leg.; MB.C.32111,
MB.C.32112.
Desc ip ion
Holo ype MB.C.32110 is a agmen showing sligh ly mo e han a qua e who l o he ph agmocone
and a poo ly p ese ed piece o he penul ima e who l (Fig. 8A). The agmen has a maximum who l
heigh o 51 mm; om his, a diame e o 105 mm can be econs uc ed. On he basis o his calcula ed
diame e , he conch pa ame e s can be de e mined. The conch is hinly discoidal and subin olu e (ww/
dm ~ 0.42; uw/dm ~ 0.21) wi h an ex emely high coiling a e (WER ~ 2.96). The who l p o ile is
weakly comp essed (ww/wh = 0.87) wi h a deeply conca e en e ha is bo de ed by a sha p, skid-like
en ola e al shoulde ; he lanks a e la ened and con e gen . Mos conspicuous is he angula umbilical
ma gin, which bo de s he weakly conca e, incu ed umbilical wall (Fig. 8B). The who l o e lap is small
(IZR = 0.14). The specimen does no show any sculp u e. The su u e line is s ongly in luenced by he
ou line o he who l p o ile; i is cha ac e is ic because o he na ow and deep, V-shaped ex e nal lobe,
which is sepa a ed om he b oadly ounded la e al lobe by a subangula en ola e al saddle (Fig. 8C).
The las qua e olu ion o he ph agmocone consis s o se en chambe s (CLI = 13); he e occu s weak
sep al app oxima ion owa d he ape u e.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
16

Rema ks
Ocunau ilus diplodocus gen. e sp. no . di e s om he o he wo species O. coelodesmus gen. e sp. no .
and O. achy ephus gen. e sp. no . om Baghuk Moun ain by he much mo e deeply conca e en e ;
acco dingly, he ex e nal lobe o O. diplodocus is also much deepe and na owe .
Fig. 8. Ocunau ilus diplodocus gen. e sp. no ., holo ype MB.C.32110 (Ko n e al. 2013 Coll.) om
he A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Ven al and la e al iews.
B. Recons uc ion o ape u al iew. C. Su u e line a ww = 42.5 mm, wh = 46.0 mm. Scale ba
uni s = 1 mm.
Table 4. Conch dimensions (in mm) and a ios o Ocunau ilus diplodocus gen. e sp. no .; econs uc ed
alues in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32110 105.0 44.5 50.9 22.5 44.0 0.42 0.87 0.21 2.96 0.14
MB.C.32111 84.0 32.2 38.2 23.3 34.4 0.38 0.84 0.28 2.87 0.10
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
17
Ocunau ilus coelodesmus gen. e sp. no .
u n:lsid:zoobank.o g:ac :FC07FF9A-4B9C-4162-8BFD-9B43A2E3052C
Fig. 9; Table 5
Diagnosis
Species o Ocunau ilus gen. no . wi h hickly discoidal, subin olu e conch (ww/dm ~ 0.55; uw/dm
~ 0.25), weakly dep essed who l p o ile (ww/wh ~ 1.10) and ex ao dina ily high coiling a e (WER
~ 3.40) a a conch diame e o 60 mm. Who l p o ile in e ed apezoidal wi h mode a ely conca e en e ,
subangula en ola e al shoulde , la ened and s ongly con e gen lanks, angula umbilical ma gin and
s eep and la umbilical wall. Who l o e lap small (IZR ~ 0.08). Wi hou sculp u e, bu ju enile conch
wi h weak, backwa dly di ec ed ibs on he inne lank, ending in low and blun nodes on he mid lank.
Su u e line wi h a a he deep, ounded V-shaped ex e nal lobe, an angula en ola e al saddle, a shallow
and b oadly ounded la e al lobe and a e y small umbilical lobe.
E ymology
F om he G eek ‘κόιλος’ = ‘hollow’ and ‘δεσμός’ = ‘band, b ace’; because o he shape o he en e .
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 9; MB.C.32113.
Pa a ypes
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; MB.C.32114.
Desc ip ion
Holo ype MB.C.32113 is an incomple e, almos ully chambe ed specimen wi h a conch diame e o
57 mm (Fig. 9B). I is p ese ed as an in e nal mould, bu shows some shell emains. The conch is, a
a conch diame e o 57 mm, hickly discoidal and subin olu e (ww/dm = 0.55; uw/dm = 0.23) wi h an
ex ao dina ily high coiling a e (WER = 3.38). The e occu s an on ogene ic dec ease o he coiling a e;
Fig. 9. Ocunau ilus coelodesmus gen. e sp. no ., holo ype MB.C.32113 (Ko n e al. 2011 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. La e al and en al iews. C. Su u e line a dm = 37.0 mm, ww = 22.2 mm, wh = 19.6 mm. Scale ba
uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
18
a a conch diame e o 37.5 mm, i has e en a alue o 3.70. The who l p o ile is weakly dep essed (ww/
wh = 1.12) wi h a weakly conca e en e ha is bo de ed by a sha p en ola e al shoulde ; he lanks
a e nea ly la and s ongly con e gen . The umbilical ma gin is subangula and he umbilical wall is
s eep and la (Fig. 9A). The who l o e lap is small (IZR = 0.08). The ew p ese ed shell emains o he
holo ype show no o namen a ion o sculp u e. The in e nal mould, howe e , has e y small ube cles on
he angula en ola e al shoulde in he p eadul s age up o a diame e o abou 35 mm. The lanks bea
e y shallow, om he umbilical ma gin backwa dly di ec ed ibs, which s eng hen in he middle o he
lank o o m e y low, b oad conical nodes. The su u e line is dependen on he c oss-sec ional shape
o he who l; i has a a he deep, ounded V-shaped ex e nal lobe, an angula saddle on he en ola e al
shoulde and a b oadly ounded la e al lobe (Fig. 9C). The las qua e olu ion o he ph agmocone has
abou se en chambe s (CLI = 13).
Rema ks
Ocunau ilus coelodesmus gen. e sp. no . occupies an in e media e posi ion be ween O. diplodocus
gen. e sp. no . and O. achy ephus gen. e sp. no . as a as he conca i y o he en e is conce ned;
in O. diplodocus he en e is much mo e deeply conca e and in O. achy ephus i is nea ly la .
Ocunau ilus coelodesmus gen. e sp. no . has e y simila conch p opo ions o O. achy ephus, bu
di e s in he conca e umbilical wall, which is weakly con ex in O. achy ephus.
Ocunau ilus achy ephus gen. e sp. no .
u n:lsid:zoobank.o g:ac :3C21BF76-5C82-4F5E-B39F-43499C76950D
Fig. 10; Table 6
Diagnosis
Species o Ocunau ilus gen. no . wi h hickly discoidal, subin olu e conch (ww/dm ~ 0.50; uw/dm
~ 0.25), weakly dep essed who l p o ile (ww/wh ~ 1.10) and ex ao dina ily high coiling a e (WER
~ 3.35) a a conch diame e o 80 mm. Who l p o ile in e ed apezoidal wi h weakly conca e en e ,
angula en ola e al shoulde , la ened and weakly con e gen lanks, angula umbilical ma gin and
s eep and weakly con ex, la ened umbilical wall. Who l o e lap e y small (IZR ~ 0.04). Wi hou
sculp u e, bu i s who l wi h low, blun nodes on he en ola e al shoulde . Su u e line wi h a e y
shallow ex e nal lobe, a subangula en ola e al saddle, a shallow and b oadly ounded la e al lobe and
a e y small umbilical lobe.
E ymology
F om he G eek ‘ταχύς’ = ‘ apid’ and ‘τρέφω’ = ‘enla ge’; because o he high coiling a e.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 10; MB.C.32115.
Table 5. Conch dimensions (in mm) and a ios o Ocunau ilus coelodesmus gen. e sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32113 57.0 31.5 28.2 12.9 26.0 0.55 1.12 0.23 3.38 0.08
MB.C.32113 37.5 21.3 19.4 9.5 18.0 0.57 1.10 0.25 3.70 0.07
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
19
Desc ip ion
Holo ype MB.C.32115 is a ully chambe ed, pa ially co oded specimen wi h a conch diame e o
85 mm (Fig. 10A). I has sligh ly mo e han wo olu ions; he i s olu ion has a diame e o abou
24 mm and an umbilical o amen o abou 4 mm diame e . The conch is, a a conch diame e o 85 mm,
hickly discoidal and subin olu e (ww/dm = 0.51; uw/dm = 0.24) wi h an ex ao dina ily high coiling
a e (WER = 3.35). Measu emen s om a posi ion hal a olu ion be o e esul in simila a ios o
conch geome y, bu he e occu s an on ogene ic dec ease o he coiling a e; a a conch diame e o
48 mm, i has a alue o abou 3.50. The who l p o ile is weakly dep essed (ww/wh = 1.08) wi h a e y
weakly conca e en e ha is bo de ed by a sha p en ola e al shoulde ; he lanks a e nea ly la and
con e gen . The umbilical ma gin is angula and he umbilical wall is s eep and weakly con ex. The
who l o e lap is e y small (IZR = 0.04). The specimen shows ha he en e is s ill con ex a a conch
diame e o 24 mm, al hough i is al eady sligh ly la ened; he en e is comple ely applana e a a conch
diame e o 48 mm (Fig. 10B). The conch is la gely ee o sculp u e and o namen . On he i s hal
o he las olu ion, he en ola e al shoulde bea s e y low and blun conical nodes. The su u e line
ex ends wi h a b oadly ounded ex e nal lobe, a subangula en ola e al saddle, a shallow la e al lobe
and a e y small umbilical lobe (Fig. 10C).
Rema ks
Ocunau ilus achy ephus gen. e sp. no . di e s om he o he wo species O. diplodocus gen. e sp. no .
and O. coelodesmus gen. e sp. no . om Baghuk Moun ain in he shape o he umbilical wall (con ex
in O. achy ephus, bu conca e in he o he species) and in he only weakly de eloped conca i y o he
en e . Ocunau ilus achy ephus has simila conch p opo ions as O. coelodesmus, bu shows a la ening
Fig. 10. Ocunau ilus achy ephus gen. e sp. no ., holo ype MB.C.32115 (Ko n e al. 2011 Coll.)
om he A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. La e al and do sal
iews. B. Recons uc ion o ape u al iew. C. Su u e line a ww = 23.5 mm, wh = 22.4 mm. Scale ba
uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
20
o he en e only a a la ge conch diame e . This la ening occu s a abou 35 mm in O. achy ephus,
bu al eady a abou 20 mm in O. coelodesmus.
Pseudo i anoce as a meniacus (Abich, 1878) has simila conch p opo ions, bu di e s om
O. achy ephus gen. e sp. no . by a dis inc ly conca e en e as well as he p esence o ube cles on
he en ola e al shoulde and umbilical ma gin. Some o he specimens wi hou ube cles men ioned by
Shimansky (1965) may belong o O. coelodesmus gen. e sp. no .
Genus Ai inau ilus gen. no .
u n:lsid:zoobank.o g:ac :C251419E-4EF4-4D59-BBAA-FEEA876287CC
New genus B – Ko n 2025: 43. — Ko n & Ghade i 2025: 30.
Type species
Ai inau ilus icanus gen. e sp. no .
Diagnosis
Genus o he amily Ocunau ilidae am. no . wi h a subin olu e conch. High coiling a e; who l p o ile
ounded ec angula , weakly dep essed wi h conca e en e . Wi hou sculp u e. Su u e line wi h shallow
ex e nal lobe and b oadly ounded la e al lobe; wi hou annula p ocess.
E ymology
F om he G eek ‘αἴφνης’ = ‘suddenly, unexpec ed’; because o he unexpec ed conch o m o a Pe mian
nau iloid.
Included species
NW I an (Ko n & Ghade i 2025): Ai inau ilus hebes Ko n & Ghade i, 2025, Wuchiapingian.
Cen al I an ( his pape ): Ai inau ilus icanus gen. e sp. no ., Wuchiapingian.
Rema ks
Ai inau ilus gen. no . is di icul o con use wi h any o he genus o Palaeozoic nau iloids because o
i s peculia conch mo phology wi h he box-shaped who l p o ile, he conca e en e , he skid-like
en ola e al shoulde , pa allel lanks and la ened umbilical wall. Ocunau ilus gen. no . di e s om
Ai inau ilus by i s clea ly con e gen lanks, esul ing in a much na owe en e ; i has an angula
en ola e al shoulde and an angula umbilical ma gin. Pseudo i anoce as also has con e gen lanks
and is cha ac e ised by a ow o nodes on he en ola e al shoulde and some imes also a he umbilical
ma gin (Shimansky 1965: 163).
The conch o Ai inau ilus gen. no . is e y simila o ha o some species o he T iassic genus
Ge manonau ilus Mojsiso ics, 1902 (e.g., Mojsiso ics 1902; Dzik 1984; Sobole 1989). The main
di e ence be ween he wo gene a is he p esence o an annula lobe in Ge manonau ilus, which is
absen in Ai inau ilus. The siphuncle has a mo e do sal posi ion in Ge manonau ilus.
Table 6. Conch dimensions (in mm) and a ios o Ocunau ilus achy ephus gen. e sp. no .; econs uc ed
alues in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32115 85.0 43.5 40.1 20.8 38.6 0.51 1.08 0.24 3.35 0.04
MB.C.32115 48.4 25.4 23.4 12.4 22.5 0.52 1.09 0.26 3.49 0.04
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
21

The La e Ca boni e ous genus Ti anoce as has a supe icially simila conch shape wi h a conca e en e
and almos pa allel lanks. Howe e , he conch o Ti anoce as has almos non-o e lapping who ls and
he sculp u e is cha ac e ised by en ola e al ube cles. Shimansky (1965: 163) has al eady sugges ed
ha he La e Ca boni e ous (Ti anoce as) and La e Pe mian o ms (Pseudo i anoce as) belong o wo
independen , homoeomo phic e olu iona y lineages and ha bo h a ose om Doma oce as.
Ai inau ilus icanus gen. e sp. no .
u n:lsid:zoobank.o g:ac :8E2C5B5B-1C6B-4492-8D76-89F957B1DCC1
Figs 11–13; Table 7
Diagnosis
Species o Ai inau ilus gen. no . wi h hickly discoidal, subin olu e conch (ww/dm ~ 0.50; uw/dm
~ 0.25), weakly dep essed who l p o ile (ww/wh ~ 1.10) and e y high coiling a e (WER ~ 2.30) a a
conch diame e o 110 mm. Who l p o ile wi h weakly conca e en e , ounded en ola e al shoulde ,
weakly con e gen lanks, ounded umbilical wall, weakly conca e umbilical wall and mode a ely deep
imp in zone (IZR ~ 0.25). Wi hou sculp u e. Su u e line wi h a e y shallow ex e nal lobe and a e y
shallow la e al lobe.
Fig. 11. Ai inau ilus icanus gen. e sp. no ., holo ype MB.C.32116 (Ko n e al. 2010 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain, en al and la e al iews. Scale ba
uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
22
E ymology
F om he G eek ‘ἱκανός‘ = ‘compe en ’; because o he obus conch.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2010; Ko n e al. leg.; illus a ed in Figs 11–12; MB.C.32116.
Pa a ypes
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; 2011; Ko n e al. leg.; illus a ed
in Fig. 13; MB.C.32117 • 3 specimens; same da a as o holo ype; 2011; Ko n e al. leg.; MB.C.32118
o MB.C.32120.
Fig. 12. Ai inau ilus icanus gen. e sp. no ., holo ype MB.C.32116 (Ko n e al. 2010 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Do sal iew. B. Recons uc ion o
ape u al iew. C. Su u e line a dm = 105 mm, ww = 53 mm, wh = 47 mm. Scale ba uni s = 1 mm.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
23
Desc ip ion
Holo ype MB.C.32116 is an almos comple e, bu pa ially co oded specimen wi h a conch diame e
o 114 mm (Figs 11, 12A). I is la gely chambe ed; only a small pa o he specimen belongs o he
body chambe . The conch is, a a conch diame e o 114 mm, hickly discoidal and subin olu e (ww/
dm = 0.48; uw/dm = 0.24) wi h a e y high coiling a e (WER = 2.31) and mode a ely wide who l o e lap
(IZR = 0.23). The who l p o ile is weakly dep essed (ww/wh = 1.09) wi h a sligh ly conca e en e ha
is bo de ed by a ounded en ola e al shoulde ; he lanks a e b oadly con ex and weakly con e gen .
The umbilical ma gin is na owly ounded and he umbilical wall is s eep and weakly incu ed. The
las who l shows a modi ica ion in he p o ile. A he beginning i is in e ed apezoidal wi h ai ly
con e gen lanks and a subangula en ola e al shoulde (Fig. 12B). The specimen does no show any
aces o sculp u e. The leng h o he ph agmocone chambe s shows conside able changes du ing he las
olu ion. The i s qua e o his who l consis s o six chambe s (CLI = 15); conside able sep al c owding
a he end o he ph agmocone causes he numbe o inc ease o abou 12 (CLI = 7.5). Especially he las
six sep a a e e y closely spaced. The su u e line shows a shallow and b oadly ounded ex e nal lobe, a
na owly ounded en ola e al saddle, a shallow la e al lobe wice as deep han he ex e nal lobe and a
e y small umbilical lobe (Fig. 12C).
Pa a ype MB.C.32117 is an incomple e, ully chambe ed specimen wi h a conch diame e o 63 mm
(Fig. 13B). I co esponds la gely o he holo ype in i s conch p opo ions and also shows a weakly
conca e umbilical wall, almos pa allel lanks, a ounded bu p onounced en ola e al shoulde and
a weakly conca e en e (Fig. 13A). The su u e line has a shallow ex e nal lobe, a na owly ounded
en ola e al saddle and a b oadly ounded la e al lobe ha is wice as deep as he ex e nal lobe
(Fig. 13C).
Rema ks
Specimens o Ai inau ilus icanus gen. e sp. no . can ha dly be con used wi h any o he species o
Baghuk Moun ain. Species o he genus Ocunau ilus gen. no . show simila i ies in conch shape, bu
he e a e some easily ecognisable di e ences. In A. icanus, he lanks a e almos pa allel, while in he
species o Ocunau ilus hey a e dis inc ly con e gen . Ano he dis inguishing c i e ion is he coiling a e,
which is conside ably lowe in A. icanus (WER ~ 2.25) han in Ocunau ilus (WER mos ly o e 3.00).
Fig. 13. Ai inau ilus icanus gen. e sp. no ., pa a ype MB.C.32117 (Ko n e al. 2011 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. La e al and en al iews. C. Su u e line a ww = 26.5 mm, wh = 25.5 mm. Scale ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
24
The e a e g ea simila i ies o A. icanus gen. e sp. no . o species o he T iassic genus Ge manonau ilus.
Fo example, A. icanus shows e y close ag eemen in conch mo phology wi h G. bido sa us
(Schlo heim, 1820), as was shown by Dzik (1984: 166, ex - ig. 64). Howe e , Ge manonau ilus di e s
om Ai inau ilus gen. no . by he p esence o an annula lobe.
Genus Aza inau ilus Ko n & Ghade i, 2025
Type species
Aza inau ilus nahidae Ko n & Ghade i, 2025; o iginal designa ion.
Diagnosis
Genus o he amily Ocunau ilidae am. no . wi h a a he small, subin olu e o sube olu e conch; who l
p o ile ly i o m, en e mode a ely o deeply conca e and bo de ed by aised keels, umbilical ma gin
b oadly ounded o subangula . Sculp u e wi h conical nodes o shallow ibs in he mid lank a ea. Su u e
line wi h na ow, ounded ex e nal lobe and b oadly ounded la e al lobe ( om Ko n & Ghade i 2025).
Included species
NW I an (Ko n & Ghade i 2025): Aza inau ilus nahidae Ko n & Ghade i 2025, Wuchiapingian.
Cen al I an ( his pape ): Aza inau ilus pho minx sp. no ., Changhsingian.
Rema ks
Aza inau ilus has an isola ed posi ion wi hin he amily Doma oce a idae because o i s conch shape
and sculp u e. A compa ably dis inc ly conca e en e has appa en ly no been desc ibed om any o he
genus o he amily; howe e , species o Fididoma oce as show a la o sligh ly conca e en e bo de ed
la e ally by low idges.
Aza inau ilus pho minx sp. no .
u n:lsid:zoobank.o g:ac :91FE758E-E2BC-45B0-BE73-90A885833281
Fig. 14; Table 8
Diagnosis
Species o Aza inau ilus wi h hickly discoidal, sube olu e conch (ww/dm ~ 0.40; uw/dm ~ 0.32),
equidimensional who l p o ile (ww/wh ~ 1.00) and e y high coiling a e (WER ~ 2.30) a a conch
diame e o 45 mm. Who l p o ile ounded ly i o m wi h s ongly con e gen , b oadly con ex lanks;
en e mode a ely conca e, umbilical ma gin subangula . Who l o e lap small (IZR ~ 0.15). Sculp u e
wi h sho ibs in he mid lank a ea. Su u e line wi h a e y shallow ex e nal lobe and a e y shallow
la e al lobe.
E ymology
F om he G eek ‘φόρμιγξ’ = he ancien G eek ly e; because o he who l p o ile.
Table 7. Conch dimensions (in mm) and a ios o Ai inau ilus icanus sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32116 114.0 55.2 50.5 27.0 39.0 0.48 1.09 0.24 2.31 0.23
MB.C.32116 75.0 40.3 38.3 18.6 – 0.54 1.05 0.25 – –
MB.C.32119 79.2 38.0 34.5 22.1 – 0.48 1.10 0.28 – –
MB.C.32117 63.0 31.0 27.4 16.6 20.6 0.49 1.13 0.26 2.21 0.25
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
25
Pa a ype
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; 2011; Ko n e al. leg.; illus a ed
in Fig. 16D–F; MB.C.32123.
Desc ip ion
Holo ype MB.C.30223 is an incomple e in e nal mould wi h a conch diame e o 57 mm; i is ully
chambe ed (Fig. 16B). The conch wid h had o be econs uc ed; i appea s ha he conch is hickly
discoidal and sube olu e (ww/dm ~ 0.48; uw/dm = 0.47) wi h a mode a ely high coiling a e (WER = 1.81).
The who l p o ile is mode a ely dep essed (ww/wh ~ 1.80) and ounded apezoidal wi h a la ly ounded
en e and a b oadly ounded en ola e al shoulde . The di e gen lanks a e sligh ly la ened; an
Fig. 16. Foo dice as eicosacan hum sp. no . om he Vedioce as beds o he Hambas Fo ma ion
a Baghuk Moun ain. A. Holo ype MB.C.30223 (Ko n e al. 2013 Coll.), econs uc ion o ape u al
iew. B. The same specimen, la e al and en al iews. C. The same specimen, su u e line a
wh = 17.2 mm. D. Pa a ype MB.C.32123 (Ko n e al. 2011 Coll.), econs uc ion o ape u al iew.
E. The same specimen, la e al iew. F. The same specimen, su u e line a dm = 50.0 mm, ww = 22.0 mm,
wh = 15.0 mm. Scale ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
32

umbilical ma gin is no p esen (Fig. 16A). The sculp u e consis s o low conical en ola e al nodes, o
which he e a e 10 on hal a olu ion. They a e en i ely con ined o he en ola e al shoulde and he
ou e lank and a e as wide as hei in e spaces. The su u e line shows a e y wide, e y shallow ex e nal
lobe and a e y shallow la e al lobe (Fig. 16C). The sep a a e a he sho in he las qua e olu ion o
he ph agmocone has abou eigh chambe s (CLI = 11); in he penul ima e olu ion, he e a e only six
chambe s on a qua e who l (CLI = 15).
Pa a ype MB.C.32123 is a pa ially co oded specimen wi h a conch diame e o 68 mm (Fig. 16E).
One who l be ween a conch diame e o 20 and 37 mm is a he well-p ese ed and allows he s udy o
he conch geome y and sculp u e. In bo h ea u es as well as in he su u e line (Fig. 16F) he pa a ype
ag ees well wi h he holo ype. The numbe o chambe s emains he same in he las olu ion; in bo h
las hal olu ions he ph agmocone has 15 chambe s each (CLI = 12). The hi d las hal olu ion has
12 chambe s (CLI = 15).
Rema ks
Foo dice as eicosacan hum sp. no . di e s om F. decacan hum sp. no . and F. asce um sp. no . in
he a he small and mo e nume ous en ola e al nodes, o which he e a e en on hal a olu ion,
while he e a e only i e e y coa se nodes on hal a olu ion in he o he wo species. Fu he mo e,
F. eicosacan hum has a mo e s ongly dep essed who l p o ile (ww/wh ~ 1.75) han F. decacan hum (ww/
wh ~ 1.25) and F. asce um (ww/wh ~ 1.00).
Foo dice as decacan hum sp. no .
u n:lsid:zoobank.o g:ac :FEB57A12-32B4-461B-968A-0037BCC865E6
Fig. 17; Table 11
Me acoce as sp. – Ko n e al. 2021: ex - ig. 17b.
Diagnosis
Species o Foo dice as wi h hinly discoidal, sube olu e conch (ww/dm ~ 0.40; uw/dm ~ 0.44), weakly
dep essed who l p o ile (ww/wh ~ 1.25) and high coiling a e (WER ~ 2.10) a a conch diame e o
60 mm. Who l p o ile apezoidal wi h di e gen , con ex lanks; en e la ened, umbilical ma gin
incipien . Sculp u e wi h abou 10 e y coa se, blun en ola e al ibs pe olu ion. Su u e line wi h a
e y shallow ex e nal lobe and a e y shallow, b oadly ounded la e al lobe.
E ymology
F om he G eek ‘δέκα’ = ‘ en’ and ‘ἄκανθα’ = ‘ ho n, spine’; because o he en en ola e al nodes pe
olu ion.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2010; Ko n e al. leg.; illus a ed in Fig. 17; MB.C.30224.
Table 10. Conch dimensions (in mm) and a ios o Foo dice as eicosacan hum sp. no . om Baghuk
Moun ain; econs uc ed alue in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32123 61.3 28.0 16.2 29.0 15.6 0.46 1.73 0.47 1.80 0.04
MB.C.30223 56.5 27.2 15.1 26.8 14.5 0.48 1.80 0.47 1.81 0.04
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
33
Desc ip ion
Holo ype MB.C.30224 is an incomple e, bu a he well-p ese ed in e nal mould wi h a diame e o
62 mm (Fig. 17B). The conch is hinly discoidal and sube olu e (ww/dm = 0.41; uw/dm = 0.44) wi h
a high coiling a e (WER = 2.08); he who l p o ile is weakly dep essed (ww/wh = 1.27) and ounded
apezoidal wi h a la ly ounded en e and a b oadly ounded en ola e al shoulde . The di e gen
lanks a e con ex; a udimen a y umbilical ma gin is de eloped (Fig. 17A). The sculp u e consis s o
abou 10 e y la ge, bu low en ola e al nodes pe olu ion; hese nodes ha e a ounded conical shape
and con inue as e y low ibs on he lank. They a e much wide han he spaces be ween hem. These
ibs a e conside ably mo e p ominen on he penul ima e who l, a a conch diame e o abou 25 mm,
han on he las who l. The su u e line shows a e y shallow ex e nal lobe and a e y shallow la e al lobe
(Fig. 17C). The las hal olu ion o he ph agmocone has abou 12 chambe s (CLI = 15).
Rema ks
Foo dice as decacan hum sp. no . di e s om F. eicosacan hum sp. no ., which possesses a simila
conch geome y, in he numbe and o ma ion o he en ola e al nodes. Foo dice as decacan hum has
only i e e y coa se nodes on hal a who l, whe eas F. eicosacan hum has en much smalle nodes.
Foo dice as decacan hum has a e y simila sculp u e o F. asce um sp. no ., bu shows a dep essed
who l p o ile (ww/wh ~ 1.25), whe eas in F. asce um his is almos equidimensional (ww/wh ~ 1.00).
Ano he c i e ion o sepa a e F. decacan hum om he a he simila F. asce um is in he wid h o he
nodes; in F. decacan hum hey a e wide han he in e spaces, while hey a e only hal as wide as hei
in e spaces in F. asce um.
Fig. 17. Foo dice as decacan hum sp. no ., holo ype MB.C.30224 (Ko n e al. 2010 Coll.) om he
Vedioce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. Do sal, la e al and en al iews. C. Su u e line a wh = 10.8 mm. Scale ba uni s = 1 mm.
Table 11. Conch dimensions (in mm) and a ios o Foo dice as decacan hum sp. no . om Baghuk
Moun ain.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.30224 61.9 25.1 19.7 27.5 19.0 0.41 1.27 0.44 2.08 0.04
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
34
Foo dice as decacan hum sp. no . di e s om he supe icially simila F. golia hus, which also
possesses e y coa se en ola e al nodes, in he much wide umbilicus. The uw/dm a io is abou 0.45
in F. decacan hum, bu only a ound 0.30 in F. golia hus.
Foo dice as asce um sp. no .
u n:lsid:zoobank.o g:ac :B448394B-F0E2-42D2-9EFA-FA09D8C8E1A7
Fig. 18; Table 12
Diagnosis
Species o Foo dice as wi h hinly discoidal, sube olu e conch (ww/dm ~ 0.33; uw/dm ~ 0.43), nea ly
equidimensional who l p o ile (ww/wh ~ 1.00) and high coiling a e (WER ~ 2.20) a a conch diame e
o 60 mm. Who l p o ile apezoidal wi h weakly di e gen , la ened lanks; en e la ened, umbilical
ma gin incipien . Sculp u e wi h abou 10 coa se, blun en ola e al ibs pe olu ion. Su u e line wi h
a e y shallow ex e nal lobe and a e y shallow, b oadly ounded la e al lobe.
E ymology
F om he G eek ‘ἀσκητός’ = ‘deco a ed’; because o he sculp u e.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 18; MB.C.32124.
Fig. 18. Foo dice as asce um sp. no ., holo ype MB.C.32124 (Ko n e al. 2011 Coll.) om he
Vedioce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. La e al iew. C. Su u e line a dm = 61.4 mm, wh = 19.6 mm. Scale ba uni s = 1 mm.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
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Desc ip ion
Holo ype MB.C.32124 is an in e nal mould wi h a diame e o 86 mm, bu he las po ion is poo ly
p ese ed (Fig. 18B). The conch is, a 62 mm diame e , hinly discoidal and sube olu e (ww/dm = 0.33;
uw/dm = 0.43) wi h a high coiling a e (WER = 2.21). The who l p o ile is weakly comp essed (ww/
wh = 0.98) and ounded apezoidal wi h a nea ly la en e and a b oadly ounded en ola e al shoulde .
The weakly di e gen lanks a e la ened; an umbilical ma gin is only ba ely isible (Fig. 18A). The
sculp u e has i e p ominen conical nodes pe hal olu ion in he ou e a ea o he lank; hese nodes
show a low ex ension on he lank owa ds he umbilicus. The nodes a e na owe han hei in e spaces.
The su u e line shows shallow, b oadly ounded lobes on en e and lanks (Fig. 18C). The las hal
olu ion o he ph agmocone has abou 16 chambe s (CLI = 11).
Rema ks
Foo dice as asce um sp. no . di e s om F. eicosacan hum sp. no . and F. decacan hum sp. no . in he
nea ly equidimensional who l p o ile (ww/wh = 1.00), which is dep essed in he o he wo species (ww/
wh = 1.75 in F. eicosacan hum and = 1.25 in F. decacan hum). Ano he c i e ion o sepa a e F. asce um
om he a he simila F. decacan hum is in he wid h o he nodes; in F. asce um hey a e only hal as
wide as hei in e spaces, while hey a e wide han he in e spaces in F. decacan hum.
Genus Ta dunau ilus Ko n & Ghade i, 2025
Diagnosis
Genus o he amily Foo dice a idae wi h e olu e conch; who l p o ile ounded iangula o ounded
apezoidal, dep essed wi h b oadly ounded en e . Sculp u e wi h one o wo ows o conical ibs nea
he en ola e al shoulde . Su u e line wi h e y shallow ex e nal lobe and b oadly ounded la e al lobe;
wi hou annula p ocess ( om Ko n & Ghade i 2025).
Type species
Ta dunau ilus nimius Ko n & Ghade i, 2025; by o iginal designa ion.
Included species
NW I an (Ko n & Ghade i 2025): Ta dunau ilus nimius Ko n & Ghade i, 2025, Wuchiapingian;
Ta dunau ilus mino Ko n & Ghade i, 2025, Wuchiapingian.
Cen al I an ( his pape ): Ta dunau ilus ape imos sp. no ., Wuchiapingian.
Rema ks
The new genus can be easily dis inguished om all o he gene a in he assemblage om Baghuk
Moun ain because o i s combina ion o a conch shape wi h a wide umbilicus, a ounded iangula o
ounded apezoidal who l p o ile and a sculp u e consis ing o conical nodes. A genus wi h a simila
mo phology is Pseudo emnocheilus, bu his is mos ly known om much smalle specimens o abou
40–60 mm in diame e . These smalle specimens ha e a much na owe umbilicus (uw/dm ~ 0.40)
compa ed o Ta dunau ilus (uw/dm ~ 0.47) and a less dep essed who l p o ile (ww/wh ~ 1.30) han
Ta dunau ilus (ww/wh ~ 1.40–1.80). The coiling a e is much lowe in Ta dunau ilus (WER below 2.00)
when compa ed o Pseudo emnocheilus (WER g ea e han 2.25).
Table 12. Conch dimensions (in mm) and a ios o Foo dice as asce um sp. no . om Baghuk Moun ain.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32124 61.6 20.5 21 26.5 20.2 0.33 0.98 0.43 2.21 0.04
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
36
Ta dunau ilus ape imos sp. no .
u n:lsid:zoobank.o g:ac :E08EF718-C2F6-44E2-9D15-A879DA51D394
Fig. 19; Table 13
Diagnosis
Species o Ta dunau ilus wi h hinly discoidal, e olu e conch (ww/dm ~ 0.35; uw/dm ~ 0.50), weakly
dep essed who l p o ile (ww/wh ~ 1.50) and low coiling a e (WER ~ 1.70) a a conch diame e o
60 mm. Who l p o ile ounded apezoidal wi h la ened en e , ounded en ola e al shoulde , s ongly
di e gen lanks and e y small imp in zone. Sculp u e wi h 20 conical nodes on he ou e lank pe
olu ion. Su u e line wi h a e y shallow ex e nal lobe and a e y shallow la e al lobe.
E ymology
F om he G eek ‘ἀπό’ = ‘ om’ and ‘ἐρῆμος’ = ‘dese ’; because o he o igin o he ma e ial.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, P o o oce as beds (ea ly
Wuchiapingian); 2012; Ko n e al. leg.; illus a ed in Fig. 19; MB.C.32125.
Desc ip ion
Holo ype MB.C.32125 is an in e nal mould wi h a conch diame e o 58 mm; i is nea ly ully chambe ed
(Fig. 19B). The conch is hinly discoidal and e olu e (ww/dm = 0.37; uw/dm = 0.53) wi h a low coiling
a e (WER = 1.70); he who l p o ile is weakly dep essed (ww/wh = 1.49) and ounded apezoidal wi h a
la ened en e , a ounded en ola e al shoulde and s ongly con ex and di e gen lanks (Fig. 19A).
The specimen shows a who l ha only sligh ly emb aces he p eceding; he umbilical seam has a posi ion
on he en e o he p eceding who l. The sculp u e consis s o 20 blun conical nodes in en ola e al
posi ion. These nodes a e sligh ly na owe han hei in e spaces. The su u e line shows e y shallow,
b oadly ounded lobes on he en e and he lank (Fig. 19C).
Fig. 19. Ta dunau ilus ape imos sp. no ., holo ype MB.C.32125 (Ko n e al. 2012 Coll.) om he
P o o oce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. Do sal and la e al iews. C. Su u e line a dm = 43.8 mm, ww = 23.8 mm, wh = 18.0 mm. Scale ba
uni s = 1 mm.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
37

Rema ks
Ta dunau ilus ape imos sp. no . di e s om he simila species T. mino o Jul a in he lowe numbe
o en ola e al nodes (20 pe who l, bu abou 25 in T. mino ) and he wide umbilicus (uw/dm ~ 0.53
in T. ape imos, bu only ~ 0.47 in T. mino ) a compa able conch diame e s. Ta dunau ilus nimius om
Jul a di e s in he b oade who ls and he double ow o en ola e al nodes.
Family Pleu onau ilidae Hya , 1900
Diagnosis
Family o he supe amily Pleu onau iloidea wi h a commonly subquad a e o weakly dep essed who l
p o ile; en e anging om con ex o weakly conca e, en ola e al shoulde and umbilical ma gin o en
p onounced, lanks usually weakly con e gen . Sculp u e wi h nume ous ibs on he lank, some imes
wi h conical ube cles and mo e a ely wi h spi al idges. An annula p ocess is p esen in he ad anced
species ( om Ko n 2025).
Included gene a
Lu onau ilus gen. no . (Pe mian); Pleu onau ilus Mojsiso ics, 1882 (T iassic); Phloioce as Hya ,
1884 (T iassic); Anoploce as Hya , 1900 (T iassic); Encoiloce as Hya , 1900 (T iassic); Enoploce as
Hya , 1900 (T iassic); Holconau ilus Mojsiso ics, 1902 (T iassic); T achynau ilus Mojsiso ics, 1902
(T iassic); Sibyllonau ilus Diene , 1915 (T iassic); Phaed ysmocheilus Shimansky & E lange , 1955
(T iassic); A c onau ilus Sobole , 1989 (T iassic); G uman oce as Sobole , 1989 (T iassic).
Rema ks
A de ailed discussion o he Pleu onau ilidae and amilies wi h simila mo phology has been gi en by
Ko n (2025).
Genus Lu onau ilus gen. no .
u n:lsid:zoobank.o g:ac :6CCC6ECD-ED65-4A11-83AC-666B45EFC1C0
New genus C – Ko n 2025: 57. — Ko n & Ghade i 2025: 7, 61, 63, ig. 33.
Type species
Lu onau ilus c a us gen. e sp. no .
Diagnosis
Genus o he amily Pleu onau ilidae wi h sube olu e conch; who l p o ile in e ed apezoidal, weakly
dep essed o equidimensional wi h b oadly ounded en e , pa allel o weakly con e gen lanks, ounded
umbilical ma gin and con ex o la ened umbilical wall. Sculp u e wi h ibs on he lanks, wi hou
nodes o ube cles. Su u e line wi h e y shallow ex e nal lobe and b oadly ounded la e al lobe; wi hou
annula p ocess.
E ymology
Named a e he Lu dese (Cen al I an), a which ma gin Baghuk Moun ain is loca ed.
Table 13. Conch dimensions (in mm) and a ios o Ta dunau ilus ape imos sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32125 57.9 21.5 14.4 30.5 13.5 0.37 1.49 0.53 1.70 0.06
MB.C.32125 44.5 17.1 11.4 21.8 – 0.38 1.50 0.49 – –
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
38
Included species
Cen al I an ( his pape ): Lu onau ilus c a us gen. e sp. no ., Wuchiapingian; Lu onau ilus elachus
gen. e sp. no ., Wuchiapingian; Lu onau ilus cymus gen. e sp. no ., Wuchiapingian.
Rema ks
Lu onau ilus gen. no . is newly in oduced he e o Pe mian species wi h a conch shape and
sculp u e simila o some T iassic species o Pleu onau ilus, such as P. mosis Mojsiso ics, 1882. The
main di e ence, howe e , is ha Lu onau ilus does no possess he annula lobe ha is p esen in
Pleu onau ilus.
Lu onau ilus gen. no . occu s wi h mo phologically simila species in he sec ions o cen al I an and
p obably also in he T anscaucasian egion. Bu i is also possible ha he genus occu s in o he egions
such as he Sal Range o Pakis an; howe e , his can only be cla i ied by e ising he co esponding
assemblages.
Lu onau ilus c a us gen. e sp. no .
u n:lsid:zoobank.o g:ac :FA260A86-6965-469D-96FE-D7D6B45B5F8A
Fig. 20; Table 14
New genus, new species – Ko n 2025: 58, ig. 29.
Diagnosis
Species o Lu onau ilus gen. no . wi h hinly discoidal, subin olu e conch (ww/dm = 0.35–0.40; uw/
dm = 0.35–0.40), nea ly equidimensional who l p o ile (ww/wh = 0.95–1.05) and high o e y high coiling
a e (WER = 2.20–2.35) a a conch diame e o 60–80 mm. Who l p o ile wi h gen ly con e gen lanks;
en e b oadly ounded, lanks sligh ly la ened, umbilical ma gin ounded, umbilical wall la ened.
Sculp u e wi h abou 25 blun la e al ibs pe olu ion; hey begin on he ou e umbilical wall and a e
s eng hened on he en ola e al shoulde o o m elonga e nodes. Su u e line wi h a shallow ex e nal
lobe and a shallow, b oadly ounded la e al lobe.
E ymology
F om he G eek ‘κρατύς’ = ‘s ong’; because o he a he obus conch and sculp u e.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 20A–D; MB.C.32126.
Pa a ypes
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; 2010; Ko n e al. leg.; illus a ed in
Fig. 20E–F; MB.C.32127 • 2 specimens; same da a as o holo ype; 2010; Ko n e al. leg.; MB.C.32128,
MB.C.32129.
Desc ip ion
Holo ype MB.C.32126 is a a he comple e, bu somewha co oded specimen wi h a conch diame e
o 88 mm (Fig. 20A, D). I is almos comple ely chambe ed and only a li le mo e han a qua e o he
las who l belongs o he body chambe . The conch is hinly discoidal and sube olu e (ww/dm = 0.33;
uw/dm = 0.38) wi h a high coiling a e (WER = 2.17). The who l p o ile is almos squa e-shaped (ww/
wh = 0.95) and shows a la ly ounded en e and a b oadly ounded en ola e al shoulde . The lanks
a e sligh ly la ened and slowly con e gen ; he umbilical ma gin is con inuously ounded and he
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
39
Fig. 20. Lu onau ilus c a us gen. e sp. no . om he Vedioce as beds o he Hambas Fo ma ion a
Baghuk Moun ain. A. Holo ype MB.C.32126 (Ko n e al. 2011 Coll.), le side la e al and do sal
iews. B. The same specimen, econs uc ion o ape u al iew. C. The same specimen, su u e line a
dm = 76.2 mm, ww = 27.0 mm, wh = 29.2 mm. D. The same specimen, igh side la e al iew. E. Pa a ype
MB.C.32127 (Ko n e al. 2010 Coll.), econs uc ion o ape u al iew. F. The same specimen, la e al
iew. Scale ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
40
umbilical wall is s eep (Fig. 20B). The sculp u e consis s o abou 25 blun ibs on he lank; he ibs and
hei in e spaces ha e almos he same wid h. These ibs al eady begin in he ou e a ea o he umbilical
wall and om he e, hey ollow a weakly backwa dly di ec ed cou se; hey ex end ac oss he lank
wi h a b oad and shallow sinus and a e s eng hened o o m adially elonga ed nodes in he a ea o he
en ola e al shoulde . The e hey suddenly weaken and diminish; no ibs a e isible on he en e . The
su u e line shows a e y shallow ex e nal lobe and a b oadly ounded la e al lobe (Fig. 20C). The las
hal who l o he ph agmocone has abou 12 chambe s (CLI = 15).
Pa a ype MB.C.32127 is a agmen o a specimen wi h a conch diame e o 76 mm (Fig. 20F). The
conch geome y is la gely he same as in he holo ype, bu he pa a ype has a sligh ly mo e comp essed
who l p o ile (ww/wh = 0.93) and a highe coiling a e (WER = 2.35). The sculp u e is be e p ese ed
han in he holo ype; i shows 16 a he sha p ibs on he las hal olu ion; hese ibs s a on he
umbilical wall and hin ou a he en ola e al shoulde . The ibs a e somewha na owe han hei
in e spaces; hey ex end ac oss he lank wi h a shallow sinus.
Rema ks
Lu onau ilus c a us gen. e sp. no . di e s om he co-occu ing L. elachus gen. e sp. no . in he
slende e who l p o ile (ww/wh = 0.95–1.05 in L. c a us bu ~ 1.15 in L. elachus), he s onge con e gen
lanks (nea ly pa allel in L. elachus) and he wide umbilicus (uw/dm = 0.35–0.40 in L. c a us, bu only
~ 0.30 in L. elachus). Fu he mo e, he ibs begin in he ou e a ea o he umbilical wall in L. c a us, bu
a he umbilical ma gin in L. elachus.
Lu onau ilus c a us gen. e sp. no . di e s om L. cymus gen. e sp. no . in he shape o he ibs, which
in L. c a us a e almos s aigh on he lanks, while hey ex end wi h a conca e a ch in L. cymus. The ibs
o L. c a us a e coa se in he mid lank a ea, while hey a e weakly de eloped in he middle o he lank
in L. cymus and s eng hened and o ming nodes on he ou e and inne lanks.
Lu onau ilus elachus gen. e sp. no .
u n:lsid:zoobank.o g:ac :7AC16980-96B5-465E-A223-3295CE49F01E
Fig. 21; Table 15
Pleu onau ilus sp. – Ko n e al. 2021: ex - ig. 17c.
Diagnosis
Species o Lu onau ilus gen. no . wi h hickly discoidal, sube olu e conch (ww/dm ~ 0.45; uw/dm
~ 0.32), weakly dep essed who l p o ile (ww/wh ~ 1.15) and e y high coiling a e (WER ~ 2.50) a
a conch diame e o 50 mm. Who l p o ile wi h nea ly pa allel lanks; en e la ened, lanks sligh ly
la ened, umbilical ma gin na owly ounded, umbilical wall la . Sculp u e wi h abou 20 blun la e al
ibs pe olu ion. Su u e line wi h a shallow ex e nal lobe and a shallow, b oadly ounded la e al lobe.
Table 14. Conch dimensions (in mm) and a ios o Lu onau ilus c a us gen. e sp. no . om Baghuk
Moun ain; es ima ed alues in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32126 87.6 29.0 30.5 33.2 28.1 0.33 0.95 0.38 2.17 0.08
MB.C.32128 80.5 30.4 30.0 30.0 28.0 0.38 1.01 0.37 2.35 0.07
MB.C.32127 76.1 26.7 28.6 28.1 26.5 0.35 0.93 0.37 2.35 0.07
MB.C.32129 61.6 24.3 22.4 24.4 21.2 0.39 1.08 0.40 2.32 0.05
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
41
- Fo ma ion o la e al ibs: la e al ibs do no occu in many o he No h Ame ican species, bu a e
p esen in species om o he egions (T. debile, T. admonens).
- Fo ma ion o he umbilical nodes: umbilical sculp u al elemen s a e p esen in some species; hey ange
om being small ube cles (T. clydense) o la ge conical nodes (T. schellbachi).
The lis o mos ly bipola cha ac e pai s shows nume ous heo e ical possible combina ions; howe e ,
co a ia ion is e y common and some o he cha ac e s o en appea simul aneously.
Tainoce as hys a um sp. no .
u n:lsid:zoobank.o g:ac :BE14C8AB-8E7D-4D26-B1BA-64AFA2B52E70
Fig. 23; Table 17
Diagnosis
Species o Tainoce as wi h hinly discoidal, e olu e conch (ww/dm ~ 0.44; uw/dm ~ 0.32), weakly
dep essed who l p o ile (ww/wh ~ 1.05) and ex emely high coiling a e (WER ~ 2.55) a a conch
diame e o 40 mm. Who l p o ile oc agonal wi h shallow longi udinal en al g oo e. Sculp u e wi h
wo ows o ube cles on he en ola e al shoulde and one ow on he umbilical ma gin. Su u e line
undula ing wi h e y shallow lobes and low saddles.
E ymology
F om he G eek ‘ὕστατος’ = ‘la es ’; because he species is he s a ig aphically la es o he nau iloids
om Baghuk Moun ain.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain sec ion H; Hambas Fo ma ion, Pa a i oli es beds (la e
Changhsingian), 0.40 m below op; 2013; Ko n e al. leg.; illus a ed in Fig. 23; MB.C.32134.
Pa a ype
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; 2011; Ko n e al. leg.; MB.C.32135.
Desc ip ion
Holo ype MB.C.32134 is a somewha co oded specimen wi h a conch diame e o 40 mm (Fig. 23A). I
is ai ly well p ese ed only on i s le side as an in e nal mould. The specimen is comple ely chambe ed.
Fig. 23. Tainoce as hys a um sp. no ., holo ype MB.C.32134 (Ko n e al. 2013 Coll.) om he
Pa a i oli es beds o he Hambas Fo ma ion a Baghuk Moun ain. A. La e al and do sal iews.
B. Recons uc ion o ape u al iew. C. Su u e line a dm = 32.1 mm, ww = 18.2 mm, wh = 12.3 mm.
Scale ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
48

The conch is hinly discoidal and sube olu e (ww/dm = 0.43; uw/dm = 0.33) wi h an ex emely high coiling
a e (WER = 2.55). The who l p o ile is gene ally oc agonal and sligh ly dep essed (ww/wh = 1.09). I has
a la ened en e wi h a shallow longi udinal dep ession; he en ola e al shoulde is subangula and he
lanks a e almos la and pa allel. The umbilical ma gin is p ominen and he umbilical wall is oblique and
la . The a eas be ween he angula ma gins o he who l p o ile a e la ened (Fig. 23B).
The sculp u e consis s o h ee ows o ube cles, one on he umbilical ma gin, one on he en ola e al
shoulde and one delimi ing he mid en al longi udinal g oo e. The i s and he las ones a e only e y
weakly de eloped; he middle one is much s onge and consis s o blun conical nodes. They a e connec ed
o he ube cles on he umbilical wall by e y low ibs. The su u e line is s ongly dependen on he who l
p o ile; i has shallow lobes a he la ened a eas and low saddles a he p onounced ma gins (Fig. 23C).
Rema ks
Tainoce as hys a um sp. no . shows a mo phology o conch and sculp u e ha can be desc ibed as
conse a i e. E en some o he La e Ca boni e ous species, such as he ype species T. quad angulum,
show an oc agonal shape o he who l p o ile wi h wo ows o ube cles each adjacen o he median
longi udinal g oo e. Howe e , T. quad angulum has coa se en ola e al nodes and a na owe umbilicus
(uw/dm < 0.30) han T. hys a um (uw/dm > 0.30).
Tainoce as hys a um sp. no . di e s by i s wide umbilicus and pa icula ly by he pa allel lanks, which
a e ei he con e gen o di e gen in many o he species o he genus Tainoce as. Ano he dis inguishing
ea u e is he shape o he ube cles, which a e much smalle in T. hys a um han in mos o he species.
Genus Epi ainoce as gen. no .
u n:lsid:zoobank.o g:ac :4F3AA6AC-9211-4B03-A581-9676BE734DAA
New genus D – Ko n 2025: 61. — Ko n & Ghade i 2025: 64, 72, 74.
Type species
Epi ainoce as lu ense gen. e sp. no .
Diagnosis
Genus o he amily Tainoce a idae wi h subin olu e o sube olu e conch; who l p o ile dep essed and
polygonal wi h a shallow o b oad mid en al longi udinal g oo e. Umbilical ma gin p onounced and
na owly ounded, lanks con e gen . Sculp u e wi h one ow o nodes on he en ola e al shoulde .
Su u e line s ongly depending on he shape o he who l p o ile, wi h ounded V-shaped ex e nal lobe
and b oadly ounded la e al lobe.
E ymology
F om he G eek ‘ἐπί’ = ‘on, abo e’; e e ing o he la e occu ence o his ainoce a id genus.
Included species
T anscaucasia (Abich 1878): Nau ilus do so plica us Abich, 1878, Wuchiapingian, Aze baijan.
Cen al I an ( his pape ): Epi ainoce as lu ense gen. e sp. no ., Wuchiapingian.
Table 17. Conch dimensions (in mm) and a ios o Tainoce as hys a um sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32134 41.2 17.7 16.2 13.6 15.4 0.43 1.09 0.33 2.55 0.05
MB.C.32135 29.5 11.5 11.0 9.2 – 0.39 1.05 0.31 – –
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
49
Rema ks
Epi ainoce as gen. no . is a genus wi h a mo phology simila o some species o Tainoce as, bu wi h a
much weake sculp u e. In con as o Tainoce as, which has a leas wo ows o nodes o ube cles in
he en ola e al egion and some imes ano he ow o nodes on he inne lank, Epi ainoce as has only
one ow o nodes, loca ed immedia ely adjacen o he median longi udinal g oo e on each side. Wi h
a simpli ica ion o he sculp u e, he new genus may be de i ed om Tainoce as. Co o ainoce as also
shows simpli ica ion o he sculp u e, bu di e s om Epi ainoce as in he absence o en ola e al nodes
and he p esence o e y coa se do sola e al nodes.
Epi ainoce as lu ense gen. e sp. no .
u n:lsid:zoobank.o g:ac :5A1405B2-C14D-4533-8089-A1C3435EBC18
Fig. 24; Table 18
Diagnosis
Species o Epi ainoce as gen. no . wi h hickly discoidal, subin olu e conch (ww/dm ~ 0.55; uw/dm
~ 0.25) and weakly dep essed who l p o ile (ww/wh ~ 1.35) and e y high coiling a e (WER ~ 2.35)
a a conch diame e o 70 mm. Who l p o ile polygonal wi h con e gen lanks; en e ec i o m wi h
b oad longi udinal mid en al g oo e, lanks la ened, umbilical ma gin na owly ounded. Sculp u e
wi h abou 15 coa se en ola e al, ans e sely elonga ed nodes pe olu ion. Su u e line wi h a small,
ounded V-shaped ex e nal lobe, a b oadly ounded, e y shallow la e al lobe and a e y shallow
umbilical lobe.
E ymology
Named a e he Lu dese (Cen al I an), a which ma gin Baghuk Moun ain is loca ed.
Fig. 24. Epi ainoce as lu ense gen. e sp. no ., holo ype MB.C.32136 (Ko n e al. 2011 Coll.) om he
A axoce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew.
B. La e al and en al iews. C. Su u e line a dm = 63.6 mm, wh = 28.2 mm. Scale ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
50
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, A axoce as beds (ea ly
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 24; MB.C.32136.
Pa a ype
IRAN – Es ahan P o ince • 1 specimen; same da a as o holo ype; MB.C.32137.
Desc ip ion
Holo ype MB.C.32136 is a co oded ph agmocone specimen wi h a diame e o 75 mm (Fig. 24B). I
allows he conch o m, sculp u e and su u e line o be s udied, bu some conch dimensions can only be
es ima ed. The conch is hickly discoidal and subin olu e (ww/dm ~ 0.55; uw/dm ~ 0.25) wi h a e y
high coiling a e (WER ~ 2.35). The who l p o ile is sligh ly dep essed (ww/wh ~ 1.35) and wides a
he na owly ounded umbilical ma gin. The en e possesses a mode a ely wide median g oo e; he
en ola e al shoulde is la ened and oblique, he lanks a e weakly la ened and con e gen and he
umbilical ma gin is na owly ounded (Fig. 24A).
Because o pa ial co osion o he specimen, he sculp u e canno be examined in de ail. I consis s o
wo ows o nodes a anged on each side o he en ola e al shoulde . The en al ow has nodes ha a e
elonga ed in a la e al di ec ion o o m sho ibs; he do sal ow consis s o much weake , ci cula nodes.
The su u e line o he holo ype has a shallow, b oadly V-shaped ex e nal lobe, a low and b oadly ounded,
sligh ly asymme ic en ola e al saddle, a shallow and e y wide la e al lobe and a e y shallow and
small lobe on he umbilical wall (Fig. 24C). The las hal olu ion has 15 ph agmocone chambe s
(CLI = 12).
Rema ks
Epi ainoce as lu ense gen. e sp. no . is simila o he T anscaucasian species E. do soplica um
comb. no ., bu di e s in ha ing a much wide mid en al g oo e and signi ican ly la ge en al nodes.
These nodes a e la e ally elonga ed in E. lu ense, whe eas hey a e simply conical in E. do soplica um.
Subo de Li oce a ina Flowe , 1955
Diagnosis
Subo de o he o de Nau ilida, in which an umbilical ma gin is o med ea ly in on ogeny; ad anced
species may eg ess his cha ac e . Conch usually pachyconic and a ely discoidal o globula ,
subin olu e o in olu e. Ju enile who l p o ile ci cula . Adul who l p o ile usually ci cula o dep essed
o al wi hou dis inc en ola e al shoulde in he phylogene ically ea ly species, showing modi ica ions
du ing e olu ion (in e ed apezoidal wi h con e gen lanks and la ened en e ). Do sal who l zone
always p esen , small o mode a ely deep. Ju enile sculp u e wi h spi al lines ha may be es ic ed o
he umbilical a ea in he phylogene ically ea ly species; adul sculp u e usually lacking excep o spi al
lines in some species. Sep a simply domed in he ea ly species; wi h do sal in lexion in ad anced species
Table 18. Conch dimensions (in mm) and a ios o Epi ainoce as lu ense gen. e sp. no . om Baghuk
Moun ain; econs uc ed dimensions and a ios in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32136 74.6 41.6 30.2 17.8 26.0 0.56 1.38 0.24 2.36 0.14
MB.C.32137 73.0 38.5 29.0 20.5 25.5 0.53 1.33 0.28 2.36 0.12
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
51
and wi h co uga ed sep a in wo de i ed clades. Su u e line depending on he who l p o ile, usually wi h
shallow lobes and low saddles; wi h dis inc lobes in wo clades ( om Ko n 2025).
Included supe amilies
Li oce a oidea Hya , 1900 (Ca boni e ous o T iassic); Ephippioce a oidea Mille & Youngquis , 1949
(Ca boni e ous o Pe mian); Clydonau iloidea Hya , 1900 (T iassic o Ju assic).
Rema ks
A discussion o he subo de Li oce a ina has been gi en by Ko n (2025).
Supe amily Li oce a oidea Hya , 1900
Diagnosis
Supe amily o he subo de Li oce a ina wi h a pachyconic and a ely discoidal o globula , subin olu e
o in olu e conch. Who l p o ile usually ci cula o dep essed o al wi hou dis inc en ola e al shoulde ;
in some species wi h a p onounced bu ounded en ola e al shoulde . Do sal who l zone usually small
o mode a ely deep. Ju enile sculp u e in he ea ly species wi h spi al lines ha may be es ic ed o he
umbilical a ea; de i ed species a e o en smoo h. Su u e line e y simple, almos s aigh ac oss lanks
and en e ( om Ko n 2025).
Included amilies
Li oce a idae Mille & Youngquis , 1949 (Ca boni e ous o T iassic); Coloce a idae Hya , 1893
[homonym, synonym o Li oce a idae Mille & Youngquis , 1949]; Pa anau ilidae Kummel in Flowe &
Kummel, 1950 (T iassic); Pe monau ilidae Ba sko & Shilo sky, 2014 (Pe mian); Plane oce a idae
Ko n, 2025 (Ca boni e ous); Jul anau ilidae Ko n & Ghade i, 2025 (Pe mian).
Rema ks
A discussion o he supe amily Li oce a oidea has been gi en by Ko n (2025).
Family Li oce a idae Mille & Youngquis , 1949
Diagnosis
Family o he supe amily Li oce a oidea wi h a usually pachyconic o globula , subin olu e o
sube olu e conch. Who l p o ile in he adul s age usually mo e o less s ongly dep essed; lanks and
en e o m a con inuous a ch in he ea ly species, he en e can be la ened o conca e in ad anced
species. Umbilical ma gin ounded; umbilical wall usually con ex. O namen usually consis ing o ine
g ow h lines; spi al lines occu in some gene a. Sep um simple in shape, conca ely domed; su u e line
e y simple, almos s aigh ac oss lanks and en e o wi h small lobes and saddles ( om Ko n 2025).
Included gene a
Solenoce as Hya , 1884 (homonym o Solenoce as Con ad, 1860, objec i e synonym o
Coelogas e oce as); Coelogas e oce as Hya , 1893 (Ca boni e ous o Pe mian); Coloce as Hya , 1893
[homonym o Coloce as Taschenbe g, 1882, synonym o Li oce as Teiche , 1940]; S ea oce as Hya ,
1893 (Ca boni e ous o Pe mian); Pe ipe oce as Hya , 1894 (Ca boni e ous o Pe mian); Po oce as
Hya , 1894 (Ca boni e ous); Nannoce as Hya , 1894 (nomen nullum, synonym o Pe ipe oce as);
Con adice as Cossmann, 1900 (objec i e synonym o Coelogas e oce as); Li oce as Teiche ,
1940 (Ca boni e ous o Pe mian); Cond aoce as Mille , Lane & Unklesbay, 1947 (Ca boni e ous
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
52
o Pe mian); Pe ip oce as Chao, 1954 (nomen nullum, synonym o Pe ipe oce as); Hemili oce as
Ruzhence & Shimansky, 1954 (Ca boni e ous o Pe mian); Bis iali es Tu ne , 1954 (Ca boni e ous);
Pseudophacoce as Tu ne , 1966 (Ca boni e ous); Neobis iali es Tucke , Mapes & A ono , 1978
(Ca boni e ous); Jianoce as Ma, 1997 (Pe mian); Nemdoce as Ba sko & Shilo sky, 2014 (Pe mian);
Pa ali oce as Ba sko & Shilo sky, 2014 (Pe mian); Ta ianau ilus Ba sko & Shilo sky, 2014
(Pe mian); Lenice as Leono a & Shchedukhin, 2020 (Pe mian); Shikhanonau ilus Leono a &
Shchedukhin, 2020 (Pe mian); Thyoce as Leono a & Shchedukhin, 2020 (Pe mian); Cele oli oce as
Ko n & Ghade i, 2025 (Pe mian); Pe unau ilus C ick & Sobole , 1994 (T iassic); Tomponau ilus
Sobole , 1989 (T iassic).
Rema ks
A discussion o he amily Li oce a idae has been gi en by Ko n (2025).
Genus Li oce as Teiche , 1940
Type species
Coloce as li a um Gi y, 1911; o iginal designa ion.
Diagnosis
Genus o he amily Li oce a idae wi h pachyconic o globula , in olu e o subin olu e conch; umbilicus
closed by a plug in some species. The i s who l is 10–20 mm in diame e wi h a e y small umbilical
o amen; he conch is apidly inc easing in heigh wi h a high coiling a e (WER usually highe han
2.50). Who ls weakly emb acing, hei p o ile anges om eni o m o nea ly ci cula . Ju enile conch
wi h longi udinal idges o lines; adul o namen wi h g ow h lines wi h a ai ly deep en al sinus and
spi al lines in some species. Sep a wi hou in lexions, sligh ly conca e. Su u e line simple, nea ly s aigh
wi h a shallow, b oadly ounded in e nal lobe. The siphuncle has a posi ion be ween he cen e o he
sep um and he en e (a e Go don 1965; Shimansky 1967).
Included Ca boni e ous species
No h Ame ica (Shuma d & Swallow 1858; Mille & Gu ley 1897; Gi y 1911; Mille e al. 1933; Newell
1936; Unklesbay 1962; Go don 1965): Nau ilus Missou iensis Swallow, 1858, Bashki ian, Missou i;
Solenochilus hen y illense Mille & Gu ley, 1897, Viséan, Indiana; Coloce as li a um Gi y, 1911,
Mosco ian, Oklahoma; Coloce as li a um a . obsole um Gi y, 1911, Mosco ian, Oklahoma; Coloce as
g eenei Mille , Dunba & Cond a, 1933, Kasimo ian, Oklahoma; Coloce as mille i Newell, 1936,
Kasimo ian, Kansas; Coloce as e icula um Mille & Owen, 1937, Kasimo ian, Oklahoma; Li oce as
pa ulum Unklesbay, 1962, Bashki ian, A kansas; Li oce as bicos a um Go don, 1965, Se pukho ian,
A kansas.
B i ish Isles (Foo d 1891; Hind 1910; Tu ne 1954; Ramsbo om & Moo e 1961): Coelonau ilus
De biensis Foo d, 1891, Viséan, De byshi e; Coelonau ilus De biensis a . globula e Foo d, 1891,
Viséan, Isle o Man; Solenocheilus globosus Hind, 1910, Bashki ian, Lancashi e; Li oce as lunense
Tu ne , 1954, Se pukho ian, Yo kshi e; Li oce as lei imense Ramsbo om & Moo e, 1961, Viséan,
I eland.
Cen al Eu ope (T enkne 1868; Hya 1894; Mille e al. 1933; Schmid 1951): Nau ilus g undensis
T enkne , 1868, Viséan, Ha z Moun ains; Coloce as globa um Hya , 1894, Viséan, Belgium; Coloce as
hya i Mille , Dunba & Cond a, 1933, Viséan, Belgium; Li oce as occluso Schmid , 1951, Viséan, Ha z
Moun ains; Li oce as schaelkense Schmid , 1951, Viséan, Rhenish Moun ains.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
53

No h A ica (Ko n & Klug 2023): Li oce as ka aouii Ko n & Klug, 2023, Viséan, An i-A las; Li oce as
e mis Ko n & Klug, 2023, Se pukho ian, An i-A las.
Wes e n Russia (Eichwald 1857; Shimansky 1967): Nau ilus excen icus Eichwald, 1857, Se pukho ian,
Wes e n Russia; Li oce as o nica um Shimansky, 1967, Se pukho ian, Wes e n Russia; Li oce as
de ja o ense Shimansky, 1967, Mosco ian, Moscow Basin.
U als (Shimansky 1967): Li oce as p aelunense Shimansky, 1967, Viséan, No h U als; Li oce as
uzhence i Shimansky, 1967, Se pukho ian, Sou h U als.
No h China (Ruan & Zhou 1987): Li oce as eni o me Ruan & Zhou, 1987, Bashki ian, Ningxia.
Included Pe mian species
No h Ame ica (Hya 1893): Coloce as globula e Hya , 1893, A inskian, Texas.
Cen al and Sou he n Eu ope (Gemmella o 1889; P ino h & Posena o 2007): Endolobus salomonensis
Gemmella o, 1889, Wo dian, Sicily; Li oce as ga denense P ino h & Posena o, 2007, Changhsingian,
Dolomi es.
Wes e n Russia, U als (Yako le 1899; K uglo 1928; Ba sko e al. 2014; Leono a & Shchedukhin
2020): Asymp oce as ko ulkense Yako le , 1899, Sakma ian, Sou h U als; Coloce as (?) sa aensis
K uglo , 1928, Sakma ian (?), Sou h U als; Coloce as abichi a . as ubense K uglo , 1928, Sakma ian
(?), Sou h U als; Li oce as olgense Ba sko & Shilo sky, 2014, Roadian, Wes e n Russia; Li oce as
shakh auense Leono a & Shchedukhin, 2020, Asselian o Sakma ian, Sou h U als.
NW I an (Ko n & Ghade i 2025): Li oce as e iculum Ko n & Ghade i, 2025, Wuchiapingian.
Cen al I an ( his pape ): Li oce as lep um sp. no ., Wuchiapingian.
Pakis an (Reed 1944): Li oce as bakhense Reed, 1944, Wuchiapingian, Sal Range.
Sou h China (Chao 1940, 1954; Xu 1977; Zhao e al. 1978; Liang 1984; Wu & Kuang 1992): Coloce as
sinense Chao, 1940, Kungu ian, Hunan; Pe ipe oce as hsueyuechiani Chao, 1954, Kungu ian, Hunan;
Li oce as o ien ale Chao, 1954, Kungu ian, Hunan; Ephippioce as hunanense Chao, 1954, Kungu ian,
Hunan; Li oce as didmyoau ise Xu, 1977, Kungu ian, Hunan; Li oce as meishanense Zhao, Liang &
Zheng, 1978, Changhsingian, Zhejiang; Li oce as chenxianense Liang, 1984, Changhsingian, Hunan;
Li oce as lichuanense Wu & Kuang, 1992, Changhsingian, Hubei.
Indo-Paci ic (Haniel 1915): Nau ilus Moleng aa i Haniel, 1915, Wuchiapingian, Timo .
Madagasca (Vaillan -Cou u ie T ea 1933): Nau ilus wa e lo i Vaillan -Cou u ie T ea , 1933,
Wuchiapingian.
Rema ks
The genus Li oce as was in oduced by Teiche (1940) o hose Ca boni e ous and Pe mian nau iloids
ha we e p e iously mos ly included in he genus Coloce as Hya , 1893. The name Coloce as had
al eady been used by Taschenbe g (1882) as a subgenus o ecen Mallophaga.
Li oce as has a s a ig aphic ange om he Viséan o he Changhsingian and is ep esen ed by species
a almos all s ages o his long in e al. A he same ime, Li oce as is geog aphically widesp ead, bo h
in he Ca boni e ous and in he Pe mian. Li oce as is also known om di e en acies a eas; Ea ly
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
54
Ca boni e ous species a e known om shallowe and deepe a eas o he shel . In he La e Pe mian,
he genus was mo e common in he shallowe a eas o he sea, as sugges ed by he occu ences a Jul a,
whe e i co-occu s wi h he mo phologically simila and closely ela ed genus Pe monau ilus.
Li oce as lep um sp. no .
u n:lsid:zoobank.o g:ac :43D3AA7B-CCE4-43FD-B6A6-D9B7A27DB882
Fig. 25; Table 19
Diagnosis
Species o Li oce as wi h hinly pachyconic, subin olu e conch (ww/dm ~ 0.68; uw/dm ~ 0.17),
mode a ely dep essed who l p o ile (ww/wh ~ 1.45) and e y high coiling a e (WER ~ 2.30) a a conch
diame e o 40 mm. Who l p o ile wi h b oadly a ched en e and lanks, b oadly ounded umbilical
ma gin, con ex umbilical wall and mode a ely deep imp in zone (IZR ~ 0.28). Wi hou sculp u e. Su u e
line wi h e y shallow and wide ex e nal, la e al and umbilical lobes.
E ymology
F om G eek ‘λεπτός’ = ‘small, weak’; because o he a he small-sizes ma e ial.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 25; MB.C.32138.
Pa a ypes
IRAN – Es ahan P o ince • 2 specimens; same da a as o holo ype; 2010; Ko n e al. leg.; MB.C.32139,
MB.C.32140.
Desc ip ion
The bes -p ese ed specimen is he obliquely de o med small holo ype MB.C.32138 wi h a conch
diame e o 41 mm (Fig. 25B). The conch geome y can only be econs uc ed. Acco ding o his, he
conch is hinly pachyconic and subin olu e (ww/dm = 0.68; uw/dm = 0.17) wi h a e y high coiling a e
(WER = 2.31) and a mode a ely wide who l o e lap (IZR = 0.28). The who l p o ile is weakly dep essed
(ww/wh = 1.44) wi h a b oadly con ex en e ha me ges con inuously wi h he lanks he lanks. The
umbilical ma gin is b oadly ounded and he umbilical wall is con ex (Fig. 25A). The su u e line is
Fig. 25. Li oce as lep um sp. no ., holo ype MB.C.32138 (Ko n e al. 2011 Coll.) om he Vedioce as
beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al iew. B. Do sal
and la e al iews. C. Su u e line a dm = 25.8 mm, ww = 21.2 mm, wh = 12.8 mm. Scale ba uni s = 1 mm.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
55
nea ly s aigh and shows e y b oad and shallow lobes on he a ea consis ing o en e and lanks and
he umbilical wall (Fig. 25C).
Rema ks
The ma e ial o he new species does no show whe he he umbilical ex ensions cha ac e is ic o he
genus was p esen in he adul s age; he e o e, assignmen o Li oce as is no en i ely ce ain. L. lep um
sp. no . di e s om he o he La e Pe mian species o he genus in he opened umbilicus and he
slende e conch; ww/dm a io o a conch diame e o 40–50 mm is below a alue o 0.70, bu in he
o he species i is abo e 0.80.
Genus Pa ali oce as Ba sko & Shilo sky in Ba sko e al., 2014
Type species
Pa ali oce as kazanicum Ba sko & Shilo sky in Ba sko e al., 2014; o iginal designa ion.
Diagnosis
Genus o he amily Li oce a idae wi h pachyconic, subin olu e conch. The conch is apidly inc easing
in heigh wi h a high coiling a e (WER = 2.30–2.90). Who ls mode a ely emb acing, who l p o ile
eni o m; umbilical ma gin p onounced, na owly ounded. Adul o namen wi h ine g ow h lines wi h
a ai ly deep en al sinus. Sep a wi hou in lexions, sligh ly conca e. Su u e line simple, nea ly s aigh
o s aigh wi h a shallow, b oadly ounded in e nal lobe. The siphuncle has a posi ion be ween he cen e
o he sep um and he en e .
Included species
Wes e n Russia (Ba sko e al. 2014): Pa ali oce as kazanicum Ba sko & Shilo sky in Ba sko e al.,
2014, Roadian.
Cen al I an ( his pape ): Pa ali oce as mac ogas e sp. no ., Wuchiapingian.
Rema ks
Pa ali oce as is simila o Li oce as and di e s mainly in he p onounced umbilical ma gin, which is
na owly ounded in Pa ali oce as bu usually b oad and e enly ounded in Li oce as. The species o
Pa ali oce as a e mo e widely umbilica e han mos known species o Li oce as.
Pa ali oce as mac ogas e sp. no .
u n:lsid:zoobank.o g:ac :0AF97520-0593-4555-9D0F-96A5FBF603A6
Fig. 26; Table 20
Diagnosis
Species o Pa ali oce as wi h hinly globula , subin olu e conch (ww/dm ~ 0.86; uw/dm ~ 0.23),
mode a ely dep essed who l p o ile (ww/wh ~ 1.90) and e y high coiling a e (WER ~ 2.35) a a conch
diame e o 60 mm. Who l p o ile wi h b oadly a ched en e and lanks, na owly ounded umbilical
Table19. Recons uc ed conch dimensions (in mm) and a ios o Li oce as lep um sp. no . om Baghuk
Moun ain.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32139 53.0 34.0 23.0 10.5 17.0 0.64 1.48 0.20 2.17 0.26
MB.C.32138 41.0 28.0 19.5 7.0 14.0 0.68 1.44 0.17 2.31 0.28
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
56
Fig. 26. Pa ali oce as mac ogas e sp. no . om he A axoce as beds o he Hambas Fo ma ion
a Baghuk Moun ain. A. Holo ype MB.C.32141 (Ko n e al. 2011 Coll.), do sal, la e al and en al
iews. B. The same specimen, econs uc ion o ape u al iew. C. The same specimen, su u e line a
dm = 54.5 mm, wh = 23.5 mm. D. Pa a ype MB.C.32142 (Ko n e al. 2011 Coll.), en al and la e al
iews. E. The same specimen, econs uc ion o ape u al iew. F. The same specimen, su u e line a
dm = 40.5 mm, ww = 35.8 mm, wh = 21.6 mm. Scale ba uni s = 1 mm.
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
57
wo gene a show a much highe coiling a e (WER abo e 3.00) han Baghuknau ilus (WER a ound 2.50)
and a much mo e p onounced subangula o angula umbilical ma gin.
Baghuknau ilus gen. no . also has simila i y o he T iassic genus Pa anau ilus (Kummel 1953; Sobole
1989), bu his di e s in he mo e do sal posi ion o he siphuncle and he p esence o an annula p ocess.
In addi ion, mos species o Pa anau ilus ha e a con ex umbilical wall.
Baghuknau ilus aplomo phus gen. e sp. no .
u n:lsid:zoobank.o g:ac :0E6D1116-BB35-4615-832A-A52849FB4934
Fig. 29; Table 22
Diagnosis
Species o Baghuknau ilus gen. no . wi h hinly discoidal, in olu e conch (ww/dm ~ 0.40; uw/dm ~ 0.10),
weakly comp essed who l p o ile (ww/wh ~ 0.65) and ex emely high coiling a e (WER ~ 2.55) a a
conch diame e o 60 mm. Who l p o ile wi h con ex en e , con ex and con e gen lanks, na owly
ounded umbilical ma gin, a s eep umbilical wall and mode a ely wide imp in zone (IZR ~ 0.30). Su u e
line wi h a e y shallow ex e nal lobe and an asymme ic, b oadly ounded la e al lobe.
E ymology
F om he G eek ‘ἁπλόος’ = ‘simple’ and ‘μορφή’ = ‘ o m’; because o he a he simple conch mo phology.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 29A–C; MB.C.32146.
Pa a ype
IRAN – Es ahan P o ince • 1 specimen; Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 29D–E; MB.C.32147.
Desc ip ion
Holo ype MB.C.32146 is an incomple e, bu on he le side a he well-p ese ed ully
chambe ed specimen wi h a conch diame e o 83 mm (Fig. 29A). Some conch pa ame e s can only
be econs uc ed. The conch is hinly discoidal and in olu e (ww/dm ~ 0.40; uw/dm = 0.10). The who l
p o ile is weakly comp essed (ww/wh ~ 0.64) wi h a con ex en e and con ex, con e gen lanks. The
umbilical ma gin is na owly ounded and he umbilical wall is s eep and con ex (Fig. 29B). The las
olu ion possesses abou 22 ph agmocone chambe s (CLI = 16.5) wi h an indica ion o c owding a he
end. The siphuncle has a do socen al posi ion. The su u e line shows a small and e y shallow lobe on
en e , a b oadly ounded asymme ic la e al lobe wi h s eepe do sal lanks and a shallow umbilical
lobe (Fig. 29C).
Pa a ype MB.C.32147 is a bila e ally co oded, ully chambe ed specimen wi h a conch diame e o
60 mm (Fig. 29E). Some conch pa ame e s on he las hal olu ion can only be econs uc ed. The
conch is hinly discoidal and in olu e (ww/dm = 0.42; uw/dm = 0.12) wi h an ex emely high coiling
a e (WER = 2.54) and a deep who l o e lap (IZR = 0.32). The who l p o ile is mode a ely comp essed
(ww/wh = 0.76) wi h a con ex en e and con ex con e gen lanks (Fig. 29D). The su u e line shows
a shallow, ounded lobe on en e and a b oadly ounded sligh ly asymme ic la e al lobe wi h s eepe
do sal lanks.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
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Fig. 29. Baghuknau ilus aplomo phus gen. e sp. no . om he Vedioce as beds o he Hambas
Fo ma ion a Baghuk Moun ain. A. Holo ype MB.C.32146 (Ko n e al. 2011 Coll.), la e al iew. B. The
same specimen, econs uc ion o ape u al iew. C. The same specimen, su u e line a dm = 72.5 mm,
wh = 38.8 mm. D. Pa a ype MB.C.32147 (Ko n e al. 2011 Coll.), econs uc ion o ape u al iew.
E. The same specimen, do sal and la e al iews. Scale ba uni s = 1 mm.
Table 22. Conch dimensions (in mm) and a ios o Baghuknau ilus aplomo phus gen. e sp. no .;
econs uc ed alues in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32146 83.4 33.0 51.5 8.2 – 0.40 0.64 0.10 – –
MB.C.32147 60.1 25.0 33.0 7.3 22.4 0.42 0.76 0.12 2.54 0.32
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
65
Rema ks
Baghuknau ilus aplomo phus gen. e sp. no . di e s om Shah ezanau ilus ghade ii gen. e sp. no . by
he conside ably slende e conch; he ww/dm a io is abou 0.40 in B. aplomo phus bu abou 0.60 in
S. ghade ii. The coiling a e is much lowe in B. aplomo phus (WER highe abou 2.50 in compa ison
wi h mo e han 3.00 in S. ghade ii).
Genus Shah ezanau ilus gen. no .
u n:lsid:zoobank.o g:ac :449ADF07-95B8-481E-8C4B-4114FD3DC347
New genus E – Ko n 2025: 69.
Type species
Shah ezanau ilus weye i gen. e sp. no .
Diagnosis
Genus o he amily Jul anau ilidae wi h hinly pachyconic, subin olu e conch. Conch apidly inc easing
in heigh wi h an ex ao dina ily high coiling a e (WER highe han 3.00). Who ls weakly emb acing,
who l p o ile weakly dep essed. Ven e b oadly ounded, umbilical wall la and s eep. Adul o namen
wi h ex emely ine g ow h lines. Sep a wi hou in lexions, sligh ly conca e. Su u e line simple wi h a
shallow ex e nal lobe and a b oadly ounded la e al lobe.
E ymology
Named a e he ci y o Shah eza, he ype locali y o he ype species.
Included species
Cen al I an ( his pape ): Shah ezanau ilus weye i gen. e sp. no ., Wuchiapingian; Shah ezanau ilus
ghade ii gen. e sp. no ., Wuchiapingian.
Rema ks
Shah ezanau ilus gen. no . di e s om Jul anau ilus in he b oadly ounded en e , which in
Jul anau ilus has a conca e shape.
Shah ezanau ilus weye i gen. e sp. no .
u n:lsid:zoobank.o g:ac :FBA54AF0-D7E6-4EBB-BBFA-2563F160A99D
Fig. 30; Table 23
Diagnosis
Species o Shah ezanau ilus gen. no . wi h a hinly pachyconic, subin olu e conch (ww/dm ~ 0.70;
uw/dm ~ 0.25), weakly dep essed who l p o ile (ww/wh ~ 1.40) and ex ao dina ily high coiling a e
(WER ~ 3.40) a a conch diame e o 60 mm. Who l p o ile wi h a weakly con ex en e , a p onounced
en ola e al shoulde , la ly con ex and weakly con e gen lanks, a subangula umbilical ma gin, a
s eep and la ened umbilical wall and a small imp in zone (IZR ~ 0.10). Su u e line wi h a b oadly
ounded and shallow ex e nal lobe and a b oadly ounded la e al lobe.
E ymology
Named a e Die e Weye (Be lin), who collec ed he holo ype o he new species.
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Shah eza; Hambas Fo ma ion, Vedioce as beds (la e Wuchiapingian);
2002; D. Weye leg.; illus a ed in Fig. 30; MB.C.32148.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
66
Pa a ype
IRAN – Es ahan P o ince • 1 specimen; Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; MB.C.32149.
Desc ip ion
Holo ype MB.C.32148 is an incomple e, ully chambe ed specimen wi h a ph agmocone diame e o
a leas 75 mm (Fig. 30A). I has been p ese ed wi hou emains o he shell. The conch is, a 57 mm
diame e , hinly pachyconic and subin olu e (ww/dm = 0.68; uw/dm = 0.26) wi h and ex ao dina ily
high coiling a e (WER = 3.38) and a small who l o e lap zone (IZR = 0.07). The who l p o ile is weakly
dep essed (ww/wh = 1.39) wi h a con ex en e , a sligh ly p onounced by ounded en ola e al shoulde
and sligh ly la ened con e gen lanks. The umbilical ma gin is e y dis inc i e and subangula ; i
delimi a es he b oad and nea ly la , almos pe pendicula umbilical wall (Fig. 30B). The su u e line
shows a shallow and b oadly ounded ex e nal lobe and a wice as wide and deep, b oadly ounded la e al
lobe (Fig. 30C). The las olu ion o he ph agmocone has sligh ly mo e han 20 sep a (CLI = 18) and
hus some indica ion o e minal sep al c owding.
Rema ks
Shah ezanau ilus weye i gen. e sp. no . di e s om S. ghade ii gen. e sp. no . in ha ing a much wide
umbilicus; he uw/dm a io is abou 0.25 in S. weye i gen. e sp. no . bu only abou 0.15 in S. ghade ii.
This di e ence pa allels a di e ence in he who l o e lap a e, which is only abou 0.07 in S. weye i, bu
0.25 in S. ghade ii and in he coiling a e, which is highe in S. weye i (WER ~3.40 compa ed o abou
3.10 in S. ghade ii).
Fig. 30. Shah ezanau ilus weye i gen. e sp. no ., holo ype MB.C.32148 (Weye 2002 Coll.) om he
Vedioce as beds o he Hambas Fo ma ion a Shah eza. A. Ven al and la e al iews. B. Recons uc ion
o ape u al iew. C. Su u e line a dm = 50.2 mm, ww = 36.3 mm, wh = 26.5 mm. Scale ba uni s = 1 mm.
Table 23. Conch dimensions (in mm) and a ios o Shah ezanau ilus weye i gen. e sp. no .
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32148 57.0 39.0 28.0 15.0 26.0 0.68 1.39 0.26 3.38 0.07
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
67
Shah ezanau ilus weye i gen. e sp. no . is di icul o con use wi h any o he La e Pe mian nau iloid
species due o i s speci ic mo phology wi h a e y high, la ened umbilical wall and an angula umbilical
ma gin. The wo species Jul anau ilus hai ape iani Ko n & Ghade i, 2025 and especially J. ashou ii
Ko n & Ghade i, 2025 show a sligh simila i y, al hough hey ha e a conca e en e .
Shah ezanau ilus ghade ii gen. e sp. no .
u n:lsid:zoobank.o g:ac :8FA187E3-F00F-44DE-A0FA-32A9E26A7B37
Fig. 31; Table 24
Diagnosis
Species o Shah ezanau ilus gen. no . wi h hinly pachyconic, in olu e conch (ww/dm ~ 0.62; uw/dm
~ 0.13), weakly dep essed who l p o ile (ww/wh ~ 1.10) and ex ao dina ily high coiling a e (WER
~ 3.10) a a conch diame e o 80 mm. Who l p o ile wi h con ex en e , con ex and con e gen lanks,
a subangula umbilical ma gin, a s eep umbilical wall and a mode a ely deep imp in zone (IZR ~ 0.25).
Su u e line wi h a e y shallow ex e nal lobe and a weakly asymme ic, b oadly ounded la e al lobe.
E ymology
Named a e Abbas Ghade i (Mashhad), in honou o his con ibu ion in s udying he Pe mian–T iassic
bounda y.
Fig. 31. Shah ezanau ilus ghade ii gen. e sp. no ., holo ype MB.C.32150 (Ko n e al. 2011 Coll.) om
he Vedioce as beds o he Hambas Fo ma ion a Baghuk Moun ain. A. Recons uc ion o ape u al
iew. B. Do sal and la e al iews. C. Su u e line a dm = 48.5 mm, ww = 32.5 mm, wh = 29.5 mm. Scale
ba uni s = 1 mm.
Eu opean Jou nal o Taxonomy 1019: 1–76 (2025)
68
Type ma e ial
Holo ype
IRAN – Es ahan P o ince • Baghuk Moun ain; Hambas Fo ma ion, Vedioce as beds (la e
Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 31; MB.C.32150.
Desc ip ion
Holo ype MB.C.32150 is a la gely chambe ed specimen wi h a conch diame e o 84 mm (Fig. 31B),
which is hea ily co oded on he las hal olu ion. Some o he conch pa ame e s on he las hal olu ion
can only be econs uc ed. The conch is hinly pachyconic and in olu e (ww/dm ~ 0.62; uw/dm = 0.13)
wi h an ex ao dina ily high coiling a e (WER = 3.09) and a mode a ely deep who l o e lap (IZR = 0.25).
The who l p o ile is weakly dep essed (ww/wh ~ 1.08) wi h a con ex en e , con ex con e gen lanks,
a na owly ounded umbilical ma gin and a s eep umbilical wall (Fig. 31A). The su u e line shows a
e y shallow, ounded lobe on en e and a b oadly ounded, nea ly symme ic la e al lobe (Fig. 31C).
Rema ks
Shah ezanau ilus ghade ii gen. e sp. no . di e s om S. weye i gen. e sp. no . in ha ing a much
na owe umbilicus; he uw/dm a io is abou 0.15 in S. ghade ii bu abou 0.25 in S. weye i. This
di e ence pa allels a di e ence in he who l o e lap a e, which is abou 0.25 in S. ghade ii, bu only
0.07 in S. weye i and in he coiling a e, which is lowe in S. ghade ii (WER ~ 3.00 compa ed o abou
3.40 in S. weye i).
Shah ezanau ilus ghade ii gen. e sp. no . has some simila i ies wi h Jul anau ilus ashou ii Ko n &
Ghade i, 2025 om Jul a. Like S. ghade ii, J. ape um has a s ou , in olu e conch (ww/dm ~ 0.65),
which is mo e umbilica ed in J. ape um (uw/dm ~ 0.25 in J. ape um, bu only 0.15 in S. ghade ii). The
main di e ence be ween he wo species is he shape o he en e , which is con ex in S. ghade ii and
conca e in J. ape um.
Discussion
The La e Pe mian Hambas Fo ma ion o sec ions a Baghuk Moun ain (Cen al I an) has yielded di e se
nau iloid assemblages. These species belong o 14 gene a, six o which a e new: Lu onau ilus gen. no .,
Epi ainoce as gen. no ., Ocunau ilus gen. no ., Ai inau ilus gen. no ., Baghuknau ilus gen. no . and
Shah ezanau ilus gen. no . Based on new ma e ial, a o al o 24 new species is desc ibed. The ma e ial
comes om ou s a ig aphic uni s, in ascending o de :
•
A axoce as beds (ea ly Wuchiapingian), nine species: Epi ainoce as lu ense gen. e sp. no .,
Se ome acoce as pen agonum sp. no ., Doma oce as myloide sp. no ., Ocunau ilus diplodocus
gen. e sp. no ., Ocunau ilus coelodesmus gen. e sp. no ., Ocunau ilus achy ephus gen. e sp. no .,
Ai inau ilus icanus gen. e sp. no ., Pa ali oce as mac ogas e sp. no . and Pe monau ilus adelphidus
sp. no .
• P o o oce as beds (ea ly Wuchiapingian), one species: Ta dunau ilus ape imos sp. no .
• Vedioce as beds (la e Wuchiapingian), 12 species: Lu onau ilus c a us gen. e sp. no ., Lu onau ilus
elachus gen. e sp. no ., Lu onau ilus cymus gen. e sp. no ., Foo dice as eicosacan hum sp. no .,
Foo dice as decacan hum sp. no ., Foo dice as asce um sp. no ., Doma oce as canonium sp. no .,
Table 24. Conch dimensions (in mm) and a ios o Shah ezanau ilus ghade ii gen. e sp. no .;
econs uc ed alues in i alics.
N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR
MB.C.32150 83.5 52.0 48.0 10.5 36.0 0.62 1.08 0.13 3.09 0.25
MB.C.32150 47.5 30.5 28.5 7.5 – 0.64 1.07 0.16 – –
KORN D. & HAIRAPETIAN V., La e Pe mian nau iloids om Baghuk Moun ain (Cen al I an)
69

Doma oce as ocomphalum sp. no ., Li oce as lep um sp. no ., Baghuknau ilus aplomo phus gen. e
sp. no ., Shah ezanau ilus weye i gen. e sp. no . and Shah ezanau ilus ghade ii gen. e sp. no .
• Pa a i oli es beds (la e Changhsingian), wo species: Tainoce as hys a um sp. no . and Aza inau ilus
pho minx sp. no .
Wi h 24 species, he assemblage om Baghuk is one o he mos di e se La e Pe mian occu ences o
coiled nau iloids. Wi h 22 Wuchiapingian species alone, i is he second species- ich assemblage o his
in e al a e he T anscaucasian egion.
Acknowledgemen s
We a e indeb ed o Lucyna Leda (Zama e) and Abbas Ghade i (Mashhad) o pa icipa ing in he ield
ips and collec ing many o he specimens desc ibed he e. We also hank Ma kus B inkmann (Be lin) o
p epa a ion o he specimens and Oska We b (Be lin) o aking pho og aphs. We g ea ly acknowledge
he e iews o an ea lie e sion by Ma ina Aub ech o á (P ague) and an anonymous e iewe . We also
acknowledge he ca e ul edi ing by K is iaan Hoedemake s and Radka Rosenbaumo á.
Funding
D.K.: Deu sche Fo schungsgemeinscha ; DFG p ojec s Ko1829/12-1, Ko1829/18-1 as well as he
TERSANE esea ch g oup FOR 2332.
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