Late Permian nautiloids from Julfa (NW Iran)

La e Pe mian nau iloids om Jul a (NW I an) Die e KORN 1,* & Abbas GHADERI 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, Facul y o Science, Fe dowsi Uni e si y o Mashhad, Azadi Squa e, 9177948974 Mashhad, I an. * Co esponding au ho s: die e [email p o ec ed], [email p o ec ed] Abs ac . The La e Pe mian Jul a and Ali Bashi o ma ions o sec ions nea Jul a (NW I an) ha e yielded di e se nau iloid assemblages. These come om he ea ly Wuchiapingian A axoce as Beds (19 species), he la e Wuchiapingian Vedioce as Beds (six species), he ea ly Changhsingian Dzhul i es Beds ( h ee species) and he la e Changhsingian Pa a i oli es Limes one ( wo species). These species belong o 20 gene a, eigh o which a e new: Fididoma oce as gen. no ., Aza inau ilus gen. no ., Se ome acoce as gen. no ., Alibashinau ilus gen. no ., Ta dunau ilus gen. no ., Co o ainoce as gen. no ., Cele oli oce as gen. no ., Jul anau ilus gen. no . A o al o 30 species a e desc ibed, o which 24 a e new: Doma oce as elegan ulum sp. no ., Doma oce as mul i ube cula um sp. no ., Pe modoma oce as hamdii sp. no ., Fididoma oce as in acos a um gen. e sp. no ., Aza inau ilus nahidae gen. e sp. no ., Ai inau ilus hebes sp. no ., Se ome acoce as cingulum gen. e sp. no ., Se ome acoce as in la um gen. e sp. no ., Se ome acoce as pa i ube cula um gen. e sp. no ., Se ome acoce as a asense gen. e sp. no ., Alibashinau ilus e us gen. e sp. no ., Alibashinau ilus ambiguus gen. e sp. no ., Ta dunau ilus nimius gen. e sp. no ., Ta dunau ilus mino gen. e sp. no ., Tainoce as admonens sp. no ., Tainoce as la ecos a um sp. no ., Tainoce as uni um sp. no ., Co o ainoce as ine me gen. e sp. no ., Tainionau ilus deinceps sp. no ., Li oce as choopani sp. no ., Cele oli oce as cele e gen. e sp. no ., Pe ipe oce as pa um sp. no ., Jul anau ilus ashou ii gen. e sp. no . and Jul anau ilus hai ape iani gen. e sp. no . The new amily Jul anau ilidae am. no . is e ec ed o accommoda e he species o Jul anau ilus gen. no . Wi h 30 species, he assemblage om he a ea a ound Jul a is one o he mos di e se La e Pe mian occu ences o coiled nau iloids. Wi h 25 Wuchiapingian species alone, i is he mos species- ich assemblage o his in e al. Keywo ds. Nau iloidea, Nau ilida, Pe mian, I an, mo phology. Ko n D. & Ghade i A. 2025. La e Pe mian nau iloids om Jul a (NW I an). Eu opean Jou nal o Taxonomy 1018: 1–113. h ps://doi.o g/10.5852/ej .2025.1018.3069 1 Eu opean Jou nal o Taxonomy 1018: 1–113 h ps://doi.o g/10.5852/ej .2025.1018.3069 eu opeanjou nalo axonomy.eu ISSN 2118-9773 2025 · Ko n D. & Ghade i A. This wo k is licensed unde a C ea i e Commons A ibu ion License (CC BY 4.0) Recei ed: 25 Augus 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:74A6C5AD-7328-444C-9478-36F290657B6E In oduc ion La e Pe mian nau iloids ha e a wide geog aphical dis ibu ion, bu species- ich occu ences a e es ic ed o a ew egions. A e he p obably i s desc ip ion o he La e Pe mian nau iloid “Nau ilus F eieslebeni” om he Cen al Eu opean Zechs ein Fo ma ion by Geini z (1841), i ook se e al decades un il almos simul aneously Mojsiso ics (1869), S ache (1877), Abich (1878), Waagen (1879) and Kayse (1883) desc ibed u he and some imes mo e species- ich occu ences om he Dolomi es (I aly), T anscaucasia (Aze baijan), he Sal Range (Pakis an) and Jiangxi (Sou h China), espec i ely. Despi e he g owing in e es in he ansi ion om he Palaeozoic o he Mesozoic since he end o he 19 h cen u y, i ook decades o publish mo e comp ehensi e s udies o La e Pe mian nau iloids. Reed (1931, 1944) e ised he assemblages om he Sal Range and desc ibed new species. Fo he si es nea he A axes Ri e in he T anscaucasus, i ook e en longe un il he 1960s (Shimansky 1962c, 1965b). In he las decades, a he di e se La e Pe mian nau iloid assemblages we e desc ibed om wo egions: (1) Sou h China: Xu (1977), Zhao e al. (1978), Liang (1984), Zheng (1984), Qin (1986), Yang e al. (1987), Wu & Kuang (1992) and Miao e al. (2021) desc ibed species- ich assemblages om a ious places and di e en acies in he Jiangxi, Zhejiang, Anhui, Hubei, Sichuan, Guizhou, Hunan and Guangxi p o inces. (2) Dolomi es: Posena o & P ino h (2004) and P ino h & Posena o (2007) desc ibed a mode a ely la ge sui e o nau iloids om he Changhsingian Belle ophon Fo ma ion and sepa a ed be ween eigh species wi hin i e gene a. In he ollowing, we desc ibe newly assembled nau iloid collec ions om he sou h side o he A as (= A axes) i e in no h-wes e n I an (Fig. 1). These new inds come om ou gene al s a ig aphic uni s, (1) he ea ly Wuchiapingian beds cha ac e ised by he ammonoid genus A axoce as Ruzhence , 1959, (2) he la e Wuchiapingian beds wi h he ammonoid Vedioce as Ruzhence , 1962, (3) he ea ly Changhsingian beds cha ac e ised by he ammonoids I ani es Teiche & Kummel, 1973, Dzhul i es She y e , 1965 and She y e i es Teiche & Kummel, 1973 and (4) he la e Changhsingian Pa a i oli es Limes one (Fig. 2). This is he mos s a ig aphically de ailed La e Pe mian nau iloid succession known om a single egion. Fig. 1. Geog aphical posi ion o Pe mian-T iassic bounda y sec ions in he T anscaucasus-NW I an egion (a e A akelyan e al. 1965); he ed do s ep esen he sec ions in es iga ed in his s udy ( om Ko n e al. 2016). Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 2 This s udy complemen s he p e ious wo k in he sec ions a ound Jul a, which was ca ied ou as pa o a p ojec suppo ed by he Ge man Resea ch Founda ion. These s udies examined he gene al sedimen a y succession and ecological changes (Leda e al. 2014; Schobben e al. 2015; Gliwa e al. 2020), palaeoclima ic changes (Schobben e al. 2014; Gliwa e al. 2022), succession and di e si y o b achiopods (Ghade i e al. 2014, 2015), ammonoids (Ko n e al. 2016; Kiessling e al. 2018; Ko n & Ghade i 2019; Ko n e al. 2019; Ghanizadeh Tab izi e al. 2021), conodon s (Isaa e al. 2016; Gliwa e al. 2020) and os acods (Gliwa e al. 2021). His o ical e iew A de ailed his o ical e iew o he ea lie s udies o nau iloids om T anscaucasia, and in pa icula om si es in he A as Valley, was gi en by Teiche & Kummel (1973). They discussed he p e ious Fig. 2. S a ig aphical posi ions o he La e Pe mian nau iloid axa om he egion o Jul a. A ax = A axoce as Beds; Vedi = Vedioce as Beds; Dzhu = Dzhul i es Beds; Pa a = Pa a i oli es Limes one. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 3 pionee ing s udies, which mainly ocused on he amous Do asham locali y, 10–16 km wes o he owns o Dzhul a (Nakhiche an P o ince, Aze baijan) and Jul a (Eas Aze baijan P o ince, I an) on he no he n side o he A as Ri e . In he i s desc ip ion o nau iloids om Do asham, Abich (1878) dis inguished be ween 18 species (Figs 3–5), six o which a e s aigh and 12 a e coiled (o iginal names and e isions): “Nau ilus excen icus Eichwald” = Pe monau ilus abichi (K uglo , 1928) “Nau ilus p opinquus, no . o m.” = Pe monau ilus abichi (K uglo , 1928) “Nau ilus pa allelus, no . o m.” = Doma oce as pa allelum (Abich, 1878) Fig. 3. Rep oduc ions o nau iloid igu es o Abich (1878). A. “Nau ilus do so plica us” Abich, 1878. B. “Nau ilus do so plica us”. C. “Nau ilus do so a ma us” Abich, 1878. D. “Nau ilus ince us” on A habe , 1900. E. “Nau ilus Pichle i” on A habe , 1900. F. “Nau ilus ube cula is” Abich, 1878. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 4 “Nau ilus con e gens, no . o m.” = Doma oce as con e gens (Abich, 1878) “Nau ilus conca us Sowe by” = Pe monau ilus abichi (K uglo , 1928) “Nau ilus dole us, no . o m.” = unknown “Nau ilus do so a ma us, no . o m.” = Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no . “Nau ilus Pichle i on Haue ” = Se ome acoce as e ae ( on A habe , 1900) gen. e comb. no . “Nau ilus ube cula is, no . o m.” = Se ome acoce as ube cula is (Abich, 1878) gen. e comb. no . “Nau ilus do so plica us, no . o m.” = Tainoce as do soplica um (Abich, 1878) “Nau ilus a meniacus, no . o m.” = Pseudo i anoce as a meniacum (Abich, 1878) “Nau ilus ince us” = Se ome acoce as e ae ( on A habe , 1900) Von A habe (1900) added he new coiled nau iloid species “Pleu onau ilus Ve ae”, now Se ome acoce as e ae ( on A habe , 1900) gen. e comb. no . and S oyanow (1910) added one new s aigh nau iloid species and one addi ional i s eco d o a coiled nau iloid. Fig. 4. Rep oduc ions o nau iloid igu es o Abich (1878). A. “Nau ilus pa allelus” Abich, 1878. B. “Nau ilus con e gens” Abich, 1878. C. “Nau ilus a meniacus” Abich, 1878. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 5 The i s monog aphic desc ip ion o he La e Pe mian nau iloids om he T anscaucasus was published by Shimansky (1965b) who desc ibed 22 species o coiled nau iloids om he A axoce as Beds, ele en om he Vedioce as Beds (bo h belonging o he Wuchiapingian), i e om he “Phisoni es Beds” (= Zal Membe ) and se en om he Pa a i oli es Limes one (bo h belonging o he Changhsingian). Fo he s aigh nau iloids, he espec i e numbe s we e 6, 2, 1 and 1. I mus be kep in mind ha 60 pe cen o he axa lis ed by Shimansky (1965b: able 5) we e held in open nomencla u e. Ne e heless, Teiche & Kummel (1973) co ec ly s a ed ha his is an imp essi e lis , which demons a es a high di e si y, pa icula ly o he lowe Jul a beds (A axoce as Beds). Teiche e al. (1973) and Teiche & Kummel (1973) in es iga ed sec ions wes o Jul a on he sou he n (I anian) side o he A as alley. They ound essen ially he same nau iloid assemblages, bu compa ison wi h he aunal lis p o ided by Shimansky (1965b) shows ha some axa we e missing, bu also epo ed he p esence o he h ee addi ional gene a Temnocheilus, Tainionau ilus and Ti anoce as in he I anian assemblages. A e his s udy, nau iloids om he en i e T anscaucasian egion we e only occasionally men ioned. Shimansky (1979b) added h ee addi ional axa (A axonau ilus nodosus Shimansky, 1979, A. sp. and Pa a hiphaeoce as p obum Shimansky, 1979) o he species lis o he La e Pe mian nau iloids o he T anscaucasus. Ko lya e al. (1983) p o ided occu ence lis s o La e Pe mian ossils om a ious places in A menia and Aze baijan including nau iloid species, bu hey did no p o ide illus a ions o hei ma e ial. Zakha o (in Ko lya e al. 1989) desc ibed he new Middle Pe mian species “Pleu onau ilus dzhagadzu ensis Zakha o in Ko lya e al., 1989”. Gliwa e al. (2020) illus a ed a selec ion o he ma e ial desc ibed in he p esen s udy. Fig. 5. Rep oduc ions o nau iloid igu es o Abich (1878). A. “Nau ilus conca us” Sowe by, 1840. B. “Nau ilus conca us”. C. “Nau ilus excen icus” Eichwald, 1857. D. “Nau ilus p opinquus” K uglo , 1928. Scale ba uni s o all igu es = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 6 Ma e ial and me hods Specimens s udied We in es iga ed a o al o 123 specimens om he Jul a and he Ali Bashi o ma ions. The majo i y o he specimens we e collec ed om loa ma e ial below he ou c ops. Howe e , based on hei li hology, all he specimens desc ibed he e can be undoub edly assigned o a membe o he Jul a and Ali Bashi o ma ions. Fou membe s ha e been dis inguished om bo om o op wi h he ollowing species ha a e ep esen ed (Fig. 2): (1) A axoce as Beds (ea ly Wuchiapingian): ligh g ey mic i ic nodula limes one (102 specimens) • Doma oce as elegan ulum sp. no . – 9 specimens • Doma oce as mul i ube cula um sp. no . – 6 specimens • Doma oce as con e gens (Abich, 1878) – 3 specimens • Pe modoma oce as hamdii sp. no . – 10 specimens • Fididoma oce as g acile (Shimansky, 1965) gen. e comb. no . – 1 specimen • Fididoma oce as in acos a um gen. e sp. no . – 2 specimens • Aza inau ilus nahidae gen. e sp. no . – 4 specimens • Ai inau ilus hebes sp. no . – 1 specimen • Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no . – 3 specimens • Se ome acoce as do ashamense (Shimansky, 1965) gen. e comb. no . – 16 specimens • Se ome acoce as e ae ( on A habe , 1900) gen. e comb. no . – 8 specimens • Se ome acoce as cingulum gen. e sp. no . – 1 specimen • Se ome acoce as in la um gen. e sp. no . – 1 specimen • Alibashinau ilus e us gen. e sp. no . – 3 specimens • Alibashinau ilus sp. – 1 specimen • Ta dunau ilus mino gen. e sp. no . – 1 specimen • Co o ainoce as ine me gen. e sp. no . – 1 specimen • Li oce as choopani sp. no . – 6 specimens • Cele oli oce as cele e gen. e sp. no . – 1 specimen • Pe monau ilus abichi (K uglo , 1928) – 24 specimens (2) Vedioce as Beds (la e Wuchiapingian): pink o ligh g ey nodula limes one (10 specimens) • Se ome acoce as pa i ube cula um gen. e sp. no . – 1 specimen • Tainoce as admonens sp. no . – 2 specimens • Tainoce as la ecos a um sp. no . – 1 specimen • Cele oli oce as sp. – 1 specimen • Pe ipe oce as pa um sp. no . – 2 specimens • Jul anau ilus ashou ii gen. e sp. no . – 1 specimen • Jul anau ilus hai ape iani gen. e sp. no . – 1 specimen • gen. inde . sp. inde . – 1 specimen (3) Dzhul i es Beds (ea ly Changhsingian): g ey nodula limes one wi h high clay con en (7 specimens) • Ocunau ilus sp. – 2 specimens • Se ome acoce as a asense gen. e sp. no . – 1 specimen • Ta dunau ilus nimius gen. e sp. no . – 1 specimen • New genus C o be desc ibed by Ko n & Hai ape ian (in p ess) – 1 specimen • Tainoce as uni um sp. no . – 1 specimen • Ti olonau ilus sp. 2 – 1 specimen (4) Pa a i oli es Limes one (la e Changhsingian): pink o da k ed nodula limes one (4 specimens) • Alibashinau ilus ambiguus gen. e sp. no . – 1 specimen • Tainionau ilus deinceps sp. no . – 1 specimen KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 7 • Ti olonau ilus sp. 1 – 1 specimen • New genus F o be desc ibed by Ko n & Hai ape ian (in p ess) – 1 specimen 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, 1965b, 1967) p oposed a 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, 1979a). This also means ha i di e s om he equen ly 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. 6). 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). The lis s o gene a and species a e no exhaus i e. When lis ing species wi hin gene a, he s a ig aphic uni and he egion o he ype ma e ial ha e been added. Species ha e been named acco ding o hei o iginal genus assignmen . Abb e ia ions used in he species desc ip ions ah = ape u al heigh dm = conch diame e IZR = imp in zone a e SD = sep al densi y (dis ance o sep a in deg ees) uw = umbilical wid h WER = who l expansion a e Fig. 6. 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. Abb e ia ion: 1. = lobe. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 8 wh = who l heigh ww = who l wid h Abb e ia ions o he eposi o ies MB.C. = Cephalopod collec ion o he Museum ü Na u kunde, Be lin, Ge many GLM = Gol a aj Ecomuseum, Jul a, I an PIN = Palaeon ological Ins i u e o he Academy o Sciences, Moscow, Russian Fede a ion LGI = Lening adskiy Go niy Ins i u , Lening ad (now Sank -Pe e bu gskiy Go niy Uni e si e , S . Pe e bu g), Russian Fede a ion 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 i e 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). 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 o 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 one clade ( 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). KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 9 Fig. 9. Doma oce as mul i ube cula um sp. no . om he A axoce as Beds o he Jul a Fo ma ion. A. Holo ype MB.C.31997 (Ko n e al. 2011 Coll.) om A as Valley, en al and la e al iews. B. The same specimen, econs uc ion o ape u al iew. C. The same specimen, who l p o iles. D. The same specimen, su u e line a ww = 26.6 mm, wh = 35.7 mm. E. Pa a ype MB.C.31999 (Ghade i 2018 Coll.) om A as Valley, who l p o ile. F. The same specimen, su u e line a ww = 13.2 mm, wh = 15.5 mm. G. Pa a ype MB.C.31998 (Ghade i 2018 Coll.) om A as Valley, la e al iew. H. The same specimen, econs uc ion o ape u al iew. I. The same specimen, su u e line a ww = 20.1 mm, wh = 24.3 mm. J. Pa a ype MB.C.32000 (Ghade i 2018 Coll.) om A as Valley, who l p o ile. K. The same specimen; su u e line a ww = 19.5 mm, wh = 26.3 mm. L. Specimen MB.C.32002 (Ghade i 2018 Coll.) om Ali Bashi 4, who l p o ile. M. The same specimen; su u e line a ww = 16.1 mm, wh = 20.0 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 16 The e is no sculp u e isible on he las who l, bu he wo who ls be o e show shallow and b oad en ola e al ube cles, which in he hi d-las who l e en shows a p olonga ion in o ibs ha a e connec ed wi h ba ely isible umbilical nodes. The specimen has a he na owly s anding sep a; on he las qua e olu ion o he ph agmocone he e a e nine sep a wi h a end o c owding a he end (SD ~ 10 deg ees in a e age). The e minal su u e line has a e y shallow ex e nal lobe, a b oadly ounded la e al lobe and a e y shallow, en ally inclined lobe on he umbilical wall (Fig. 9D). Pa a ype MB.C.31998 is, hough incomple e, one o he bes in he a ailable ma e ial (Fig. 9G). I is a ully sep a e specimen wi h an es ima ed diame e o abou 73 mm and allows he s udy o wo who ls o he ph agmocone. The es ima ed ww/dm and uw/dm a ios a e 0.31 and 0.37, espec i ely. The who l p o ile is weakly comp essed (ww/wh = 0.79) wi h a nea ly la en e , a p onounced subangula en ola e al shoulde and la ened, weakly con e gen lanks. The who ls a e wides a he na owly ounded umbilical ma gin (Fig. 9H). The las who l o he ph agmocone appea s o be ee o sculp u e, bu he penul ima e who l shows a he coa se, low en ola e al conical nodes, which a e a anged in dis ances o abou 15 deg ees. The su u e line has a e y shallow ex e nal lobe, a subangula en ola e al saddle (wi h a posi ion a he en ola e al shoulde ) and a b oadly ounded, shallow la e al lobe. Addi ionally, a e y shallow, small and ounded lobe is isible on he umbilical wall (Fig. 9I). Bo h he las and he penul ima e who l show six chambe s on a qua e o a olu ion (SD ~ 15 deg ees). A comple e su u e line is isible in he agmen a y pa a ype MB.C.32000. I shows a a he na ow and shallow ex e nal lobe, a b oadly ounded la e al lobe ha has h ee imes he dep h o he ex e nal lobe. The specimen shows a a he na ow in e nal lobe o h ee qua e s he dep h o he la e al lobe (Fig. 9K). The smalle agmen a y pa a ypes MB.C.31999 (Fig. 9E–F) and MB.C.32002 (Fig. 9L–M) show he a ia ion o he dep h o he ex e nal lobe. Rema ks Doma oce as mul i ube cula um sp. no . di e s om D. elegan ulum sp. no . in he conical en ola e al nodes o he p eadul who ls, he sligh ly mo e e olu e and s ou e conch, he less comp essed who l p o ile and he na owly ounded umbilical ma gin. Doma oce as mul i ube cula um sp. no . di e s om D. con e gens by he less s ongly con e gen lanks and he much wide who l p o ile (ww/wh ~ 0.80 in con as o D. con e gens wi h ~ 0.60) a a conch diame e o 75 mm. The en e is much wide in D. elegan ulum sp. no . when compa ed o D. con e gens. Table 2. Conch dimensions (in mm) and a ios o Doma oce as mul i ube cula um sp. no .; 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.31997 100.9 27.2 37.3 38.6 35.6 0.27 0.73 0.38 2.39 0.05 MB.C.31997 – 23.8 30.3 – – – 0.79 – – – MB.C.31997 – 15.0 17.2 – – – 0.87 – – – MB.C.31998 75.9 21.8 27.6 28.5 26.1 0.29 0.79 0.38 2.32 0.05 MB.C.32000 – 20.4 26.4 – – – 0.77 – – – MB.C.32002 – 16.3 20.1 – – – 0.81 – – – MB.C.32001 – 15.1 18.8 – – – 0.80 – – – MB.C.31999 – 13.3 15.4 – – – 0.86 – – – KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 17 Doma oce as con e gens (Abich, 1878) Fig. 10 Nau ilus con e gens Abich, 1878: 17, pl. 3 ig. 2. Doma oce as con e gens – Shimansky 1965a: 41, pl. 15 ig. 9. Diagnosis Species o Doma oce as wi h ex emely discoidal, sube olu e and weakly comp essed, in e ed apezoidal who l p o ile (ww/wh ~ 0.60) a a conch diame e o 75 mm. Who l p o ile wi h la , e y na ow en e , subangula en ola e al shoulde , con e gen , la ened lanks and b oadly ounded umbilical ma gin. Su u e line wi h a e y na ow and e y shallow ex e nal lobe and a much la ge and deepe , b oadly ounded la e al lobe. Ma e ial examined IRAN – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 10; MB.C.32003 • 1 specimen; same da a as o p eceding; 2018; Ghade i leg.; MB.C.32004. – Eas Aze baijan • 1 specimen; Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2010; Ko n e al. leg.; MB.C.32005. Desc ip ion Specimen MB.C.32003 is, hough agmen a ily p ese ed, sui able o iden i ica ion as i shows some speci ic cha ac e s (Fig. 10A). I has a who l heigh o 26 mm and a ww/wh a io o 0.57. The en e is la and bo de ed by an angula en ola e al shoulde ; he lanks a e weakly con ex and con e ge om he ounded umbilical ma gin (Fig. 10B). The su u e line shows a small and e y shallow ex e nal lobe and a e y wide, b oadly ounded la e al lobe (Fig. 10C). Rema ks Abich (1878: 17, pl. 3 ig. 2) desc ibed and illus a ed, as “Nau ilus con e gens, no . o m.”, a agmen (less han a qua e o a olu ion) o an in e nal mould o 40 mm who l heigh . F om his, he conch shape can be econs uc ed a he well. The in e ed apezoidal who l p o ile is comp essed (ww/wh = 0.62) and e y weakly in lec ed do sally by he p eceding who l. I is wides nea he umbilical ma gin. The lanks con e ge a he apidly and a e sepa a ed om he la ened en e by a subangula en ola e al shoulde . The e is a shallow longi udinal g oo e on he ou e po ion o he lank jus adjacen o he en ola e al shoulde . No o namen can be seen. The su u e line shows a b oad ounded lobe on he lank and a e y shallow lobe on he en e . Fig. 10. Doma oce as con e gens (Abich, 1878), specimen MB.C.32003 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a A as Valley. A. La e al iew. B. Who l p o ile. C. Su u e line a ww = 14.5 mm, wh = 26.0 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 18 Doma oce as pa allelum (Abich, 1878) Nau ilus pa allelus Abich, 1878: 17, pl. 2 ig. 2. Rema ks “Nau ilus pa allelus, no . o m.” was desc ibed and illus a ed by Abich (1878: 17, pl. 2 ig. 2). This illus a ion shows a agmen o less han hal a olu ion o a specimen wi h a conch diame e o abou 120 mm. Only he las who l is p ese ed and his appea s o be de o med o co oded on one side. The e o e, he conch shape canno be p ecisely desc ibed. Howe e , i can be seen ha he umbilicus is a he wide (uw/dm ~ 0.45) and ha he lanks a e almos pa allel. Specimens wi h his mo phology ha e no been collec ed om NW I an. Genus Pe modoma oce as Ruzhence & Shimansky, 1954 Type species Pe modoma oce as apezoidale Ruzhence & Shimansky, 1954; o iginal designa ion. Diagnosis Genus o he amily Doma oce a idae wi h a subin olu e o sube olu e conch. High o ex emely high coiling a e; who l p o ile weakly comp essed o weakly dep 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. 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 species Sou h U als (K uglo 1928; Ruzhence & Shimansky 1954; Ba sko e al. 2014): Doma oce as F ede icksi K uglo , 1928, A inskian, Sou h U als; Pe modoma oce as apezoidale Ruzhence & Shimansky, 1954, A inskian; Pe modoma oce as pe mianum Ba sko & Shilo sky in Ba sko e al., 2014, Roadian; Pe modoma oce as ma ielense Ba sko & Shilo sky in Ba sko e al., 2014, Roadian. T anscaucasia ( his pape ): Pe modoma oce as hamdii sp. no ., Wuchiapingian, NW I an. Himalayas (Diene 1903): Nau ilus hunicus Diene , 1903, Wuchiapingian. Timo (Haniel 1915): Disci es A habe i Haniel, 1915, Kungu ian. Rema ks Acco ding o Ruzhence & Shimansky (1954: 95), Pe modoma oce as is dis inguished om Doma oce as by i s mo e angula and lowe who ls; he a io o who l wid h o heigh is be ween 0.90 and 1.10 in Pe modoma oce as and only abou 0.70 in Doma oce as. In addi ion, he ex e nal lobe is less well de eloped in Pe modoma oce as. Pe modoma oce as is hough o be close o he Ame ican gene a Penascoce as and Pa apenascoce as in e ms o who l p o ile, bu is clea ly dis inguished om hem by a g ea e ampli ude o su u e elemen s and longe chambe s. Pe modoma oce as hamdii sp. no . u n:lsid:zoobank.o g:ac :C3FB68E4-18D0-4B64-946A-75FE355723EA Fig. 11; Table 3 Doma oce as hunicum – Shimansky 1965b: 161, pl. 15 ig. 12. — Teiche & Kummel 1973: 421, pl. 1 igs 7–8, pl. 2 igs 3–4. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 19 Diagnosis Species o Pe modoma oce as wi h ex emely discoidal, sube olu e conch (ww/dm ~ 0.30; uw/dm ~ 0.40), weakly comp essed who l p o ile (ww/wh ~ 0.80) and e y high coiling a e (WER ~ 2.45) a a conch diame e o 80 mm. Who l p o ile wi h weakly conca e en e , subangula en ola e al shoulde , gen ly con e gen , la ened lanks and na owly ounded umbilical ma gin. O namen wi h ine g ow h lines, wi hou ibs o nodes. Su u e line wi h a na ow and e y shallow ex e nal lobe and a much la ge and deepe , b oadly ounded la e al lobe. E ymology Named a e he la e Bahaeddin Hamdi (1935–2019), he p omo e o palaeon ology in I an. Type ma e ial Holo ype IRAN – Eas Aze baijan • Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 11; MB.C.32006. Pa a ypes IRAN – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32007 • 1 specimen; same da a as o p eceding; 2011; Ko n e al. leg.; MB.C.32008 • 1 specimen; same da a as o p eceding; 2013; Ko n e al. leg.; MB.C.32009 • 2 specimens; same da a as o p eceding; 2018; Ko n e al. leg.; MB.C.32010 o MB.C.32011. – Eas Aze baijan • 1 specimen; Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32012 • 1 specimen; same da a as o p eceding; 2010; Ko n e al. leg.; MB.C.32013 • 1 specimen; same da a as o p eceding; 2011; Ko n e al. leg.; MB.C.32014 • 1 specimen; Zal; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32015. Desc ip ion Holo ype MB.C.32006 is a ph agmocone agmen wi h a who l heigh o 30 mm (Fig. 11A) and allows o examina ion o he do sal who l a ea. The shape o he conch was econs uc ed using his agmen ; his esul s in a diame e o 80 mm. The who l p o ile o he specimen is comp essed (ww/wh = 0.80) and apezoidal wi h a e y weakly conca e en e , subangula en ola e al shoulde s, la ened and con e gen lanks, a ounded umbilical ma gin and a e y shallow do sal zone (Fig. 11B). The do sal zone shows ha he penul ima e who l also had a weakly conca e en e . The comple e su u e line is exposed. I shows ha he ex e nal lobe is e y shallow and ha he en ola e al saddle is subangula . The la e al lobe is b oadly ounded and con inues, a he umbilical seam, in o a e y shallow in e nal Fig. 11. Pe modoma oce as hamdii sp. no ., holo ype MB.C.32006 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a Ali Bashi 4. A. La e al iew. B. Who l p o ile. C. Su u e line a ww = 23.3 mm, wh = 27.6 mm. Scale ba uni s = mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 20 lobe, in which a e y low and ounded saddle is aised (Fig. 11C). The sep a a e closely spaced; he e a e abou en chambe s on a qua e o a olu ion (SD ~ 9 deg ees). Some o he o he specimens, such as pa a ype MB.C.32015, which is no illus a ed, show a ans o ma ion o he en ola e al shoulde om a subangula o a igh ly ounded shape in he adul s age. In his specimen he shape change akes place a abou 28 mm who l heigh . Rema ks I is mos likely ha he specimens illus a ed by Shimansky (1965b) and Teiche & Kummel (1973) unde he name “Doma oce as hunicum” belong o his species. Howe e , P. hunicum di e s om he new species in ha ing sep a in much la ge dis ances (abou i e sep a pe qua e olu ion) han P. hamdii sp. no . (10 sep a pe qua e olu ion). Ano he di e ence is he a e on who l o e lap, which is a he g ea in P. hunicum (IZR ~ 0.15), while i is e y low in P. hamdii (IZR ~ 0.05). The new species has a na owe who l p o ile (ww/wh = 0.80) han he ype species o he genus. Howe e , as i s conch shape wi h he almos ec angula , a he wide who l p o ile clea ly dis inguishes i om he species o he genus Doma oce as, i is classi ied he e as Pe modoma oce as. Genus Fididoma oce as gen. no . u n:lsid:zoobank.o g:ac :FE3F69BC-FA94-4E56-9253-D9583FCD3BB2 New genus A – Ko n 2025: 39. Type species Fididoma oce as in acos a um gen. e sp. no . Diagnosis Genus o he amily Doma oce a idae wi h a small o mode a ely la ge, subin olu e conch; who l p o ile ly i o m, en e la o conca e and bo de ed by aised keels, umbilical ma gin na owly ounded. Sculp u e missing o wi h blun ibs on he lanks in he ju enile s age. Su u e line wi h na ow, ounded ex e nal lobe and b oadly ounded la e al lobe. E ymology Combina ion o he La in ‘ idis’ (noun, .) = ‘ly e’ and Doma oce as; because o he ly i o m who l p o ile. Included species T anscaucasia (Shimansky 1965b; his pape ): Fididoma oce as g acile (Shimansky, 1965) gen. e comb. no ., Wuchiapingian, Aze baijan; Fididoma oce as in acos a um gen. e sp. no ., Wuchiapingian, NW I an. Table 3. Conch dimensions (in mm) and a ios o Pe modoma oce as hamdii sp. no .; 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.32006 80.0 24.4 30.5 31.0 29.0 0.31 0.80 0.39 2.46 0.05 MB.C.32009 – 30.6 35.3 – – – 0.87 – – – MB.C.32010 – 28.0 35.0 – – – 0.80 – – – MB.C.32015 – 29.5 33.3 – – – 0.89 – – – MB.C.32007 – 25.5 28.6 – – – 0.89 – – – MB.C.32013 – 21.8 25.4 – – – 0.86 – – – KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 21 Rema ks The new genus di e s om mos o he gene a in he amily Doma oce a idae by i s na ow umbilicus (uw/dm = 0.25–0.30), which in he o he gene a is usually much wide (uw/dm = 0.35–0.40). The mos impo an dis inguishing ea u e, howe e , is he shape o he en ola e al shoulde , which in he o he gene a is igh ly ounded o simply subangula o angula , bu in Fididoma oce as gen. no . is accen ua ed by a shallow in lexion o he ou e lank. The ly i o m who l p o ile in Fididoma oce as is also he main di e ence o he o he wise simila La e Ca boni e ous genus S enodoma oce as. Fididoma oce as g acile (Shimansky, 1965) gen. e comb. no . Fig. 12; Table 4 Doma oce as g acile Shimansky, 1965b: 160, pl. 16 ig. 1. Diagnosis Species o Fididoma oce as gen. no . wi h hinly discoidal, subin olu e conch (ww/dm ~ 0.35; uw/dm ~ 0.25), weakly comp essed who l p o ile (ww/wh ~ 0.70) and ex emely high coiling a e (WER ~ 3.05) a a conch diame e o 40–55 mm. Who l p o ile in e ed apezoidal wi h con e gen lanks; en e and lanks la ened, umbilical ma gin b oadly ounded. Su u e line wi h b oadly ounded, mode a ely deep ex e nal lobe and sligh ly la ge and deepe , b oadly ounded la e al lobe. Type ma e ial Holo ype AZERBAIJAN • Do asham 1; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); illus a ed by Shimansky (1965b: pl. 16 ig. 1); PIN 1572/211. Ma e ial examined IRAN – Eas Aze baijan • 1 specimen; Ali Bashi N; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ko n e al. leg.; illus a ed in Fig. 12; MB.C.32016. Fig. 12. Fididoma oce as g acile (Shimansky, 1965) gen. e comb. no ., specimen MB.C.32016 (Ko n e al. 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a Ali Bashi N. A. La e al iew o le side. B. Recons uc ion o ape u al iew. C. La e al iew o igh side. D. Su u e line a dm = 58.0 mm, ww = 17.0 mm, wh = 21.0 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 22 Desc ip ion Specimen MB.C.32016 is a ai ly comple e, bu a he poo ly p ese ed specimen wi h a conch diame e o 58 mm (Fig. 12A, C). On he igh side i is hea ily enc us ed by a abula co al colony. The conch is hinly discoidal and subin olu e (ww/dm = 0.35; uw/dm = 0.26) wi h a comp essed who l p o ile (ww/ wh = 0.70). The en e is weakly la ened and bo de ed by a subangula en ola e al shoulde . The umbilical ma gin is ounded, om he e he sinuous lanks con e ge o he en e (Fig. 12B). The su u e line has a ela i ely la ge and deep, b oadly ounded ex e nal lobe, a na owly ounded en ola e al saddle and a b oadly ounded, shallow la e al lobe ha occupies he en i e lank (Fig. 12D). Rema ks Shimansky (1965b) discussed he ex e nal simila i y o Fididoma oce as g acile gen. e comb. no . and he La e Ca boni e ous species S enodoma oce as moo ei (Mille , Dunba & Cond a, 1933). He s a ed ha di e ences be ween hese species mainly ega d e y di e en conch sizes and he much mo e in olu e conch o S. moo ei. He hus ega ded his as a case o homeomo phy. Howe e , he size o he specimens can ha dly be used o dis inguish species o e en gene a. The holo ype o F. g acile is only 40 mm in diame e and is p obably a p eadul specimen. The di e ence in umbilical wid h is a be e sepa a ing c i e ion; he holo ype o S. moo ei has, a a conch diame e o 40 mm, a uw/dm a io o 0.21, whe eas his is 0.28 in he holo ype o F. g acile. A hypo he ical c i e ion o di e en ia ion be ween Fididoma oce as gen. no . and S enodoma oce as could be he posi ion o he siphuncle; his is close o he en e in S. moo ei and also in S. kleihegei (Mille , Lane & Unklesbay, 1947). Howe e , he posi ion o he siphuncle in F. g acile gen. e comb. no . is no known. S enodoma oce as kleihegei and S. ga di (Mu phy, 1970) possess a conca e en e bo de ed by a sligh ly aised en ola e al shoulde , which is na owly ounded o subangula in F. g acile. Because o hese di e ences, bu mainly because o he la ge s a ig aphic dis ance in occu ence o he h ee known La e Ca boni e ous No h Ame ican species o S enodoma oce as and F. g acile, we ollow he in e p e a ion o Shimansky (1965b) he e and do no classi y F. g acile in S enodoma oce as. Ins ead, we place i in he new genus Fididoma oce as desc ibed he e. Fididoma oce as g acile gen. e comb. no . can be easily dis inguished om mos o he species o he amily Doma oce a idae om T anscaucasia by he na owe umbilicus, he mo e widely emb acing who ls and he highe coiling a e (WER > 3.00 in F. g acile, bu usually 2.25–2.50 in he o he species). Fididoma oce as g acile di e s om F. in acos a um gen. e sp. no . in he absence o ibs in he ju enile s age and in he less dis inc en ola e al shoulde , which is na owly ounded in F. g acile bu angula in F. in acos a um. Fu he mo e, F. g acile has a highe coiling a e (WER > 3.00) han F. in acos a um (WER < 2.75). Fididoma oce as in acos a um gen. e sp. no . u n:lsid:zoobank.o g:ac :3DDD3943-57AC-4E12-9036-70812D1C1220 Fig. 13; Table 5 Diagnosis Species o Fididoma oce as gen. no . wi h hinly discoidal, subin olu e conch (ww/dm ~ 0.40; uw/ dm ~ 0.27), weakly comp essed who l p o ile (ww/wh ~ 0.90) and ex emely high coiling a e Table 4. Conch dimensions (in mm) and a ios o Fididoma oce as g acile Shimansky, 1965 gen. e comb. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32016 58.4 20.5 29.2 15.4 25.0 0.35 0.70 0.26 3.06 0.14 PIN 1572/211 40.0 15.0 20.5 10.0 17.0 0.38 0.73 0.25 3.02 0.17 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 23 (WER ~ 2.55) a a conch diame e o 30 mm. Who l p o ile in e ed apezoidal o ly i o m wi h con e gen lanks; en e la ened, lanks la ened o sligh ly conca e in he ou e a ea, umbilical ma gin na owly ounded. Sculp u e in he ju enile s age wi h blun ibs on he lanks. Su u e line wi h e y na ow ex e nal lobe and sligh ly la ge and deepe , b oadly ounded la e al lobe. E ymology Combina ion o he La in ‘in a’ (p epos.) = ‘inne ’ and ‘cos a um’ (adj., n.) = ‘ ibbed’; because o he ju enile sculp u e. Type ma e ial Holo ype IRAN – Eas Aze baijan • Ali Bashi N; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ko n e al. leg.; illus a ed in Fig. 13A–B; MB.C.32017. Pa a ype IRAN – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 13C–E; MB.C.32018. Desc ip ion Holo ype MB.C.32017 is he be e p ese ed o he wo compa a i ely small specimens (Fig. 13A). I has a conch diame e o 32 mm and is hinly discoidal and subin olu e (ww/dm = 0.42; uw/dm = 0.26). The coiling a e is ex emely high (WER = 2.61) and he imp in zone is small. The who l p o ile is weakly comp essed (ww/wh = 0.89) and shows a na owly ounded umbilical ma gin and a s eep, sligh ly la ened umbilical wall. The conch is wides nea he umbilical ma gin; om he e he lanks slowly con e ge owa ds he angula en ola e al shoulde , which is p ese ed as a weakly ele a ed idge. The en e becomes la only in he las hal o he olu ion, beginning a a conch diame e o abou 22 mm. Be o e his, he en e is b oadly ounded (Fig. 13B). Shell emains a e isible on la ge a eas o he specimen, hey show pe iodically s eng hened g ow h lines ex ending wi h a b oad a ch in backwa d di ec ion ac oss he lanks and o m a deep, angula en al sinus. Two hi ds o he las olu ion belong o his s age, while he p e ious on ogene ic s age be o e possesses shallow, backwa dly di ec ed ibs on he lanks. Fig. 13. Fididoma oce as in acos a um gen. e sp. no . A. Holo ype MB.C.32017 (Ko n e al. 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a Ali Bashi N, la e al and ape u al iews. B. The same specimen, econs uc ion o ape u al iew. C. Pa a ype MB.C.32018 (Ghade i 2018 Coll.) om A as Valley, la e al and ape u al iews. D. The same specimen, econs uc ion o ape u al iew. E. The same specimen; su u e line a ww = 8.4 mm, wh = 7.0 mm. Scale ba uni s = mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 24 Pa a ype MB.C.32018 has a conch diame e o 31 mm (Fig. 13C). I is hinly discoidal and subin olu e (ww/dm = 0.39; uw/dm = 0.30) wi h a comp essed who l p o ile (ww/wh = 0.87). The en e is ounded up o a conch diame e o 27 mm and becomes la ened he ea e (Fig. 13D). This means ha i eaches he s age wi h p onounced en ola e al shoulde s and he applana e en e a a la ge conch diame e han he holo ype. The su u e line shows a e y shallow, b oadly ounded ex e nal lobe and a sligh ly deepe , ounded la e al lobe (Fig. 13E). Rema ks Fididoma oce as in acos a um gen. e sp. no . can ha dly be con used wi h ano he nau ilid species o T anscaucasia because o i s e y cha ac e is ic mo phology. Fididoma oce as in acos a um di e s om F. g acile gen. e comb. no . in he p esence o ibs in he ju enile s age and in he mo e dis inc en ola e al shoulde , which is angula in F. in acos a um bu na owly ounded in F. g acile. Fu he mo e, F. in acos a um has a lowe coiling a e (WER < 2.75) han F. g acile (WER > 3.00). New amily Ko n & Hai ape ian (in p ess) Diagnosis Family o he supe amily G ypoce a oidea wi h a usually discoidal, subin olu e conch. Who l p o ile in he adul s age weakly comp essed o weakly dep essed; lanks and en e usually sepa a ed by a dis inc en ola e al shoulde , en e mo e o less conca e. Umbilical ma gin usually subangula o angula , a ely ounded; umbilical wall s eep, o en la ened. O namen usually consis ing o ine g ow h lines. Sep um simple in shape, conca ely domed; su u e line depending on who l p o ile wi h shallow o V-shaped ex e nal lobe and shallow la e al lobe ( om Ko n & Hai ape ian in p ess: 14). Included gene a Pseudo i anoce as Shimansky, 1965 (Pe mian); new genus A o be desc ibed by Ko n & Hai ape ian (in p ess) (Pe mian); new genus B o be desc ibed by Ko n & Hai ape ian (in p ess) (Pe mian); Aza inau ilus gen. no . (Pe mian). Rema ks A de ailed discussion o he new amily will be gi en by Ko n & Hai ape ian (in p ess). Genus Aza inau ilus gen. no . u n:lsid:zoobank.o g:ac :CD71EEC0-54D7-47EC-A234-F505CEDED059 New genus B – Ko n 2025: 43. Type species Aza inau ilus nahidae gen. e sp. no . Diagnosis Genus o he new amily Ko n & Hai ape ian (in p ess) 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, Table 5. Conch dimensions (in mm) and a ios o Fididoma oce as in acos a um gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32017 32.0 13.3 15.0 8.4 12.2 0.42 0.89 0.26 2.61 0.19 MB.C.32018 30.3 11.7 13.5 9.0 11.2 0.39 0.87 0.30 2.52 0.17 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 25 Genus Pseudo i anoce as Shimansky, 1965 Type species Nau ilus a meniacus Abich, 1878; o iginal designa ion. Included species Nau ilus a meniacus Abich, 1878, Wuchiapingian, Aze baijan. Rema ks The ype species o he genus is a he poo ly known. Al hough he e has been a b ie e ision by Shimansky (1965b) on he basis o new ma e ial, he e is s ill no clea di e en ia ion om o he species. Shimansky (1965b) discussed he a ia ion in he sculp u e o his specimens; howe e , his may be due o he possibili y ha he also included specimens o he new genus A o be desc ibed by Ko n & Hai ape ian (in p ess). Pseudo i anoce as a meniacum (Abich, 1878) Nau ilus a meniacus Abich, 1878: 24, pl. 2 ig. 5. Pseudo i anoce as a meniacum – Shimansky 1965b: 162, pl. 16 igs 5–6. Type ma e ial Holo ype AZERBAIJAN • Do asham 1; A axoce as Beds (ea ly Wuchiapingian); illus a ed by Abich (1878: pl. 2 ig. 5); LGI 14/99. Rema ks “Nau ilus a meniacus, no . o m.” was desc ibed and illus a ed by Abich (1878: pl. 2 ig. 5). I is a ph agmocone agmen wi h a who l heigh o abou 38 mm. I is de o med bu shows some ea u es o he conch shape and sculp u e. Acco ding o Abich’s desc ip ion, he lanks a e sligh ly conca e and he en e is mo e clea ly conca e; hey a e sepa a ed by a na owly ounded en ola e al shoulde . The en ola e al shoulde bea s closely spaced nodes, which ha e he same wid h as he in e spaces and also co espond app oxima ely o he dis ances be ween he sep a. The su u e line shows b oadly ounded lobes on he lanks and en e . Subo de Tainoce a ina Shimansky, 1957 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 in he ad anced species. Conch usually discoidal, subin olu e o e olu e. Ju enile who l p o ile dep essed o al o ci cula . Adul who l p o ile dep essed o al o eni o m in he ea ly species, showing nume ous modi ica ions du ing e olu ion (in e ed apezoidal, apezoidal o polygonal who l p o iles o wi h en al dep ession). Do sal who l zone always p esen , bu usually e y small. Ju enile sculp u e wi h adial ibs on he lank; adul sculp u e wi h adial ibs on he lank, en ola e al nodes o se e al ows o nodes in de i ed species. Sep a simply domed; wi h do sal in lexion in ad anced species. Su u e line depending on he who l p o ile, wi h shallow lobes and low saddles ( om Ko n 2025). Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 32 Included supe amilies Tainoce a oidea Hya , 1883 (Ca boni e ous o T iassic); Pleu onau iloidea Hya , 1900 (Ca boni e ous o T iassic). Rema ks A de ailed discussion o he subo de Tainoce a ina has been gi en by Ko n (2025). Supe amily Pleu onau iloidea Hya , 1900 Diagnosis Supe amily o he subo de Tainoce a ina wi h a discoidal, subin olu e o sube olu e conch. Who l p o ile in ea ly species subquad a e wi h dis inc en ola e al shoulde and dis inc umbilical ma gin. De i ed species show a a ia ion o modi ica ions including apezoidal, in e ed apezoidal o hexagonal who l p o iles wi h a less angula en ola e al shoulde and umbilical ma gin. Who l o e lap is always e y small. Sculp u e in ea ly species wi h ans e se ibs and en ola e al nodes, in de i ed species o en wi h ibs and se e al ows o nodes. Sep a simply domed, in de i ed species wi h do sal in lexion ha p oduces an annula p ocess. Su u e line wi h b oadly ounded la e al lobe and shallow lobe o low saddle on he en e ( om Ko n 2025). Included amilies Pleu onau ilidae Hya , 1900 (Pe mian o T iassic); Gzheloce a idae Ruzhence & Shimansky, 1954 (Ca boni e ous o Pe mian); Mosquoce a idae Ruzhence & Shimansky, 1954 (Ca boni e ous o Pe mian); Ak ubonau ilidae Ruzhence & Shimansky, 1954 (Pe mian); Rhiphaeoce a idae Ruzhence & Shimansky, 1954 (Pe mian); Me acoce a idae Ko n, 2025 (Ca boni e ous o Pe mian); Foo dice a idae Ko n, 2025 (Pe mian). Rema ks A de ailed discussion o supe amily Pleu onau iloidea has been gi en by Ko n (2025). Family Me acoce a idae Ko n, 2025 Diagnosis Family o he supe amily Pleu onau iloidea wi h an equidimensional o mo e commonly weakly dep essed, apezoidal o in e ed apezoidal who l p o ile. Ven e usually la ened, bu anging om sligh ly con ex o sligh ly conca e. Ven ola e al shoulde o en p ominen , anging om b oadly ounded o subangula . Flanks weakly con e gen , pa allel o weakly di e gen , usually la ened and anging om weakly con ex o weakly conca e. Umbilical ma gin usually p onounced, usually subangula in he in e media e g ow h s age. Sculp u e wi h en ola e al conical nodes, o en wi h do sola e al nodes and low ibs on he lank. Su u e line wi h shallow lobes and low saddles. In e nal lobe e y shallow, wi hou annula p ocess ( om Ko n 2025). Included gene a Me acoce as Hya , 1883 (Ca boni e ous o T iassic); Mojs a oce as Hya , 1883 (T iassic); Huanghoce as Yin, 1933 (Pe mian); ? Shansinau ilus Yabe & Mabu i, 1935 (Pe mian;); Coope oce as Mille , 1945 (Pe mian); Epime acoce as Lib o i ch, 1946 (Ca boni e ous) (nomen nudum); Pseudo oo dice as Ruzhence & Shimansky, 1954 (Pe mian); Pseudo emnocheilus Ruzhence & Shimansky, 1954 (Pe mian); Tanchiashani es Zhao, 1954 (Pe mian); Mahoningoce as Mu phy, KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 33 1974 (Ca boni e ous); Lichuanoce as Xu, 1977 (Pe mian); Sino i anoce as Pan, 1983 (Pe mian); An hodiscoce as Qin, 1986 (Pe mian); Se ome acoce as gen. no . (Pe mian). Rema ks A de ailed accoun o he esea ch his o y o Me acoce as and gene a wi h simila mo phology has been gi en by Ko n (2025). The e o e, only he di e ences be ween he new genus Se ome acoce as gen. no . and Me acoce as will be discussed he e. Genus Se ome acoce as gen. no . u n:lsid:zoobank.o g:ac :28DA9B4A-3B7E-421E-A054-779F73B20EA9 New genus C – Ko n 2025: 50. Type species Pleu onau ilus Ve ae on A habe , 1900. Diagnosis Genus o he amily Me acoce a idae wi h a subin olu e o sube olu e conch; who l p o ile equidimensional o mo e o less s ongly dep essed, usually apezoidal wi h weakly di e gen lanks. Ven e usually weakly con ex o la ened; en ola e al shoulde na owly o b oadly ounded. Umbilical ma gin p onounced and subangula in he in e media e g ow h s age, ounded in he adul s age. Sculp u e wi h conical nodes on he en ola e al shoulde o on he umbilical ma gin o bo h, some imes connec ed by low ibs on he lank. Su u e line wi h shallow ex e nal lobe o e y low ex e nal saddle and b oadly ounded la e al lobe; wi hou annula p ocess. Siphuncle small wi h subcen al posi ion en ad o sep um cen e. E ymology F om he La in ‘se us’ (adj., ne .) = ‘la e’; because o he high s a ig aphic posi ion and he simila i y o Me acoce as. Included species T anscaucasia and NW I an (Abich 1878; on A habe 1900; Shimansky 1965b; Ko lya e al. 1989; his pape ): Nau ilus do so a ma us Abich, 1878, Wuchiapingian, Aze baijan; Nau ilus ube cula is Abich, 1878, Wuchiapingian, Aze baijan; Nau ilus ince us Abich, 1878, Wuchiapingian, Aze baijan; Nau ilus Ve ae on A habe , 1900, Wuchiapingian, Aze baijan; Me acoce as do ashamense Shimansky, 1965, Wuchiapingian, Aze baijan; Pleu onau ilus dzhul ensis Shimansky, 1965, Wuchiapingian, Aze baijan [synonym o Se ome acoce as e ae gen. e comb. no .]; Pleu onau ilus cos alis Shimansky, 1965, Wuchiapingian, A menia; Pleu onau ilus dzhagadzu ensis Zakha o in Ko lya e al., 1989, Capi anian, Aze baijan; Se ome acoce as cingulum gen. e sp. no ., Wuchiapingian, NW I an; Se ome acoce as in la um gen. e sp. no ., Wuchiapingian, NW I an; Se ome acoce as pa i ube cula um gen. e sp. no ., Wuchiapingian, NW I an; Se ome acoce as a asense gen. e sp. no ., Changhsingian, NW I an. Cen al I an (Ko n & Hai ape ian in p ess): new species I o be desc ibed by Ko n & Hai ape ian (in p ess), Wuchiapingian. Pakis an (Waagen 1879; Reed 1931, 1944): Nau ilus la issimus Waagen, 1879, Wuchiapingian, Sal Range; Gy oce as Medlico ianum Waagen, 1879, Wuchiapingian, Sal Range; Me acoce as wa chense Reed, 1931, Wuchiapingian, Sal Range; Me acoce as chi idilense Reed, 1944, Wuchiapingian, Sal Range; Pa ame acoce as enus um Reed, 1944, Wuchiapingian, Sal Range. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 34 Sou h China (Xu 1977; Zheng 1984; Ma 1997): Me acoce as hunanense Xu, 1977, Changhsingian, Hunan; Pleu onau ilus changxingensis Zhao, Liang & Zheng, 1978, Changhsingian, Zhejiang; Pleu onau ilus zhongyingensis Zheng, 1984, Changhsingian, Guizhou; Pleu onau ilus magnus Zheng, 1984, Changhsingian, Guizhou; Pleu onau ilus an uensis Ma, 1997, Wuchiapingian, Jiangxi; Pleu onau ilus cu a us Ma, 1997, Wuchiapingian, Jiangxi; Pleu onau ilus obus us Ma, 1997, Wuchiapingian, Jiangxi. Rema ks The T anscaucasian ma e ial appea s o ep esen a mo phocline anging om o ms wi h a ela i ely simple sculp u e consis ing only o en ola e al nodes o o ms wi h ibs and a ew ows o nodes on he lank. All o hese o ms ha e p e iously been assigned o he gene a Me acoce as and Pleu onau ilus Mojsiso ics, 1882 (Kummel 1953; Shimansky 1965b; Teiche & Kummel 1973). Howe e , Shimansky (1965b: 158) al eady no ed some unce ain ies ega ding he exac delimi a ion o he wo gene a on he basis o he specimens om Dzhul a. A di ision o he mo phocline ep esen ed in he ma e ial om Jul a in o he wo gene a Me acoce as and Pleu onau ilus would ha e consequences o he phylogene ic econs uc ion o he subo de Tainoce a ina. Species p e iously assigned o Pleu onau ilus a e al eady known om Ea ly Pe mian s a a, such as he Leona d Fo ma ion o Texas. These we e placed in he genus Foo dice as Hya , 1893 by Mille & Youngquis (1949) and in Pseudo oo dice as by Ruzhence & Shimansky (1954). Kummel (1953: 12) discussed a leng h he phylogene ic ela ionships wi hin he amily Tainoce a idae (as he unde s ood i ) and concluded ha hese species belong o Pleu onau ilus, a iew ha was suppo ed by Shimansky (1967). Howe e , his would mean ha he o igin o he genus Pleu onau ilus was in he Ea ly Pe mian o e en ea lie . Assigning pa s o he T anscaucasian mo phocline o Pleu onau ilus would mean ha his genus is polyphyle ic o ha e y simila mo phologies occu in he gene a Me acoce as and Pleu onau ilus. Dzik (1984: 161) al eady assumed ha he genus Pleu onau ilus, as p e iously used by many au ho s, is a polyphyle ic axon con aining se e al e olu iona y lineages. In con as o p e ious au ho s (Kummel 1953; Ruzhence & Shimansky 1954), who p oposed a ela i ely simple e olu iona y scena io s a ing wi h he main genus Me acoce as, om which mos o he o he o he ainoce a id gene a b anched o , Dzik (1984, p. 162, ig. 62) p oposed a much mo e complex pic u e wi h a numbe o independen e olu iona y lineages. In his phylogene ic econs uc ion, Me acoce as does no play a cen al ole and is no conside ed o be ances al o gene a such as Tainoce as Hya , 1883 and Pleu onau ilus. Also, he La e Pe mian species known om T anscaucasia, which Shimansky (1965b) and Teiche & Kummel (1973) e e ed o as Me acoce as and Pleu onau ilus, we e hough o ha e e ol ed independen ly o Me acoce as. In he ollowing, we place he mo phocline wi h “Nau ilus do so a ma us” and “Pleu onau ilus e ae” comple ely in he new genus Se ome acoce as gen. no . Some ep esen a i es o hese La e Pe mian o ms ha ha e p e iously been placed in Pleu onau ilus ac ually ha e a sculp u e e y eminiscen o Pleu onau ilus, bu hey lack an annula p ocess. Such o ms a e desc ibed by Ko n & Hai ape ian (in p ess) as belonging o he new genus C o be desc ibed by Ko n & Hai ape ian (in p ess). Se ome acoce as gen. no . di e s om Me acoce as in he shape o he who l p o ile, which in Se ome acoce as is dep essed and usually apezoidal wi h weakly di e gen lanks, while Me acoce as no mally has con e ging lanks. A u he di e ence lies in he o ma ion o he sculp u e, which is composed o coa se ibs in Se ome acoce as. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 35 Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no . Figs 3C, 17; Table 9 Nau ilus do so a ma us Abich, 1878: 20, pl. 4 ig. 1. Pleu onau ilus do so-a ma us – on A habe 1900: 215. Pleu onau ilus (Pleu onau ilus) do so-a ma us – Kummel 1953: 36. Me acoce as do soa ma um – Shimansky 1965a: 41, pl. 14 ig. 5. — Teiche & Kummel 1973: 417, pl. 3 igs. 5–6. Diagnosis Species o Se ome acoce as gen. no . wi h hickly discoidal, sube olu e conch (ww/dm = 0.45–0.50; uw/dm = 0.40–0.45), weakly dep essed who l p o ile (ww/wh = 1.15–1.40) and e y high coiling a e (WER = 2.30–2.50) a a conch diame e o 50 mm. Who l p o ile weakly apezoidal wi h gen ly di e gen lanks; en e and lanks la ened. Sculp u e wi h 10–12 low en ola e al nodes pe olu ion; he nodes show a sho ex ension owa d he mid lank. Su u e line wi h a b oad and shallow ex e nal lobe and a sligh ly deepe , b oadly ounded la e al lobe. Ma e ial examined IRAN – Eas Aze baijan • 1 specimen; Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 17A–C; MB.C.32026. – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 17D–F; MB.C.32027 • 1 specimen; same da a as o p eceding; 2012; Ko n e al. leg.; illus a ed in Fig. 17G; MB.C.32028. Desc ip ion Specimen MB.C.32026 is he la ges specimen a ailable; i is a somewha co oded agmen o an in e nal mould wi h a who l heigh o 38 mm, co esponding o a conch diame e o abou 115 mm (Fig. 17A). Only one hi d o he who l is p ese ed, showing emains o he las h ee ph agmocone chambe s and a pa o he body chambe . The who l p o ile is apezoidal and sligh ly dep essed (ww/ wh = 1.19) wi h a b oadly ounded en e and a p ominen , na owly ounded en ola e al shoulde . The la ened lanks a e weakly di e gen and he umbilical ma gin is b oadly ounded (Fig. 17B). The sculp u e consis s o e y shallow ibs beginning on he inne lank and becoming coa se owa ds he ou e lank whe e hey e mina e in ans e sely elonga ed, low and blun en ola e al nodes. They become much weake in he las p ese ed pa o he body chambe . I appea s ha hese nodes do no coincide be ween he wo sides o he conch, bu a he al e na e in posi ion. The su u e line o he specimen shows a e y b oad and la ex e nal lobe, a b oadly ounded en ola e al saddle and a la , b oadly ounded la e al lobe (Fig. 17C). Specimen MB.C.32027 (Fig. 17D) is an in e nal mould agmen o a specimen wi h a conch diame e o abou 48 mm, consis ing o a qua e o a who l wi h he las ph agmocone chambe s and pa o he body chambe . I has a subquad a e, weakly dep essed who l p o ile (ww/wh = 1.17) wi h a la ly ounded en e , a na owly ounded en ola e al shoulde , sligh ly conca e and weakly di e gen lanks and a aised and hus a he p onounced, bu ounded umbilical ma gin (Fig. 17E). The en ola e al shoulde bea s sho adially elonga ed nodes ha ex end only a sho dis ance owa ds he lank, we e hey a e isible only as e y low idges. The umbilical ma gin bea s ba ely isible, e y small ube cles. The su u e line has a e y shallow ex e nal lobe and a b oadly ounded la e al lobe (Fig. 17F). The sep a a e sepa a ed by an a e age o abou 13 deg ees. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 36 Specimen MB.C.32028 is ex e nally co oded and was sec ioned o he s udy o he inne who ls; i is sligh ly de o med and was digi ally e-de o med o he illus a ion (Fig. 17G). The c oss sec ion shows some on ogene ic changes in he shape o he who ls, s a ing wi h a ci cula p o ile in he ea ly on ogene ic s age, ollowed by an in e ed apezoidal shape a a conch diame e o 11.5 mm. Hal a olu ion la e , a 19 mm diame e , he who l p o ile is ounded-hexagonal wi h a p onounced, sligh ly aised umbilical ma gin and pa allel lanks. In he cou se o hal a olu ion om a conch diame e o 30 o 45 mm, he lanks become clea ly di e gen and sligh ly conca e. A 39 mm diame e , he sec ion shows a p onounced en ola e al node ex ending abou 3 mm beyond he en ola e al shoulde . Finally, Fig. 17. Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no . om he A axoce as Beds o he Jul a Fo ma ion. A. Specimen MB.C.32026 (Ghade i 2018 Coll.) om Ali Bashi 4, la e al iew. B. The same specimen, who l p o ile. C. The same specimen, su u e line a ww = 45.0 mm, wh = 38.5 mm. D. Specimen MB.C.32027 (Ghade i 2018 Coll.) om A as Valley, la e al iew. E. The same specimen, who l p o ile. F. The same specimen, su u e line a ww = 23.5 mm, wh = 18.5 mm. G. Specimen MB.C.32028 (Ko n e al. 2012 Coll.) om A as Valley, c oss sec ion and econs uc ion o ape u al iew. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 37 he who l p o ile o he las olu ion shows a b oadly ounded umbilical ma gin and lanks ha me ge con inuously in o he umbilical wall. Only mino on ogene ic changes in he ca dinal conch pa ame e s can be de ec ed, bu i mus be aken in o accoun ha he ww/dm and ww/wh a ios can be in luenced by he posi ion o he c oss sec ion ela i e o he ibs. Whe he he sec ion hi s a node o in e space is c i ical o he shape o he who l p o ile. Howe e , i can be seen clea ly ha he uw/dm a io slowly inc eases om a alue o 0.33 a a conch diame e o 11.5 mm o 0.42 a 68 mm diame e . Rema ks Abich (1878: pl. 4 ig. 1) illus a ed a specimen wi h a diame e o 53 mm unde he name “Nau ilus do so a ma us, no . o m.”; his agmen allows a ai ly accu a e desc ip ion o he mo phology o he conch and he sculp u e (Fig. 3C). The conch is sube olu e (uw/dm ~ 0.40) wi h a weakly dep essed who l p o ile (ww/wh ~ 1.25). The en e appea s o be la ened and he lanks a e bo de ed by a ounded en ola e al shoulde and a ounded bu dis inc umbilical ma gin. The sculp u e consis s o i e la e ally di ec ed nodes on hal a who l, which a e abou hal as wide as hei in e spaces. They a e somewha elonga ed owa ds he umbilicus and ex end almos o he middle o he lank. The penul ima e who l bea s abou eigh shallow adial ible s on he lank on hal a olu ion. The su u e line ex ends wi h shallow and ounded lobes ac oss en e , lanks and umbilical wall. Shimansky (1965b: pl. 14 ig. 5) showed a mo e comple e specimen wi h a conch diame e o 72 mm; his ag ees well wi h Abich’s o iginal specimen. The close esemblance o hese wo specimens o he newly collec ed ma e ial om I an allows a ai ly good iden i ica ion o he new specimens. Se ome acoce as do soa ma um gen. e comb. no . is e y simila o S. do ashamense gen. e comb. no ., bu di e s in ha ing conside ably sho e ibs, con ined o he ou e lank. In con as o S. do ashamense, S. do soa ma um has no umbilical nodes. Ano he , albei small, di e ence lies in he wid h o he umbilicus. In S. do soa ma um, he uw/dm a io is g ea e han 0.40 and in S. do ashamense only a ound 0.37. Se ome acoce as do ashamense (Shimansky, 1965) gen. e comb. no . Fig. 18; Table 10 Me acoce as do ashamense Shimansky, 1965b: 157, pl. 14 ig. 3. Pleu onau ilus sp. inde . 1 – Teiche & Kummel 1973: 418, pl. 1 igs 3–4. Pleu onau ilus sp. – Gliwa e al. 2020: ex - ig. 17b. non Me acoce as do ashamense – Teiche & Kummel 1973: 416, pl. 4 igs 5–6. Table 9. Conch dimensions (in mm) and a ios o Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no .; 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.32026 115.0 45.4 39.0 47.0 37.0 0.39 1.16 0.41 2.17 0.05 MB.C.32027 48.2 22.8 19.5 19.4 18.0 0.47 1.17 0.40 2.55 0.08 MB.C.32028 67.9 29.8 23.6 28.8 22.7 0.44 1.26 0.42 2.26 0.03 MB.C.32028 45.2 22.6 15.6 18.4 15.3 0.50 1.45 0.41 2.29 0.01 MB.C.32028 29.8 15.3 11.2 11.0 10.8 0.51 1.37 0.37 2.45 0.04 MB.C.32028 19.1 9.4 7.7 6.4 7.4 0.49 1.23 0.34 2.66 0.04 MB.C.32028 11.7 5.9 5.0 3.8 5.0 0.51 1.18 0.33 3.06 0.00 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 38 Diagnosis Species o Se ome acoce as gen. no . wi h hickly discoidal, sube olu e conch (ww/dm = 0.45–0.50; uw/dm = 0.35–0.40), weakly dep essed who l p o ile (ww/wh = 1.20–1.30) and e y high coiling a e (WER = 2.30–2.50) a a conch diame e o 50 mm. Who l p o ile nea ly ec angula , usually wi h gen ly di e gen lanks; en e and lanks la ened. Sculp u e wi h 10–12 low en ola e al nodes pe olu ion; he nodes a e connec ed by sligh ly cu ed ibs wi h weak nodes on he umbilical ma gin. Su u e line wi h a b oad and shallow ex e nal lobe and a usually sligh ly deepe , b oadly ounded la e al lobe. Type ma e ial Holo ype AZERBAIJAN • Do asham; A axoce as Beds (ea ly Wuchiapingian); illus a ed by Shimansky (1965b: pl. 14 ig. 3); PIN 1572/246. Ma e ial examined IRAN – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2013; Ko n e al. leg.; illus a ed in Fig. 18A–C; MB.C.29347 • 1 specimen; same da a as o p eceding; 2018; Ko n e al. leg.; illus a ed in Fig. 18D–F; MB.C.32029 • 1 specimen; same da a as o p eceding; 2018; Ghade i leg.; illus a ed in Fig. 18G–I; MB.C.32030 • 1 specimen; same da a as o p eceding; 2018; Ghade i leg.; illus a ed in Fig. 18J–L; MB.C.32031 • 5 specimens; same da a as o p eceding; 2018; Ghade i leg.; MB.C.32032 o MB.C.32036 • 1 specimen; same da a as o p eceding; 2011; Ko n e al. leg.; MB.C.32037 • 2 specimens; same da a as o p eceding; Ko n e al. 2013, 2018 Coll.; MB.C.32038, MB.C.32039. – Eas Aze baijan • 2 specimens; Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32040, MB.C.32041 • 1 specimen; Ali Bashi main alley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32042 • 1 specimen; Zal; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32043. Desc ip ion Specimen MB.C.29347 is a 120-deg ee long segmen consis ing o pa o he body chambe and wo chambe s o he ph agmocone. The o al diame e can be es ima ed a app oxima ely 60 mm (Fig. 18B). The specimen has a subquad a e who l p o ile wi h a la ened en e , a ounded en ola e al shoulde , sligh ly di e gen and weakly conca e lanks and a ounded umbilical ma gin (Fig. 18A). The sculp u e consis s o coa se ibs ha de ia e sligh ly om he adial di ec ion owa ds he ape u e. They eme ge on he inne lank nea he umbilical ma gin wi h a sligh ly ube cula hickening, become mo e p ominen in he middle o he lank and, a e ape ing u he , end close o he en ola e al shoulde in a blun and elonga ed ube cle. Ribs and nodes a e asymme ically a anged and no a anged in exac co espondence on bo h sides o he conch. Specimen MB.C.32029 is a ph agmocone segmen o 120 deg ees leng h (Fig. 18E). I s maximum who l heigh is 20 mm, which co esponds o a conch diame e o abou 50 mm. The who l p o ile is weakly dep essed and subquad a e (ww/wh = 1.30) and almos wides a he ounded umbilical ma gin, om whe e he lanks e y slowly di e ge owa ds he ounded en ola e al shoulde (Fig. 18D). The en e is la ened and possesses a e y shallow conca e dep ession a he beginning o he who l segmen . The sculp u e shows i e p ominen ibs on he segmen . These ibs o igina e a he umbilical ma gin and ex end adially ac oss he lanks; in he ou e lank a ea hey s eng hen signi ican ly o o m la ge and ounded nodes. The su u e line has a a he small and shallow ex e nal lobe and a la ge ounded la e al lobe wi h asymme ic shape ( he en al side is much longe han he do sal) and a e y shallow, small lobe on he umbilical wall (Fig. 18F). KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 39 Fig. 18. Se ome acoce as do ashamense (Shimansky, 1965) gen. e comb. no . om he A axoce as Beds o he Jul a Fo ma ion. A. Specimen MB.C.29347 (Ko n e al. 2013 Coll.) om A as Valley, 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 ww = 19.8 mm, wh = 20.4 mm. D. Specimen MB.C.32029 (Ko n e al. 2018 Coll.) om A as Valley, who l p o ile. E. The same specimen; la e al and en al iews. F. The same specimen, su u e line a ww = 20.8 mm, wh = 15.8 mm. G. Specimen MB.C.32030 (Ghade i 2018 Coll.) om A as Valley, en al and la e al iews. H. The same specimen, who l p o ile. I. The same specimen, su u e line a ww = 21.5 mm, wh = 19.5 mm. J. Specimen MB.C.32031 (Ghade i 2018 Coll.) om A as Valley, la e al iew. K. The same specimen, who l p o ile. L. The same specimen; su u e line a ww = 24.2 mm, wh = 18.9 mm. Scale ba uni s = 1 mm. Table 10. Conch dimensions (in mm) and a ios o Se ome acoce as do ashamense Shimansky, 1965 gen. e comb. no .; 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.29347 60.0 24.2 21.5 19.0 20.5 0.40 1.13 0.32 2.31 0.05 MB.C.32031 57.1 26.4 22.2 21.1 20.5 0.46 1.19 0.37 2.44 0.08 MB.C.32030 54.3 27.4 21.2 19.8 19.2 0.50 1.29 0.36 2.39 0.09 MB.C.32029 48.7 23.6 18.2 18.0 17.0 0.48 1.30 0.37 2.36 0.07 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 40 The agmen a y specimen MB.C.32030 (Fig. 18G; 21 mm who l heigh ) has a simila shape as specimen MB.C.32029, bu is less dep essed (ww/wh = 1.19). The ibs a e no as coa se as in he o he specimens and he su u e line possesses a deepe ex e nal lobe (Fig. 18I). The sep a a e densely a anged a in e als a e aging abou 10 deg ees only. Specimen MB.C.32031 (Fig. 18J) is a agmen wi h 22 mm who l heigh and possesses a subquad a e who l p o ile (ww/wh = 1.19) wi h weakly di e gen lanks (Fig. 18K). I s sculp u e consis s o coa se and sha p ibs and he su u e line shows a compa a i ely na ow and deep ex e nal lobe (Fig. 18L). The sep a a e spaced a in e als a e aging abou 15 deg ees. Rema ks Se ome acoce as do ashamense gen. e comb. no . was es ablished by Shimansky (1965b) mainly on he basis o a a he small holo ype wi h a conch diame e o only 35 mm. The ype specimen s ill shows some ea ly on ogene ic ea u es, such as he dis inc ly angula umbilical ma gin and he a he coa se ibs. Al hough he new ma e ial om I an only consis s o agmen s, hei ju enile mo phology can also be seen in he inne who ls o se e al o he la ge specimens. The e o e, an assignmen o his ma e ial o S. do ashamense is ai ly ce ain. The la ge specimens om I an show weakening o he ibs and nodes. Se ome acoce as do ashamense gen. e comb. no . di e s om S. do soa ma um gen. e comb. no . in possessing a he coa se ibs on he lanks and small ube cles a ound he umbilicus. The umbilicus is somewha na owe in S. do ashamense (uw/dm = 0.35–0.40 when compa ed o 0.40–0.45 in S. do soa ma um). Ano he species wi h simila mo phology is S. e ae gen. e comb. no . bu his di e s om S. do ashamense gen. e comb. no . in he mo e clea ly de eloped umbilical nodes and he occasional p esence o an addi ional ow o nodes in he mid lank egion. A u he dis inguishing ea u e is he shape o he en e ; in S. do ashamense i is la ened and in S. e ae b oadly ounded. Se ome acoce as e ae ( on A habe , 1900) gen. e comb. no . Figs 19–20; Table 11 Pleu onau ilus Ve ae on A habe , 1900: 216, pl. 18 ig. 4. Pleu onau ilus e ae – Shimansky 1965a: 41. Pleu onau ilus (Pleu onau ilus) e ae – Kummel 1953: 36. Pleu onau ilus dzhul ensis Shimansky, 1965b: 158, pl. 15 igs 5–6. Nau ilus Pichle i – Abich 1878: 21, pl. 4 ig. 2. Pleu onau ilus sp. inde . 2 – Teiche & Kummel 1973: 418, pl. 4 igs 3–4. Diagnosis Species o Se ome acoce as gen. no . wi h hickly discoidal, sube olu e conch (ww/dm ~ 0.50; uw/dm ~ 0.35), weakly dep essed who l p o ile (ww/wh ~ 1.25) and e y high coiling a e (WER = 2.40–2.50) a a conch diame e o 50 mm. Who l p o ile nea ly ec angula , usually wi h gen ly con e gen lanks; en e b oadly con ex, lanks la ened. Sculp u e wi h abou 15 low en ola e al nodes pe olu ion; he nodes a e connec ed by sligh ly cu ed ibs wi h weak nodes on he umbilical ma gin, some imes wi h an addi ional ow o nodes on he mid lank. Su u e line nea ly s aigh on he en e and wi h a b oadly ounded la e al lobe. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 41 Type ma e ial Holo ype IRAN – Eas Aze baijan • Zal; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 23; MB.C.32052. Desc ip ion Holo ype MB.C.32052 is a a he poo ly p ese ed specimen wi h a conch diame e o 38 mm (Fig. 23A). I is hinly pachyconic (ww/dm = 0.70) and sube olu e (uw/dm = 0.34) wi h a nea ly ec angula , mode a ely dep essed who l p o ile (ww/wh = 1.73) and a e y high coiling a e (WER = 2.40). The who ls a e wides a he umbilical ma gin, om whe e he weakly con ex umbilical wall app oaches he umbilical seam. The lanks con e ge slowly owa ds he p onounced, na owly ounded en ola e al shoulde s; he en e is b oadly a ched (Fig. 23B). The main elemen o he sculp u e consis s o delica e la e al ibs ending in a se ies o small ube cles on he en ola e al shoulde . The su u e line is isible only on he lanks and he umbilical wall, bo h o which ha e e y shallow, b oadly ounded lobes. Rema ks Se ome acoce as pa i ube cula um gen. e sp. no . di e s om he o he species o he genus in he e y small and mo e nume ous en ola e al ube cles. Se ome acoce as a asense gen. e sp. no . u n:lsid:zoobank.o g:ac :05175D5D-1AC6-4527-AEF5-57CDC0E9EDC3 Fig. 24; Table 14 Pleu onau ilus sp. – Gliwa e al. 2020: ex - ig. 17d. Diagnosis Species o Se ome acoce as gen. no . wi h hinly discoidal, subin olu e conch (ww/dm ~ 0.45; uw/dm ~ 0.28), nea ly quad a e who l p o ile (ww/wh ~ 0.90) and ex emely high coiling a e (WER ~ 2.70) a a conch diame e o 80 mm. Who l p o ile la ened, pa allel lanks; en e b oadly ounded, lanks sligh ly la ened, umbilical ma gin subangula . Sculp u e wi h abou 15 sho la e al plica ions pe olu ion. E ymology Named a e he ype locali y a he A as Ri e (Wes Aze baijan, I an). Type ma e ial Holo ype IRAN – Wes Aze baijan • A as Valley; Zal Membe o he Ali Bashi Fo ma ion (9.50 m below op) (ea ly Changhsingian); 2011; Ko n e al. leg.; illus a ed in Fig. 24; MB.C.29349. Desc ip ion Holo ype MB.C.29349 is a he comple e and has a conch diame e o 82 mm (Fig. 24B). I is p ese ed wi h adhe en shell ma e ial bu does no show he su u e line. The conch is hinly discoidal and Table 13. Conch dimensions (in mm) and a ios o he holo ype o Se ome acoce as pa i ube cula um gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32052 38.2 26.7 15.4 13.0 13.6 0.70 1.73 0.34 2.41 0.12 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 48 subin olu e (ww/dm ~ 0.43; uw/dm = 0.28); he who l p o ile is sligh ly comp essed and subquad a e (ww/wh = 0.92) wi h la ened, pa allel lanks, a p onounced en ola e al shoulde and a b oadly ounded en e (Fig. 24A). The umbilical ma gin is subangula in he las hal o a olu ion wi h an obliquely o ien ed umbilical wall. In con as , he penul ima e olu ion (up o abou 30 mm dm) has a e y p onounced umbilical ma gin and an almos e ical umbilical wall. The sculp u e o he las olu ion is domina ed by weakly p o ac ed adial plica ions. They begin a he umbilical ma gin, whe e hey bea small ube cles, and gain in s eng h ac oss he lanks. They e mina e a he en ola e al shoulde whe e hey a e coa ses and bea ano he ow o ube cles. The umbilical opening allows he s udy o he sculp u e in he p eceding who l, in which he ibs a e much coa se han in he las and he umbilical ma gin is mo e p onounced and bea s a idge occupied wi h small ube cles. Rema ks Se ome acoce as a asense gen. e sp. no . di e s om S. cos ale (Shimansky, 1965) gen. e comb. no . in he shape o he who l p o ile; in S. a asense he ww/wh a io is abou 0.90 and in S. cos ale only 0.60– 0.70 (Shimansky 1965b). In addi ion, S. a asense has signi ican ly coa se adial ibs han S. cos ale. Se ome acoce as a asense gen. e sp. no . di e s om he o he species o he genus Se ome acoce as gen. no . o Jul a by he p ominen umbilical ma gin in he adul s age. Fig. 24. Se ome acoce as a asense gen. e sp. no ., holo ype MB.C.29349 (Ko n e al. 2011 Coll.) om he Zal Membe o he Ali Bashi Fo ma ion (9.50 m below op) a om A as Valley. A. Recons uc ion o ape u al iew. B. La e al iew. Scale ba uni s = 1 mm. Table 14. Conch dimensions (in mm) and a ios o he holo ype o Se ome acoce as a asense gen. e sp. no .; es ima ed alues in i alics. N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.29349 81.5 35.5 38.4 22.8 32.0 0.43 0.92 0.28 2.70 0.17 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 49 Family Rhiphaeoce a idae Ruzhence & Shimansky, 1954 Diagnosis Family o he supe amily Pleu onau iloidea wi h a small, slende i s who l. Who l p o ile weakly dep essed, ellip ical o apezoidal. Ven e b oad and weakly con ex, lanks con ex o sligh ly la ened and umbilical ma gin b oadly ounded o absen . Sculp u e wi h sho ibs on he lank. Su u e line wi h a low ex e nal saddle, some imes wi h a shallow ex e nal lobe, a e y shallow la e al and a a he deep unnel-shaped in e nal lobe; wi hou annula p ocess (a e Ruzhence & Shimansky 1954). Included gene a Rhiphaeoce as Ruzhence & Shimansky, 1954 (Pe mian); Pa a hiphaeoce as Ruzhence & Shimansky, 1954 (Pe mian); Sholakoce as Ruzhence & Shimansky, 1954 (Pe mian); Rhiphaeonau ilus Ruzhence & Shimansky, 1954 (Pe mian); Eximioce as Shchedukhin, 2022 (Pe mian); Alibashinau ilus gen. no . (Pe mian). Rema ks The amily Rhiphaeoce a idae can be dis inguished om o he Pe mian nau iloids by hei su u e line wi h i s a he deep in e nal lobe. The only excep ions a e he ep esen a i es o he amily Ak ubonau ilidae, bu hese di e in ha ing a much la ge ju enile conch. Genus Alibashinau ilus gen. no . u n:lsid:zoobank.o g:ac :BD4CEDE2-0145-467A-BD60-AD2C0216959E New genus D – Ko n 2025: 55. Type species Alibashinau ilus e us gen. e sp. no . Diagnosis Genus o he amily Rhiphaeoce a idae wi h sube olu e o e olu e conch; who l p o ile ci cula o sligh ly comp essed o dep essed wi h b oadly ounded en e and lanks. Sculp u e wi h low, ounded adial ibs on he lanks o wi hou ibs. Su u e line wi h shallow ex e nal lobe and b oadly ounded la e al lobe; in e nal lobe a he deep, wi hou annula p ocess. E ymology Combina ion o Ali Bashi and Nau ilus; because o he ype locali y o he ype species. Included species NW I an ( his pape ): Alibashinau ilus e us gen. e sp. no ., Wuchiapingian; Alibashinau ilus ambiguus gen. e sp. no ., Changhsingian. Rema ks The new genus comp ises species wi h a simple conch mo phology. These a e widely umbilica e o ms wi h a ci cula o weakly comp essed who l p o ile and a low coiling a e. This mo phology dis inguishes hem om he as majo i y o La e Ca boni e ous and Pe mian nau ilids. The simple mo phology o hese species makes i challenging o de e mine hei genus and amily. I is unclea whe he hey belong o a conse a i e e olu iona y lineage o simple o ms o i hey a e a esul o mo phological simpli ica ion om mo e complex o ms. The placemen o he new genus in he amily Rhiphaeoce a idae is based on he shape and sculp u e o he shell. Alibashinau ilus gen. no ., like Rhiphaeoce as, has a a he simple o al, ounded who l Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 50 p o ile. Howe e , he ju enile conch is no p ese ed in he Jul a ma e ial, so i is no known whe he he i s who l is small, as in he Rhiphaeoce a idae, o la ge, as in he Ak ubonau ilidae. The e o e, he assignmen o Alibashinau ilus can only be made wi h ese a ions. Alibashinau ilus e us gen. e sp. no . u n:lsid:zoobank.o g:ac :4C7D84B1-4455-4069-A54B-3033FEB384A0 Figs 25–27; Table 15 Diagnosis Species o Alibashinau ilus gen. no . wi h hinly discoidal, e olu e conch (ww/dm ~ 0.35; uw/dm ~ 0.50), equidimensional o weakly dep essed who l p o ile (ww/wh ~ 1.10–1.30) and mode a ely high coiling a e (WER ~ 1.85) a a conch diame e o 100–140 mm. Who l p o ile nea ly ci cula wi h con inuously ounded en e and lanks; who l o e lap e y small. Sculp u e wi h sho , coa se ibs on he ou e lank. Su u e line wi h wide and e y shallow ex e nal lobe, e y shallow la e al lobe and a he na ow, deep in e nal lobe. Fig. 25. Alibashinau ilus e us gen. e sp. no ., holo ype MB.C.32053 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a A as Valley; la e al and en al iews. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 51 E ymology F om he La in ‘ e us’ (adj., m.) = ‘old’; since he species has a mo phology simila o ha o much olde nau iloids. Type ma e ial Holo ype IRAN – Wes Aze baijan • A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Figs 25, 26A–B; MB.C.32053. Fig. 26. Alibashinau ilus e us gen. e sp. no . A. Holo ype MB.C.32053 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a A as Valley, econs uc ion o ape u al iew. B. The same specimen, su u e line a dm = 94.0 mm, ww = 32.5 mm, wh = 26.5 mm. C. Pa a ype MB.C.32054 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a Ali Bashi, main alley, la e al iew. D. The same specimen, who l p o ile. E. The same specimen, su u e line a ww = 35.4 mm, wh = 30.2 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 52 Pa a ypes IRAN – Eas Aze baijan • 1 specimen; Ali Bashi, main alley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 26C–E; MB.C.32054 • 1 specimen; Ali Bashi; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); illus a ed in Fig. 27; GLM#GH1006. Desc ip ion Holo ype MB.C.32053 is an incomple e in e nal mould wi h a diame e o almos 140 mm (Fig. 25). The s udy o conch shape and sculp u e is only possible o abou h ee qua e s o he las olu ion, as he inne who ls a e poo ly p ese ed due o ec ys allisa ion. The conch is hinly discoidal and e olu e (ww/dm = 0.30; uw/dm = 0.50) and he who l p o ile is nea ly ci cula and sligh ly dep essed (ww/ wh = 1.09). The o e lap zone o he who ls is e y small (Fig. 26A) and he coiling a e is mode a ely high (WER = 1.85). Fig. 27. Alibashinau ilus e us gen. e sp. no ., pa a ype GLM#GH1006 om he A axoce as Beds o he Jul a Fo ma ion om Jul a, en al and la e al iews. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 53 The sculp u e consis s o p ominen ibs, which a e coa ses in he en ola e al egion and g adually dec ease in s eng h owa ds he umbilicus. On he body chambe , hese ibs become weake , and a he g ea es diame e o he specimen, only shallow adial olds emain. Hal a olu ion has hi een ibs. The su u e line shows shallow undula ion wi h b oadly ounded lobes and saddles and a a he na ow, ounded in e nal lobe (Fig. 26B). Pa a ype MB.C.32054 is a who l segmen o nea ly hal a olu ion and comp ising o wo chambe s o he ph agmocone and a long po ion o he body chambe (Fig. 26C). I s conch diame e can be es ima ed wi h abou 132 mm. The conch is se pen iconic wi h a wide umbilicus (uw/dm ~ 0.52) and a weakly dep essed subci cula who l p o ile (ww/wh = 1.28). The coiling a e is mode a e (WER ~ 1.91) and he who l o e lap zone e y small (Fig. 26D). The sculp u e is composed o low ibs con ined o he lanks. They a e coa ses in he en ola e al a ea and become con inuously weake ac oss he lanks be o e hey disappea nea he umbilicus. The siphuncle has a subcen al posi ion on he do sal side a abou a qua e o he who l heigh. Rema ks Alibashinau ilus e us gen. e sp. no . can be easily dis inguished om A. ambiguus gen. e sp. no . by he coa se ibs on he ou e lank. O he di e ences a e he g ea e wid h o he umbilicus (uw/dm ~ 0.50 in A. e us, bu only ~ 0.37 in A. ambiguus) and he lowe coiling a e (WER ~ 1.85 in A. e us in con as o ~ 2.30 in A. ambiguus). Alibashinau ilus ambiguus gen. e sp. no . u n:lsid:zoobank.o g:ac :581650DC-6AA3-4D6E-9DA5-5409F4945D92 Fig. 28; Table 16 Diagnosis Species o Alibashinau ilus gen. no . wi h hinly discoidal, sube olu e conch (ww/dm ~ 0.35; uw/dm ~ 0.35), weakly ci cula 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 120 mm. Who l p o ile nea ly ci cula wi h con inuously ounded en e and lanks; who l o e lap e y small. O namen wi hou ibs. Su u e line wi h wide and e y shallow ex e nal lobe and e y shallow la e al lobe. E ymology F om he La in ‘ambiguus’ (adj.) = ‘ques ionable’; because o he unclea assignmen o he species. Type ma e ial Holo ype IRAN – Eas Aze baijan • Ali Bashi 4; Pa a i oli es Limes one (1.50 m below op) o he Ali Bashi Fo ma ion (la e Changhsingian); 2011; Ko n e al. leg.; illus a ed in Fig. 28; MB.C.32055. Table 15. Conch dimensions (in mm) and a ios o Alibashinau ilus e us gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32053 139.4 42.4 38.5 69.5 37.0 0.30 1.10 0.50 1.85 0.04 MB.C.32053 101.4 34.2 28.4 48.4 26.5 0.34 1.20 0.48 1.83 0.07 MB.C.32054 132.0 49.2 38.5 69.0 36.5 0.37 1.28 0.52 1.91 0.05 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 54 Desc ip ion Holo ype MB.C.32055 has a conch diame e o 120 mm and su e ed om some co osion (Fig. 28B). Howe e , i allows he s udy o he conch shape and p opo ions ac oss one olu ion. The who l p o ile is nea ly ci cula wi h a e y small o e lap upon he p eceding olu ion (Fig. 28A). The e is no sculp u e isible; he conch appea s o be smoo h. Almos he en i e specimen is chambe ed; he su u e line is almos s aigh . Despi e o co osion, i is clea ha he e a e no ibs o o he sculp u e. Rema ks Alibashinau ilus ambiguus gen. e sp. no . is dis inguished om A. e us gen. e sp. no . by he lack o ibs on he ou e lank. O he di e ences a e he na owe umbilicus (uw/dm ~ 0.37 in A. ambiguus, bu ~ 0.50 in A. e us) and he highe coiling a e (WER ~ 2.30 in A. ambiguus in con as o ~ 1.85 in A. e us). Fig. 28. Alibashinau ilus ambiguus gen. e sp. no ., holo ype MB.C.32055 (Ko n e al. 2011 Coll.) om he Pa a i oli es Limes one (1.50 m below op) o he Ali Bashi Fo ma ion a Ali Bashi 4. A. Recons uc ion o ape u al iew. B. La e al iew. Scale ba uni s = 1 mm. Table 16. Conch dimensions (in mm) and a ios o Alibashinau ilus ambiguus gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32055 120.3 43.1 42.7 44.3 41.0 0.36 1.01 0.37 2.30 0.04 MB.C.32055 79.3 30.8 31.1 26.5 – 0.39 0.99 0.33 – – KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 55 Alibashinau ilus sp. Fig. 29 Ma e ial examined IRAN – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 29; MB.C.32056. Desc ip ion Specimen MB.C.32056 is a body chambe agmen (Fig. 29A) wi h a who l heigh o 30 mm and a who l wid h o 26 mm. The who l p o ile is almos pe ec ly o al and is only in luenced by a e y small who l o e lap zone (Fig. 29B). The specimen shows qui e coa se and b oadly ounded, s aigh ibs on he lank. Rema ks The single specimen mos p obably ep esen s an own, independen species, bu i s agmen a y p ese a ion does no p o ide su icien da a o a clea cha ac e isa ion. I is hus kep in open nomencla u e. Family Foo dice a idae Ko n, 2025 Diagnosis Family o he supe amily Tainoce a oidea wi h a apezoidal who l p o ile; en ola e al shoulde ounded, lanks s ongly di e gen . Sculp u e wi h en ola e al conical nodes, some imes wi h low ibs on he lank. Su u e line wi h shallow lobes and low saddles. In e nal lobe e y shallow, wi hou annula p ocess ( om Ko n 2025). Included gene a Foo dice as Hya , 1893 (Pe mian); A axonau ilus Shimansky, 1979 (Pe mian); Ta dunau ilus gen. no . (Pe mian). Rema ks A de ailed discussion o Foo dice as and gene a wi h simila mo phology has been gi en by Ko n (2025). Fig. 29. Alibashinau ilus sp., specimen MB.C.32056 (Ghade i 2018 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a A as Valley. A. La e al iew. B. Who l p o ile. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 56 Genus Ta dunau ilus gen. no . u n:lsid:zoobank.o g:ac :A29DB483-82CF-4C67-AC41-D6FBE9C79244 New genus E – Ko n 2025: 56. Type species Ta dunau ilus nimius gen. e sp. no . 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. E ymology F om he La in ‘ a dus’ (adj., m.) = ‘slow’; because o he low coiling a e o he conch, and ‘nau ilus’ because o he ela ionship. Included species NW I an ( his pape ): Ta dunau ilus nimius gen. e sp. no ., Wuchiapingian; Ta dunau ilus mino gen. e sp. no ., Wuchiapingian. Cen al I an (Ko n & Hai ape ian in p ess): new species J o be desc ibed by Ko n & Hai ape ian (in p ess), Wuchiapingian. Rema ks The new genus can be easily dis inguished om all o he gene a in he assemblage o Jul a 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 gen. no . (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). Ta dunau ilus nimius gen. e sp. no . u n:lsid:zoobank.o g:ac :6A04A10E-B527-425A-8438-848513FDD2E5 Figs 30–31; Table 17 Diagnosis Species o Ta dunau ilus gen. no . wi h hickly discoidal, e olu e conch (ww/dm ~ 0.53; uw/dm ~ 0.47), mode a ely dep essed who l p o ile (ww/wh ~ 1.80) and mode a ely high coiling a e (WER ~ 1.95) a a conch diame e o 150 mm. Who l p o ile ounded apezoidal wi h s ongly di e gen lanks; en e b oadly ounded, a ea o lanks and umbilical wall weakly conca e. Sculp u e wi h wo ows o en ola e al nodes in he in e media e g ow h s age. Su u e line wi h e y b oad and shallow ex e nal lobe, sligh ly deepe , b oadly ounded la e al lobe and b oadly ounded in e nal lobe. E ymology F om he La in ‘nimius’ (adjec i e, m.) = ‘ e y la ge’; because o he la ge size o he conch. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 57 nodes, in some species wi h ows o nodes on he lank. Sep a simply domed; su u e line depending on he who l p o ile, usually wi h shallow lobes and low saddles ( om Ko n 2025). Included gene a Tainoce as Hya , 1883 (Ca boni e ous o Pe mian); Tainionau ilus Mojsiso ics, 1902 (Pe mian o T iassic); Ti olonau ilus Mojsiso ics, 1902 (Pe mian); Tylonau ilus P ingle & Jackson, 1928 (Ca boni e ous o ?Pe mian); Aulame acoce as Mille & Unklesbay, 1942 (Pe mian); Hexagoni es Hayasaka, 1947 (Pe mian); Hunanoce as Chao, 1954 (Pe mian); He engnau ilus Xu, 1977 (Pe mian); Cla inau ilus Zhao, Liang & Zheng, 1978 (Pe mian); Eulomacoce as Zhao, Liang & Zheng, 1978 (Pe mian); Li ome acoce as Zhao, Liang & Zheng, 1978 (Pe mian); Neo ainoce as Zhao, Liang & Zheng, 1978 (Pe mian;); Pa a ainonau ilus Zhao, Liang & Zheng, 1978 (Pe mian); Sei onau ilus Zhao, Liang & Zheng, 1978 (Pe mian); Neocla inau ilus Liang, 1984 (Pe mian); Nodonau ilus Liang, 1984 (Pe mian); Nodopleu oce as Zheng, 1984 (Pe mian); Meixianlingi es Qin, 1986 (Pe mian); Pa a ainoce as Qin, 1986 (Pe mian); Siamnau ilus Ishibashi e al., 1994 (Pe mian); Gujiaonau ilus Miao e al., 2019 (Pe mian); new genus D o be desc ibed by Ko n & Hai ape ian (in p ess) (Pe mian); Co o ainoce as gen. no . (Pe mian). Rema ks A de ailed discussion o amily Tainoce a idae has been gi en by Ko n (2025). Genus Tainoce as Hya , 1883 Type species Nau ilus quad angulus McChesney, 1860; o iginal designa ion. Diagnosis Genus o he amily Tainoce a idae wi h a subin olu e o sube olu e conch; who l p o ile mo e o less s ongly dep essed, anging om subquad a e and hexagonal o polygonal wi h a dis inc mid en al longi udinal g oo e. Umbilical ma gin usually p onounced and subangula in he in e media e s age, ounded in he adul s age. Sculp u e usually wi h wo ows o conical nodes on he en e and addi ional ows on he lank. Sep a simply domed, su u e line s ongly depending on he shape o he who l p o ile, usually wi h shallow ex e nal lobe and b oadly ounded la e al lobe. Siphuncle small wi h subcen al posi ion en ad o sep um cen e. Included Ca boni e ous species No h Ame ica (McChesney 1860; Mille e al. 1933; Mille & Unklesbay 1942; Lin z 1958; Tucke & Mapes 1978; S u geon e al. 1982): Nau ilus quad angulus McChesney, 1860, Gzhelian, Illinois; Nau ilus nodoca ina us McChesney, 1860, Gzhelian, Illinois; Tainoce as monili e Mille , Dunba & Cond a, 1933, Gzhelian, Texas; Tainoce as o unda um Mille , Dunba & Cond a, 1933, Gzhelian, Texas; Tainoce as mu ayi Mille & Unklesbay, 1942, Gzhelian, Neb aska; Me acoce as ma ylandica Lin z, 1958, Gzhelian, Ma yland; Tainoce as sexlinea um Tucke , 1976, Kasimo ian, Illinois; Tainoce as collinsi S u geon, Windle, Mapes & Hoa e, 1982, Gzhelian, Ohio. Done s Basin (De no 2024): Tainoce as luxae e na De no , 2024, Kasimo ian. Wes e n Russia (Waagen 1879): Nau ilus T au scholdi Waagen, 1879, Gzhelian, Moscow Basin. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 64 Included Pe mian species No h Ame ica (Swallow 1860; Hya 1891, 1893; Mille e al. 1933; Mille & Thomas 1936; Mille & Unklesbay 1942; Mille & Kemp 1947; Mille & Youngquis 1949): Nau ilus occiden alis Swallow, 1860, A inskian, Kansas; Tainoce as ca a um Hya , 1891, Asselian, Texas; Tainoce as Du oni Hya , 1893, Asselian, New Mexico; Tainoce as neb ascense Mille , Dunba & Cond a, 1933, A inskian, Neb aska; Tainoce as wyomingense Mille & Thomas, 1936, Asselian, Wyoming; Tainoce as schellbachi Mille & Unklesbay, 1942, Kungu ian, A izona; Tainoce as clydense Mille & Kemp, 1947, Kungu ian, Texas; Tainoce as unklesbayi Mille & Youngquis , 1949, Kungu ian, Texas. Alps and Sou he n Eu ope (Gemmella o 1889; Simić 1933; Sch é e 1974; P ino h & Posena o 2007): Pleu onau ilus Toulai Gemmella o, 1890, Wo dian, Sicily; Tainoce as zmaje a ense Simić, 1933, Wuchiapingian, Se bia; Tainoce as bükkense Sch é e , 1974, Changhsingian, Bűkk Moun ains; Tainoce as c assicos a um Sch é e , 1974, Changhsingian, Bűkk Moun ains; Tainoce as bales ense P ino h & Posena o, 2007, Changhsingian, Dolomi es; Tainoce as malsine i P ino h & Posena o, 2007, Changhsingian, Dolomi es. NW I an ( his pape ): Tainoce as admonens sp. no ., Wuchiapingian; Tainoce as la ecos a um sp. no ., Wuchiapingian; Tainoce as uni um sp. no ., Changhsingian. Cen al I an (Ko n & Hai ape ian in p ess): new species N o be desc ibed by Ko n & Hai ape ian (in p ess), Changhsingian, Cen al I an. Pakis an (Reed 1931, 1944): Tainoce as Noe lingi a . subglobosa Reed, 1931, Wuchiapingian, Sal Range; Tainoce as comp um Reed, 1944, Changhsingian, Sal Range; Tainoce as debile Reed, 1944, Changhsingian, Sal Range; Tainoce as imuense Reed, 1944, Wuchiapingian, Sal Range. Sou h China (Kayse 1883; Chao 1954; Zheng 1984): Nau ilus mingshanensis Kayse , 1883, Wuchiapingian, Jiangxi; Nau ilus o ien alis Kayse , 1883, Wuchiapingian, Jiangxi; Tainoce as changlingpuense Chao, 1954, Roadian, Hunan; Tainoce as hunanense Chao, 1954, Roadian, Hunan; Tainoce as gibbosum Zheng, 1984, Changhsingian, Guizhou; Tainoce as guizhouense Zheng, 1984, Changhsingian, Guizhou; Tainoce as la e onodosum Zheng, 1984, Changhsingian, Guizhou. Japan (Hayasaka 1957, 1962; Ehi o & A aki 1997): Tainoce as abukumense Hayasaka, 1957, Capi anian; Tainoce as ki akamiense Hayasaka, 1962, Roadian; Tainoce as ca ina um Ehi o & A aki, 1997, Capi anian. Rema ks Tainoce as is he ypical ep esen a i e o he amily, supe amily and subo de named a e i and is conside ed one o he ca dinal gene a wi hin he o de Nau ilida. Abou 40 species o Tainoce as ha e been desc ibed so a and he genus has a long s a ig aphic ange ex ending om he la es Ca boni e ous o he la es Pe mian. Howe e , only a ew e o s ha e been made o clea ly de ine he genus in i s mo phological ange. While Mille e al. (1933: 147) and Mille & Youngquis (1949: 80) ga e a de ailed cha ac e isa ion o he genus, Shimansky (1962b: 121) and Kummel (1964: K413) cha ac e ised he genus Tainoce as wi h jus one sen ence: “Like Me acoce as bu wi h a double ow o nodes on he en e .” S u geon e al. (1997: 29) we e mo e p ecise: “Simila o Me acoce as bu possessing wo en al ows o nodes o ibs sepa a ed by a median sulcus.” They ga e a mo e de ailed ou line o he cha ac e s ypically p esen in Tainoce as. I is appa en ha a simple de ini ion is no adequa e o de ine he a he complex genus, especially in espec ing he ac ha o he ainoce a id gene a ha e been es ablished in ecen decades. A p ecise KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 65 mo phological delinea ion and axonomic in e p e a ion o Tainoce as equi es a discussion o se e al ques ions: (1) Wha a e he mo phological cha ac e s ha can be used o clea ly dis inguish Tainoce as om o he gene a? (2) A e he ainoce a ids (Tainoce as and de i ed gene a o he La e Pe mian) eally a monophyle ic uni ? (3) Did long- anging e olu iona y lineages wi h s able mo phology exis wi hin Tainoce as, o did simila conch shapes and sculp u es eme ge i e a i ely and independen ly? The i s ques ion is no easy o answe . The p e iously used cha ac e o a double ow o en al nodes canno be used uni e sally as a ca dinal sepa a ing cha ac e because some species ha e only one ow o en al nodes o no en al nodes a all in he adul s age (e.g., he La e Ca boni e ous T. collinsi and T. ma ylandicum and he La e Pe mian T. bales ense). Ins ead, o he suppo ing cha ac e s, such as he p esence o he mid en al g oo e, mus be used. I should be made clea ha in Tainoce as his g oo e incu es a b oadly ounded en e . This is in con as o gene a such as Me acoce as, in which a conca e en e , i p esen a all, always occu s as a shallow dep ession o he en i e en e . The second ques ion is easie o answe . Al hough he mo phological spec um o Tainoce as is a he b oad and somewha a iable, he combina ion o se e al mo phological cha ac e s, such as he p esence o he mid en al g oo e, he ows o nodes on he en e and en ola e al shoulde and he p onounced umbilical ma gin, sugges s a monophyle ic se ies o o ms. Due o he complexi y o he mo phology, a polyphyle ic o igin o Tainoce as can be excluded. To answe he hi d ques ion, i is necessa y o e alua e he cha ac e s o conch geome y and sculp u e wi h ega d o hei a ia ion wi hin he genus Tainoce as. The ollowing cha ac e s ha e p o en o be pa icula ly a iable (wi h some ep esen a i e examples): - Gene al shape o he who l p o ile: i can ange om ec angula (T. neb ascense) o oc agonal o polygonal (T. clydense, T. admonens sp. no .); he ww/wh a io can ange om app oxima ely equidimensional (T. ca a um) o weakly dep essed (T. schellbachi, T. admonens sp. no .) and mode a ely dep essed (T. du oni). - Gene al shape o he en e : in all o he species, he en e is mo e o less ipa i e, bu he deg ee o ipa i ion a ies om weak wi h nea ly con ex en e (T. ca a um) o e y s ong wi h clea ly de ined en ola e al applana ion o ming a ec i o m en e (T. clydense, T. du oni). - A angemen and shape o he lanks: he lanks can be di e gen (T. quad angulum, T. admonens sp. no .), pa allel (T. quad angulum) o con e gen (T. du oni). They can be la ened (T. ca a um) o weakly conca e (T. du oni, T. admonens). - Shape o he umbilical ma gin and umbilical wall: while he umbilical ma gin is usually na owly ounded (T. neb ascense) o subangula (T. du oni), he umbilical wall anges om oblique (T. ca a um, T. clydense) o s eep (T. du oni) and om weakly con ex (T. neb ascense) o la ened (T. du oni, T. admonens sp. no .). - Wid h and dep h o he mid en al g oo e: he g oo e can a y om na ow (T. neb ascense, T. clydense) o wide (T. schellbachi, T. admonens sp. no .) and om shallow (T. collinsi, T. ca a um, T. admonens) o deep (T. du oni, T. wyomingense). - Fo ma ion o he en al nodes: en al sculp u al elemen s may appea as sho plica ions (T. collinsi), small ube cles (T. neb ascense, T. monili e , T. nodoca ina um, T. admonens sp. no .), o also la ge conical, blun nodes (T. clydense) o poin ed nodes o spines (T. quad angulum, T. schellbachi). Some species possess coa se en al ans e se ibs (T. unklesbayi). The en al ube cles may be a anged symme ically (T. neb ascense) o al e na ing on he igh and le sides o he mid en al g oo e (T. monili e ). Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 66 - 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 sp. no .). - Fo ma ion o 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 ically 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 admonens sp. no . u n:lsid:zoobank.o g:ac :962E20FB-58E7-40B0-B125-55893A5F1495 Fig. 34; Table 20 Tainoce as (?) sp. – Gliwa e al. 2020: ex - ig. 17c. Diagnosis Species o Tainoce as wi h hinly pachyconic, sube olu e conch (ww/dm ~ 0.65; uw/dm ~ 0.32), weakly dep essed who l p o ile (ww/wh ~ 1.45) and ex emely high coiling a e (WER ~ 2.85) a a conch diame e o 50 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 weakly conca e, umbilical ma gin na owly ounded. Sculp u e wi h abou 12 ain ibs on he lank pe olu ion, s eng hened o o m small, longi udinally elonga ed ube cles on he en ola e al shoulde . Su u e line wi h a shallow, ongue-shaped ex e nal lobe and a b oadly ounded, e y shallow la e al lobe. E ymology F om he La in ‘admonens’ ( e b in pa iciple) = ‘ eminding’; because o he simila conch geome y wi h he La e Ca boni e ous species o he genus. Type ma e ial Holo ype IRAN – Wes Aze baijan • A as Valley; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 34D–F; MB.C.29348. Pa a ype IRAN – Wes Aze baijan • 1 specimen; same da a as o holo ype; illus a ed in Fig. 34A–C; MB.C.32060. Desc ip ion Holo ype MB.C.29348 is a pa ly co oded specimen, bu ne e heless allows examina ion o he conch geome y, sculp u e and su u e line (Fig. 34D). I has a conch diame e o 48 mm and is ully chambe ed. The conch is hinly pachyconic and sube olu e (ww/dm = 0.65; uw/dm = 0.32) wi h an ex emely high coiling a e (WER = 2.80) and a e y small who l o e lap zone. The who l p o ile is weakly dep essed who l p o ile (ww/wh = 1.44) and polygonal wi h di e gen , conca e lanks; he umbilical ma gin is na owly ounded and he umbilical wall is la ened. The en ola e al shoulde is p onounced and subangula ; i delimi s he b oad en e , which is ounded- ec i o m in c oss sec ion and possesses h ee conca e zones, o which he cen al is he deepes (Fig. 34E). The sculp u e consis s o ain , longi udinally elonga ed conical ube cles wi h a posi ion on he en ola e al shoulde . The e a e abou 15 o such ube cles pe olu ion. The en e bea s one ow o e y low, blun ube cles on each side adjacen o he longi udinal g oo e. These ube cles do no co espond o he ible s on he lank and a e KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 67 mo e nume ous. The su u e line is undula ed wi h e y shallow lobes in he conca e pa s o he conch (Fig. 34F). Pa a ype MB.C.32060 is a a he s ongly co oded specimen, bu ne e heless allows examina ion o he conch geome y, sculp u e and su u e line. I has a conch diame e o 56 mm and is ully chambe ed (Fig. 34A). The conch is hinly pachyconic and sube olu e (ww/dm = 0.67; uw/dm = 0.32) wi h an ex emely high coiling a e (WER = 2.89) and a e y small who l o e lap zone. The who l p o ile is mode a ely dep essed who l p o ile (ww/wh = 1.50) is gene ally ounded polygonal wi h slowly di e gen , weakly conca e lanks; he umbilical ma gin is na owly ounded and he umbilical wall is la ened. The en ola e al shoulde is p onounced and delimi he b oad en e , which is con ex in c oss sec ion, bu possesses h ee conca e zones, o which he cen al is he deepes (Fig. 34B). The sculp u e consis s o low, longi udinally elonga ed conical ube cles wi h a posi ion on he en ola e al shoulde . The e a e abou 12 o such ibs pe olu ion. The en e bea s one ow o e y low, ba ely isible nodes on bo h sides o he longi udinal g oo e. The su u e line is undula ed wi h e y shallow lobes in he conca e pa s o he conch (Fig. 34C). Fig. 34. Tainoce as admonens sp. no . A. Pa a ype MB.C.32060 (Ko n e al. 2011 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a A as Valley, la e al and ape u al iews. B. The same specimen, econs uc ion o ape u al iew. C. Su u e line a dm = 57.5 mm, ww = 36.0 mm, wh = 22.6 mm. D. Holo ype MB.C.29348 (Ko n e al. 2011 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a A as Valley, la e al and ape u al iews. E. The same specimen, econs uc ion o ape u al iew. F. Su u e line a dm = 32.6 mm, ww = 22.8 mm, wh = 10.7 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 68 Rema ks Tainoce as admonens sp. no . has, because o i s e y weak sculp u e, a ma ginal posi ion in he mo phological spec um o he genus. Pa icula ly he weak ibs on he lank and he p esence o only one ow o en al nodes dis inguishes he new species om he o he s o he genus. The gene al conch shape and he sculp u e a e simila o he s a ig aphically olde species o he genus, such as he La e Ca boni e ous ype species T. quad angulum, bu he new species di e s in he conside ably weake de eloped en ola e al and en al ube cles. The Ea ly Pe mian T. clydense is mo e simila , especially in he non-co esponding en al and en ola e al ube cles. Howe e , he en al nodes in T. clydense a e much highe han in T. admonens sp. no . In addi ion, he umbilicus in T. clydense is na owe (uw/dm ~ 0.25) han in T. admonens (uw/dm ~ 0.30). Tainoce as la ecos a um sp. no . u n:lsid:zoobank.o g:ac :69764943-0130-453D-9EB0-B478EC4E6515 Fig. 35; Table 21 Diagnosis Species o Tainoce as wi h discoidal, sube olu e conch (ww/dm ~ 0.45; uw/dm ~ 0.33) and weakly dep essed who l p o ile (ww/wh ~ 1.15) a a conch diame e o abou 100 mm. Who l p o ile hexagonal wi h gen ly 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 20 sha p, coa se ibs on he lank pe olu ion; en e wi h one ow o hump-like, low nodes adjacen o he longi udinal g oo e. Su u e line wi h a a he deep, ongue-shaped ex e nal lobe and a b oadly ounded la e al lobe. E ymology Combina ion o he La in ‘la e’ (ad .) = ‘b oadly’ and ‘cos a um’ (adj., n.) = ‘ ibbed’; because o he coa se ibs on he lanks. Type ma e ial Holo ype IRAN – Eas Aze baijan • Ali Bashi 4; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 35; MB.C.32061. Desc ip ion Holo ype MB.C.32061 is a agmen o a ph agmocone wi h an es ima ed conch diame e o 95 mm (Fig. 35B). I is sligh ly ec onically dis o ed, bu ne e heless allows he conch geome y o be ecognised qui e well; i can be es ima ed ha he wid h o he umbilicus is abou one hi d o he conch diame e . The who l p o ile is weakly dep essed wi h a who l wid h o 45 mm and a who l heigh o 39 mm; he en e is con ex and has a median longi udinal g oo e wi h a wid h o abou 12 mm. The weakly con e gen lanks a e la ened and bo de ed by a ounded en ola e al shoulde and a ounded umbilical ma gin (Fig. 35A). Table 20. Conch dimensions (in mm) and a ios o Tainoce as admonens sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32060 56.4 37.6 25.0 17.9 23.2 0.67 1.50 0.32 2.89 0.07 MB.C.32060 33.2 23.2 15.0 9.6 – 0.70 1.55 0.29 – – MB.C.29348 48.0 31.2 21.6 15.5 19.3 0.65 1.44 0.32 2.80 0.11 MB.C.29348 27.7 19.6 11.0 8.7 – 0.71 1.78 0.31 – – KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 69 The sculp u e consis s o a combina ion o ibs and nodes. On he lank, i e e y s ong ibs a e p esen on a qua e o a olu ion; hese begin a a sho dis ance om he umbilical ma gin and ex end in sligh o wa d di ec ion wi h a shallow la e al sinus ac oss he lank. In he en ola e al egion hey inc ease in s eng h and o m sha p idges ending in en ola e al nodes. These con inue a sho dis ance owa ds he la e al side o he en e , whe e hey a e di ec ed backwa ds. Ve y low and blun ly ounded hump-like conical nodes occu immedia ely adjacen o he median longi udinal g oo e; hese nodes co espond o he ibs on he lanks. The su u e line shows a ongue-shaped ex e nal lobe and a b oadly ounded la e al lobe (Fig. 35C). Rema ks A new species is es ablished he e, al hough only one agmen a y specimen is a ailable. The eason o his is he peculia sculp u e o Tainoce as la ecos a um sp. no ., which consis s o a combina ion o ibs on he lanks and one ow o e y low, hump-like nodes on he en e . I is especially he coa se ibs ha gi e he new species a unique posi ion in he genus; o he species possess conical nodes on he lanks ins ead. In la e al iew, wi h he na owly ounded umbilical ma gin, he la ened lanks and he somewha conca e ibs, he new species bea s some esemblance o he species o he genus Se ome acoce as gen. no ., bu di e s in he en al longi udinal g oo e and he en al nodes. Fig. 35. Tainoce as la ecos a um sp. no ., holo ype MB.C.32061 (Ghade i 2018 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a Ali Bashi 4. A. Who l p o ile. B. The same specimen, la e al and en al iews. C. The same specimen, su u e line a wh = 27.0 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 70 Tainoce as uni um sp. no . u n:lsid:zoobank.o g:ac :1F60FF8F-3ABB-4B98-ABED-853626335165 Fig. 36 Diagnosis Species o Tainoce as wi h hinly pachyconic, sube olu e conch and nea ly quad a e who l p o ile (ww/wh ~ 1.10) a a conch diame e o abou 80 mm. Who l p o ile wi h weakly con e gen lanks; en e la ened wi h a b oad longi udinal mid en al g oo e, lanks sinuous, umbilical ma gin na owly ounded. Sculp u e wi h i e coa se ibs pe qua e olu ion, ex ending om he umbilical ma gin o he mid en al g oo e, s eng hened o o m small, longi udinally elonga ed nodes on he en ola e al shoulde and he en e . Su u e line wi h a e y shallow ex e nal lobe and a b oadly ounded, e y shallow la e al lobe. E ymology F om he La in ‘uni um’ ( e b in pa iciple) = ‘uni ed’; because o he amalgama ed en ola e al and en al nodes. Type ma e ial Holo ype IRAN – Wes Aze baijan • Zal; Zal Membe o he Ali Bashi Fo ma ion (ea ly Changhsingian); 2018; Ghade i leg.; illus a ed in Fig. 36; MB.C.32062. Desc ip ion The agmen a y holo ype MB.C.32062 has a who l heigh o 30 mm and hus may belong o a specimen wi h a conch diame e o abou 80 mm (Fig. 36A). I s who l p o ile is almos quad a e bu sligh ly dep essed (ww/wh = 1.06) wi h na owly ounded umbilical ma gin, la ened slowly con e ging lanks, ounded en ola e al shoulde s and a la ened en e wi h a shallow and b oad longi udinal g oo e (Fig. 36B). The sculp u e o he agmen shows i e ounded ibs on he who l segmen o 90 deg ees. These ibs a e sligh ly conca e in hei cou se; hey begin on he inne lanks and become mo e p ominen o e mina e in p onounced en ola e al nodes, which ha e a posi ion on he ou e lank. On he en e , ano he ow o nodes is p esen adjacen o he en al g oo e. The en ola e al and en al nodes a e co esponding and connec ed by a blun ib. The su u e line is weakly undula e wi h e y shallow, b oadly ounded ex e nal and la e al lobes (Fig. 36C). Rema ks Tainoce as uni um sp. no . has a simila conch shape and sculp u e o ha o T. la ecos a um sp. no ., bu di e s in he blun lank ibs (which a e sha p in T. la ecos a um) and he much hicke en al nodes, which a e e y weakly de eloped in T. la ecos a um. Ano he di e ence be ween he wo species is he ib-like connec ion o he en ola e al and en al nodes in T. uni um, whe eas hese a e sepa a e in T. la ecos a um. Tainoce as uni um sp. no . di e s om many o he species o he genus in he p esence o lank ibs. In addi ion, he almos comple e usion o he en ola e al and en al nodes is no de eloped in he o he species. Table 21. Es ima ed conch dimensions (in mm) and a ios o Tainoce as la ecos a um sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32061 95 45 39 – – 0.47 1.15 – – – KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 71 New genus D Ko n & Hai ape ian (in p ess) Type species New species O o be desc ibed by Ko n & Hai ape ian (in p ess); o iginal designa ion. 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 ( om Ko n & Hai ape ian in p ess). Included species T anscaucasia (Abich 1878): Nau ilus do so plica us Abich, 1878, Wuchiapingian, Aze baijan. Cen al I an (Ko n & Hai ape ian in p ess): new species O o be desc ibed by Ko n & Hai ape ian (in p ess), Wuchiapingian. Rema ks Acco ding o Ko n & Hai ape ian (in p ess), he new genus D is a genus wi h a mo phology ha is simila o ha o some species o Tainoce as, bu wi h a much less de eloped sculp u e. In con as o Tainoce as, which usually has 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, he new genus D has only one ow o nodes, loca ed on each side immedia ely adjacen o he median longi udinal g oo e, bu lacks ano he ow in he en ola e al egion. 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 he new genus D in he absence o en ola e al nodes and he p esence o e y coa se do sola e al nodes. Fig. 36. Tainoce as uni um sp. no ., holo ype MB.C.32062 (Ghade i 2018 Coll.) om he Zal Membe o he Ali Bashi Fo ma ion a Zal. A. Ven al and la e al iews. B. Who l p o ile. C. Su u e line a ww = 26.5 mm, wh = 26.0 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 72 New genus D do soplica um (Abich, 1878) Fig. 3A–B Nau ilus do so plica us Abich, 1878: 23, pl. 2 ig. 6, pl. 3 ig. 1, pl. 4 ig. 8. Coelonau ilus do soplica us – on A habe 1900: 217. Tainoce as do soplica um – Shimansky 1965a: 41, pl. 14, ig. 1. Rema ks Abich (1878) illus a ed wo agmen s as “Nau ilus do so plica us, no . o m.”. The i s o hese (Abich 1878: pl. 4 ig. 8) has a who l heigh o abou 27 mm. I shows ounded, s ongly con e gen lanks and a en e wi h a longi udinal mid en al g oo e abou 15 mm wide (Fig. 3B). This g oo e is bo de ed on bo h sides by a sligh ly aised ma gin ha is deco a ed wi h la ge, blun nodes. Ven ola e al nodes a e ob iously lacking. The su u e line has a ai ly na ow, ounded lobe on he en e and a b oad, ounded lobe on he lank. The second specimen (Abich 1878: pl. 2 ig. 3, pl. 3 ig. 1) has a who l wid h o abou 35 mm (Fig. 3A) and co esponds o he la ge one in conch shape and sculp u e. The specimen has a b oadly ounded umbilical ma gin. I shows ha he e a e nei he en ola e al no umbilical nodes; he lank appea s o be smoo h. New genus D do soplica um occupies a mo phologically isola ed posi ion in he amily Tainoce a idae due o he p esence o only a single ow o en al nodes. The species is p obably no ep esen ed in he new collec ions om NW I an. Genus Co o ainoce as gen. no . u n:lsid:zoobank.o g:ac :F415AD8C-A2F7-4191-809F-FDC1796B0974 New genus F – Ko n 2025: 61. Type species Co o ainoce as ine me gen. e sp. no . Diagnosis Genus o he amily Tainoce a idae wi h sube olu e conch; who l p o ile dep essed and polygonal wi h a shallow mid en al longi udinal g oo e. Umbilical ma gin b oadly ounded, lanks con e gen . Sculp u e wi hou nodes on he en e and he en ola e al shoulde , bu wi h coa se umbilical nodes. Su u e line s ongly depending on he shape o he who l p o ile, wi h shallow, ounded V-shaped ex e nal lobe and b oadly ounded la e al lobe. E ymology Combina ion o he La in ‘co ona’ (noun) = ‘w ea h’ and Tainoce as; because o he co ona e umbilicus. Included species NW I an ( his pape ): Co o ainoce as ine me gen. e sp. no ., Wuchiapingian. Rema ks The gene a o he amily Tainoce a idae a e gene ally dis inguished on he basis o hei sculp u e. The new genus di e s om all o he gene a o he amily in he absence o en al and en ola e al nodes o ube cles. In con as o mos o he membe s o he amily Tainoce a idae, he ype species o Co o ainoce as shows e y coa se umbilical nodes. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 73 Desc ip ion Specimen MB.C.32066 is a badly co oded and hus poo ly p ese ed indi idual wi h a diame e o 93 mm (Fig. 40B). I can, howe e , because o i s cha ac e is ic longi udinal nodes on he en e on bo h sides o he deep median g oo e (Fig. 40), be a ibu ed o he genus Ti olonau ilus. These nodes ex end, a leas a he beginning o he las who l, as low ibs owa ds he mid lank, whe e hey wedge ou . Rema ks The conch o m and sculp u e o he specimen om Jul a bea s some esemblance o T. sebedinus om he Dolomi es (P ino h & Posena o 2007). Howe e , he ibs on he lank a e no iceably sha pe in T. sebedinus and ex end owa ds he umbilical ma gin. Un o una ely, he poo p ese a ion o he specimen om Ali Bashi p ecludes a mo e de ailed discussion. 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 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 Fig. 40. Ti olonau ilus sp. 2, specimen MB.C.32066 (Ko n e al. 2011 Coll.) om he Pa a i oli es Limes one (2.80 m below op) o he Ali Bashi Fo ma ion a Ali Bashi 4. A. La e al iew. B. Recons uc ion o ape u al iew. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 80 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 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 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 am. no . (Pe mian). Rema ks A de ailed discussion o he 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 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 81 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 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 gen. no . (Pe mian); Pe unau ilus C ick & Sobole , 1994 (T iassic); Tomponau ilus Sobole , 1989 (T iassic). Rema ks A de ailed discussion o he 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 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 82 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. 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 ( his pape ): Li oce as choopani sp. no ., Wuchiapingian. Cen al I an (Ko n & Hai ape ian in p ess): new species P o be desc ibed by Ko n & Hai ape ian (in p ess), 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. Indopaci 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. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 83 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 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 choopani sp. no . u n:lsid:zoobank.o g:ac :47A75088-13DD-4168-B8B0-07AEAA7A93E5 Figs 41–42; Table 24 Li oce as sp. inde . – Teiche e al. 1973: 401, pl. 8 ig. 9. Li oce as sp. – Gliwa e al. 2020: ex - ig. 17e. Diagnosis Species o Li oce as wi h hinly globula , in olu e conch (ww/dm ~ 0.85–0.90; uw/dm ~ 0.05), mode a ely dep essed who l p o ile (ww/wh ~ 1.50–1.65) and high o e y high coiling a e (WER ~ 2.00–2.35) be ween a conch diame e o 75 and 135 mm. Who l p o ile wi h b oadly a ched en e and lanks, b oadly ounded umbilical ma gin and con ex umbilical wall and deep imp in zone (IZR ~ 0.40). O namen wi h coa se, con ex g ow h lines and coa se spi al lines. Su u e line nea ly s aigh . E ymology Named a e Hadi Choopan, o me I anian M Olympia, e e ing o he mos obus nau iloid ound in he Jul a assemblage. Type ma e ial Holo ype IRAN – Eas Aze baijan • Zal; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; illus a ed in Fig. 41A; MB.C.32067. Pa a ypes IRAN – Eas Aze baijan • 1 specimen; same da a as o holo ype; illus a ed in Fig. 41B–C; MB.C.32068 • 1 specimen; Ali Bashi 4; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2018; Ghade i leg.; MB.C.32069 • 1 specimen; same da a as o holo ype; 2010; Ko n e al. leg.; MB.C.32070 • 1 specimen; Ali Bashi; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); illus a ed in Fig. 42; GLM#GH1005. – Wes Aze baijan • 1 specimen; A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2013; Ko n e al. leg.; illus a ed in Fig. 41D–E; MB.C.29350. Desc ip ion Holo ype MB.C.32067 is an incomple e specimen wi h a conch diame e o app oxima ely 133 mm (Fig. 41A). The p opo ions o he conch can only be es ima ed; he specimen was appa en ly globula Fig. 41 (opposi e page). Li oce as choopani sp. no . om he A axoce as Beds o he Jul a Fo ma ion. A. Holo ype MB.C.32067 (Ghade i 2018 Coll.) om Zal, la e al and ape u al iews wi h do sal iew econs uc ion. B. Pa a ype MB.C.32068 (Ghade i 2018 Coll.) om Zal, c oss sec ion. C. The same specimen, su u e line a ww = 84.5 mm. D. Pa a ype MB.C.29350 (Ko n e al. 2013 Coll.) om A as Valley, la e al and en al iews. E. The same specimen; econs uc ion o ape u al iew. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 84 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 85 (ww/dm ~ 0.90) wi h an almos closed umbilicus and a mode a ely high coiling a e (WER ~ 2.10). The who l p o ile is semiluna e and wides a he b oadly ounded umbilical ma gin; i has a deep imp in zone (IZR ~ 0.40). Flanks and en e a e con inuously ounded o o m a hemisphe e. Shell emains show e y coa se g ow h lines, which ex end wi h a con ex a ch ac oss he lanks and o m a deep, subangula en al sinus. A second elemen o o namen a e coa se spi al lines, which a e almos o he same s eng h as he g ow h lines, oge he o ming a e icula ed o namen . Pa a ype MB.C.32068 wi h a conch diame e o 112 mm was sec ioned o s udying he conch on ogeny (Fig. 41B), bu he inne who ls a e ec ys allised o de o med. Howe e , i shows an almos isome ic on ogeny be ween 48 and 113 mm diame e . Du ing his in e al o one and a hal olu ions, he conch changes om hickly pachyconic (ww/dm = 0.82 a 48.5 mm dm) o hinly globula (ww/dm = 0.87 a 113 mm dm). The umbilicus is almos closed (uw/dm dec eases om 0.06 o 0.02). The su u e line o he specimen is almos s aigh wi h a e y shallow ex e nal lobe and an also e y shallow la e al lobe (Fig. 41C). The smalle agmen a y pa a ype MB.C.29350 wi h a conch diame e o 50 mm had o be econs uc ed o i s conch p opo ions (Fig. 41E), bu i is e iden ha i is sligh ly slende e han he wo la ge specimens (ww/dm ~ 0.78). This specimen is p ese ed wi h shell ma e ial ha shows hy hmically ein o ced g ow h lines ex ending wi h a b oadly ounded la e al p ojec ion and a deep subangula en al sinus. Rema ks Li oce as choopani sp. no . is one o he ew species in he species- ich genus wi h an almos closed umbilicus and is he e o e easy o dis inguish om many o he species, which usually ha e a uw/ dm a io o abou 0.20. Among he Middle and La e Pe mian species wi h a nea ly closed umbilicus, Fig. 42. Li oce as choopani sp. no ., pa a ype GLM#GH1005 om he A axoce as Beds o he Jul a Fo ma ion om Jul a, do sal and la e al iews. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 86 L. meishanense and L. hsuechuechiani (Chao, 1854) a e slende (ww/dm ~ 0.57 and 0.67, espec i ely), L. ga denense and L. didmyoau ise a e s ou e (ww/dm > 0.90) a compa able conch diame e s. La ge specimens o L. choopani sp. no . ha e a coa se o namen a ion consis ing o g ow h lines and spi al lines. Such an adul o namen is no known om o he species, al hough only a ew species wi h a conch diame e o o e 100 mm a e known. Genus Cele oli oce as gen. no . u n:lsid:zoobank.o g:ac :DDB55D63-9584-47C5-B8B4-BD69879FDB4F New genus G – Ko n 2025: 66. Type species Cele oli oce as cele e gen. e sp. no . Diagnosis Genus o he amily Li oce a idae wi h globula , in olu e o subin olu e conch. The conch is apidly inc easing in heigh wi h an ex ao dina ily high coiling a e (WER highe han 3.50). Who ls e y weakly emb acing wi h nea ly ci cula p o ile. Adul o namen e y weak. Sep a wi hou in lexions, sligh ly conca e. Su u e line simple, nea ly s aigh o s aigh wi h shallow lobes. E ymology Combina ion o he La in ‘cele e’ (adjec i e, n.) = ‘ as ’ and Li oce as; because o he conch geome y wi h he ex ao dina ily high coiling a e. Included species NW I an ( his pape ): Cele oli oce as cele e gen. e sp. no ., Wuchiapingian. Rema ks Cele oli oce as gen. no . is a genus ha di e s om Li oce as in i s ex ao dina ily high coiling a e o he conch. The ype species has a who l expansion a e o o e 3.60, a alue a abo e ha o mos species o Li oce as. The who l expansion a e o li oce a ids is usually a ound 2.50. The new species is he e o e placed in i s own genus. The conch o Cele oli oce as gen. no . is e y eminiscen o ha o he genus Solenochilus Meek & Wo hen, 1870, which is also cha ac e ised by an excep ionally high who l expansion a e. Howe e , he posi ion o he siphuncle, cen al in Cele oli oce as and ma ginally en al in Solenochilus, indica es ha bo h gene a belong o phylogene ically dis an e olu iona y lineages. Table 24. Conch dimensions (in mm) and a ios o Li oce as choopani sp. no .; 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.32067 133.0 118.0 71.0 7.0 41.0 0.89 1.66 0.05 2.09 0.42 MB.C.32068 113.0 98.2 66.5 2.5 38.8 0.87 1.48 0.02 2.32 0.42 MB.C.32068 74.2 66.4 44.0 3.3 25.7 0.89 1.51 0.04 2.34 0.42 MB.C.32068 48.5 39.7 26.9 2.9 – 0.82 1.48 0.06 – – MB.C.29350 49.5 38.5 28.0 1.0 17.0 0.78 1.38 0.02 2.32 0.39 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 87 Cele oli oce as cele e gen. e sp. no . u n:lsid:zoobank.o g:ac :B1A2F8FC-0603-4F60-A7BA-5E5735C44FF2 Fig. 43; Table 25 Li oce as sp. – Gliwa e al. 2020: ex - ig. 17g. Diagnosis Species o Cele oli oce as gen. no . wi h hinly globula , subin olu e o in olu e conch (ww/dm ~ 0.85–0.90; uw/dm ~ 0.15), weakly o mode a ely dep essed who l p o ile (ww/wh ~ 1.40–1.60) and ex ao dina ily high coiling a e (WER ~ 3.60) be ween a conch diame e o 30 and 60 mm. Who l p o ile wi h b oadly a ched en e and lanks, b oadly ounded umbilical ma gin and con ex umbilical wall and small imp in zone (IZR ~ 0.15). Shell su ace smoo hs. Su u e line nea ly s aigh . E ymology F om he La in ‘cele e’ (adjec i e, n.) = ‘ as ’; because o he conch geome y wi h he ex ao dina ily high coiling a e. Type ma e ial Holo ype IRAN – Wes Aze baijan • A as Valley; A axoce as Beds o he Jul a Fo ma ion (ea ly Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 43; MB.C.29352. Desc ip ion Holo ype MB.C.29352 is ully sep a e wi h a conch diame e o 57 mm (Fig. 43A). I is sligh ly de o med bu ela i ely well p ese ed, allowing he s udy o conch geome y, su u e line and, wi h limi a ions, o namen a ion. The ema kable conch geome y is mainly caused by he ex ao dina ily high coiling a e (WER = 3.60). The conch is globula and in olu e (ww/dm = 0.88; uw/dm = 0.14) and he ww/wh a ion amoun s 1.60. The who l p o ile is c escen -shaped wi h a con ex umbilical wall and a ounded umbilical ma gin om which he lanks con e ge s ongly owa ds he b oadly ounded en e (Fig. 43B). The Fig. 43. Cele oli oce as cele e gen. e sp. no ., holo ype MB.C.29352 (Ko n e al. 2011 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a A as Valley. A. La e al and ape u al iews. B. Recons uc ion o ape u al iew. C. Su u e line a dm = 57.0 mm, ww = 46.5 mm, wh = 27.2 mm. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 88 who l o e lap zone is e y small (IZR = 0.15). I appea s ha he who l wid h inc eases as e han he who l heigh on he las olu ion; he ww/wh a io inc eases om 1.43 o 1.60 du ing he las hal olu ion. The e a e some shell emains a ached o he ph agmocone; hese a e almos smoo h wi h no aces o o namen a ion. The su u e line is almos s aigh , bu has a e y la and e y wide ex e nal lobe (Fig. 43C). Rema ks Cele oli oce as cele e gen. e sp. no . is e y di e en om all o he La e Pe mian nau iloids. I is clea ly sepa able om he co-occu ing Li oce as choopani sp. no . by i s ex ao dina ily high coiling a e (WER = 3.60 in con as o 2.30 in L. choopani) and he much smalle who l o e lap a e. Cele oli oce as sp. Fig. 44; Table 26 Ma e ial examined IRAN – Wes Aze baijan • 1 specimen; A as Valley; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 44; MB.C.32071. Desc ip ion Specimen MB.C.32071 is a agmen o a ju enile specimen wi h a conch diame e o 35 mm (Fig. 44A). I shows a hickly pachyconic, in olu e conch (ww/dm = 0.82; uw/dm = 0.12) wi h a weakly dep essed o al who l p o ile (ww/wh = 1.43). The coiling a e is ex ao dina ily high (WER = 3.35). Rema ks The specimen is oo poo ly p ese ed o a mo e de ailed desc ip ion. Because o he conch geome y and he ex ao dina ily high coiling a e, i p obably belongs o Cele oli oce as gen. no . Table 25. Conch dimensions (in mm) and a ios o Cele oli oce as cele e gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.29352 57.1 50.5 31.6 7.8 27.0 0.88 1.60 0.14 3.60 0.15 MB.C.29352 32.6 28.5 19.9 5.7 – 0.87 1.43 0.17 – – Fig. 44. Cele oli oce as sp., specimen MB.C.32071 (Ko n e al. 2011 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a A as Valley. A. Ven al and la e al iews. B. Recons uc ion o ape u al iew. C. Su u e line a dm = 24.5 mm, ww = 22.0 mm, wh = 13.2 mm. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 89 uw/dm = 0.14). The who l p o ile su e ed om de o ma ion, bu he symme y o he specimen sugges s ha he en e was p obably conca e nea he e minal ape u e. On bo h sides o he specimen he e a e emnan s o la e al umbilical p ocesses, hemisphe ical in sec ion and abou 22 mm in diame e . As hey a e b oken i is impossible o de e mine hei o iginal leng h. The shell emains appea smoo h bu show aces o ine g ow h lines, which a e almos s aigh in hei cou se ac oss he lank. The sep a a e c owded a he end o he ph agmocone a a conch diame e o abou 94 mm; he su u e line has an almos s aigh cou se (Fig. 46C). The smalle specimens show ha in aspeci ic a ia ion in conch geome y is a he limi ed. Specimen MB.C.32075 may se e as a cha ac e is ic example (Fig. 47A–B). I is a agmen wi h a conch diame e o 56 mm and has a globula and subin olu e shape (ww/dm = 0.88; uw/dm = 0.19) wi h a mode a ely dep essed who l p o ile (ww/wh = 1.95) and a e y high coiling a e (WER = 2.26). Howe e , he specimen di e s om he o he s in ha i is he only indi idual o show emnan s o a coa se o namen , isible only in a small a ea as imp essions on he in e nal mould. I appea s ha he specimen has a he coa se g ow h lines, spaced abou one millime e apa , wi h a a he shallow and wide sinus ex ending ac oss he en e . The su u e line is almos s aigh (Fig. 47C). Specimens MB.C.29351 (34 mm dm; Fig. 47G) and MB.C.32077 (51 mm dm; Fig. 47F) gi e an imp ession o he on ogene ic de elopmen o he conch, in which no majo changes can be seen. The conch is hinly globula (ww/dm = 0.95 and 0.88 espec i ely) and he who l wid h is app oxima ely wice he who l heigh . Fig. 48. Pe monau ilus abichi (K uglo , 1928) om he A axoce as Beds o he Jul a Fo ma ion o Ali Bashi. A. Specimen GLM#GH1003, la e al and en al iews. B. Specimen GLM#GH1001, en al and do sal iews. Scale ba uni s = 1 mm. Table 28. Conch dimensions (in mm) and a ios o Pe monau ilus abichi (K uglo , 1928). N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32074 134.0 99.0 64.5 19.0 39.5 0.74 1.53 0.14 2.01 0.39 MB.C.32074 94.5 74.0 43.0 – – 0.78 1.72 – – – MB.C.32088 84.5 66.0 40.5 16.5 28.0 0.78 1.63 0.20 2.24 0.31 MB.C.32076 78.0 58.7 34.3 12.0 27.0 0.75 1.71 0.15 2.34 0.21 MB.C.32075 56.4 49.8 25.6 10.7 18.9 0.88 1.95 0.19 2.26 0.26 MB.C.32080 56.0 46.1 25.0 10.5 19.5 0.82 1.84 0.19 2.35 0.22 MB.C.32077 50.5 44.5 22.0 10.8 17.0 0.88 2.02 0.21 2.27 0.23 MB.C.29351 33.6 32.0 16.5 – – 0.95 1.94 – – – MB.C.32091 28.2 31.5 12.9 6.3 10.6 1.12 2.44 0.22 2.57 0.18 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 96 Specimen MB.C.32076 (78 mm dm; Fig. 47D) has a simila mo phology bu is slende e (ww/dm = 0.75). In his specimen he sep a o wo consecu i e who ls can be seen. They show a displaced siphuncle wi h a diame e o 0.15 o he ape u e heigh . A cha ac e is ic isible in nea ly all he specimens is he p esence o a mid en al longi udinal line on he in e nal mould, being p oduced by a hin in e nal g oo e in he shell. The cha ac e was al eady igu ed by Abich (1878: pl. 3 ig. 3) and desc ibed unde he species “Nau ilus conca us”. Rema ks Abich (1878) desc ibed he ollowing species, which may belong o Pe monau ilus abichi: • “Nau ilus excen icus Eichwald”. – The ully chambe ed specimen wi h a conch diame e o 30 mm desc ibed and illus a ed by Abich (1878: pl. 1 ig. 4) has an almos sphe ical shape wi h a a he wide umbilicus (uw/dm = 0.40). Flanks and en e a e b oadly ounded. The su u e line is almos s aigh bu o ms a shallow lobe on he en e . • “Nau ilus p opinquus, no . o m.”. – Abich (1878: pl. 3 ig. 6) desc ibed and illus a ed only a small, chambe ed specimen wi h a conch diame e o abou 20 mm. I shows a na owly umbilica e conch wi h a clea umbilical ma gin and appa en ly b oadly ounded lank and en e . The su u e line is almos s aigh . This specimen is p obably no sui able o cha ac e ise a species. • “Nau ilus conca us Sowe by”. – Abich (1878: pl. 3 igs 3–4) illus a ed wo agmen s wi h abou 50 mm and a conch diame e o 40 mm, espec i ely. The smalle specimen is be e p ese ed and shows he who l p o ile o an inne who l. A a diame e o 16 mm, he conch is globula (ww/ dm = 1.03) wi h a e y na ow umbilicus (uw/dm = 0.10) and a dep essed who l p o ile (ww/wh = 1.90). The who l p o ile shows a shallow do sal in lexion. Al eady on A habe (1900) sugges ed ha i is no clea whe he he h ee species desc ibed by Abich (1878), “Nau ilus excen icus”, “Nau ilus p opinquus” and “Nau ilus conca us”, belong o one single species. He synonymised hese h ee species wi h he Middle Pe mian Nau ilus co nu us Golo kinsky, 1869 om he Volga Basin o Russia. In a e ision o his occu ence, K uglo (1928) discussed “Nau ilus excen icus” o Abich and ga e i he new name “Coloce as Abichi”. K uglo (1933) la e placed his species in his new genus Pe monau ilus. Shimansky (1962c) hen used his species o ound he new subgenus Pe monau ilus (Alexand onau ilus), which should be dis inguished om he nominal genus by he p esence o hin ans e se ibs. In a la e accoun (Shimansky 1965a), howe e , he did no use his anymo e. Ba sko e al. (2014), when e ising he Middle Pe mian cephalopods o he Volga- U al egion, also did no use he subgenus. Ins ead, hey sepa a ed he genus Pe monau ilus and hei new genus Nemdoce as om he Li oce a idae, o which hey in oduced he new amily Pe monau ilidae ha is cha ac e ised by hei spiny umbilical p ojec ions. The p esence o umbilical shell p ocesses in he ma e ial om Jul a was al eady sugges ed by on A habe (1900: 212), bu he specimen he ci ed ( on A habe 1900: pl. 18 ig. 1) is p obably oo small o ha e de eloped such p ojec ions. The la ge specimen MB.C.32074 now allows us o clea ly assign he ma e ial o Pe monau ilus. Ou ma e ial di e s om he species om he Volga-U al egion mainly by he na owe umbilicus (uw/dm is abou 0.15, in con as o 0.20–0.30 in he species om he Volga). Teiche & Kummel (1973) did no lis Pe monau ilus among hei collec ion om Ali Bashi, al hough Shimansky (1965a) men ioned 28 specimens in his collec ion om he neighbou ing si es no h o he A as Ri e , meaning ha i is a commonly occu ing genus. I is mos likely ha he ma e ial a ibu ed o Li oce as sp. inde . by Teiche & Kummel (1973) in ac belongs o Pe monau ilus. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 97 Family Jul anau ilidae am. no . u n:lsid:zoobank.o g:ac :9B985B5C-9C06-4713-A6C1-763F6D4B4AA4 new amily – Ko n 2025: 65, 69, ig. 35. Type genus Jul anau ilus gen. no . Diagnosis Family o he supe amily Li oce a oidea wi h a usually pachyconic, subin olu e o in 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 usually sepa a ed by dis inc en ola e al shoulde , en e la ened o conca e. Umbilical ma gin subangula o angula ; umbilical wall s eep, la ened. O namen usually consis ing o ine g ow h lines. Sep um simple in shape, conca ely domed; su u e line wi h shallow lobes on en e and lank. E ymology Named a e he ype genus Jul anau ilus gen. no . Included gene a Jul anau ilus gen. no . (Pe mian); new genus E o be desc ibed by Ko n & Hai ape ian (in p ess) (Pe mian); new genus F o be desc ibed by Ko n & Hai ape ian (in p ess) (Pe mian). Rema ks The amily Jul anau ilidae am. no . is cha ac e ised by a combina ion o cha ac e s no ound in any o he amily o Palaeozoic nau ilids. This is he combina ion o a a he s ou conch wi h a e y p onounced umbilical ma gin and also a some imes p onounced en ola e al shoulde . While he i s cha ac e sugges s a placemen in he supe amily Li oce a oidea, he second and hi d cha ac e s show a close mo phological ela ionship o he supe amilies Pleu onau iloidea and G ypoce a oidea. Un o una ely, he ea ly on ogene ic de elopmen o he conch in he species o he Jul anau ilidae is no known. Howe e , he ma e ial shows ha he p onounced umbilical ma gin is p esen ea ly in on ogeny and ha his ea u e can he e o e be conside ed apomo phic, whe eas he en ola e al shoulde does no assume a subangula shape un il a la e on ogene ic s age, i a all. The e o e, hese o ms a e included he e as a new amily o he supe amily Li oce a oidea. Genus Jul anau ilus gen. no . u n:lsid:zoobank.o g:ac :D5ED78D7-D4E4-43D0-B9E9-193D3554B4AB new genus H – Ko n 2025: 69, ig. 35. Type species Jul anau ilus ashou ii gen. e sp. no . Diagnosis Genus o he amily Jul anau ilidae am. no . wi h hinly pachyconic, in olu e o subin olu e conch. Conch apidly inc easing in heigh wi h an ex emely o ex ao dina ily high coiling a e (WER highe han 2.50). Who ls weakly emb acing, who l p o ile weakly dep essed. Ven e weakly conca e, umbilical ma gin usually na owly 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 shallow ex e nal lobe and b oadly ounded la e al lobe. E ymology Combina ion o he name o he ype egion and Nau ilus. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 98 Included species NW I an ( his pape ): Jul anau ilus ashou ii gen. e sp. no ., Wuchiapingian; Jul anau ilus hai ape iani gen. e sp. no ., Wuchiapingian. Rema ks Jul anau ilus gen. no . has some simila i ies o Coelogas e oce as, pa icula ly in he pachyconic conch and he shape o he en e wi h he shallow median g oo e. Howe e , he specimens o Coelogas e oce as ha e a ounded umbilical ma gin, which di e s om he angula e o subangula umbilical ma gin in Jul anau ilus. Since he shape o he umbilical ma gin is e y s able in nume ous Ca boni e ous and Pe mian nau iloids, his is also assumed o Jul anau ilus. The simila i y in en e shape be ween Coelogas e oce as and Jul anau ilus is he e o e in e p e ed as a case o con e gen mo phological e olu ion. Jul anau ilus ashou ii gen. e sp. no . u n:lsid:zoobank.o g:ac :B2ABF480-DEB4-43BA-97D1-80FFF3C2B11B Fig. 49; Table 29 New genus H, new species – Ko n 2025: 70, ig. 35. Diagnosis Species o Jul anau ilus gen. no . wi h a hinly pachyconic, subin olu e conch (ww/dm ~ 0.65; uw/dm ~ 0.25), weakly dep essed who l p o ile (ww/wh ~ 1.30) 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 a weakly conca e en e , a subangula Fig. 49. Jul anau ilus ashou ii gen. e sp. no ., holo ype MB.C.32095 (Ko n e al. 2011 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a A as Valley. A. La e al and ape u al iews. B. Recons uc ion o ape u al iew. C. Su u e line a dm = 76 mm, ww = 55 mm, wh = 42 mm.Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 99 en ola e al shoulde , la ly con ex and weakly con e gen lanks, a na owly ounded 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 Ali Reza Ashou i (Mashhad), o acknowledge his suppo in he p ojec . Type ma e ial Holo ype IRAN – Wes Aze baijan • A as Valley; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2011; Ko n e al. leg.; illus a ed in Fig. 49; MB.C.32095. Desc ip ion Holo ype MB.C.32095 is a ully chambe ed conch wi h a diame e o 87 mm (Fig. 49A); sugges ing a o al diame e o app oxima ely 150 mm including he body chambe . I is hinly pachyconic (ww/ dm = 0.62) and subin olu e (uw/dm = 0.24) wi h an ex ao dina ily high coiling a e (WER = 3.10). The e is almos no who l o e lap. The who ls a e wides a he p onounced, na owly ounded umbilical ma gin; he umbilical wall is s eep and la ened. The lanks con e ge owa ds he p onounced en ola e al shoulde , which sepa a es hem om he b oad and weakly la ened en e . The mid en e possesses a e y shallow longi udinal dep ession. Shell emains a e no p ese ed. The su u e line has a shallow, b oadly ounded ex e nal lobe and a b oad and shallow la e al lobe o equal dep h (Fig. 49C). Rema ks Jul anau ilus ashou ii gen. e sp. no . di e s om J. hai ape iani gen. e sp. no . in he open umbilicus, which is closed in J. hai ape iani and in he highe coiling a e (WER ~ 3.10 in J. ashou ii bu only ~ 2.60 in J. hai ape iani). Jul anau ilus hai ape iani gen. e sp. no . u n:lsid:zoobank.o g:ac :AF3620C5-D1F3-401D-8B29-8BB929B610E2 Fig. 50; Table 30 Diagnosis Species o Jul anau ilus gen. no . wi h a hinly pachyconic, in olu e conch (ww/dm ~ 0.65; uw/dm ~ 0.05), weakly dep essed who l p o ile (ww/wh ~ 1.15) and ex emely high coiling a e (WER ~ 2.60) a a conch diame e o 80 mm. Who l p o ile wi h a weakly conca e en e , a p onounced en ola e al shoulde , la ly con ex and mode a ely con e gen lanks, a na owly ounded umbilical ma gin, a s eep and la ened umbilical wall and a mode a ely deep imp in zone (IZR ~ 0.30). Su u e line wi h b oadly ounded and shallow ex e nal lobe and b oadly ounded la e al lobe. E ymology Named a e Vachik Hai ape ian (Is ahan), o acknowledge his suppo in he esea ch p ojec on he Pe mian–T iassic bounda y. Type ma e ial Holo ype IRAN – Eas Aze baijan • Ali Bashi 4; Vedioce as Beds o he Jul a Fo ma ion (la e Wuchiapingian); 2010; Ko n e al. leg.; illus a ed in Fig. 50; MB.C.32096. Table 29. Conch dimensions (in mm) and a ios o he holo ype o Jul anau ilus ashou ii gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32095 87.4 54.4 42.2 21.0 37.8 0.62 1.29 0.24 3.10 0.10 Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 100 Desc ip ion Holo ype MB.C.32096 has a conch diame e o 83 mm and is ully chambe ed (Fig. 50A). I is hinly pachyconic (ww/dm = 0.63) wi h an almos closed umbilicus and an ex emely high coiling a e (WER = 2.58). The who l p o ile shows ha he p eceding who l is s ongly emb aced (IZR = 0.31). The conch is wides a he p onounced, na owly ounded umbilical ma gin, om whe e a s eep, b oadly ounded umbilical wall app oaches he umbilical seam. The lanks con e ge owa ds he b oadly ounded en ola e al shoulde s; he en e is b oadly ounded wi h a shallow and wide median dep ession. Shell emains a e no p ese ed. The su u e line has a shallow, b oadly ounded ex e nal lobe and a much la ge and b oadly ounded la e al lobe, which is wice as deep as he ex e nal lobe (Fig. 50C). Rema ks Jul anau ilus hai ape iani gen. e sp. no . di e s om J. ashou ii gen. e sp. no . in he closed umbilicus (uw/dm = 0.24 in J. ashou ii) and he lowe coiling a e (WER ~ 2.60 in J. hai ape iani bu ~ 3.10 in J. ashou ii). Fig. 50. Jul anau ilus hai ape iani gen. e sp. no ., holo ype MB.C.32096 (Ko n e al. 2010 Coll.) om he Vedioce as Beds o he Jul a Fo ma ion a Ali Bashi 4. A. La e al and ape u al iews. B. Recons uc ion o ape u al iew. C. Su u e line a dm = 70.5 mm, ww = 52.5 mm, wh = 44.5 mm. Scale ba uni s = 1 mm. Table 30. Conch dimensions (in mm) and a ios o he holo ype o Jul anau ilus hai ape iani gen. e sp. no . N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32096 83.2 52.2 45.2 2.5 31.4 0.63 1.15 0.03 2.58 0.31 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 101 New genus F Ko n & Hai ape ian (in p ess) Type species New species R o be desc ibed by Ko n & Hai ape ian (in p ess); o iginal designa ion. Diagnosis Genus o he amily Jul anau ilidae am. no . wi h discoidal, in olu e conch. The conch is apidly inc easing in heigh wi h a e y high o ex emely high coiling a e (WER usually highe han 2.50). Who ls mode a ely s ongly emb acing, hei p o ile anges om comp essed o weakly dep essed. 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, nea ly s aigh o s aigh wi h a low ex e nal saddle, b oadly ounded in e nal lobe. The siphuncle has a do socen al posi ion ( om Ko n & Hai ape ian in p ess). Included species Cen al I an (Ko n & Hai ape ian in p ess): new species R o be desc ibed by Ko n & Hai ape ian (in p ess), Wuchiapingian. Rema ks Ko n & Hai ape ian (in p ess) compa ed he new genus wi h o he simila gene a; in e ms o conch mo phology, he new genus shows some simila i ies o he gene a Jul anau ilus gen. no . and he new genus E o be desc ibed by Ko n & Hai ape ian (in p ess). This is pa icula ly ue o he shape o he who l p o ile wi h sligh ly la ened, con e ging lanks and he sligh ly la ened umbilical wall. Howe e , he o he wo gene a show a much highe coiling a e (WER abo e 3.00) han he new genus (WER a ound 2.50) and a much mo e p onounced subangula o angula umbilical ma gin. New genus F Ko n & Hai ape ian (in p ess) sp. Fig. 51; Table 31 Ma e ial examined IRAN – Wes Aze baijan • 1 specimen; A as Valley; Pa a i oli es Limes one o he Ali Bashi Fo ma ion (la e Changhsingian); 2011; Ko n e al. leg.; illus a ed in Fig. 51; MB.C.32097. Fig. 51. New genus F Ko n & Hai ape ian (in p ess) sp., specimen MB.C.32097 (Ko n e al. 2011 Coll.) om he Pa a i oli es Limes one o Ali Bashi. A. Recons uc ion o ape u al iew. B. La e al and en al iews. Scale ba uni s = 1 mm. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 102 Desc ip ion Specimen MB.C.32097 is he agmen o a hal who l, which belongs en i ely o he body chambe (Fig. 51A). The conch has a diame e o abou 53 mm and is discoidal and in olu e (ww/dm = 0.45; uw/ dm = 0.06). Flanks and en e s a e ounded. The su ace o he in e nal mould appea s o be comple ely smoo h. Rema ks The single specimen om he Pa a i oli es Limes one is oo poo ly p ese ed o be assigned o a speci ic species. Subo de inde . Genus and species inde . Fig. 52 Ma e ial examined IRAN – Eas Aze baijan • 1 specimen; Ali Bashi 4; lowe Jul a Fo ma ion (ea ly Wuchiapingian); 2013; Ko n e al. leg.; illus a ed in Fig. 52; MB.C.32098. Desc ip ion Specimen MB.C.32098 is he agmen o a qua e who l, which belongs en i ely o he ph agmocone (Fig. 52A). The cu a u e o he who l sugges s ha he conch was subin olu e. The who l p o ile has a heigh o 24.5 mm and a wid h o 31.6 mm and is dep essed ellip ical wi hou an umbilical ma gin o a en ola e al en e ; he who l o e laps he p eceding o a e y low deg ee (Fig. 52B). The sep a a e a anged in sho dis ances o less a 10 deg ees. The su u e line is nea ly s aigh (Fig. 52C). Rema ks The single specimen does no allow p ecise iden i ica ion due o he agmen a y p ese a ion and he small numbe o cha ac e is ic ea u es. Howe e , because o he unusual combina ion o cha ac e s, such as he weakly dep essed ellip ical who l p o ile, he a he na ow umbilicus and especially he e y dense sep a, i is b ie ly desc ibed and illus a ed he e. Table 31. Recons uc ed conch dimensions (in mm) and a ios o he new genus F Ko n & Hai ape ian (in p ess) sp. N . dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR MB.C.32097 53 24 29 323 0.45 0.83 0.06 3.12 0.21 Fig. 52. gen. inde . sp., specimen MB.C.32098 (Ko n e al. 2013 Coll.) om he A axoce as Beds o he Jul a Fo ma ion a Ali Bashi 4. A. Recons uc ion o ape u al iew. B. La e al and en al iews. C. Su u e line, a wh = 24 mm. Scale ba uni s = 1 mm. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 103 Discussion The La e Pe mian Jul a and Ali Bashi o ma ions o sec ions nea Jul a (NW I an) ha e yielded di e se nau iloid assemblages. These species belong o 20 gene a, eigh o which a e new: Fididoma oce as gen. no ., Aza inau ilus gen. no ., Se ome acoce as gen. no ., Alibashinau ilus gen. no ., Ta dunau ilus gen. no ., Co o ainoce as gen. no ., Cele oli oce as gen. no ., Jul anau ilus gen. no . Based on new ma e ial, a o al o 30 species is desc ibed, o which 24 a e new. The ma e ial comes om ou s a ig aphic uni s, in ascending o de : A axoce as Beds (ea ly Wuchiapingian) – 19 species and one in open nomencla u e: Doma oce as elegan ulum sp. no ., Doma oce as mul i ube cula um sp. no ., Doma oce as con e gens (Abich, 1878), Pe modoma oce as hamdii sp. no ., Fididoma oce as g acile (Shimansky, 1965) gen. e comb. no ., Fididoma oce as in acos a um gen. e sp. no ., Aza inau ilus nahidae gen. e sp. no ., Ai inau ilus hebes sp. no ., Se ome acoce as do soa ma um (Abich, 1878) gen. e comb. no ., Se ome acoce as do ashamense (Shimansky, 1965) gen. e comb. no ., Se ome acoce as e ae ( on A habe , 1900) gen. e comb. no ., Se ome acoce as cingulum gen. e sp. no ., Se ome acoce as in la um gen. e sp. no ., Alibashinau ilus e us gen. e sp. no ., Alibashinau ilus sp., Ta dunau ilus mino gen. e sp. no ., Co o ainoce as ine me gen. e sp. no ., Li oce as choopani sp. no ., Cele oli oce as cele e gen. e sp. no ., Pe monau ilus abichi (K uglo , 1928). Vedioce as Beds (la e Wuchiapingian) – six species: Se ome acoce as pa i ube cula um gen. e sp. no ., Tainoce as admonens sp. no ., Tainoce as la ecos a um sp. no ., Pe ipe oce as pa um sp. no ., Jul anau ilus ashou ii gen. e sp. no ., Jul anau ilus hai ape iani gen. e sp. no . Dzhul i es Beds (ea ly Changhsingian) – h ee iden i ied species and h ee in open nomencla u e: Tainoce as uni um sp. no ., Ti olonau ilus sp. 2, Se ome acoce as a asense gen. e sp. no ., new genus C o be desc ibed by Ko n & Hai ape ian (in p ess) sp., Ta dunau ilus nimius gen. e sp. no ., new genus A o be desc ibed by Ko n & Hai ape ian (in p ess) sp. Pa a i oli es Limes one (la e Changhsingian) – wo iden i ied species and wo in open nomencla u e: Alibashinau ilus ambiguus gen. e sp. no ., Tainionau ilus deinceps sp. no ., Ti olonau ilus sp. 1, Baghuknau ilus sp. Wi h 30 species, he assemblage om he a ea a ound Jul a is one o he mos di e se La e Pe mian occu ences o coiled nau iloids. Wi h 25 Wuchiapingian species alone, i is he mos species- ich assemblage o his in e al. The succession o nau iloids in he Jul a sec ions shows a signi ican decline in bo h he numbe o specimens and he numbe o species abo e he lowe A axoce as Beds. This decline is almos diame ically opposed o he abundance and di e si y o ammonoids, which a e pa icula ly common and di e se in he Pa a i oli es Limes one (Ko n e al. 2016). The decline o nau iloids is accompanied by a deepening o he habi a (Leda e al. 2014; Gliwa e al. 2020); he nau iloids in he lowe A axoce as Beds occu in associa ion wi h species- ich b achiopod assemblages (Ghade i e al. 2014). A compa ison o he nau iloids om Jul a wi h he assemblages om o he egions wi h ossil-bea ing La e Pe mian sedimen a y ocks is di icul o se e al easons. (1) A p ecise s a ig aphic co ela ion is o en di icul o impossible. (2) Facies di e ences be ween he a ious occu ences make i di icul o compa e simila posi ions in he sedimen a ion basins and hus he habi a s o he nau iloids. (3) La i udinal di e ences and hus di e en seawa e empe a u es could ha e in luenced he nau iloid associa ions. Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 104 Time-equi alen associa ions o nau iloids om cen al I an (Ko n & Hai ape ian in p ess) show simila composi ions a he amily and genus le el, bu he e a e p obably no common species om bo h egions. Al hough he wo egions a e sepa a ed by only abou 1000 kilome es and ha e e y simila sedimen a y sequences (Leda e al. 2014; Gliwa e al. 2020; Ko n e al. 2021; Heue e al. 2022), he e a e clea di e ences in he nau iloid communi ies. The genus Se ome acoce as gen. no ., which is well- ep esen ed in he Jul a sec ions, is e y a e in he cen al I anian sec ions, and he opposi e is ue o he new gene a A and C o be desc ibed by Ko n & Hai ape ian (in p ess). Acknowledgemen s We a e indeb ed o he A as F ee Zone O ice (Jul a) o hei suppo o he ield sessions. We hank Lucyna Leda (Zama e) and Nahideh Ghanizadeh Tab izi (Tab iz) o assis ance du ing he ield wo k and o con ibu ing nume ous specimens, Ma kus B inkmann (Be lin) o p epa a ion o he specimens and Jenny Huang and Jamie Lembke (Be lin) o aking pho og aphs. Fo access o he specimens in he Gol a aj Ecomuseum, we hank Eskanda Ebdali (Jul a). We acknowledge he e iews o an ea lie e sion by He wig P ino h (S Ul ich) and an anonymous e iewe . We also acknowledge he ca e ul edi ing by K is iaan Hoedemake s and Na acha Beau. Funding Die e Ko n: 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. Abbas Ghade i: Fe dowsi Uni e si y o Mashhad, g an 1/62381 and suppo by he Vice P esiden o Resea ch and Technology o he Fe dowsi Uni e si y o Mashhad. Re e ences Abich H. 1878. Geologische Fo schungen in den kaukasischen Lände n. Theil I. Eine Be gkalk auna aus de A axesenge bei Djoul a in A menien. Hölde , Wien. Agassiz L. 1847. An In oduc ion o he S udy o Na u al His o y: In a Se ies o Lec u es Deli e ed in he Hall o he College o Physicians and Su geons. G eeley & McEl a h, New Yo k. A akelyan R.A., G un T.A. & She y e A.A. 1965. K a kiy s a ig a icheskiy oche k. T udy paleon ologicheskogo Ins i u a Akademiya Nauk SSSR 108: 20–25. [In Russian.] Ba sko I.S., Leono a T.B. & Shilo sky O.P. 2014. Middle Pe mian cephalopods o he Volga-U al Region. Paleon ological Jou nal 48: 1331–1414. h ps://doi.o g/10.1134/S0031030114130012 Bisa W.S. 1930. On he gonia i e and nau iloid auna o he middle coal measu es o England and Wales. Summa y o P og ess o he Geological Su ey o G ea B i ain 1929: 75–89. B ühwile T., Buche H., Wa e D., Schneebeli-He mann E., Hochuli P.A., Roohi G., U -Rehman K. & Yaseen A. 2012. Smi hian (Ea ly T iassic) ammonoids om he Sal Range, Pakis an. Special Pape s in Palaeon ology 88: 5–114. Cane a G. 1906. La auna del Calca e a Belle ophon. Con ibu o alla conoscenza dei limi i pe mo- iassici. Bolle ino della Socie à Geologica i aliana 25: 427–452. Chao K.-K. 1940. Uppe Paleozoic cephalopods om cen al Hunan, China. Jou nal o Paleon ology 14 (1): 68–73. Chao K.-K. 1954. Pe mian cephalopods om Tanchiashan, Hunan. Ac a Palaeon ologica Sinica 2: 1–58. Chi a R., Vasle D. & Le Nind e Y. 2006. Nau iloids o he Pe mian–T iassic Khu Fo ma ion, cen al Saudi A abia. GeoA abia 11: 81–92. h ps://doi.o g/10.2113/geoa abia110181 KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 105 Teiche C. & Kummel B. 1973. Nau iloid cephalopods om he Jul a Beds, Uppe Pe mian, No hwes I an. Bulle in o he Museum o Compa a i e Zoology, Ha a d Uni e si y 144: 409–434. Teiche C., Kummel B. & Swee W.C. 1973. Pe mian–T iassic s a a, Kuh-e-Ali Bashi, No hwes e n I an. Bulle in o he Museum o Compa a i e Zoology, Ha a d Uni e si y 145 (8): 359–472. Tichy G. 1975. Übe das E s au e en on Pe monau ilus aus de Fold ik C eek Fo ma ion (Obe pe m) on Os g önland. Neues Jah buch ü Geologie und Paläon ologie, Mona she e 1975: 693–703. T enkne W. 1868. Palaeon ologische No i ä en om no dwes lichen Ha ze. I. Ibe ge Kalk und Kohlengebi ge on G und. Abhandlungen de na u o schenden Gesellscha zu Halle 10: 123–182. Tucke J.K. 1976. A coiled nau iloid auna om he Ma oon Fo ma ion (Pennsyl anian) o Illinois. T ansac ions o he Illinois S a e Academy o Science 69: 57–77. Tucke J.K. & Mapes R.H. 1978. Coiled nau iloid cephalopods om he Wol Moun ain Shale (Pennsyl anian), no h-cen al Texas. Jou nal o Paleon ology 52: 596–604. Tucke J.K., Mapes R.H. & A ono S.M. 1978. New coiled nau iloids om he Wewoka Fo ma ion (Pennsyl anian) o Oklahoma. Jou nal o Paleon ology 52: 67–72. Tu ne J.S. 1954. New Ca boni e ous nau iloids om he No h o England. T ansac ions o Leeds Geologis s Associa ion 6: 219–226. Tzwe ae M. 1888. Golo onogiya e khnego ya usa s edne usskago kamenougol’nago iz es nyaka. T udy geologicheskogo Komi e a 5: 1–58. [In Russian.] Tzwe ae M. 1898. Nau ilidy i ammonei nizhnyago o dela S edne usskago kamennougol’nago iz es nyaka. T udy geologicheskogo Komi e a 8: 1–46. [In Russian.] Unklesbay A.G. 1962. Pennsyl anian Cephalopods o Oklahoma. Bulle in o he Oklahoma Geological Su ey 96: 1–150. Vaillan -Cou u ie T ea I. 1933. Paléon ologie de Madagasca . XIX. Le Pe mo-T ias ma in. Annales de Paléon ologie 22: 39–96. on A habe G. 1900. Das jünge e Paläozoicum aus de A axes-Enge bei Djul a. Bei äge zu Paläon ologie und Geologie Ös e eich-Unga ns und des O ien s 12: 209–302. Waagen W. 1879. Sal Range ossils, 1. P oduc us Limes one ossils. Palaeon ologia Indica 1: 1–85. Wo hen A.H. & Meek F.B. 1875. Desc ip ions and illus a ions o in e eb a e ossils om he Paleozoic o ma ions. Illinois Geological Su ey 6: 69–154. Wu S. & Kuang W. 1992. S udy o nau iloids in he Uppe Pe mian Changxingian ee s om Lichuan, wes Hubei. Jou nal o China Uni e si y o Geosciences 17: 289–294. Xu G. 1977. Cephalopoda, Fossil A las o Sou h-Cen al China, Pa 537–582. Geological Publishing House, Beijing. Yako le N. 1899. Fauna neko o ykh e khnepaleozoyskikh o lozheniy Rossii. 1. Golo onogie i b akhiopody. T udy Geologicheskiy Komi e a 15: 1–150. [In Russian.] Yang Z., Yin H., Wu S., Yang F., Ding M. & Xu G. 1987. Pe mian–T iassic bounda y s a ig aphy and auna o Sou h China. Geological Memoi s 2: 1–379. Zhao J., Liang X. & Zheng Z. 1978. La e Pe mian cephalopods om Sou h China. Palaeon ologia Sinica, Se ies B 12: 1–194. [In Chinese.] Zheng Z. 1984. La e Pe mian nau iloids om wes e n Guizhou. Ac a Palaeon ologica Sinica 23: 239– 253. [In Chinese.] Eu opean Jou nal o Taxonomy 1018: 1–113 (2025) 112 P in ed e sions o all pape s a e deposi ed in he lib a ies o ou o he ins i u es ha a e membe s o he EJT conso ium: Muséum na ional dʼHis oi e na u elle, Pa is, F ance; Meise Bo anic Ga den, Belgium; Royal Museum o Cen al A ica, Te u en, Belgium; Royal Belgian Ins i u e o Na u al Sciences, B ussels, Belgium. The o he membe s o he conso ium a e: Na u al His o y Museum o Denma k, Copenhagen, Denma k; Na u alis Biodi e si y Cen e , Leiden, he Ne he lands; Museo Nacional de Ciencias Na u ales-CSIC, Mad id, Spain; Leibniz Ins i u e o he Analysis o Biodi e si y Change, Bonn – Hambu g, Ge many; Na ional Museum o he Czech Republic, P ague, Czech Republic; The S einha d Museum o Na u al His o y, Tel A i , Is aël. KORN D. & GHADERI A., La e Pe mian nau iloids om Jul a (NW I an) 113