Nautiloid Cephalopods from the Julfa Beds, Upper Permian, Northwest Iran

NAUTILOID CEPHALOPODS FROM THE JULFA BEDS, UPPER PERMIAN, NORTHWEST IRAN

CURT TEICHERT' AND BERNHARD KUMMEL"

Aii.sTRACT. Sexenteen species of niiutiloid ceph- Abich (1878), who described the following alopods, from the Upper Permian Julfa Beds of the Araks Araxes west species from ( ) Gorge northwestern Iran are descrihed and discnssed. of Soviet Dzhulfa (the modern generic as- Tlie\ belong to t\\el\e genera. Eight of the species is in : are assigned to, or compared with, pre\iously de- signment placed parentheses ) scrihed species; the remainder is left nnnamed. This Nautilus excentricus Eichwald appears to be one of the richest nantiloid faunas known from beds of very Late Permian age. {Permonautilus) Nautilus propiiKjuus Abich INTRODUCTION (Liroceras) Nautiloicls of Late Permian age are ex- Nautilus parallelus Abich tremely imeommon. In coimeetion with our {Doniatoccras) study of tlie Ali Bashi Formation'' at Kuli-e- Nautilus convergens Abich Ali Ha.slii near Julfa, northwestern Iran (Stenopoceras) (Teichert, Kimimel, and Sweet, in press; Nautilus concavus Sowerby Kummel and in Teichert, 1971, and press), (Liroceras) we were al^le to collect a fairK' large fauna Nautilus clolerus Abich of nautiloids from the underhing Julfa (Doniatoceras) Beds. This is report a description and dis- Nautilus dorsoannatus Abich cussion of this fauna. (Metacoceras) We wish to acknowledge assistance re- Nautilus picJileri v. Hauer ceived from Victoria Kohler, at Harvard (Pleuronautilus) University, and William L. Fisher, at the Nautilus incertus Abich University of Kansas. The material \\'as (Pleuronautilus) collected in 1968, during field work sup- Nautilus tubercularis Abich ported bv National Science Foundation Grant GA-996. (Pleuronautilus) Nautilus (lorsoplicatus Abich

PREVIOUS WORK ( Tainoceras)

The first record of nautiloids from the Nautilus armeniacus Abich Late Permian of the Transcaucasus was by ( Pseudotitanoceras) Nautilus cornutus Golovinski ^ University of Kansas. (Permonautilus) " Harvard University'. Oiihoceras annulatum ^ Sowerby This formation is being described by Teichert, ( ?Neocycloccras) Kummel, and Sweet (in press). It conesponds Orthoceras cribrosum Geinitz essentially to the "Transition beds" plus "Paiatiro- lites beds" et al. . of Stepanov ( 1969 ) (?)

Bull. Mus. Comp. ZooL, 144(6): 409-434, March, 1973 409 410 BiiUetin Museum of Comparative Zoology, Vol. 144, No. 6

Orthoceras transversum Abich '"Domatoceras atypicum Shimanskiy (?) ^•'^Domatoceras hunicum (Diener) Orthoceras hicinctum Abich "Domatoceras gracile Shimanskiy k-t:- (") Domatoceras parallehim (Abich) Orthoceras margaritatum Abich Domatoceras cf. gracile Shimanskiy 1 ( ?Neocyc!oceras ) ^^Domatoceras sp. Orthoceras turriteUum Abich ^Pseudotitanoceras arrneniacum (Abich) (?) Permonautihis abichi (Kruglov) Permonautilus sp. Regardless of whether or not the species *'^-?Stearoceras sp. are narrowly defined, for a late Paleozoic ^^Paranautilus pcrcgrimis (Waagen) fauna this an impressively diverse list. In a "Styrioiiaufilus sp. rcstudy of the Soviet Dzhulfian fauna Art- '^*Neocycloceras obUqueannulatum haber (in Freeh and Arthaber, 1900) intro- ( Waagen ) duced only one new species, namely, Pleuro- cf. nautihts verae Arthaber. Even the review of *^?NeocycIoceras cyclophorum ( Waagen ) the stratigraph}' and of these paleontolog)- Abich beds 1910 resulted in **?NeocycIoceras marg(iritatu)n ( ) by Stoyanow ( ) only ?Neocycloceras cf. margaritatum two new species of nautiloids—Orthoceras (Abich) lopingense Stoyanow and Nautihis himiciis lopingense (Stoyanow) Diener. In fact these are the only nautiloids "^Lopingoceras Lopingoceras cf. lopingense (Stoya- described by Stoyanow in that report. In the comprehensive monograph on now) hicinctum the Permo-Triassic formations of So\det ^^'Cycloceras (Abich) Dzhulfa, edited V. E. Ruzhentsev and by Only six of these species were com-

T. G. 1965 , the nautiloids were Sarycheva ( ) pletely described and illustrated in the text. studied Shimanski\', who the by recognized These are indicated by a single asterisk in from the Araxoceras and following species the above list. A number of species were Vedioceras beds: illustrated on plates but were not described "^Tainoceras in the text, and these are indicated in the dorsopUcatum ( Abich ) above list a double asterisk. A few '^^Metacoceras dorsoarmatum (Abich) by general comments on this impressive list of Metacoceras sp. 1 (ex. gr. dorsoarma- are in order. Pseudotemnocheilus tum Abich) species we regard as a synonym of Temnocheilus. ^^Pseudotemm:>cJ}eihis sp. The illustration by Shimanskiy (in Ruz- '^'^Foordiceras sp. 1 hentsev and 1965, ]}\. 14, fig. 7) ^^Foordiceras cf. grypoceroides (Reed) Sar)'cheva, of a specimen of Pseudotemnocheilus is very "'^Foordiceras cf. flemingiamim (Kon- poor, but it seems to be possibly conspecific inck ) with what we describe here as Temno- ""^Tirolonautihis sp. " * cheilus sp. indet. The illustrations of the ?Hexago) i ites sp . various of Foordiceras to be ^"Pleuronaiitihis incertus (Abich) species appear similar to we to Pleuro- Pleuronautihis ci. incertus (Abich) specimens assign Pleuronaiitilus verae (Arthaber) nautilus sp. indet. 2. Tirolonautilus sp. and '^^Pleuronautilus tuhercularis (Abich) Hexagonites sp. were both based on single, "Pleuronautihis dzJndfensis Shimanskiy \'ery fragmentary specimens. On the basis "Pleuroiiautihis costalis Shimanskiy of the illustrations of these specimens, we Pleuronautilus sp. do not believe that they are generically ^"?Phaedrysmocheilus sp. identifiable. The generic identification of ^-^Domatoceras convergens (Abich) Phaedrysmocheilus and Stearoceras were Upper Permian Nautiloids, Iran • Tcidieii and Kummcl 411

(juestioned by Shimanskiy, and. on the basis does not include Pernwnautilus. Faranau- of the ilkistrations he presented, we agree tiJii.s, and Styriotwutihi.s. that these identifications are doubtful. Ex- In an analysis of the nautiloid fauna of the ckiding the orthoccrids it appears that this Julfa Beds it is important to consider the nautiloid fauna of Soviet Dzhulfa includes fauna of the overlying Ali Bashi Fonnation. nine genera, which, as late Paleozoic nau- It is these strata that are of prime impor- tiloid faunas go, is a large number. tance in consideration of the Permian- Triassic boundary of this region. Shimanski}' PRESENT INVESTIGATIONS (in Ruzhentse\- and Sarycheva. 1965: 41) listed the from the The specimens described in the present following species equivalents of the Ali Bashi Formation at So\'iet paper were collected by us from that part Dzhulfa beds beds 1-4 of of the section which Stepano\- et ul. (1969) {Phisonitc.s plus the "Induan called Julfa Beds. These correspond to the stage"): Dzhulfian of Stage ( Dzhulfinskiy Yarns) Tainuceras sp. 1 (ex. gr. cluingliiigpiiciise Grunt, and Ru- Arakclyan, Shevyrev {in Chao) zhentsev and 1965: 23). .\11 col- Sarycheva. Tainocero.s sp. 2 (ex. gr. changlingpuense lections were obtained in a small side valley, Chao) to the main access \'alle\' to Kuh- parallel Tainoccra.s sp. 3 8 of and c-Ali Bashi, about km west Julfa Metacoceras sp. 2 (ex. gr. dorsoainuitinn about 200-300 m upstream from our .\li Abich)

Bashi locality 4 (Teichert, Kummel, and PFoordice ras s p . 2 Fleuronautilus dieneri v. Sweet, in press). Our collections contain sp. (ex. gr. the following genera and species: Krafft) PFIeuronautilus sp. Lopiuiioccras Iopingen.se (Stoyanow) Tainionautilus sp. cf. L. Lopingoceras sp. lopingense (Stoy- Domatoceras sp. 2 anow ) PSyringomiutihis vagus Shimanskiy Ne cloce ras cf. N. ocy sp. ohJiqueannulatum ?Faranautilus sp. (\\'aagen) PNeocycloceras sp. Tainoceras sp. indet. Onl\- Metacoceras- (lorasluimeme Shimanskiy PSyringonautilus vagus Shimanski)' was described and illustrated in Ruzhen- Metacoceras dorsoarnuitum (Abich) ( tsev and 1965: 163, 16, PlcuroiuiutiJiis sp. indet. 1 Sarycheva, pi. fig. Most of the were Fleuronautilus sp. indet. 2 9). remaining species illustrated b\' Tainiondutihis sp. indet. photographs, unaccompanied The to TeninoclicUus sp. indet. by descriptions. specimen assigned Doiiuitoccra.s Jninicmn (Diener) PSyringonautilus. to judge from the illustra- Domatoceras paralleJuni (Abich) tion, appears to be more likeK' to represent Donmtoceras sp. indet. a species of Stearoceras. Stenopoceras sp. indet. Teichert, Kummel, and Sweet (in press) Titanoccros indet. sp. describe the following species of nautiloids Pscudotitanoccra.s armeniacum (Abich) from the Ali Bashi Formation at Kuh-e-Ali Lirocera.s indet. sp. Bashi:

This fauna is essentialh" the same as that PDoIorthoceras sp. from the beds at Soviet corresponding Neocycloceras sp. Dzhulfa for the of Teni- except presence Lopingoceras sp. nocheihis, Tainionautilu.s, and Titonoceras. Tainoceras sp. indet. .\t the same time the Kuh-e-Ali Bashi fauna Metacoceras sp. indet. 412 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6

Table I DlSTRIBUTIOX OF NaUTILOID GeNERA IX JuLFA Beds axd Ali Bashi Formatiox Upper Permian Nautiloids, Iran • Teichert and Kummel 413

Figure 1. Diagram showing inferred phylogeny and geologic distribution of late Paleozoic and Triassic coiled nautiloids. (Adapted from Kummel, 1953.) Vol. No. 6 414 Bulletin Museum of Comparative Zoology, 144,

estab- MCZ 9776 4, unfigured speci- placed Lopingoceras in his newly (PI. fig. 11), with- men KU 34435. lished family, Pseudocycloceratidae, out stating reasons. Lop/ngoceras sp. cf. L. lopingense Lopingoceras lopingense (Stoyanow) (Stoyanow) Plate 4, figures 13, 14 Plate 4, figures 9, 1 1 Our collections contain at Ortlioccras lopiii^oisc Stovanow, 1910: 70, pi. 7, Description. of that figs. 2, 3. least four fragments phragmocones - Shiman- Lopingoceras lopingense (Stoyanow), are similar to those here placed in Lop- 1965: 41, skiy, in Ruzhentsev and Saiycheva, the number ingoceras lopingense, except that 42, pi. 16, fig. 10. of annulations per chamber is normally two. at Description. Our collections contain In addition the conch tends to be compressed least four fragmentary specimens that we in cross section rather than circular. On When annu- assign to Lopingoceras lopingense. the more narrowly rounded sides the 1910: he established this species, Stoyanow ( lations form narrow lobes, while the sutures cited of outline of the annu- The 70 ) angularity remain straight (Pi. 4, fig. 14). length lations as its most important distinguishing of the chambers is 2.8 to 3.0 mm, or about a formal feature, but did not really give the same as th(^ length of normal chambers it. Since the is known which description of species of L. lopingense. In one piece, and only from fragmentary phragmoconc^s consists of five camerae, or parts thereof, is living chambers, a complete description and of the basal part of the living chamber, difficult to is 2.8 give. the IcMigth of four of the chambers mm, and Obx'iously, the conch is longiconic but that of the last camera is 2.0 mm, and it The sutures are and the straight. straight bears only one ring. chambers (juite short, each chamber bearing Rcnuirks. A character which these speci- that thus runs to one annulation parall(>l mens share with typical Lopingoceras lop- the sutures. The on Plate specimen figured ingeme is the angularity in perpendicular 4, figure 11, is a fragmentarx' phragmocone section of the annulations. No similar forms of three entire and two fragmen- Dzhulfian beds consisting are as yet known from ihc chambers tary chambers. Of the three entire ucar Soviet D/hnlfa. one is Their two are short and long. lengths Repository. MCZ 9775 (Pi. 4, figs. 13, the third 34431. are 2.8, 3.0, and 5.6 mm. Thus, 14), unfigured specimen KU chamber is about t\\ icc> as long as each of Genus Flower & Caster 1935 the first two and it carries two rings, instead Neocycloceras we are here concerned of one. Apparently, Type species, Neocyc/oceras ob//quum with one of the rare cases where constmc- Flower and Caster, 1935 tion of a septum was skipped. Teichert Flower and Caster (1935: 15) stated that has Teichert et ol, 1964: K31, K52) "other {in this genus was separated from from the literature some cited previously annulated forms by the oblique sinuous kind. described cases of this sutures arid the eccentric nummuloidal Remarks. listed Stoyanow (1910: 103) siphuncle." It is further characterized by from a bed he called annu- Orthoceras lopingense "large, sinuous and slightly oblique the "Cephalopoda-zone," 11 to 14 m above the lations." Flower and Caster placed base of his Dzhulfa section. Shimanskiy genus questionably in the family Orthocera- 1965: tidae Flower 309) placed (in Ruzhentsev and Sarycheva, 42) Hyatt. (1939: in the Pseudortho- listed the species only from the Araxoceras it equally questionably in Zone, the lowest of the Dzhulfian zones. ceratidae, and maintained this position 1941: Repository. MCZ 9774 (PI. 4, fig. 9), subsequent publications (Flower, Upper Permian Nautiloids, Iran • Teichert and Kummel 415

475; 1943: 107; 1945: 685), whereas Flower cross the opposite broad side of the shell and Kummel (1950: 610) included the almost horizontalh'. Only one-half of the genus without query in that family. Both last suture of the phragmocone is preserved; in Ruzhentse\- it in direction as the annu- Shimanski)' ( and Sarycheva, slopes the same 1965) and' Sweet {in Teichert et a/., 1964) lations. placed Neocycloceras in the family Pseudor- We have three additional fragments of thoceratidae. Sweet, however, with reserva- living chambers of comparable size, but tions. of poorer preservation. As far as one can The range of the genus is stated to be judge, their features are essentially the same Upper De\'onian to Lower Carboniferous as those of the li\ing chamber described by Sweet (in Teichert et «/., 1964), and above. Devonian to Lower Carboniferous and In addition, our collections contain five Upper Permian 1)\' Shimanskiy (in Ruzhen- fragments of considerably smaller living tsev, 1962). chambers with oblique annulations, the largest of these having long and short shell Neocycloceras sp. cf. N. obliqueannulafum diameters of 10.5 and 9.5 mm respectively. (Waagen) Two of these havt> fragments of camerae Plate 4, figures 10, 12 attached to them which allow one to observe that the sutiu-es are In all cf. Orthoccras ohlique-annulatiim Waagen, 1879: oblique. speci- of the shell is or 69, TO. pi. 6, figs. 9, 10. mens the cross section more less elliptical. Whether or not these smaller Description. A number of fragmentarx specimens belong to the same species as the conch parts, ranging from about 4 mm to larger ones described above could presum- about 20 mm in conch diameter, are almost ably be decided on the basis of a larger certainl)- assignable to the genus Neocyclo- sampl(>. ceras and ma\' be related to the species The siphuncle cannot be observed in any described by Waagen as Orthoceras obUqiie- of the specimens. (iniuilatu)n. .Ml our specimens are more or Remarks. Tlie reascjn h)r (jur uncertainty less fragmentary li\'ing chambers and two of in the proper taxonomic assignment of the them have parts of camerae attached to Julfa specimens lies in the deficiencies of their bottom. Waagen's description of the type material, The is a largest specimen (Pi. 4, fig. 12) which came from the Chhidru Formation li\ing chamber that is probably complete, ("upper Productus limestone") of the Salt with one camera attached at the bottom. Range of West Pakistan. Waagen (1879: The camera is 6.5 mm and about 18 long that his few 69 ) himself admitted fragments mm wide and it seems to bear one annula- "seem barel\' sufficient for the foundation tion. However, only one-half of the outside of a species," although the oblique annu- of the camera is The preser\ed. living lations seemed to him such a characteristic chamber is 24 The has mm long. aperture feature that he did in fact establish the an elliptical outline, having diameters of species. Unfortunately, he seemed to be 19 and 14 The mm respectiveh'. living unable to observe the sutures, for he does chamber bears five annulations, the last not mention them. one situated close to the apertural margin. Miller and Unklesbay (1942) referred The annulations are at least strong, rising \^^aagen's species questionably to the genus 1 the between mm above depressions them, Brachycycloceras. However, the annulations and these are wider than the annulations. in that genus are not oblique. The annulations cross one of the broad The species has been recorded and in sides of the shell in a low arch, slope part illustrated from the Dzhulfian beds adapically across the two narrow sides, and near Dzhulfa (Arthaber, in Freeh and 416 Biilletin Museum of Comparative Zoology, Vol. 144, No. 6

Arthaber, 1900: 210; Shimanskiy, in Ruz- this fragment does not seem justified. The hentsev and Sarycheva, 1965: 41), from illustrations and descriptions of Nautilus Loping in Kiangsi, China (Freeh, 1911: dorsoplicatus Abich (1S7S: 23, pi. 2, fig. lOS), and from the Aghil" Mountains in 6; pi. 3, fig. 1; pi. 4, fig. 8) are most Tibet (Renz. 1940: 67). unsatisfactory. The specimen illustrated by Repositon/. KU 34411 (PL 4, fig. 10), Abich on his plate 4, figure 8 does show MCZ 9777 (PI. 4, fig. 12), unfigured"^ speci- marked similarities to that in our collec- mens MCZ 9778, KU 34412. tion. Even so it seems unwise to assign a specific name to our specimen. Shimanskiy Order NAUTILIDA Agassiz, 1847 also listed four specimens assigned to three Superfamily TAINOCERATACEAE indeterminate spcx-ies of Tainoceras from Hyatt, 1883 the Ali Bashi Formation of Soviet Dzhulfa. Family TAINOCERATIDAE Hyatt, 1883 Two of these fonns were illustrated but not Genus 7a;noceras Hyatt, 1883 described. The illustrations clearly show tlK> specimens to be species of Tainoceras, Type species. Nautilus quadrangulus but the incompleteness of the specimens McChesney, 1860 and their poor preservation make specific Tainoceras sp. indet. identification impossible. Our collections Plate 4, figures 1, 2 from the Ali Bashi Fonuation at Kuh-e-Ali Bashi ecmtain one fragmentary specimen Discussion. This record is based on a of Tainoceras. fairly \vell-preser\'ed phragmocone 74 mm In a re\iew of the genus Tainoceras, in diameter. The widtli of the most adoral Kummel (1953: 24) recorded 23 species camera is 51.5 mm and the heiglit 36 mm. assignable to the genus. It has a range from The concli is moderately invohite with an Middle Pennsylvania!! to Late Permian in oval whorl section. The \enter is broadlv age and one species, Trematodiscus klip- rounded with a prominent median furrow. sfcini Mojsisovics (1882) from the Karnian The ventrolateral shoulders are broadK' of the Alps, is also included in this genus. rounded, merging imperceptibh into con- Repository. MCZ 9752 (Pi. 4, figs. 1, 2). vex lateral areas. The umbilical shoulders are more abruptly rounded and the umbil- Genus Mefacoceras Hyatt, 1 883 ical wall steep; the umbilicus has a diam- Nautilus (Discus) eter of approximately 22 mm. The orna- Type species. sangamon- ment consists of a row of nodes on the ensis Meek and Worthen, 1861 venter each side of the ventral adjoining A^etococeras dorashamense Shimanskiy furrow and another row of nodes on the Plate 4, figures 5, 6 ventrolateral shoulder. The nodes are gener- Metacoceras dorashamense Shimanskiy, in Ru- ally located on the adoral part of each zhentsev and Sarycheva, 1965: 157, pi. 14, fig. of a camera. The suture consists deep 3. ventral lobe in the ventral furrow and a Discussion. Our collections contain nine shallo\\' lateral lobe on the flanks. of The in specimens mainly poor preservation. Shimanskiy ( Ruzhentsev and Sary- only complete specimen (Pi. 4, figs. 5, 6) cheva, 1965: 41) lists Tainoceras dorsopli- is a phragmocone measuring 39 mm in catum (Abich) from the Araxoceras beds diameter, 23 mm for the width of the adoral of Soviet Dzhulfa. His table indicates that whorl, and 17 mm for the height. Shiman- he had three but no formal specimens, skiy distinguished his species primarily on one description was given and only speci- the presence of nodes on both the \'entro- men, a small fragment of phragmocone, lateral and umbilical shoulders, the latter was illustrated. Specific identification of being very small, and on ribs between the Upper Permian Nautiloids, Iran • Teichert and Kinnmel 417

nodes increasing in prominence to\\ arc! the Repository. MCZ 9753 (Pi. 4, figs. 5, 6), \entrolateral node. The umbiHcal nodes are unfigured specimens MCZ 9754. not apparent on the one illustration Shiman- in Metacoceras dorsoarmatum (Abich) skiy ( Ruzhentse\' and Saryche\a, 1965, Plate pi. 14, fig. 3b) published. The ventrolateral 3, figures 5, 6 nodes are and the so- very conspicuous Nautilus doisoaniiatus Abich, 1878: 20, pi. 4, lateral these called ribs are extensions from figs. 1, la. - nodes. This is the general nature of the Metacoceras dorsoarmatum. Shimanski\, in Ru- zhentsev and 1965: 5. ornament pattern in our specimens. Saryche\ a, 41, pi. 14, fig. In a review of the Mefaco- general genus Discussion. This species differs from Kummel noted that: "The cc'ras\ (1953) Metacoceras dorashamense in haxing a more of variation and icw or high degree poorly sub(iuadrate whorl section rather than a a preserved specimens pre\'ent thorough depressed whorl section. Likewise there understanding of most of the species. There are no nodes on the umbilical shoulder. are more than twice as of many species The prominent ventrolateral nodes taper off Metacoceras as of other (50) any Pennsyl- up the lateral areas, giving the appearance vanian and Permian nautiloid." Shimanskiy of incipient ribs. The largest specimen in had two of his new only speciiuens species our collection (PI. 3. figs. 5, 6) is a phrag- and the illustiated is the type surely phragmocone 37.5 miu in diameter; the most niocone of a juvenile. Our specimens are of adoral whorl has a width of 17.7 mm and limited in the understand- help broadening a height of 16 mm. The remaining six of this because of their ing species frag- specimens are fragments of phragmocone. nientar\ nature and generally poor preser- General comments on the genus Metaco- vation. As there are no data on the range ceras and the species from the Julfa Beds of xariation in ornament for of any species are offered in the preceding section. Metacoceras, direct of species comparison Repositonj. MCZ 9755 (PI. 3, figs. 5, 6), is The onl\' of the meaningless. significance unfigured specimens MCZ 9756. large number of species assigned to this is an indication of the genus widespread Genus Pleuronautilus Mojsisovics, 1882 distribution and general abundance of Type species, Pleuronautilus trinodosus Metacoceras in marine seaways of Pennsyl- Mojsisovics, 1882 vanian and Pennian age. In addition to Metacoceras dorashamense, Di.scussion. This is another of the most Shimanski\"s fauna also includes Metaco- common genera of nautiloids in the Julfa ceras dorsoarmatum (Abich). The latter Beds. The relatixe abundance of specimens species was said to differ in having onI\" and the fact that they are ornamented have \ entrolateral nodes. Our collections also led to a proliferation of species names, as contain specimens assignable to M. dor- is generalK- the case. Abich (1(S78) de- soarmatum. but the really significant dif- scribed three species that can readily be ference from -A/, dorashamense is in the assigned to this genus, e. g., Nautdus tuher- more or less quadrate whorl section. Our cularis Abich, A', pichleri Hauer, and N. collections from the Ali Bashi in ox'erlying incertus Abich. Shimanskiy ( Ruzhentsev Formation contain one fragment of a phrag- and Sarycheva, 1965: 41) recognized five mocone, which, on the basis of its depressed species from the Dzhulfian beds of Soviet whorl section, appears to be a representa- Dzhulfa. e.g., Fleuronautihis incertus ti\e of M. dorashamense. In Soviet Dzluilfa (Abich), P. verae (Arthaber), P. tuhercu- Shimanski)' listed from the beds with PJiiso- laris (Abich), P. dzJudfensis Shimanskiy, nites one specimen attributed to Metaco- and P. co.stalis Shimanskiy. In addition, ceras sp. 2 (ex. gr. dorsoarmatum Abich). Shimanskiv listed three distinct intermediate 418 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6

species of Pleuronautihis. The differences Among the various species of this genus these between species are stated to be recognized by Shimanskiy from the Julfa primarily in ornament pattern. However, Beds, our specimen most resembles Pleuro- it needs to be emphasized that all are based nautihis costaUs Shimanskiy. The resem- on relatively few and generally fragmentar\' blance is largcK' confined to the general specimens. The significance of the differ- shape and proportions of the whorl section. ences in ornament pattern is completely Our specimen lacks the additional rows of nnknown. Knmmel (1953), in a review of nodes on the ribs which are said to charac- the genus Pleuronautihis, recorded 24 Per- terize P. costalis. However, as mentioned mian species and 34 Triassic species. Most above, the significance of these differences of these are likewise based on \'ery few in ornament is not known. Our specimen specimens. Because most species of pleuro- is also very similar to P. verae Arthaber nautilids have been narrowly conceived, (m Freeh and Arthaber, 1900, pi. IS, fig.4). especially those from the Julfa Beds, we All other pleuronautilids from the Julfa Beds conclude it is inadvisable to assign specific display a more rapid growth or inflation of names to our specimens. the whorls, and, at the diameter of this Within our collection we have two distinct specimen, already ha\e robust, depressed forms: the first, consisting of onlv one whorl sections. In addition, the ornament specimen, has a more compressed whorl pattern is highly variable. section and the whorls increase in dimen- Repository. KU 34384. sions slowly; the second form includes 15 Pleuronaufilus 2 specimens and is characterized by a robust, sp. indet. Plate depressed whorl section. 4, figures 3, 4, 7, 8 Our collections from the Ali overlying Discussion. The remaining specimens of Bashi Formation contain two fragmentary pleuronautilids in our collection are grouped which resemble the specimens specimens under this heading. Most specimens are described here as Pleuronautihis sp. indet. fragments, but two are fairly complete. The 2, and they are possibly conspecific. most characteristic feature is the depressed whorl section. The lateral areas bear ribs, Pleuronaufilus sp. indet. 1 and in this feature there is much variation. Plate 1, figures 3, 4 The large specimen of Plate 4, figures 3, 4 Discussion. Among the 16 specimens of has ribs that originate on a low node on the pleuronautilids in our collection one is ventrolateral shoulder, decrease in strength, distinctly different from all the others. This and completely disappear across the mid- specimen is all phragmocone, 59 mm in area of the flanks. In other specimens the diameter, 17 mm in width of adoral whorl, ribs extend fully to the umbilical shoulder. and 19.2 mm in height. The umbilicus is There is likewise much variation in general

25 mm in diameter. Both the venter and the prominence of the ribs, that is, in some lateral areas are slightlv convex. Tlie ventro- specimens they are very weakly expressed lateral shoulders are sharply rounded, the and in others thev are verv noticeable. umbilical shoulders more broadly so, and The specimen of Plate 4, figures 3, 4 the umbilical wall slopes to the umbilical has the following dimensions: diameter 87 seam at an angle of about 45 degrees. The mm, height of adoral whorl section 31.5 mm, suture has a shallow ventral lobe and a width of adoral whorl section 40 mm, dia- broad lateral lobe occupying the whole meter of umbilicus 28.5 mm. These pro- flank. The lateral areas bear prominent portions are typical of all the remaining radial ribs that extend from the umbilical fragments included here. shoulder to the ventrolateral shoulder where It is possible to match one or more of our they terminate in a node. fragments with Pleuronautilus tubercularis Upper Permian Nautiloids, Iran • Teiclicii and Kummel 419

(Abich), P. inccrtus (Abich), or P. dzhiil- nautilus trachyceras Freeh from the Mian- fensis Shimanskiy. Howe\'er, as mentioned wali Formation of the Salt Range also has above, all these species are very incom- been included in tliis genus. Our specimen pletely known because they are based on is most similar to the type species, T. transi- few and generally poorly preser\'ed speci- torius, but whether it is conspecific cannot mens. Study of our suite of mainly frag- be determined because of its fragmentary mentary specimens leads us to conclude nature. tliat this is nothing more than a single, Shimanski)' ( in Ruzhentsev and Sary- somewhat \^ariable species. It seems doubt- cheva, 1965: 15S. pi. 15. figs. 8a, b) iden- ful that there are possibly more than t%vo tified one specimen from the Dzhulfian speci(\s of Pletironautihis in the Julfa Beds' strata at Baisal as Tainionautilus sp. The fauna. specimen is small (34 mm diameter) and has Repositonj. MCZ 9757 (PI. 4, figs. 3, 4), conspicuous forward-projecting lateral ribs. MCZ 9758 "(PI. 4. figs. 7, 8), unfigured The precise relationship of this specimen specimens MCZ 9759. KU 343S9, 34414. w ith ours is obscure because of poor preser- 34415. 34419. 34427. 34428. xation and the fragmentarv nature of our specimen. Genus Tainionautilus Mojsisovics, 1902 Rcpositon/. MCZ 9760. unfigured specimens KU 34391, 34413. Type species, Nautilus trans'iforius Waagen, 1879 Family KONINCKIOCERATIDAE Hyatt, Tainionautilus sp. indet. mZittel, 1900 Plate 3, figures 9, 10 Genus 7emnoc/ie;7us McCoy, 1844 Disctissioii. This identification is made on Type species. Nautilus (Temnocheilus) a single fragment of phragmocone consisting coronatus McCoy, 1844 of only one-third \olution. The whorl section is depressed, subquadrate, meas- Temnocheilus sp. indet. uring 41 mm in width and approximately Plate 3, figures 1-4,7, 8 30 mm in height. The flanks are flattened Discussion. This very distincti\e late and converge slighth- toward the venter. Paleozoic genus is in our collec- The venter is broad and flattened. Both represented tion one the \-entrolateral and mnbilical shoulders by nearly complete phragmocone and sLx fragments. The best specimen (Pi. are sharph' rounded. The venter bears a 3. figs. 3, 4) has a diameter of 61 mm, an conspicuous, smooth, deep median furrow adoral whorl width of 27 mm (measured aligned on both sides by broad diagonal on the nodes), and a whorl of 15.5 ribs that extend to the ventrolateral shoulder. height mm. Tlie umbilicus has a diameter of 30.5 The lateral areas do not appear to bear ribs, mm. The whorl cross section is subtrape- but this is uncertain because of poor preser- zoidal \\'ith a broad, only slighth- convex vation. The sutiu'e forms a deep \'entral lobe venter, rounded ventrolateral shoulders, and on the venter and a broad rounded lobe on convex lateral areas that converge toward the lateral area. the dorsum. The \entrolateral area bears Only five species have been referred to prominent nodes spaced about the width this genus, Tainionautilus fugax Mojsiso- of one camera apart. The suture has a broad \'ics, from the Bellerophon Limestone, the shallow ventral lobe and a similar lobe on type species T. transitorius (Waagen), T. the lateral areas. A smaller fragment uynnei (Waagen), T. multicostatus (Reed). with a whorl width of 17 mm is illustrated and T. sp. Reed from the Chhidru Formation on Plate 3, figures 7, 8. Three of the re- of the Salt Range, West Pakistan. Tainio- maining fragmentary specimens are portions 420 Bulletin Mu.s(in)i of Coinpurativc Zoology, Vol. 144, No. 6

of larger individuals. One specimen is a at Kuh-e-Ali Bashi are species of the genus fragment of body chamber and measures Domatoceras. We have 35 specimens of 39 mm in width and 27.5 mm in height. which only a few are essentially complete. Another fragment is a phragmocone con- The same predominance prevails in corre- sisting of four camerae that measures 43 mm lative strata of Soviet Dzhulfa where Shi- in width 21.5 in in Ruzhentsev and mm height. The larger manskiy ( and Sarycheva, fragments ha\'e the same whorl shape, orna- 1965: 41) reported 36 specimens. In addi- ment pattern, and suture as the more com- tion to being represented by large numbers plete specimen of Plate 3, figures 3, 4. of specimens, there is also a great diversity Temnocheilus is fairly common in the in form. The differences in form are Pennsylvanian and Permian. Kummel expressed primarily in the shape of the

1953 : 18 in a review of the whorl section. from ( ) , general Shimanskiy recognized genus, listed 23 described species from the th(^ Araxoccras beds the following species: U.S.A. and Eurasia. Within the family D. convergcns (Abich), D. atypicuui Shi- Koninckioccratidae Temnocheilus is a long- manskiy, D. hunicum (Diener), D. gracile ranging genus that appears to have given Shimanskiy, and D. parallclum (Abich). In rise to onlv one other form—FoorcUceras. the overlying Vedioceras beds Shimanskiy This is in sharp conti'ast to Metacoceras, records D. cf. iiracile and D. sp. 1, and in which gave rise to a numbc^r of late Paleo- the equivalents of th(> Ali Bashi Formation, zoic genera (Kummel, 1953). What is of D. sp. 2. interest is that the two genera represent Nautilus convergens Abich we believe to parallel evolutionary lines. be a species of Stenopoceras rather tlian Shimanskiy {in Ruzhentsev and Sarycheva, Domatoceras. Domatoceras (itij))icum Shi- 1965: 41) lists Pseudotemnocheihts from thc^ manskiy is an inflated form with a narrow Araxoccras and Vcdioceras bcxls in Soviet conea\e \(Miter and convergent convex Dzhulfa; however, he neither discussed nor Hanks; t)ur collections include two specimens illustrated these in of this that specimens. Kummel ( species. We believe D. gracile Teichert et al., 1964) placed Psetidotemno- Shimanskiy is a synonym of D. parallclum cheilus in the synonymy of Temnocheilus. (Abich); our collections contain eight It would thus appear that this Soviet region specimens of this species. The most abun- includes in its nautiloid fauna a temnochei- dant species, represented by 23 specimens, lid possibly similar to that of Kuh-e-Ali is D. hunicum (Diener), which has a sub- Bashi. rectangular whorl section. Finally, our Repositon/. MCZ 9761 (Pi. 3, figs. 1, 2), collections contain a fragmentary specimen MCZ 9762 (PI. 3, figs. 3, 4), KU 34394 (PI. consisting of one camera and a portion of of 3, figs. 7, 8), unfigured specimens MCZ body chamber, making up one-quarter 9763^ a volution of conch, and this seems to represent a very unique new genus of Superfamily TRIGONOCERATACEAE domatoceratid. The primary feature is a Hyatt, 1884 fastigate venter with a smooth sharp midline and the remaining ventral area convexly Family GRYPOCERATIDAE Hyatt, curving to merge with smooth flanks. To in Zittel, 1900 the best of our knowledge no other domato- a section. The Genus Domofoceros Hyatt, 1 891 ceratid has such whorl presence of this form emphasizes the statement Type species, Domafoceras umbilicafum regarding the diversity of the domatocer- Hyatt, 1891 atids in the Julfa Beds. Because our speci- Discussion. The predominant coiled nau- men is so fragmentary we refrain from tiloids in our collections from the Julfa Beds introducing a new generic name, but that it Upper Permian Nautiloids, Iran • Tcicheii and Kiimmcl 421

represents a new genus we have no ques- Dzhulfian fauna. The specimen of Doinato- tion. ceras gracile illustrated by Shimanskiy {in Ruzhentsev and Sarycheva, 1965, pi. 16, fig. Domatoceras hunicum (Diener) 1) is identical to our specimen illustrated Plate 1, figures 7, 8; Plate 2, figures 3, 4 on Plate 2, figures 9, 10. We believe these specimens to be ju\'enile forms; a mature Nautilus {Domatoceras?) hunicus Diener, 1903: is that illustrated on Plate 5, pi. 1, figs. la-c. phragmocone Gn/poceras (Domatoceras) liunicum, - Kunnnel, 2, figures 1, 2. In the more mature whorls 1953: 55. the flanks are more parallel than in the Domatoceras - in Ruzlientse\- hunicum, Shimanskiy, immature whorls. Another difference is that and .Sarycheva, 1965: 41, 15, fig. 12. pi. the ^'{M^trolateral and umbilical shoulders Discussion. This species is represented of immature whorls are sharply rounded, in our collections by 24 fragiiiciitar\' speci- though much less so than those of mature mens, most of which are body chambers. whorls. In conjunction with this change The species is characterized by an evolute, the venter on the inner whorls is flat but smooth conch with a siibqnadrate whorl becomes slightly convex on the outer whorls. section. The illustrated b()d\- chamber (PI. Our collections contain nine specimens of 2, figs. 3, 4) has a whorl height of 27 lum this compressed form, but we conclude that and a width of 23 mm. The venter is D. gracile is a synonym of D. parallelum flattened and the flanks are only slightly (Abich). convex, converging gently toward the venter. Repository. MCZ 9767 (Pi. 2, figs. 9, 10), Tlie ventrolateral and imibilical shoulders KU 34387 (Pi. 2, figs. 1, 2), unfigured speci- are rounded. The siphuncle is central. The mens MCZ 9768. suture has a ventral lobe that occupies most of the venter and a lateral lobe that Doma/oceras sp. indet. occupies nearly all of the flanks. Plate 2, figures 7, 8 This species differs from the associated Discus.?ion. This species is represented species in the fauna primarily in the shape b\' two fragments of phragmocone. The of the whorl section. Domafoceras para]- illustrated .specimen (Pi. 2, figs. 7, 8) has leliiin has a much nrore compressed whorl a whorl height of approximately 28.5 mm section and D. a more inflated atypicum and a width of 34.5 mm. The venter is whorl section with a relatively narrow slighdy concave and has a width of 15 mm. venter. The ventrolateral shoulders are rounded MCZ 9764 Repository. (PI. 2, figs. 3, 4), and the flanks broadly convex, diverging 9765 MCZ (PI. 1, figs. 7, 8), unfigured markedly toward the dorsum. The width specimens MCZ 9766, KU 34393, 34395, of the whorl at the umbilical shoulder is 34429, 34430. nearly twice the width of the venter. Tlie umbilical shoulders are sharply rounded. Domafoceras parallelum (Abich) The suture has a ventral lobe occupying the Plate 2, figures 1,2, 9, 10 venter and a lateral lobe occupying most of the flank. Nautilus parallelus Abich, 1878: 17, pi. 3, fig. 2. - Grijpoceras (Domatoceras) parallelus, Kummel, These specimens are considerably differ- 1953: 55. - ent from any domatoceratid described or Domatoceras parallelus, Shimanskiy, in Ruzhentsev illusti'ated from Soviet and Sarycheva, 1965: 41, pi. 15, fig. 10. by Shimanskiy Domatoceras gracile Shimanskiy, in Ruzhentsev Dzhulfa. It most resembles Domatoceras and Sarycheva, 1965: 160, pi. 16, fig. 1. atypicum Shimanskiy. However, in that Discussion. This is the compressed form species the whorls are said to be higher among species of Domatoceras in the than wide and the sutures to be straight 422 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6

across the venter. The venter was de- shoulder. Ventral shoulder protrudes scribed as narrow but is convex ratlier than and is carinate, umbilical shoulder concave. The shape of the whorl section almost rectangular. Sculpture consists of our specimens is unique among domato- of oval tubercles stretched out along ceratids described to date. Unfortimately, the \'entral shoulder and of sparse oval the specimens are too incomplete for com- tubercles on the umbilical shoulder. plete analvsis. Sutures broad, with deep lateral and Repository. MCZ 9769 (Pi. 2, figs. 7, 8), ventral lobes and a smaller and nar- unfigured specimen MCZ 9770. rower dorsal one. A very small lobe occurs on the umbonal wall. Domatoceratid n. gen., n. sp. further stated that (translated) : Plate 2, figures 5, 6 Shimanskiy On the basis of the of Discussion. One specimen in oiu^ collec- general shape the shell, the of the \entral tion is unique and clearly belongs to an concavity side and the of oval tubercles undescribed new genus of domatoceratids. presence the ventral shoulder the new However, it consists of only one camera and along resembles Titanoceras Hyatt. a small portion of phragmocone. It is clearly genus The of the cross section is a too incomplete to be made the type of a shape feature new genus. The unique feature of the distinguishing (being trapezoidal in our and convex specimen is thc^ acute x'cnter with arching genus having lateral sides in and also ventral shoulders blending smoothh* with Titanoceras) , tlie occurrence of tubercles the lateral flanks. The suture begins witli pre-umbonal in Fseiidotitanoceras. a pointed saddle on the acute venter, followed a shallow concave lobe by occupying In spite of this resemblance Shimanskiy all of the ventral then an acute slioulder, concluded that the two genera were unre- saddle followed by a broad concave lobe lated homeomorphs and that Fseiidotitano- the whole flank. occupying ceras might belong to tlie family Mosquo- The acute venter and the suture clearly ceratidae. indicate that this specimen represents a unique, as yet undescribed new genus Pseudofifanoceras armeniacum (Abich) related to Domatoceras. Plate 1, figures 5, 6 Repository. MCZ 9773 (PI. 2, figs. 5, 6). Nautilus armeniacus Abich, 1878: 24, pi. 2, fig. 5. Pscudotitanoceras armeniacum, - Shimanskiy, in Genus Pseudotitonoceros in Shimanskiy, Ruzhentsev and Sarycheva, 1965: 162, pi. 16, and Ruzhentsev Sarycheva, 1965 figs. 5, 6.

Type species. Nautilus armeniacus Abich, Discussion. Our collections include one 1878 specimen that can readily be assigned to this It is a of Discussion. When he established the species. large fragment phragmocone of one-third volution. Only the genus Pseudotitanoceras, Shimanskiy (in venter and one lateral side are preserved. Ruzhentsev and Sarycheva, 1965: 162) gave The most adoral part of the specimen most the following diagnosis (translated from probably preserves a portion of the living Russian : ) chamber. Our specimen has a length of 120 Shell large, evolute, discoidal, whorl mm; we estimate the total diameter is in section trapezoidal; height of whorls excess of 200 mm. The ventral area is wide almost equal to width. Ventral side and markedly concave. The ventrolateral conca\'e, lateral sides flat, diverging shoulders are acutelv rounded and aligned considerably towards the umbilical by elongate, low nodes. The lateral sides Upper Permian Nautiloids, Iran • Teichert and Kummcl 423

converge strongly toward the ventrolat- too incomplete for meaningful comparison eral shoulders and are flattened except near with the known species of the genus. Again the ventrolateral shoulder where they it should be pointed out that all species are become slightly concave. The camerae are knowai from very few, often fragmentary 12 mm high measured along the \'enter specimens. except the most adoral two camerae, which Stenopoceras is a compressed involute arc 9 mm and 10 mm high. The suture has evolutionary offshoot of Domatoceras. In a deep V-shaped ventral lobe, a narrowh- the Triassic, Gnjpoceras is the direct lineal rounded saddle centered on the ventrolat- descendant of Domatoceras and itself gave eral shoulder, and a broad, deep, rounded off a compressed involute line, Gryponau- lat(>ral lobe occupying the whole lateral area. tihis. Our specimen is approximately twice the Repository. KU 34434 (Pi. 1, figs. 9, 10). size of the largest recorded b\' Shimanskiy fiom Soviet Dzhulfa, but there is no reason Genus Titanoceras Hyatt, 1884 to doubt that they are conspecific. This is species, Noufilus ponderosus Meek, the as to P.seudo- Type only species assigned yet 1872 titanoccras and is known only from the Dzluilfian strata in the vicinity of the Aras Tifanoceras sp. indet. River. Even here, however, it is rare, as Plate 1, figures 1, 2 Shimanskiy had only four specimens and Discussioti. Our collections contain one we managed to collect onlv one. large specimen that represents an indeter- Repository. KV 34392 '(PI. 1. figs. 5, 6). minate species of this genus. The specimen is all phragmocone, consisting of one-third Genus Stenopoceras Hyatt, 1893 \-olution. The specimen had a diameter of Type species, Phacoceras dumblei Hyatt, at least 140 mm. The adoral camerae have 1891 a height of 59 mm and a width of approxi- 6r. The suture has a narrow, acute \'enter, but the lateral area is occupied by ventral saddle and a broad deep lateral a broad lateral lobe. The camerae along the lobe. These are the diagnostic features of ventrolateral shoulder in the adoral portion the genus Stenopoceras. of the phragmocone are 15 mm in width; in it is 4 Kummel ( 1953 ) reviewed the taxonomic the two adoral camerae mm. and evolutionary data on Stenopoceras and Titanoceras is not a particularly common listed eight species of the genus. Most of late Paleozoic nautiloid. Previously it has the species are known from Upper Carbon- been reported only from Upper Carbonif- iferous and Permian formations of the erous and Lower Permian formations of

United States. One species, S. rouillieri (de Nebraska and western Australia. Tliere is Koninck) occurs in the Upper Carbonif- no reason to doubt the generic assignment erous of central Russia. Our specimen is of our specimen, but specific comparison 424 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6

with previously described species is of lim- REFERENCES ited value because of its incompleteness. Abich, H. W. 1878. Geologische Forschungen MCZ 9771 1. 1. Repositonj. (PI. figs. 2). in den kaukasischen Landern. Th. 1, Eine Bergkalkfauna aus der Ara.xesenge bei Djoulfa Superfamily CLYDONAUTILACEAE Hyatt, in in Armenien. Wien: Alfred Holder. 126 pp. I. S. 1963. Sistema i Zittel, 1900 Bahskov, filogeniya psev- dortotseratid. B\ull., Moskov. Obshch. Isp- Family LIROCERATIDAE Miller and Young- ^•tat. Prirodi. N.s', Vol. 68, Otdel GeoL, 38(4) : 149-150. quist, 1949 DiEXER, C. 190.3. Permian fossils of the Central Himalayas. Paleont. Ser. Genus /./roceros Teichert, 1940 Indica, 15, 1(5): 1-204. R. H. 1939. of the Pseudor- Type species, Co/oceras liratum Girty, 191 1 Flower, Study thoceratidae. Palaeontographica Amer., 2: Liroceras sp. indet. 245^60, 9 pis.

~~. 1941. Plate 3, figures 11, 12 Revision and internal structures of Leurocycloceras. Amer. Tour. Sci., 239: Discussion. This is one of thc> more 469-488, 3 pis.~~

. 1943. Studies of Paleozoic Nautiloidea. common nantil(jid genera in the Julfa Beds Annulated orthoceraconic of at Kuh-e-Ali Bashi. Our collections contain VII, genera Paleo- zoic nautiloids. Bull. Amer. Paleont., 28 16 most of which arc specimens, fragmentary (109): 102-140. and crushed. The slightly largest specimen, . 1945. Classification of Devonian nau- consisting of a phragmocone, had a diameter tiloids. Amer. Midland Naturalist, 33: 675- of at least 100 mm. 724, 5 pis. AND K. E. , Caster. 1935. The Liroceras is one of the simplest of late stratig- raphy and paleontology of northwestern Paleozoic nautiloids. It is characterized bv Pennsylvania. Part II, Paleontology. Section a imohite smooth, globular, conch with a A, The cephalopod fauna of the Conewang depressed whorl section and an essentially Series of the Upper Devonian in New York and Bull. Amer. 22 straight suture. E\en with well-prescrxed Pennsylvania. Paleont., (75): 1-74, 8 pis. material .specific identification is most diffi- -, .\ND B. KuMMEL. 1950. A classification cult. Our material is not well enough of the Nautiloidea. Jour. Paleont., 24: 604- preserved for this. 616. In x'iew of the relatixe abundance of Frech, F. 1911. Abschlies.sende palaontologische Liroceras in our collections, it is of interest Bearbeitung der Samuihingen F. von Richt- die weiterer fossiler that the genus is not listed by Shimanskiy hofens, Untersuchung Reste aus den von ihm bereisten Pro\'inzen (in Ruzhentsev and Sarycheva, 1965: 41) sowie ein Entvvurf einer erdgeschichtlichen as a member of the nautiloid fauna in Uebersicht Chinas. In Richtliofen, F. F. von, Soviet Dzhulfa. At the same time the speci- 1877-1912, China. Berlin: Dietrich Reimer, Vol.5: 1-289. mens Abich ( 1878 ) described and illustrated

, AXD G. VOX Arthaber. 1900. tjber das as Nautilus concavus and Nautilus propin- Palaozoicum in Hocharmenien und Persien qtius Abich appear to be conspecific with mit einem Anhang iiber die Kreide von Sirab (Hu- forms. Our collections from the over- in Persien. Beitr. Palaont. Geol. Ost.-Ung. lying Ali Bashi Formation contain two Orient., 12: 161-308. specimens of Liroceras for which there is no GmxY, G. H. 1911. The fauna of the Moore- field shale of Arkansas. Bull. U. S. Geol. reason to believe that they are not conspe- Surv., 439: 1-148. cific with those recorded here from the Hyatt, A. 1883. Genera of fossil cephalopods. Julfa Beds. Proc. Boston Soc. Nat. Hist., 22: 25.3-272.

. 1884. Genera of fossil Repository. KU 34386 (PI. 3, figs. 11, 12); cephalopods. Proc. Boston Soc. Nat. 22: 273-338. 10 Hist., unfigured specimens MCZ 9772 ( speci- . 1891. Carboniferous cephalopods. Ann. KU 343S5. 34420. mens), 34418, 34424, Rept. Texas Geol. Surv., No. 2: 327-356. 34426. . 1893. Carboniferous cephalopods, se- Upper Permian Nautiloids, Iran • Teidieii and Kummcl 425

cond paper. Ann. Rept. Te.xas Geol. Surv., Rexz, C. 1940. Die palaeozoischen Faunen \on Xo. 4(1892): 377^74. 1929-30. Wissensch. Ergebnisse der Nieder- Klmmel, B. 1953. American Triassic coiled landischen Expedition in dem Karakorum, vol. nautiloids. U. S. Geol. Surv., Prof. Paper 250: 3, pt. 1, Geologic, Abt. 1. Leiden, pp. 9-68. 1-104, 19 pis. RuzHEXTSEV, V. E. (ed.) 1962. Osno\y Paleon- 1971. Pemiian- , AND C. Teichert. tologii. Molh uski-Golovonogie I. Izdatel. Triassic boundar>' beds in middle Tetliys .\kad. Nauk SSSR, Moscow. 438 pp.

). Int. Permian-Triassic Conference, AXD T. G. Sarycheva. 1965. Raz\'itie i (Abst. , Calgar>-, 1971, Program witli Abstracts. Uniw smena morskikh organizmo\- na rubezhe Paleo- Sur\-. Bull. Calgary, Geol. Canada. Canad. zoya i Mezozo>a. (Development and change Petrol. Geol, 19(2): 336-337. of marine organisms at the Paleozoic-Mesozoic

~~AXD . Pennian-Triassic boundar\' -, boundarw ) Trudy Paleont. Inst. Akad. Nauk beds in middle Tethys. Can. Sci. Jour, (in SSSR. 108: 1-431.~~

press ) . Stepanov, D. L., F. Golshaxi, axd J. Stocklix. McChesxey, J. H. 1860 (1859). Descriptions 1969. Upper Permian and Permian-Triassic of new of fossils from the Paleozoic species boundary in North Iran. Geol. Surv. Iran, rocks of the western states. 1- Chicago, pp. Rept. No. 12: 1-72. 76. Stoyaxgw, a. a. 1910. On the character of the McCoy, F. 1844. A sx-nopsis of the characters boundary of Paleozoic and Mesozoic near of the Carboniferous Limestone fossils of Djulfa. Verhandl. Russ.-Kaiserl. Mineral. Ireland. Dublin: Univ. Press. 207 pp., 29 pis. Gesell., St. Petersburg, Ser. 2, 47: 61-135, Meek, F. B. 1872. Report on the paleontolog\ pis. 6-9. of eastern Nebraska \\ ith some remarks on the C. 1940. Carboniferous rocks of that district. In F. \'. Teichert, Actinosiphonate cephalo-

pods ( CyTtoceroidea ) from the De\onian of Ha>den, Final report of the United States Australia. Joiu". Royal Soc. W. Australia, 26: Geological Sur\ey of Nebraska. Final Rept., 59-75, 4 pis., 2 figs. Pt. 2, Paleont., pp. 83-239, pis. 1-11. et al. 1964. fea- .WD A. H. 1861. , Cephalopoda-general , WoRTHEx. Descriptions of new Carboniferous fossils from Illinois and tures, Endoceratoidea-Actinoceratoidea-Nauti- other western states. Proc. Acad. Nat. Sci., loidea, Bactritoidea. In Moore, R. C. (ed.). Philadelphia, I860: 447-472. Treatise on Invertebrate Paleontology, Pt. K, A. A. G. 1942. .Miller, M., axd Uxklesbay. Mollusca 3. Geol. Soc. Amer., Univ. Kansas Pemiian nautiloids from western United Press, 519 pp., 361 figs. States. Jour. Paleont., 16: 719-738. — B. .\XD , KuMMEL, W. Sweet. Permian- Mojsisovics, E. vox. 1882. Die Cephalopoden Triassic strata, Kuh-e-Ali Bashi, northwestern der mediterranen Triasprovinz. Abhandl. Geol. Iran. Bull. Mus. Zool. Reichsanst. Wien, 10: 1-322. Comp. (in press). W. 1879. Salt Productus . 1902. Das Gebirge urn Hallstatt. Ab- Waagex, Range fossils; handl. Geol. Reichsanst. \\'ien, Suppl. H., 6: limestone fossils. Mem. India Geol. Surv.,

~~175-356. 1 : Paleont. Indica, Ser. 13, 1 ( ) 1-72. 426 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6~~

~~PLATE 1. TITANOCERAS, PLEURONAUTILUS, PSEUDOTITANOCERAS, DOMATOCERAS, STENOPOCERAS~~

~~Figures Page~~

~~1, 2. Titanoceras sp. indet., MCZ 9771. X0.5. 423~~

~~3, 4. Pleuronautilus sp. Indet. 1, KU 34384. XL 418 5, 6. Pseudotitanoceras armeniacum (Abich), KU 34392. XO.5. 422~~

~~7, 8. Domatoceras hunicum (Diener), MCZ 9765. XO.5. . 421~~

~~9, 10. Stenopoceras sp. indet., KU 34434. 9, XI; 10, X%. 423 Upper Permian Nautiloids, Iran • Teichert and Kummel 427~~

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~~10 428 Bulletin Museum of Comparative Zoology, Vol. 144, No. 6~~

~~PLATE 2. DOMATOCERAS~~

~~Figures Page~~

~~1, 2. Domafoceros parallelum (Ablch), KU 34387. X 1. 421~~

~~3, 4. Domofoceros hunicum (Abich), MCZ 9764. XI- 421~~

~~5, 6. Domatoceratid, n. gen., n. sp., MCZ 9773. XI- 422~~

~~7, 8. Domofoceros sp. indet., MCZ 9769. XI- 421~~

~~9, 10. Domofoceros parallelum (Abich), MCZ 9767. XI- 421 Upper Permian Nautiloids, Iran • TeicJiert and Kummel 429~~

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~~PLATE 3. TEMNOCHEILUS, METACOCERAS, TAINIONAUTILUS, LIROCERAS~~

~~Figures Page~~

~~1,2. Temnocheilus sp. indef., MCZ 9761. XI- 419~~

~~3, 4. Temnocheilus sp. indet., MCZ 9762. XL 419~~

~~5, 6. A^efococeras dorsoarmatum (Abich), MCZ 9755. XI- 417~~

~~7, 8. Temnocheilus sp. indet., KU 34394. X 1 419~~

~~9, 10. Ja'mionaufilus sp. indet., MCZ 9760. XI 419~~

~~11, 12. Liroceras sp. indet., KU 34386. XI- :• 424 Upper Permian Nautiloids, Iran • Teicheii and Kummel 431~~

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~~PLATE 4. TAINOCERAS, PLEURONAUTILUS, METACOCERAS, LOPINGOCERAS, NEOCYCLOCERAS~~

~~Figures Page~~

^16 1, 2. To/noceros sp. indet., MCZ 9752. XL "llS 3,4. Pleuronaulilus sp. indet. 2, MCZ 9757. X%- 5, 6. Metacoceras dorashamense Shimanskiy, MCZ 9753. XI- '^^^ '118 7, 8. Pleuronautilui sp. indet. 2, MCZ 9758. XI 9. Lopingoceras /op/ngense (Stoyanow), MCZ 9774. Xl-5 414

~~10. Neocyc/oceros sp. cf. N. obllqueannulalum (Waagen), KU 34411. XI 415~~

~~11. (.op/ngoceros /op/ngense (Stoyanow), MCZ 9776. X 2. -- 414 12. Neocyc/oceros sp. cf. N. obliqueannulatum (Waagen), MCZ 9777. XI- 415~~

~~13. 14. Lopingoceros sp. cf. L /op/ngense (Stoyanow), MCZ 9775. X1-5. - 414 Upper Permian Nautiloids, Iran • Teichcrt and Ktimmcl 433~~

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