a! 9, Phylum: Mollusca 2. Subphylum: Conchijerg Fig, 91. Median section of a Recent nautilus, shell mantle hood buccal mass radula\ upper jaw. inner ring of tentacles \, outer ring of tentacles cirri - lower jaw funnel valve hyponome “ cerebral ganglion — pedal ganglion / visceral ganglion gill \ kidney ‘osphradium Summary of Recent cephalopods Among the Recent cephalopods two groups are distinguished which differ considerably in terms of diversity: A. Tetrabranchiata (Ectocochlia): cephalopods with two pairs of gills, numerous arms and an external shell. Nautilus (‘pearly nautilus’) is the only representative of this group. B. Dibranchiata (= Coleoidea, Endocochlia): cephalopods with one pair of gills. The shell is inside the soft body, if and when it is developed at all. This group includes: 1. The ‘Decapoda’ (squids, Decabrachia with the Orders Teuthida and Sepiida) with ten arms of which two are differentiated into tentacles, 2. The ‘Octopoda’ (octopuses, Octobrachia) with eight arms and a sac-like body.ISS 'CSsSs;s ~~~ 6, Class: Cephalopoda a tof their common ancestry and similar radulae, the ammonoids and accoun blished as closely related sister groups, as opposed to the coleoids are esta Speirs group of the archaic Nautilus relatives, In this book the former are Hs combined as the Infraclass Neocephalopoda and the latter as the Infraclass palcephalopoda (= ‘nautilids’), Infraclass: Palcephalopoda (nautilids in the wider sense) The first three subclasses of the cephalopods are here combined as the palcephalopoda: the Nautiloidea, Endoceratoidea and Actinoceratoidea. The morphology of fossil Palcephalopoda (nautilids) The fossil relatives of the present-day Nautilus also had an external shell. The animals lived in the most recently formed part of the shell, the so-called body chamber, which was open to the outside. The older part of the shell, the shragmocone or chambered part, is divided into numerous chambers by dividing walls known as septa. The very first chamber to be formed is known as the protoconch (initial or apical chamber). A membranous tube, the siphuncle, traverses all the chambers from the protoconch to the body chamber. The significance of the siphuncle Studies of Recent cephalopods with shells have revealed that the chambers contain a gas mixture resembling atmospheric ait without oxygen, i.e. consisting mainly of nitrogen, with a gas pressure of about 0.8 atmospheres. Newly formed chambers are initially filled with fluid. Only when the new septum is sufficiently strong to resist the water pressure will the siphuncle pump the water out, sometimes against a substantial external pressure. The siphuncle contains arterial and venous blood vessels. Its tissues are capable of selectively excreting water and salts, rather like kidney tissue. The shells of fossil nautilids may be rolled into a logarithmic spiral like the Nautilus, but most of them, especially the geologically oldest ones from the Cambrian to Silurian, are more or less straight. The siphuncle may occupy different positions varying from central to peripheral. Using the present-day Nautilus as a reference, the siphuncle is regarded 4S a ventral organ, Accor (a) endogastric (siphuncle : f the whorl); ( on the outside of the who: J me9. Phylum: Mollusca 2, Subphylum: Conchiferg >» Fi o 135 itudi f part of an actinoceratoig tudinal section of pa : tip led fen represent the areas filled by prima stip \ ry Carel during the lifetime of the animal, siphuncle radial canal air chamber SePtal deposits endosiphonal Geposits Perispatium, SUPPOTtINg ring septum shell wait the next septum, the siphuncle is Surrounded by a membranous Siphonal laye (= connecting ting, ectosiphon) (Fig. 92), The siphuncle may be iphonal calcareous secretions (Fig, 93). According to the structi » Stenosiphonate (with a si wide siphuncle) forms are Partly or completely filled with endosi; ‘ure of the siphuncle. narrow siphuncle) and eurysiphonate (with a distinguished, In addition to the si Phuncle, the air chambers may also be more or less extensively filled with calcareous deposits,6, Class: Cephalopoda cyrtocone: slightly curved gyrocone: coiled into a loose spiral nautilicone: completely coiled The orthocone and cyrtocone shells are either brevicone Jongicone (elongated, slender). In contrast to the jaws of all other Recent cephalopods, the jaws of the Recent Nautilus are partly calcified. The radula has 13 teeth per transverse row and is thus fairly wide (‘latiradulate’), With very few exceptions, the dividing walls (septa) are simple and at most slightly arched. This applies particularly to the marginal parts which are fused with the outer shell wall (the so-called suture line), An indentation in the initial chamber of some nautilids which sometimes looks like a healed injury is known as cicatrix (‘scar’). It does not occur in the ammonoids. Its origin and function are unknown, but they are probably linked with the initial part of the siphuncle, the caecum, In the Upper Ordovician the Palcephalopoda achieved their maximum size with up to 9m in the endoceratids. With the exception of the Recent Archi- teuthis, the cephalopods have never surpassed this size. In terms of the number of species, their climax of development was in the period between the Upper Ordovician and the Lower Devonian. Coiling first began in the Ordovician and prevailed until the Triassic, with the concomitant decline of the orthocone, cyrtocone and gyrocone forms. Only coiled nautilids are known in the post-Triassic Period. (short, squat) or Palaeoecology and biostratonomy As the only Recent form, the present-day Nautilus is predominantly distributed in the Indo-Pacific area. The shells may have carried over great distances after death, In experiments, even slightly damaged empty Nautilus shells floated for about one month. In fossil forms the length of the body chamber is important. An empty Endoceras will float provided the body chamber is no} less than one-half is i 1 of the phragmocone (less than ‘halt not longer than one-quarte!2. Subphylum: Conchifera 9, Phylum: Mollusca 13g mechanism, which is not certain in all cases, then the funnel and body ch, must have been in a roughly horizontal position: otherwise one could only imagine a slow creeping movement or an equally slow vertical movement, Early ellesmeroceratids with very narrowly spaced septa will certainly haye been part of the mobile epibenthos. In their case, equilibrium in the water would have been guaranteed by a little buoyancy in the chambers, The following shell types predominate: 1, More or less complete coiling, stability provided by ventral POsition of the body chamber. 2. Heavy cameral and siphonal deposits, particularly in the ventral ang apical regions; stable horizontal position (Fig. 94), 3. Ontogenetic loss of older parts of the shell by truncation (decollation), air chambers in the dorsal part of the teleoconch: stable horizontal position. 4. Formation of brevicone shell: stable vertical position because the heavy body is in the body chamber at the bottom, Any preserved colouring is distributed accordingly: on one side only in longicone shells and more or less evenly all round in brevicone shells, Fossil nautilids may have had ink sacs, but there is still no unequivocal evidence to this effect. Fig. 94. Longitudinal sections of various nautilid shells of the Lower Palaeozoic. Stippled areas, body chamber and siphuncle; black areas, calcareous secretions inside the chambers and siphuncle. “Orthoceras’ body chamber Actinoceras6. Class: Cephalopoda 139 ions of the maximum diving depths, b S ert and the shell, have yielded the flowing ese oe of the ae 450m , Plectronoceras 250m Orthoceras 400m Naseihus 800m Bactrites 550m Spirula 1600 m These are maximum values, and the actual d ve been about half the values shown, as ill aa fossil nautilids were thus inhabitants of the shallow sh, hand, the water could not be too shallow, oth would have been jeopardised by far-reaching ie erwise the lar, Pths inhabited will Probably only ustrated by the example of Nautilus. elves. On the other g¢ forms especially lifestyle may have been preferred by small f wave Movements. A pl anktonic to present-day Spirula. Larger forms were P iL: Subclass: Nautiloidea Since the (? Lower) Upper Cambrian. commonly straight at first; geologically youn: The position of the siphuncle is variable; it is cyrtochoanitic (Fig. 95). development ensued in th ¢ Lower Ordovician. The Orthocerida have Straight shells. O Orthoceras (Fig. 96A , 3), Michelinoceras. and the time of their occurrence have still Orms such as the bac tobably Part of the tritids, similar benthos. A very diverse central group; ger forms are Strongly involute. thin, orthochoanitic or ccurrence: Ordovician-Triassic, Their division from similar genera not been satisfactorily resolved. Fig. 95. The most important types of cephalopod siphuncle.fo 9, Phylum: Mollusca 2. Subphylum: Conchifery Ce Fig. 964. Nautiloidea: (1) Volborthetla (x 5); (2) Peg M4 (3) Orthoceras (x0.8);(4) Ascoceras (0:4); (3) Ones nrg b: x 2); (6) Gomphoceras (X 0.3). Ceras (4. xe ‘44 JW! The Ascocerida discarded the oldest, orthochoanitic part of the shell (so-called decollation); the remaining part had a cyrtochoanitic siphuncle and septa extending well to the front. Occurrence: Ordovician-Silurian. Ascoceras (Figs. 94 and 964, 4; Silurian). The Oncocerida are exogastric cyrtocones, often with a narrow aperture and actinosiphonate deposits. Occurrence: Middle Ordovician-Lower Carboniferous. Examples: Oncoceras (Ordovician), Gomphoceras (Silurian) (Fig. 964, 5, 6). : In contrast, the externally similar Discosorida are predominantly endogasti¢ with thick complex connecting rings or endosiphonal deposits and often with 4 narrow aperture. Example: Phragmoceras (Fig. 96B, 3; Silurian). The Tarphiycerida are initially coiled, but the later stages are un‘ varying degrees, with the position of the siphuncle changing, The ql 4 conspicuous hyponomic sinus, Occurrence; Ordovician-Silurianps di nda i | 6, Class: Cophalope wautitoidea: (1) Lituites (x02) (2) Ophiogera (1) : 1.0.2); (4) Nephriticeras (x 0.44); (5) Aturia (x 0.13); Fig. 96. Oo ceras @) PmmanonautTus 0.2); (7) Trochoceras (x 0.2). (@” includes the genera Lituites and Ophioceras (Fig. 96B, 1, 2) which are guide fossils of the Lower Ordovician. The Barrandeocerida are coiled to cyrtocone, their siphuncle is thin-walled and lacks deposits, orthochoanitic to secondarily cyrtochoanitic. Occurrence: Ordovician-Devonian, The group includes Nephriticeras (= 2 Cyrtoceras) (Fig. 96B, 4). The Nautilida embrace the majority of nautiloids from the Devonian to the present day. They are curved to coiled; the siphuncle is simple, orthochoanitic * and variable in position, There is a large break at the end of the Triassic which only the Superfamily Nautilaceae survived: A turia (Fig. 96B, 5; Palaeocene- Some of the Palaeozoic and Miocene) with slightly differentiated suture lines, Triassic Nautitida are strongly sculpted and the whorl section is sometimes rectangular. Examples: Germ 2 a Fain) (Fig. 968, 6, 7),f “em Wes, 9. Phylum: Mollusca 2. Subphylum: Conchiferg 2: Subclass: Endoceratoidea ly Usually orthocone and holochoanitic (Fig. 95) with i siphuncle with conical endosiphonal deposits (stacked one inal e They are directly derived from the Ellesmerocerida and grew iF, the ; to 9m, Occurrence: Ordovician-Silurian, Examples: £ length fe cian) ndoce Cyrtendoceras (Ordovician), Leurocycloceras (Ordovician-situ (Crdoy, 1-3). Tian) (Fig. 97 3 Subclass: Actinoceratoidea Orthocone, thick siphuncle bulging between the septa, end, canal system and very strong obstruction rings (annulosiphonate) Occ 1onal Ordovician-Carboniferous. Examples: Aetinoceras, Armenoceras (yea Ordovician-Lower Silurian) (Fig. 97, 5, 4). Fig. 97. Endoceratoidea: (1) Endoceras (x 0.2; a, b: x n doceras (x 0.25); (3) Leurocycloceras (X 0.7). Actinoessg ge) OT (4) Armenoceras (X 0.3); (5) Actinoceras (x 0.3) Be6. Class: Cephalopoda 5 Infraclass: Neocephalopoda 143 Comprising the ammonoids and coleoids and thei the bactritids. “iF mutual ancestorg Q Subclass: Bactritoidea Ordovician forms with a marginal si Bactroceras. The bactritids embrace the ancestray forms of the belemnites and are thus grouped together e ammonites and with them. Occurren, Permian. ce: Ordovician. In Bactrites (Fig. 98, 1) (Silurian-U; from such forms. oval, but in Cyrtobactrites of the Low Pper Permian) the cro; -section is slightly er Devonian (Fig. 98, 2) it is well flattened, Alateral lobe is added to the extemal siphonal lobe. Fig. 98. Bactritoidea: with the left part of the fig a ventral view; x 3); (2) Cy; (1) Bactrites (the figure represents a combination, ure showing a lateral view and ‘rtobactrites (x 1), the right part Parabactrites (Carboniferous-Permian) is brevicone and cyrtochoanitic with tarroly spaced septa, The belemnites can be derived from similar early forms. 5. Subclass: Ammonoidea ‘ This subclass includes the goniatites, clymeniids, ceratites and ammonites ‘S such. The term Ammonoidea should be carefully distinguished from the term ammonites which refers to Stratigraphically the A CSat 9. Phylum: Mollusca 2. Subphylum: Conchiferg v the suture is usually complicated; ' ; there is predominantly spiral coiling and abs, ence of forms. Primitivaty Nighy Differentiation of the suture line » Suture line: line of contact between the septum and the shey Prosuture: suture between the protoconch and the first gp. “al Primary suture: suture between the first and second | nber Lobe: part of the suture which is con nbers cave with respect to Saddle: part of the suture which is convex with tespect toe a External (ventral), lateral and internal (dorsal) lobes appear _ in and phylogeny; they are thus known as primary lobes Theeg rmtopeny ide of the shell Where the two preceding whorls meet (Fig. 99), Fig. 99. Evolution of the ammonoid Primary suture. three-lobed | fourlobed five-lobed [> sitobedl Cretaceous tw we [oe heteromorphs — x - Jurassic ORE 0 iH Triassic ceratites Devor Lobes can be incre; rise to umbilical (U) tise to adventitious ( arising in the extemal . formation of median (M) lobes, Sutural lobes (S-lobes) 4s formed in the Uslobe on i -f the umbilical6, Class: Cephalopoda 145 Fig. 100. Evolution of the ammonoid prosut ure, Vee rate Development 28 the primary suture (Fig. 99): in the Devonian the primary suture consists of internal, lateral and extemal lobes and the corresponding saddles, and is thus three-lobed (trilobate). In the Permian and Triassic the primary suture is four-lobed (quadrilobate) by the addition of a U-lobe; in the Jurassic and Cretaceous the primary suture is four-, five- or six-lobed (with one to three U-lobes). The prosuture and primary suture develop quite independently; the former is a larval structure. The terminology presented is genetic and suitable for the recognition and illustration of phylogenetic relationships. It is commonly used by German research workers and, in a modified version, by Russian workers. In the English-speaking areas the morphographic terminology set out in the Treatise on Invertebrate Paleontology (Moore & Teichert, 1952ff) is commonly used, It is easier to use for simple (not interpreting) descriptions. The Ammonoidea originated from the bactritids which themselves are derived from orthocone nautiloids (namely sphaerorthoceratids, themselves derived from michelinoceratids). Morphographic terms for the Ammonoidea are shown in Figs. 101 and 102. Fig. 101. Morphographic terms for the elements of the ammonoid suture line, ventral (median) saddle6. Class: Cephalopoda Fig. 103. Order Anarcestida: a) : : CL) Anetoceras (x 0.5): it oan (3) Anarcestes (0.5); (4) MaaneaerCCR Eris oud AWG) Protobltes (015); (7) Beloceras (x 0,3), 1 en oeeres Anarcestes (Fig. 103, 3): Parallel line to agoniatitids, with wider whorls and slower increase in the whorl height. Biconvex growth lines. Occurrence: Lower to Middle Devonian. Maenioceras (Fig. 103, 4); Flattened flanks; almost parallel external9. Phylum: Mollusca 2. Subphylum: beri ifera The following dominated in the Lower Upper Devon; nian: Manticoceras (Fig. 103, 5): Wide sad (= M-lobe); this distinguishes it from Anarces Explosive development of the manticoceratid dle on the 4, ‘ank, EB. tes, Occurrence: Upsated s in the Per Dey forms such as Upper Devonian Livi Beloceras (Fig. 103, 7): Disc-shaped, with largess of lobes, Set increas in the 2: Order: Clymeniida In this rapidly and highly specialised orde siphuncle is always internal except in the very early juvenile stages when i is external. The prosuture is latisellate. During the migration of fe a from the outside to the inside, the animal must have maintained its a (ventral side towards the outside). The clymeniids experienced i ae 4 the Upper Upper Devonian of Europe and North Africa in particular Fig. 104), 1 the retrochoanitie Fig. 104. Clymeniid and goniatite zone fossils of the Upper Devonian, series and stage zone fossils | |Wocklumerts vi Wocklumeria ieee Kalloclymenia 3 Dasbergian Vv Clymenia Upper © | Hembergian IN| Platyetymenia Devonian < | Nehdenian " Cheiloceras § < 3 i | Adorfian ' Acanthoclymenia from the Upper Devonian I of New York is regarded as the earliest genus. Lobe differentiation in the clymeniids: primary lobes E, LI or similar anarcestid as the ancestral form) gave rise to three lines nantly disc-shaped, an extremely wide umbili IS, § Examples: + clea Gonioclymenia: fronMere rccern, e, e 6. Class: Cephalopoda 149 Fig, 105, Phylogenetic development of the Suture line in the clymeniids, C ' ESA Ce Oer { : : Gonioclymeniace’ 2 2, Fig. 106. Order Clymeniida: (1) Gonioclymenia (x. 0.5); (2) Hexacly- menia (x 0.5); (3) Wocklume: ria (Left: x 0.5; right: x 0.75); (4) Platy- clymenia (a: X 0.8; b: x 1.3); (5) Clymenia (x 0.25); (6) Parawocklumeria (x 0.65).