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function of the ammonite fluted septal margins, The Lewy, Zeev ABSTRACT-The fluted margins of ammonite septa were thought to resist the hydrostatic pressure upon the phragmocone while the ammonoid dived. However, ammonoids probably did not dive deeper than the extant nautilids, whose conchs, with the simple septa, sustain pressure correlative to depth of about 800 m. The backward and forward stretching lobes and saddles actually provide resistance to pressure perpendicular to the septum. Ammonoids lived for about three to five years, and septa were precipitated in intervals of nearly two weeks to two days, which explain the small dimensions of the scars of the adductor muscles, which were periodically detached and reattached. The weak hold between these small muscles and the buoyant conch was compensated for by the backward branching and expanding folds (forming the sutural lobes), into which the soft tissue penetrated and stiffened for a required period to firmly anchor the body to the conch throughout its whole circumference. The greater the complexity of the septa marginal fluting, the better the ammonoid could withstand the dragging force between the body and the buoyant conch, and hence the more aggressively the ammonoid predated and competed with other creatures. INTRODUCTION HE AMMONOIDS evolved in the Devonian, comprising evolute to involute planispirally coiled conchs quite similar to that of the contemporaneous nautiloids. However, in the ammonoid conch the siphuncle lies in a ventral position (dorsal in Upper Devonian clymeniids), and the septal margins develop more folds than in nautiloids (more sutural elements). The main evolutionary trend within the Ammonoidea is evidenced by the increase in the number of septal marginal folds and by their complexity, in contrast to the rather simple bowl-shape septum in most of the nautiloids. Since their origin in the Devonian, ammonoids highly diversified and became abundant in the Mesozoic oceans, whereas nautiloids decreased in diversity to about five genera towards the end of the era (Moore, 1964). There is no doubt among paleontologists that the great advantage of the ammonoids over the nautilids was achieved through the fluting of the septa margins; however, the actual function of this marginal fluting is still disputed. Kennedy and Cobban (1976, p. 23; with references) evaluate the functions attributed to the septal marginal fluting. They mention the strengthening of the phragmocone against implosion under hydrostatic pressure (Buckland, 1836). The enlarged surface of the similarly folded posterior part of the body may improve physiological functions (Newell, 1949). The enlarged surface of the folded septa increases the overall weight of the conch that may contribute to buoyancy control (Reyment, 1958). The marginal folding secures body attachment to the shell during the secretion of new septa (Spath, 1919). Saunders (1995) analyzed Paleozoic ammonites suture patterns and raised doubts on Buckland's (1836) hypothesis on the mechanical support by the septal fluting. Alternatively, Saunders (1995) suggests that this marginal fluting forms septal recesses where camera] liquid is stored to improve buoyancy adjustment, However, Buckland's (1836) hypothesis was firmly advocated with mathematical calculations and hydrostatic experiments (e.g., Westermann, 1971, 1975; Hewitt and Westermann, 1986, 1987, 1997) as the functional explanation for septal marginal fluting. Nevertheless, such calculations may be wrongly interpreted. For example, trochospirally coiled gastropods are most resistant to pressure along their coiling axis, which does not mean that this mode of coiling evolved to resist such pressures. In light of Buckland's theory the observed evolutionary increasing complexity in the fluting of the septal margins (sutural complexity) implies a gradual shift of ammonoid activity into the dark, deep marine water, where ammonoid prey is scarce. This contradicts the abundance of ammonites in shelf sediments not as shoreward drifted conchs (Kennedy and Cobban, 1976, p. 36). Actually, most of the ammonoids are regarded epipelagic creatures (up to 250 m water depth), even by the main supporters of Buckland's theory (e.g., Batt, 1991; Westermann, 1990; Westermann and Tsujita, 1999). The marginal folds of the septa strongly bend forward and backward, approaching the inner wall of the ammonite at a low angle. The thickness of a pachydiscid ammonite septum decreases from 0.9 mm in the center to less than 0.3 mm at the margin and to about 0.1 mm in the folioles and lobules (Westermann, 1975; Westermann and Tsujita, 1999). This construction and the marginally thinning of the septa [Westermann, 1975; later suggested to act "like a spring or shock absorber"; (Westermann, 1999b, p. 120)] actually reduces the resistance to external hydrostatic pressure (e.g., Daniel et al., 1997). Therefore, Saunders et al. (1999) suggested an opposing hydrostatic control on ammonoid bathymetric range, whereby more complexly sutured ammonites had shallower depth limits. The marginal fluting of the septa consists mainly of backward pointing tubes in a dendritic pattern that form buttresses resembling columnal arches of Gothic churches. These structures improve the resistance of the ammonite septum to pressure upon its main surface in both adapical (strongest pressure; see Kennedy and Cobban, 1976, p. 24) and adoral directions. Well-preserved ammonites with sutures of variable complexities occur in rather shallow marine carbonate platform sediments, as if suture complexity did not differentiate ammonites to habitats of different bathymetries. For example, the Upper Cenomanian Neolobites with a pseudogoniatitic suture is associated with acanthoceratids and the hoplitid Forbesiceras, which have much more complex sutures (e.g., Lewy and Raab, 1978). The assumption that the fluted margins of the septa significantly improve the resistance of the phragmocone to external pressure implies that all ammonoids lived in or dived into deeper waters than inhabited by nautilids. Mesozoic nautilids thrived in shallow and moderately deep waters like the ammonoids. However, from the Cretaceous onward nautilids weakened relative to more efficient marine predators, and in Late Cenozoic times the surviving nautilids withdrew into deeper waters where competition was minor. Extant nautilids live in a restricted bioprovince, diving during the day to 400-600 m depth (Westermann, 1999a). Comparison of fossil and extant nautilids does not show a significant change in septa thickness or configuration during the last 70 m. y. when the nautilids migrated into greater bathymetries

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and hence hydrostatic pressure. Actually only their physiology changed to adapt to waters with a low oxygen content (O'Dor et al., 1993; Boutilier et al., 1996). Extant nautilids can safely dive to about 600 m depth (off Fiji; Westermann, 1999a). The conchs of the various species can resist hydrostatic pressure before imploding, which correlates to depths of 760-830 m (Kanie et al., 1980; Hewitt and Westermann, 1990; Westermann, 1999a). This maximal depth is suggested to be the limit for the distribution of ammonites, most of which are supposed to have lived in shallower depths (e.g., Westermann, 1993; Westermann and Tsujita, 1999). Consequently the variability in shape and intensity of the folding of ammonite septal margins did not evolve to facilitate ammonoids to adapting to a greater bathymetric range than that which the nautilids could sustain (see Oloriz and Palqvist, 1995; Oloriz et al., 1999; Lehmann, 1981, p. 73). Another function must have been carried out by the marginal folding, which made the ammonoids so successful in the Mesozoic oceans. Among all the functions related to the septal marginal fluting, the one suggested by Spath (1919, p. 30) is emended and elaborated herein as the most reasonable one. Spath (1919) indicated that before forming a new phragmocone chamber, the cochleate cephalopod detaches the muscles from the conch and moves forward. Thereby the connection between the body and the conch is temporarily by the ventral siphuncle and hence very weak. "The folded posterior portion of the ammonite animal with its lobes and saddles afforded the means of holding on, a function performed in Nautilus by the strong central siphuncle." However, when the ammonoid moved forward to secrete a new septum it could not have hold itself onto the folded margins of the former septum. Nevertheless, the present analysis emends Spath's (1919) view by suggesting that the temporary means of body-to-conch attachment operated whenever it was needed in addition to the adductor muscles, regardless of the short-lasting state of muscle detachment. THE MEMBRANE AND TIE-POINTS THEORY Henderson (1984, p. 461) analyzed the complexly folded septa of Pseudophyllites indra (Forbes) and concluded that "they were not related to phragmocone strength but facilitated the attachment of adductor muscles." This was earlier suggested by Blind (1975, 1980; along the mural ridge; more references in Hewitt and Westermann, 1997). "The pattern of suture can therefore be regarded as expressing organization of the musculature; its complexity and diversity among ammonites reflects evolution of the longitudinal muscle system" (Henderson, 1984, p. 475). This means that the posterio-lateral part of the body formed the shape of the septa marginal folds (suture pattern), contradicting the hypothesis that the septum was precipitated on the interface between two fluids (or between the soft tissue and a cameral fluid) of different viscosity (Checa and Garcia-Ruiz, 1996). However, this physical process cannot result in the simplification of septal folding in gerontic conchs. The interaction of pressure of cameral fluid from behind a membrane (or the soft tissue) attached to the inner wall at numerous tie-points was suggested to have formed the complex shape of the template on which the septum was precipitated (Seilacher, 1975). Both the purely physical process (e.g., Checa and Garcia-Ruiz, 1996), as well as the membrane-tie points mechanism (Seilacher, 1975), can imitate goniatitic and phylloid suture patterns with adorally rounded elements, but not ammonitic sutures. Actually the membrane theory of Seilacher (1975) requires the ammonoid to define the relative position of the tie-points for the accurate reproduction of the suture pattern of the mature stage with minor changes throughout further growth of each species. These hypothetical tie-points are supposed to form the acute tips of the lobules while the free part of the membrane inflates into rounded saddlets. However, since this theory needs a control on the position of the tie points, it does not contradict a possible attachment of the membrane along a tie band [e.g., mural ridge (Blind, 1975, 1980); posterior aponeurosis or Haftband (Westermann, 1975)], which would facilitate the creation of all suture patterns. Nevertheless, in both suggested ways of membrane attachment the posterior margins of the mantle should fold into the required configuration of the tie-points or tie-band. MANTLE MUSCULATURE Coleoid cephalopods in general, and octopods in particular, display a great variety of body postures, and skin pattern and coloration for concealment, defense, or courtship (Packard and Sanders, 1969; Forsythe and Hanlon, 1988). The body musculature, which consists of fibers running in all three main directions in space, allows great postural flexibility (flamboyant pattern; Packard and Sanders, 1969; Forsythe and Hanlon, 1988; Hanlon, 1988). The octopod can develop a papillate skin that stiffens for a short period and mimics various structures as a camouflage within its surrounding (e.g., hermatypic coral; Voss, 1971). The skin patterns illustrated in the literature suggest that the posteriomarginal part of the octopod body can stretch along a conch inner wall in the pattern of the ammonoid septa marginal fluting (suture pattern). This kind of musculature was probably already developed in the cochleate ammonoids. The body flexibility of ammonoids was probably like that of extant octopods (e.g., Voss, 1971), and the posterior marginal soft tissue of the ammonoid could penetrate and exit the tiniest features of complexly fluted ammonite septa. It also could form the shape of the fluted septum and stiffen it as a template for the secretion of a new septum. Temporary disturbances in muscle contraction might have formed the inverted suture pattern (Ward and Westermann, 1976), while the weakening of the mantle musculature in the gerontic stage should have caused the simplification of the marginal folding of the few last septa. PSEUDOSUTURES The plasticity of the soft tissue enables the posterior part of the ammonoid to exit through the "bottle neck" of tree-shaped (dendroid omega; Fig. 1) lobes in complexly fluted septal margins, and reshape before the secretion of a new septum. In this case the body moves to its new position in the body chamber in a single step. Ammonites with a rather simple suture pattern with open elements lack the bottle neck feature. The posterior part of the ammonoid can exit the septal marginal folds and move forward in the body chamber while the soft tissue maintains the configuration of the septum. The body creeps forward in the stiffened morphology of the septum, and can temporarily stop for a while, as suggested by

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suture-like striation in between genuine septa in a few ammonite internal casts. The suture-like striations are the imprint of a narrow, slightly elevated, acute band. Perhaps it was the attachment band of the membrane, or the beginning of the secretion of a new septum upon the inner wall of the conch. Accordingly these pseudosutures represent the initial stage of septum secretion during a short pause in the forward movement of the body before it reached the proper position. They occur on certain segments of the ammonite internal casts, whereas other segments lack them (Hewitt et al., 1991; Landman et al., 1993). These incidentally triggered, multiple, unsuccessful attempts of septa secretion seem to reflect a stress in growth and living settings, during which the posterior part of the body is contracted in the shape of the septum and alert for an immediate septum construction. AMMONOID LONGEVITY Bucher et al. (1996) presented a comprehensive review of the literature concerning the longevity of ammonoids. They concluded that ammonoids had a much shorter life span than extant nautilids, which become sexually mature at an age of 10-15 yr and die some five years later. Ammonoids were thought to live a few years after reaching sexual maturity, which in some genera was their terminal growth stage (e.g., Westermann, 1990, p. 465). Bucher et al. (1996) also refer to ammonoid's maturity, which Lewy (1996) suggested that in many cases it represented the breeding stage close before death. According to Bucher et al. (1996, p. 450) most shallow-water ammonoids attained maturity in about five years, though maturity after one or two years as in coleoids cannot be excluded. COMPARISON BETWEEN AMMONOIDS AND OCTOPODS Lewy (1996) demonstrated the similarity in breeding strategies between ammonoids and extant octopods. In many ammonites the terminal body chamber is relatively large, inflated, and with a constricted aperture or apertural appendages. Most heteromorph ammonites have a hooked terminal body chamber. The boat-like floating orientation of these modified body chambers was with the aperture facing upward. It restricted lateral movement, hunting, and most other life functions, especially in the case of a constricted aperture. These modifications at the terminal growth stage, which deteriorated the life quality of the adult creature, were suggested to ensure the development of the brood in a specially constructed egg case (Lewy, 1996). "Several thousand minute ammonite shells were discovered in the body chambers and adhering matrix of 15 mature specimens of Scaphites ferronensis Cobban" (coil. WA. Cobban; Landman, 1985, p. 1). These ammonitellas range from less than 1 mm up to 2 mm in diameter, representing young ammonoids just before and close after hatching (similar Carboniferous ammonitella concentrations analyzed by Tanabe et al., 1995). It seems that the egg-case forming, terminal body chamber of these scaphitid macroconchs sunk to the sea floor with the nearly hatched to just hatched young. Concentrations of tiny spheres less than a millimeter in diameter found in a few ammonites, resemble the tiny argonautid eggs, which the female lays in a specially constructed calcitic egg case precipitated by expansions of two of her arms. The tiny eggs develop while the egg case floats and drifts with the currents (pelagic mode of breeding). The various argonautid egg cases remarkably resemble Upper Cretaceous ammonites, though these structures are complicated to secrete, are highly fragile, and thus do not offer any special security to the young. Lewy (1996) suggested that the similarities between octopods and ammonoids are of phylogenetic significance, and that octopods are nude ammonoids, having lost their conch in Jurassic times. Ammonoids, like the extant octopods, had two modes of breeding: the stationary and the pelagic mode. Some females laid large, yolk-rich eggs in bundles attached to a substrate. The female and the male cared for the breed until (or nearly) the young hatched. During this brood-care extant octopod parents do not eat and usually die of starvation or exhaustion at an age of two to three years (e.g., Heller, 1990). The conchless ancestral argonautids looked for a floating empty ammonite for the pelagic mode of breeding, which they physiologically had to carry out. In each bioprovince they occupied the most common empty conchs, and slightly modified the shape of the body chamber for the egg-case purpose. With time the argonautids may have been able to repair and enlarge conchs by secreting calcite from two expanded arms. These arms secreted a complete egg case in Cenozoic times when the cochleate ammonoids were absent, and no floating conchs (apart of a few nautiloids) were available for the pelagic mode of breeding. The egg case secreted by the Cenozoic argonautids was shaped like Cretaceous ammonites, which their ancestors used to occupy and emend. In spite of the 65 m.y. of evolution under varying ecological settings, all extant argonautid egg cases resemble Upper Cretaceous ammonites. Their complex shape and ornament do not improve the safety of the brood in the fragile egg-case, and seem to reflect phylogenetic relationships between octopods and ammonoids rather than the convergent evolution of their egg cases (Lewy, 1996). These relationships suggest that ammonoids had a rapid growth-rate, as well as a short life span of two to three years (as deduced on the basis of other criteria; Bucher et al., 1996) before starving to death during the brood-care period. The gigantic Entroctopus dofleini (Wulker) reaches an arm span of up to 9.6 m and a weight of 272 kg in about two to three years. Both male and female stop eating and die after the reproductive period, but males may perhaps live one or two years longer if they do not reproduce (Heller, 1990, p. 295; Hartwick, 1983). The large dimensions of this octopod species are comparable to the largest ammonite hitherto found (Upper Cretaceous Parapuzosia seppenradensis (Landois) with a preserved diameter of 1.70 m, reconstructed to about 2.50 m (Summesberger, 1979; Kennedy and Kaplan, 1995). Apart from a few unusually large ammonites, many of an average size that lived for two to four years, precipitated a new septum at intervals of about two weeks to one or two days (e.g., Weitschat and Bandel, 1991). AMMONOID MUSCULATURE The comprehensive study of ammonite muscle scars and imprints of other anatomical features by Jordan (1968) is updated by Doguzhaeva and Mutvei (1996) who recorded all the muscle scars observed on different ammonite species; so not all muscle types actually occur in a single species. Well-preserved Upper Cretaceous pachydiscid ammonites show a pair of dorsal muscle scars of a moderate size, a small ventral scar that is

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associated with a mid-ventral attachment band, and a longitudinal band that probably reflect the attachment of the mantle (Landman et al., 1999). Kennedy et al. (1999) described muscle scars of Baculites. Common are paired dorsal scars immediately in front of the last suture on the dorsal or dorsolateral side of the body. Paired lateral scars are found in some ammonites on each side of the body chamber. A mid-dorsal scar occurs in the internal lobe of the suture. Similarly a single ventral scar occurs in front of the ventral lobe. Apart of the dorsal muscles scars all other single or paired scars are small, and could not avoid the detachment of the body from the conch by a predator. Actually the whole mode of attachment of the ammonoid to its conch is weak compared to the musculature of extant nautilids (e.g., Jordan, 1968). Henderson (1984, p. 475) suggested that "the pattern of suture can therefore be regarded as expressing organization of the musculature; its complexity and diversity among ammonites reflects evolution of the longitudinal muscle system," which is similar to the interpretation of Blind (1975, 1980). The above mentioned short interval between the precipitation of successive septa requires a rapid detachment of the adductor muscles before the ammonoid moves forward, and their rapid reattachment at the new position (discussed by Henderson, 1984, p. 472). The small scars of the adductor muscles observed on ammonites (Jordan, 1968; Doguzhaeva and Mutvei, 1996) reflect the small dimensions of these muscles, which, therefore, could be rapidly detached and reattached to the conch. However, these narrow muscles could not efficiently function as adductor muscles to anchor the predating creature to its buoyant conch. THE ANCHORAGE FUNCTION OF THE SEPTA MARGINAL FOLDS The backward stretching and branching septal folds are confined to a narrow marginal zone of the septa. The dendritic pattern of these folds, as represented by the lobes of the suture, has an omega-like (12) shape with a narrow opening, which branches into numerous tubes with closed tips (Fig. 1). It is suggested that the soft tissue of the posterior margins of the ammonoid mantle (compared to that of an octopod) could penetrate into these branching tubes along the inner periphery of the conch, and stiffen there. The backward expanding dendritic shape turned the stiffened tissue into strong anchors (Fig. 1) in the number of the sutural lobes. Thereby the whole circumference of the body firmly held to the conch, even stronger than by the pair of large mid-lateral adductors of extant nautilids. The septal marginal fluting has a bilateral symmetry, whereby between correlative sutural lobes on both flanks the septum is adorally convex in a perfect arch (Fig. 2). Combined with the thickening of the septum from its margins to the center as described from a Cretaceous Pachydiscus (Westermann, 1975), this structure resists pressures on its surface in an adapical (backward) direction (Fig. 2). Such a force is comparable to a dragging force between the buoyant conch and the ammonoid anchored to the latter along the septa marginal folds. The lobules in complexly marginal fluting branch and expand to form deep lobes. The septal wall of these lobules joins the inner wall of the conch at a low angle, whereby it cannot support the resistance of the phragmocone wall to withstand external pressure (Fig. 2). However, the more complexly the marginal fluting, the better is the anchorage of the soft tissue to the buoyant conch. The resulting adorally arching of the septa, combined with their thickening toward the center, together improve the resistance of the ammonoid to strong dragging forces between the organism and its buoyant conch. Such forces are inflicted on the septum when the ammonoid struggles with prey while the stiffened soft tissue is anchored in the septal marginal folds. Since the development of the omega-shaped anchoring site in the Ammonoidea, they improved their anchoring function by further branching and expanding while the opening of the omega narrowed. However, this advantageous evolutionary trend seems to have gone beyond the functional needs as exemplified by the highly complex sutures of many Middle and Upper Cretaceous Desmoceratacea (Fig. 1). Nevertheless, among them are the largest ammonites such as the lower Lower Campanian Parapuzosia seppenradensis (Landois) reconstructed to nearly 2.50 m in diameter (Summesberger, 1979; Kennedy and Kaplan, 1995), P. bradyi Miller and Youngquist (1946) 1.80 m, and P. bosei Scott and Moore (1928) 1.44 m in diameter. The related pachydiscids attain large dimensions as well, and are most common in outer shelf and pelagic sediments. It suggests that these ammonoids lived and successfully competed with other predators in the open marine settings, eating well to rapidly grow in a short time to gigantic dimensions (compared to the extant Enteroctopus dofleini). The more complicated the marginal fluting of the ammonite septum, the stronger the anchorage of the animal to the buoyant conch, and hence the ability of the creature to cope with larger and stronger prey. The complexity of the suture pattern reflects the aggressiveness of the ammonoid, and not its capability to resist hydrostatic pressure. Therefore, ammonites with sutures of different complexities co-occur. This mode of anchorage of the body to the buoyant conch operated only when needed. It substituted for the adductor muscles, which their position and small size could not resist the strong dragging forces. The shape of the saddles of the suture result from that of the lobes with some modification of the central part of the saddle, which forms a small anchoring point in between the lobes. The soft tissue of the posterior part of the ammonoid not only penetrated and exited the septal marginal folds, but actually formed the shape of these complex folds for the precipitation of a new septum. The septal configuration in two internal molds of the Upper Campanian Libycoceras is slightly shifted towards the umbilicus, whereby the umbilical flank of each lobe overlay the adjacent saddle (Fig. 3). This phenomenon is viewed from two sides of the septum. The specimen in Figure 3.1, 3.2 shows a frontal view, and another specimen (Fig. 3.3, 3.4) looks on the posterior septal surface of the internal mold. The seemingly truncated tips of some lobes (Fig. 3.4) are well seen on the posterior surface of the septum (Fig. 3.3), where the tips of the lobes are deflected over the former septum without any truncation or deformation (see Holder, 1952). This means that the septum is neither shaped under physical stresses (e.g., Checa and Garcia-Ruiz, 1996), nor precipitated on a membrane connected to tie-points (e.g., Seilacher, 1975), all of which would have contorted the delicate features of the sutural pattern. Consequently the complexly shaped septum is precipitated by the modified posterio-marginal soft tissue, which can bend without changing the shape of the delicate folds. CONCLUSION Ammonoids grew rapidly, frequently secreting a new septum, which required the detachment and reattachment of the adductor muscles every

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few days. Therefore, these muscles were small and restricted, and thus useless as a firm hold between the animal and the conch. The marginal folds of ammonite septa form backward branching and expanding tubes, which the soft tissue can enter and temporarily stiffen there in order to hold firm to the conch. Thus the backward expanding marginal folds of the septa provide circumferential anchorage sites that firmly hold the body to the buoyant conch in addition to the few, small adductor muscles. These folds do not improve the resistance of the phragmocone to external pressure as previously advocated, but actually resist the adoral dragging force, which is inflicted between the body of the predating ammonoid and the buoyant conch. The complexity of the marginal folding reflects the strength of the connection between the body and the conch, and thus the aggressiveness of the ammonoid (in contrast to Lehmann, 1981, p. 75), though its evolution may have exceeded its functional needs. ACKNOWLEDGEMENTS The suture line in Figure 1 is based on a drawing published by Cobban and Kennedy (1993) in U.S. Geological Survey Professional Paper 1533, for which I am thankful. R. Beckman, N. Shragai, M. Dvorchek, and Y Levy are thanked for their technical assistance. REFERENCES BATT, R. J. 1991. Sutural amplitude of ammonite shell as a paleoenvironmental indicator. Lethaia, 24:219-225. BLIND, W. 1975. Ober die Entstehung and Funktion der Lobenlinie bei Ammonoideen. Palaontologische Zeitschrift, 49:254-267. BLIND, W. 1980. Ober Anlage and Ausformung von Cephalopoden-Septen. Neues Jahrbuch fur Geologie and Palaontologie, Abhandlungen, 160:217-240. BOUTILIER, R. G., T G. WEST, G. H. POGSON, K. A. MESA, J. WELLS, AND M. J. WELLS. 1996. Nautilus and the art of metabolic maintenance. Nature, 382:534-536. BUCHER, H., N. H. LANDMAN, S. M. KLOFAK, AND J. GUEX. 1996. Mode and rate of growth in ammonoids, p. 407-461. In N. H. Landman, K. Tanabe, and R. A. Davis (eds.), Ammonoid Paleobiology: Topics in Geobiology 13. Plenum Press, New York. BUCKLAND, W. 1836. Geology and mineralogy considered with reference to natural theology. William Pickering, London, volume 1, 599 p.; volume 2, 128 p. CHECA, A. G., AND J. M. GARCIA-RUIZ. 1996. Morphogenesis of the septum in ammonoids, p. 253-296. In N. H. Landman, K. Tanabe, and R. A. Davis (eds.), Ammonoid Paleobiology. Topics in Geobiology 13. Plenum Press, New York. COB BAN, W. A., AND W. J. KENNEDY. 1993. The Upper Cretaceous dimorphic pachydiscid ammonite Menuites in the Western Interior of the United States. U.S. Geological Survey Professional Paper 1533, 14 p. DANIEL, T. L., B. S. HELMUTH, W. B. SAUNDERS, AND P. D. WARD. 1997. Septal complexity in ammonoid cephalopods increased mechanical risk and limited depth. Paleobiology, 23:470-481. Doc,UZHAEVA, L., AND H. MUTVEI. 1996. Attachment of the body to the shell in ammonoids, p. 43-63. In N. H. Landman, K. Tanabe, and R. A. Davis (eds.), Ammonoid Paleobiology. Topics in Geobiology 13. Plenum Press, New York. FORSYTHE, J. W., AND R. T HANLON. 1988. Behavior, body patterning and reproductive biology of Octopus bimaculoides from California. Malacologia, 29:41-55. HANLON, R. T. 1988. Behavioral and body patterning characters useful in taxonomy and field identification of cephalopods. Malacologia, 29: 247-264. HARTWICK, B. 1983. Octopus dofleini, p. 277-291. In P R. Boyle (ed.), Cephalopod Life Cycles. Academic Press, Oxford. HELLER, J. 1990. Longevity in molluscs. Malacologia, 31:259-295. HENDERSON, R. A. 1984. A muscle attachment proposal for septal function in Mesozoic ammonites. Palaeontology, 27:461-486. HEWITT, R. A., AND G. E. G. WESTERMANN. 1986. Function of complexly fluted septa in ammonoid shells, L, Mechanical principles and functional models. Neues Jahrbuch fur Geologie and Palaontologie, Abhandlungen, 172:47-69. HEWITT, R. A., AND G. E. G. WESTERMANN. 1987. Function of complexly fluted septa in ammonoid shells. II. Septal evolution and conclusions. Neues Jahrbuch fair Geologic and Palaontologie, Abhandlungen, 174:135-169. HEWITT, R. A., AND G. E. G. WESTERMANN. 1990. Nautilus shell strength variance as an indicator of habitat depth limits. Neues Jahrbuch fur Geologie and Palaontologie, Abhandlungen, 179:71-95. HEWITT, R. A., AND G. E. G. WESTERMANN. 1997. Mechanical significance of ammonoid septa with complex sutures. Lethaia, 30:205-212. HEWITT, R. A., A. CHECA, G. E. G. WESTERMANN, AND P. M. ZABORSKI. 1991. Chamber growth in ammonites inferred from colour markings and naturally etched surfaces of Cretaceous vascoceratids from Nigeria.

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