[Palaeontology, Vol. 50, Part 6, 2007, pp.

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AMMONOID SHELL STRUCTURES OF PRIMARY ORGANIC COMPOSITION

HWILER* , DIETER KORN , by CHRISTIAN KLUG* , THOMAS BRU NTER SCHWEIGERT , ARNAUD BRAYARD* and JOHN TILSLEY GU
ontologisches Institut und Museum der Universita t Zu rich, Karl Schmid-Str. 4, CH-8006 Zu rich, Switzerland; *Pala e-mails: chklug@pim.uzh.ch; bruehwiler@pim.uzh.ch; abrayard@pim.uzh.ch r Naturkunde der Humboldt-Universita t zu Berlin, Invalidenstr. 43, D-10115 Berlin, Germany; e-mail: dieter.korn@museum.hu-berlin.de Museum fu r Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany; e-mail: schweigert.smns@naturkundemuseum-bw.de Staatliches Museum fu oenvironnements et Pale obiosphe ` re, Universite Claude Bernard Lyon 1, 2 rue Dubois, F-69622 Villeurbanne Cedex, France; UMR-CNRS 5125, Pale e-mail: arnaud.brayard@University-lyon1.fr Ansell Road, Shefeld S11 7PE, UK; e-mail: tilsley@tiscali.co.uk Typescript received 18 May 2006; accepted in revised form 30 November 2006

Abstract: Palaeozoic

and Mesozoic cephalopod conchs occasionally reveal dark organic coatings at the aperture. A number of these coatings, including still unrecorded examples, are described, gured and interpreted herein. On the basis of elemental analysis, actualistic comparison and a comparison with Triassic bivalves, some of these coatings are shown to consist of apatite and primarily probably of conchiolin (and also probably melanin). In several Mesozoic ammonoid genera such as Paranannites, Psiloceras, Lytoceras, Phylloceras, Harpoceras and Chondroceras, some of these coatings (recorded herein for most of these taxa for the rst time) are interpreted as a structure similar to the black band, which was previously known only from Recent Allonautilus and Nautilus. In contrast to these nautilid genera, however, the organic material of some Mesozoic ammonoids was not deposited on the inside of the shell but externally, albeit positioned at the terminal aperture as in Recent nautilids. Some ammonoids of Carboniferous and Triassic age show

several such bands at more or less regular angular distances on the ultimate whorls and at the aperture, e.g. Nomismoceras, Gatherites, Owenites, Paranannites, Juvenites and Melagathiceratidae gen. et sp. nov. Triassic material from Oman shows that the black coating was probably secreted from the inside, because the position of this organic deposit changes from interior to exterior in an anterior direction (i.e. adaperturally). This structure has previously been referred to as a false colour pattern and is here interpreted as having been formed at an interim aperture or megastria (alter Mundrand). All structures discussed in the paper are considered to have been secreted by a single organ and to have been initiated by some form of stress or adverse conditions. Thus, certain environmental parameters and growth anomalies appear to have inuenced their formation.
Key words: Ammonoidea,

mature modications, body

chamber, growth, taphonomy.

Among fossil invertebrates, molluscs probably represent the most useful group for studying ontogeny and growth. Growth changes are recorded in most molluscan shells and usually are preserved even on internal moulds. In some cases, these growth changes permit interpretations with respect to changes in mode of life and habitat (e.g. da 2003; Klug and Korn 2004). A fascinat tzel and Fry Nu ing topic is the morphogenetic countdown (Seilacher and Gunji 1993; Seilacher and LaBarbera 1995) that has been postulated for some ammonoids. Terminal growth of ammonoids is remarkable because it is occasionally associated with drastic changes in shell morphology (e.g. the ultimate whorl of many heteromorph taxa; see Davis et al. 1996 and references therein). Among Recent nautilids (compare Collins and Ward 1987; Ward 1987; Text-g. 1), mature specimens show

(1) a shell growth band (shell thickening at the apertural edge, up to 25 mm wide and 1 mm thick); (2) a black band at the aperture (this character is not expressed in all adults according to Ward 1987); (3) a deepening of the ocular sinuses; (4) a reduction of whorl height by a decrease in whorl expansion rate; (5) a reduction of whorl width by a decrease in whorl width expansion rate and a simultaneous formation of a more rounded venter; (6) septal thickening (the last septum is up to 30 per cent thicker than the penultimate septum); (7) septal crowding; (8) a distinct maximum shell diameter (a poor character because of variability); (9) a white ventral area; (10) an increase in body chamber length (caused by narrowing of the whorl section); and (11) a reduction of cameral liquid (probably to compensate for additional shell material at the aperture and a longer body chamber). Among

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these modications, septal crowding is probably the most widely known phenomenon that has also been described for many fossil chambered ectocochleate cephalopods. Except for 9 and 11, all the phenomena listed (although often slightly differing in one aspect or another) are also known in ammonoids (compare Sun 1928; Davis 1972; Davis et al. 1996). Some of these, however, are quite rare. For example, questionable fossil remains of the black band have, to our knowledge, been documented just once (Klug 2004). For it to be fossilized, organic matter needs to be preserved because originally it consisted of conchiolin and melanin. Conchiolin (or conchin) is made up of a chitinous albuminoid and, as such, has a moderately low preservation potential (Allison 1988; Briggs et al. 1993; Kear et al. 1995; Briggs and Wilby 1996). In specimens described by Klug (2004), remains of the black band and black aperture had transformed to apatite. The black band was rst described for Recent nautilids (Saunders and Spinosa 1978; Doguzhaeva and Mutvei 1986; Collins and Ward 1987; Ward 1987; Mutvei et al. 1993; Mutvei and Doguzhaeva 1997). It is usually represented by a dark line of varying width (15 mm) and thickness (< 1 mm) that surrounds the inside of the aperture in mature specimens. In most of the ammonoids described below, this dark material apparently lies on the outside of the conch, leading to the question as to whether this structure is really the same as in Recent nautilids. Here we describe and illustrate some examples of ammonoids from the Carboniferous of Great Britain, the

Triassic of Oman, Nevada, South China and Germany, and the Jurassic of Germany that preserve black bands and related structures. The origin and formation of these structures are discussed and interpreted.

TERMINOLOGY
The black band The black band (Saunders and Spinosa 1978; Collins and Ward 1987; Ward 1987; black border of Stenzel 1964) is a structure at the terminal aperture in Recent nautilids consisting of conchiolin and melanin (Comfort 1950). It surrounds the adult aperture in a band of varying width and thickness and passes gradually into the black layer (Stenzel 1964). Although most of the organic matter is found in the interior of nautilids and externally in several ammonoids, this structure is here referred to as the black band for simplicity. In Recent nautilids, it does not display the dark coating in all adult specimens (Ward 1987), which might in part account for the fact that this structure is rare among ammonoids. We may speculate that a combination of terminal growth and environmental stress is required for the formation of this structure. The function of the mantle adhesive layer in Recent nautilids has been discussed by Mutvei et al. (1993) and Mutvei and Doguzhaeva (1997). According to these authors, the anterior mantle margin was attached to the apertural mantle attachment layer with vertical pores (Mutvei and Doguzhaeva 1997, text-g. 10); the latter contained nger-like epithelial extensions. Apparently, the apertural mantle attachment is rather weak in Recent nautilids and appeared to function in preventing water from entering between the mantle and shell, particularly during swimming (Mutvei et al. 1993, p. 11).

The black layer The black layer (Ward 1987; Keupp 2000; Kulicki et al. 2001; Klug and Lehmkuhl 2004; Klug et al. 2004; black lm of Stenzel 1964 and black deposit of Mutvei and Doguzhaeva 1997) is an organic part of the dorsal shell. In Recent nautilids and in some ammonoids, it projects dorsally out of the body chamber. Behind the aperture, increasingly thick layers of lustrous shell cover the organic material, causing an increasingly light colour. In Recent nautilids, the mantle margin secretes both the black layer and the black band. According to Pruvot-Fol (1937), these black structures represent excretions of metabolic waste like cephalopod ink which all contain melanins (Comfort 1950). This fact also questions the interpreta-

TEXT-FIG. 1.

Nautilus pompilius (Linnaeus, 1758), in section PIMUZ 7806; Recent; locality unknown; with black band rich). (photograph Heinz Lanz, Zu

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tion of organic matter in the body chamber of a Triassic ammonoid as fossil ink by Doguzhaeva et al. (2004). This material might equally represent remains of the black layer. Adverse life conditions as well as injuries may also cause deposition of conchiolin, darkening the conch more or less locally or at an interim or terminal aperture (Landman and Cochran 1987; Saunders et al. 1987; Ward 1987; Keupp and Riedel 1995; Rein 2000, 2002; Klug and Lehmkuhl 2004). On the basis of shape, Klug (2004) referred to these structures as black stripes (a spiral stripe caused by a punctiform injury) or black apertures (supercially resembling a black band, but much wider).

True colour patterns Colour patterns have been recorded from numerous fossil cephalopod shells including various Palaeozoic nautiloids and CarboniferousCretaceous ammonoids (Mapes and Davis 1996). Presumably, these patterns consisted of pigments (mainly melanin and porphyrins) stored in either the periostracum or the outer part of the shell as in Recent molluscs (Hollingworth and Baker 1991). These structures are not discussed herein as numerous articles on this subject are available (for references, see Mapes and Davis 1996).

The black aperture The black aperture (Klug 2004) is a black line of varying width formed at an interim aperture caused by adverse conditions or injuries (see, e.g. Keupp and Riedel 1995). Like the black band, it encircles the aperture, yet not necessarily the terminal one, and also consists of conchiolin.

Siphuncle As the connecting rings of ammonoids are also of organic composition, these have to be listed here, too. This part has also been examined and discussed frequently (for a review and references, see Tanabe and Landman 1996). Remarkably, details of soft-tissues within the siphuncle, including blood vessels, have been discovered and described by Tanabe et al. (2000).
Institutional abbreviations: MHI, Muschelkalkmuseum Hagdorn, ontologisches Institut und Museum, Ingelngen; PIMUZ, Pala rich University; SMNS, Staatliches Museum fu r Naturkunde, Zu Stuttgart; Zx, British Geological Survey, Nottingham.

The black stripe The black stripe (Klug 2004) is a black spiral band, starting at a minor shell injury which occurred at a former apertural edge. For this structure, the terms Rippenscheitelung, forma verticata, forma pseudocarinata and forma semiverticata have been introduced for ammo lder 1956, 1977; Keupp 1979, 2000; Hengsbach noids (Ho 1996), although the black deposits that occasionally accompany these injuries are rarely preserved. These structures represent the regenerative reaction of molluscs to a punctiform injury of the apertural epithelium, often caused by arthropod attacks (see Keupp 2000 and references therein). It is very likely that at least some of these healed injuries were originally accompanied by a black line now no longer preserved.

DESCRIPTION OF SPECIMENS
Carboniferous Carboniferous strata have yielded surprisingly well-preserved ammonoid specimens which often show highly unusual details. For instance, the rst radulae to be discovered were in specimens of Eoasianites from the Carboniferous of Uruguay (Closs and Gordon 1966; Closs 1967; Bandel 1988) and the oldest aragonitic goniatitids come from the Pennsylvanian (Desmoinesian) Buckhorn Shale of Oklahoma (Smith 1938; Kulicki et al. 2002). Two specimens of Nomismoceras vittiger (Phillips, 1836) with coarsely recrystallized shells of Asbian (Early Carboniferous) age from Derbyshire are available (Zx2212, 2288; Text-g. 2), measuring 19 and 26 mm in diameter and collected at Treak Cliff near Castleton (Korn and Tilsley 2006). Both display roughly equidistant dark lines that are clearly symmetric, and which can be traced back on two to three younger whorls in the umbilicus. In both specimens, the dark bands lie within zones of thickened shell. It is important to note that the anterior edge of each of these dark bands is more clearly dened than the posterior one. This is because shell

False colour patterns False colour patterns (Mapes and Davis 1996) include more or less regular transverse patterns that can be associated with constrictions, megastriae and pseudoconstrictions, as well as with spiral lines formed by the umbilical seam of a missing subsequent whorl which cross the false colour patterns. Normally, they are parallel to growthrelated structures such as those listed above (see Mapes and Sneck 1987). Usually, more than one such line occurs within a single whorl.

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thickness between the black material and shell exterior increases posteriorly. About two to three such bands can be seen per whorl; they form a little projection on the umbilical edge, followed by a strongly asymmetrical sinus and an adjacent parabolic ventrolateral projection. On the venter, the bands form moderately deep U-shaped sinuses. EDAX analyses have yielded no data on the origin of the black material; possibly it is still covered by a very thin layer of calcite.

Triassic Remarkably, Triassic deposits worldwide have yielded ammonoids with organic remains preserved. Much that is currently known about ammonoid palaeobiology was gained from Triassic material. One of the most spectacular examples are the phosphatized ammonoids from Spitsbergen which preserve, in addition to delicate remains of septa, signicant portions of the digestive tract, beaks with radula and even gill remains (Lehmann and Weitschat 1973; Lehmann 1985; Weitschat 1986; Weitschat and Bandel 1991, 1992). Diversity, and preservation of colour patterns and black layers in part account for the fame of Triassic ammonoids from Fossil Hill or Crittenden Springs in Nevada (Smith 1932; Kummel and Steele 1962; Mapes and Davis 1996; Keupp 2000). Some specimens collected there also show the black band. In Oman, Early Triassic ammonoids occur in exotic blocks of Hallstatt facies in deep-water strata of the Hawasina unit (Blendinger 1991, 1995; Tozer and Calon 1990). Although most of these are difcult to extract and prepare, preservation is sometimes excellent. In rare cases, remains of false colour

patterns have been found. In contrast, Early Triassic ammonoids from South China are numerous and easily collected. However, so far no colour patterns have been documented, except for some rare, recently discovered, specimens that display intriguing false colour patterns and other kinds of black structures. Finally, ammonoids from the Muschelkalk (Middle Triassic) of the Germanic Basin have received some attention, not just in view of their utility as stratigraphical tools but also for their supercially poor preservation, which make them interesting study objects for taphonomic analyses (e.g. Kumm 1927; Seilacher 1963, 1966, 1968, 1971; Mayer 1968; Mundlos 1970; Aigner 1975; Duringer 1982; Hagdorn and Mundlos 1983; Mundlos and Urlichs 1990; Maeda and Seilacher 1996; Klug 2001; Zeeh and Hagdorn 2002; Klug et al. 2004, 2005a, b). These ammonoids have not been fully appreciated previously, which is demonstrated by the fact that in recent years, several specimens that preserve originally organic structures such as the black layer, the black band, black apertures, organic membranes in the phragmocone and elsewhere have been found and described (Rein 1993, 1995, 2005; Klug 2004; Klug et al. 2004, in press). These are structures that are rarely preserved otherwise (e.g. Weitschat and Bandel 1992). From the Early Triassic Thaynes Formation of Nevada, USA, two well-preserved specimens of Paranannites slossi (Kummel and Steele, 1962) (Jenks Coll. nos. 269C, 499C) and one of Juvenites septentrionalis Smith, 1932 (Jenks Coll. no. 499C; see Text-g. 3) were put at our disposal by J. Jenks (Salt Lake City), who collected them from the Euemingites romunderi Zone at Crittenden Springs (Elko County, Nevada). The Paranannites specimens measure 27 mm (No. 499C) and 31 mm

C A B

TEXT-FIG. 2.

The Early Carboniferous ammonoid Nomismoceras vittiger (Phillips, 1836) from Treak Cliff near Castleton, Derbyshire (UK). A, Zx2212. BC, Zx2288. Note the black lines on the outer whorls visible at irregular distances.

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(No. 269C) in diameter. All three retain calcitic replacement shell and display more or less regular radial, dark bands. On account of the densely spaced growth lines and apertural shell thickening, the two specimens of Paranannites were apparently adult. These two also have remains of the black layer and wrinkle layer (Runzelschicht; Walliser 1970; Bayer 1974). One of these specimens (499C) shows a darkened terminal aperture (Text-g. 3A). Well-preserved Early Triassic ammonoids are also available from Oman from an exotic block of Hallstatt facies at Wadi Musjah (75 km SSW of Musqat). Two specimens of Paranannites sp. with false colour patterns (Textg. 4AE) were collected from a bed also yielding Owenites koeneni Hyatt and Smith, 1905 and Inyoites oweni Hyatt and Smith, 1905 of Smithian age. They measure 19 mm (PIMUZ 26262) and 20 mm (PIMUZ 26263) in diameter. Because of the relatively dense spacing of growth lines and constrictions near the aperture, PIMUZ 26262 was apparently adult; most representatives of this species are roughly the same size and thus most of them were probably adult too (including PIMUZ 26263). Both specimens have remains of the wrinkle layer. They show several black lines that extend symmetrically from the umbilical shoulders around the venter and coincide with shell thickenings posterior of constrictions. The width of these lines does not increase towards the venter but remains constant. The anterior edge of the lines follows the shell constrictions. The posterior edge is more or less parallel to the anterior one and therefore crosses growth lines. As in the Carboniferous material, the anterior edge of these dark bands is more clearly delimited than the posterior edge. Posterior of the constrictions, the dark material is located internally in the thickened shell, and the outermost shell layer is not coloured (Text-g. 4E). Towards the aperture, the dark material crosses the shell from within and surfaces so that the entire shell is coloured at the constrictions. A well-preserved Early Triassic representative of the Melagathiceratidae (gen. et sp. nov.; PIMUZ 25900) from Jinya (Guangxi, South China; Flemingites rursiradiatus beds) also displays intriguing remains of dark material (Text-g. 4FJ). Interpretations of these are ambiguous. Indeed, the specimen clearly displays symmetrical black lines extending from the umbilicus to the venter, coinciding with constrictions as in the Omani specimens of Paranannites. The dark material of the lines is also concentrated only at the constrictions. At the aperture, probable equivalents of the black band and black layer are visible on the inside of the shell. A more questionable dark pattern is seen on the umbilical wall, which generates a black spiral line. It is difcult to identify the origin of this structure, which rather appears as a dark calcitic crust covering the umbilical wall.

Only three cephalopod specimens from the German rttemberg Upper Muschelkalk of northern Baden-Wu (southern Germany) that display remains of the black band or black aperture are available. One of these (SMNS 65424) is a specimen of Ceratites postspinosus Riedel, 1916 measuring 139 mm in diameter and thus more or less adult, which is corroborated by a slight increase in umbilical width charn acteristic of adult Ceratites. It was collected from the Scho and Hippelein quarry at Neidenfels from Upper Muschelkalk marls (postspinosus Zone, Ladinian). This specimen displays irregular and poorly preserved black remains, which extend radially over 48 mm and spirally over 176 mm. As in the following specimen, the black material does not surround the entire aperture, demonstrating incomplete preservation, partial sediment lling only and pressure solution. This and the next two specimens have previously been discussed by Klug (2004). Ceratites meissnerianus Penndorf, 1951 (SMNS 25397-3; leg. A. Lehmkuhl and M. Urlichs) shows remains of the black layer. Collected from a quarry near Unterohrn rttemberg, Germany) in the Upper (northern Baden-Wu Muschelkalk (semipartitus Zone, Ladinian), it probably represents a more or less adult individual; this is indicated by its large diameter (309 mm), septal crowding and the eccentric shape of the umbilicus. At its aperture, it preserves black remains that measure over 52 mm radially and 33 mm in maximum width. These black remains do not surround the whole aperture and thus probably represent only a small portion of the entire structure, which suffered from pressure solution as well as mechanical wear and breakage. As seen in the black band of Recent Nautilus, these dark layers become lighter coloured posteriorly (adapically). A specimen of the nautilid Germanonautilus bidorsatus (von Schlotheim, 1820) showing a black line at a former aperture was collected by H. Hagdorn at a quarry near rttemberg; compressus Garnberg (northern Baden-Wu Zone, Anisian; MHI 919). This specimen was described and gured by Klug and Lehmkuhl (2004; it displays the large muscle attachment scar of the cephalic retractor as well as faint traces of the black layer). It is mentioned here for comparison because it preserves a narrow black line surrounding a former aperture (not a megastria or alter Mundrand; see also the discussion of false colour patterns below). Judging by the abnormal depth of the hyponomic sinus, this aperture probably formed following injury. Additionally, it continued to grow after the formation of the black line and thus, this was not the terminal aperture. Growth increments within the hyponomic sinus, however, display a faint but widely spaced ribbing probably reecting a high growth rate. The data obtained from EDAX analyses performed on the black substance of the black line were quite interesting. Both silica and probably apatite occur (Klug 2004), which is additionally corrobo-

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A D

KLUG ET AL.: AMMONOID SHELL STRUCTURES

T E X T - F I G . 4 . Early Triassic ammonoids from Oman and South China with false colour patterns and other possible organic remains. AE, Paranannites sp., Owenites koeneni beds (Smithian), from Wadi Musjah (75 km SSW of Musqat, Oman). AB, PIMUZ 26262, lateral and ventral views. CE, PIMUZ 26263. C, lateral view. D, ventral view. E, detail of D showing crosssection of shell with two dark stripes at constrictions; note that the dark material crosses the shell in an anterior direction. FJ, Melagathiceratidae gen. et sp. nov. (PIMUZ 25900), Flemingites rursiradiatus beds (Smithian), Jinya (Guangxi, South China). F, lateral view. GH, apertural view. IJ, view of umbo showing black material on umbilical wall forming a spiral; note that Paranannites clearly displays false colour patterns; in contrast, dark remains of Melagathiceratidae gen. et sp. nov. are more difcult to interpret in terms of organic deposits.

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C B

rated by comparable analyses of a coating of ceratitid muscle attachment structures which also revealed apatite (Klug et al. in press).

Jurassic A number of Jurassic ammonoids from Germany, Great Britain and Switzerland display black lines at the aper-

tures. Most of these are of Early Jurassic age, which is not surprising as the conditions of preservation were occasionally excellent at this stratigraphic interval, for instance tten (for reviews, see Bottjer et al. in some Fossillagersta 2002 and Selden and Nudds 2004). 1. The oldest material is of the index of the basal ammonoid zone of the Early Jurassic, Psiloceras planorbis (J. de C. Sowerby, 1824). Two crushed specimens on a slab of claystone from early Hettangian sediments at Blue Anchor

T E X T - F I G . 3 . Triassic ammonoids from southern Germany and the USA with black band and false colour patterns. AC, Paranannites slossi (Kummel and Steele, 1962) and Juvenites septentrionalis Smith, 1932 (Jenks Coll. no. 499C); Euemingites romunderi Zone (Thaynes Formation, Early Triassic), Crittenden Springs (Elko County, Nevada). A, detail of C, aperture with black band, dorsal shell and false colour patterns. B, lateral view. C, ventral and apertural view. D, Ceratites postspinosus Riedel, 1916 (SMNS 65424); rttemberg; note remains of the black band postspinosus Zone (Ladinian, Upper Muschelkalk), Neidenfels, northern Baden-Wu bingen); modied after Klug (2004). EF, Ceratites meissnerianus (Penndorf, 1951) (SMNS 25397-3); (photograph Wolfgang Gerber, Tu rttemberg; note remains of a questionable black semipartitus Zone (Ladinian, Upper Muschelkalk); Unterohrn, northern Baden-Wu band and septal crowding. E, detail of F showing remains of the black band. F, lateral view of entire specimen with the black band bingen). DF modied after Klug (2004). (photograph Wolfgang Gerber, Tu

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(Watchet, Somerset, UK; PIMUZ 6519; Text-g. 5) were rich); one is nearly complete collected by G. Kahn (Zu and measures 49 mm in diameter. Both preserve the aragonitic shell and the terminal aperture. Anterior to the aperture, the black layer is clearly visible on the venter of the penultimate whorl of specimen A. Additionally, the other specimen (B) shows a trace of the black layer at the umbilical seam. The most interesting structure, however, is the black band, which is well preserved in both specimens. Its colour is slightly lighter than that of the black layer and its position is 12 mm behind the aperture in PIMUZ 6519 (leaving a narrow white stripe) and directly at the aperture in other specimens (e.g. PIMUZ 12581; same locality and strata). This structure was found in many specimens from this locality. A poorly preserved individual that also showed this bingen black band was subjected to EDAX analyses at Tu University. These element analyses, however, did not yield any sign of preservation of originally organic material. Elements identied indicate the presence of calcium carbonate and clay minerals. 2. An additional lot of three specimens collected from the Posidonienschiefer Formation (Posidonia Shale, Toarcian, Early Jurassic) of southern Germany are on display in museums, which is why we did not take samples for elemental analyses. Phosphatized, originally organic, structures are well known from this stratigraphic unit; thus, we assume the black structures seen in these specimens to be phosphatized as well. This conclusion is supported by the similar appearance of associated, primarily chitinous, ammonoid beaks.

The rst specimen, identied as Phylloceras (Phylloceras) heterophyllum (J. Sowerby, 1820), and collected at Ohmden from the Fleins Bed (tenuicostatum Zone, semicelatum Subzone), is on exhibit at Stuttgart (SMNS 26462; Textg. 6). It measures 87 cm in diameter and was thus most likely adult at death, which is also evident from coarser growth lines, large size and presence of the black band. At its terminal aperture, it displays a black line of varying width (almost 10 mm). In addition to this, it preserves the lower and upper beak in the body chamber. This is of interest because the beaks and the black band apparently show the same type of preservation, indicating a similar, originally chitinous composition. Usually, only the periostracum of the ammonoid shell was left, as demonstrated by Seilacher et al. (1976). From this it might be concluded that the black band was part of the periostracum, and thus probably external as in specimens from other localities. Another large, and most likely adult, specimen (SMNS 26465; Text-g. 7) from the Posidonienschiefer Formation, assigned to Lytoceras ceratophagum (Quenstedt, 1885), is also from Ohmden, having been collected from the Unterer Schiefer Bed (falciferum Zone, elegantulum Subzone). It measures 41 cm in diameter, and shows the black line very clearly, delimited posteriorly by an alter Mundrand (megastria) and anteriorly by the terminal aperture. The distance between these two is 30 mm; nearly 20 irregular growth lines or lirae are seen therein. This black band was probably part of the periostracum. The third specimen is an adult Harpoceras (Harpoceras) falciferum (J. Sowerby, 1820) from Holzmaden (falciferumbifrons zone, falciferumlower commune subzones),

T E X T - F I G . 5 . Psiloceras planorbis (J. de C. Sowerby, 1824) (PIMUZ 6519); planorbis Subzone (planorbis Zone, Hettangian, Early Jurassic), Blue Anchor (Watchet, Somerset, UK). Note the black band, black layer and the trace it leaves at the umbilical seam. rich). Natural size (photograph Thomas Galfetti, Zu

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TEXT-FIG. 6.

Phylloceras (Phylloceras) heterophyllum (J. Sowerby, 1820), SMNS 26462, Posidonienschiefer Formation (Fleins Bed, rttemberg, Germany). Note the upper and tenuicostatum Zone, semicelatum Subzone; Toarcian, Early Jurassic), Ohmden (Baden-Wu lower beak in the body chamber and the black band.

housed at the Urweltmuseum Hauff and previously gured by Selden and Nudds (2004, text-g. 150). At an overall diameter of 24 cm, it shows narrowly crowded ribs and a black band whose width does not exceed 5 mm at its aperture (Text-g. 8). In addition to the crowded ribs and its large size, the slightly widened umbilicus indicates that this was a mature individual. 3. The only specimen from the Middle Jurassic (Bajocian) is a Chondroceras sp. (PIMUZ 16339) from Lupfen near Talheim (south-west Germany; Text-g. 9). Among seven specimens from a small collection it is the single one that preserves a black structure and a complete terminal aperture. In this specimen, adulthood is indicated by eccentric coiling of the last whorl; a radial constriction; a strong, prorsiradiate rib; a triangular constriction which is restricted to the umbilical wall, and three short anterior projections (one on each ank and one on the venter). Remains of the black band are preserved only on the left side, being at a very low angle to the crest of the strong rib on the anterior ank of it and sweeping forwards on the venter, forming part of a ventral projection. As in the above-mentioned Psiloceras, there is a light-coloured shell portion between the black band and the actual aperture.

It appears certain, however, that it is part of the terminal apertural modication. 4. The last fossil (PIMUZ 7527) is of Late Jurassic (Oxfordian) age; it represents the only specimen in which the black structure is probably secondary, i.e. not a true black band (Text-g. 9D). This specimen probably belongs to the microconchiate genus Glochiceras. Collected at the gern, a mountain ridge north of Zu rich, this internal La mould measures 23 mm in diameter, which is characteristic of adult specimens of this taxon. It also displays typically adult apertural modications, a constriction and lateral apertural apophyses. The deeper portion of this apertural modication, i.e. the constriction and the median furrow of the right apertural apophysis, contains dark grey matter that resembles some dendritic deposits well known from Late Jurassic deposits in southern Germany; magnesium oxide secondarily lled the void which was originally occupied by shell. It differs from the true black bands described above in composition and colour. The presence of such secondary deposits can easily be explained by increased shell thickness within the constriction: after dissolution of the aragonitic shell, the fairly broad void often witnessed secondary mineral deposition.

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TEXT-FIG. 7.

Lytoceras ceratophagum (Quenstedt, 1885), SMNS 26465; Posidonienschiefer Formation (Unterer Schiefer Bed, rttemberg, Germany). Note the black band. falciferum Zone, elegantulum Subzone; Toarcian, Early Jurassic), Ohmden (Baden-Wu

We suggest referring to this feature as a pseudo-black band.

irregularly distributed pore canals which have not been documented in the fossil specimens.

Recent For comparison, a shell of Recent Nautilus pompilius (Linnaeus, 1758) PIMUZ 7806 was examined (Textg. 1). At a diameter of 183 mm, it shows the black band directly behind the aperture. Especially the posterior margin is very irregular and the black matter fades out over approximately 10 mm. The exact distribution of the black material at the apertural margin is somewhat unclear because in many museum specimens of this taxon, the apertural edge was removed, polished or damaged during post-mortem transport. It turned out that the black material is thickest near the umbilical plug (c. 1 mm) and thins towards the venter to approximately 03 mm. On the venter and in the umbilical plug, the black material actually covers part of the surface. The black band passes into the black layer without interruption. This feature is not visible in fossils available to us, but we expect that there are no differences between ammonoids and nautilids in this respect. As described by Mutvei and Doguzhaeva (1997), the area of the black band also displays the

TERMINAL APERTURE
All structures described here undoubtedly are related to growth processes, except for the pseudo-black band in Glochiceras. In other Jurassic specimens, and in Paranannites from Nevada, the formation of a black band at the aperture is clearly linked to terminal growth. Some details of the growth patterns, however, appear to differ from those of terminal growth in Recent nautilids (Textg. 10). The main difference between the terminal aperture and the distribution of the conchiolin in Jurassic ammonoids and Recent nautilids is the fact that in several ammonoids the organic material is present mainly on the shell exterior, perhaps as part of the periostracum as in the Posidonia Shale material (compare Seilacher et al. 1976; Text-g. 10) whereas in nautilids it can be found mainly internally. In all of the studied ammonoids, the black band rather represents a thickened organic coating (like a thickened periostracum) on the outer shell surface. Similar black lines occur behind the aperture in juvenile ammonites (annulare Linie; Richter 2002, pp. 1314, pl.

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black band

1 cm

TEXT-FIG. 8.

Harpoceras (Harpoceras) falciferum (J. Sowerby, 1820), photograph courtesy of Urweltmuseum Hauff; Posidonienschiefer Formation (Posidonia Shale, falciferumbifrons zones, falciferumlower commune subzones; Toarcian, Early rttemberg, Germany). Jurassic), Holzmaden (Baden-Wu

B A

C D
T E X T - F I G . 9 . AC, Chondroceras sp. (PIMUZ 16339); Middle Jurassic (Bajocian), Lupfen near Talheim (Germany); note the black gern (north of Zu rich, Switzerland); note the band on the last rib. D, ventral view of Glochiceras sp. (PIMUZ 7527), Oxfordian, La pseudo-black band.

11, g. 1; pl. 21, gs 4, 11), which possibly represent interim or black apertures. In nautilids, it appears very likely that the black band is somehow related to the aper-

tural mantle attachment, which served to prevent water from entering between shell and mantle (Mutvei et al. 1993). Since the black band appears to be external in am-

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A B C D E

F G
T E X T - F I G . 1 0 . Schematic cross-sections through the shell; in BG at the adult aperture, in A roughly half a whorl prior to the terminal aperture. Towards the aperture is on the right, towards the outside is to the top of the illustration. In BF the shell thickness is estimated because the aragonitic shell is either not preserved or potentially incomplete. A, cross-sections through megastriae in Paranannites sp. (PIMUZ 26263) containing conchiolin at an alte Mundrand. B, cross-section through terminal aperture of Paranannites slossi (Kummel and Steele, 1962) (Jenks Coll. no. 499C), displaying a thin dark layer immediately at the aperture on the shell surface. CD, two specimens of Psiloceras planorbis (J. de C. Sowerby, 1824) (PIMUZ 6519 and 12581); in these specimens, the black band is in one case (C) immediately behind, and in the other (D) directly in front of the terminal shell thickening. E, several ammonite species from the Toarcian Posidonia Shale at Ohmden (Germany) displaying the thin, dark black band. Since only the periostracum is preserved, the band appears to be part of the periostracum and thus, this structure was probably on the outside of the shell. F, Chondroceras sp. (PIMUZ 16339); note the black band on the surface of the last rib. G, terminal aperture of Nautilus pompilius (Linnaeus, 1758) (PIMUZ 7812) with the thickness relationship of the aragonitic shell and the conchiolin of the black band.

monoids, it might either have resulted from gerontic growth lacking a soft-tissue attachment function or, alternatively, it might be explained by the mantle that extended out of the aperture.

FALSE COLOUR PATTERNS AND MEGASTRIAE

In Carboniferous and Early Triassic material from Nevada, Oman and South China, black lines are formed at irregular and, apparently aperiodic, angular distances (cf. e.g. Bucher 1997). In all of these specimens, the anterior edge of these black lines is clearly delimited whereas the lines fade out posteriorly. A specimen from Oman (PIMUZ 26262) has helped us to understand this phenomenon. A piece of shell had broken off and enabled the study of shell cross-section at two of these bands. It turned out that the black material covered the inside of a former aperture, because it traverses the shell from the inside to the outside with an adoral tilt. Possibly the structures thus formed were identical to the black band in Recent nautilids, at least in its distribution and formation. This shows that these black lines were formed during a growth halt. Mapes and Davis (1996) included these structures in their category of false colour patterns, and stated that, in many cases, these correlated with the formation of constrictions, pseudo-constrictions, varices and megastriae. The distribution of the black material in the

Omani specimen demonstrates that these lines were formed at a growth interruption and, thus, can be interpreted as megastriae, previously also referred to as, for nder, demarcation lines or tranexample, alte Mundra sitional mouth borders (Pompeckj 1884; von Mojsisovics hner 1894; Diener 1895; Matsumoto et al. 1972; 1886; Wa Bucher and Guex 1990; Matsumoto 1991; Tozer 1991; for a review of this structure and additional references, see Bucher et al. 1996). It is quite conceivable that this structure was also related to a temporary apertural soft-tissue attachment as in the terminal aperture of Recent nautilids. As a secondary effect, these false colour patterns may have had similar functions (e.g. camouage) as true colour patterns, which are conned to the surface of the ammonoid shell.

BLACK APERTURE
The Middle Triassic cephalopod remains of Germany do not allow an unequivocal interpretation. The smaller of the two ammonoid specimens (Ceratites postspinosus) is not of a size typical of adults of the species. It is unclear, however, whether this falls within the intraspecic variability. Septal crowding is seen in this specimen but this can easily be produced under environmental stress, as can a black band (see Arnold 1985; Ward 1987 for Recent nautilids). The irregular distribution of the black substance within the black line at the aperture might be some

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kind of artefact. The fact that it wedges out posteriorly is reminiscent of the state of the black band in Recent nautilids. For this specimen, it appears reasonable to assume that the black structure represents a combination of mature modication and a structure formed under adverse environmental conditions. In contrast, the black line found in Germanonautilus can denitely be interpreted as a black aperture. In addition to terminal growth itself, the rarity of preservation of this structure invites a discussion on the age pyramid of an ordinary ammonoid. When studying ammonoid taphocoenosis, usually two cases are encountered. Either only more or less adult ammonoids of approximately the same size are found and juveniles are exceedingly rare, or representatives of almost all growth stages are preserved, although only very rarely representing the actual age pyramid. This phenomenon can be explained to a large extent by taphonomic bias, i.e. early dissolution and or breakage of fragile shells of small specimens or sorting by currents, respectively (e.g. Maeda 1991). Remarkably, the black band is rarely preserved even in ammonoid taphocoenoses of the rst type. This can probably be explained mainly by the organic nature of this structure and its poor preservation potential. Additionally, it shows that truly fully grown ammonoids are rare for many taxa. Furthermore, not every mature ammonoid will have secreted a black band; after all, some mature Recent nautilids also lack this feature (Ward 1987). This ts the interpretation of ammonoids as rstrategists, which would correspond to a type III survivorship curve.

As the black bands of the ammonoids depicted herein are located on the external shell surface, they clearly differ from the state in Recent nautilids, in which the area of the black band serves the attachment of the apertural mantle. This functional interpretation appears unlikely for ammonoids. In ammonoids, the black band might simply be a gerontic artefact.
Acknowledgements. This work beneted from a research stay in Berlin (DE-TAF) in July 2006, supported by Synthesys, where the Carboniferous material was examined. It is a contribution to Swiss National Science Foundation project no. 200020-113554 to rich). For providing material and photographs Hugo Bucher (Zu of specimens we thank Jim Jenks (Salt Lake City), Gilbert M. rich), Hans Hagdorn (Ingelngen) and Rolf Bernhard Kahn (Zu Hauff (Holzmaden). Some of the photographs were taken by rich), Heinz Lanz, Thomas Galfetti and Rosemarie Roth (all Zu bingen). We acknowledge the and by Wolfgang Gerber (Tu EDAX analyses run by Michael Montenari and Hartmut Schulz bingen), and appreciate the thorough revision of the (both Tu manuscript by John W. M. Jagt (Maastricht) as well as the constructive reviews of Ryoji Wani (Tokyo) and an anonymous referee.

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CONCLUSIONS
As all of the above structures primarily consisted of conchiolin and are located at the terminal or at a former aperture, and as they are preserved in several specimens from the German Posidonia Shale (where most ammonoids display periostracum preservation), it is reasonable to assume that most of these structures were formed by the mantle fold, which also secreted the periostracum. Terminal growth of the specimens, like any kind of environmental stress or injury, might have represented internal stress. Thus, we suggest that all these structures were formed under adverse conditions in a broad sense which perturbed the regular formation of periostracum and shell. Therefore, the formation of all of the structures presented herein has the same underlying cause, i.e. some kind of stress, but with varying results: injury (black stripe), adverse conditions at any growth stage (black aperture), interim growth stops (megastriae; false colour bands) and terminal growth (black band).

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