Izmirella cretacea nov . gen . , nov . sp . , a complex alga from the Lower Cretaceous , NE of Bornova-Izmir / Turkey

In the Izmir region (Western Turkey) near Bornova, Mesozoic rocks are represented by the “wildfl ysch” zone of the Taurids consisting of a Maastrichtian-Danian sandy-shaly matrix with Late Triassic to Late Cretaceous carbonate megablocks. Previous work on the Mesozoic carbonates has been carried out by AKDENIZ et al. (1986), ERDOĞAN (1990) and ISINTEK et al. (2000). AKDENIZ et al. (1986) noted a thick carbonate sequence of Late Cretaceous age, tectonically overlying the Late Cretaceous clastics. ERDOĞAN (1990) reported that the Late Triassic to Late Cretaceous limestone masses in question were allochthonous blocks in the so-called “Bornova Melange” of Maastrichtian-Danian age. According to ERDOĞAN (1990) the carbonate block lying between the village of Naldöken and Çiçekli consists of Jurassic to Upper Cretaceous deposits, including an Albian bauxite lens north of Naldöken (Fig. 1). Recently the presence of a second bauxite bearing horizon in the Barremian part of the upper limestone unit has been documented NE of Naldöken (Fig. 2). The Izmirella-bearing limestones are exposed in the socalled “Bornova-block” located between Naldöken and Çiçekli (1:25.000, Izmir – L18 – b1 sheet; Figs. 1, 2). The studied Cretaceous deposits, north of Naldöken, are represented by a thick sequence of lagoonal, subtidal to supratidal limestones, Barremian–Lower Aptian in age and including a bauxite horizon. The type section of the Izmirella-bearing limestones is located 250 m east of the Hendilagili Ridge, 2.7 km to the northeast of Naldöken (Figs. 1b, 2), and consists of a grey, thick bedded limestone sequence about 30 m thick (Fig. 3). The basal part of the section (about 10 m thick) consists of thick-bedded peloidal micrite and/or peloidal sparry facies, including laminated fenestral structures, intercalated with terrestrial lime clays and palaeosols. The middle part of the section is made up of well bedded or nodular micritic and/or sparitic limestones, sometimes laminated, with pebbly or clayey micrite associated with early erosional surfaces. The clayey micritic limestones include rare foraminifera, gastropods, ostracods, faecal pellets, SalpingoBaba Senowbari-Daryan1 and Ismail Isintek2


INTRODUCTION
In the Izmir region (Western Turkey) near Bornova, Mesozoic rocks are represented by the "wildfl ysch" zone of the Taurids consisting of a Maastrichtian-Danian sandy-shaly matrix with Late Triassic to Late Cretaceous carbonate megablocks.Previous work on the Mesozoic carbonates has been carried out by AKDENIZ et al. (1986), ERDOĞAN (1990) and ISINTEK et al. (2000).AKDENIZ et al. (1986) noted a thick carbonate sequence of Late Cretaceous age, tectonically overlying the Late Cretaceous clastics.ERDOĞAN (1990) reported that the Late Triassic to Late Cretaceous limestone masses in question were allochthonous blocks in the so-called "Bornova Melange" of Maastrichtian-Danian age.According to ERDOĞAN (1990) the carbonate block lying between the village of Naldöken and Çiçekli consists of Jurassic to Upper Cretaceous deposits, including an Albian bauxite lens north of Naldöken (Fig. 1).Recently the presence of a second bauxite bearing horizon in the Barremian part of the upper limestone unit has been documented NE of Naldöken (Fig. 2).
The Izmirella-bearing limestones are exposed in the socalled "Bornova-block" located between Naldöken and Çiçekli (1:25.000,Izmir -L18 -b1 sheet; Figs. 1, 2).The studied Cretaceous deposits, north of Naldöken, are represented by a thick sequence of lagoonal, subtidal to supratidal limestones, Barremian-Lower Aptian in age and including a bauxite horizon.The type section of the Izmirella-bearing limestones is located 250 m east of the Hendilagili Ridge, 2.7 km to the northeast of Naldöken (Figs. 1b,2), and consists of a grey, thick bedded limestone sequence about 30 m thick (Fig. 3).The basal part of the section (about 10 m thick) consists of thick-bedded peloidal micrite and/or peloidal sparry facies, including laminated fenestral structures, intercalated with terrestrial lime clays and palaeosols.
The middle part of the section is made up of well bedded or nodular micritic and/or sparitic limestones, sometimes laminated, with pebbly or clayey micrite associated with early erosional surfaces.The clayey micritic limestones include rare foraminifera, gastropods, ostracods, faecal pellets, Salpingo-Baba Senowbari-Daryan 1 and Ismail Isintek 2

AB STRA CT
Izmirella cretacea nov.gen., nov.sp., a complex alga, forming microbioherms is described from Barremian -Aptian platform carbonates of the "Bornova block", an allodapic unit derived from the northern margin of the Menderes Massif, included in Maastrichian -Danian fl ysh deposits, called the "Bornova melange".The verticillate thallus of this alga consists of strongly imbricated bowl-to funnel-shaped whorls having tightly stacked laterals with two layers.The fi rst is composed of radially and fan-like running segments, appearing as large cells in section parallel to the growth direction.The second is composed of multibranched tubes, running perpendicular to the segments of the fi rst layer.Besides its organizational traits, this has original calcite mineralogy and an aggregative behaviour leading to the formation of laminated build-ups closely resembling stromatolitic bioconstructions or even stromatoporoids.porella (Hensonella?)dinarica RADOIČIĆ, large (2-25 cm) fragments of Izmirella microbioherms.Salpingoporella (Hensonella?)dinarica RADOIČIĆ is a paleobiogeographic marker of the Mediterranean Southern Tethyan margin (PEYBERNES & CONRAD, 1979), and has been reported from beds ranging from the Valanginian(?)(SCHINDLER & CONRAD, 1994) or from the Tithonian?-Berriasian(GRANIER, 2002) to Late Aptian (VELIĆ & SOKAČ, 1983).Its acme is in the Barremian-Lower Aptian (LUPERTO SINNI & MASSE, 1993).

Geologia Croatica Geologia Croatica
The upper part of the type section with Izmirella cretacea nov.gen., nov.sp. is characterized by bioturbated, thick bedded or laminated micritic and/or intra-pelsparitic limestone with laminated fenestral structures, and thick bedded nodular micritic limestone.

Derivatio nominis:
After the city of Izmir, close to which the alga was found.
Diagnosis: Thallus segmented, composed of strongly imbricated bowl-to funnel-shape whorls, each with tightly stacked, septate laterals.Laterals, appearing as an oval or rectangular cell layer in sections, are divided radially several times towards the periphery of the whorls.A multibranched cell layer originates from the upper surface of the laterals, forming a second layer of whorls.A third layer consisting of "chaotic" tubes is located between the individual layers of the thallus.Calcifi cation was originally calcitic.The organism grew with successive rows of closely packed individuals forming domal, laminated thalli resembling stromatolites or even stromatoporoids.
Type species: Izmirella cretacea nov.sp.Material: Four microbioherms contain numerous hemispherical laminae.From one microbioherm containing the holotype (Pl.4, Fig. 1) three large thin sections (10´15 cm) and from the second microbioherm two large thin sections were made.

Description of microbioherms
Izmirella cretacea nov.gen., nov.sp.forms bioconstructed bodies up to 22 cm high which resemble stromatolites or stromatoporoid sponges with distinct laminations (Pl. 1,Pl. 2,Pl. 3,Fig. 1).The individual laminae (layers) of the outside of the bioherms correspond internally to the individual rows, which are formed by numerous thalli of Iz mirella cretacea arranged one beside the other (Pl.2, Figs.1-2).In thin sections, individual laminae are separated by dark horizons, usually thinner than the layers of Izmirella rows (Pl.2, Figs.1-2).These dark horizons are composed of small and multibranched tubes of "chaotic" arrangement (Pl. 5,Figs. 5,8) resembling Girvanella.These tubes will be described in detail under the description of Izmirella.
The microbioherm containing the holotype exhibits the well-developed lamination on one side only but the paratype, illustrated in Pl. 1 shows well developed lamination on both sides.The base of this microbioherm is bulbous and shows several circular shaped initial parts of individual thalli of Izmirella, having a diameter of approximately 2 mm.Each basal bulb consists of the individual thalli of Izmirella cut in cross section (see Pl. 2, Fig. 1 upper part of the photograph).
The outer surfaces of all four available microbioherms are lamellar.The height of individual laminae is almost constant (about 3 mm, rarely varying between 2 mm and 5 mm).The laminae in the basal part of two microbioherms extend laterally, parallel to the substrate but in the younger part they are almost sub-spherical.In the third microbioherm the laminae at the top also extend parallel to the substrate (Pl. 1, Figs. 1-2).This specimen seems to be broken at both sides.The top of all microbioherms are vaulted.
Three large thin sections (10x15 cm, two parallel and one transverse to the growth direction) were cut into the microbioherm containing the holotype.Two large thin sections were made from the second microbioherm.
As mentioned above, the darker horizons between the individual laminae (Pl.2, Figs.1-2; Pl. 3, Fig. 1) are composed of a "chaotic" cell layer, similar to Girvanella (Pl.4, Fig. 7; Pl. 5,Figs. 5,8).As observed in several specimens, the edge of the last whorl becomes trumpet-like from which the "chaotic" Fi gu re 4: Section through three layers of laterals appearing white in thin sections (1) and three layers of tubes of second layers appearing dark in thin sections (2).The upper right corner of the photograph corresponds to the upper surface of laterals, from which the tubes of second layer (2) originate.The centre of the thallus is located in the NE, outside of the drawing.Redrawn from Pl. 5, Fig. 2.
Fi gu re 5: Section exhibiting the radiating branching pattern of the laterals from the axial stem.From seven laterals at the base (from the "axial stem"), about 40 laterals originate in the periphery of the whorl.
View of the "front-side" of a paratype showing the well-developed wavy laminae oriented parallel to the substrate (for magnifi cation of laminae see fi g. 4).
View of the other side ("back-side") of the same specimen illustrated in fi g. 1.
Side-view of the same specimen illustrated in fi g. 1 or fi g. 2.
Magnifi ed part from Fig. 1 (quadrangle marked with white line) shows details of the lamination.
1 Magnifi cation from thin section 1q (transverse to oblique section) showing three layers (correspond to the laminae, separated by dark area containing "chaotic tubes") in which the individual thalli are arranged in rows one beside the other.In the lower part of the photograph the thalli are cut in longitudinal section exhibiting the individual soup-plate-, bowl-or funnel-shaped whorls arranged one above the other.At the upper part of the photograph the thalli are cut obliquely or transversely, exhibiting the circular outline of the whorls in cross section.
2 Magnifi cation from thin section 1/l showing the rows (layers or laminae), in which the individual thalli are arranged one beside the other.The dark areas between the layers of adjacent thalli contain small "chaotic tubes".The youngest whorl of the thallus is usually fi lled with calcite cement.
1 Longitudinal section (parallel to the growth direction) of a microbioherm containing the holotype (see Pl. 4, Fig. 1) exhibits the individual layers (laminae) composed of numerous thalli of Izmirella arranged in rows one beside the other in horizontal direction (thin section 1/l).
2 Magnifi cation from fi g. 1 showing several thalli with bowl-or funnel-shaped whorls.The interior of the last whorl is usually fi lled with calcite cement.Thin section1/1.
3 Magnifi cation from fi g. 1.View of a thallus showing the plate-like and relatively broad whorls at the lower part (right in photograph) and V-shaped whorls at the upper part (left in photograph) of a thallus.Thin section 1/1.
4 Section through the younger part of a thallus that exhibits the cup-shaped last whorl.From the upper part of the youngest whorl there appears to arise a new whorl or a new thallus(?).The interior of the last whorl is covered by micrite of organic origin; the remainder is fi lled with calcite cement (for details see Fig. 10).
5 Longitudinal section through a thallus showing the "laterals" of the fi rst order (small arrows), branching to the "laterals" of the second and higher orders.The large triangular arrows indicate the radially arranged "laterals" of the fourth order.The medium-sized triangular arrows indicate the polygonal outline of the "tubes of the second layer" running perpendicular to the "laterals" of fi rst and second order.The small triangular arrow indicates the axial stem connecting the individual whorls.The large arrows at the left of the photograph indicate the branching pattern of the "tubes of the second layer" from the upper part of the "laterals".Note the wavy (zig-zag-like) wall between the "laterals" of fi rst and second orders (compare fi g. 7).Thin section 1/1.
6 Similar to fi g. 5. Longitudinal section through four whorls exhibiting alternating "laterals" and the polygonal outline of the "tubes of the second layer" which are cut in transverse section and run perpendicular to the "laterals".Note the wavy (zig-zag-like) wall between the "laterals".Thin section l/q.PLATE 4 1-7: Izmirella cretacea nov.gen., nov.sp.
All fi gures are from the same microbioherm as the holotype (thin section 1/l, Pl. 3, Fig. 1).
Scale in all fi gures 1 mm.
1 Holotype.Axial longitudinal section showing numerous soup-plate to bowl-to funnel-shaped whorls.The narrow axial stem is cut in the upper and lower parts of the whorl.Thin section 1/1.
2 Section through a thallus exhibiting several whorls with the axial stem cut in the centre of the last four whorls.Tubes of the second order, originating from the edges of the last whorl, extend bridge-like between the edges of the last whorl.The interior of the last whorl is covered by micritic layer (black) of organic origin."Up" is toward the left.
3 Magnifi cation of several whorls showing the "laterals" (appearing as large cells) passing to the small, branched tubes ("tubes of second layer") which have arisen from the laterals of fi rst order (arrow)."Up" is toward the left.
4 Magnifi cation of a thallus showing the bowl-like whorls at the lower part (right in photograph) and funnel-or Vshaped at the upper part (left in photograph) of a thallus.The small arrow indicates the radially arranged and fanlike multibranched tubes of the fi rst order of "laterals" (longitudinal section)."Up" is toward the left.
1 Section through two thalli each with numerous whorls.The interior of the last whorl in the thallus on the left is fi lled with calcite cement.The axial stem is cut in the lower part of the thallus on the right.For details see fi g. 2.
2 Magnifi cation of the area in fi g. 1, marked with white rectangle, showing numerous whorls with the laterals appearing as rectangular cells (lower part) adjacent to the axial stem.The axial stem passes through the whorls and branches at the upper most part (see fi g. 3).
1 Section through three layers of laterals and tubes in a second layer.White arrows indicate the tube-like laterals.
Tubes of the second layer originate from the "pores", in the upper surface of laterals.For magnifi cation of the area marked with rectangle marked see fi g. 2. "Up" is toward the left.
Girvanella-like tube layer originates (Pl. 3,Fig. 4;Pl. 5,Figs. 5,8;Pl. 6,Figs. 4,7;Fig. 8).The edges of the last whorl in some specimens are connected by a layer of tubes, forming a lid (appearing as a bridge between the edges) covering the top of the whorl (Pl. 3,Fig. 4;Pl. 4,Fig. 2;Pl. 6,.Apparently the repetitive settlement of the alga results from these tube layers originating from the edges or lids of the last whorl.In most cases one or several calcite-fi lled cavities were observed within the clusters of the "chaotic" Girvanella-like tubes (Pl.5, Fig. 8).Interpretation of these cavities as conceptacles is very speculative.
The most peculiar part of this alga is the axial stem connecting the whorls of the thallus (Pl. 4,5;Pl. 5,Fig. 3;Pl. 6,(10)(11)13).The axial stem is very narrow (less than 1 mm; usually 0.40-0.80mm).Numerous sections show that the axial stem is a part of this alga and can not be interpreted as a boring or as an interaction between the alga and another organism (symbiosis or commensalism; like a worm, etc.).The following criteria support this interpretation: a) the axial stem is always in the centre of the whorls and the whorls are arranged symmetrically around it; b) the cements of the axial stem continue into the whorls and there is no recognizable boundary between them (Pl. 6,(11)(12)(13); c) the laterals of the last whorl originated from the branching of the axial stem and the axial stem does not possess its own wall (Pl.6, Figs.2-3, 4-5, 10-13); d) the axial stem never steps out over the last whorl.
The holotype (Pl.4, Fig. 1) exhibits numerous bowl-to V-shaped whorls in longitudinal section, showing only some of the characteristics of Izmirella cretacea summarized from the paratypes.The holotype is composed of at least 20 whorls.The axial stem is cut through several whorls.A reconstruction of a half-whorl of Izmirella cretacea nov.gen., nov.sp. is given in Fig. 6.
Rejuvenation of the thalli usually takes place at the boundary between two laminae (Pl. 2,Pl. 3,Figs. 2,4,Pl. 4,Fig.4).This boundary is marked by a dark horizon where individual whorls, at the base of the thalli, are usually developed as a soup-plate pile arranged densely one above the other.

DISCUSSION
Izmirella cretacea nov.gen., nov.sp. is a representative of fossil plants, most probably algae.However, because of the totally different characteristics, its systematic classifi cation within the known groups of algae is uncertain.
In longitudinal sections, (parallel sections to the growth direction), through the thalli of Izmirella-buildups, the individual laterals appears as cell rows, resembling some red algae (Rhodophycea) with "large cells", e. g. representatives of the genus Lithoporella (FOSLIE) or Titanoderma NÄGELI (compare WRAY, 1977, fi g. 58;RASSER, 1994;PILLER & RASS-ER, 1999).Izmirella may be ascribed to such algae if its thallus is fractured (Pl.5, Fig. 6).In thin-sections of Izmirella, the "large-cell"-like structures (similar to Lithoporella), are in reality the radially arranged (similar to representatives of Modern Acetabulariaceae) "laterals" cut in cross section.Cells with different sizes and arrangement may occur in some red algae, e. g. in crustose coralline red algae Phymatolithon calcareum and Lithothamnium coralloides (ADEY & MCKIBBIN, 1970) or in recent genus Tenarea (WRAY, 1977, fi g. 54).Laterals in Izmirella are, however, segments that are arranged as in the cups of Recent dasycladales Acetabularia -radially around the axial stem, which is not known from red algae.
The radially arranged "laterals" and the axial stem connecting the successive whorls is a common feature of the Dasycladales, family Acetabulariaceae (see BERGER & KAE-VER, 1992).In this group, individuals are usually isolated; the gregarious potential and the ability to build small bioherms, as in Izmirella are very rare in the fossil record.Neo- showing the radially arranged "laterals" that are subdivided internally and peripherally to laterals of the second and higher orders (compare also Fig. 7).From the pores of the upper surface (shown in third lateral from left) open into the cross-sections of "tubes of the second layer" which are oriented perpendicular to the laterals of fi rst and higher orders.Schematic, not to scale.teutloporella BASSOULETT et.al. is the only representative (BASSOULETT et. al., 1978) of gregarious (numerous thalli growing one beside the other) dasycladales occuring in the Oxfordian-Tithonian according to GRANIER & DELOFFRE (1993) or Oxfordian-Berriasian according to SCHLAGINT-WEIT & EBLI (1999).For instance in the Upper Jurassic reef facies of Sicily, this alga is very abundant in nodular microbioherms up to 30 cm in diameter (SENOWBARI-DARYAN et al., 1994).
The following features might support the affi liation of Izmirella to the Dasycladales: a) Each thallus possesses an axial stem which passes through the individual whorls (Pl. 4,5;Pl. 5,Fig. 3;Pl. 6,13).The axial stem with a diameter of 40-80 µm, however, is extremely narrow to bear the weight of laterals with a diameter of about 3 mm, but the Recent Acetabularia also possesses a narrow stalk.It may be assumed that the arrangement of the thalli in rows and their growth strategy in Izmirella, i. e. one beside the other, could help to support the individual thalli in growth position.
b) The arrangement of the "laterals" is comparable with the rays (laterals) from caps of Recent Acetabularia, Polyphysa or Chalmasia, radiating around the axial stem (see BERG-ER & KAEVER 1992, fi gs. 3.56, 3.81, 3.91), c) The segments, i.e. whorls, are arranged vertically around the axial stem, one above the other.
d) The microbioherms are composed of individual stacked thalli of Izmirella.
Notwithstanding these apparent similarities, the verticillate organisation of the whorls around the axial stem and the septate nature of the laterals, do not conform to the features of the Dasycladales (BERGER & KAEVER, 1992).The following features of Izmirella argue against its assignment to Dasycladales: a) Calcite mineralogy of skeleton: The primary skeletal mineralogy of the Dasycladales is aragonite; primary calcite in Dasycladales is extremely rare.Minor amounts of calcite do occur in some Dasycladales e. g. in Bornetella (FLAJS, 1977), and the membranes around the laterals of Clypeina jurassica (FAVRE) are composed of Mg-calite (CONRAD & VAROL, 1990).Similarly, the sporangia within the axial stem in Diplopora iranica (SENOWBARI-DARYAN & HAMEDANI, 2000) were primary Mg-calcite, but a primary Mg-calcite mineralogy for the thallus of Dasycladales is not known.The systematic position of some dasyclad type organisms, e. g.Palaeozoic Psedo vermiporella ELLIOT, as Dasycladales is uncertain (FLÜGEL, 2004).Therefore, the primary calcite mineralogy of Izmirella does not support its classifi cation as a dasycladalean.
b) The branching pattern of the "tubes of the second layer" is similar to porostromate cyanophyceans like Cayeuxia, Hedstroemia or Halysis mentioned above.However, such structures, originating from the radially arranged laterals around the axial stem, are not known from the Dasycladales.Ramification of laterals to a higher order and the formation of the "chaotic tube" in Izmirella are not characteristic for Acetabulariaceae.Gametangia, occurring in the laterals of e. g.Acetabularia, have not been observed in the laterals of Izmirella.
c) Also the "chaotic" (Girvanella-like) tube layer between the individual thalli of Izmirella rows does not support its interpretation as a Dasycladacean.
An alternative classifi cation to the Dasycladales might be the red algae.Actually a placement of Izmirella in the Peyssonneliacea could be envisaged.In this view the "stacked superimposed whorls" might be interpreted as lamellar thalli growing on some non-calcifi ed erect fi laments (= "axial stem"); Figure 9: Section through the last cup of specimen illustrated in Pl. 6, Fig. 8 showing the large "cells" of the laterals, numerous tubes of the lid (appearing as bridge between the edges of the cup), micritic fi lling within the bowl (dotted area) and the small spherical elements (gametangia?)within the micritic fi lling.The white area within the micritic fi lling is sparry calcite cement.
this peculiar mode of life may be refl ected by the wavy "bowlshaped" aspect.Nevertheless, this hypothesis looks inadequate if one considers the absence of a well-defi ned cellular network with a hypothallus and perithallus (DENIZOT, 1968) which typifi es the Peyssonneliacea.
The cell-like structure (oval or rectangular) of the laterals in sections directly around the axial stem (Pl. 4,Figs. 3,6;Pl. 5,Pl. 7,Figs. 2,5,8), also are similar to some red algae (e. g.Lithoporella, see WRAY, 1977, fi g. 56), but they are tube-like in Izmirella.Different cell sizes, developed as two tube layers of Izmirella, is known from other red algae, e. g. in the crustose coralline red algae Phymatolithon calcareum and Lithothamnium coralloides (ADEY & MC KIBBIN, 1970).The cells are signifi cantly elongated during winter growth.Also the mineralogy of the cells (i.e. content of Mg) in modern calcareous red algae varies from summer to winter (MIL-LIMAN et al., 1971;KOLESAR, 1978).Seasonal cell sizes and variation in Mg-content in calcite of the cell walls is also known from fossil solenoporacean algae (WRIGHT, 1985;WENDT, 1993).The different cell sizes, and possibly different content of Mg in the cell wall of laterals and "tubes of the second layer" in Izmirella could also refl ect seasonal variation, but these features do not justify classifying Izmirella as a solenoporacean red algae.
The fan-like branching pattern of the fi rst order of tubes and of the "chaotic" tube layer of Izmirella could support its attribution to the cyanophyceans.Such branching patterns of cells is common in cyanophyceans, but the axial stem of Izmirella and the arrangement of laterals around the axial stem are features contrary to the affi liation of Izmirella with this group.
Izmirella nov.gen. is surely an alga but its taxonomic position within the algae remain uncertain.

Fi gu re 1 :
Location map of the study area in Western Turkey; (a) geological map of the Bornova area; (b) the position of the Izmirella nov.gen.bearing site (simplifi ed after ERDOĞAN, 1990).Fi gu re 2: Detailed geological map of the Mesozoic carbonates of the "Bornova block" in the area of the type locality of Izmirella nov.gen.

Figure 6 :
Figure6: Reconstruction of a half whorl of Izmirella cretacea nov.gen., nov.sp.showing the radially arranged "laterals" that are subdivided internally and peripherally to laterals of the second and higher orders (compare also Fig.7).From the pores of the upper surface (shown in third lateral from left) open into the cross-sections of "tubes of the second layer" which are oriented perpendicular to the laterals of fi rst and higher orders.Schematic, not to scale.

Figure 7 :
Figure 7:The branching pattern of the laterals of the fi rst, second and further orders (6 orders, see numbers) of the whorls.The branching doesn't occur at exactly the same level.The wall between the tubes is zig-zag-like (wavy).Schematic, not to scale.

Figure 8 :
Figure 8: Redrawn from Pl. 3, Fig. 7 (quadrangle) showing the edge of a trumpet-like broadened lateral, from which the tubes of the second layer originate.

Figure 10 :
Figure 10: Section through two cups exhibiting the laterals (1) and the tubes of second layers (appearing dark between the laterals).The inside of the cup is covered with a thin micritic layer (dotted area).The white area in the centre of the cup is sparry calcite cement.Between the edges of the cup the bridge-like appearing tubes (lid) are developed, from which the "new" and multibranched laterals originated (rejuvenation).Redrawn from specimen illustrated in Pl. 3, Fig.4.