GeolCarp_Vol65_No5_339

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, OCTOBER 2014, 65, 5, 339—364 doi: 10.2478/geoca-2014-0024

Introduction

Upper Triassic sediments, especially of the Carnian age, form
the major element within the Taurus Mountains. The area
around A ag˘

yaylabel has already been investigated by Özgül

& Arpat (1973), Dumont & Kerey (1975), Monod (1977),
Poisson (1977), Gutnic et al. (1979), Robertson (1993, 2000),

enel (1997), Gindl (2000), Robertson et al. (2003), Lukeneder

et al. (2012) and Lukeneder & Lukeneder (2014).

The Carnian section at A ag˘

yaylabel displays a lithologi-

cal change from pure carbonatic to more siliciclastic sedi-
mentation (Lukeneder et al. 2012). The facies change occurs
exactly at the Lower—Upper Carnian boundary ( = Julian—Tu-
valian boundary), and represents the beginning of the so-
called Carnian Pluvial Event in the section at A ag˘

yaylabel.

Facies interpretations change from open platform margin
conditions, through deeper shelf margin conditions, to finally
open marine-influenced basinal conditions (Lukeneder et
al. 2012).

During the Carnian time, the sediments around the area of

A ag˘

yaylabel were deposited within an intrashelf area on

the western end of the Cimmerian System (Gindl 2000;
Stampfli & Borel 2002; Lukeneder et al. 2012). Sedimento-
logical and paleontological investigations show a delayed
carbonate factory collapse during that time.

Taphonomic implications from Upper Triassic mass flow

deposits: 2-dimensional reconstructions of an ammonoid

mass occurrence (Carnian, Taurus Mountains, Turkey)

ALEXANDER LUKENEDER and SUSANNE MAYRHOFER

Natural History Museum, Geological-Palaeontological Department, Burgring 7, A-1010 Wien, Austria;

alexander.lukeneder@nhm-wien.ac.at; susanne.mayrhofer@nhm-wien.ac.at

(Manuscript received January 22, 2014, accepted in revised form October 7, 2014)

Abstract: Ammonoid mass occurrences of Late Triassic age were investigated in sections from A ag˘

yaylabel and

Yukar

yaylabel, which are located in the Taurus Platform-Units of eastern Turkey. The cephalopod beds are almost

monospecific, with > 99.9 % of individuals from the ceratitic genus Kasimlarceltites, which comprises more than hun-
dreds of millions of

ammonoid specimens. The ontogenetic composition of the event fauna varies from bed to bed,

suggesting that these redeposited shell-rich sediments had different source areas. The geographical extent of the mass
occurrence can be traced over large areas up to 10 km

. Each of the Early Carnian (Julian 2) ammonoid mass occur-

rences signifies a single storm (e.g. storm-wave action) or tectonic event (e.g. earthquake) that caused gravity flows and
turbidity currents. Three types of ammonoid accumulation deposits are distinguished by their genesis: 1) matrix-supported
floatstones, produced by low density debris flows, 2) mixed floatstones and packstones formed by high density debris
flows, and 3) densely ammonoid shell-supported packstones which result from turbidity currents. Two-dimensional calcu-
lations on the mass occurrences, based on sectioning, reveal aligned ammonoid shells, implying transport in a diluted
sediment. The ammonoid shells are predominantely redeposited, preserved as mixed autochthonous/parautochnonous/
allochthonous communities based on biogenic and sedimentological concentration mechanisms ( = in-situ or post-mortem
deposited). This taphonomic evaluation of the Kasimlarceltites beds thus reveals new insights into the environment of
deposition of the Carnian section, namely that it had a proximal position along a carbonate platform edge that was
influenced by a nearby shallow water regime. The Kasimlarceltites-abundance zone is a marker-zone in the study area,
developed during the drowning of a shallow water platform, which can be traceable over long distances.

Key words: Kasimlarceltites, ammonoid mass occurrence, taphonomy, Triassic, Taurus Mountains, Turkey.

Lower Carnian faunal elements, exclusively detected at

A ag˘

yaylabel and characterized as Kasimlarceltites krystyni,

Klipsteinia disciformis and Anasirenites crassicrenulatus
(Lukeneder & Lukeneder 2014) indicate a rather isolated but
still connective paleoceanographic position of the intrashelf
area on the western end of the Cimmerian System.

The present study examines deposits representing an acme

zone, registered within the Upper Triassic (Carnian) Kartoz
and Kas

mlar formations, which crop out at A ag˘

yaylabel

( = Kartoz) and Yukar

yaylabel ( = Karap

nar). This acme

zone  is  characterized  by  several  beds  yielding  ammonoid
mass  occurrences  of  the  Carnian  ammonoid-genus  Kasim-
larceltites. The aim of the present work is to detail the distri-
bution  and  taphonomy  of  the  Kasimlarceltites  mass
occurrence at the lowermost part of the Kas

mlar Formation

within the Julian 2. Results on taphonomy and environmental
processes, obtained by the two dimensional analyses of sec-
tions, thin sections, as well as by conclusions from outcrop
logs and block data, are presented. This leads to a more de-
tailed  picture  of  the  sedimentological  dynamics,  hence  to  a
better understanding of the taphonomy and sedimentology of
such  Upper  Triassic  shell  beds  (e.g.  ammonoids).  Dynamic
processes for specific mass flow deposits (e.g. debris flows,
grain  flows  or  turbidity  currents;  Middleton  &  Hampton
1973,  1976;  Flügel  1978,  2004;  Lowe  1982;  Brown  &

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Loucks  1993;  Stow  &  Mayall  2000;  Mulder  &  Alexander
2001;  Kawakami  &  Kawamura  2002;  Potter  et  al.  2005;
Nichols  2009;  SEPM  2014),  also  termed  gravity  flows  or
density currents, will be discussed in respect to resedimenta-
tion of sediments and biogenic components. The determina-
tion  of  specific  mechanisms  in  terms  of  the  genesis
concerning event bed deposition, storm deposition, shell ac-
cumulation and the consequences for the facies and bioclas-
tic  fabric  were  reported  and  discussed  in  Aigner  (1982a,
1985),  Einsele  &  Seilacher  (1982),  Kreisa  &  Bambach
(1982),  Brett  &  Baird  (1986),  Kidwell  (1986,  1991a,b,
1993a,b),  Kidwell  et  al.  (1986),  Tucker  &  Wright  (1990),
Brett & Seilacher (1991), Chuanmao et al. (1993), Einsele et
al. (1991a,b), Seilacher & Aigner (1991), Soja et al. (1996),
Hips  (1998),  Fürsich  &  Pandey  (1999),  Martin  (1999),
Storms  (2001),  Lukeneder  (2003a,b,  2004a,b),  Fernández-
López  (2007),  Montiel-Boehringer  et  al.  (2011),  and  Pérez-
Lopéz & Pérez-Valera (2012).

The result is a detailed succession of abundance or accumu-

lation layers (i.e. distinct layers with ammonoid mass occur-
rences) within an acme zone in the Upper Triassic of the

A ag˘

yaylabel section. Such ‘ammonoid-beds’ are the result of

bio-events, which are often manifested by the abundance or
mass occurrence of ammonoids (Lukeneder 2001, 2003b). The
presented paper is a first step and the initial point for the lateral
correlation of such ammonoid mass occurrences and establish-
ment of ammonoid abundance zones within the Taurus Moun-
tains. Trigger mechanisms and potential scenarios, causing the
accumulation of such ammonoid shell beds, are discussed.

Geographical setting

The A ag˘

yaylabel (AS) sections (i.e. AS I—AS IV) are lo-

cated in southwest Turkey, about 90 km northeast of Antalya
and approximately 70 km southeast of Isparta (Figs. 1—3).
A ag˘

yaylabel is accessible from the two major cities of the

region, Eg˘irdir and Bey ehir, located 50 km and 40 km
away, respectively. The locality adjoins the small village
A ag˘

yaylabel (1000 m above sea level) on the northern

slope of an east-west trending ridge, between 1050 m to
1100 m at N 37°33’05” and E 31°18’16”. The former

Fig. 1. Locality map of the investigated area showing the outcrops of Upper Triassic sediments around the area of A ag˘

yaylabel ( = Kartoz)

and Yukar

yaylabel ( = Karap

nar) within the the Anamas Dag˘ carbonate platform in the Taurus Mountains (southwest Turkey). Investigated

sections are indicated as AS I, II, III and IV and KA I, II and III.

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name of A ag˘

yaylabel was Kartoz, after which the Kartoz

Formation was named (Dumont 1976; Gindl 2000; Lukeneder
et al. 2012; Lukeneder & Lukeneder 2014).

A particular depositional situation is marked by the small,

Triassic (Late Carnian – Julian 2) blocks containing am-
monoid mass occurrences, which are included within a Ceno-
zoic conglomerate fan (Deynoux et al. 2005), mainly formed
by Triassic sediments from the nearby area at AS III (Figs. 1, 3).

The Yukar

yaylabel ( = Karap

nar, KA) sections (i.e. KA I—

KA III) are located 5 km to the north of the A ag˘

yaylabel

(AS) sections (Figs. 1, 2, 4) near a small road through the vil-
lage. The village is situated at 1328 m to 1340 m above sea
level with N 37°57’69” and E 31°29’12”.

Geological setting

Geologically the area is located on the Anamas Dag˘ car-

bonate platform or Anamas-Akseki Autochthonous. The Ana-
mas Dag˘ is part of the so-called Taurus-Platform-Units
between the Antalya Suture in the South and the I

zmir-Ankara

Suture  in  the  North,  south  of  the  Isparta  Angle  (Robertson
1993;  enel 1997; Andrews & Robertson 2002; Robertson et
al.  2003).  The  Anamas-Akseki  Autochthonous  ( = Karaca-
hisar-Autochthonous)  includes  Middle  to  Upper  Triassic
limestones,  marlstones  and  shales  of  up  to  500 m  thickness
(Gindl  2000).  The  geology  of  southwestern  Turkey  and  the
Anamas Dag˘ carbonate platform has been extensively inves-
tigated by Özgül & Arpat (1973), Dumont & Kerey (1975),
Monod (1977), Poisson (1977), Gutnic et al. (1979), Robert-
son (1993, 2000),  enel (1997), and Robertson et al. (2003).

The deposits of this area belong to two formations, the strati-

graphically older Kartoz Formation (earliest Carnian), and the
younger Kas

mlar Formation (Carbonate, Marlstone and Shale

member; Lukeneder et al. 2012), which reaches from Early to
Late Carnian age (Julian 2 – Tuvalian 1). The fossil fauna
(Lukeneder & Lukeneder 2014) reported within this work de-
rives from the Kas

mlar Formation with Lower Carnian to Up-

per Carnian sediments (Julian 2 – Tuvalian 1; Fig. 2).

The paleogeographic domain of the Anatolian System

(Taurus  Mts,  Turkey)  was  characterized  during  Triassic
times  by  microplates  located  in  the  middle  of  the  western
Tethys Ocean. The investigated succession was deposited in
an intra-shelf basin of equatorial paleolatitude at the western
end  of  the  ‘Cimmerian  terranes’  or  ‘Cimmerian  blocks’
( engör et al. 1984; Scotese et al. 1989; Dercourt et al. 1993,
2000;  Scotese  1998,  2001;  Gindl  2000;  Stampfli  &  Borel
2002; Stampfli et al. 2002; Lukeneder et al. 2012; Lukeneder
& Lukeneder 2014). This area was located between the ‘old’
Paleotethys in the North and the Neotethys in the South dur-
ing the Late Triassic (Carnian, 228—216 Ma – Gindl 2000;
Gradstein  et  al.  2012).  While  the  Paleotethys  Ocean  under-
went  subduction  along  the  southern  margin  of  Eurasia,  the
young Neotethys Ocean (southern branch of Neotethys, sensu

engör & Y

lmaz 1981) was widened between the African

continent and the Cimmerian terranes, which consisted of Tur-
key, Iran, Afghanistan, Tibet, and Malaysia (Golonka 2004).
In the North of these Cimmerian terranes the I

zmir-Ankara

Ocean (northern branch of the Neotethys, sensu engör &

lmaz 1981) and the ‘old’ Paleotethys were still open to the

East (Tekin et al. 2002; Golonka 2004; Tekin & Göncüog˘lu
2007; Göncüog˘lu et al. 2010).

Lithology and facies

The Triassic succession of A ag˘

yaylabel starts with an an-

gular  unconformity  above  Carboniferous  rocks  (Dumont
1976;  Gindl  2000).  The  main  formations  are  the  Middle  to
Upper  Triassic  Kartoz  Formation  (Late  Carnian)  and  the
Kas

mlar Formation (uppermost Lower Carnian to Upper

Carnian). The Kartoz Formation consists of shallow-water
platform carbonates with thick-shelled bivalves (megalo-
donts) and corals. In contrast, the overlying Kas

mlar Forma-

tion starts disconformably (i.e. hiatus) with an 8-m-thick pile
of  deeper-water  limestones;  this  precedes  12 m  marlstone-
part into shales (Fig. 2). The section is dated based on con-
odonts,  ammonoids,  and  halobiids  (Lukeneder  et  al.  2012;
Lukeneder  &  Lukeneder  2014).  A  detailed  age  assignment
follows  Krystyn  et  al.  (2002)  and  Gallet  et  al.  (2007).  The
strata dip approximmately 50° towards the Northeast.

The studied successions at A ag˘

yaylabel ( = Kartoz) and

Yukar

yaylabel ( = Karap

nar) start with shallow-water lime-

stones  of  the  Kartoz  Formation,  with  thick-shelled  bivalves
and corals (Lukeneder et al. 2012; Lukeneder & Lukeneder
2014; Fig. 2). This phase ends with a corroded and iron oxide-
stained  dissolution  surface  (without  any  traces  of  boring),
pointing  probably  to  subaerial  exposure  or  sedimentological
omission.  The  Kartoz  Formation  represents  a  drowned  car-
bonate  platform  and  is  disconformably  overlain  by  deeper-
water,  hemipelagic,  black  limestones  of  the  Kas

mlar

Formation, which includes, at its base (i.e. 1.8—16.0 m) levels
of thin ammonoid floatstone- and packstone layers (Kasimlar-
celtites beds = acme range zone – Fig. 2). This lower part is
followed by a thick slump breccia (e.g. preserved at AS I and
AS IV), containing up to meter-sized patch reef blocks (inter-
preted  as  Cipit-boulders  by  Lukeneder  et  al.  2012)  together
with  comparably  small  ammonoid  coquinas  and  filament
limestone components. The Lower to Upper Carnian bound-
ary is marked by a change to grey limy marlstones, with rare
ammonoid-  and  pelagic-bivalve-bearing  layers,  passing  up-
wards into a thick pile of sterile dark shale with thin silty and
rare  siliciclastic  interbeds.  Microfacies  analyses  identify  the
Carnian  depositional  system  around  A ag˘

yaylabel and

Yukar

yaylabel as an intrashelf platform environment grading

upwards into deeper zones, influenced by pelagic conditions
(Lukeneder et al. 2012).

All sections where the ammonoid mass occurrence of

Kasimlarceltites exists start with shallow water carbonates of
the Kartoz Formation (Figs. 2, 3). Coral-bafflestones, megalo-
dontid limestones and shallow water breccias predominate.
This facies is characterized by a dense abundance of in-situ
corals and megalodontid shells (Lukeneder et al. 2012),
trapped by a cortoid grainstone matrix representing an open
platform environment.

Directly above an unconformity, the younger Kas

mlar

Formation is divided into the Carbonate member (units A, B,
and C), the Marlstone member, and the Shale member. The

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lowermost  Carbonate  member  appears  at  the  base  with  the
Kasimlarceltites beds (acme zone). The Kasimlarceltites beds
are bioclastic pelagic wackestones, which were deposited in a
deep  shelf  margin  or  mid  ramp  position  (Lukeneder  et  al.
2012;  see  Flügel  1978,  2004  for  facies  types).  The  faunal
spectrum  mirrors  two  different  source  areas  of  the  bioclastic
input: thin-shelled bivalves (halobiids), which are most com-
mon,  represent  the  autochthonous  component  (deeper  shelf
margin), whereas the original habitat of the benthic foramin-
ifera  (lagenids),  the  thick-shelled  bivalves  (megalodontiids),
and  the  large,  low-spired  gastropods  was  situated  on  a  fore
slope  or  on  a  shallow  marine  ramp.  Tilted  geopetal  fills  of
gastropods  and  ammonoids  (e.g.  Kasimlarceltites),  together
with clasts of eroded semilithified sedimentary layers (‘plas-
ticlasts’),  prove  episodic  erosion,  downward  transport,  and
resedimentation. Bioturbation has mostly obliterated the origi-
nal arrangement of autochthonous fine grained sediments (to-
gether  with  thin-shelled  bivalves  and  radiolaria)  alternating
with coarser-grained ‘tempestitic’ or ‘turbidtitic’ layers. When
the original layering is preserved, bedding planes are strongly
affected  by  stylolites.  Common  authigenic  pyrite  (mostly
well-developed as cubic crystals) in different levels of the sec-
tion is probably of synsedimentary origin. As indicated by the
high  abundance  of  halobiid  bivalves  and  the  dark-coloured
sediment, it could point to temporary dysaerobic conditions of
the bottom waters and/or the pore-fluids of the unconsolidated
sediments (Flügel 1978, 2004; McRoberts 2010).

Material and fossil assemblage

The fossil material was collected by the authors (1995—2011),

by Mathias Harzhauser and Franz Topka (both Natural History
Museum Vienna), and on earlier field excursions (1980—1997)
by  Leopold  Krystyn,  Andreas  Gindl  and  Philip  Strauss  (ex-
cursion  organized  by  the  University  of  Vienna).  All  speci-
mens  described  within  this  study  have  been  extensively
collected  from  the  Kas

mlar Formation at the sections

A ag˘

yaylabel and Yukar

yaylabel, which consists of a Lower

Carnian (Austrotrachyceras austriacum Zone) ammonoid
fauna (Lukeneder & Lukeneder 2014). The ammonoid fauna
contains ammonoids of all ontogenetic stages.

The ammonoids are well preserved, phragmocones are

mostly filled with secondary calcite and the shell is neomor-
phically  replaced  by  secondary  calcite.  Due  to  the  fact  that
draught  filling  ( = draft  filling;  Seilacher  1968;  Maeda  &
Seilacher  1996;  Olivero  2007)  is  absent,  which  led  to  a  pre-
served  primary  phosphatic  siphuncle,  a  fast  deposition  and
sedimentation after death can be assumed. Only a few speci-
mens of the genera Kasimlarceltites, Klipsteinia, Anasirenites
and  Megaphyllites  show  suture  lines.  A  total  of  479  am-
monoid specimens, two nautiloid specimens and four coleoid
specimens  have  been  collected  (Lukeneder  &  Lukeneder
2014). The ammonoid assemblages consist of 12 ammonoid
genera  with  Kasimlarceltites,  Spirogmoceras,  Sandlingites,
Klipsteinia,  Neoprotrachyceras,  Sirenites,  Anasirenites,
Paratropites,  Trachysagenites,  Proarcestes,  Megaphyllites,
Joannites,  Simonyceras,  containing  13  species,  a  single  co-
leoid genus (Atractites), and a single nautiloid species.

The ammonoids are clearly dominated by the genus

Kasimlarceltites with more than 100 million specimens
(counting method: see Mayrhofer & Lukeneder, in prep.),
due to the fact that it is the main faunal element of the mass
occurrences, followed by Sirenites with 56 specimens.

The matrix contains mainly juvenile to adult halobiid bi-

valves  (i.e.  Halobia  rugosa),  megalodontid  shells,  gastro-
pods (i.e. Omphaloptycha type), chaetitids, corals, calcareous
sponges,  sponge  spicules,  foraminifera,  radiolaria,  dasycla-
daceaes,  cyanobacteria,  peloids,  and  planktic  crinoids  (e.g.
Osteocrinus). Lower Carnian conodonts are present with Gla-
digondolella and Metapolygnathus.

Detailed stratigraphic sections of the Lower to Upper

Carnian interval were measured and described from
A ag˘

yaylabel ( = Kartoz, AS) and Yukar

yaylabel ( = Karap

nar,  KA).  325  thin-sections  of  200  layers,  collected  in  2007
and 2012, were made and used for petrographic studies. Ad-
ditional  sectioning  and  polishing  was  performed  on  block
samples  in  longitudinal,  horizontal  and  tangential  (90°)  di-
rections for reconstructions of the orientation and alignment
of  the  ammonoids  (see  Potter  &  Pettijohn  1977;  Futterer
1982; Olivero 2007) within the mass occurrence. The corre-
sponding bed numbers are indicated by the abbreviation AS
(for A ag˘

yaylabel) and KA (for Yukar

yaylabel = Karap

nar),

by the corresponding succession number (as there are several
successions at each locality AS I—AS IV resp. KA I—KA III)
and the corresponding bed number (e.g. AS I/1 = sample from
A ag˘

yaylabel, succession I, bed 1). Additional facies inves-

tigations were conducted under a dissecting microscope (Zeiss
Discovery V20) with attached digital camera (AxioCam
MRc5). Sectioning and photographing were done at the Natu-
ral History Museum in Vienna (NHMV = NHMW).

Detailed petrographic analyses were done using a petro-

graphic  polarization  microscope  from  Leica  (Leica
DDM4500P)  and  a  digital  camera  (Leica  DFC4420).  Sec-
tioning and photographing were done at the Natural History
Museum  Vienna  and  at  the  University  of  Vienna  (Depart-
ment of Petrography).

The material is stored within the collection of the Geologi-

cal-Paleontological Department of the NHMV. The inventory
numbers  of  blocks  are:  for  AS I  NHMW  2012/0133/0551—
0561, for NHMW AS II 2014/0094/0001, for AS III NHMW
2014/0095/0001,  for  AS IV  NHMW  2014/0091/0001—0007,
for  KA I  NHMW  2014/0092/0001—0003,  for  KA  II  NHMW
2014/0093/0001—0004, and for KA III 2014/0096/0001.

Biostratigraphy

The sections at A ag˘

yaylabel (AS I—AS IV) and the bio-

chrono-stratigraphically equivalent sections at Karap

nar

(KA  I—KA  IV)  comprise  about  50 m  of  essentially  calcare-
ous beds passing into marlstones (in part marly limestones)
and  shaly  beds  with  considerable  siliciclastic  input  at  the
top. The lowermost part is represented by the Kartoz Forma-
tion,  an  Upper  Triassic  (earliest  Carnian;  Fig. 2)  light-grey,
shallow-water carbonate succession. It comprises corals and
thick-shelled  bivalves.  Neoprotrachyceras  sp.,  found  in  the
lower part of bed AS I/2, dates at least the top of the Kartoz

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Fig. 3. Localities and detailed views on the Kasimlarceltites layers within shallow water carbonates from the Kas

mlar Formation of the

sections around A ag˘

yaylabel (AS I, AS II, AS III, AS IV). A – The locality A ag˘

yaylabel 1 (AS I); B – Detail of the Kasimlarceltites

acme zone at AS I, base of the Kas

mlar Formation; C – The locality A ag˘

yaylabel 2 (AS II); D – Detail of the Kasimlarceltites acme

zone at AS II; E – The locality A ag˘

yaylabel 3 (AS III) within Cenozoic (middle Miocene) conglomerates of the Köprüçay Formation;

F – Detail of the conglomerates comprising Triassic blocks with Kasimlarceltites accumulations; G – The locality A ag˘

yaylabel 4 (AS IV);

H – Detail of the Kasimlarceltites acme zone at AS IV, base of the Kas

mlar Formation.

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Formation as Julian 2 (Austrotrachyceras austriacum Zone).
A hiatus between this top and the overlying Kas

mlar Forma-

tion cannot be excluded for all sections.

The overlying Kas

mlar Formation can be divided into

three ‘members’: a carbonate member, a marlstone member,
and a shale member (Lukeneder et al. 2012). The carbonate
member starts with dark-grey to black, thin-bedded lime-
stones containing ammonoid-rich beds with a nearly mono-

specific assemblage of Kasimlarceltites and very rare Sireni-
tes,  represented  by  floatstones  of  latest  Early  Carnian  age
(Julian  2/II;  Fig. 2).  Towards  the  top  thick  slump  breccias
follow; they contain up to meter-sized patch-reef blocks to-
gether with small ammonoid coquinas and filament-limestone
components. The overlying well-bedded (cm—dm thick) peloi-
dal filament-wackestone with Lower Carnian conodonts (e.g.
Gladigondolella  tethydis  and  Metapolygnathus)  and  an  am-

Fig. 4. Localities and detailed views of the Kasimlarceltites layers within shallow water carbonates from the Kas

mlar Formation of the

sections around Karap

nar (KA I, KA II, KA III). A – The locality Karap

nar 1 (KA I); B – Detail of the Kasimlarceltites acme zone at

KA I, base of the Kas

mlar Formation; C – The locality Karap

nar 2 (KA II); D – Detail of the Kasimlarceltites acme zone at KA II, mid-

dle part of the Kas

mlar Formation; E – The locality Karap

nar 3 (KA III); F – Detail of the Kasimlarceltites acme zone at KA III, base

of the Kas

mlar Formation.

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monoid fauna dominated by the genera Neoprotrachyceras
and Anasirenites are of latest Early Carnian age (Anasirenites
level of Julian 2/IIb). The bioturbated matrix contains juve-
nile halobiid bivalves, sponge spicules, radiolaria, peloids,
and planktic crinoids ( = Osteocrinus sp.).

The interval with the comprising Kasimlarceltites acme

zone (lowermost Carbonate member) represents the latest Early
Carnian (Julian 2 – Lukeneder & Lukeneder 2014; Fig. 2).

The Kasimlarceltites mass occurrence

The Kasimlarceltites mass-occurrence resp. acme zone is

quite common in the area around A ag˘

yaylabel and Karap

nar.

Fig. 5. A – Surface of an accumulation layer of Kasimlarceltites, uncoated, NHMW 2012/0133/0558; B – Same specimen coated with
ammonium chloride, NHMW 2012/0133/0558; C – Top-to-bottom thin section of the same layer with densely packed ammonoid shells,
NHMW 2012/0133/0559; D – Polished slice of an ammonoid accumulation layer at section AS IV – block 1—slice C (89 mm, frontal
view), NHMW 2014/0091/0001; E – Ammonoid accumulation surface from section AS IV, NHMW 2012/0133/0558; F – Kasimlarceltites
krystyni, lateral view, holotype, NHMW 2012/0133/0014. Kas

mlar Formation, Carbonate member Unit A, Austrotrachyceras austriacum

Zone (Julian 2). Each scale bar represents 1 cm.

Fig. 6. Terminology and orientation of frontal and orthogonal sec-
tions measured from the locality A ag˘

yaylabel I, with indicated

present-days cardinal directions for blocks AS I, AS II, AS III and
AS IV.

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For a detailed data set of the A ag˘

yaylabel area (AS Ia,

AS Ib, AS II, AS III) and the Karap

nar area (KA I, KA II,

KA III) on lithology, microfacies, geochemistry, geophysics,
fossil  content  and  environment  see  Table 1.  The  standard
Microfacies  zones  (i.e.  SMF  zones)  are  in  accordance  with
Flügel  (1978,  2004;  see  also  Carrillat  &  Martini  2009)  and
Lukeneder et al. (2012).

The A ag˘

yaylabel (Kartoz) area

Locality: A ag˘

yaylabel Ia, AS Ia (Figs. 1, 2, 3A—B; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Debris flow, event bed

Locality: A ag˘

yaylabel Ib, AS Ib (Figs. 1, 2, 3A; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Debris flow

Locality: A ag˘

yaylabel II, AS II (Figs. 1, 2, 3C—D; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Debris flow, event bed

Locality: A ag˘

yaylabel III, AS III (Figs. 1, 3E—F; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Primary debris flow, secondary fan-delta
conglomerates

Locality: A ag˘

yaylabel IV, AS IV (Figs. 1, 2, 3G—H; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Turbidite, event bed

The Karap

nar (Yukar

yalabel) area

Locality: Karap

nar I, KA I (Figs. 1, 2, 4A—B; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Debris flow, event bed

Locality: Karap

nar II, KA II (Figs. 1, 2, 4C—D; Table 1)

Lithology: Dark grey to black limestone
Interpretation: Debris flow, event bed

Locality: Karap

nar III, KA III (Figs. 1, 2, 4E—F; Table 1)

Lithology: Dark grey to black limestones
Interpretation: Debris flow, event bed.

The Kasimlarceltites mass occurrence as an

abundance zone

An abundance zone or acme zone is a stratum or body in

which the abundance of a particular taxon or specified group
of  taxa  is  significantly  greater  than  is  usual  in  the  adjacent
parts  of  the  section  (Salvador  1994;  Murphy  &  Salvador
1999). Its boundaries are made of biohorizons and the name
is  given  by  the  abundant  taxon  or  taxa  (Lukeneder  2003b).
Biohorizons  are,  for  example,  characterized  by  a  sharp  and
significant  biostratigraphic  change  within  the  fossil  assem-
blage  and/or  the  change  of  frequency  of  its  members  (see
Salvador 1994; Steininger & Piller 1999). The latter authors

recommended the term biohorizon to be used instead of the
terms  ‘surface’,  ‘level’,  ‘marker’  and  ‘datum  planes’.  Such
biohorizons are of great importance for lateral correlation over
wide distances. Densely fossiliferous layers were also termed
as  ‘shell  beds’,  ‘coquinas’,  ‘concentration-Lagerstätten’,
‘lumachelles’,  ‘bioclastic  limestones’  and  ‘bioclastic  beds’
(Brett & Seilacher 1991; Kidwell 1991a,b; Martin 1999).

Patterns which lead to beds with a notable abundance of

ammonoid shells have been called ‘inter-regional mass oc-
currences’ or ‘stray occurrences’ (Kemper et al. 1981). Such
abundance zones are of exceptional value for intra-regional
correlation in the Triassic.

An Upper Triassic monotonous ammonoid assemblage

representing a thickness of at least one single bed up to a few
meters,  were  reported  and  taxonomically  described  by
Lukeneder et al. (2012 – ‘ammonite floatstones with Ortho-
celtites’;  Orthoceltites = old  synonym  of  Kasimlarceltites)
and  Lukeneder  &  Lukeneder  (2014  –  ‘Kasimlarceltites
mass occurrence’).

At the investigated A ag˘

yaylabel (AS I—IV) and Kas

mlar

(KA I—III) sections the Kasimlarceltites ammonoid abun-
dance zone (characterized by abundance or mass occurrence
of ammonoids) could be detected for the first time. The
names of the separated beds were given following the domi-
nating genus Kasimlarceltites.

The Kasimlarceltites acme zone starts at every section

from A ag˘

yaylabel and Karap

nar directly above the shal-

low water carbonates of the Kartoz Formation (Fig. 2). At
AS Ia and AS IV, the Kasimlarceltites acme zone is capped
by an interval of debris flow deposits comprising Cipit boul-
ders (Lukeneder et al. 2012). At the Karap

nar sections, the

upper edge of the sections is not visible, hence ending within
the Kasimlarceltites beds. The acme zone appears with devi-
ant ranges or thickness in distinct localities (Fig. 2; Table 1).
It  occurs  at  the  sections  AS Ia  with  1.8 m,  at  AS  Ib  with
8.0 m,  at  AS II  in  single  blocks,  at  AS III  within  Cenozoic
conglomerates  (Deynoux  et  al.  2005)  as  reworked  blocks,
and at AS IV with 4.0 m. The acme zone sections at Karap

nar appear at KA I with 6.0 m, at KA II with 16.5 m and at
KA III with 5.0 m. The most undisturbed and entire section
occurs at KA II. Other sections (e.g. AS Ia, AS IV, KA I,
KA III) are characterized by tectonics or seem to be affected
by more or less strong shearing mechanisms.

Biostratinomy and taphonomy

The biostratinomy is defined as the sum of environmental

factors that affect organic remains between death and the final
burial or embedding (Müller 1963; Brett & Baird 1986; Mar-
tin 1999). Biostratinomy is thus a very important part of work
in taphonomy, the study of the entire post mortem history of
organic remains resulting in fossil material (Fernández-López
& Fernández-Jalvo 2002; Fernández-López 2007).

The taphonomic investigations of fossil cephalopod as-

semblages  provide  insight,  not  only  into  the  autecology  of
these  organisms,  but  also  into  their  paleoenvironment  and
paleocommunity  structure  (Brett  &  Baird  1986;  Allison  &
Briggs 1991; Brett & Seilacher 1991; Bottjer et al. 1995).

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Table 1: Parameters of the Kasimlarceltites Abundance Zone and measured blocks at different localities in the A ag˘

yaylabel and Karap

nar

area. Data are measured within this study for the first time.

Sediments of the lower Upper Triassic (Carnian) of the

Kas

mlar Formation appear in the lower part with well pre-

served  and  mostly  entire  ( > 95 %)  ammonoid  specimens
(Fig. 5).  The  distinct  ammonoid  shell  concentration  beds
within  single  logs  and  different  localities  (i.e.  A ag˘

yaylabel

and Karap

nar) differ in lateral distribution, thickness, sedi-

mentological features, packing of shells, diversity of organ-

isms, alignment and taphonomic processes (Figs. 6, 7, 9, 10;
Tables 1, 2, 3; see also Fürsich & Pandey 1999).

The ammonoids are tiny with a maximum diameter of

33 mm. They show an involute more spheroidal shape and are
accumulated  in  the  beds AS Ia–  beds 4,  6;  AS IV  –  bed 8;
KA I  –  beds 10,  12;  KA II  –  beds 2,  11,  122;  KA III  –
bed 8.  Less  than  5 %  of  those  accumulated  ammonoids  are

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preserved  only  with  fragmentation.  Small  accumulated  am-
monoids  of  all  ontogenetic  stages  (i.e.  Kasimlarceltites)  as
well  as  shell  fragments  in  body  chambers  of  adjacent  large
ammonoids  can  be  found  within  the  above  mentioned  beds,
which might hint at the effect of agglomeration and comminu-
tion  by  dense  sediment  flows  with  a  laminar  internal  flow.
Both,  straight  shells  (e.g.  Atractites)  and  also  planispiral
coiled  shells  (e.g.  Kasimlarceltites,  Sirenites,  Neoprotrachy-
ceras) are present within the Kasimlarceltites acme zone. Ero-
sional features, as described by Fernández-López (2007) and
encrustation  of  ammonoids  are  absent.  Whilst  in  some  thin
and  distinct  layers  ammonoids  are  accumulated  in  masses
with an almost horizontal alignment (Figs. 8—10), ammonoids
are  less  abundant  in  the  limestone  beds  between  the  am-
monoid mass occurrence beds (Figs. 5, 6, 9).

In accumulated mixed assemblages (i.e. autochthonous,

parautochthonous and allochthonous; Martin 1999), detected
at the A ag˘

yaylabel outcrops AS Ia (Lukeneder et al. 2012),

AS II—IV and the Karap

nar KA I—III sections, females

( = macroconchs)  and  corresponding  males  ( = microconchs)
are  found  together.  In  most  cases  even  ammonitellae,  juve-
nile  and  adult  stages  are  detected  in  the  same  layers.  Some
ammonoid  specimens  show  different  sediment-infillings
within  the  body  chamber  (coarser  material)  compared  to  the
surrounding  finer  limestone  chamber  (‘normal’  sedim-
entation = ambient sediment). All of these facts might point to
post-mortem, biostratinomic mixing of ecologically age-sepa-
rated populations as discussed by Olóriz (2000) and Olóriz &
Villaseñor (2010). Due to the long body chamber (i.e. mesod-
ome-longidome)  in  Kasimlarceltites,  geopetal  structures  (i.e.
sparry  calcite  on  top)  are  also  observable  in  numerous  body
chambers (see Olivero 2007; cf. Seilacher 1968), hence body
chambers were not entirely filled. Geopetal structures are gen-
erally aligned in almost identical directions. No serious mix-
ture and dislocation of geopetal alignments occurs (Figs. 7, 9).
Furthermore, the mixed assemblages comprise a considerable
amount of bivalves, gastropods, sponges and corals from shal-
lower  environments  from  a  nearby  platform  or  upper  ramp
(Lukeneder et al. 2012).

Fig. 7. Thin-section photographs of accumulation layers and facies from the Kas

mlar Formation at A ag˘

yaylabel and Karap

nar. A—H – Note

the well preserved shell geometry indicating very early cementation (i.e. early selective lithification): A – Bioclastic floatstone with the abun-
dant ammonoid Kasimlarceltites and halobiids, accompanied by rare gastropods. The ammonoids and gastropods are filled with coarse sparry
calcite floating in micritic matrix, base of Kasimlarceltites Formation, AS I – bed 6, NHMW 2012/0133/0560; B—C – Very irregular
packing = “injection” of heterometric bioclasts (B), and the same but with larger heterometric range (C); B – Bioclastic floatstone-packstone,
characteristic sponge-accentuated Cipit facies dominated by Kasimlarceltites and sponges, accompanied by Sirenites, gastropods and corals,
Kas

mlar Formation, AS I – bed 18, NHMW 2012/0133/0561; C – Bioclastic floatstone-packstone, sponge-accentuated Cipit facies domi-

nated by Kasimlarceltites and sponges, accompanied by gastropods and corals, peloidal matrix, Kas

mlar Formation, AS II, NHMW 2014/

0094/0001; D – Allochthonous block with mixed packstone (left) and floatstone (right) areas, dominated by Kasimlarceltites and sponges,
accompanied by Sirenites, gastropods and corals; found within the Cenozoic (middle Miocene) conglomerates, Köprüçay Formation, AS III,
NHMW 2014/0095/0001; shelter effects favouring very dense packing; E – Bioclastic packstones with the abundant ammonoid Kasimlarcel-
tites above laminated, peloidal packstone layers, base of Kas

mlar Formation AS IV – bed 8, NHMW 2014/0091/0008; sheltering by fragmen-

ted body chamber infilled by a very fine comminute matrix; F – Bioclastic packstones with the abundant ammonoids Kasimlarceltites and
Sirenites, peloidal matrix, base of Kas

mlar Formation, KA I, NHMW 2014/0092/0003; comminute, bioclastic matrix and reworked ammonoids;

G – Bioclastic wackestone with a redeposited, floated Kasimlarceltites shell (note dislocated geopetal structure, different infilling and colour)
at the base, matrix contains radiolarian, ammonitellae and juvenile ammonoids, middle Kas

mlar Formation, KA II, NHMW 2014/0093/0004;

very fine matrix with diagenetic patches; H – Bioclastic wackestone with floated Kasimlarceltites shells in the middle, and peloidal packstone
at the base, peloidal matrix contains radiolarian, ammonitellae and juvenile ammonoids, base of Kas

mlar Formation, KA III, NHMW 2014/

0096/0001; geopetals indicate ammonoid reworking. Thin-sections are orientated in upright position. Each scale bars represent 1 mm.

Fig. 8.  A  –  Primary  angles  in  shell  axes  (i.e.  2°—3°)  caused  by
the  ontogenetic  variation  of  whorl  breadth  in  mid-aged  and
adult  Kasimlarceltites  (adult = holotype,  NHMW 2012/0133/0014);
B – Shell axes (black pies, white arrow) and geopetal axes (grey
pies, black arrow) of Kasimlarceltites and Sirenites within a sample
KA II – block 4 (sections D—E—F).

Ammonoid shell alignment

Angles and orientation of shells and body chambers (both in

respect to the horizontal block surface) from 20 blocks with 34

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polished  slices  from  4  different  sites  (i.e.  AS Ia,b,  AS IV,
KA I, KA II) were measured in addition with maximal visi-
ble  diameters  on  the  slices  and  thin  sections.  Additional
measurements  were  made  on  visible  geopetal  structures.
2367 ammonoid specimens were analysed on slices and sec-
tions.  Ammonoid  shell  directions  and  alignment  were  not
measured in chaotic, transported ‘Cipit’ boulders from debris
flow  deposits  at  AS Ib  –  bed 18  and  within  Triassic  boul-
ders and blocks incorporated in Cenozoic gravels (Deynoux
et al. 2005) of AS III, because they do not mirror the primary
mechanism of orientation and transport of ammonoid shells,
due to their repeated transport and reorientation.

The most reliable data were obtained from AS I where

blocks could be taken orientated (Fig. 6). The front section of
the AS I blocks (as taken in the field) is orientated with a di-
rection from NE to SW (NE is left on the block, orientated to-
ward the valley). On a horizontal line, marking the base and
surface of the layers and blocks, 0° is located at the right
(SW), 90° at the top, 180° at the left (NE) and 270° at the base

(Figs. 9, 10). In addition, where possible, slices at 90°  ( = or-
thogonal) to the frontal view were performed to get ideas on
the  three  dimensional  orientation  of  the  ammonoids  in  the
blocks.  To  gain  the  true  dip  direction  (Nichols  2009)  of  the
ammonoid shell ‘plane’ the calculation needs two different ap-
parent dip measurements (i.e. ammonoid shell axes; Fig. 11).
This implies a direction (i.e. only AS I blocks) of the later slic-
es rotated by 90° to a direction of NW (front) to SE (back).

A clear orientation and alignment (Potter & Pettijohn

1977) can be detected in the accumulation layers.

The predominant orientation of shell axes (only AS Ia

with cardinal direction) versus the layer base/surface of
blocks in frontal and orthogonal slices was analysed at
A ag˘

yaylabel (Figs. 6, 10, 11, Tables 2, 3).

F r o n t a l
AS Ia, AS IV
O r t h o g o n a l
AS Ia, AS IV

Fig. 9. Ammonoid shell alignment in Carnian blocks from A ag˘

yaylabel and Karap

nar, with indicated accumulation layers and measure-

ments. A – AS I – block 1, slice B (orthogonal view), NHMW 2012/0133/0554; B – AS IV – block 1, slice C (89 mm, frontal view),
NHMW 2014/0091/0003; C – KA I – block 1, slice A (frontal view), NHMW 2014/0092/0001; D – KA II – block 4, slice A,B,C
(0 degree, frontal view), NHMW 2014/0093/0003. Primary angles in shell axes indicated by arrows, arrowheads directed to the body cham-
ber ( = maximum whorl breadth). Sketches (below sections) of the corresponding shell axes (black arrows) of each ammonoid specimen
within the event layer (grey shading). Scale bar represents 1 cm.

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Within accumulation layers

from the Karap

nar area the

orientation of shell axes (not in
cardinal  direction)  appears  in
frontal  and  orthogonal  slices
(Fig. 10,  Tables 2,  3).  Layers
in  KA I  can  be  separated  in
two parts (accumulation = event
layer  and  above  with  normal
sedimentation), KA II (geopetal
axe in body chambers varies in
angle section).

F r o n t a l
KA I, KA II
O r t h o g o n a l
KA I, KA II

The internal fabric of the ac-

cumulation  layers  (i.e.  am-
monoid  shells)  ranges  from
densely  packed,  ammonoid
shell  supported  packstones
(e.g. AS IV – bed 8; KA I –
bed 10)  to  ammonoid-float-
stones  (AS Ia  –  beds 4,  6;
Figs. 7).  The  event  layers  are
heterogeneous,  with  cases  of
shell  imbrication  in  parts  (see
imbrication in turbidite Bouma
interval T

, Bouma et al. 1982;

Eberli  1991)  and  cases  of
loosely  packed  or  floating
specimens  in  different  parts.
Even in single blocks, the den-
sity  diverges  from  the  base,
mid  part  and  top  of  beds
(Fig. 9).  The  matrix  infills  the
body  chambers  and  the  space
between shells. The imbricated
shells  show  a  horizontal  or  at
least  low-angle  deposition.
Vertical  or  perpendicular  am-
monoid  specimens  are  ex-
tremely  rare  in  accumulation
layers  of  A ag˘

yaylabel and

Karap

nar. An alignment of the

biogenic  particles  (i.e.  am-
monoid  shells)  was  observed
in the majority of the accumu-
lation layers (AS I – beds 4, 6;
most beds at KA I, KA II). Al-
though  the  alignment  within
most

accumulation

layers

(AS I  –  beds 4,  6;  most  beds
at KA I, KA II) took place with-
in  the  sediment,  in  blocks 1
and  2  (corresponding  to  the
same layer) in AS IV and KA II

Table 2:

Results on the biofabric,

orientation and size classes of

Kasimlarceltites

krystyni

at different localities in the

yaylabel

and Karap

nar

area,

frontal

view.

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Table 3:

Results

the

biofabric,

orientation

and

size

classes

Kasimlarceltites

krystyni

different

localities

the A

yaylabel

and

Karap

nar

area,

orthogonal

view.

(e.g.  blocks 1,  2)  the
aligned  ammonoids  are
accumulated  in  a  shell
coquina on the top of lam-
inated, peloidal ‘silts’.

The body chambers are

filled  with  the  same  ma-
trix,  which  is  formed  by
peloids  from  a  shallower
environment,  represent-
ing  the  primary  deposi-
tional area. The shells are
densely

packed,

well

sorted and in contact with
numerous other shells.
The densely packed accu-
mulation layer marks an
ammonoid

packstone.

Siphonal  structures  are
preserved, hindering sedi-
ment to fill the phragmo-
cones (see Olivero 2007),
hence rapidly buried after
death exhibiting the final,
hollow  ‘particles’  em-
bedded  in  the  peloidal
layers.  The  matrix  above
and below this event layer
differs  distinctly  as  it
consists of fine mud with
only  rare  and  smaller
ammonoid

specimens,

deposited under ‘normal’
conditions. An additional
block  from  AS IV  (i.e.
block 3)  marks  a  ‘nor-
mal’  calm  deposition  at,
or near the habitat of the
ammonoids,  appearing
with  rare,  entire  am-
monoid  shells  with  al-
most  undisturbed  and
horizontal

alignment

(Fig. 9, Table 1). The
shell

axis

block

AS IV – block 3 shows
a  clear  picture  with
dominant  intervals  at
160—180°/340—360°  with
75.0 %  and  even  more
expressed in the interval
160—200°/340—020°  with
97.2 %.

The acme zone of the

genus

Kasimlarceltites

ranges  due  to  tectonics
from  1.8 m  at  AS Ia  to
16.5 m  at  KA II,  and  is
intercalated by accumula-

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tion event beds (Figs. 2, 3, 4, 7). On the basis of the geological
time scale of Gradstein et al. (2012) a stratigraphical range from
one third to one half of the Austrotrachyceras austriacum Zone
can be assumed, which approximately equals a duration of
200 ky (at AS I, AS IV)—500 ky (at KA II) for the Kasimlar-
celtites abundance zones. This time span is calculated without
consideration of any hiatus or time-averaging, which might
occur. The overall quantity of more than 100 million ammon-
oid specimens is estimated by the data gained from numerous
localities from A ag˘

yaylabel and Karap

nar (e.g. layer thick-

ness, number of mass occurrence layers and specimens etc.)
and the geographical distribution (over 10 km

) of the Kasim-

larceltites abundance zone. Single blocks from both localities
(A ag˘

yaylabel and Karap

nar) with 15

×15×8 cm contain up

to 3500 specimens (Mayrhofer & Lukeneder in prep.).

The calculated mean angle of all combined sections and

specimen  axes  (n 1696)  in  frontal  view  is  almost  horizontal
with 172°/352° (Fig. 10) indicating a shallow orientation (low
angle)  of  ammonoid  shells  throughout  the  various  localities
(Tables 1, 2, 3). Within the polyspecific KA II – block 4 an
increased angle with 164°/344° was observed, as well as an
increased number of steeper orientations in body chambers.
In contrast the mean angle of shell axes in KA II – block 3
is shallow (low angle – almost planar) with 178°/358°. The
measured  angle  is  influenced  by  the  ontogenetic  stage  and
the  morphological differences in whorl height. Adult  speci-
mens show an angle of 2° whereas juveniles appear with 3°
owing to the different whorl expansion rates.

The same situation can be observed in the combined ortho-

gonal data (Fig. 10, Table 3). The mean angle of all specimen
axes  (n 1696)  in  orthogonal  view  is  177°/357°  (172°/352°
frontal),  depicting  a  very  shallow  orientation  of  ammonoid
shells  throughout  the  various  localities.  Anyhow,  an  in-
creased angle of 15°/195° in KA II – block 4, where Kasim-
larceltites occurs together with some specimens of Sirenites,
was  detected.  As  already  observed  in  frontal  view,  the  or-
thogonal  data  strengthen  the  contrasting  picture  with  an  al-
most  horizontal,  very  shallow  mean  angle  of  shell  axes  in
KA II – block 3 by 179°/359° (Fig. 10, Table 3). The lower
accumulation  layer  in  KA I  –  block 1a  (Fig. 10)  shows
mean  angles  of  004°/184°  whereas  the  ‘normal’  sedimenta-
tion part above (KA I – block 1n; Fig. 10) appears with an
increased angle of 018°/198°.

Size groups in Kasimlarceltites

In general the ceratitid genus Kasimlarceltites Lukeneder

&  Lukeneder  (2014)  is  a  small  sized,  1.0  to  3.3 cm,  almost
smooth  ammonoid.  The  abundance  of  different  ontogenetic
stages within the occurring size classes (Fig. 10, Tables 2, 3)
varies  from  locality  to  locality,  even  from  layer  to  layer  in
the  same  locality.  Densely  spaced  sutures  at  the  phragmo-
cone/body  chamber  boundary  of  some  specimens  mark  the
adult  stage  in  numerous  ammonoids.  For  more  detailed  de-
scription  see  Lukeneder  &  Lukeneder  (2014,  fig. 4—5).  At
different localities minimum and maximum range size values
of Kasimlarceltites differs markedly (Fig. 10, Tables 2, 3).

Size ranges and dominant diameters (in mm) are given for

all accumulation layers, in frontal and orthogonal direction

for comparison of potential spatial alignments (Fig. 10, Ta-
bles 2, 3).

A ag˘

yaylabel

F r o n t a l (Fig. 10, Table 2)
AS Ia, AS IV
O r t h o g o n a l (Fig. 10, Table 3)
AS Ia, AS IV

Karap

nar

F r o n t a l (Fig. 10, Table 2)
KA I, KA II
O r t h o g o n a l (Fig. 10, Table 3)
KA I, KA II

The overall impression fixed by the data (i.e. angles and

diameters)  of  all  specimens  measured  (n 1696)  in  frontal
views  shows  a  unimodal  positively  skewed  distribution  of
size  classes.  For  better  comparison  of  distribution,  curve
shapes  and  visualization  of  size  classes,  distribution  curves
were  set  to  100 %  (Fig. 10).  A  clear  dominance  of  small
forms from 2.1—12.5 mm is evident (Fig. 10). The maximum
size class appears at 4.2—6.2 mm. Maximal size classes from
25.2—29.3 mm are rare to absent in most localities. From the
taxonomical  work  by  Lukeneder  &  Lukeneder  (2014)  on
prepared specimens from the same localities, a maximal dia-
meter of Kasimlarceltites is known with 33 mm. The differ-
ence  in  maximal  diameters  of  about  37 mm  shows  the
uncertainty  and  imprecision  in  numerical  measurements  in
slices  and  sections,  since  the  majority  of  excavated  am-
monoids from the sections miss the aperture area and there-
fore fail to reflect the maximum size.

Specimens from AS Ia – block 4, AS Ia – block 3 and

KA I  –  block 1  show  their  maximum  size  in  frontal  view
with  unimodal  curves  (positively  skewed)  at  2.1—10.4 mm.
Somewhat  shifted  curves,  increasing  in  mean  size,  are
present  in  KA II  –  block 2  (bimodal)  and  KA II  –  block 3
with positively skewed, unimodal curves around a maximum
of 4.2—14.6 mm. Exceptional, broad distribution curves occur
in AS IV – block 3 with 2.1—23.0 mm and KA I – block 1
with 2.1—20.9 mm.

Analogously, the picture of all specimens measured (n 675)

in orthogonal views show a unimodal positively skewed dis-
tribution of size classes. The abundance peak also lies within
the  class  4.2—6.2 mm  as  seen  in  the  frontal  view  (Fig. 10).
A  clear  dominance  of  small  forms  from  2.1—10.4 mm  is
established  (Fig. 10).  The  maximum  size  class  appears  at
4.2—6.2 mm.  Maximal  size  classes  from  25.2—27.2 mm  are
rare to absent in some localities.

Specimens from AS Ia – blocks 1, 2, 3 and KA II – block 2

show the maximum size in frontal view with unimodal curves
(positively skewed) at 2.1—10.4 mm. Somewhat shifted
curves, increasing in mean size, with a slightly bimodal mode
are present in AS IV – block 3 and KA I – block 1a (accu-
mulation layer) with positively skewed curves around a maxi-
mum of 4.2—14.6 mm. Exceptional, broad distribution curves
occur in KA II – block 3 with a wide range of 2.1—18.8 mm.

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Variation in diameters and size classes, as reported above for

angles, is also present within single layers, as seen in KA I –
block 1 (a – accumulation, n – normal). The lower accumu-
lation layer (KA I – block 1a; Fig. 10, Tables 2, 3) shows an
abundance of the mean size class 4.2—12.6 mm, whereas the
normal sedimentation part above (KA I – block 1n) appears
with a decreased diameter class of 0.0—6.2 mm. Both distribu-
tion curves are positively skewed but KA I – block 1a shows
weak bimodal distribution. Hence the latter ammonoid assem-
blage comprises juvenile and adult specimens. While juvenile
specimens are compressed in shape, adult specimens are much
more depressed. This might be interpreted as mating commu-
nity, environmental forcing or transport mechanism (e.g. cur-
rent or drift selection of different size or morphology classes).

Discussion

The Kasimlarceltites mass occurrence as an abundance zone

The trigger mechanisms and possible processes, which

cause the abundance and accumulation of ammonoids and the
formation of such widespread ammonoid mass occurrences or

‘event’ layers, are discussed. Current induced alignment, win-
nowing and earth quakes or storm events (Aigner 1985; Brett
& Baird 1986; Seilacher & Aigner 1991; Hips 1998; Radley
&  Barker  1998;  Storms  2001;  Pérez-Lopéz  &  Pérez-Valera
2012) are discussed as trigger factors for such gravity forced
‘event  beds’.  The  described  examples  are  thus  autocyclic
(Einsele et al. 1991b), formed by nonperiodic mass flows and/
or  turbidites  endemic  to  the  tectonic  setting  (Bouma  et  al.
1982; Cook et al. 1982; Howell & Normark 1982; Tucker &
Wright 1990; Potter et al. 2005; Hornung 2008). Final deposi-
tion took place on tectonically unstable slope areas, shown by
the  presence  of  frequent  neptunian  dykes  (see  Flügel  2004;
Črne et al. 2007) within the Kasimlarceltites acme zone.

In the majority of such fossil accumulation beds, several

reasons amplify the primary signal (Kidwell 1986; Kidwell
et al. 1986). Pérez-Lopéz & Pérez-Valera (2012) presented a
tripartite model for storm influenced beds and near platform
carbonate environments with the pot/gutter casts, the tem-
pestite beds and the storm winnowed deposits. According to
Aigner (1985) storm processes can be distinguished in three
distinct physical categories with barometric effects, wind ef-
fects, and wave effects. The causes and features of such rap-
idly deposited ‘event’ sediments are extensively reviewed by

Fig. 10. Compilation of size classes, shell axes and body chamber orientations from Kasimlarceltites in frontal and orthogonal view of all
measured blocks of A ag˘

yaylabel (AS I, AS II, AS III, AS IV) and Karap

nar (KA I, KA II), Kas

mlar Formation, Kasimlarceltites acme

zone, Lower Carnian, Upper Triassic. See text for description and details.

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Einsele & Seilacher (1982) and Einsele et al. (1991a), and
the depositional environment of sandy turbidites and other
mass flows deposits are summarized by Einsele (1991).

As noted by Aigner (1985), storm-generated tempestites

show a clear gradation with fine sediments that is extremely
subtle or lacking in distal tempestite fronts (difficult to sepa-
rate from distal turbidites), which is not seen in the Carnian
layers at A ag˘

yaylabel and Karap

nar. Specifically, he noted

that proximal storm-event layers contain more mixed faunas,
with  allochthonous  specimens,  in  contrast  to  distal  layers,
where  autochthonous  and  parautochthonous  assemblages
predominate,  having  been  reworked  in-situ  (see  also  Gole-
biowski 1991).

Progressing gravity, debris flows show typical transition

to turbidity currents, due to changes in different transporta-
tion phases when variable degrees of debris dilution occur.
Therefore the transportation of debris might indicate net re-
working (pers. comm. Olóriz 2014).

The dynamic history and the influence of sediment transport

to that area during the Carnian can also be traced by the fre-
quent Cipit boulders (directly above the Kasimlarceltites acme
zone), derived from the shallow-platform edge and transported
by gravity down across the slope, as well as by the frequently
silty, turbidite layers in the Kas

mlar shales in the Tuvalian.

Geochemical and geophysical data (Lukeneder et al. in

prep.) will confirm or contradict the idea of drastic changes
in ecological factors (e.g. anoxic events, methane eruptions,
for toxic events see also Noe-Nygaard et al. 1987) during the
sedimentation of the Kasimlarceltites acme zone. Such eco-
logical factors can act as trigger mechanisms for potentially
catastrophic ammonoid mass mortality of the ammonoid de-
posits from Turkey. Such traces are not observed in the rede-
posited layers (episodic deposits in Brett & Baird 1986) and
in  the  ‘normal’,  background  sedimentation  beds  ( = ‘host’
beds)  where  Kasimlarceltites  rarely  occurs.  In  contrast  to
this,  Adamíkova  et  al.  (1983)  discussed  mass  natality  as  a
possible trigger for a lower Barremian (Cretaceous) mass oc-
currence of ammonoids (Adamíkova et al. 1983).

The presence of ammonoid abundance zones (‘ammonoid-

beds’;  characterized  by  abundance  or  mass-occurrence  of
ammonoids)  seems  to  be  related  to  sea-level  rises  or  falls
(Kidwell  1988,  1991a,b,  1993a,b;  Martin  1999;  Lukeneder
2001,  2003b).  Most  probably  ammonoid  mass  occurrences
reflect  transgressive  phases  (see  also  Hoedemaeker  1994;
Aguirre-Urreta & Rawson 1998, 1999). Anyhow, Fernández-
López et al. (2002) argued that ammonoid mass occurrences
deposited within shallow water environments reflect regres-
sive  phases,  whilst  ammonoid  mass  occurrences  deposited
within  deeper  water  environments  reflect  transgressive
trends (Fernández-López et al. 2002). The observation of dif-
ferent preservational features (e.g. sedimentary infilling, en-
crustation, abrasion, bioerosion, reorientation and dispersal)
might  hint  to  the  right  interpretation  (a  shallow  or  deeper-
water environment – Fernández-López et al. 2002).

Biostratinomy and taphonomy

The nearly entire ( > 95 %) well preserved ammonoid spec-

imens of the lower Upper Triassic (Carnian) of the Kas

mlar

Formation,  suggest  no-to-moderate,  non-destructive  trans-
port mechanisms and therefore favour a more autochthonous
to  parautochthonous  nature  of  the  specimens.  Judging  from
internal structures of the limestone beds and the alignment of
the fossil content (Potter & Pettijohn 1977), water saturated
gravity  flows  ( = liquified  flows  –  Mulder  &  Alexander
2001; SEPM 2014) are induced at a medial position between
a  proximal  (near-source)  and  a  distal  depositional  develop-
ment  (see  Aigner  1985).  Hence,  moderate  transportation  of
at  least  some  intraclasts,  plasticlasts  and  bioclasts  such  as
gastropods and in parts ammonoids, is presumed.

The fragmentation (e.g. broken body chamber) of less than

% of the ammonoids provides evidence for no, gentle or

very  weak  post  mortem  transport,  without  breakage  on  the
sea  floor  through  current  effects,  and/or  consequences  of
predation  (Lukeneder  2004a,b).  Anyway,  the  tiny  shells
(max.  33

mm) of Kasimlarceltites were probably resistant

against  breakage  by  impact  of  shells  with  other  bioclasts
transported by the currents. Ruptures reflecting post mortem
histories, caused by current-induced transport before embed-
ding, are absent. It may also reflect the enhanced stability of
ammonoid shells with smaller size, more ‘spheroidal’ shape
and  suggests  that  involute  morphologies  are  more  resistant
to damage compared to other ammonoid shell morphologies
(e.g.  broken  body  chamber  of  Simonyceras;  specimen  fig-
ured on Fig.

11A in Lukeneder & Lukeneder 2014). Further-

more, buccal masses with preserved beak apparatus (e.g.
aptychi like jaws) are completely missing in the Turkish ma-
terial, hinting at more parautochthonous than autochthonous
depositional conditions.

Accumulated small ammonoids (i.e. Kasimlarceltites) of

all ontogenetic stages (e.g. ammonitellae, juveniles to adults)
and shell fragments in body chambers of adjacent large am-
monoids, in combination with cephalopod alignment in sev-
eral single layers (e.g. AS

Ia – beds

4, 6; AS

IV – bed

I – beds

10, 12; KA

II – beds

2, 11, 122; KA

III –

bed

8) also suggest transport-effects on deposition (e.g. mass

flow,  bottom  currents,  winnowing  –  Potter  &  Pettijohn
1977;  Flügel  2004;  Potter  et  al.  2005;  Fernández-López
2007; Nichols 2009).  The accumulation  of the  shells  which
show almost horizontal alignment (Figs.

8, 9, 10) in thin and

distinct layers, is probably due to episodes of reworking and
current-induced removal of sediment (i.e. winnowing). Hori-
zontally aligned specimens were most probably secondarily
re-orientated and aligned during a first gravity flow sedimen-
tation phase.

The absence of any erosional feature (i.e. mechanical- or

bio-erosion;  see  Fernández-López  2007)  or  encrustation  on
any  side  of  the  ammonoids  suggest  calm  environments  with
relatively fast burial processes. The shell transport took place
during single events or phases of slow gravity sediment flows,
as  is  reflected  by  the  occurrence  in  several  separated  am-
monoid layers up to 10

cm in thickness (Figs.

3, 4, 5, 7).

The ‘normal’ occurrence range, with rare specimens of

Kasimlarceltites, clearly shows the inhabitation of that pa-
leogeographical area during the Carnian (Late Triassic) over
a hundred thousand years (Lukeneder & Lukeneder 2014;
for numerical age see Gradstein et al. 2012). In contrast, ac-
cumulation in masses occurs only within single event beds,

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caused by huge storms or tectonically induced earthquakes
( = seismic activity in Einsele 1991; see slope failures in
Potter et al. 2005).

Mixed assemblages are defined in Lukeneder (2004a,b) as

comprising allochthonous elements (e.g. gastropods, sponges,
corals)  transported  from  the  shallower  shelf  to  upper  slope
and autochthonous benthic and parautochthonous pelagic el-
ements  (e.g.  ceratitid  ammonoid  Kasimlarceltites)  from
deeper  marine  environments.  See  also  the  discussions  by
Olóriz  &  Villaseñor  (2010)  on  post-mortem  and  possible
biostratinomic mixing of ecologically age-separated popula-
tions. Assuming an analogous situation for the A ag˘

yaylabel

sections as for the Karap

nar sections, this would mean that

the  presence  of  undamaged  macroconchs  accompanied  by
intact  microconchs  within  the  same  bed  points  to  a  similar
derivation  of  the  ammonoids  from  the  same  source,  in  this
case  the  pelagic  fauna  of  the  water  column  above,  without
assuming any depth. This points to a huge problem in the un-
derstanding of ammonoids life and habitats. The question, if
ammonoids  in  general,  and  Kasimlarceltites  or  Sirenites
from Turkey in detail, were planktonic drifters, nektic swim-
mers  or  nektobenthic  swimmers  is  beyond  the  field  of  the
study  here.  Nonetheless,  the  answer  to  this  question  would
make a serious difference to the study of biostratinomic fea-
tures  in  chambered  cephalopods,  both  fossil  (e.g.  am-
monoids, nautiloids) and recent (e.g. Nautilus, Spirula).

The post mortem history, hence the subsequently compli-

cated  drifting  mechanisms  (i.e.  waterlogging  and  sinking
versus  surfacing  and  floating;  see  Seilacher  1968;  Olivero
2007)  in  ammonoid  shells  depends  mostly  on  the  water
depth  and  hydrostatic  pressure  (Maeda  &  Seilacher  1996)
when the animal dies. This interpretation is contrasted to the
occurrence  of  some  specimens  showing  different  sediment-
infillings of the body chambers as compared to the surround-
ing  and  embedding  finer  limestone.  Reasons  for  these
differences in sediment infilling reflect the difference in sed-
iment and shell transportation history. Specimens filled with
coarser material (i.e. shallower water relicts) are redeposited
from shallower areas. Shells are preserved without fragmen-
tation (i.e. body chamber present), mostly with almost simi-
lar  alignment  and  rare  infilling  of  phragmocones  by
sediment  (see  Olivero  2007).  Observed  geopetal  structures,
aligned in almost identical directions, within the accumulated
ammonoids give evidence for a fast burial history of almost
the  entire  contingent  of  ammonoids.  Ammonoid  specimens
shown  by  Olivero  (2007)  from  the  Santonian—Lower  Cam-
panian  mass  flow  deposits  of  Antarctica  also  exhibit  pre-
served siphuncle tubes preventing the infill of phragmocones
with sediment.

The cephalopods of the A ag˘

yaylabel and Karap

nar sec-

tions  thus  constitute  a  mixed  autochthonous/parautochtho-
nous/allochthonous  (Martin  1999)  fauna.  This  effect  is
enhanced by the fact that gravity currents, submarine mass-
flows (mud supported and water saturated) may already con-
tain  a  mixed  shelf  and  slope  assemblage  by  picking  up
bioclasts  from  different  bathymetric  zones  along  their  way
(Einsele & Seilacher 1991). The Carnian mixed assemblages
comprise  a  considerable  amount  of  bivalves,  gastropods,
sponges  and  corals  from  shallower  environments  from  a

nearby  located  platform  or  upper  ramp  (Lukeneder  et  al.
2012; see Eberli 1991). The term ‘mixed’ assemblage is used
in the sense of Kidwell & Bosence (1991). The latter authors
described  a  mixed  assemblage  as  the  addition  of  shells  of
one assemblage to the members of another assemblage. For
classification  and  reviews  on  taphonomic  processes  of  ma-
rine  shelly  faunas  see  also  Norris  (1986),  Kidwell  et  al.
(1986),  Brett  &  Seilacher  (1991),  Kidwell  (1991a,b),  Kid-
well & Bosence (1991), and Speyer & Brett (1991).

Ammonoid shell alignment

The ammonoid shell orientation (i.e. rose diagrams of

shell axes, body chamber orientation; Figs. 9, 10, 11) within
the  accumulation  layers  was  measured  to  gain  information
on  the  prevailing  conditions  and  mechanisms  (e.g.  bottom
water currents, debris flows, turbidites, storms etc.; Goldring
1991)  at  the  time  of  deposition  (Early  Carnian,  Austrotra-
chyceras austriacum Zone).

A more detailed picture of the three-dimensional alignment

of specimens from AS I—block 1 (i.e. 150

×45×140 mm

block, 70 slices with 2 mm distance) is presented in Lukene-
der et al. (2014). Lukeneder et al. (2014) show that the inter-
nal,  dominant  orientation  of  specimens  in  fossil  mass
occurrences can be exploited as a useful source of informa-
tion  about  the  flow  type  and  direction  determining  the  pre-
cise conditions for their transportation and accumulation. A
series  of  studies,  using  different  kind  of  fossils,  especially
those with elongated shape (e.g. elongated gastropods), deal
with  their  orientation  and  the  subsequent  reconstruction  of
the  depositional  conditions  (e.g.  paleocurrents,  transport
mechanisms).  However,  disk-shaped  fossils  like  planispiral
cephalopods or gastropods were used, up to now, with cau-
tion  for  interpreting  paleocurrents.  Moreover,  most  studies
just deal with the topmost surface of such mass occurrences,
due to its easier accessibility. Within Lukeneder et al. (2014)
the exact spatial shell orientation was determined for a sam-
ple of 675 ammonoids, and the statistical orientation analy-
sed with a NW/SE-orientation. The study of Lukeneder et al.
(2014)  from  the  A ag˘

yaylabel mass occurrence combines

classical  orientation  analysis  with  modern  3D-visualization
techniques,  and  establishes  a  novel  spatial  orientation  anal-
ysing method, which can be adapted to any kind of abundant
identifiable object. Such a spatial alignment with imbrication
in  a  gravity  flow  was  detected  by  Hladil  et  al.  (1996)  for
Lower Devonian tentaculite shells from the Czech Republic.
There,  the  analysed  fossils  show  a  similar  oblique  orienta-
tion  (i.e.  upward  and  downward)  due  to  gravity  transport
(Hladil  et  al.  1996),  caused  by  a  rapid  consolidation  of  the
host sediment. It should be noted that the natural, post mortem
orientation angle of a particular ammonoid shell (i.e. Kasim-
larceltites) after sinking onto the sea floor, is almost horizon-
tal  with  approximately  2—3°  (Fig. 8),  depending  on  the
ontogenetic stage. This phenomenon is caused by the maximal
whorl breath at the body chamber near the aperture. The hori-
zontal shell orientation dominates over oblique alignment in
layers formed during undisturbed backround sedimentation.

The Triassic assemblages differ significantly from a Mid-

dle Devonian example of nautiloid mass occurrences ( = con-

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centrates in Soja et al. 1996), which was interpreted as being
a combination of mass spawning and mass mortality events
(biological  process;  see  Lukeneder,  in  print)  with  storm-re-
lated accumulations (physical process). In contrast to the Tri-
assic  accumulation  layers  from  Turkey,  the  densely  packed
Devonian cephalopod mass occurrence (i.e. nautiloid grain-
stone) shows no orientation, alignment or imbrication of the
shells (Soja et al. 1996). Goniatites beds from Upper Devo-
nian of Poland (Niechwedowicz & Trammer 2007) were de-
scribed  as  post  mortem  shell  accumulation,  not  transported
very  far,  deposited  within  the  habitat  realm  and  interpreted
as the result of condensation, which has often been noted for
cephalopod limestones. Niechwedowicz & Trammer (2007)
interpreted the shell alignment due to wave or current trans-
port in shallower environments.

At least some of the abundant ammonoid specimens seem

to have been redeposited from shallower shelf regions into a
slope  environment.  Similar  occurrences  were  reported  from
low  density  gravity  flows  (e.g.  slump  deposits  and  turbid-
ites) from Lower Triassic redeposited ammonoid accumula-
tions  of  East  Russia  (Maeda  &  Shigeta  2009).  The  authors
note  sporadically  intercalated  ammonoid  beds  (with  nearly
horizontal  alignment  of  ammonoid  shells)  within  otherwise
‘normal’ mudstone sedimentation. Maeda & Shigeta (2009)
identified  an  allochthonous  source  (i.e.  primary  shelf  edge
biotope)  for  ammonoids  deposited  and  transported  in  such
fossiliferous,  turbidite  layers.  Taxonomical  related  findings
of celtitids were presented by Manfrin et al. (2005) from ad-
jacent basinal series, surrounding the Middle Triassic Later-
mar  Platform  (N Italy).  The  faunas  from  these  ammonoid
layers were interpreted by Manfrin et al. (2005) as storm de-
posits,  due  to  the  mixture  of  pelagic  and  platform  derived
fossils  and  the  partly  perpendicular  ( = non-equilibrium  in
Manfrin et al. 2005) alignment of ammonoids within the co-
quinas.  It  is  evident  that  such  layers  consist  of  transported,
mixed faunal elements (i.e. pelagic ammonoids and benthic
gastropods – Manfrin et al. 2005), deposited by distinct and
short  events.  This  is  almost  identical  to  the  assemblages
found  at  A ag˘

yaylabel, but with the remarkable difference

of chaotic shell-alignment, enhanced shell-fracturing and
variation of sedimentological features.

Mass flow transport is evident from the orientation of

shells within the layers (Middleton & Hampton 1973, 1976;
Brown  &  Loucks  1993;  Potter  et  al.  2005;  Nichols  2009),
not only at the top of beds as is the case when water current
causes  mass  occurrence  or  turbidite  accumulation.  A  some-
how  mixed  or  transitional  mechanism  of  debris  flow  (i.e.
laminar flow) and turbidity current (i.e. turbulent flow) dep-
osition is assumed for floatstones of AS Ia and AS II. A low
density,  less  concentrated  and  water  saturated  debris  flow
( = liquefied debris flow) with transitional features into a tur-
bidite  transport  (pers.  comm.  Michael  Wagreich  2013;  see
Lowe  1982;  Einsele  1991;  Hladil  et  al.  1996;  Mulder  &
Alexander 2001; Olivero 2007) is highly presumable. A hor-
izontal  or  at  least  low-angle  deposition  of  imbricated  shells
is  caused  by  low-density  transports  (Hladil  et  al.  1996).  A
positive  enhancement  of  an  existing  inclination  can  be
forced by slipping or sliding processes of the not consolidated
sediment layers, down the slope or ramp.

The mostly erosive base of the accumulation layers, fur-

thermore undulated with fragmented ammonoid shells, re-
flects sudden and punctual reworking phases at the base of
the sediment flow to some extent (see Sepkoski et al. 1991).

The majority of the accumulation layers (AS I – beds 4, 6;

most  beds  at  KA I,  KA II)  were  formed  by  water  saturated
debris flow deposition which caused the alignment of the bio-
logical particles (i.e. ammonoid shells) within the sediment.
A  contrasting  exception  can  be  observed  in  section  AS IV
(e.g. bed 8; Figs. 10, 11, Tables 1, 2, 3) and KA II within the
blocks 1 and 4. In blocks 1 and 2 (corresponding to the same
layer)  in  AS IV  and  KA II  (e.g.  blocks 1,  2)  a  clear  short-
term  turbidite  transport  took  place.  Ammonoid  shells  seem
to  float  on  the  peloidal-silt  level,  most  probably  due  to  hy-
drodynamic  sorting.  Hence,  low  density,  empty  ammonoid
shells  (1.1—1.2 g/cm

– Maeda 1999; Maeda & Shigeta

2009), subsequently filled by calcite, remain suspended in
the water column during or shortly after the transport event.

All of the distinct ammonoid event layers represent only

episodic  (i.e.  formed  during  days  to  weeks),  punctual  thin
horizons (see Brandner et al. 2012) which interrupt the ‘nor-
mal’  background  sedimentation  phase,  consisting  of  lime-
stone  beds  with  only  rare  occurrences  of  ammonoids  (e.g.
Kasimlarceltites,  Klipsteinia,  Sirenites).  Brett  &  Baird
(1986)  distinguished  two  kind  of  deposition,  characterized
by  the  mode  of  sedimentation  rate  with  the  long-lasting
background  deposition  (1—10 cm/10

yr) and the punctual

episodic sedimentation (1—50 cm/10

—2

yr).

The almost horizontal alignment of the shells from the am-

monoid-rich  horizons  reflects  the  normal  long-term  sedi-
mentation  and  no-to-weak  transport  of  shells  before
embedding. The observed angle is similar to angles detected
when  single  specimens  are  lain  on  a  horizontal  ground
(Fig. 8). Contrastingly, the increased angle in the polyspecific
KA II – block 4 with 164°/344° (frontal data) resp. 15°/195°
(orthogonal  data)  is  interpreted  as  caused  by  a  more  turbu-
lent transport and the mixture with specimens of the bigger
Sirenites (strong ribs and spines), on which Kasimlarceltites
(almost  smooth)  specimens  often  ‘lean’.  The  increased  ob-
liqueness in axes is also caused by the increased number of
steeper orientations in body chambers with a mean direction
of 111° in KA II – block 4 (Fig. 10).

The increased angle seems to reflect bioturbation and sec-

ondary  dislocation  of  the  small  objects  (i.e.  0.5—8.0 mm).
Such  small  sized  skeletal  objects  are  easily  orientated  by
common  depositional  processes  (e.g.  bottom  currents,  tur-
bidites, storms, mass flows) but not obliged to be aligned re-
lated  to  their  shell  axes,  owing  to  the  almost  globular
morphology of small objects.

A more indistinct and imprecise picture is seen when inter-

preting the body chamber orientation in both, frontal and or-
thogonal  two-dimensional  slices  (Fig. 10,  Tables 2,  3).  As
the  body  chamber  length  in  Kasimlarceltites  varies  from
three-fourths  to  an  entire  whorl  (i.e.  mesodome  to  longi-
dome;  see  Westermann  1996),  measurements  and  data  can
only  show  the  direction  of  the  biggest  part  of  the  outer
whorl, not assuming to show the definite, final apertural di-
rection.  This  problem  should  be  solved  with  3D  sectioning
and  reconstruction  (Lukeneder  et  al.  2014).  Hence  the  data

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show unimodal (only one direction of body chambers) to bi-
modal (two opposite directions) distribution roses (Fig. 10).
As shown above, the data on axes orientations, hence the
transport induced directions and alignments are more reli-
able within our two-dimensional based work.

Size groups in Kasimlarceltites

The intraspecific variation of size classes and the differ-

ence  within  layers  and  localities  is  the  expression  of  either
different  sources  (i.e.  environments,  depth  range)  of  the
transported  sediment  (i.e.  including  ammonoids  shells),  or
the  result  of  sexual  (i.e.  females = macroconchs  –  M,
males = microconchs  –  m)  or  ontogenetic  (i.e.  ammonitel-
lae,  juveniles  and  adults)  separation  of  ammonoids  habitats
during life.

The difference in abundance and resulting distribution

curves  from  frontal  and  orthogonal  size  classes  in  several
blocks of different localities, once again strengthens the con-
cept  of  an  alignment  of  ammonoid  shells  after  intense  and
detectable  transport.  This  important  issue  can  be  noticed  in
KA II – block 2 with frontal values of 8.4—10.4 mm versus
smaller  orthogonal  values  between  4.2—6.2 mm,  KA I  –
block 1 smaller frontal values (4.2—6.2 mm) versus orthogo-
nal values (8.4—10.4 mm), and AS IV – block 3 with a fron-
tal  wide  range  (6.3—18.8 mm)  versus  a  narrow  orthogonal
range  of  6.3—8.3 mm  and  a  second  peak  at  25.2—27.2 mm.
Frontal and orthogonal size class distribution is almost iden-
tical in AS Ia – blocks 1, 2, 3 (Fig. 10).

Conclusions

The Upper Triassic macrofauna of A ag˘

yaylabel and

Karap

nar (Taurus Mountains, Turkey) is represented espe-

cially by ammonoids (i.e. ceratitids) and bivalves (i.e. halo-
biids).  The  whole  section  yielded  over  2300  ammonoids,
extracted  and  prepared  as  well  as  embedded  in  blocks  and
sections.  The  fauna  can  be  assigned  to  the  Lower  Carnian
Austrotrachyceras austriacum Zone (Lukeneder & Lukeneder
2014)  and  contains  ammonoids  of  all  ontogenetic  stages,
from ammonitallae to adults (for more detail see Lukeneder
& Lukeneder 2014).

The invertebrate fauna (e.g. ammonoids, bivalves, gastro-

pods, sponges, corals) is accumulated in isolated and distinct
single-event layers. The cephalopod shells are aligned and
concentrated in particular levels and some show, to some ex-
tent, current-induced orientation. Alignment of shells into
diverse orientations suggests mass flow currents or other tur-
bulent bottom-water currents.

The ammonoid fauna at A ag˘

yaylabel and Karap

nar were

deposited  together  with  sediments  formed  by  gravity  in-
duced flows and turbidites under the influence of winnowing
and  bottom  currents.  The  sediments  were  partly  reworked
and transported in suspension for some distance, from shal-
lower areas at the platform edge to the upper slope onto the
deeper parts of the slope and basin. The sediments were ini-
tially deposited on the platform shelf close to the slope edge
(shelf  break  area),  which  is  also  near  the  final  embedding

place of the ammonoid remains. In this area, unstable marine
sediment  accumulations  create  the  prerequisite  conditions
for remobilization by gravity flows and/or turbidity currents
(Einsele 1991; Potter et al. 2005). Those flows and currents
then  built  up  the  floatstones  and  packstones  comprising  the
ammonoid  mass  occurrences.  The  final  deposition  of  the
floatstones to packstones from A ag˘

yaylabel and Karap

nar

took place on tectonically unstable slope areas during condi-
tions of relatively high sedimentation rates. Successions with
abundant or accumulated ammonoid-layers are widespread
over a 15 km

area.

Debris flows and turbidity current, or a combination of

both, were triggered either by storm wave activity or by forc-
ing other physical events such as earthquakes, tsunamis and
less probably sediment overloading, which led to the forma-
tion of event beds and ammonoid accumulation layers on the
upper slope to basin of the Carnian (Austrotrachyceras austria-
cum Zone) from the Taurus Mountains. The ammonoid accu-
mulation layers (and therefore the lower part of the Kas

mlar

Formation) are almost monospecific, dominated by the cer-
atitid genus Kasimlarceltites with up to 99.9 %. The thickness
of the Kasimlarceltites acme zone ranges from 1.8 m at
A ag˘

yaylabel (section AS I) and 16.5 m at Karap

nar II (sec-

tion KA II). The position, and hence the exact geographical
cardinal direction of the source area, is unknown.

The orientation measurements (e.g. angles of axes and

body chambers) of the ammonoids also point to origination
by  water-saturated,  liquefied  debris  flows,  resulting  in  bio-
genic  floatstones  or  packstones  (i.e.  matrix  supported  and
ammonoid shell supported; see Flügel 1978, 2004). The two-
fold picture clearly points to various transport mechanisms,
hence  a  change  of  source  areas  or  transport  history  during
the Julian—Tuvalian in Carnian times (Late Triassic). An in-
creasing water depth, either due to a sea level rise or a tec-
tonic drop of the carbonate platform is also evident for that
Anatolian  area,  clearly  detectable  in  the  sedimentological
and  paleontological  record.  Subsequently,  current  systems
changed  during  that  time  of  paleo-oceanographic  modifica-
tion  and  restructuring,  resulting  in  unstable  conditions  and
thus redeposited accumulation layers.

The Kasimlarceltites event layers are intercalated with

‘normal’  sedimentation  beds,  which  are  represented  by
wackestones with only rare, floating Kasimlarceltites speci-
mens. Small ammonoids (i.e. juvenile Kasimlarceltites) and
shell  fragments  in  the  body  chambers  of  somewhat  larger
ammonoids  also  support  the  assumed  effect  of  agglomera-
tion and comminution by dense sediment flows with a lami-
nar  internal  flow.  The  accumulation  of  ammonoid  layers
indicates either on-site deposition at short, favourable ‘time-
intervals’,  or  reworked  accumulation-layers  after  gravity
flow  transport  (slow  debris  flow).  Most  ammonoid  speci-
mens  are  not  fragmented  and  do  not  show  bioerosion  (i.e.
boring, encrustation). This suggests a short transport history
of  the  sediment  masses  and  rapid  incorporated  shells  (e.g.
ammonoids,  bivalves,  gastropods)  as  well  as  relatively  fast
burial.  Moreover,  because  most  body  chambers  of  the  am-
monoid  specimens  are  filled  with  debris,  they  were  already
dead at the time when they were transported. These are thus
true,  redeposited  accumulations,  and  their  initial  accumula-

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Fig. 11. Calculation scheme for true dip direction in ammonoid
shells from A ag˘

yaylabel (AS I, AS I – blocks 1, 2, 3, 4). True dip

direction of the ammonoid shell ‘plane’ needs two different (frontal
and orthogonal) apparent dip measurements of ammonoid shell
axes. For more details on this method and three dimensional calcu-
lations see Lukeneder et al. (2014).

tion  in  shallow  water  must  have  also  been  characterized  by
fairly  rapid  burial,  to  avoid  postmortem  bioerosion.  The
presence  of  ammonoids  bearing  intraclasts  (i.e.  different
colour  due  to  different  source  areas),  of  sponges  associated
with  corals  and  gastropods,  of  slightly  dislocated  shell  an-
gles in ammonoids, of dislocated geopetal structures (i.e. in
ammonoid  shells)  and  in  cases  of  a  cover  of  broken  am-
monoid shells and bivalve coquinas (Kidwell 1991a,b) sig-
nify a turbulent transport and redeposition of sediments and
fossils.

The assemblages within the described accumulation layers

depict an indefinite mixture of autochthonous ( = indigenous
in  Kidwell  1991b),  parauthochthonous,  and  allochthonous
( = exotic  in  Kidwell  1991b)  sedimentological  and  biogenic
elements.  Concentrations  of  ammonoid  shells  are  mainly
based  on  primary  biogenic  (sensu  Kidwell  et  al.  1986)  and
sedimentological concentration mechanisms (see Kidwell et
al. 1986; Meldahl 1993). The combination of both (biogenic
and  sedimentological  skeletal  concentrations)  suggest  inner
shelf and inner ramp environments (Kidwell et al. 1986).

The enormous quantity of ammonoid shells in these thin

beds also suggests a possible gregarious life style of the cer-
atitid  Kasimlarceltites,  at  least  during  times  of  mating  and
spawning  (see  also  Soja  et  al.  1996;  Lukeneder,  in  print).
Buccal  masses  with  preserved  beak  apparatus  (e.g.  aptychi
like  jaws)  are  completely  missing  in  the  Turkish  material.
Isolation  took  place,  either  through  transport,  due  to  differ-
ent  behaviour  within  the  water  column,  or  through  current-
induced grain differentiation during accumulation. The latter
scenario  leads  to  different,  and  unknown,  places  of  deposi-
tion for these two cephalopod elements of the same animal.

A highly variable sea floor morphology is induced by the

lithological and sedimentological deviations within the adja-
cent sections. Bottom physiography produced different accu-
mulation models and ammonoid shell bed types.

This leads consequently to the question of the time during

which Kasimlarceltites dominated the fauna in this Late Tri-
assic (i.e. Carnian – Julian 2) area. A stratigraphical range
(Gradstein  et  al.  2012)  from  one  third  to  one  half  of  the
Austrotrachyceras  austriacum  Zone  (approximately  200 ky
at AS I and AS IV – 500 ky at KA II) is calculated for the
Kasimlarceltites  acme  zone,  not  considering  any  hiati  or
time averaging, which might have occurred. A possible geo-
graphical differentiation into habitats from sexual dimorphic
pairs (i.e. females, males and juveniles), and hence the origi-
nal  water  depths  inhabited  by  different  size  and  morpho-
groups,  bears  the  potential  to  change  the  picture  of  the
formation  mechanisms  and  habitats  in  ceratitid  ammonoids
and to produce such mass occurrences or event beds in par-
ticular. The obtained information encourages future research
focused  on  the  preservation  history  and  processes  causing
ammonoid accumulations.

Acknowledgments: This study benefited from Grants from the
Austrian Science Fund (FWF) within the Project P 22109-B17.
The authors highly appreciate the help and support from the
General  Directorate  of  Mineral  Research  and  Exploration
(MTA, Turkey) and are thankful for the digging permission
within  the  investigated  area.  Special  thanks  go  to  Ye im
Islamog˘lu  (MTA,  Ankara)  for  organizing  and  guiding  two
field trips. The authors thank Leopold Krystyn and Andreas
Gindl  (both  University  of  Vienna),  Mathias  Harzhauser  and
Franz  Topka  (both  Natural  History  Museum  Vienna)  and
Philipp  Strauss  (Austrian  Oil  Exploration  Company  OMV,
Vienna),  who  provided  material  from  previous  field  trips.
Simon  Schneider  (CASP,  Cambridge,  United  Kingdom)  and
Thomas Hofmann (Geological Survey of Austria, Vienna) are
acknowledged  for  their  support  in  collecting  literature.  We
thank  Simon  Schneider  (CASP,  Cambridge,  United  King-
dom)  for  explanations  of  some  details  concerning  the  new
established  method.  Photographs  of  ammonoid  specimens
were  taken  by  Alice  Schumacher  (Natural  History  Museum
Vienna). We kindly acknowledge Susan Kidwell (University
of  Chicago),  Federico  Olóriz  (Universidad  de  Granada)  as
well as Jozef Michalík (Slovak Academy of Science) for their
comments, which greatly improved the quality of this manu-
script. We are grateful to Michael Wagreich (University Vien-
na,  Vienna)  for  discussions  and  helpful  comments.  We,  the
authors,  dedicate  the  paper  to  Eva  Chorvátová  (Bratislava)
who passed away much too early in 2014. She supervised the
first steps of our submission to Geologica Carpathica.

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