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Introduction
In 1968 ELLIOTT described a new dasycladalean ge-
nus and species as Harlanjohnsonella annulata from
transgessive, basal Upper Cretaceous, possibly Ceno-
manian of “Tetrebovo, Dlaglica SE of Zlatibor massif,
SW Serbia” with the following diagnosis of the genus
“Weakly calcified thin-walled tubular and annular
dasyclad, with successive verticils showing numerous
swollen primaries, the presumed secondaries not be-
ing calcified”. The species is typified 1993 in
GRANIER & DELOFFRE Discussing the relationships,
ELLIOTT concluded “that the form now described was,
in fact, a dasyclad of very similar plant morphology to
D. (Dissocladella) savitriae, but more weakly calci-
fied” and because “there is no direct fossil evidence of
secondary branch-structure the species cannot cor-
rectly be referred to Dissocladella.”
Based on material sampled at the type locality (RA-
DOIČIĆ 1995), from which new thin sections have been
prepared, it is documented that the genus Harlan-
johnsonella ELLIOTT represents a younger synonym of
Dissocladella PIA confirming ELLIOTT’s presumption.
The scope of the present paper is a taxonomic revision
and detailed illustratation of Dissocladella annulata
(ELLIOTT) nov. comb.
Geological setting
The Cretaceous deposits of the southern (Tetrebo-
vo, Vis, Fig. 1) and of the central Zlatibor Mountains
are rare remnants of a Cretaceous (Albian–?Lower
Senonian) cover resting upon ultramaphic rocks. First
data on the Cretaceous strata of southern Zlatibor
were given by ELLIOTT (1968, “possibly Cenoma-
Observations on Dissocladella annulata (ELLIOTT, 1993) nov. comb.
(Calcareous algae, Dasycladales) from the Cenomanian of west Serbia
RAJKA RADOIČIĆ1& FELIX SCHLAGINTWEIT2
Abstract. Based on material from the type area at Tetrebovo in the Zlatibor massif of W Serbia, the Ceno-
manian dasycladalean alga originally described as Harlanjohnsonella annulata by ELLIOTT (1968, typified
1993 in: GRANIER & DELOFFRE), is emended and revisited as Dissocladella annulata (ELLIOTT) nov. comb.
The evidence of tufts of short secondaries arising at the top of the drop-like primaries allows its transfer to the
genus Dissocladella PIA, 1936. This species displays a different degree of skeleton calcification which is de-
scribed in detail. The monospecific genus Harlanjohnsonella ELLIOTT becomes invalid, as being a junior syno-
nym of Dissocladella.
Key words. Dasycladales, Dissocladella annulata (ELLIOTT) nov. comb., systematic taxonomy, Cenoma-
nian, Serbia.
Апстракт. Дазикладацејску врсту из ценоманских слојева Тетребова (јужни Златибор) описао је EL-
LIOTT (1968, типификована 1993 у: GRANIER & DELOFFRE)каоHarlanjohnsonella annulata nov. gen., nov. spec.
Преиспитивањем узорака из типског локалитета добијени су детаљнији подаци о грађи ове врсте – до-
кументовано је постојање секундарних огранака. Овај податак указао је на њену припадност роду Dis-
socladella PIA, а на основу тога моноспецифични род Harlanjohnsonella је инвалидизиран. Дијагноза врсте
Dissocladella annulata (ELIOTT, 1968) nov. comb. допуњена је уз детаљне илустрације о структури скелета.
Кључне речи: Dasycladales, Dissocladella annulata (ELLIOTT) nov. comb., систематска таксономија,
ценоман, Србија.
GEOLO[KI ANALI BALKANSKOGA POLUOSTRVA
ANNALES GÉOLOGIQUES DE LA PÉNINSULE BALKANIQUE 71 53–71 BEOGRAD, decembar 2010
BELGRADE, December 2010
1Kralja Petra 38, 11000 Beograd, Serbia. E-mail rradoicic@sezampro.rs
2Lerchenauerstr. 167, 80935 München, Germany. E-mail ef.schlagintweit@t-online.de
DOI: 10.2298/GABP1071053R
nian”, based on RAMPNOUX’s data and fossil material)
and RAMPNOUX (1974). The presence of Turonian
strata was documented by RADOIČIĆ (1995) at Vis
(Fig. 1), in the central Zlatibor and Ravni area in the
eastern part of Zlatibor. Regrettably, these data was
ignored in the context of geological mapping for the
Prijepolje sheet (ĆIRIĆ et al. 1977), and by subsequent
researchers (e.g., DIMITRIJEVIĆ et al. 2002: Geology of
Zlatibor).
The Tetrebovo succession (Figs. 1, 2), overlies pe-
ridotites. According to RAMPNOUX (1970, p. 275, fig.
145; 1974, p. 63), it starts with a Lower Cretaceous
weathering crust. In detail the succession consists of:
– basal conglomerates,
– sandy limestones with gastropods, at places lime-
stones with “Harlanjohnsonella” annulata ELLIOTT,
and,
– limestones with Chondrodonta joannae CHOFAT,
Radiolites lusitanicus PARONA and R. peroni DOU-
VILLE.
This succession was ascribed to the Turonian by
RAMPNOUX in 1970, afterwards in 1974 to the Ce-
nomanian–Turonian as being equal to that one at Rav-
ni (East Zlatibor). In the meantime, the rudist lime-
stones in the Ravni area, dated as Turonian by PEJOVIĆ
& PASIĆ (1958) were revised as being Cenomanian in
age (RADOIČIĆ 1995). It should be mentioned, that this
does not correspond to the same Ravni succession
dated as Turonian by RADOIČIĆ. Afterwards, these
sparsely outcropping Tetrebovo Cretaceous deposits
were sampled only at places (RADOIČIĆ 1995). One of
the oldest observed beds is represented by a marly
limestone containing gastropods and large specimens
of Dissocladella annulata (EL-
LIOTT) nov. comb., which are
well visible without lens. Up-
ward in the section, the lime-
stone contains rare benthic
foraminifera, the dasycladale-
an algae Heteroporella lepina
PRATURLON, Terquemella sp.
and a few fragments of Disso-
cladella annulata. They are
followed by limestones with
frequent foraminifera, respecti-
vely Marssonella turris (D’OR-
BIGNY), Rotalia mesogeensis
TRONCHETTI, Pseudorhipidio-
nia casertana (DECASTRO),
Pseudocyclammina rugosa (D’
ORBIGNY) and Praealveolina
cf. iberica REICHEL. Between
these beds and youngest ob-
serveds skeletal calcarenites
(middle-upper Cenomanian),
the limestone with relatively
frequent Pseudorhapydionina
dubia DECASTRO is sampled.
The limestone with Dissocladella annulata con-
tains numerous large skeleton fragments, small and
minute debris, different gastropods, molluscan shells
RAJKA RADOIČIĆ & FELIX SCHLAGINTWEIT
54
Fig. 1. Geological map of the Tetrebovo area, according to
RAMPNOUX (1970, fig. 145), and Geological map sheet
Prijepolje 1:100000, ĆIRIĆ et al. (1977). Legend: 1, Upper
Triassic limestones; 2, Serpentinites; 3, Habzburgites; 4,
Weathering crust, Lower Cretaceous; 5, Upper Cretaceous
limestones (?Albian–Cenomanian).
Fig. 2. Tetrebovo hill, northern view, Cretaceous deposit are covered by forest.
and rare crustacean fragments, also mentioned by
ELLIOTT (1968). In the 25 thin sections studied, only
one specimen of Pseudorhipidionina casertana and a
few small foraminifera were observed. Clearly, Disso-
cladella annulata obviously populated shallow-water
environments, probably of low salinity. At the type
locality, the limestone with Dissocladella annulata
can be ascribed to the lowermost Cenomanian.
Systematic taxonomy
Division Chlorophyta
Order Dasycladales PASCHER
Family Triploporellaceae (PIA, 1920)
Tribus Dissocladelleae ELLIOTT, 1977
Genus Dissocladella PIA, 1936 in: RAMA RAO and
PIA, 1936
(Synonym Harlanjohnsonella ELLIOTT, 1968)
Dissocladella annulata (ELLIOTT, 1968),
nov. comb., revisited
Pls. 1–5, Pl. 6, Figs. 1–16
1968 Harlanjohnsonella annulata nov. gen., nov.
sp. – ELLIOTT, p. 494, pl. 93, figs. 1-2, pl. 94,
figs. 1-2.
1978 Harlanjohnsonella annulata ELLIOTT – BASSO-
ULLET et al., p. 120, pl. 12, figs. 8-9.
non 1978 Harlanjohnsonella annulata ELLIOTT – LAU-
VERJAT & POIGNANT, p. 123, pl. 2, figs. 1, 5-6.
1995 Harlanjohnsonella annulata ELLIOTT – RADOI-
ČIĆ, pl. 1, fig. 1.
Material. Twenty-five thin sections from the sam-
ple 022070, R. RADOIČIĆ collection RR4579 – 4584/9
deposited at the Geological Institute, Beograd.
Diagnosis. Elongated cylindrical thallus exhibiting
a large central stem with moderately spaced horizon-
tal whorls. The whorls consist of numerous laterals:
drop-like primaries which, at the top, bear tufts of 5–6
thin phloiophorous secondaries. Primary calcification
generally weak, stronger or only somewhat thicker
around the proximal area of the primaries, becoming
thinner outwards, especially at tip and around the
secondaries. Possible presence of fertile and sterile
individuals.
Description. Being rather variable in size, the skele-
ton of this species is rather thin with smooth inner sur-
face. The primary calcification is diagenetically over-
grown in variable degrees. Different degrees of recrys-
tallization can be observed even within the same whorl.
The weakly calcified distal part of the whorls, if not
early diagenetically recrystallized, is more or less dis-
solved or abraded. Therefore, secondary laterals are
preserved only in very rare cases, while poorly pre-
served secondaries are discernable as pores or open
pores on the surface of many recrystallized skeletons
(Pl. 1, Figs. 2, 4; Pl. 2, Figs. 3, 4; Pl. 3, Fig. 7; Pl. 4, Fig.
9; Pl. 5, Figs. 1–6). Only in some specimens, the mem-
brane of the central stem can be recognized as a dark
thin micritic line (Pl. 2, Figs. 1, 2, 5; Pl. 3, Fig. 7). The
thin calcareous encrustation of the membrane is rarely
preserved; it can be recognized only between two pri-
maries of successive whorls, visible in some sections
(Pl. 1, Fig. 4, arrows; Pl. 3, Fig. 4, left; Pl. 6, Figs. 4, 5).
A thin-walled calcareous tube encloses the pores of pri-
maries (Pl. 2. Fig. 1), Pl. 3, Fig. 6; Pl. 4, Fig. 1) or more
frequently, the thin wall on the surface bears open pores
of primaries (Pl. 1, Fig. 1; Pl. 2, Fig. 5; Pl. 3, Figs. 2–5).
In rare specimens, the skeleton is dissolved so that it
consists of a thin calcareous layer with irregular exter-
nal surface, on which some parts of the basal calcifica-
tion of the primaries can be recognized (Pl. 1, Fig. 3; Pl.
2, Figs. 6, 7).
The laterals are arranged in a plane; they are rarely
slightly overlapping as shown in the specimen illus-
trated in Pl. 5, Fig. 8. In successive whorls, laterals do
not alternate regularly, but occasionally alternation
can be observed in a few successive whorls. In the
proximal part the primaries display a regular funnel-
like form; they communicate with the central stem by
means of minute pores. In deep tangential sections,
small basal pores can be seen gradually increasing
from the center to the periphery (Pl. 4, Figs. 1).
Transversal sections of primaries are circular in shape
(Pl. 4, Figs. 1, 2; Pl. 5, Fig. 7, 8). In both, transversal
and longitudinal sections of the skeleton, the pores are
often secondarily enlarged or diminished and/or more
or less deformed.
Besides more or less large specimens, the analyzed
limestones of Tetrebovo contain numerous small and
especially minute fragments of disintegrated skele-
tons. These have particular value for the recognition
of the structure of this species and the processes of the
skeleton alteration. Minute and small fragments as
those shown in Pl. 1, Fig. 4 (arrows), Pl. 3, Fig. 3 (ar-
rows), and Pl. 6, Figs. 1–7 indicate that some skele-
tons are characterized by a somewhat stronger pri-
mary calcification only of the shorter proximal part of
the whorls, along with an especially thin calcification
of the main axis membrane. Therefore, such a kind of
calcification facilitated skeleton disintegration, more
probably early post-mortem and becoming preserved
as small or minute fragments. A further abiogenic
stage of calcification resulted in the overgrowth of the
primary calcification leading to irregular thallus coat-
ings to variable degrees that is completely or partly
coverage even within the same whorl.
Calcification and mode of preservation (Figs. 3
and 4) Dissocladella annulata is characterized by two
types of primary calcification, shown in the drawings
on Fig. 3/1 and 3/3; Figure 3/4 illustrates the relation-
ship between these two types, on Fig. 4. are given dif-
ferent calcification types of the laterals and of the
preservation of skeleton.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 55
First type, skeletons of group A (A skeleton): rela-
tively strong proximal calcification usually covering
1/3 of their length, and then distally gradually becom-
ing thinner. Distally, delicate parts of the skeleton,
including the secondaries (Fig. 4/1A), were not pre-
served in the studied material. The characteristic fea-
ture of the group A skeleton is that the specimens dis-
play a smaller size of the primary laterals (Fig. 3/1,
Fig. 4/1A-B, and 2A ), resulting in a primarily non-
calcified space between the calcified whorls and an
annulation of the skeleton (referring to the species
name annulata). The best examples are illustrated in
Pl. 4, Figs. 3, 5 also in Pl. 6, Figs. 1–7, 10 and 11, and
also in Fig. 3/1, 4/1A and 2A.
Second type, skeletons of group B (B skeleton):
this type is characterized by a) larger primary laterals
which are, also in both transversal and longitudinal
sections, in slight contact in the largest middle part of
the lateral’s length and b) by the stronger proximal
calcification which, between successive whorls is
compact - cf. collective calcified skeleton sensu DE
CASTRO (1997, “guaine calcificata collettiva”). Se-
condarily altered, this skeleton part is formed by cal-
cite mosaic trimmed by smaller grains. In the studied
material the B skeleton is often preserved as non
annulated relatively thin calcareous tubes (proximal
area) with open pores of primaries (Pl. 3, Figs. 2–5).
It has to be mentioned that in some recrystallized
skeletons including secondaries, the “annulations” are
reflected on the surface only as shallow feeble canals
between the whorls (Pl. 4, Fig. 9; Pl. 5, Fig. 2).
Therefore, the space between them is not calcified and
as a consequence of this, the strong proximal primary
calcification ends in this area (Fig. 5/3; Pl. 4, Figs. 3
left wall, and 4). Hence, this skeleton type is general-
ly preserved as calcareous tubes mainly with open
pores of the primaries at the surface.
Dimensions. The dimensions given by ELLIOTT
(1968) are indicated between brackets.
Longest observed specimen (L): 12 mm
RAJKA RADOIČIĆ & FELIX SCHLAGINTWEIT
56
Fig. 3. Two types of calcification of Dissocladella annula-
ta (longitudinal sections): 1, 2 – Skeleton A: relatively
strong calcified proximal part of primaries which generally
occur in the rock as dispersed elements of disintegrated
skeleton; examples: Pl. 6, Fig. 1; Pl. 1, Fig. 4, arrows; Pl.
3, Fig. 3, arrows. 3– B skeleton; example: Pl. 6, Fig. 9.
4– Relation between A and B skeletons.
Fig. 4. Dissocladella annulata - different calcification
types of the laterals and of the preservations of skeleton
(longitudinal sections): 1A – Skeleton A: primary calcifica-
tion of primaries and secondaries with non-calcified space
between the whorls; 1B – More or less calcified distal part
of primaries with advanced recrystallization in the space
between the whorls; example: Pl. 4, Fig. 1 and 4, left; 1C –
Usual preservation of type B skeleton (collective calcified
skeleton); example: Pl. 3, Figs. 2, 3, Pl. 6, Fig. 9. In the
material studied, these parts of the skeleton are preserved
as a mosaic trimmed by small calcite grains; 1D – Recry-
stallized type B skeleton with only primary pores and with
preserved secondaries, too; example: Pl. 2, Fig. 1 and Pl. 6,
Fig. 13. 2A – Completely recrystallized primaries in the
annulated skeleton with encrusted stem membrane; exam-
ple: Pl. 4, Fig. 3, right arrow. 2 A/B – Different grades of
recrystallized space between whorls, with or without pre-
served pores of primaries, type A skeleton; example: Pl. 4,
Figs. 3, left arrow. 2C – Calcification type of the primaries
corresponding to the collective calcified skeleton sensu DE
CASTRO (1997).
External diameter (excluding small form in Pl. 2,
Fig. 5) (D): 1.18–3.10 mm (up to 2.25 mm)
Central stem diameter (d): 0.940–2.590 mm
d/D: 71% – 89.5% (about 74 %)
Thickness of the calcareous wall (e): 0.098–0.247
mm, maximum up to 0.330 mm (recrystallized –
skeletons into secondaries) (0.26 mm)
Distance between successive whorls (h):
0.198–0.210 mm (0.19–0.25 mm)
Diameter of primary pores (p): 0.098–0.123 mm
(0.13–0.14 mm)
Thickness of central stem membrane 0.015– 0.024 mm
Number of laterals in a whorl (w): 35–70 (48–50)
The distance between the whorls represents a fairly
constant value, while the most variable biometric
parameter is the main stem diameter.
Relationships As a consequence of the species
emendation and new combination, Dissocladella an-
nulata (ELLIOTT) is placed in the Tribus Dissocla-
delleae ELLIOTT, 1977. The genus Harlanjohnsonella
ELLIOTT, 1968 (so far monospecific) becomes invalid
as representing a synonym of Dissocladella PIA, 1936.
Mention should be made, that the possible existence
of secondary laterals was already assumed by ELLIOTT
(1968) and integrated in the genus diagnosis. ELLIOTT
anticipatorily remarked the similarity of “Harlan-
johnsonella” annulata with Dissocladella savitriae
PIA, 1936 (type-species of the genus) from the Ma-
astrichtian–Danian of India showing some similar
dimensional parameters (d, d/D, p) and both dis-
playing typical thallus annulation. Dissocladella an-
nulata (ELLIOTT) may show more variable and relati-
vely larger external diameters and higher number of
laterals per whorl (w about 40 in D. savitriae). Apart
from this, the special type of calcification and diffe-
rent degree of preservation (due not only to diagene-
sis) seems to be a species-specific feature of D. annu-
lata, not reported from D. savitriae with fully calci-
fied ring-like elements enclosing both primaries and
secondaries. Curiously, BASSOULLET et al. (1978, p.
92) mention an internal thallus undulation, though not
mentioned in the original description. In any case, D.
annulata lacks any internal undulation.
Remarks. In the generic discussion, ELLIOTT inclu-
ded the Carboniferous Coelosporella, the Permian
Epimastopora and Pseudoepimastopora. From annu-
lar forms such as the Cretaceous Neomeris cretacea
DELMAS & DELOFFRE non STEINMANN (DELMAS &
DELOFFRE, 1962), Dissocladella annulata “differs in
the apparently simple branch-structure”. Furthermore
ELLIOTT concluded that “a closer comparison can be
made with Dissocladella, especially the Paleocene
Dissocladella savitriae (PIA, 1936).”
BASSOULLET et al. (1978, p. 120) essentialy refer to
affinities with Pseudoepimastopora: “Le genre Pseu-
doepimastopora parait très voisin du genre Harlan-
johnsonella et les differences n´apparaissent pas évi-
dentes” and....... “cette espèce pourrait appartenir au
genre Pseudoepimastora”. BASSOULLET et al. (1978)
furthermore express doubts on the existence of “annuli
or rings” in Harlanjohnsonella (ELLIOTT 1968, p. 494).
Also JAFFREZO et al. (1980) describing Pseudoepima-
stopora pedunculata were discussing affinities/diffe-
rences to the genus Harlanjohnsonella. Pseudoepima-
stpora, however, cannot be considered in the discussion
as it represents a nomen nudum (e.g. GRANIER &
DELOFFRE 1993; GRANIER & GRGASOVIĆ 2000). In the
“New taxonomy of Dasycladale Algae” presented by
DELOFFRE (1988), Harlanjohnsonella ELLIOTT, 1968 is
treated as a synonym of Paraepimastopora ROUX, 1979
although Harlanjohnsonella was established more than
ten years earlier. Paraepimastopora is included by
DELOFFRE (1988) in the Mastoporea PIA with aspondy-
le thalli, whereas in the original description ELLIOTT
placed it in the tribus Thyrsoporelleae. DRAGASTAN
(1975) reported Harlanjohnsonella sp. from the Lower
Cretaceous of Romania, a form later included tentative-
ly in the synonymy of Anisoporella? cretacea (DRAGA-
STAN 1967) by BUCUR (1995). From the Valanginian of
Greece, DRAGASTAN & RICHTER (2003) described
Harlanjohnsonella fuechtbaueri as a new species cha-
racterized by a head-and-peduncle thallus morphology:
the peduncle bearing “only primary vesiculiferous ra-
mification with two shapes: a proximal tubular and the
distal part globulous, like vesicle. The cylindrical pe-
duncle is continued by a ”head” made up of euspondy-
le verticils with vesiculiferous ramification”. As shown
in the present paper, Harlanjohnsonella represents a
junior synonym of Dissocladella; therefore the generic
position of the dasycladalean alga described by DRA-
GASTAN & RICHTER (2003) remains open. The authors
also allege that Dissocladella annulata (ELLIOTT)
should exhibit a head-and-peduncle type thallus, a view
that must be rejected due to the studied abundant mate-
rial. Concerning the section designated as holotype, in
Pl. 4, Fig. 3, it has to be mentioned that the presence of
a “head” is not sure (it may be the section of another
specimen in a densely packed algal limestone).
Acknowledgements
We thank to reviewers FILIPPO BARATTOLO (Napoli) and
MARC A. CONRAD (Genève) for comments and helpfull
sugestions, to MARC A. CONRAD also for improvement
english language.
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Montenegro orientale (Yougoslavie). Mémoires de la
Societé Géologique de France, Nouvelle serie, 52
(1973), 119: 6–99.
Резиме
Опсервације о врсти Dissocladella
annulata (ELLIOTT, 1993) nov. comb.
(Dasycladales) из ценомана западне
Србије
Кредне наслаге јужног (Тетребово, Вис, сл. 1) и
централног Златибора остаци су кредног покрова
(алб–?доњи сенон) на ултрамафитима. Први пода-
так о креди на јужном Златибору дао је ELLIOTT
(1968), а на основу података и материјала RAMP-
NOUX-a. Присуство туронских седимената доку-
ментовано је знатно доцније на узвишењу Вис (сл.
2), на централном,иуобласти Равни на источном
Златибору (RADOIČIĆ, 1995). Подаци о креди на
јужном Златибору игнорисани су од стране тима
који је радио гелошку карту листа Пријепоље
1:100 000 (ĆIRIĆ et al. 1978), као и од потоњих ауто-
ра студије о креди Златибора (DIMITRIJEVIĆ et al.
2002).
Систематика
Ред Dasyladales PACHER
Фамилија Triploporellaceae (PIA, 1920)
Триба Dissocladelleae ELLIOTT, 1977
Род Dissocladella PIA, 1936, in RAMA RAO and PIA,
1936
(Синоним Harlanjohnsonella ELLIOTT, 1968)
Dissocladella annulata (ELLOTT, 1968),
nov. comb., revisited
Табле 1–5; Таб. 6, сл. 1–16.
Дијагноза. Издужен цилиндричан талус са
пространом централном стабљиком која носи
раздвојене хоризонталне пршљенове. Пршљенови
се састоје од бројних субсферичних примарних
огранака који, на врху, носе 5–6 тањих флоио-
форних секундарних огранака. Примарна калци-
фикација је релативно слаба, јача или само нешто
дебља у проксималном дијелу примарних oгра-
нака, отанчавајући дистално, особито у врху и око
секундарних огранака. Могуће је постојање фер-
тилних и стерилних индивидуа.
Врсту Dissocladella annulata карактерише зна-
тна варијабилност димензија, скелет је релативно
танак, глатке унутрашње површине. Примарна
калцификација прекривена секундарно и прекри-
сталисала у различитом степену, често неуједна-
чено у истом пршљену. Секундарни огранци ри-
јетко су очувани. Особеност ове врсте је различита
калцификација – потојање два типа (тип А и Б, сл.
3). Скелет типа А има нешто ситније примарне
огранке, те међу пршљеновима остаје примарно
RAJKA RADOIČIĆ & FELIX SCHLAGINTWEIT
58
некалцифициран простор што је узрок анулације
скелета али и склоности ка десинтеграцији. Ске-
лет типа Б има нешто крупније примарне огранке,
јаче калцифициран проксимални дио, те компакну
калцификацију међу пршљеновима (тип “колектив-
не калцификације скелета” у смислу DECASTRO-а,
1997). Очуваност скелета овог типа је другачија –
то су кречњачке цјевчице са порама примарних
огранака или са отвореним порама уколико зид
није потпуније очуван. У скелету рано прекриста-
лисалих јединки изгубљени су елементи унутарње
грађе, али са отисцима секундарних огранака на
површини скелета.
Dissocladella annulata била је настањена у
плитководном ареалу, највјероватније у условима
смањеног салинитета. Кречњаци са Dissocldella
annulata у типском локалитету приписани су нај-
нижем ценоману.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 59
RAJKA RADOIČIĆ & FELIX SCHLAGINTWEIT
60
PLATE 1
Dissocladella annulata (ELLIOTT, 1993) nov. comb., emended, aspects of different skeleton preservation.
Fig. 1. Relatively well preserved oblique section of type B skeleton with calcification reaching from the main stem
to p.p. distal parts of the primary laterals; note, that in the topmost whorl, the primary laterals are in slight
contact. Thin section RR4584/7.
Fig. 2. Oblique section of type B skeleton altered by endolithic activity; in parts of the wall secondaries are
discernible. Thin section RR4583.
Fig. 3. Oblique section of a dissolved skeleton with only a very thin remnant of the proximal part being preserved.
Note some primary pores in the upper part of the figure. Thin section RR4583/5.
Fig. 4. Slightly oblique transverse section of a completely recrystallized type B skeleton with denticulated outer
(moulds of secondaries) and smooth inner surface. Arrows: minute fragments of primaries (type A skeleton),
in two of which parts of the encrusted stem membrane are preserved. Thin section RR4583/2.
Scale bar for all figures = 0.50 mm
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 61
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PLATE 2
Dissocladella annulata (ELLIOTT, 1993) nov. comb., emended.
Fig. 1. Relatively well preserved, slightly oblique trnsverse section with main stem preserved as a thin micritic line;
arrow: somewhat thicker part of the skeleton wall with a few poorly preserved open pores of secondaries.
Thin section RR4584/8.
Figs. 2, 3. Transverse sections of poorly preserved, recrystallized and more or less dissolved skeletons with, in the
upper part, slightly visible micritic main stem membrane. Thin sections RR4583/1 and 4583/7.
Fig. 3. Slightly oblique transverse section of a recrystallized type B skeleton, partly dissolved, with denticulate outer
and smooth inner surface. Thin section RR4583/7.
Fig. 4. Oblique section of a large fragment, partly recrystallized, altered by endolithic activity and showing some rel-
atively well visible open pores of secondary laterals. Thin section RR4583.
Fig. 5. Slightly oblique transverse section with only the proximal part of the skeleton and the main stem membrane
as a thin micritic line being preserved. Thin section RR4583.
Figs. 6, 7. Transverse and longitudinal section of skeletons in nearly last stadium of dissolution; in both the inner sur-
face is smooth. In Fig. 6, left, note the fragment of longitudinal wall section with three drop-like pores of pri-
mary laterals; Thin sections RR4584/8 and 4583/5.
Scale bar for all figures = 0.50 mm.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 63
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PLATE 3
Dissocladella annulata (ELLIOTT, 1993) nov. comb., emended.
Fig. 1. Longitudinal section of a prevailing recrystallized calcareous tube (9 mm in length) in which some
subsequently recrystallized pores of primaries are discernible. Thin section RR4584.
Figs. 2, 3. Longitudinal-oblique sections of thin type B skeleton’s wall showing open pores of primaries on the surface.
Arrows in Fig. 3: small fragments; note on the left: calcification of group A, versus those of group B skeletons;
detail shown in Fig.4. Thin sections RR4584 /5 and 4584/1.
Fig. 4. Detail of the section in Fig. 3, A versus B type skeletons; note the white lines: the relationship of the distance
between the whorls (c-c): in group A (left) and B (right). Thin section RR4584/1.
Fig. 5. Transverse section. Thin secttion RR4583/4.
Fig. 6. Oblique section of the smallest skeleton observed. Thin section RR4584/2.
Fig. 7. Recrystallized skeleton, note (arrow), two pores of primaries arising from the micrite main stem (micrite line)
(lower arrow). In the upper part a few pores of secondaries are slightly discernible (upper arrow). Thin scetion
RR4583/5.
Scale baar for all figures = 0.50 mm.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 65
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PLATE 4
Dissocladella annulata (ELLIOTT, 1993) nov. comb., emended
Figs. 1, 4. Longitudinal-oblique sections of annulated, selectively altered skeletons. Only the space between the whorls
in the proximal area is recrystallized, while the primary calcification is preserved in their distal parts (left in
both figures). Thin sections RR4584/8 and 4584/5.
Fig. 2. Longitudinal-oblique section of type B skeleton. Thin section RR4584/1.
Fig. 3. Longitudinal-oblique section of an annulated skeleton in which, in contrast to that one in Fig. 2, the whorls
with the primaries are completely recrystallized (Fig. 4/2A), while the space between the whorl is not filled;
arrows: thin encrusted stem membrane (right) and (left) recrystallized basal part of the space between the
whorls. Thin section RR4582.
Fig. 5. Tangential section corresponding to the skeleton shown in Fig. 2. Thin section RR4584/5.
Fig. 6. Slightly deformed longitudinal-oblique section of type A skeleton similar to that one shown in Fig. 3, poorly
preserved and slightly deformed. Thin section RR4584/2.
Fig. 7. Fragment of a longitudinal-oblique section of a type A skeleton; note the encrusted main stem membrane on
the right. Thin section RR4584.
Fig. 8. Oblique section of type A skeleton with encrusted stem membrane between the whorls. Thin section
RR4584/6.
Fig. 9. Oblique section of a recrystallized skeleton altered by endolithic activity; secondaries discernible in the upper
part (arrow). Thin section RR4583/3.
Scale bar for all figures = 0.50 mm.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 67
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PLATE 5
Dissocladella annulata (ELLIOTT, 1993) nov. comb.
Fig. 1. Fragment of a tangential section with pores of secondaries in the upper part. Thin section RR4583/1.
Figs. 2–6. Poorly preserved recrystallized oblique sections and fragments with pores of secondaries on the surface. Thin
sections RR4584/2, 4584/2, 4584/4, 4584/3 and 4584/9.
Figs. 7, 8. Tangential-oblique and shallow tangential section; “pores“ between whorls in Fig. 7 are in fact pseudopores.
Thin sections RR4584/2 and 4584/4.
Fig. 9. Oblique deep tangential section. Thin section RR4584/7.
Fig. 10. Tangential section with pseudopores (= not uniformly calcified space between whorls). Thin section
RR4583/5.
Scale bar for all figures = 0.50 mm.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 69
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PLATE 6
Dissocladella annulata (ELLIOTT, 1993) nov. comb., emended (Figs. 1–16), and associated organisms (Figs. 17–21).
Figs. 1–8, 10.Longitudinal and longitudinal-slightly oblique sections of different minute and small fragments of type A
skeletons. Thin sections RR4584, 4583/2, 4584/5, 4584/2, 4581, 4584/8, 4584/7, 4584/1, 4583/2 and 4584.
Fig. 9. Longitudinal section, fragment of type B skeleton. Thin section RR4583/4.
Fig. 11. Tangential section of a type A skeleton, corresponding to longitudinal section of skeleton in Fig. 10. Thin
section RR4583/2.
Fig. 12. Fragment, longitudinal section of a recrystallized type B skeleton with few open pores of secondaries. Thin
section RR 4583/4.
Fig. 13. Fragment of a slightly oblique transverse section with three pores of secondaries. Thin section RR4583/5.
Fig. 14. Fragment of recrystallized transverse section with denticulate surface. Thin section RR4584/4.
Fig. 15. Oblique section, note pores of secondaries . Thin section RR4586/5.
Figs. 16. Oblique section of a recrystallized skeleton affected by endolithic activity within the primaries. Thin section
RR 4583.
Figs. 17, 18. Microgastropods. Thin section RR4583/6.
Figs. 19–21. Sections of crustaceans – Carpathocancer SCHLAGINTWEIT & GAWLICK (former Carpathiella MISIK, SOTAK &
ZIEGLER). Thin sections RR4583/2, 4584/6 and 45 84/3.
Figures 1–15: scale bar = 0.25 mm; figures 16–21: scale bar = 0.50 mm.
Dissocladella annulata (ELLIOTT, 1993) nov. comb. (Calcareous algae, Dasycladales) from the Cenomanian of west Serbia 71
