GFF volume 120 (1998), pp. 365–371.
Article
Sediments and stromatoporoid morphotypes in Ludfordian (Upper Silurian) ree-
fal sea stacks on Gotland, Sweden
OLOF SANDSTRÖM
Sandström, O., 1998: Sediments and stromatoporoid morphotypes in
Ludfordian (Upper Silurian) reefal sea stacks on Gotland, Sweden.
GFF, Vol. 120 (Pt. 4, December), pp. 365–371. Stockholm. ISSN 1103-
5897.
Abstract: A study of the sedimentology and stromatoporoid morphol-
ogies in the Upper Silurian of southeastern Gotland reveals a strong re-
lationship between the depositional environment and the morphological
range of stromatoporoids. In high-energy environments, characterised
by packstones and grainstones, low-profile forms dominate, whereas
in calmer settings, characterised by wackestones and packstones, high-
profile stromatoporoids dominate. Four rock units are recognised, rep-
resenting two major facies: Reef-like limestones and Detrital grain-
stones. Reef-like limestones comprise a variety of lithologies including
wackestones, packstones, boundstones, and framestones. All units are
shallow marine, but the occurrence of crystal silt may indicate a vadose
situation on top of the reef-like facies. In the reef-like units, hard sub-
strates for stromatoporoid growth were probably provided by microbial
cementation. Sedimentation rates were generally low although episodic
high sedimentation occurred as indicated by raggedness in stromatopo-
roid skeletons. The stromatoporoid fauna is considered a low-diversity
assemblage dominated by Parallelostroma typicum.
Keywords: Stromatoporoid morphologies, carbonate petrography, ree-
fal carbonates, Silurian, Gotland, Hemse Beds Sweden
O. Sandström, Dept. of Geology, Lund University, Sölvegatan 13, SE-
223 62 Lund, Sweden, e-mail olof.sandstrom@geol.lu.se. Manuscript
received 11 July 1997. Revised manuscript accepted 2 November 1998.
Investigations of stromatoporoids and their shapes have shown
that several growth forms may have been induced by environ-
mental factors (Kershaw & Riding 1978; Bjerstedt & Feldmann
1985; Kano 1989, 1990; Kershaw 1990, 1997), enabling their
use in facies interpretations. Some species are confined to certain
growth forms by genetic controls, and are consequently confined
to certain sedimentary environments (cf. Kershaw 1990, 1997).
Petrographic studies of the sedimentary rocks enclosing the stro-
matoporoids give general and specific clues to the depositional
history. Stromatoporoid shapes and growth strategies may add
information regarding the nature of depositional patterns.
This paper is an attempt to highlight relationships between the
depositional history and stromatoporoid growth in a reefal, stro-
matoporoid-rich sea stack field called the Folhammarn nature
reserve (Fig. 1C) in the Upper Silurian Hemse Beds unit d (cf.
Fredholm 1988a, 1988b) of Gotland belonging to the Polygnath-
oides siluricus conodont biozone (Jeppsson 1983). Comparisons
are made with other known mid- to upper Hemse Beds reefal
localities.
The island of Gotland, situated in the Baltic Sea east of the
Swedish coast, consists of Silurian rocks ranging from Upper
Llandovery to Upper Ludlow and are dominated by limestones
and marls (mudstones with a large carbonate content), with a to-
tal thickness of about 500 m. The Silurian strata of the island are
currently subdivided into 13 topostratigraphical units (Fig. 1B),
mainly following the stratigraphy of Hede (1921, 1925) with the
oldest parts in the NW and the youngest in the S giving a gentle
dip towards the SE. The strata have not been subjected to any
greater compaction or heating.
Laufeld & Bassett (1981) described the overall Gotland geol-
ogy as a carbonate platform of limestone wedges with interrup-
tions by shaly marls. Later studies (Frykman 1989; Jacobsson
1997) envisage a ramp setting for the platform. Frykman (1989)
based this idea on the fact that no platform margin or strict sub-
division of shallow/deep-water facies can be recognised. Instead
there is a gradual transition from shallow to deep water facies.
Jacobsson (1997) describes a prograding ramp development
from the Wenlockian Slite Beds with initially clays and marls,
continuing with storm deposits and increasing carbonate content
as progradation and shallowing continues. The reefs and reef like
deposits developed most extensively in the most shallow parts of
the ramp. As progradation progressed several reef bands devel-
oped (cf. Fig. 1A) giving the present range and geographical ex-
tension of the reefal limestones. Jeppsson (1990) suggested a cy-
clic oceanic model which includes the distribution of marls and
reef limestones based on climate changes affecting deep sea bot-
tom currents. He divided the Gotland strata into P- and S- states,
where the P-state represents humid periods with shales and marls
and temperature stratified oceans, and the S-state represents dry
periods with limestones and reefs and black shales in basin ar-
eas elsewhere reflecting salinity stratification. The Fol-hammarn
area was probably deposited at the later stages of an S-episode
(Etelhem Secundo episode according to Jeppsson 1998).
Methods
Since the Folhammarn sea stack field is a nature reserve, only
a total of 30 samples with a maximum weight of 150 g/sample
was allowed. Thirteen are of the stromatoporoids, one is a coral
and 16 samples are of the surrounding sediments. Four sea stacks
(R1–4; Fig. 1C), reflecting three different facies, were selected
along the field where sampling and measuring took place. Thin
sections were made from all samples for petrographic and textur-
al analysis as well as species determination of stromatoporoids.
Limestone classification follows Dunham (1962). On each sea
stack a sketch of the rock surface at 1/5 reduction was drawn
using a 1×1 m frame with a 5×5 cm grid. Point counting was
carried out on these (1600 pts/sketch) and on thin sections of
the matrix (500 pts/sample). In this way both macro- and mi-
cro-components were obtained. To avoid over-representation
of medium-sized particles, only grains of 2 cm in diameter and
larger were sketched and, consequently, on thin sections only
grains smaller than 2 cm in diameter were counted. This method
was introduced by Kano (1989), and the results he obtained from
Gotland reefal deposits are summarised in Kano (1994) which
366 Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian GFF 120 (1998)

Fig. 1. A. Map of Gotland with the major

facies distribution of marls (grey), limestones
(white), silt and sand stones (light grey) and
reef and reefal rocks (black). (After Hadding
1941; Manten 1971; Eriksson & Laufeld
1978.) B. Hedeʼs (1921) topostratigraphical
units of the Silurian of Gotland. C. Detailed
map of the study area Folhammarn sea stack
field. The four studied sea stacks are marked
R1–R4. Locality names refer to Laufeld
(1974) and the Appendix.

also includes a discussion on the reliabil-

ity of the method.
From the four sea stacks, the base di-
ameter (B) and height (V) of a total of
87 whole stromatoporoid skeletons were
measured for morphotype classification
and plotted in V/B diagrams (cf. Kershaw
1990) in order to interpret the different
facies. To be able to measure a specimen
in the field, it must be displayed so that the
full basal diameter and height is visible.
All measured stromatoporoids are whole
and not eroded. Since measurements of
the skeletons were made in field, and due
to the limited amount of sampling, species
confined to certain shapes were not dis-
tinguished. For morphotype classification
Kershaw & Ridingʼs (1978) terminology
is used.

Results
Based on stromatoporoid shapes and pet-
rographic characteristics, four rock units
were distinguished: large stromatoporoid
packstones (LP), anastomosing and lami-
nar stromatoporoid packstones (ALP), fa-
vositid/stromatoporoid grainstones (FG)
and detrital grainstones and rud-stones
(DG). Of the four sea stacks, two expose
LP (R2 and R4), one ALP (R1) and one
FG (R3). DG is distinguished in the up-

Fig. 2. Photographs of the different facies and

special features of the Folhammarn area.
A. Crinoid stem parts in upright position (LP).
Photograph taken from above. B. Reef cav-
ity-filling including a cephalopod orthocone
(LP). C. Inter-reef packstones between small
stro-matoporoid buildups (b) (LP) and capped
by detrital grainstones (DG). D. Stromatopo-
roid-favositid bearing grainstones exhibiting
an eastern dip (FG). E. Detail of D, showing
that layering is marked by bands of stromat-
oporoids and favositids separated by coarse
bio-clastic grainstones (FG). F. Anastomos-
ing laminar stromatoporoid packstone (ALP)
that laterally grades into large stromatoporoid
packstone (LP). G. Framestone of anastomos-
ing laminar stromatoporoids (ALP).
GFF 120 (1998) Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian 367

Fig. 3. Photomicrographs of petrographic and

other features of the four units at Folham-
marn. A. Packstone with bands of rhodolithic
growth, unit LP. B. Packstone displaying
type B stromatactics at centre of image, unit
ALP. C. Rudaceous grainstone, unit DG. D.
Gir-vanella encrusting a stromatoporoid,
unit ALP. E. Rhodolithic intraclast, unit LP.
F. Vadose silt, partly dolomitized, unit ALP.
Scales equal 0.5 mm.

permost parts of all sea stacks except for

R3. Facies LP and ALP seem to be dis-
tributed in a patch like manner over the
entire sea stack field, although a vertical
zonation may be distinguished in places,
ranging from LP, ALP to DG in the lo-
cality Fågelhammar 1. Further north, in
Fågelhammar 2 ALP is absent. FG occurs
in one specific area in Fågel-hammar 5
forming an elongated limestone hill sepa-
rating the northern sea stack field from the
southern one.

Facies distribution
There are two main facies types repre-
sented in the Folhammarn area; reef-like
limestones and detrital grainstones. The
reef-like limestones are commonly fol-
lowed by grainstones at the top of most
sea stacks (cf. Fig 2C). However, the lat-
eral variation is more complex and each
unit displays lateral variations and com-
plex interactions and may even grade into
each other. Reef-like limestones are the
units LP and ALP. The LP unit shows a
variety of subfacies like extremely large
stromatoporoid colonies, small ʻpatch
reefsʼ (boundstones; Fig. 2C), inter-reef
packstones, reef cavity fillings (Fig. 2B;
intra-reef packstones) and crinoid mead-
ows (Fig. 2A; encrinitic packstones). The
ALP unit differs from LP only in that the
reefs are of laminar and repeated laminar
stromatoporoids (framestones; Fig. 2G).
Matrix and inter-reef sediments are here
packstones and grainstones. LP and ALP
are lateral equivalents and grade into each
other in a patch like manner (cf. Fig. 2F).
ALP is, however, confined to the southern
sea stack field (Fågelhammar 1). Fig. 2C
shows an example from R2, where small
stromatoporoid boundstone piles (b) are
separated by inter-reef packstones (unit
LP) which is overlain by rudaceous grain-
stones (DG). This succession is the most
common in the area.
Detrital grainstone facies are FG and
DG. FG comprises stromatoporoids and
favositids of autochthonous origin and
some are even displayed in situ giv-
ing a rather distinct layering that dips a
few degrees to the east (Fig. 2D–E). DG
comprises grainstones and rudstones of
predominantly broken stromatoporoids
and favositids together with crinoid frag-
ments. Layering may be present, but the
unit may also show diffuse characters and
may in places appear massive.
Large stromatoporoid packstones (LP). –
Greenish to bluish packstone consisting
of a micritic matrix, in places neomorph-
ically altered to microspar and granular
sparite. Grains are skeletal fragments of
mainly stromatoporoids, crinoids, corals
and bryozoans (Fig. 3A and E). Sporadic
grains of pyrite occur. Algae occur but are
not common.
In unit A stromatoporoid morpholo-
gies range from low domical to extended
domical (cf. Fig 4). Very large colonies
occur at a distance of 1–20 m from each
other. These colonies are between 1 and
4.5 m in base and up to 2.5 m high. They
display initial coalescence and do often
have ragged margins. Coral intergrowth
is common. Laminae may be enveloping
and non-enveloping within the same skel-
eton. Raggedness of the laminae as well
as sediment inclusions indicate frequent
episodic pulses of high sedimentation that
may have partly buried the colonies. In a
few colonies, large favositids are incorpo-
rated with the skeleton. Fig. 5 shows an
example of a typical giant stromatoporoid
colony with coalescence in its most initial
parts. The coalescent growth indicates a
simultaneous budding and an initial lat-
eral mode of growth, that later passed
into vertical growth. The uppermost parts
of the skeleton display non-enveloping
laminae. Thus, in the lower parts, growth
styles were mostly low domical and be-
came successively vertically extended in
its upper parts. As a result, the giant speci-
mens are generally high domical.
In places, dense accumulations of
domical stromatoporoids and favositids
occur forming small patch reefs (Fig. 2C).
These accumulations are distanced in a
similar manner as for the large solitary,
coalesced colonies. Within the accumu-
lations, cavity dwellers of trilobites and
bivalves as well as cephalopods (Fig. 2B)
368 Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian
Fig. 4. V/B plots of measured stromatoporoid skeletons from the three stromatoporoid bearing
subunits LP, ALP and FG.
may be found. Abundant occurrences of
crinoid holdfasts in the packstones (Fig.
2A) as well as locally rich accumulations
of crinoid stem parts (encrinites) indicate
occurrences of what was probably crinoid
meadows situated between large stromat-
oporoids and stromatoporoid accumula-
tions.
Anastomosing and laminar stromatopo-
roid packstones (ALP). – The ALP-unit
displays packstones (Fig. 3B, D, F), and
in a few samples, grainstones. The mic-
ritic matrix of the packstones is in places
neomorphically altered to micro-spar,
pseudospar and granular sparite. The col-
our of the rocks is greenish and reddish.
In thin section skeletal grains may be
coated with a hematite-impregnated ce-
ment. Skeletal particles are mostly from
crinoids, stromatoporoids, corals and, not
so commonly, algae and brachiopods. Py-
rite grains occur sporadically.
Stromatoporoid morphologies range
from low domical to laminar (cf. Fig. 4)
although some large high domical forms
also occur (cf. LP unit). Laminar forms
may locally exhibit anastomosing char-
acters forming framestones (Fig. 2G).
Almost all forms show ragged margins
and sediment inclusions. None of the
studied samples (N=23) exhibited coral
inter-growth or incorporation of tabulates.
Crinoid stem parts are common as well as
atrypid brachiopods.
Favositid/stromatoporoid grainstones
(FG) and detrital grainstones and rud-
stones (DG). – These units are grainstones
(Fig. 3C), grey to white, in places rud-
aceous with large skeletal debris. Layer-
ing is present in FG marked by bands of in
situ, mostly small (base diameter <1 m),
stromatoporoids and colonies of tabulate
corals (commonly favositids; Fig. 2D–E).
In thin sections from both units, particles
are mainly crinoid fragments although
fragments of stromatoporoids and cor-
als may be common. Algae are rare and
fragmented. The cement is of syntaxial
sparite around the crinoid fragments as
well as prismatic calcite. Within coral and
stromatoporoid skeletons, drusy spar and
equant spar are about equally distributed.
The stromatoporoids occurring within
FG are all small low profile forms (low
GFF 120 (1998)
domical and laminar; cf Fig. 4). They do
not display ragged margins or sediment
inclusions. They occur together with fav-
ositid tabulate corals in bands separating
different depositional events.
Petrography
Macro- and microscopic constituents of
the four units differ mainly on a higher
scale, i.e., reef-like limestones and detri-
tal grainstones (Fig. 6). Within the reefal
limestones stromatoporoids are the main
fossil constituent of the LP unit, whereas
in the ALP unit corals are a significant
skeletal constituent. The amounts of mic-
rite and sparite in these two units are com-
parable. The detrital units are dominated
by their high sparitic content, the skeletal
content is dominated by stromatoporoids
and crinoids. In the FG unit, a significant
portion is favositid corals.
Although the amount of algae was too
small to be seen in the point counting re-
sults, it is still important when interpreting
the petrographic analysis. The algae are of
two kinds and occur mostly in the reef-
like limestones as encrusters, intra-clasts
and binders. Encrusters have Gir-vanella
structure and binding forms are probably
rhodolithic algae (Fig. 3A, D, E). Both
types also occur as intraclasts and as pel-
loids. Girvanella is seen attached to stro-
matoporoids and is often found as a rim.
In thin section, rhodolithic microbials are
commonly seen as darker micritic molds
capping and binding organisms into intra-
clastic aggregates.
In the ALP-unit, internal sediment is
present deposited in small cavities (Fig.
3F) forming geopetal characters. The size
of the internal grains varies but most are
silt-sized although larger dolomite grains
occur. The sediment is here interpreted as
crystal silt and has so far only been found
within this unit.
Stromatoporoid fauna
Of the 13 stromatoporoid samples, 11
could be classified (two were strongly di-
agenetically altered). Three species were
identified; Parallelostroma typicum (8
samples), Stromatopora bekkeri (2 sam-
ples), and Pycnodictyon densum (1 sam-
ple). Eight samples were from LP, 3 from
ALP and 2 from FG. The stromatoporoid
species were distributed as follows; P.
typicum in units LP and ALP, S. bekkeri
in unit FG, and P. densum in unit ALP.
S. Kershaw (pers. comm. 1998) found a
specimen of S. bekkeri in unit LP as well.
GFF 120 (1998) Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian 369

Kano (1990) investigated unit ALP with respect to the stromat-

oporoid assemblage and found that it was dominated by P. typi-
cum with minor occurrences of Stromatopora venukovi, Petridi-
ostroma convictum and Ecclimadictyon robustum. Mori (1970)
noted P. typicum and E. robustum from this area as well. In total,
only six species is so far noted from the Folhammarn area, and
must therefore be considered a low diversity assemblage domi-
nated by one or two species (P. typicum and S. bekkeri). For each
unit alone, the diversity seems to be even lower.

Fig. 5. An idealised sketch of a very large stromatoporoid colony from

Interpretation and discussion unit A. The colony shows a three-stage growth with coalescent growth
initially. The second stage shows a lateral growth style, often exhibiting
Depositional environment ragged marigins and sediment inclusions. The final stage has a more
Occurrences of rhodolithic and girvanellid textures show that upright vertical growth style often with non-overlapping laminae.
sedimentation was very low between sedimentation events, in-
dicating a protected shallow environment. Also the fine-grained
micritic sediment, together with scattered occurrences of what
seem to be type B stromatactis, indicate a very quiet environment
with low wave and current action close to the sea floor (cf. Flajs
& Hüssner 1993). However, wave action may have been pro-
found at a somewhat higher level, indicated by the occurrence of
stemmed crinoids and higher-profile stromatoporoids. Bioturba-
tion of the sediment seems to have been low. Low bioturbation,
absence of microbial grazers and occurrences of stromatactis
plus high-profile stromatoporoids and stemmed crinoids may be
explained by quiet waters and salinity or oxygen stratification
closest to the sea floor.
Watts (1988) showed that inter-reef sediments from the Wen-
lockian Högklint Formation of Gotland displayed proximal and
distal parts with decreasing grain-sizes farther from the reefs. The
proximal parts display packstones and grainstones. Distal units
display wackestones and packstones. For the reef-like facies of Fig. 6. Constituents of the stromatoporoid-bearing units occurring at
Folhammarn. The upper bar represents macroscopic point counting of
Folhammarn, a grading is possible in the southern field (Fågel- sketched surfaces and the two lower represent microscopic point count-
hammar 1) with packstones and occasional grainstones, and in ings of the matrix of the units. S = stromatoporoids, T = tabulates, C =
the northern field (Fågelhammar 2) one sample is a wackestone corals, B = bryozoans, Ç = crinoids, b = brachiopods, M = molluscs, Ic
and the others are packstones. This suggests a southward proxi- = incertae. Black = micrite, grey = sparite, white = skeletal components
mality trend, supported by the fact that unit ALP occurs only in (only middle bar).
the southern field. This is the only unit that displays framestones.
In unit LP microbials are mostly found in situ in the southern
field and as transported and rounded intraclasts and aggregates in
the northern field. This further supports a southward proximality back-reef environment, such episodes of high sedimentation are
trend and a transport direction towards the N of material from a triggered by storms and hurricanes (e.g. Hubbard et al. 1991;
presumed larger body of organic buildups. Hubbard 1992; Li et al. 1997; Blanchon et al. 1997).
Kano (1990, 1994) suggested a stable substrate for the growth Vadose silt was recorded in the Klinteberg Formation reefal
of stromatoporoids in unit ALP, based on karstic erosion of unit environments (Frykman 1985). It is interpreted as a cavity accu-
LP. The present study, however, does not support a subaerial ex- mulation of eroded fines from the movement of meteoric waters
posure of the LP-unit. Indications of a vadose or karstic situation through the vadose zone (cf. Dunham 1969). Vadose silt is there-
were not found. Instead, cementation and lithification of the sub- fore formed at a distance below the zone of aerial contact and is
strate may have been induced by microbial activity of rhodolithic only an indirect evidence. The sediment in which it was formed
nature. Since such activity was not uniform over a distance, this may not have been subjected to aerial exposure. At Folhammarn,
could also explain the patchy nature of the occurrence and dis- vadose silt was found within unit ALP and this indicates that
tribution of boundstones, framestones and extremely large stro- the upper portions of that unit may have been subaerially ex-
matoporoid colonies, i.e., there was a mosaic of lithified and soft posed for some time before the re-entry to marine conditions
substrates and, consequently, limitations of the available hard with deposition of detrital grainstones making up a shallow shoal
substrates needed for colonization. However, there are several complex or a beach deposit.
localities in the Ludfordian of Gotland where a karstic situation
must have prevailed (e.g., Cherns 1982; Kano 1989; Keeling &
Kershaw 1994). Stromatoporoids
Total sedimentation rates must have been low to moderate, The shape and growth style of stromatoporoid colonies may give
as suggested by Kano (1990, 1994). However, there were epi- clues to the nature of the physical processes acting on the envi-
sodes of high sedimentation rates yielding ragged margins and ronment in which they lived (Kano 1989, 1990; Kershaw 1990,
sediment inclusions in stromatoporoid skeletons. In a protected 1997; Kershaw & Keeling 1994). Since some species are con-
370 Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian
fined to certain growth forms, these are also confined to environ-
ments suitable for that style of growth. Either the stromatoporoid
will have adapted to a change in the environment, or it will be
replaced by organisms more suitable for the altered situation.
Some features of the shape of stromatoporoids are always in-
dicative of physical processes acting on the environment (e.g.
ragged margins, sediment inclusions; Broadhurst 1966; Kershaw
& Riding 1978; Stearn 1982; Kano 1990), others are more un-
certain indicators due to species confinement and/or ontogenetic
behaviour of the individual (e.g. laminar, domical, etc.; Krebs
1974; Bjerstedt & Feldmann 1985; Kano 1989, 1990; Kershaw
1990, 1997).
The use of stromatoporoid morphologies in facies interpreta-
tion seems to work well for certain processes. The bulk morpho-
types will yield information about sedimentation and the general
water agitation. Together with specific characters of individu-
al skeletons and petrographical and textural data of the rocks,
coarse environmental reconstructions may be obtained.
There are several factors indicating periodic sedimentation
stress on the stromatoporoid populations occurring in reef-like
facies. Firstly, the occurrence of ragged margins and repeated
laminar forms together with sediment inclusions, secondly, the
scarcity of a reef framework, thirdly many specimens grew coa-
lesced and developed into very large colonies. The colonies often
grew separated from each other and dense clusters are scarce.
Finally, only six stromatoporoid species have been found, Paral-
lelostroma typicum (dominating), Pycnodictyon densum, Strom-
atopora venukovi, S. bekkeri, Petridiostroma convictum, and Ec-
climadictyon robustum (cf. Mori 1970; Kano 1990; S. Ker-shaw
pers. comm. 1998). Low diversity faunas dominated by only one
or a few species may indicate stressed habitats (Copper 1994).
The Kuppen area on Gotland shows signs of low sedimentation
rates (Kershaw 1993) and yields a more diverse fauna of 16
stromatoporoid species, where c. 40% of the specimens are of
one species (Kershaw 1990). This is, however, still considered
a low-diversity assemblage because many other reefs exhibit far
greater diversity.
Each of the sediment pulses may have partly buried the active
parts of the skeleton and the post-pulse phase is characterised
by lateral growth of the skeleton expanding over the new event
layer giving rise to ragged margins and/or repeated lamination.
However, recent research has shown that some sclerosponges are
able to expand their skeletons into free space (Stearn & Pickett
1994) and, therefore, raggedness may not always be an indica-
tion of sediment stress. Whether an individual will have survived
in a situation like this is dependent on three factors: ability to re-
move sediment from active tissue, the size of the sedimentation
pulse, and the duration between pulses. If the stromatoporoid
is unable to remove particles from the living tissue, even small
sedimentation pulses may be fatal. Ability to remove sediments
enhances the survival rate in habitats experiencing sedimenta-
tion stresses.
At Folhammarn, sedimentation rate and water energy are the
major controlling factors for the forming of morphotype varia-
tions and frequencies within a unit. Other factors may include
plasticity of the shape of the species within the investigated unit,
demography and preservation of whole specimens. The demo-
graphic pattern of the studied assemblage is important since most
species did not take their final form until they were fully grown
(cf. Stearn 1975). If the death rate is high after settlement and
during the growth phase this will tend to skew the assemblage
GFF 120 (1998)
toward lower V/B-values. High death rates before settlement
will not directly contribute to the assemblage since these are not
measurable in the field and may not have secreted calcite. Also,
the assemblage will be affected in an environment where frag-
mentation and/or truncation of the specimens is common since
such specimens are unmeasurable. Still, it is the environment that
will yield changes to the stromatoporoid assemblage that in turn
is shown by the frequency and distribution of morphotypes.
Coral intergrowth can be seen in, amongst other localities, the
Kuppen biostrome belonging to the middle Hemse Beds, and
the Ljugarn sea stack field of the same age as the Folhammarn
sea stack field. Whether coral intergrowth is a true symbiosis
or not is yet to be clarified (Kershaw 1987). At Folhammarn
coral inter-growth occurred only in the large stromatoporoid
specimens, and therefore it could be tempting to interpret it as
a growth enhancement mechanism for the stromatoporoids and
corals as well. However, coral intergrowth at Kuppen (Kershaw
1987) occurs in smaller specimens than in Folhammarn, and in
the Ljugarn sea stack field occurrences are from both small and
large stromatoporoid skeletons (own observations). Due to this
and the fact that there are no clear and direct advantages of inter-
growth for the stromatoporoids as seen in the fossil record, one
cannot conclude a true symbiosis.
Conclusions
The stromatoporoid fauna of Folhammarn is dominated by Par-
allelostroma typicum and is considered a low diversity assem-
blage. Stromatoporoid morphotypes differ significantly between
units and is probably due to differences in the sedimentary pat-
tern as well as wave energy (stress).
Very large stromatoporoids (>1 m in base) show signs of a
two-stage growth, beginning with a lateral mode of growth in-
terrupted by episodic sedimentation followed by a second stage
with vertical growth and non-overlapping laminae.
Coral intergrowth is common in larger specimens, but does not
seem to reflect a growth enhancement mechanism since inter-
growth has been reported from small specimens in other locali-
ties like Kuppen and Ljugarn.
Four different units, LP, ALP, FG, and DG were recognised
and grouped into reef-like (LP and ALP) and detrital (FG and
DG) facies. Three of the units are stromatoporoid bearing (LP,
ALP, and FG). All units are considered shallow marine. Geo-
graphically, the Folhammarn complex was marine and protected
and not far from shore, although it is not to be considered la-
goonal. Occurrence of vadose silt in the ALP-unit suggests a
subaerial contact at the top of the unit or above. Hard substrates
for stromatoporoid growth may have been provided by microbial
cementation instead of earlier suggested karstic processes (cf.
Kano 1990, 1994). Sedimentation rates were low in general, but
the influence of storms and hurricanes yielded episodic pulses
of high sedimentation indicated by the ragged margins and sedi-
ment inclusions of the stromatoporoid skeletons.
Acknowledgements. – Field work was partially financed through grants from Lunds Geolo-
giska Fältklubb. The Allekvia field station on Gotland is acknowledged for accommodation
support during field work. This work was accomplished during my time as a research student
at the Dept. of Geology, Historical Geology and Paleontology, Lund University. I wish to
thank Dr. Stephen Kershaw at Brunel University, London, for all support and ongoing dis-
cussions on stromatoporoid morphologies and Gotland reefs. Thanks also to Länsstyrelsen,
Gotlands län for permission to collect samples at the Folhammarn nature reserve. Prof. Kent
Larsson, Lennart Jeppsson and Anita Löfgren, all at Lund University, read and commented
on early drafts of the manuscript. Drs. Akihiro Kano and Stephen Kershaw refereed the
manuscript and made useful suggestions to the final appearance of this article.
GFF 120 (1998) Sandström: Sediments and stromatoporoid morphotypes in the Ludfordian
References
Bjerstedt, T.W. & Feldmann, R.M., 1985: Stromatoporoid paleosynecology in the Lu-
cas Dolostone (Middle Devonian) on Kelleys island, Ohio. Journal of Pale-ontol-
ogy 59, 1033–1061.
Blanchon, P., Jones, B. & Kalbfleisch, W., 1997: Anatomy of a fringing reef around
Grand Cayman: Storm rubble, not coral framework. Journal of Sedimentary Re-
search 67, 1–16.
Broadhurst, F.M., 1966: Growth forms of stromatoporoids in the Silurian of southern
Norway. Norsk Geologisk Tidsskrift 46, 401–404.
Cherns, L., 1982: Palaeokarst, tidal erosion surfaces and stromatolites in the Silurian
Eke Formation of Gotland, Sweden. Sedimentology 29, 819–833.
Copper, P., 1994: Reefs under stress: the fossil record. Courier Forschungsinstitut
Senckenberg 172, 87–94.
Dunham, R.J., 1962: Classification of carbonate rocks according to depositional tex-
ture. In W.E. Ham (ed.): Classification of carbonate rocks, 108–121. American
Association of Petroleum Geologists, Memoir 1.
Dunham, R.J., 1969: Early vadose silt in Townsend mound (reef), New Mexico. In
G.M. Friedman (ed.): Depositional environments in carbonate rocks, 139–181. So-
ciety of Economic Paleontologists and Mineralogists, Special Publication 14.
Eriksson, C.O. & Laufeld, S., 1978: Philip structures in the submarine Silurian of
northwest Gotland. Sveriges Geologiska Undersökning C 736, 1–30.
Flajs, G. & Hüssner, H., 1993: A Microbial Model for the Lower Devonian Stromatac-
tis Mud Mounds of the Montagne Noire (France). Facies 29, 179–194.
Fredholm, D., 1988a: Vertebrates in the Ludlovian Hemse Beds of Gotland, Sweden.
Geologiska Föreningens i Stockholm Förhandlingar 110, 157–179.
Fredholm, D., 1988b: Vertebrate biostratigraphy of the Ludlovian Hemse Beds of
Gotland, Sweden. Geologiska Föreningens i Stockholm Förhandlingar 110, 237–
253.
Frykman, P., 1985: Subaerial exposure and cement stratigraphy of a Silurian bio-herm
in the Klinteberg Beds, Gotland, Sweden. Geologiska Föreningens i Stockholm
Förhandlingar 107, 77–88.
Frykman, P., 1989: Carbonate ramp facies of the Klinteberg Formation, Wenlock-
Ludlow transition on Gotland, Sweden. Sveriges Geologiska Undersökning C 820,
1–79.
Hadding, A., 1941: The pre-Quaternary sedimentary rocks of Sweden. IV. Reef lime-
stones. Lunds Universitets Årsskrift N.F. 2, 37(10), 1–137.
Hede, J.E., 1921: Gottlands silurstratigrafi. Sveriges Geologiska Undersökning C 305,
1–100.
Hede, J.E., 1925: Beskrivning av Gotlands silurlager. In H. Munthe, J.E. Hede & L.
von Post: Gotlands geologi, en översikt, 13–30. Sveriges Geologiska Undersök-
ning C 331.
Hubbard, D.K., 1992: Hurricane-induced sediment transport in open-shelf tropical
systems – an example from St. Croix, U.S. Virgin Islands. Journal of Sedimentary
Petrology 62, 946–960.
Hubbard, D.K., Parsons, K.M., Bythell, J.C. & Walker, N.D., 1991: The effects of
Hurricane Hugo on the reefs and associated environments of St. Croix Islands – A
preliminary assessment. Journal of Coastal Research 8, 33–48.
Jacobsson, C.M., 1997: Storm deposition on a Silurian prograding carbonate ramp,
Slite Beds, Gotland. GFF 119, 199–206.
Jeppsson, L., 1983: Silurian conodont faunas from Gotland. Fossils and Strata 15,
121–144.
Jeppsson, L., 1990: An oceanic model for lithological and faunal changes tested on the
Silurian record. Journal of the Geological Society, London, 147, 663–674.
Jeppsson, L., 1998: Silurian oceanic events: Summary of general characteristics. In E.
Landing & M.E. Johnson (eds.): James Hall Centennial Volume. Silurian cycles:
Linking dynamic stratigraphy with atmospheric and oceanic changes, 233–251.
New York Geological Survey, State Museum Bulletin 491.
Kano, A., 1989: Deposition and palaeoecology of an upper Silurian stromatoporoid
reef on southernmost Gotland, Sweden. Geological Journal 24, 295–315.
Kano, A., 1990: Species, morphologies and environmental relationships of the Lud-
lovian (Upper Silurian) stromatoporoids on Gotland, Sweden. Stockholm Contribu-
tions in Geology 42, 85–121.
Kano, A., 1994: Quantitative compositions and reef development of the Silurian
limestones of Gotland, Sweden. Courier Forschungsinstitut Senckenberg 172,
141–146.
371
Keeling, M. & Kershaw, S., 1994: Rocky shore environments in the Upper Silurian of
Gotland, Sweden. GFF 116, 69–74.
Kershaw, S., 1987: Stromatoporoid-coral intergrowth in a Silurian biostrome. Lethaia
20, 371–380.
Kershaw, S., 1990: Stromatoporoid paleobiology and taphonomy in a Silurian bio-
strome on Gotland, Sweden. Palaeontology 33, 681–705.
Kershaw, S., 1993: Sedimentation control on growth of stromatoporoid reefs in the
Silurian of Gotland, Sweden. Journal of the Geological Society, London 150,
197–205.
Kershaw, S., 1997: Palaeoenvironmental change in Silurian stromatoporoid reefs,
Gotland, Sweden. Boletín de la Real Sociedad Española de Historia Natural (Sec-
ción Geológica) 91, 329–342.
Kershaw, S. & Keeling, M., 1994: Factors controlling the growth of stromatoporoid bi-
ostromes in the Ludlow of Gotland, Sweden. Sedimentary Geology 89, 325–335.
Kershaw, S. & Riding, R., 1978: Parameterization of stromatoporoid shape. Lethaia
11, 233–242.
Krebs, W., 1974: Devonian carbonate complexes of central Europe. In L.F. Laporte
(ed.): Reefs in time and space, 155–208. Society of Economic Paleontologists and
Mineralogists, Special Publication 18.
Laufeld, S., 1974a: Reference localities for paleontology and geology in the Silurian
of Gotland. Sveriges Geologiska Undersökning C 705, 1–172.
Laufeld, S., 1974b: Silurian chitinozoa from Gotland. Fossils and Strata 5, 1–130.
Laufeld, S. & Bassett, M.G., 1981: Gotland: The anatomy of a Silurian carbonate
platform. Episodes 1981 (2), 23–27.
Li, C., Jones, B. & Blanchon, P., 1997: Lagoon-shelf sediment exchange by storms-
Evidence from foraminiferal assemblages, east coast of Grand Cayman, British
West Indies. Journal of Sedimentary Research 67, 17–25.
Manten, A.A., 1971: Silurian reefs of Gotland. Elsevier. 539 pp.
Mori, K., 1970: Stromatoporoids from the Silurian of Gotland II. Stockholm Contribu-
tions in Geology 22, 1–152.
Stearn, C.W., 1975: The stromatoporoid animal. Lethaia 8, 89–100.
Stearn, C.W., 1982: The shapes of Paleozoic and modern reef-builders: A critical re-
view. Paleobiology 8, 228–241.
Stearn, C.W. & Pickett, J.W., 1994: The stromatoporoid animal revisited: Building the
skeleton. Lethaia 27, 1–10.
Watts, N.R., 1988: Carbonate particular sedimentation and facies within the Lower
Silurian Högklint patch reefs of Gotland, Sweden. Sedimentary Geology 59, 93–
113.
Appendix: Localities
The descriptions follow the recommendations by Laufeld (1974a). Further in-
formation on the localities Fågelhammar 1 and 2 is given in Laufeld (1974a and
1974b). Coordinates are given in the Swedish National Grid (Rikets Nät), printed
on the topographical map sheets.
FÅGELHAMMAR 1, 167661 636170, ca. 7550 m SW of Gammelgarn church.
Topographical map sheet 56D Ljugarn. Geological map sheet Aa 170 Kattham-
marsvik. Reference point: the pill-box. Hemse beds, upper part.
FÅGELHAMMAR 2, 167672 636210, ca. 7170 m SW of Gammelgarn church.
Topographical map sheet 56D Ljugarn. Geological map sheet Aa 170 Kattham-
marsvik. Hemse beds, upper part.
FÅGELHAMMAR 5, 167665 636203, ca. 7500 m SW of Gammelgarn church.
The cliff section between the northern and the southern sea stack areas ca. 255 m
NNE of the triangulation point at Folhammar. Topographical map sheet 56D Lju-
garn. Geological map sheet Aa 170 Katthammarsvik. Hemse beds, upper part.